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HARVARD UNIVERSITY
©
Library of the
Museum of
Comparative Zoology
Volume 135 MCZ 1992
Bonn LIBRARY
JAN 2 7 1993
Tijdschrift
voor
Entomologie
A journal of systematic and evolutionary
entomology since 1858
Published by the Nederlandse Entomologische Vereniging
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long
tradition in the publication of original papers on insect taxonomy and systematics.
The editors particularly invite papers on the insect fauna of the Palaearctic and
Indo-Australian regions, especially those including evolutionary aspects e.g.
phylogeny and biogeography, or ethology and ecology as far as meaningful for
insect taxonomy. Authors wishing to submit papers on disciplines related to
taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied
entomology, are requested to contact the editorial board before submitting.
Usually, such papers will only be published when space allows.
Editors
E. J. van Nieukerken (elected 1986) and J. van Tol (1985)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
Advisory board
M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (London).
The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.
Editorial address
c/o National Museum of Natural History,
Postbus 9517, 2300 RA Leiden, The Netherlands.
Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:
Nederlandse Entomologische Vereniging
c/o Instituut voor Taxonomische Zoölogie
Plantage Middenlaan 64
1018 DH Amsterdam
The Netherlands
Subscription price per volume Hfl. 300,— (postage included).
Special rate for members of the society. Please enquire.
Instructions to authors
Published with index of volume 135 (1992).
Graphic design
Ontwerpers B.V., Aad Derwort, ’s-Gravenhage
Tijdschrift voor Entomologie
Contents of Volume 135
Articles
141
145
163
169
27
43
87
S. J. Brooks & S. J. Richards
A new species of Oreagrion (Odonata: Coenagrionidae): montane damselflies from
New Guinea.
Chen, P. & N. Nieser
Gerridae, mainly from Sulawesi and Pulau Buton (Indonesia). Notes on Malesian
aquatic and semiaquatic bugs (Heteroptera), Ill.
Chen, P.: see Nieser
Decae, A. E.: see Helsdingen
Heller, K.-G. & K. Reinhold
A new bushcricket of the genus Poecilimon from the Greek Islands (Orthoptera:
Phaneropterinae).
Heller, K.-G.: see Willemse
Helsdingen, P. J. van & A. E. Decae
Ecology, distribution and vulnerability of Macrothele calpeiana (Walckenaer)
(Araneae, Hexathelidae).
Lansbury, I.
Notes on the marine-freshwater gerrid genus Rheumatometroides (Hemiptera,
Gerridae) of Papua New Guinea.
Nieser, N. & P. Chen
Revision of Limnometra Mayr (Gerridae) in the Malay Archipelago. Notes on
Malesian aquatic and semiaquatic bugs (Heteroptera), Il.
Nieser, N.: see Chen
Noreika, R. & R. Puplesis
Salicaceae feeding Gracillariidae (Lepidoptera) of Central Asia.
Pape, T.
Phylogeny of the Tachinidae family-group (Diptera: Calyptratae).
Pérez, T. M.
Feather mites (Acarina, Xolalgidae) in the uropygial gland tuft of Aratinga holochlora
(Sclater) (Aves, Psittacidae).
Puplesis, R.: see Noreika
ili
179 Ratcliffe, B. C.
New species and country records of Brazilian Cyclocephala (Coleoptera: Scarabaeidae:
Dynastinae).
Reinhold, K: see Heller
Richards, S. J. : see Brooks
191 Schouten, R. T. A.
Revision of the genera Euchromius Guenée and Miyakea Marumo (Lepidoptera:
Crambidae: Crambinae).
91 Smit, H.
Water mites from New South Wales and Queensland, Australia (Acari, Hydrachnellae).
113 Stock, J. H.
Pycnogonida from southern Brazil.
275 Vondel, B. J. van
Revision of the Palaearctic and Oriental species of Peltodytes Régimbart (Coleoptera:
Haliplidae).
299 Willemse, F. & K.-G. Heller
Notes on systematics of Greek species of Poecilimon Fischer, 1853 (Orthoptera:
Phaneropterinae).
Book Announcements and reviews
42 Block, W., 1992. An annotated bibliography of antarctic invertebrates (terrestrial and
freshwater). []. van Tol] * Heie, O. E., 1992. The Aphidoidea (Hemiptera) of
Fennoscandia and Denmark. IV. — Fauna Entomologica Scandinavica 25. [J. van Tol]. ©
Hansen, M., 1991. The Hydrophiloid beetles. Phylogeny, classification and a revision of
the genera (Coleoptera, Hydrophiloidea). [J. Huijbregts].
140 Skou, Peder, 1991. Nordens Ugler. Handbog over de i Danmark, Norge, Sverige,
Finland og Island forekommende arter af Herminiidae og Noctuidae (Lepidoptera). —
Danmarks Dyreliv 5. [E. J. van Nieukerken] * Cherepanov, A. |, 1991. Cerambicidae
of Northern Asia. Volume 3. Lamiinae. Part III.
Referees for volume 135
T. G. Brom (Amsterdam), E. Gittenberger (Leiden), M. Holmen (Helsinge), J. Huijbregts
(Leidschendam), R. de Jong (Leiden), P. Oosterbroek (Amsterdam), D. Vane-Wright
(London).
Dates of Publication
Volume 135 (1), pages 1-140, 15 July 1992
Volume 135 (2), pages 141-316, | December 1992
© Nederlandse Entomologische Vereniging, Amsterdam ISSN 0040-7496
volume 155, no. 1, 1772 ISSN 0040-7496
Tijdschrift
voor
i Entomologie
A journal of systematic and evolutionary
entomology since 1858
Published by the Nederlandse Entomologische Vereniging . sal u di
Tijdschrift voor Entomologie
A journal of systematic and evolutionary entomology since 1858
Scope
The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a long
tradition in the publication of original papers on insect taxonomy and systematics.
The editors particularly invite papers on the insect fauna of the Palaearctic and
Indo-Australian regions, especially those including evolutionary aspects e.g.
phylogeny and biogeography, or ethology and ecology as far as meaningful for
insect taxonomy. Authors wishing to submit papers on disciplines related to
taxonomy, e.g. descriptive aspects of morphology, ethology, ecology and applied
entomology, are requested to contact the editorial board before submitting.
Usually, such papers will only be published when space allows.
Editors
E. J. van Nieukerken (elected 1986) and J. van Tol (1985)
Co-editors
A. W. M. Mol (1990) and R. T. A. Schouten (1990)
Advisory board
M. Brancucci (Basel), N. E. Stork (London) and M. R. Wilson (London).
The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the
‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),
Amsterdam.
Editorial address
c/o National Museum of Natural History,
Postbus 9517, 2300 RA Leiden, The Netherlands.
Correspondence regarding membership of the society, subscriptions and
possibilities for exchange of this journal should be addressed to:
Nederlandse Entomologische Vereniging
c/o Instituut voor Taxonomische Zoölogie
Plantage Middenlaan 64
1018 DH Amsterdam
The Netherlands
Subscription price per volume Hfl. 300,— (postage included).
Special rate for members of the society. Please enquire.
Instructions to authors
Published with index of volume 134 (1991).
Graphic design
Ontwerpers B.V., Aad Derwort, ’s-Gravenhage
I. LANSBURY
MCZ
LIBRARY
Oxford University Museum, United Kingdom
NOTES ON THE MARINE-FRESHWATER GERRID
GENUS RHEUMATOMETROIDES (HEMIP TERA,
SEP 1 4 1999
HARVARD
UNIVERSITY
GERRIDAE) OF PAPUA NEW GUINEA
Lansbury, I., 1992. Notes on the marine-freshwater gerrid genus Rheumatometroides
(Hemiptera, Gerridae) of Papua New Guinea. — Tijdschrift voor Entomologie 135: 1-10, figs.
1-43, tables 1-4. assN 0040-7496]. Published 15 July 1992.
The marine-freshwater gerrid Rheumatometroides Hungerford and Matsuda from Madang
Province, Nagada Harbour were studied in February-March 1990 and the results compared
with data from West New Britain, Kimbe area. Rheumatometroides serena sp. n. is described
from Negada Harbour, and compared with À. brown: Hungerford and Matsuda, originally de-
scribed from Solomon Islands and now recorded from West New Britain. Variation between
females of R. browni from West New Britain and Solomon Islands is commented upon. Notes
are given on the habitat preferences of both species in Papua New Guinea and their localised
distribution compared with other marine bugs (Gerridae and Veliidae).
I. Lansbury, Hope Entomological Collections, University Museum, Oxford, OX1 3PW,
United Kingdom.
The genus Rheumatometroides Hungerford and
Matsuda (1958) was described from the Solomons
Islands, Kolombangara. No comment was made on
the habitat of this genus. E. S. Brown, who collected
the original series, describes the habitat in his field
notes as ‘surface of river running into lagoon (Jack
Harbour). Slow flowing’. Specimens were collected
on the Ist and 14th October, 1954. Matsuda (1960)
redescribed the genus comparing it with other genera
of the Trepobatinae Matsuda. Andersen and
Polhemus (1976) refer briefly to Rheumatometroides
and a closely related genus Stenobates Esaki, com-
menting that such records as there are indicate that
both are marine occurring in estuaries and nearshore
localities. Lansbury (1989) briefly summarises textual
references to date.
In 1990 I visited the Christensen Research
Institute (cri), Madang, Papua New Guinea and the
north coast of West New Britain, Kimbe area, to
study marine bug. A new species of
Rheumatometroides was found at Nagada Harbour
(cri) and À. browni Hungerford and Matsuda was
found in both freshwater and the intertidal zone in
West New Britain, and what appears to be a form
identical with that from Nagada, offshore in West
New Britain.
24 Christensen Research
Madang, Papua New Guinea.
' Publication nr. Institute,
Samples were collected both by handnet during
daylight and other samples were obtained using a
light trap run from the end of the CRI jetty.
Two immature stages of À. browni and a late instar
of the new species are briefly described.
SYSTEMATICS
Rheumatometroides serena sp. n.
(figs. 1-20)
Type material. — Holotype male: Papua New Guinea,
Madang Province, Madang, Nagada Harbour, collected
between 22.ii and 16.iii.1990, collected at light (mvlt) (in
OXUM). — Paratypes 39 d 32 © at mvlt, 43 d 16 9 under
jetty and in coral rubble close inshore (in oxuM, Forest
Research Institute, Lae, coll. Christensen Research Institute,
Madang, and RMNH Leiden)
Additional material. — West New Britain, Tamari, 100 m
offshore, 30.iv.1989. R. N. B. Prior, 1 d; West New
Britain, Tamari Beach, June 1989, R. N. B. Prior, 5 d 2 9.
— Immature stages: Nagada Harbour, 23.i1-14.i11.1990, 86
specs. at mvlt; 14 in coral rubble and under jetty; West New
Britain, Tamari, 100 m offshore, 30.iv.1989. 1 spec;
Tamari Beach, June 1989, R. N. B. Prior, 1 spec.
Description
Adult apterous. Males 2.72-3.1 mm long, maxi-
mum width 1.24-1.32 mm, females 2.68-3.0 mm
long, maximum width 1.44-1.60 mm.
Coloration. — Male: Head mesially with a broad
longitudinal ‘anchor-shaped’ bluish-black stripe ex-
tending from anterior margin almost to posterior
margin between eyes. Posterior margin adjacent to in-
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
LANSBURY: Rheumatometroides of Papua New Guinea
Figs. 10-20. Rheumatometroides serena sp. n. — 10-16, paratypes. — 10, suranal plate and genital segment, d ; 11, genital scler-
ites side view, d; 12, ventral aspect of sclerites, d; 13, ventral aspect of surannal plate processes, d ; 14, ovipositor side view,
9; 15, ventral aspect 2nd gonapophyses, ®; 16, side view 2nd gonopophyses, 9. Abbreviations Ga gonangulum, Pr base of
proctiger. Scale line 0.5 mm. — 17-20. Immature. — 17, dorsal aspect; 18, front leg; 19, antenna; 20, hind leg. Scale line 1
mm.
ner margin of eyes bluish black. Remainder of head
orange yellow. Pronotum bluish-black with a variably
shaped orange yellow blotch mesially. Mesonotum
bluish-black with two (1+1) orange yellow blotches,
mesially mesonotum appearing rather paler.
Metanotum and tergites bluish-black evanescent.
Eighth tergite yellowish brown, laterally bluish-black.
Suranal plate pale yellow, processes distally black.
Connexival segments 1-5 orange brown, remainder
bluish black.
Head. Ventrally yellow. Rostrum pale to dark
brown. Antennae black, proximal ventral margin of
Figs. 1-9. Rheumatometroides serena sp. n. paratypes. — 1, male dorsal aspect; 2, female dorsal aspect; 3, male abdomen ven-
trally, 4, male side view; 5, male (cleared in KOH and lactic acid); 6, male front leg; 7, male antenna; 8, female front leg; 9,
female antenna. Scale line 1 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
first segment pale brown. Pronotum laterally dark
brown-black, propleura and ventrally pale yellow.
Mesopleura bluish-black graduating ventrally to pale
yellow. Metathoracic spiracle orange. Dark abdomi-
nal coloration fading laterally, sternites pale yellow.
Metacoxal plates bluish-black.
Legs. Front leg, inner margin of femur and tibia
yellowish brown, outer margins of femur and tibia
black. Middle and hind legs black.
Female: Superficially resembling male, dark area
on head reduced. Tergites and connexivum tend to be
uniformly dark brown to black. Depending upon the
angle of filtered light on dry mounted specimens, the
dark brown-black coloration especially of the prono-
tum, mesonotum, metanotum and adjacent tergites
are partially covered by areas of pale bluish evanescent
coloration.
Structure. — Male: Elongate parallel sided (fig. 1).
Head length slightly greater than maximum width
across almost straight posterior margin between eyes.
Narrowest interocular space about 0.75 X posterior
width. Head sometimes slightly dilated in front of
eyes. Maximum eye width less than length and
0.66 X head length. Inner margin of eyes sinuate.
Rostrum 8 : 13 : 9. Antenna male (fig. 7) segment 1-
4322314615)
Pronotum 0.25 X longer than head. Antero-lateral
margins divergent, straight, postero-lateral margins
convergent, hind margin straight. Mesonotum just
over 2 median pronotal length, mesially with a shal-
low longitudinal depression becoming obsolescent
anteriorly, posterior margin concave. Metanotum
mesially about 0.4 X length of mesonotum with a
faint Y-shaped figure. First visible tergite clearly long-
er than remainder, ratios 5: 3: 1.5: 1.5 : 1.5. Eighth
tergite as long as preceding five, lateral margins
fringed with long hairs. Suranal processes curving
round segment, directed cephalad. Metacetabula par-
allel from posterior of mesonotum. Connexivum
strongly reflexed and folded onto dorsum.
Metasternum about 1.2 length of mesosternum.
Omphalium prominent, located midway between an-
terior and posterior margins (fig. 3). A male partially
cleared in KOH and examined in a 50% lactic acid
solution has extended part of thorax and abdomen.
Tergites are clearly visible and abdominal spiracles are
aligned pleurally. The intersegmental membranes are
narrow compared with tergal plates. The eighth seg-
ment bears a prominent spinose projection ventrally.
This segment is normally aligned so that dorsal sur-
face is continuous with 7th tergite and ventral projec-
tion partially encloses suranal plate segment (fig. 5).
Legs: Front femur broad, slightly curved (fig. 6),
flattened on inner surface, tibia more strongly flat-
tened with a prominent projection on inner distal
margin. Distal ventral margin of femur and outer dis-
Table 1. Proportions of leg segments of Rheumatometroides
serena sp. n.
Femur Tibia Tarsus I Tarsus II
Male front leg 35 31.5 3 13
Female front leg 35 30.5 3.5 15
Male middle leg 90 101 40 35
Female middle leg 87 102 40 35
Male hind leg 94 35 6 10
Female hind leg 91 35 6 11
tal margin of tibia with clusters of longer hairs. Tarsi
flattened, first very short, 2nd shining. Middle and
hind femora moderately robust. Proportions of leg
segments (table 1).
Genitalia: Suranal plate and processes large, dorsal-
ly plate pilose. Genital capsule partially enclosed by
lateral processes (fig. 10). Ventral aspect of suranal
plate and genitalia (fig. 13). Internal structures a con-
fusing array of paired sclerites (figs. 11, 12). Base of
phallus, a heavily sclerotised plate with slightly pro-
jecting rounded corners. Parameres small and sym-
metrical.
Female: Robust trapezoidal form (fig. 2). Head
length equalling maximum width across slightly co-
nave margin between eyes. Narrowest space 0.6 X
posterior width. Maximum eye width less than length
and about 0.35 X head length. Antennal segments 1-
AS 9) 27m ole 16:
Pronotum about 0.5 X longer than head, lateral
margins slightly rounded and diverging, posterior
margin evenly shallowly rounded. Mesonotum 3 X
median pronotal length, mesially with a prominent
longitudinal sulcus. Inner lateral margins of mesono-
tum curved inwardly, reflexed margins with numer-
ous hairs increasing in length and density distally.
Metanotum about 0.5 or less median length of meso-
notum, posterior margin shallowly concave.
Metanotum midway with a suture-like emarginate
line extending from inner margin of connexivum.
Visible tergites more or less subequal in length.
Metacetabula sharply convergent anteriorly.
Omphalium not conspicuous. Sternites increasing in
lengthrdistally,/3) 3.954 45h yor 317.
Genitalia (figs. 14-16). First gonocoxa triangular,
inner margin lightly sclerotised and reflexed inwards.
Gonangulum long originating near spiracle below
Table 2. Proportions of leg segments of Rheumatometroides
serena sp. n. Immature stages.
Femur Tibia Tarsus
Front leg 25 23 16
Middle leg 74.5 84 65
Hind leg 74.5 30 16
LansBury: Rheumatometroides of Papua New Guinea
Figs. 21-28. Rheumatometroides browni. — 21-24. Male Tamari Creek. — 21, dorsal aspect; 22, ventral aspect; 23, front leg; 24,
antenna. Scale line 1 mm. — 25-28. Female, Kapiura. — 25, ovipositor side view; 26, second gonapophysis; 27, side view of
first gonapophysis; 28, ventral aspect of first and second gonapophyses. Abbreviations Ga gonangulum; Pr proctiger; sp 8 8th
spiracle; Gx gonocoxa; ra ramus; s stylus; Gol Go2 first and second gonopophyses. Scale line 0.5 mm.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
proctiger, attached to first gonapophyses, which has
two (1+1) stylus-like projections basally, upper mar-
gin of first gonapophyses sclerotised. Second gona-
pophyses are longer than first and lightly sclerotised,
widely separated and joined by a membrane which is
raised lobe-like distally. Lateral margins reflexed bear-
ing on outer margin a row of blunt spines (figs. 15,
16). Attached to proximal end of 2nd gonapophyses,
a pair of lightly sclerotised ramus curving upwards
and expanded into a pair of lobes joined by a mem-
brane (fig. 16).
Immature stage 4th or Sth instar (figs. 17-20):
Head dark brown with a narrow pale yellow U-
shaped figure close to inner margin of eyes and across
posterior margin of head. Pronotum with two (1+1)
dark brown rectangular blotches. Mesonotum pale
yellowish brown with two (1+1) large almost oval
dark brown patches each with elongate pale yellow
blotch closely resembling adult pattern. Metanotum
yellow, laterally with an almost triangular dark brown
patch. Anteriorly to insertion of middle and hind legs
with variable triangular dark brown patches. Tergites
with pairs of dark brown patches. Abdominal spira-
cles placed medially. Mesonotum 3X median length
of pronotum and 2 X length of metanotum. Anterior
margin of first tergite deeply emarginate, tergites 2-5
subequal in length, tergites 6 and 7 clearly longer (fig.
17).
Front leg, femora more or less parallel sided resem-
bling that of adult female, tibia strongly produced
with a tuft of longer hairs distally (fig. 18). Hind fe-
mur moderately robust with an interrupted row of
spine-like projections, ventral margin with four tri-
chobothria-like hairs arising from circular areas de-
void of fine short hairs. Tibia and tarsus to a lesser ex-
tent with short fine hairs (fig. 20). Antennae (fig. 19):
first segment basally pale, remainder of first and oth-
er segments brown, segment 1-4 19 : 13.5 : 11 :
13.25. See also table 2.
Etymology. — À noun in apposition. This species is
dedicated to Mrs Serena Jebb, wife of the Director,
Dr. M. Jebb of the Christensen Research Institute,
Madang, P. N. G.
Remarks. — R. serena with R. agaaga Lansbury and
R. metawa Lansbury (Solomon Islands) form a dis-
tinct group based on the male front tibia, which is
strongly produced, whereas that of R. browni is not so
produced. The omphalium of À serena is in the cen-
tre of the metasternum as are all the foregoing insects.
According to Matsuda (1960) the omphalium of the
closely allied Stenobates is more conspicuous and lo-
cated at the intersegmental suture between the meso
and metasternum. Andersen (1982) distinguishes
Stenobates from Rheumatometroides by the former
having the metasternum distinctly produced, whereas
the latter is not produced.
Rheumatometroides serena is distinguished from À.
agaaga and À metawa (only males known) by the
colour pattern. The 3rd and 4th antennal segments of
R. metawa are almost subequal as are those of À. sere-
na, wheras R. agaaga has the 4th segment longer.
Rheumatometroides serena also resembles R. metawa.
However, R. serena has as shorter mesonotum 2 me-
dian pronotal length compared with R. metawa, mes-
onotum 2.75 X longer than median pronotal length.
The anterior suture between the meso and metaster-
num of R. metawa is antero-mesially produced, in R.
serena it is evenly rounded.
Rheumatometroides browni Hungerford and
Matsuda
(figs. 21-43)
Rheumatometroides browni Hungerford and Matsuda, 1958:
203-206. — Matsuda 1960: 351-353, Lansbury 1989: 63-
65.
Description
Adult apterous. Males 3.2-3.48 mm long, maxi-
mum width 1.2-1.36 mm, females 3.2-3.48 mm
long, maximum width 1.28-1.56 mm.
Coloration. — Male: Head, interocular space with a
median longitudinal shining black stripe, inner mar-
gins of eyes and front of head black, remainder pale
yellowish brown. Pronotum black with a median yel-
low blotch, lateral margins variably narrowly pale yel-
low. Mesonotum black with a large pale yellow W-
shaped area. Metanotum black with silvery
pubescence. Tergites black with silvery pubescence.
Eighth tergite basally broadly pale yellowish brown,
lateral margins and distally graduating to shining
black. Suranal plate dorsally brown, lateral processes
paler with tips darker. Propleura pale yellow, upper
margins of mesopleura black with slight silvery pu-
bescence, distal lower margin with brown stripe.
Metacetabula dark brown to black with silvery pubes-
cence and scattered pale yellow. blotches.
Connexivum pale brown basally, upper margins with
black markings (fig. 21). Underside of head, proster-
num and metasternum pale creamy yellow, sternites
like-colored, lateral margins suffused with brown-
black (fig. 22). Rostrum basally pale yellow graduat-
ing to black. Base of first antennal segment pale yel-
low, remainder of segments black. Legs, proximal
underside of front coxae, femur, middle and hind fe-
mora pale yellow, remainder dark brown-black.
Female: Not differing in broad details from male.
Tergites tend to be more densely covered with silvery
pubescence. Connexivum uniformly brown with
patches of silvery pubescence along outer margins.
Sternites uniformly pale creamy yellow.
Structure. — Male: Elongate parallel-sided (fig. 21).
Head length subequal to widest interocular space dis-
tally. Maximum eye width 0.75 X eye length and
about half median head length. Rostrum 12: 12: 10.
Pronotum 0.3X longer than median head length,
lateral margins rounded, median length about 2 later-
al margins. Posterior margin slightly emarginate.
Mesonotum mesially slightly depressed longitudinal-
ly. Metanotum depressed sloping caudad, less than
0.5 X median length of mesonotum, lateral margins
sinuate. Posterior margin with a narrow transverse
band of silvery pubescence. Tergites short with trans-
verse bands of silvery pubescence. Eighth tergite long-
er than metanotum, lateral margins sinuate and pilose
(fig. 32). Suranal plate short with a pair of processes
directed cephalad. Connexivum strongly reflexed and
folded over dorsum. Metacetabula diverging from
posterior margin of mesonotum. Mesosternum long,
metasternum 0.9X length of mesosternum.
Omphalium located centrally, lateral margins of met-
asternum pilose (fig. 22).
Genitalia: Abdominal segment 8 dorsally and later-
ally covered with fine hairs. Suranal plate densely pi-
lose, dorsally processes large, inner margin almost
straight. Ventrally segment enclosing capsule distad
pilose. Phallic organ is partially inflated (fig. 33).
Layout of structure broadly agrees with Brooks
(1960) interpretation (fig. 33). Sclerotised structure
within inflated organ is assumed to be dorsal plate of
vesica. Endosoma figured by Brooks (1960) is broad-
ly similar to fig. 34. Ventral plate attached to basal
plate and connected to dorsal sclerite of vesica.
Parameres symmetrical, small and rounded (fig. 35).
Female: Pronotum 0.7 longer than median head
length, lateral margins slightly rounded, short about
0.3 median length, posterior lateral margins sinuate.
Mesonotum 3 median pronotal length, postero-me-
dian margins straight, postero-lateral margins direct-
ed obliquely caudad against outer margin of metacet-
abula. Mesonotum mesially sulcate. Metanotum
triangular, broad distal margin covered with silvery
pubescence. Metanotum about 0.5 X length of mes-
onotum. Tergites with distal margins more or less
straight, five distal segments with variable areas of sil-
very pubescence. Metacetabula diverging from poste-
rior margin of mesonotum. Mesonotum length 7
length of metasternum. Omphalium close to anterior
margin of metasternum.
Genitalia (figs. 25-28). (Kapiura form): First geno-
coxa plate-like elongate, inner margin evenly curved,
outer margin slightly sinuate. Tergal margin nar-
rowed dorsally pilose (fig. 25). Gonangulum narrow,
heavily sclerotised, originating near dorsal margin of
gonocoxa. First gonapophyses extending beyond 2nd
gonapophyses, distally bluntly serrate along latero-
ventral margins. First gonapophyses with a pair of
LansBuRY: Rheumatometroides of Papua New Guinea
long stylus-like projections (figs. 25, 28). Second
gonapophyses basally attached to first gonapophyses
which has a spatulate projection ventrally (figs. 25,
27). Membrane between 2nd gonapophyses distally
turned upwards and slightly carinate mesially.
Proximally 2nd gonapophyses with a pair of elongate
ramus terminating in two (1+1) lobes (fig. 26).
Female genitalia (figs. 29-31) (form from
Kolombangara, Solomon Islands): First gonocoxa
plate-like, appearing almost parallel sided, inner mar-
gin reflex and heavily sclerotised, tergal margins even-
ly curved (fig. 29). Gonangulum narrow, heavily
sclerotised, attached to gonocoxa by an elongate
membrane. First gonapophyses sclerotised, proximal-
ly with two (1+1) stylus-like projections (fig. 29).
Second gonapophyses sclerotised, distally with a se-
ries of blunt projections. The two lobes of 2nd gona-
pophyses joined by a strong convoluted membrane,
distally bluntly acuminate. Proximally attached to
membrane, a pair of ramus terminating in two lobes
(figs. 30, 31).
Legs (table 3): Male front femur, lower margin
straight, inner surface of femur and tibia slightly flat-
tened, tibia slightly produced distally, tarsi flattened
(fig. 23). Middle femora more robust than hind fe-
mora. Antennal segments 1-4 male (fig. 24) 32 : 19.5
20.22 1-5; female 3): 192722)
Immature stages: Third (?) instar, partially cleared
and slide mounted (figs. 36-39): Head pattern most-
ly dark brown with two pale areas anteriorly, posteri-
or margin medially broadly pale brown. Antennae
light brown, internodial pieces visible between seg-
ments 2-4 with a continuous fringe of hairs along
outer margin (fig. 38). Pronotum largely dark brown,
eyes very slightly overlapping antero-lateral margins
of pronotum. Mesonotum with a complex pattern of
light and dark brown linear stripes. Metanotum with
two rectangular dark brown patches. First four ter-
gites with pairs of dark brown blotches, distal seg-
ments uniformly pale brown (fig. 36).
Legs: Front leg uniformly pilose, tibia not pro-
duced distally (fig. 37). Ventral margin of femur and
lateral margins of tibia with rows of prominent
spines. Hind leg (fig. 39) femur moderately robust
with a row of minute spine-like projections. Ventral
margin with four trichobothria-like hairs arising from
circular areas devoid of short fine hairs. Tibia and to
a lesser extent tarsi covered with short fine hairs.
Fourth-5th instar, partially cleared and slide
mounted (figs. 40-43): Posterior margin of head
broadly pale yellowish brown, anteriorly with two
(1+1) large triangular yellow blotches, remainder of
head dark brown. Eyes not overlapping antero-lateral
angles of pronotum. Distal lateral margins of head de-
pressed with two (1+1) trichobothrial pits. Antennae
brown, basal segment with scattered spines, fine hairs
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 29-43. Rheumatometroides browni. — 29-31. Female, Kolombangara. — 29, ovipositor side view; 30, 2nd gonapophysis;
31, ventral aspect of 2nd gonapophysis. Scale line 0.5 mm. — 32-35. Male, Kapiura. — 32, terminal segments side view; 33,
aedeagus; 34, internal sclerites; 35, paramere tilted aspect. Scale line 0.5 mm. — 36-43. Immature, Dami Creek. — 36, 40, dor-
sal aspect; 37, 41, front legs; 38, 42, antenna; 39, 43, hind legs. Scale line 1 mm.
Table 3. Proportions of leg segments of Rheumatometroides
browni Hungerford and Matsuda.
Femur Tibia Tarsus I Tarsus II
Male front leg 34.5 32 4 15
Female front leg 37 31 4 1177
Male middle leg 90 111 35 32
Female middle leg 94 118 37 36
Male hind leg 96 34 525 12
Female hind leg 102 40 7 13
Table 4. Proportions of leg segments of Rheumatometroides
browni Hungerford and Matsuda. Immature stages.
Femur Tibia Tarsus
Front leg 26 23 17
Middle leg 71 85 60
Hind leg 71 28 16
encircling distal end of segment. Remaining segments
covered with fine hairs. Internodial pieces not con-
spicuous as in earlier instar (fig. 42). Antennal ratios
1-4 are 20 : 12 : 12 : 14. Pronotal shape similar to
adult with two dark brown rectangles. Mesonotum
tending towards pattern of adult. Metacetabula
groove present along anterior margin close to posteri-
or margin of metanotum. Metathoracic spiracle on
lateral margin indicated by a dark brown spot.
Metanotum large and pale brown, antero-laterally
narrowly dark brown, posterior margin deeply con-
cave. Abdominal segments narrowly marked with two
(1+1) dark brown rectangular bars anteriorly. Legs:
front femur curved and pilose, distally with a cluster
of longer hairs. Tibia and tarsi covered with short fine
hairs, tibia slightly produced distally (fig. 41). Hind
leg moderately robust with four long trochobothria-
like hairs arising from circular areas devoid of short
fine hairs. Femur with a row of spine-like projections
(fig. 43). Proportions of leg segments, see table 4.
Remarks. — The series from West New Britain
compared with those from the Solomon Islands ap-
pear to be very similar externally, especially the colour
pattern and general external morphology. The differ-
ences noted in the female ovipositor and ancilliary
structures of the two populations (shape of gonocox-
al plate, length of paired styli, distribution of blunt
serrations and apparent absence of a ventral spatulate
projection on 2nd gonapophyses, as figs. 27, 29),
principally may indicate that the form from West
New Britain may be distinct from the Solomon
Islands form. The male genitalia appear to offer little
to support this hypothesis at present.
LANSBURY: Rheumatometroides of Papua New Guinea
MISCELLANEOUS BIOLOGICAL NOTES
Between 23 February and 16 March, R. serena was
collected on a number of occasions. The first speci-
men was recorded on 22 February, when a female was
collected from beneath the CRI jetty walkway at 5
pm. Subsequent searches for a number of days
starting at 7 am through to late afternoon failed to
find any more specimens in the intertidal zone and
more exposed areas, although occasional specimens
were seen most days in late afternoon in the shade of
the walkway. Scattered adult Halobates Eschscholtz
(Gerridae) and aggregations of Halovelia Bergroth s.
l. (Veliidae) (adults and immatures) were frequently
seen. Eventually numbers of À. serena were found in
the interstices of a coral rubble wall built to prevent
erosion (8-9 March). This species appears to shun di-
rect sunlight and exposed areas. A number of samples
were collected at light (mvlt) between 25 February
and 16 March. The light was run between 8-9.30 pm
from the end of the jetty over water always in excess
of two m deep. The largest number was found on 27
February (11 males, 6 females, 28 immatures), low ti-
de 1.2 m, high tide 1.45 m, and on 1 March (10 ma-
les, 14 females, 10 immatures), low tide 1 m, high ti-
de 1.5 m. It is not clear if the samples taken at light
came from the populations living amongst the coral
rubble, or another population which was living bene-
ath the main landing stage. This structure is sup-
ported by a series of piers made of tyres filled with
concrete. Between each tyre a layer of concrete was
used, which has numerous crannies. The area around
the landing stage provides shelter for many Halobates
and veliids. A mvlt sample collected on 13 March in-
cluded approximately 70 adult and 360 immature
Halobates and about 200 veliids. An aggregation ta-
ken on 1 March during daylight within one m of sho-
re included 200 immature Halobates, 500 Halovelia
adults and 1700 immature, but no
Rheumatometroides.
Colonies of Halovelia were set up in large tanks to
study their behaviour. Inadvertently small numbers
of R. serena were also introduced. Veliid mortality was
extremely high under these conditions, whereas the
gerrids survived. Adult À. serena were watched
grooming, the tip of the rostrum is ‘cleaned’ with the
tarsi of the front legs, longer middle legs are used to
groom the hind legs. At rest, the gerrid supports itself
on the tarsi. The first antennal segment is held curv-
ing outwards, the second and third are always aligned
in a more or less straight line, the fourth curved over
at internodial piece, the alignment forming a shallow
sickle-like outline. Live gerrids which were inadver-
tently transferred to diluted sea water quickly suc-
cumbed and drowned in the laboratory.
Rheumatometroides browni appears to be far more tol-
erant of lower salinities. Specimens were found up to
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
200 m upstream in West New Britain, Tamari Creek.
Habitat preferences from both regions (Solomon
Islands and West New Britain) seem to be similar, À
browni occurring in the zone where freshwater flows
into the sea. At Dami Creek, at low tide the creek ap-
pears not to receive any inflow of sea water, but the
freshwater continues to flow down the beach into the
sea (pers. obs.).
Rheumatometroides tends to skate across the surface
in a series of short fast spurts. Unlike Halobates, it
does not leap across the water to avoid the net. The
colony living amongst the coral rubble proved diffi-
cult to collect as individuals would run across the rub-
ble hiding in small cracks or getting behind the rub-
ble where the substrate was moist. Between 22
February and 16 March no pairs were seen in copula
and no females were seen ovipositing.
ACKNOWLEDGEMENTS
This work was carried out during the tenure of a
Fellowship from the Christensen Research Institute,
Madang, P. N. G. I especially wish to thank Dr.
Matthew Jebb, Director cri for his assistance during
10
my stay. Additional funds were provided via Prof.
David S. Smith, Hope Professor of Entomology ena-
bling me to visit West New Britain as a guest of Dr.
R. N. B. Prior, Kimbe to whom I am extremely grate-
ful for samples collected prior to my short visit.
REFERENCES
Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera
Gerromorpha). — Entomonograph 3: 1-455.
Andersen, N. M. & J. T. Polhemus, 1976. Water striders
(Hemiptera: Geriidae, Veliidae etc.). p. 187-224. - In: L.
Cheng (ed.), Marine insects. North Holland Publishing
Co.
Hungerford, H. B. & R. Matsuda, 1958. A new genus of the
Gerridae from the Solomon Islands. — Pan-Pacific
Entomologist 34: 203-206.
Lansbury, I., 1989. Notes on marine Rheumatometroides
from the Solomon Islands (Hem., Gerridae). —
Entomologist's Monthly Magazine 125: 63-71
Matsuda, R., 1960. Morphology, evolution and a classifica-
tion of the Gerridae (Hemiptera - Heteroptera). —
Science Bulletin University of Kansas 41 (2): 25-632.
Received: 15 October 1991
Accepted: 25 November 1991
NICO NIESER' & PINGPING CHEN’
'Tiel, The Netherlands
* Biological Control Laboratory, Chinese Academy of Agricultural Sciences, Beijing, China
REVISION OF LIMNOMETRA MAYR (GERRIDAE) IN
THE MALAY ARCHIPELAGO
Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), II
N. Nieser & P. Chen, 1992. A revision of Limnometra Mayr (Gerridae) in the Malay
Archipelago. Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), II. — Tijdschrift
voor Entomologie 135: 11-26, figs. 1-60, table 1 [ISSN 0040-7496]. Published 15 July 1992.
A key to the species of Limnometra in Indonesia and E. Malaysia and new distributional data
are presented. Nine new species of Limnometra are described: L. aploa, L. arachnis, L. lepta and
L. melanochroa from Sulawesi, L. genitalis from Pulau Buton, L. pseudoinsularis from
Kalimantan, L.hysterema from New Britain, L. monochroma from Guadalcanal and L. poliakan-
thina from Queensland (Australia).
Correspondence: Dr. N. Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.
Key words. — Sulawesi; Buton; Kalimantan; Papua New Guinea; Solomon Is.; Australia;
Gerridae; Limnometra; key; new species.
Representatives of the gerrine genus Limnometra
are restricted to tropical Asia, although three speci-
mens have been recorded from Tanzania (Hun-
gerford & Matsuda 1958). We presume that the spec-
imens from Africa are incorrectly labelled, since they
belong to a large species never collected in that area
again.
The present paper is mainly based on material re-
cently collected in Sulawesi, where several species oc-
cur as a frequent component of the surface fauna of
streams. When we started working up the material it
soon turned out that we had several new species at
hand. The examination of unidentified material from
Malesia in several museum collections revealed even
more species, so that we decided to report on this ge-
nus separately. A paper on the remaining Gerridae
from Sulawesi is in preparation.
MATERIAL AND METHODS
Specimens from Sulawesi were mainly collected by
J.P. Duffels (zma), J. van Tol (RMNH) and N. Nieser.
Localities sampled by Nieser have numbers starting
with N89 followed by two digits. All were taken in
Sulawesi Tenggara (SE Sulawesi) and Buton in 1989.
Details of localities can be found in Nieser & Chen
(1991). In addition, specimens from G. Zimmer-
mann's collection (zc), collected within the fra-
mework of Project Wallace 1985, and by Dr. I.
Lansbury (oxum) from other places in Southeast Asia
have been studied.
Although the number of species in the genus
Limnometra is apparently large, there are very few dis-
tinctive characters. The colour pattern on head and
thorax, which is usually very striking, is, apart from a
few exceptions, very uniform. The parameres are all
of the same shape, small, rod-like with the base slight-
ly broader than the apex. The general shape of the
genital segments is also quite similar in most species.
In various cases identification has to rely on relative
measurements, pilosity of the legs and annulation on
legs and antennae. In addition, we found that the en-
dosomal sclerites, especially the lateral pair, provide
specific characters in some cases. For this purpose the
endosoma has to be cleared, for which we used hot
10% KOH, and made free from the basal plate and
phallotheca which enclose it. As the endosoma has to
be studied from various angles, permanent micro-
scopic slides are less appropriate than keeping it in a
drop of glycerin in a microvial. There are usually four
sclerites, a medio-dorsal and a ventral one, which are
usually seen as one sclerite forming a wide arc within
the endosoma. Secondly, there is a symmetrical pair
of smaller lateral sclerites. We consider the open end
of the arc as posterior and the thicker posterior Y-
shaped part of the arc as dorsal, which corresponds to
the active condition of the endosoma.
Measurements are in millimeters, presented as the
range or the average value of five specimens (of each
sex). Measurements were taken from specimens of the
sample containing the holotype. In case of small sam-
ples additional specimens from other sites were used.
The width of the head is measured across the eyes, the
thoracic width is measured across the mesoacetabula.
11
TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Depositories
NC Nieser collection, Tiel, The Netherlands.
OXUM Oxford University Museum, England.
RMNH National Museum of Natural History,
Leiden, The Netherlands.
snow Snow Entomological Collections, Lawrence,
Kansas, U. S. A.
ZC Zimmermann collection, Marburg, B.R.D.
ZMA Zoological Museum, Institute of Taxonomi-
cal Zoology, Amsterdam, The Netherlands.
A pair of reference specimens of the Sulawesi spe-
cies was sent to Museum Zoologi Bogor (Indonesia)
and J. T. Polhemus collections, Englewood,
Colorado (U. S. A.), when sufficient material was
available.
Acknowledgements
Thanks are due to R. Brooks (snow), J. Duffels
(ZMA), I. Lansbury (oxum), J. van Tol (RMNH) and G.
Zimmermann (zc) for the loan of specimens.
‘TAXONOMIC PART
Limnometra Mayr belongs to the Gerrinae and
contains medium-sized to large species, which are
commonly seen on streams in Southeast Asia. Some
workers, e.g. Hungerford & Matsuda (1958), consi-
der it to be a distinct genus, while Matsuda (1960)
and Andersen (1982) rank it as a subgenus in
Tenagogonus Stal. Although Limnometra and
Tenagogonus are structurally very close, due to reduc-
tion of the abdomen in Tenagogonus, they can be kept
apart at first sight. For our present purpose the actual
taxonomic status is not important. Limnometra and
Tenagogonus can be distinguished from other SE
Asian Gerrinae by the following set of characters
(adapted from the key by Andersen 1982): Length
less than 25 mm. Pronotum with a dark median stri-
pe, venter generally pale. Third segment of rostrum
reaching mesosternum. Metasternum without lateral
scent channels (from scent orifice to metacetabula),
metathoracic spiracle located much more than its
own length away from base of fore wing.
The difference between Limnometra and
Tenagogonus is as follows (adapted from Hungerford
& Matsuda 1958):
1. Connexival spines normal, as in fig. 37 (except L.
minuta Mayr from Nicobar Islands). Male abdo-
men not reduced. Last ventral abdominal seg-
ment plus genital segments shorter than preced-
ing four segments. Hind coxae of male rarely
extending beyond middle of second ventral ab-
dominaliseement ea nee. Limnometra
— Connexival spines reduced. Male abdomen re-
duced. Ventral abdominal segments short, last
12
segment plus the genitalia at least as long as pre-
ceding four segments, usually longer. Hind coxae
of male nearly reaching or surpassing posterior
margin of second ventral abdominal segment .....
EL II rd, Tenagogonus
Key to males of Malesian Limnometra
1. Eight large black spots on venter (character holds
for female too), connexival spines shorter than
hrs genitaliserment. ee L. octopunctatus
— Usually only one black spot in front of meso-
acetabula, connexival spines longer ................. 2
2. Moderately small species. Head width usually
under 93 MANU NN A eee EN 3
— Larger species, head width usually over 1.8 mm
RR A ROS alice Fonsi 6
3. Front femur as slender as middle femur .......... 4
RAS AS atone Mor RU ee ate L. borneensis
4. Middle femur with a fringe of cilia about half as
Wide AS EM UE et a eee en L. bruneiensis
— Middle femur without a fringe of cilia (small
Spimes andisparse hairs present) i... ito 5
5. Lateral endosomal sclerite parallel-sided (fig. 44)
en di Nes rene Dda Re L. pseudoinsularis
— Lateral endosomal sclerite bulky in central part
(fig.cO) ern E. ae element. L. insularis
6. Antennae, middle and hind femur unicolorous,
thelattennotipalérneantipe®" ne ee ee: 7
— Antennae, middle and hind femur not unicolor-
ous, hind femur pale near tip, or some antennal
segments at least partly whitish ....................... 8
7. Ventro-posterior margin of first genital segment
distinctly produced laterally. Middle femur with
a short black line dorsally near base (holds in fe-
male too, but may be inconspicuous in specimens
withrdarksfemmun) set: oe eee ENS L. ciliata
— Rear margin of first genital segment with only a
faint production laterally. Middle femur without
the dorsobasal black line .................. L. cursitans
8. Middle femur with a long and curved blunt spine
at short distance from apex, two rows of small
PERS TE se teeters shaw tial: L. femorata
— Middle femur without a large curved spine near
apex (not to be confused with smaller sharply
Pointedrapicalispurs)er er ee en 9
9. Both middle and hind femur with a conspicuous
fringeroficilia atmen en unie BELA 10
— Hind femur without a conspicuous fringe of
Gilia: med. Rat a nein toa 11
10. Fore femur slender, ratio L/W over 20, without
subapical indentation, ventrally with a dense
fringe of cilia two thirds the width of femur … …
watt nennen tnt Ae L. lepta
— Fore femur broader, ratio L/W about 16, with a
distinct subapical indentation, ventral fringe of
cilia about half the width of femur .... L. pulchra
11. Fore femur not broader than middle femur ... 12
— Fore femur broader (1.3 X or more) than middle
feu ore 16
12. Fore femur ventrally with conspicuous dense pu-
bescencens dee aten etat ereen ede 14
— Fore femur without conspicuous pubescence 13
13. Segments of fore tarsus of subequal length (spe-
cies keyed out twice as its head width centres
around SMI) e nn L. bruneiensis
— First segment of fore tarsus distinctly (1.2X)
longernthan second) rer nn L. kallisto
14. General colour medium to dark brown, with pale
yellowish spots dorsally at sutures of connexival
segments; apical row of small pegs ranging over
more than half the middle femur ........................
A a 4 L. melanochroa
— General colour yellowish to light brown, sutures
of connexiva not marked by light spots; apical
row of small spines on middle femur short .... 15
15. Hind femur slightly (1.03 X ) longer than middle
femur, both longer than body ...... L. nigripennis
— Hind femur distinctly (1.1 X) longer than mid-
dle femur, which is slightly shorter than body ....
Ee ne fo L. submarginalis
16. Second antennal segment subequal (up to 1.1 X
asslongeas) swidthrofilead tse een adderen: 1%
— Second antennal segment longer (1.2X) than
widthiofheadbese e et L. arachnis
17. In ventral view first genital sternite (S8) about 1.5
times as long as second (pygophore, Pg): in dorsal
view S8 covering Pg completely, only tip of proc-
tigerä(Br)kvisible jes en. L. genitalis
— In ventral view S8 subequal to or shorter than Pg,
dorsally both Pg and Pr visible ...................... 18
18. Fore femur relatively slender, its length 10 times
its width, width of fore femur about 1.4X the
width of middle femur ............... L. annulicornis
— Fore femur stouter, its length about 7 times its
width, width of fore femur about 1.7 X the width
of middle femur … nn L. aploa
Limnometra annulicornis (Breddin)
(fig.1)
Gerris annulicornis Breddin, 1901: 83-85.
Limnometra annulicornis. — Hungerford & Matsuda 1958:
420-422, pl. 4 fig. 24, pl. 15, fig. 26.
Material. — Sulawesi Utara: Dumoga Bone NP.
Tumpah River, 900m, 23.11.1985, 3 9, J.P. Duffels, project
Wallace stat. 28 (zMa); Dumoga Bone N.P., waterfall creek,
tributary of Tumpah, UTM-XL0064, c. 225m asl,
23.iv.1985-B, 2 d,2 9, J. van Tol; same Tumpah river nr.
Edwards Subcamp, UTM-WL9365, c. 590 asl, 30.iv.1985,
1 9; same, 664m, at light, 3.vi.1985-A, 2 ©, J. van Tol;
same, brooklet c. 2 km W of Edwards subc., UTM-
WL9265, 4.vi.1985-A, 1 d 1 ®, J. van Tol (RMNH);
NIESER & CHEN: Revision of Limnometra
Dumoga Bone N.P., Toraut R., Maze, 7.xi.1985 3 d 1 9,
same 16/17.xi.1985, 6 d 6 9, G. Zimmermann; Tumpah
R., Sandinsel und Stromschnelle, 19.x.1985, 1 d 2 2; wa-
terfalls near Tumpah, 21.x.1985, 8 4, G. Zimmermann; 50
Km N Kotamobagu, thermal Quellen, 26.x.1985, 5 d 6 9,
G. Zimmermann (zc, NC ). Sulawesi Selatan: Maros 1 d 1
2 (RMNH).
Distribution. — Sulawesi.
Remarks. — Structurally very similar to L. arachnis
sp. n. which is, however, distinctly larger. For further
differences see under L. arachnis. The length of an-
tennal segment related to width of head is rather fluid
in this species. We have a male from Dumoga Bone
NP, Toraut R., Maze, 16.ix.1985 with width of head
2.0, 2nd antennal segment of left antenna 1.9 and of
right antenna 2.3 mm. The holotype has only one
row of small pegs at apex of middle femur. Most spec-
imens have, however, two short peg rows at apex of
middle femur of male.
Limnometra borneensis Hungerford & Matsuda
(fig. 2)
Limnometra borneensis Hungerford & Matsuda, 1958: 407-
408, pl. 2 fig. 12, pl.12 fig. 9.
Material examined. — Sarawak, Borneo, Mt. Pol, leg.
Mjöberg 1 d 1 ® paratypes. In addition: Mt. Tibang,
1200m, 1 ?, leg. Mjôberg 1925 (ZMA).
Distribution. — Previously only known by the type
series from Sarawak.
Remarks. — The female from Mt Tibang is slightly
larger than the paratypes, but seems to fit best with
this species. The spool-shaped fore leg in the male is
diagnostic. The lateral endosomal sclerites are also pe-
culiar in that they are posteriorly broadened and in-
cised (fig. 2). The posterior triangular part of prono-
tum seems to be somewhat shorter than in the very
similar L. insularis Hf. & Mats. We need, however,
longer series of fresh specimens of both species to ver-
ify whether this is a specific difference or is related to
development of indirect flight muscles.
Limnometra bruneiensis Miyamoto
(fig. 3)
Limnometra bruneiensis Miyamoto, 1967: 229-232, figs. 24-
29.
Material. — C. Borneo: Sg. Pijau, 1 d, leg. Mjöberg 1925;
Malaysia: E. Sabah, 70 km S Lahad Datu, Danum Valley,
main trail West 0 North 1-5, 150m, 7-9.xii.1989, narrow
creeks in rainforest, 2 d 1 @, leg. MJ. & J.P. Duffels sam-
ple Sabah 60 (zma).
Distribution. — N. and C. Borneo.
Remarks. — The types in Entomological Labora-
13
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Illasi ci 12
Figs. 1-10, Endosomal sclerites in Zimnometra, upper: lateral view of median arc and lateral sclerite, lower: different views of
lateral sclerite, scales 0.5 mm. 1, L. annulicornis, 2. L. borneensis (paratype), 3. L. bruneiensis, 4. L. ciliata, 5. L. femorata, 6. L.
insularis (holotype), 7. L. kallisto, 8. L. nigripennis, 9. L. octopunctata (one view of lateral sclerite only), 10. L. pulchra. — 11-
16: L. aploa (paratypes), 11-14 apex of abdomen, scale 2 mm, 11-12 ventral view, 11 d, 12 9, 13-14 lateral view, 13 d, 14
2; 15-16 endosomal sclerites, 15 median arc and lateral sclerite, left dorsal view, right lateral view, scale 0.5 mm, 16 differ-
ent views of lateral sclerite, scale 0.25 mm.
tory of Kyushu University were, unfortunately, not
available for study.
Limnometra ciliata Mayr
(fig. 4)
Limnometra ciliata Mayr, 1865: 44. — Hungerford &
Matsuda 1958: 409-412, pl. 3 fig. 14, pl. 13 fig. 22 (re-
description); Andersen 1964: 333.
14
Material. — + Viet Nam: Saigon (Than-Po Ho Chi
Minh), 1.vii.1924, 1 2 E. le Moult (RMNH). — ® Indonesia.
Sulawesi Tenggara: Sungai Anggoro, 20.ii, N8901, 1 9;
Kendari, 21.11, N8903 5 d 3 ®; Pool near Sungai Sampara,
22.ii, N8910 2 d 3 ® (2 ® apt); small stream near S.
Sampara, 22.11, N8912 1 9; Jalan Asera, 24.ii, N8916 9 d
17 ® (48 6 © apt); Jalan Paku Jaya, 24.11, N8920 2 9 (1
apt.); Tamborasi, 1.iii, N8925 1 d 1 2; Road to Mowewe,
3.111, N8933 3 8 1 2 (2 d 1 © apt); Teluk Kendari, 13.iii,
N8946 1 d 2 ®, 3 lvV; Teluk Kendari, 13.iii, N8949 2 9.
— P. Buton: Road to Lawele, 9.iii, N8939 3 ® apt. — Bali:
garden ponds, 8.viii.1983, leg. N. Nieser 2 d 1 9 apt. (NC
). — N.E. Sumatra tusschen Serdang en Tobameer (between
Serdang and Danau Toba), 2 ®, Dr. B. Hagen; Sum{atra],
Laula Slm., viii.1913, 2 2 apt., E. Jacobson; Sum. Kalung,
xii.1913, 3 d 5 @ apt, 1 2 macr., Le Moult; S. Coast,
Palabuhan Ratu, 4.xii.1976, 1 d 2 9, P. H. van Doesburg.
— NE Borneo, 1 d apt. — * Malaysia. Sabah, 16 km NE
Tenom Agr.Res.Stat. 115°59'E 5°11'N, TL, 180m,
11.vi.1986, 1 ® apt, leg. J. Huisman. — + Papua New
Guinea: West New Britain, Tamari, rain water tank,
14.1.1989, 2 d; West New Britain, Buluma nr. Dami, rain
water pit, 17.1.1989, 2 d 2 2; West New Britain, Blalla
Apulpu village, freshwater creek, 27.1.1989, 1 G ; Milne Bay
prov. , South of Alotau, Pandanus, shade, 29.ix.1989, 1 G,
all leg. R. N. B. Prior (oxum, Nc ). Macropterous unless oth-
erwise stated, most females with eggs.
Distribution. — Widespread. Thailand and
Malaysia through Indonesia and Philippines to
Guam, Solomon and Fiji Islands.
Remarks. — The males from Bali are even smaller
than the small specimens recorded by Andersen
(1964), body length 11.0 and 11.5 respectively.
Limnometra cursitans (Fabricius)
Gerris cursitans Fabricius, 1794: 192.
Limnometra cursitans. - Hungerford & Matsuda 1958: 413,
pl.5 fig.28, pl.14, fig.23.
Material. — Irian Jaya, Bernhard camp, 50m, vii.1938, 1
9, J. Olthof (Neth. Ind-American New Guinea Expedit.) (-
RMNH); Teluk Bintuni, Babo, 13.viii.1941, 1 2, E. Linquist
(NC ).
Distribution. — New Guinea and Australia
(Queensland).
Limnometra femorata Mayr
(fig. 5)
Limnometra femorata Mayr, 1865: 443. — Hungerford &
Matsuda 1958: 413-415, pl. 5 fig.27, pl. 13 fig. 21 (rede-
scription), Cheng & Fernando 1969: 106-107, fig. 72-
83.
Material. — NE Sumatra: Kuala Simpang, lowland forest,
iv.1954, 2 2; Seleh Kuala Simpang, lowland forest, iv.1954,
1 d 4 9, A. Sollaert (RMNH), all macropterous.
Distribution. — A widespread species: Malaysia
(Perlis, Selangor), Philippines, Borneo and Banguey.
New record for Sumatera.
Limnometra insularis Hungerford & Matsuda
(fig. 6)
Limnometra insularis Hungerford & Matsuda, 1958: 404-
NIESER & CHEN: Revision of Limnometra
405, pl. 4 fig. 21, pl. 11 fig. 17. - Cheng & Fernando
1969: 105-106, fig. 60-71.
Material. — We have studied the holotype from Sumatera
(Serdang, Tanjong Morawa, Hagen) and a paratype female
from Jawa (Arjeano) from sNow.
Distribution. — W. Malaysia, Sumatera and Jawa.
Two reported females, one from an island near
Borneo and one from Myanma may belong to a dif-
ferent species.
Remarks. — A small species, male length 11.8 mm,
width of head 1.64 mm. General colour light brown
with an orangeish tinge, with a striking pale yellow
band all along the lateral margin of pronotum.
Similar to L. borneensis, which has, however, a differ-
ent shape of front femur and lateral endosomal scler-
ites (figs. 3, 6). L. pseudoinsularis is also very similar,
only the form of lateral endosomal sclerites (fig. 44) is
quite distinct.
Limnometra kallisto (Kirkaldy)
(fig. 7)
Gerris kallisto Kirkaldy, 1899: 506.
Limnometra kallisto. — Hungerford & Matsuda 1958: 422-
424, pl.5 fig.31, pl.16 fig.27 (redescription)
Material. — ® Indonesia: Irian Jaya, Bernhard Camp B.,
100m, 13.iv.1939, L. J. Toxopeus (Neth. Ind.-American
New Guinea Exp.) 7 d 4 @ (RMNH); N. N. Guinea,
Pionierbivak, vi-vii.1920, W. C. van Heurn, 1 d; Beaufort
14.11.1913, Versteeg, 1 9 (ZMA). — * Papua New Guinea:
Milne Bay prov., South of Alotau, Pandanus, shade,
29.ix.1989, 1 d 1 2 leg. R. N. B. Prior (oxuM, Nc).
Distribution. — Misool (Mysol), New Guinea,
New Britain and Solomon Islands. Hungerford &
Matsuda (1958) place Misool in the Maluku.
However, biogeographically as well as both under
Dutch and Indonesian administration it belongs (c.
q. belonged) to New Guinea.
Limnometra nigripennis Mayr
(fig. 8)
Limnometra nigripennis Mayr, 1865: 443. — Hungerford &
Matsuda 1958: 415-418, pl.4 fig.26, pl.14 fig.24;
Andersen 1967: 262.
Material. — We have seen 1 d 1 ?, apterous from
Palawan (Philippines), reported by Andersen (1967).
Distribution. — Widespread in the Philippine
Islands, including the southern part of Mindanao. To
be expected in the islands north of Sulawesi.
15
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Limnometra octopunctata Hungerford
(fig. 9)
Limnometra octopunctata Hungerford, 1955: 67-68. —
Hungerford & Matsuda 1958: 400-401, pl. 4 fig. 22, pl.
10 fig. 13; Andersen 1964: 329; Cheng & Fernando
1969: 107-108, figs. 84-93.
Material. — Sulawesi Tenggara: Near Sungai Sampara,
22.11, N8912 1 2; Tamborasi, 1.iii, on the sea, between
boulders, N8925 1 9.
Distribution. — W Malaysia, NE Sumatera and
Sulawesi Tenggara (new record).
Limnometra pulchra Mayr
(fig. 10)
Limnometra pulchra Mayr, 1865: 443-444. — Hungerford &
Matsuda 1958: 418-419, pl. 4 fig. 23, pl. 15 fig. 25.
Material. — Maluku: Bacan Wayana, logged forest and
camp, alt. c 50m, 05/16.vi.1985, J. Huijbregts, 4 d 2 9 (-
RMNH); Buru, Station 13, 28.viii-4.ix.1921, L. J. Toxopeus
161 (zma).
Distribution. — Maluku. The record ‘Celebes;
Halmaheira’ by Hungerford & Matsuda (1958) is
confusing, as Halmaheira is not on Sulawesi, but one
of the islands of Maluku (the Moluccas). Actually
there are two records outside Maluku (Hungerford &
Matsuda 1958): the type locality ‘Java’ and 2 d 2 9
from ‘Jawa Gunung Gede(h), Frühstorfer’. The label
of the type seems to be a mistake of Mayr (1865) as
the type series bears labels ‘Amboina, Dr. Doleschal
1859’. Frühstorfer's labels are reported to be quite of-
ten incorrect. It is unlikely that this species occurs on
Jawa because it should have been collected more of-
ten.
Limnometra submarginalis (Miyamoto)
Tenagogonus (Limnometra) submarginalis Miyamoto, 1967:
226-229, figs. 18-23 [not available for study].
This species is only known by the type series from
N. Borneo. None of our specimens agrees with the
description.
Limnometra aploa sp. n.
(figs. 11-16)
Type material. — Holotype d: Sulawesi Tengah, SW
Luwuk, between Desa Seseba and Singsing Camp (+1°09'S
122°31'E), 80m, narrow stream in lowland rainforest,
14.x.1989, leg. J. P. Duffels (Sul12) (ZMA). — Paratypes,
same data as holotype, 10 4 9 2 (including allotype, 3 4
brach) (ZMA); Taki Lalang, small brooklet, 20.x.1989, 2 à 1
2, leg. J. vanTol. — Sulawesi Tenggara: Road Lalimboe km
4, 21.11.1989, N8903, 2 d 1 ©; Jalan Paku Jaya, 24.ii.1989,
16
N8920 4 d 3 2 (1 d 1 2 snow); Sungai Kolaka,
27.11.1989, N8921B, 1 d 1 ®; rd. Kolaka Kendari km 20,
stream, 3.11.1989, N8934 1 8 %; Sungai Mokowu
17.x.1989, 2 d 5 9, same, 30.x.1989, 1 à, leg. J.van Tol
(89JvT24); NE Kolaka, nr. G. Watuwila, Mokowu Camp
along Mokowu river, 200m, disturbed lowland rainforest, 3
3 4 2 (1 d brach), leg. J. P. Duffels (Sul.24). Mac-
ropterous unless otherwise specified.
Description
Macropterous specimens. — Dimensions. Length
3 10.8-12.0, 2 10.5-11.6; width of head & 1.80-
1.97, 2 1.80-1.92; humeral width of pronotum dé
1.95-2.20, 2 1.90-2.02; thoracic width d 3.15-3.50,
9 2190-3226:
Colour. Basic colour dark yellow to light dull
brown, head and thorax with the normal pattern of
blackish stripes. Antennae medium brown, apical
third of second segment yellowish, not very contrast-
ing. Fore femur and tibia light brown with variable
longitudinal medium brown stripes on inner and out-
er faces, tarsus darker. Middle and hind legs light to
medium brown, femora with yellow apical ring, not
very contrasting. Hemielytra medium brown, veins
dark brown to blackish. Venter pale yellow, whitish
ciliation dense on thoracic, loose on abdominal vent-
er.
Structural characters. Length of antennal segments
I: IT : III: IV, male 2.4 : 1.55 : 2.7 : 3.4, female 2.05
: 1.3 : 2.2 : 3.2. Second antennal segment shorter
(0.9) than width of head in male, more distinctly so
in female. Fore femur in male broader (1.5) than
middle femur, slightly flattened ventrally, no conspic-
uous pubescence, with wide, shallow subapical inden-
tation. Fore femur in female only slightly broader
than middle femur. Middle femur of male with a
fringe of cilia less (0.8) wide than femur, apically with
two short rows of small pegs, inner row with 4-6, out-
er row with 2-4 pegs. Middle femur in female and
hind femur in both sexes with short cilia only.
Thoracic and abdominal venter, including male geni-
tal segments with long cilia, more dense and obvious
in male than in female, looser on abdomen than on
thorax.
Male. Middle and hind femur slightly (0.90 and
0.94 respectively) shorter than body. Hemielytra just
reaching beyond the apices of connexival spines in
most specimens. Connexival spines more or less par-
allel and horizontal, reaching to or up to 0.25 of their
length beyond apex of genital segments. Median
length of last two abdominal sternites subequal,
shorter (0.85) than median length of first genital seg-
ment in ventral view. Endosomal sclerites figs. 15-16.
Female. Hind border of pronotum with a promi-
nent fringe of dark hairs (a few shorter ones present at
apex of pronotum in males). Middle and hind femur
distinctly shorter than body. Hemielytra reaching
halfway connexival spines. Last abdominal sternite
along median line longer (1.2) than penultimate ster-
nite and twice as long as genital sternites. Genital seg-
ments small, short, reaching at most halfway connex-
ival spines. Connexival spines long, distinctly
pointing dorsally (fig. 14), somewhat converging.
Brachypterous form. As macropterous, except for
slightly reduced pronotum, humeral width 1.8 and
hemielytra onto abdominal tergite 5.
Etymology. — Aploos (Greek adjective meaning
simple) refers to the lack of striking characters in this
species.
Comparative notes. — At first sight L. aploa looks
like small specimens of L. genitalis sp. n. Apart from
the smaller average dimensions, the genital segments
and female connexival spines are relatively smaller
and middle and hind femur shorter than length of
body in L. aploa. L. aploa differs from L. annulicornis
Breddin, also known from Sulawesi, by its smaller
size, the fourth antennal segment distinctly longer
(1.3 X longer than the first (subequal in L. annulicor-
nis), the connexival spines in the female pointing dor-
sally, and the shape of the endosomal sclerites (figs. 1,
15-16).
Limnometra arachnis sp. n.
(figs. 17-22)
Type material. — Holotype 4, Sulawesi Tenggara, rd.
Kolaka Kendari km 20, stream, 3.iii.1989, leg. N. Nieser
N8934 (RMNH). Paratypes, same data as holotype 9 d 7 9
(NC, 1 2 allotype RMNH); Sungai Mokowu, 30.x.1989, J.
van T'ol 1 d 3 2 (RMNH); NE Kolaka, nr. Gng. Watuwila,
Mokowu Camp along Mokowu River, 200m, disturbed
lowland rainforest, 29-31.x.1989, J.P. Duffels, Sul. 24, 1 d
3 2; Centipede Camp, (c. 3°49'S 122°31'E), nr. Gng.
Watuwila, 1100m, undisturbed hilly rainforest, narrow
stream, 2-5.xi.1989, J. P.Duffels, Sul. 30, 1 d; Sulawesi
Tengah: Sungai Batui, 17.x.1989, J.v.Tol 1 d 4 9;
P.Buton: Jismil camp, 18.X1.1989, J. vanTol, 1 d 2 ® (-
RMNH).
Additional specimens. — P. Buton, Jismil camp,
13.xi.1989, J. vanTol 1 d 1 2 (not made paratypes because
third antennal segment is also whitish in apical half and the
male is 15 mm long). All macropterous, most females with
eggs.
Description
Dimensions. Length d 16.5-17.5, 2 13.5-14.8;
width of head d 2.3-2.5, 9 2.08-2.12; humeral
width of pronotum d 2.8-3.0, 2 2.3-2.4; width
across mesoacetabula d 4.1-4.6, 9 3.6-3.8.
Colour. Basic colour dark yellow to light orange-
ish, posterior lobe of pronotum light brown, hemiely-
tra medium brown, head and thorax with the normal
pattern of blackish stripes. Antennae and legs light to
medium brown, fore femur and tibia with a pair of
NIESER & CHEN: Revision of Limnometra
distinct darker longitudinal stripes. Apical half of sec-
ond and apical five sixth of fourth antennal segments
and apices of middle and hind femurs and tibiae very
pale yellowish to whitish. Venter pale yellow, with
dense short whitish ciliation becoming longer and
more loose on abdominal venter, less distinct in fe-
male than in male.
Structural characters. Length of antennal segments
I: Il: Il: IV, male 4.1 : 2.6 : 3.5 : 3.6, female 2.7 :
1.7 : 2.5 : 2.9. Second antennal segment longer (1.08)
than width of head in male, shorter (0.8) in female.
Fore femur broader (1.3) than middle femur, slightly
flattened ventrally, with dense short pubescence and
wide, shallow subapical indentation (slightly less ob-
vious in female). Middle femur of male with a fringe
of cilia which is in the middle part about as wide as fe-
mur, apically with two short rows of small black pegs,
inner row with 9-12, outer row with 4-8 pegs. Middle
femur in female and hind femur in both sexes with
short cilia only. Hemielytra reaching beyond the api-
ces of connexival spines in males, just reaching con-
nexival apices in female. Thoracic and abdominal
venter, including male genital segments with quite
long cilia, more dense and obvious in male than in fe-
male, looser on abdomen than on thorax. Abdominal
venter with a distinct median carina over its whole
length. Connexival spines long, reaching well beyond
genital segments.
Male. Middle and hind femur slightly (1.05 and
1.1 respectively) longer than body. Connexival spines
more or less parallel and horizontal, reaching one
third of their length beyond apex of genital segments.
Median length of last abdominal sternites subequal to
first genital, shorter (0.65) than median length of pe-
nultimate abdominal sternite in ventral view.
Endosomal sclerites figs. 21-22.
Female. Middle and hind femur distinctly shorter
(0.8) than body. Last abdominal sternite along me-
dian line slightly longer (1.05) than penultimate ster-
nite and more than twice as long as the small, short,
genital segments. Connexival spines reaching half
their length beyond apex of genital segments, slanting
upwards by about 30°, (fig. 20), somewhat converg-
ing.
Etymology. — Arachnis (greek noun in apposition
meaning spider) refers to the spidery impression these
animals make when floating on the water.
Comparative notes. — Similar to L. annulicornis
Breddin also from Sulawesi which is, however, small-
er, has in male second antennal segment shorter than
width of head and middle and hind femur slightly
shorter than body. Females of L. annulicornis have the
genital segments relatively larger.
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 17-22. L. arachnis (paratypes), 17-20 apex of abdomen, scale 2 mm, 17-18 ventral view, 17 d, 18 ®, 19-20 lateral view,
19 d, 20 2; 21-22 endosomal sclerites, scale 0.5 mm, 21 median arc and lateral sclerites, above lateral view, below dorsal
view, 22 different views of lateral sclerite. — figs. 23-28. L. genitalis (paratypes), 23-28 apex of abdomen, scale 2 mm, 23-24
ventral view, 23 6, 24 ®, 25-26 lateral view, 25 d, 26 2; 27-28 endosomal sclerites, 27 median arc and lateral sclerites, scale
0.5 mm, above dorsal view, below lateral view, 28 different views of lateral sclerite, scale 0.25 mm. — figs. 29-34: L. lepta, 29-
32 apex of abdomen, scale 2 mm, 29-30 ventral view, 29 d (holotype), 30 9 (paratype), 31-32 lateral view, 31 d, 32 9; 33-
34 endosomal sclerites (paratype), 33 median arc and lateral sclerites, scale 0.5 mm, above dorsal view, below lateral view, 34
different views of lateral sclerite, scale 0.25 mm.
Limnometra genitalis sp. n.
(figs. 23-28)
Type material. — Holotype brachypterous 4 P. Buton,
cascade stream, 10.iii.1989, leg. N. Nieser N8942 (RMNH).
— Paratypes, same data as holotype: 5 & brach., 5 d 6 ©
macr. (1 2 allotype RMNH, 1 d 1 @ macr. ZMA); Buton, rd.
Bunga stream, 8.iii.1989, N8935 4 d brach., 10 d 8 ©
18
macr. (1 d macr. RMNH); small cascading stream near sea,
9.11.1989, N8941 1 d brach., 5 d 10 2 macr.; small
stream in wet forest, 10.111.1989, N8944, 9 & brach., 6 2
macr., all leg. N. Nieser (Nc, 1 d zma, 1 d 1 9 JTPC).
Description
Macropterous specimens. — Dimensions. Length
4 13.2-14.4, 2 12.1-13.4; width of head d 2.1-2.2,
® 1.9-2.1; humeral width of pronotum d 2.2-2.4, 9
2.1-2.3; width across mesoacetabula d 3.5-3.8, 9
D SD. 7e
Colour. Basic colour dark yellow with an orangeish
tinge, head and thorax with the normal pattern of
blackish stripes. Antennae medium brown, apical
third of second segment yellowish. Fore femur yellow
with longitudinal medium brown stripes on inner
and outer faces, tibia and tarsus medium brown.
Middle and hind femur light brown with yellow api-
cal ring. Middle tibia and tarsus medium brown, api-
cal quarter of middle femur usually lighter.
Hemielytra dull light brown, veins blackish. Venter
pale yellow with rather long and dense whitish cilia-
tion on thorax and abdomen, more prominent in
male than in female.
Structural characters. Length of antennal segments
IAE PIV male 2222; 31:35 female 5:
1.6 : 2.4 : 3.0. Second antennal segment slightly
shorter (0.95) than width of head in male, more dis-
tinctly so in female. Fore femur in male broader (1.5)
than middle femur, flattened ventrally, with thick-
ened pubescence, with wide, shallow apical indenta-
tion. Fore femur in female only slightly broader (1.1)
than middle femur, apical indentation indistinct.
Middle femur with a fringe of cilia about as wide as
femur in male, half as wide in female. Middle femur
of male apically with two short rows of 4-6 small pegs
each. Middle and hind tibia about as long as body or
hind femur slightly longer. Hemielytra just reaching
the apices of connexival spines in male, reaching half-
way in female.
Male. Connexival spines more or less parallel and
horizontal, reaching to or up to 0.25 of their length
beyond apex of genital segments. Median length of
last two abdominal sternites subequal or penultimate
slightly longer. Median length of first genital segment
in ventral view about 1.5 times median length of last
abdominal sternite (fig. 23). Endosomal sclerites figs.
27-28.
Female. Hind border of pronotum with a promi-
nent fringe of dark hairs (a few shorter ones present at
apex of pronotum in males). Last abdominal sternite
along median line longer (1.15) than penultimate
sternite and about four times as long as genital ster-
nites. Genital segments very small. Connexival spines
long, distinctly pointing dorsally (fig. 26) and con-
verging with tips touching or nearly touching in most
specimens.
Brachypterous form (males only). — Essentially as
macropterous but slightly smaller. Mean length mac-
ropterous 13.7, brachypterous 13.2. Humeral width
of pronotum just narrower than width of head (in
macropterous just broader). Hemielytra reaching
slightly over base to halfway last abdominal tergite.
NIESER & CHEN: Revision of Limnometra
Middle and hind femur slightly longer than body.
Etymology. — Genitalis (a latin adjective meaning
fertile, marked out to propagate) refers to the large
genital segments in male.
Ecology. — N89412, N8942 and N8944 are small
streams at shaded hilly sites in forests. L. genitalis was
found on quiet pools. N8935 is a larger more open
stream (over 10m wide), also in forest, with a small
artificial waterfall (probably to retain some water dur-
ing dry periods).
Comparative notes. — This species runs to L. annu-
licornis and L. kallisto with the key by Hungerford &
Matsuda (1958), but does not seem to be related. The
long first genital segment in males and the strongly
converging and upward pointing connexival spines in
females separate this species from other Limnometra.
Limnometra lepta sp. n.
(figs. 29-34)
Type material. - Holotype à, Sulawesi Utara, Dumoga
Bone N.P., Bank Toraut R. near Base Camp, 2.11.1985,
stat.6, lowland rainforest MV light trap, J. P. Duffels & J.
D. Holloway (Project Wallace). — Paratypes 5 2 same data
as holotype (ZMA, NC). All macropterous. Additional para-
types, Sulawesi Tengah, Sg Batui, 15.x.1989, J. vanTol,
89JvT024, 1 © (RMNH); Dumoga Bone N.P., Toraut R.,
Maze, 7-17.xi.1985, leg. G. Zimmermann 10 d 17 9
(ZC,NC, RMNH, SNOW). Strasse nach Malibagu Restaurant,
Bach, 18.xi.1985, 1 2 (zc).
Description
Dimensions. Length d 15.1-18.4, 9 13.5-14.7;
width of head d 2.3-2.6, 2 1.98-2.01; humeral
width of pronotum d 2.3-2.6, 2 1.95-2.2; width
across mesoacetabula d 3.6-4.3, 3.1-3.4.
Colour. Basic colour dark yellow to light dull
brown, thorax with the normal pattern of blackish
stripes. Dark M-shaped figure on head incomplete,
only lateral legs distinct. Antennae medium brown,
third and fourth segment largely pale yellowish with a
black spot on their articulation. Fore femur light to
medium brown, longitudinal darker stripes indis-
tinct. Middle and hind femur light tot medium
brown, lighter apical ring not apparent. Hemielytra
medium to dark brown, veins lighter, no costal or sub
costal light band. Venter pale yellow, in male with
whitish ciliation dense on thoracic, loose on abdomi-
nal venter.
Structural characters. Length of antennal segments
I: II: III : IV, male 4.9 : 4.0 : 5.1 : 5.0, female 2.8 :
2.2 : 3.0 : 3.3. Second antennal segment distinctly
longer (1.6) than width of head in male, less distinct-
ly so (1.1) in female. Fore femur without subapical
indentation, about as broad as middle femur, in male
ventrally, with dense ciliation, about half as wide as
19
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
femur. Fore femur in female, middle and hind femur
in both sexes, without obvious ciliation. Middle fe-
mur in male apically with a short row of 5-9 small
pegs at border of inner and dorsal surface. Hemielytra
reaching to or somewhat beyond the apices of con-
nexival spines.
Male. Middle and hind femur longer (1.1 and 1.2
respectively) than body. Connexival spines very
slightly diverging, horizontal or slightly pointing dor-
sad, reaching about half their length beyond apex of
genital segments. Median length of last abdominal
and first genital sternites subequal, shorter (0.75)
than penultimate abdominal sternite. Endosomal
sclerites figs. 33-34.
Female. Middle and hind femur distinctly (0.7)
shorter than length of body. Connexival spines par-
allel and horizontal to very slightly diverging and/or
pointing upward, reaching about one fifth of their
length beyond apex of abdomen. Median length of
last and penultimate abdominal sternite subequal and
about one third as long as genital sternites.
Etymology. — Leptos, a Greek adjective meaning
slender, refers to the relatively slender body of this
species.
Comparative notes. — In the key by Hungerford &
Matsuda (1958) this species runs to L. nigripennis,
but it is a much more slender species. From other
similar species L. lepta differs by characteristics of
width of fore femur and leg ciliation as indicated in
the key. The apex of the abdomen of the male is very
similar to that of L. annulicornis Mayr (Hungerford
& Matsuda 1958, pl.15 fig. 26), with which it forms
mixed populations in Dumoga Bone National Park.
L. annulicornis differs, however, in having a stouter
fore femur, middle femur distinctly ciliated and apex
of middle and hind femur with distinct light bands.
Limnometra melanochroa sp. n.
(figs. 35-40)
Type material. — Holotype brachypterous d Sulawesi
Tengah: Palu, 60 km SE: Lore Lindu N.P, Danau Tambing
and brooklets, UTM SJ95, 1600m asl, Pandanus,
7.xii.1985, J.van Tol (RMNH). — Paratypes same data as hol-
otype 5 d 5 ® (including allotype) brach.; Lore Lindu N.P.,
Rano Rano, 10 km NE Gimpu, 1600m, lower montane for-
est, 15.111.1985, 4 6 3 @ brach., 1 d 1 @ macr. J. P. & M.
J. Duffels, stat 42 (zma). — Sulawesi Tenggara: NE of
Kolaka, Centipede Camp, c. 3°49'S 121°40'E nr Gng
Watuwila, 1100m, undisturbed hilly rainforest, narrow
stream, 2-5.xi.1989, 4 d macr., leg. J.P. Duffels, stat. Sul.30
(ZMA).
Description
Brachypterous specimens. — Dimensions. Length
d 14.6-18.5, 2 13.6-16.0; width of head & 2.1-2.4,
20
9 1.9-2.1; humeral width of pronotum d 2.3-3.0, ®
2.2-2.5; width across mesoacetabula d 3.8-5.0, ©
3.5-4.0.
Colour. Basic colour medium to dark brown, head
and thorax with the pattern as usual in genus.
Antennae brown, without obvious lighter annulation.
Legs medium brown with darker stripes indistinct.
Middle femur with distinct, quite narrow, yellow ap-
ical ring. Apical part of tibiae and tarsi of middle and
hind legs yellowish. Hemielytra duil light brown,
veins with blackish margins and lighter centre.
Connexiva with pale yellowish patches dorsally at seg-
mental sutures. Thoracic venter yellowish, abdominal
venter light brown.
Structural characters. Length of antennal segments
I: IL: III: IV, male 3.8 : 2.5 : 3.35 : 3.2, female 2.7:
1.6 : 2.2 : 2.7. Second antennal segment subequal to
or longer than (1.0-1.2) width of head in male, short-
er (0.85) in female. Fore femur parallel sided, as wide
as middle femur, hardly flattened ventrally. Fore fe-
mur in male with a shallow apical indentation. Fore
and middle femur in male with dense long ciliation
ventrally, half as wide as femur on fore, as wide as fe-
mur on middle. Hind femur in both sexes and fore
and middle femur in female with short pubescence
only. Middle and hind femur slightly longer (1.1)
than body in male, shorter (0.85) in female. Middle
femur slightly longer than hind femur in most large
and slightly shorter in most small specimens. Middle
femur in male with a row of small pegs on apical two
thirds of inner surface, towards apex this row be-
comes double. Thoracic and abdominal venter with
dense longer cilia in male, short pubescence in fe-
male. Hemielytra reaching halfway tergite 7 in male
and halfway tergite 6 to anterior margin of 7 in fe-
male.
Male. Connexival spines medium sized, horizontal
and slightly divergent, reaching just beyond apex of
genital segments. Median length of last abdominal
sternite longer (1.4) than first genital sternite and
shorter (0.8) than penultimate abdominal sternite.
Endosomal sclerites figs. 39-40.
Female. Connexival spines medium sized, parallel
and hardly pointing dorsally caudally, usually reach-
ing to apex of proctiger. Median length of last ab-
dominal sternite subequal to penultimate sternite and
about two times as long as first gonocoxa.
Macropterous form. — Essentially as brachypterous
except for well developed prothorax, humeral width
of pronotum 1.3 times the width of head. Hemielytra
reaching beyond the apex of abdomen, leaving tips of
connexival spines uncovered in both sexes.
Etymology. — Melanochroos (a greek adjective
meaning dark skinned) refers to the dark general
colour compared to other species in the genus.
Ecology. — So far only known from altitudes over
1000m apparently in more or less undisturbed forest
in SE and Central Sulawesi.
Comparative notes. — Due to its dark colour and
pale markings on connexiva, L. melanochroa is, at first
sight, similar to some species of Aquarius, e.g. À. ven-
tralis (Fieber). Closer examination of colour pattern
and the position of the metathoracic spiracle, howev-
er, assign this species to Limnometra. By its coloration
it is not similar to any other Limnometra.
Limnometra pseudoinsularis sp.n.
(figs. 41-44)
Type material. — Holotype 4, C. Borneo, Sg. Birang,
Mjöberg 1925 (ZMA).
Description
Based on unique holotype d. — Dimensions.
Length 10.4; width of head 1.73; humeral width of
pronotum 1.68; width across mesoacetabula 2.62.
Colour. Basic colour pale brown. Dark pattern on
head indistinct. Dark pattern on thorax as usual in ge-
nus. Antennae with segments I, II, basal third of III
and IV light brown, apical parts of III and IV whitish.
Legs light to medium brown. Middle and hind femur
with rather indistinct yellow apical ring. Hemielytra
greyish brown, veins darker. Venter pale yellow.
Structural characters. Length of antennal segments
I: I: Ill: IV, 2.8 : 2.3 : 3.2 : 3.0. Second antennal
segment longer (1.3) than width of head. Fore femur
parallel sided, slightly narrower (0.9) than middle fe-
mur, hardly flattened ventrally, with short denser pu-
bescence without apical indentation. Middle and
hind femur with short pubescence only. Middle fe-
mur apically with 2 rows of 5-6 small pegs on inner
surface. Middle femur slightly shorter (0.96) than
body, hind femur subequal in length with body.
Venter with sparse longer cilia on last abdominal ster-
nite and genital segments only. Hemielytra reaching
well beyond the apices of connexival spines.
Connexival spines long, parallel and horizontal reach-
ing a third of their length beyond the apex of genital
segments. Median length of last and penultimate ab-
dominal sternite subequal slightly longer (1.08) than
first genital sternite. Endosomal sclerites figs. 43-44.
Etymology. — Pseudoinsularis a Latin adjective
meaning ‘false-insularis’ refers to the similarity to L.
insularis.
Comparative notes. — Very similar to L. insularis.
The most distinct difference is in the lateral endoso-
mal sclerite, which is parallel-sided in L. pseudoinsu-
laris, and with a thickened central part in L. insularis
(figs. 44, 6).
NIESER & CHEN: Revision of Limnometra
Limnometra monochroma sp.n.
(figs. 51-56)
Type material. — Holotype d Papua New Guinea: West
New Britain, Buluma nr. Dami, rain water pit, 17.1.1989,
leg. R. N. B. Prior (oxum). — Paratypes, 1 d 1 9 same data
as holotype (Nc); West New Britain, Von River, Banaule
Village, torrential habitat, 21.11.1990, leg. I. Lansbury, 2 9
(OXUM).
Description
Dimensions. Length d 11.2-11.9, 2 11.2-11.4;
width of head d 1.83-1.84, 2 1.78-1.88; humeral
width of pronotum d 1.90-1.92, 2 1.83-2.0; width
across mesoacetabula d 2.82-2.87, 9 2.9-3.1.
Colour. Basic colour yellow to light brown, dorsum
of abdomen reddish. Head and thorax with the nor-
mal pattern of blackish stripes. Dark M-shaped figure
on head incomplete, only lateral legs distinct (except
for 1 2 from Von River, which is much darker than
the other specimens, having a complete M-figure).
Antennae medium brown, without whitish parts. Fore
femur light brown with a dorsal and ventral longitudi-
nal medium brown, stripe. Middle and hind femur
medium brown, rather narrow lighter apical rings dis-
tinct. Tibiae and tarsi brown, tibiae yellowish in apical
part. Hemielytra medium brown with darker veins, no
costal or sub costal light band. Venter yellow.
Structural characters. Length of antennal segments
I: I: III : IV, male 2.5 : 1.7 : 2.3 : 3.2, female 2.2:
1.4 : 1.8 : 2.9. Second antennal segment distinctly
shorter ( d 0.9, 2 0.8)) than width of head. Fore fe-
mur of male broader (1.6) than middle femur, with a
wide, shallow, subapical indentation and with short,
dense pubescence ventrally. Fore femur in female
slightly (1.3) wider than middle femur, with an indis-
tinct subapical indentation, without obvious cilia-
tion. Hemielytra reaching about halfway the apices of
connexival spines. Ventral abdominal carina distinct
on sternites 3-7 in male, and on 2-6 in female.
Male. Middle femur slightly shorter (0.9) than
body, with a fringe of cilia about half as wide as femur
and two short rows of 4-6 small pegs at border of in-
ner and dorsal surface. Hind femur subequal to body
length. Connexival spines parallel, virtually horizon-
tal, reaching about half their length beyond apex of
genital segments. Median length of last abdominal
shorter (0.8) than penultimate abdominal sternite.
Genital sternites subequal, shorter (0.6) than last ab-
dominal sternite. Endosomal sclerites figs. 55-56.
Female. Middle and hind femur distinctly (0.8)
shorter than length of body. Connexival spines par-
allel and slightly pointing upward, reaching about
one third of their length beyond apex of abdomen.
Median length of last and penultimate abdominal
sternite subequal and somewhat (1.1) longer than
genital sternites.
21
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
22
Etymology. — Monochromos, a Greek adjective
meaning unicolourous, refers to the unicolourous an-
tennae of this species.
Comparative notes. — In the key by Hungerford &
Matsuda (1958) and our key this species runs to Z.
annulicornis, from which it differs by having the an-
tennae unicolourous. Males differ, moreover, by the
second antennal segment relatively shorter compared
to the width of head and hind femur longer than mid-
dle femur (subequal in L. annulicornis). From other
similar species L. monochroma differs by characteris-
tics of width of fore femur and leg ciliation as indicat-
ed in the key.
Limnometra poliakanthina sp. n.
(figs. 57-60)
Type material. — Holotype d: Australia, Queensland,
Tulatten, rainforest creek, 21.v.1979, 0/0138, I. Lansbury
(oxum). — Paratype: 1 d, same data as holotype (Nc).
Description
Dimensions. Length d 17.0-17.9; width of head
2.41-2.42; humeral width of pronotum 2.78-2.96;
width across mesoacetabula 4.33-2.46.
Colour. Basic colour yellowish to light brown.
Interoculus unicolourous, without M-shaped figure.
Thorax with the normal pattern of blackish stripes,
which is not very dark. Antennae medium brown,
without whitish parts. Venter and legs about the same
colour as rest of body, darker stripes indistinct.
Middle and hind femur with lighter apical annula-
tion, which is not strongly contrasting. Hemielytra
dull medium brown with shining castaneous veins,
no costal or sub costal light band, darker hind wings
shining through.
Structural characters. Length of antennal segments
I: IT : III: IV, 3.5 : 2.25 : 3.1 : 3.8. Second antennal
segment distinctly shorter (0.9) than width of head.
Fore femur broader (1.3) than middle femur, with a
shallow, subapical indentation, ventrally flattened
without obvious pubescence. Hemielytra reaching
NIESER & CHEN: Revision of Limnometra
beyond the apices of connexival spines. Ventral ab-
dominal carina indistinct. Middle and hind femur
subequal, longer (1.07) than body. Middle femur
with short cilia and setae and a double row of small
blackish pegs along its entire length except for basal
0.15 part, where there is a single row. Connexival
spines parallel to slightly diverging, virtually horizon-
tal, reaching about half their length beyond apex of
genital segments. Median length of last abdominal
shorter (0.85) than penultimate abdominal sternite.
First genital sternite shorter (0.7) than last abdominal
sternite and shorter (0.75) than last genital sternite.
Endosomal sclerites figs. 59-60.
Etymology. — Poliakanthinos, a Greek adjective,
meaning with many spines, referring to the double
row of pegs on the middle femur.
Comparative notes — With the key by Hungerford
& Matsuda (1958) and our key this species runs to L.
annulicornis, which has, however, short rows of pegs
on the middle femur. The only other species with a
long partly double peg row on the middle femur is L.
melanochroa, which has, however, a different general
colour pattern and is more solidly built.
Limnometra hysterema sp.n.
(figs. 45-50)
Type material. — Holotype d: Solomon Islands,
Guadalcanal, Tsarivonga R., 22.10.1955, E. S. Brown (ox-
UM). — Paratypes: Guadalcanal, Honiara District, Mt.
Parsten, 13.vi.1964, E. S. Brown, 2 2 (oxuM).
Description
Dimensions. Length d 15.0, 12.8-13.0; width
of head d 2.14, 2 1.96-2.00; humeral width of pro-
notum d 2.40 2 2.17-2.20; width across mesoaceta-
bula/d35042.39255332:
Colour. Basic colour light brown, dorsum of abdo-
men with a reddish tinge. Head and thorax with the
normal pattern of blackish stripes. Dark M-shaped
figure on head partly indistinct. Antennae unicolor-
Figs. 35-40. L. melanochroa (paratypes), 35-38 apex of abdomen, scale 2 mm, 35-36 ventral view, 35 d, 36 2, 37-38 lateral
view, 37 6, 38 2; 39-40 endosomal sclerites, 39 median arc and lateral sclerites, scale 0.5 mm, above lateral view, below dor-
sal view, 40 different views of lateral sclerite, scale 0.25 mm. — figs. 41-44. L. pseudoinsularis, 3 (holotype), 41-42 apex of ab-
domen, scale 2 mm, 41 ventral, 42 lateral view; 43-44 endosomal sclerites, 43 median arc and lateral sclerites, scale 0.5 mm,
above lateral, below dorsal view, 44 different views of lateral sclerites, scale 0.25 mm. — figs. 45-50: Z. hysterema, 45-48 apex
of abdomen, scale 2 mm, 45-46 ventral view, 45 d (holotype), 46 9 (paratype), 47-48 lateral view, 47 8,48 9 ; 49-50 en-
dosomal sclerites (holotype), 49 median arc and lateral sclerites, scale 0.5 mm, above lateral view, below dorsal view, 50 dif-
ferent views of lateral sclerite, scale 0.25 mm. — figs. 51-56. L. monochroma, 51-54 apex of abdomen, scale 2 mm, 51-52 ven-
tral view, 51 & (holotype), 52 9 (paratype), 53-54 lateral view, 53 5, 54 2; 55-56 endosomal sclerites (paratype), 55 median
arc and lateral sclerites, scale 0.5 mm, above lateral view, below dorsal view, 56 different views of lateral sclerite, scale 0.25
mm. — figs. 57-60: L. poliakanthina, 57-58 apex of abdomen of d holotype, scale 2 mm, 57 ventral, 58 lateral view; 59-60
endosomal sclerites of d paratype, 59 median arc and lateral sclerites, scale 0.5 mm, left lateral, right dorsal view, 60 differ-
ent view of lateral sclerites, scale 0.25 mm.
23
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
ous, medium brown. Legs light to medium brown
longitudinal stripes indistinct. Middle and hind fe-
mur with lighter apical rings distinct. Middle and
hind tibiae yellowish in apical part. Hemielytra grey-
ish brown with lighter veins, no costal or sub costal
light band, hind wings darker. Venter pale yellow, in
male with sparse lighter ciliation on thorax and base
of abdomen.
Structural characters. Length of antennal segments
I: I] : III: IV, male 3.35 : 2.3 : 2.5 : 3.0, female 2.4:
1.63 : 2.15 : 3.0. Second antennal segment longer
(1.1) than width of head in male, shorter (0.8) in fe-
male. Fore femur of male broader (1.4) than middle
femur, with a subapical indentation which has a very
small tooth near its apical edge and with dense pubes-
cence ventrally. Fore femur in female hardly (1.07)
wider than middle femur, with an indistinct subapical
indentation, without obvious pubescence. Middle fe-
mur in both sexes shorter than hind femur.
Hemielytra reaching about halfway the apices of con-
nexival spines. Connexival spines parallel, virtually
horizontal to pointing somewhat dorsad, reaching
about half their length beyond apex of genital seg-
ments. Ventral abdominal carina distinct on sternites
3-7 in both sexes.
Male. Middle femur slightly longer (1.07) than
body, with a fringe of cilia about half as wide as femur
and without a row of small pegs apically. The tiny
black setae on middle femur become, however, peg
like in basal part, suggesting several rows of very small
pegs in the basal part of middle femur. Hind femur
without apical pegs and with normal setae. Median
length of last abdominal shorter (0.8) than penulti-
mate abdominal sternite. Genital sternites subequal,
shorter (0.6) than last abdominal sternite, second
genital sternite slightly damaged. Endosomal sclerites
figs. 49-50.
Female. Middle and hind femur distinctly (0.85
and 0.9 respectively) shorter than length of body.
Median length of last sternite slightly longer (1.08)
than penultimate abdominal sternite subequal and
longer (1.3 and 1.45 respectively) than genital ster-
nites.
Etymology. — Hysterema, a Greek noun meaning ‘-
that what is lacking’, refers to the absence of small peg
rows in apical part of middle femur in males of this
species.
Comparative notes. — Similar to L. annulicornis, L.
monochroma and L. arachnis. The latter is much larg-
er and more solidly built. All three differ in having
one or two short rows of small pegs apically on the
middle femur of male and the black setae in basal part
of the middle femur not modified.
24
DISCUSSION
Although we have not seen all species of the genus,
some remarks on species groups can already be made.
Limnometra includes several apparently isolated spe-
cies, e.g. L. femorata Mayr with aberrant colour
pattern, a peculiar midfemoral spur in the male and
elongate lateral endosomal sclerites (fig. 5), L. octo-
punctata Hungerford with peculiar ventral markings
similar to Tenagogonus quinquemaculatus Miyamoto,
but not to any species of Limnometra, and an aberrant
lateral endosomal sclerite (fig. 9). A further example is
L. melanochroa with a colour pattern reminding so-
mewhat of L. rossi Hungerford & Matsuda. The latter
species falls outside the scope of this paper. It is one of
the smallest species in the genus, has lateral endoso-
mal sclerites of the L. insularis type, and a relatively
more stout build.
L. ciliata and L. cursitans form a small group, char-
acterised by uniformly coloured antennae and legs
and a tendency of having broad dark sublateral bands
on the pronotum. Unfortunately we have not seen
the male of L. cursitans, so we are unable to comment
upon the endosomal sclerites.
L. insularis, L. borneensis and L. pseudoinsularis
form a group of slender species with a remarkably
similar colour pattern, of which the lemon coloured
caudolateral margin of the pronotum is striking. In
addition, they have rather broad lateral endosomal
sclerites without a twisted tip (figs. 2, 6, 44). L. bru-
neiensis is associated with this group by its lateral
sclerites, but its more solid build reminds of the next
group.
Finally there is a large group of medium-sized to
large species: L. annulicornis, L. arachnis, L. aploa, L.
genitalis, L. hysterema, L. kallisto, L. lepta, L. mono-
chroma and L. pulchra. They have in common a very
uniform general build (except for size) and colour
pattern. In addition, the apex of the lateral endosomal
sclerites is twisted (figs. 1, 7, 10, 16, 22, 28, 34, 50,
56). This group has so far only been found on
Sulawesi where the genus has its highest species diver-
sity eastward to the Solomon Islands.
Up to now, we have studied two genera, Enithares
(Notonectidae) and Limnometra, with several appar-
ently closely related species in Sulawesi and Buton. In
Enithares we have found four closely related species
occurring in Sulawesi and Buton, with restricted
ranges. E. horvathi Kirkaldy in SW Sulawesi, E. cae-
saries Nsr. & Chen in Central Sulawesi, E. lansburyi
Nsr. & Chen in SE Sulawesi and £. skutalis Nsr. &
Chen on Buton (Nieser & Chen 1991). In
Limnometra we have found in the same area five
closely related species, viz. L. annulicornis in N
Sulawesi (and ? S Sulawesi), Z. lepta in N and NE
Sulawesi, L. aploa in E Sulawesi, L. arachnis in E
Sulawesi and on Buton and L. genitalis endemic to
Buton. A similar pattern has been found in the gerrid
genus Prilomera Amyot & Serville (Polhemus &
Polhemus 1986, cited in Whitten et al. 1988). The
genus Rhagovelia Mayr (Veliidae) seems to have a
more complex distributional pattern in Sulawesi
(Polhemus & Polhemus 1988), with a greater num-
ber of ecologically separated species in a given area.
The actual distributional ranges of the species of
the three genera Enithares, Limnometraand Ptilomera
show considerable similarity, possibly characterising
areas of endemism for aquatic Heteroptera. We sup-
pose that the presence of watersheds rather than a his-
torical separation of Sulawesi Selatan with slightly
higher sea level, was the most important causing fac-
tor for speciation in stream-dwelling insects. The lat-
ter phenomenon has been used in part for the expla-
nation of speciation and subspeciation in macaques
(Macaca, Cercopithecidae cited in Whitten et al.
1988), which also have similar distributional pat-
terns. At present our knowledge of the distribution of
aquatic Heteroptera is insufficient to make further
comments. Some of these restricted ranges may ap-
pear artificial after more intensive collecting in unex-
plored parts of Southwest Sulawesi and the eastern is-
lands as Banggai, Sula and Talaud.
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321-324, figs. 1-18.
Andersen, N. M., 1967. A contribution to the knowledge of
Philippine semiaquatic Hemiptera-Heteroptera. -
Entomologiske Meddelelser 35: 260-282, figs. 1-29.
Andersen, N. M., 1982. The semiaquatic bugs (Hemiptera,
Gerromorpha). — Entomonograph 3: 1-455, figs. 1-638.
Breddin, G., 1901. Die Hemipteren von Celebes. Ein
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turforschenden Gesellschaft Halle 24: 1-213, 1 plate.
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Cheng, L., & Fernando, C. H., 1969. A taxonomic study of
the Malayan Gerridae (Hemiptera: Heteroptera) with
notes on their biology and distribution. — Oriental Insects
3: 97-160, figs. 1-306, maps. 1-2.
Fabricius, J. C., 1794. Entomologia Systematica emendata
et aucta 4. Hafniae.
Hungerford, H. B., 1955. A new Limnometra from
Sumatra. — Journal of the Kansas Entomological Society
28: 67-68.
Hungerford, H. B. & Matsuda, R., 1958. The Tenagogonus-
Limnometra complex of the Gerridae. — Kansas
University Science Bulletin 39: 371-457, pls. 1-16, 1 text
fig
Kirkaldy, G. W., 1899. Aquatic Rhynchota in the collection
of the Royal Museum of Belgium. — Annales de la Société
Entomologique de Belgique 43: 42-47.
Lundblad, O., 1933. Zur Kenntnis der aquatilen und semi-
aquailen Hemipteren von Sumatra, Java und Bali. —
Archiv für Hydrobiologie, Supplement 12: 1-195, 263-
489, pls. 1-21, figs. 1-142.
Matsuda, R., 1960. Morphology, evolution and a classifica-
tion of the Gerridae (Hemiptera-Heteroptera). —
University of Kansas Science Bulletin 41: 25-632, figs. 1-
SE
Mayr, G. L., 1865. Diagnosen neuer Hemipteren II. —
Verhandlungen Zool.-Bot. Gesellschaft Wien 15: 429-
446.
Miyamoto, S., 1967. Gerridae of Thailand and North
Borneo taken by the joint Thai-Japanese biological expe-
dition 1961-62. — Nature and life in Southeast Asia 5:
217-257, figs. 1-92.
Nieser, N., & P. P. Chen, 1991. Naucoridae, Nepidae and
Notonectidae, mainly from Sulawesi and Pulau Buton
(Indonesia). — Tijdschrift voor Entomologie 134: 47-67,
figs. 1-21.
Polhemus, J. T. & D. A. Polhemus, 1988. Zoogeography,
ecology and systematics of the genus Rhagovelia Mayr
(Heteroptera: Veliidae) in Borneo, Celebes and the
Moluccas. — Insecta Mundi 2: 161-230, figs. 1-233.
Whitten, A. J., Mustafa, M. & Henderson, G., 1988. The
ecology of Sulawesi. — Gadjah Mada University
Press, Yogyakarta: xxi + 779, 64 pls.
Received: 28 August 1991
Accepted: 30 December 1991
25
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 1. Leg measurements in Limnometra in millimeters.
femur
L. aploa male
fore leg 4.1
middle leg 10.4
hind leg 10.8
L. aploa female
fore leg 3.6
middle leg 9.1
hind leg 92
L. arachnis male
fore leg 6.3
middle leg 17.5
hind leg 18.2
L. arachnis female
fore leg 4.4
middle leg 11.6
hind leg 11.4
L. genitalis male
fore leg De
middle leg 133
hind leg 157
L. genitalis female
fore leg 4.2
middle leg 10.2
hind leg 10.0
L. lepta male
fore leg 6.9
middle leg 195
hind leg 20.8
L. lepta female
fore leg 4.3
middle leg 113
hind leg 11.6
Ue)
tibia
32
9.0
6.7
3.8
10.4
26
tars]
0.55
tars2
femur
L. melanochroa male
fore leg 6.3
middle leg 17.9
hind leg 17.7
L. melanochroa female
fore leg 4.6
middle leg 12.4
hind leg 12.4
L. pseudoinsularis male
fore leg SW
middle leg 10.0
hind leg 10.6
L. hysterema male
fore leg 5.5
middle leg 16.0
hind leg 17.4
L. hysterema female
fore leg 4.4
middle leg 11.0
hind leg ID
L. monochroma male
fore leg 4.1
middle leg 10.7
hind leg 181657
L. monochroma female
fore leg 3.4
middle leg 2
hind leg 95
L. poliakanthina male
fore leg 6.1
middle leg 18.8
hind leg 18.8
tibia
5.4
15.2
15.6
4.0
10.9
10.2
3.2
8.2
6.3
Se
12.8
10.2
SE
10.1
6.9
55)
95
6.7
3.0
8.2
5.8
5.1
17.2
16.3
REMIGIJUS NOREIKA & RIMANTAS PUPLESIS
Zoologijos katedra VPI, Vilnius, Lithuania
SALIGACE AM EEEDING GRAGILEARIIDAE
(LEPIDOPTERA) OF CENTRAL ASIA
Noreika, R. & Puplesis, R., 1992. Salicaceae feeding Gracillariidae (Lepidoptera) of Central
Asia. Tijdschrift voor Entomologie 135: 27-41, figs. 1-31 [issn 0040-7496]. Published:
15 July 1992.
Seven Phyllonorycter species, two Cameraria and one Caloptilia species feeding on members of
Salicaceae in Central Asia are reviewed. Phyllonorycter dentifera sp. n., mining on Populus prui-
nosa Schrenk, is described. Phyllonorycter obliquifascia (Filipjev) and P. saliciphaga (Kuznetzov)
are transferred to Cameraria Chapman. A checklist and review of literature of all Central Asian
Gracillariidae is provided. A key to the species, descriptions and figures of the imagos, male
genitalia and mines of all species reviewed are provided.
Correspondence: Minological Research Laboratory, Zoologijos katedra VPI, g. Studentu 39,
Vilnius 34, Lithuania.
Keywords. — Gracillariidae, Caloptilia, Phyllonorycter, Cameraria, new species, Salicaceae,
Central Asia, checklist.
The Gracillariidae form a large family of leaf-min-
ing moths, known from all continents except
Antarctica. About 600 species of this family occur in
the Palaearctic (Kuznetzov 1981).
As far as known, the larvae of most Gracillariidae
are leaf-miners of woody and herbaceous angiosperms
belonging to 56 families of Dicotyledones (or
Magnoliopsida; Takhtajan 1987). Only one species,
Spulerina corticicola Kumata, 1964, has been record-
ed as a bark-miner of members of Pinaceae
(Gymnospermae). The majority of species feed on
Fagaceae and Rosaceae (almost 45-50% of the
Palaeartic species). Further many species are special-
ized on Fabaceae, Betulaceae and Salicaceae. Most
species are oligophagous. As an example, all species
(except Caloptilia stigmatella (Fabricius)) feeding on
Salix and Populus are restricted to Salicaceae. Only C.
stigmatella has also been noted in addition on
Betulaceae and Myricaceae (Kumata 1982).
The study of the Gracillariidae of Central Asia was
initiated by Gerasimov (1930, 1931, 1932). He not-
ed 13 species from Central Asia, including eight new
taxa. Later, one new species was added by Danilevsky:
Parornix persicella Danilevsky, 1955 and many more
species, including seventeen new species were added
due to Kuznetzov’s investigations (1956, 1975, 1978,
1979, 1981, 1985). Furthermore, some papers con-
taining faunistic or ecological data on Gracillariidae
from Tadzhikistan were published by Sherniyazova
(1975, 1982, 1984, 1988).
In the course of our own investigations in
Tadzhikistan, Turkmeniya, Uzbekistan and southern
Kazakhstan from 1986 to 1991, ten species were re-
corded for the first time from Central Asia. One of
them, Dialectica scalariella (Zeller), had earlier been
known from southern Europe, North Africa and the
Middle East (Wapshere & Kirk 1977). Eight species
were described as new (Noreika 1991; Noreika and
Puplesis in press).
The gracillarid fauna of the southern part of
Central Asia is poorly known. Five species have been
found in Iran (Deschka 1979; Triberti 1985, 1986,
1989) and four species are known from Afghanistan.
Two species are known from northern Pakistan:
Phyllonorycter montanella Bradley, 1980 (Bradley
1980) and Parornix compressa Triberti, 1989 (Triberti
1989). No species are known from Mongolia or the
centralasiatic part of China. Most of these species
mine leaves of various plant-families: Caprifoliaceae,
Rosaceae, Plantaginaceae, Fabaceae, Fagaceae,
Betulaceae, Viburnaceae and other ones. As yet, no
Salicaceae feeding species have been recorded from
these countries.
A vegetation type, dominated by Salicaceae, is
widespread in Central Asia and forms a characteristic
aspect of its vegetation. It is often predominant along
rivers and streams, forming a forest type called
“Tugai’. As many as ten species out of the 72 species
of Gracillariidae known from Central Asia are record-
ed as feeding on Salicaceae and apart from one spe-
cies, restricted to this family. Only one species,
Caloptilia stigmatella (Fabricius, 1781), appears to be
disjunct oligophagous, since in other regions this spe-
cies has also been noted on Myricaceae and
Betulaceae (Kumata 1982).
The ten Salicaceae feeding species of the centrala-
27
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Os
i
)
AT
=. | | |
a | =]
\)
|
|
2
\
\N
a)
ZI,
\N
—
Figs. 1-5. Adult Gracillariidae (males). - 1, Caloptilia stigmatella, 2, Cameraria saliciphaga; 3, C. obliquifascia; 4, Phyllonorycter
populifoliella; 5, Ph. populi. Scale 1 mm.
28
siatic fauna are the topic of this paper. The treatment
of species feeding on related host plants follows a tra-
dition in Gracillariidae (compare Kumata 1963,
Kuznetzov 1981, Ermolaev 1982, 1988) and is espe-
cially useful for diagnostic purposes. However, the
polyphyly of such an ‘ecological group’ is evident.
MATERIAL AND METHODS
Methods for the preparation of the genitalia large-
ly follow Falkovitsh & Stekolnikov (1978). The ge-
nitalia were studied with the help of a Biolam micro-
scope and a MBS-10 stereomicroscope.
The material studied has been collected during ex-
peditions (from 1986 to 1991) in various regions of
Central Asia, including previously unexplored areas
(Kugitangtau mountains in eastern Turkmeniya and
the valley of Vakhsh in southern Tadzhikistan). In
addition, the collection of the Zoological Institute of
the Russian Academy of Sciences in St. Petersburg
(ZIAS) was studied.
In the genera Phyllonorycter and Cameraria the fe-
male genitalia are not described, since in these genera
they have little diagnostic value, and are rarely used
for taxonomy or identification.
Locality names are spelled in accordance with the
Times Atlas of the World (comprehensive edition
1975 and later).
Botanic nomenclature follows Cherepanov (1981)
and Takhtajan (1987). Under ‘Material examined”
only centralasiatic material is listed.
The type specimens of the new species are deposit-
ed in the collection of the Minological Research
Laboratory of the Zoological Department of the
Pedagogical Institute, Vilnius, Lithuania (ver), and in
the National Museum of Natural History, Leiden,
The Netherlands (RMNH).
Checklist of the Gracillariidae of Central Asia
Genus Micrurapteryx Spuler, 1910
1. M. kollariella (Zeller, 1839)
2. M. sophorella Kuznetzov, 1979
3. M. fumosella Kuznetzov & Tristan, 1985
4. M. tortuosella Kuznetzov & Tristan, 1985
5. M. sophorivora Kuznetzov & Tristan, 1985
6. M. bidentata Noreika, 1992a
7. M. minima Noreika, 1992a
Genus Polymitia Triberti, 1986
8. P. eximipalpella (Gerasimov, 1930)
9. P. laristana Triberti, 1986
Genus Gracillaria Haworth, 1828
10. G. loriolella Frey, 1881
Genus Caloptilia Hübner, 1825
11. C flava (Staudinger, 1870)
12. C. stigmatella (Fabricius, 1781)
Noreika & Purresis: Gracillariidae
13. C. semifascia (Haworth, 1828)
14. C. acerivorella (Kuznetzov, 1956)
15. C. acericolella Kuznetzov, 1981
16. C. fribergensis (Fritzsche, 1871)
17. C. populetorum (Zeller, 1839)
18. C. cuculipennella (Hiibner, 1796)
19. C. roscipennella (Hübner, 1796)
Genus Cupedia Klimesch & Kumata, 1973
20. C. cupediella (Herrich-Schäffer, 1855)
Genus Aspilapteryx Spuler, 1910
21. A. tringipennella (Zeller, 1839)
22. A. limosella (Duponchel, 1843)
23. A. magna Triberti, 1985
Genus Calybites Hübner, 1822
24. C. phasianipennella (Hübner, 1813)
Genus Eucalybites Kumata, 1982
25. E. aurogutella (Stephens, 1835)
Genus Acrocercops Wallengren, 1881
26. A. iraniana Triberti, 1989
Genus Dialectica Walsingham, 1897
27. D. scalariella (Zeller, 1850
Genus Leucospilapteryx Spuler, 1910
28. L. omissella (Stainton, 1848)
Genus Parornix Spuler, 1910
29. P. subfinitimella Kuznetzov, 1956
30. P. hastata Triberti, 1989
31. P. compressa Triberti, 1989
32. P. kugitangi Noreika, 1991
33. P. asiatica Noreika, 1991
34. P. torquilella Zeller, 1850
35. P. petiolella Frey, 1863
36. P. cotoneasterella Kuznetzov, 1978
37. P. turcmeniella Kuznetzov, 1956
38. P. amygdalella Kuznetzov, 1978
39. P. persicella Danilevsky, 1955
Genus Phyllonorycter Hübner, 1822
40. Ph. medicaginella (Gerasimov, 1930)
41. Ph. fabaceaella (Kuznetzov, 1978)
42. Ph. insignitella (Zeller, 1846)
43. Ph. schreberella (Fabricius, 1781)
44. Ph. millierella (Staudinger, 1871)
45. Ph. emberizaepennella (Bouche, 1834)
46. Ph. montanella Bradley, 1980
47. Ph. lonicerifaga Noreika, 1992a
48. Ph. caspica Noreika, 1992a
49. Ph. raikhonae Noreika, 1992b
50. Ph. pastorella (Zeller, 1846)
51. Ph. populifoliella (Treitschke, 1833)
52. Ph. populi (Filipjev, 1926)
53. Ph.populicola Kuznetzov, 1975)
54. Ph. asiatica (Gerasimov, 1931)
55. Ph. connexella (Zeller, 1846)
56. Ph. dentifera Noreika, sp.n.
57. Ph. pruinosella (Gerasimov, 1931)
58. Ph. turcomanicella (Kuznetzov, 1956)
59. Ph. aceripestis (Kuznetzov, 1978)
29
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
li), >
ES
\ | |
9
Figs. 6-10. Phyllonorycter species (males). - 6, Ph. populicola, 7, Ph. pastorella; 8, Ph. pruinosella; 9, Ph. asiatica; 10, Ph. den-
tifera, holotype. Scale 1 mm.
30
60. Ph. aceriphaga (Kuznetzov, 1975)
61. Ph. juglandicola (Kuznetzov, 1975)
62. Ph. infirma Deschka, 1974
63. Ph. platani (Staudinger, 1870)
64. Ph. pyrifoliella (Gerasimov, 1933)
65. Ph. malella (Gerasimov, 1931)
66. Ph. cerasicolella (Herrich-Schäffer, 1855)
67. Ph. iranica Deschka, 1979
68. Ph. corylifoliella (Hübner, 1796)
69. Ph. turanica (Gerasimov, 1931)
70. Ph. hissarella Noreika, 1992b
Genus Cameraria Chapman, 1902
71. €. obliquifascia (Filipjev, 1926) comb.n.
72. C. saliciphaga (Kuznetzov, 1975) comb.n.
SYSTEMATIC PART
Key to the male Salicaceae feeding Gracillariidae of
Central Asia (based on genitalia)
l. Uncus and transtilla absent; valva strongly broad-
ened distally, with very long setae (fig. 11)..........
es OLA. Caloptilia stigmatella
- Uncus and transtilla present; valva not broadened
distallyawıthshortsetaen. een D
Dea alyaeısymmetnicalkr re eee tees che sees 3
= Valvaerasymmetrical sta nee 7
3. Aedeagus half as long as valva, without apical
hook and without bulge basally … … … … … … … … 4
— Aedeagus longer than valva, with apical hook, but
wathoutioulge balle #27" enna 5
4. Valva strongly narrowed in apical third; 8th ster-
nite wathydistal invagination, Ci.
er NN Cameraria saliciphaga
— Valva gradually narrowed at tip only; 8th sternite
without distal invagination .......... C. obliquifascia
5. Aedeagus more than twice as long as valva; valva
distallyawıderr. nenn. Ph. populifoliella
— Aedeagus almost as long as valva or slightly long-
er than valva; valva narrowed distally … … … … … 6
6. Valva not abruptly tapered in distal half and with
additional small tooth above middle and tiny one
LADEN cena ho nn Ph. populi
— Valva abruptly tapered in distal half and without
additonal tooth above middle and with more or
lessjlongionelauapesm ren na Ph. pastorella
7. Valvae almost equal in length and width; transtil-
[APS NM EN CA EEE Een Ph. asiatica
— Right valva distrinctly more slender than left one
8
8. Vinculum large, more or less rectangular; only
left valva with long distal process (right valva with
extremelyasmallétoothvorly,) ee
a EN DS Ph. pruinosella
— Vinculum small, more or less triangular; both
valvae with weakly sclerotized but more or less
large, tooth-like distal process.......... Ph. dentifera
Noreika & Puptesis: Gracillariidae
Descriptions
Caloptilia stigmatella (Fabricius, 1781)
(figs. 1, 11, 22)
Tinea stigmatella Fabricius, 1781: 297.
Gracillaria stigmatica Matsumura, 1931: 1101.
Caloptilia stigmatella Inoue 1954: 1-26, Ermolaev 1977:
108, Kuznetzov 1981: 182-183, 192, 201, Kumata 1982:
32-34, 147, 157, Patzak 1986: 140, 149, 158
Diagnosis. — Can immediately be separated exter-
nally from the other treated species by the shape of
the costal blotch of the forewing. From similar orien-
tal C. chrysolampra (Meyrick) it differs by larger size
as well as broader and longer aedeagus.
External characters (fig. 1). — Wingspan 12-16
mm. Face whitish. Vertex and labial palpi greyish
brown. Maxillar palpi pale grey. Scapus grey with
some ferruginous scales. Flagellum grey annulated
with dark grey. Thorax greyish brown. Forewing
greyish brown with distinct white triangular costal
blotch, slightly irrorate with ferruginous scales. Cilia
pale grey with black cilia-line. Hindwing and cilia sil-
very grey.
Male genitalia (fig. 11). — Symmetrical. Vinculum
triangular. Saccus weakly developed. Valva curved,
slightly dilated apically, straight on terminal margin.
Aedeagus slender, almost pointed distally, without
cornuti.
Female genitalia. — See Kumata (1982).
Biology. — Host plants: Populus spp. and Salix spp.,
in Central Asia on S. blakii Goerz. Mine (fig. 22)
starts as underside epidermal blotch; later the larva
rolls the leaf up to more than half length, producing
pyramid-like case.
Distribution. — Throughout Europe, Asia Minor,
Central Asia, Siberia, Eastern Asia (including Japan
and China), North America.
Material examined: 56, 42. — Turkmeniya: 1d,
Ashkhabad, env. Firyuza, 9.v.1991 larva on Salix, E.
Kokanova (coll. Kokanova). — Tadzhikistan: 14, 30 km N
Dushanbe, env. Varzob (loc. Kondara), 20.vii.1990, R.
Noreika (vei); 1d, the same locality, 6.ix.1990, larva on
Salix blakii, N5069, ex p. 10.x.1990, R. Noreika (vei); 24,
49, same locality, 12.vi.-7.vii.1991, V. Sruoga (vPI).
Cameraria saliciphaga (Kuznetzov, 1975)
comb. n. (figs. 2, 12, 23)
Lithocolletis saliciphaga Kuznetzov 1975: 418-420, 1981:
283, 287, 288.
Diagnosis. — Similar to C. obliquifascia (Filipjev),
but differs by shape of medial fascia and especially by
paler appearance of forewing (in C. obliquifascia it is
irrorate with blackish scales) and strong bulbose
broadening of medial part of valva.
External characters (fig. 2). — Wingspan 5.0-7.0
al
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Fig. 11-14. Male genitalia.-11, Caloptilia stigmatella, 12, Cameraria saliciphaga; 13, G. obliquifascia, 14, Phyllonorycter popu-
ifoliella. Scale 0.2 mm.
92
mm. Face white, sometimes with central orange spot.
Labial palpi white. Tuft on head consists of orange-
ferruginous and white, piliform scales. Scapus grey-
ish, with few ferruginous scales. Flagellum greyish,
annulated with brownish black. Thorax orange-ferru-
ginous with white margins. Forewing orange-ferrugi-
nous with four oblique fasciae, distally edged with
fuscous or brown scales. Basal fascia short and not
reaching costa. Second fascia inwards sharply curved
towards costa. Medial fascia usually bifurcate towards
costa, but occasionally only curved. Apical fascia sin-
uate and bifurcate distally; lower branch distinctly
narrower than upper. Apical spot absent. Cilia grey.
Hindwing and cilia grey. Abdomen dark ferruginous
brown at upperside, dark grey laterally.
Male genitalia (fig. 12). — Symmetrical. Uncus
shorther than distal process of valva. Vinculum
broad, but invaginated distally. Valva simple, but it’s
medial part strongly broadened (bulbose). Apical
parts of valva strongly and abruptly narrowed and el-
ongate. Transtilla symmetrical. Sternite 8 broad,
shorter than valva and slightly invaginated distally.
Aedeagus tube-like in shape, almost twice as short as
valva, broadened in basal part, narrowed apically.
Biology. — Host plants: Salix excelsa S. G. Gmel.
and S. triandra L. Mine (fig. 23) on leaf upperside, as
elongate blotch, usually occupying nearly one-fourth
of width of leaf, and situated in various parts of leaf.
Pupation in mine.
Distribution. — Turkmeniya, Uzbekistan and
Tadzhikistan.
Material examined: 72d , 64%. — Turkmeniya: 5d, env.
Ashkhabad, Geok — Tepe, 12.v.1986, R. Puplesis (vei); 14,
lolotan, 13.viii.1988, R. Puplesis (vei); 3d, 29, Tedzhen,
14-15.viii.1988, V. Sruoga et R. Puplesis (ver); 66,
59, env. Svintsovyy Rudnik, 11.viii.1989, larvae on Salix
triandra, R. Noreika (ver); 1d, 19, the same locality,
26.viii.1990, R. Noreika (vei). — Uzbekistan: 34, 19, env.
Dzharkurgan, 31.viii.1990, R. Noreika (vei). -
Tadzhikistan: 38, 52 (holotype and paratypes), 30 km N
Dushanbe, env. Varzob (loc. Kondara), 27.vii.1970, larvae
on Salix excelsa, R. Sherniyazova (z1as); 6d, 19, same local-
ity, 27.vi.-20.vili.1986, R. Puplesis (ver); 176, 239, same
locality, 17-23.viii.1989, R. Puplesis (vri); 1d, same local-
ity, 4.xi.1989, larva on Salixsp., N 5042, ex p. 1.iii.1990, R.
Puplesis (vei); 7d, 122, same locality, 14.viii.2.ix.1990, R.
Noreika (vei); 138, 109, same locality, 20.vii-9.viii.1990,
V. Sruoga; 4d, 49, same locality, 5-8.viii.1990, J.
Bajarunas (vri); 2d, env. Nurek (Zardolu), 6.viii.1990, V.
Sruoga (vei).
Cameraria obliquifascia (Filipjev, 1926) comb. n.
(figs. 3, 13, 24)
Lithocolletis obliquifascia Filipjev 1926: 284 Gerasimov
1932: 237-239, Kuznetzov 1981: 283, 287, 288.
Diagnosis. — Externally similar to C. saliciphaga
Norerka & PupLesis: Gracillariidae
(Kuznetzov), but considerably larger. In contrast to
C. saliciphaga, forewing irrorate with blackish and
brownish scales. InC. obliquifascia second fascia of fo-
rewing more oblique than in C. saliciphaga and not
sinuate. In male genitalia it differs by lack of bulbous,
medial broadening of valva.
External character (fig. 3). — Wingspan 7.0-8.1
mm. Face and labial palpi white. Tuft on head con-
sists of pale yellowish ferruginous, piliform scales.
Scapus white with mixture of brown-ferruginous
scales. Flagellum yellowish ferruginous, annulated
only on upperside with brown. Thorax ochreous with
mixture of brownish scales in centre. Forewing ochre-
ous, irrorate with black and brown scales, and with
four white strongly oblique fasciae, with diffuse distal
black edging. Basal fascia short, not reaching costa.
Second fascia sometimes slightly interrupted just be-
fore reaching costa. Medial fascia slightly sinuate, and
bifurcate near costa. Fourth (distal) fascia curved an-
teriorly. Forewing apex darker than rest of forewing,
without apical spot. Line just proximal to cilia (not
cilia line!) is narrow and fuscous. Cilia pale yellowish.
Hindwing and cilia pale greyish. Abdomen dorsally
dark grey, and laterally greyish.
Male genitalia (fig. 13). — Symmetrical. Uncus less
than half length of valva. Vinculum broad, trapezi-
um-shaped, twice as short as aedeagus. Valva simple,
without abrupt broadening, rounded at apex, and
bent ventrally. Transtilla symmetrical, with posterior
(distal) margin. Aedeagus tube-shaped almost half as
long as valva. Basal part of aedeagus flattened dorso-
ventrally, and broadened laterally. Tube of aedeagus
slightly curved medially.
Biology. — Host plants: Salix spp. and Populus af-
ghanica (Aitch. et Hemsl) Schneid, P. alba L. Mine
(fig. 24) on upperside of leaf, occasionally on under-
side, variable in shape, sometimes elongate, and situ-
ated in various places of leaf. In centre of mine, round
spot present with light surroundings along frass-line.
Distribution. — Turkmeniya, Uzbekistan and
Tadzhikistan.
Material examined: 21 d, 35d.- Tadzhikistan: 30 km N
Dushanbe, env. Varzob (Kondara), 28.vi-21.viii.1986, R.
Puplesis (vei); 5d, 19, same locality, 17-23.viii.1989, R.
Noreika (vei); 7d, 149, same locality, 14-20.viii.1990, R.
Noreika (vpi); 56, 59, same locality, 15.viii.1990, larvae
on Populus afghanica, N 5054 and N 5071, R. Noreika
(vei); 2d, 159, same locality, 9.vii-9.viii.1990, V. Sruoga
(vei); 1d, same locality, 20.vii.1990, R. Puplesis (vei); 16,
same locality, 8.viii.1990, J. Bajarunas (ver).
Phyllonorycter populifoliella (Treitschke, 1833)
(Figs. 4, 14, 25)
Lithocolectis populifoliella Treitschke, 1833: 188. Gerasimov
1932: 242-244, Kuznetzov 1981: 284, 285.
33
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 15-18. Male genitalia of Phyllonorycter. - 15, Ph. populi, Sayat, Turkmeniya, 28.vii.1990; 16, Ph. populi, 8th sternite of
other specimen, same data; 17, Ph. populicola; 18, Ph. pastorella. Scale 0.2 mm.
34
Phyllonorycter populifoliella (Treitschke); Buhl et al. 1983: 5-
8.
Diagnosis. — Externally similar to Ph. sagitella
(Bjerkander) and Ph. apparella (Herrich-Schäffer),
but differs by indistinct forewing pattern. In male
genitalia resembling Ph. populi (Filipjev), but differ-
ing by longer vinculum and aedeagus.
External characters (fig. 4). — Wingspan 7.0-8.0
mm. Face and palpi creamy white. Tuft creamy white
with mixture of brownish scales. Scapus creamish.
Flagellum brownish annulated with brown. Thorax
white, slightly irrorate with brown. Forewing creamy
white with variable, brown pattern. Five to six costal
and four dorsal strigulae present, variably developed.
Costal strigulae larger and more distinctive than dor-
sal strigulae. Apical spot brown. Line at apex of fo-
rewing creamy. Cilia creamy. Hindwing and cilia
brownish creamy. Abdomen dark creamy.
Male genitalia (fig. 14). — Symmetrical. Uncus
reaching slightly beyond apex of valva. Vinculum
longer than width of valva, proximally narrowed.
Valva simple, slightly broadened distally with small
tooth at apex. Transtilla U-shaped. Sternite 8 trian-
gular at apex and with slight distal invagination.
Aedeagus more than twice as long as valva, with small
hook at apex.
Distribution. — Throughout Europe, the Caucasus,
Ural, Western Siberia, northern part of Central Asia.
Biology. — Host plants: Populus tremula L., P. nigra
L., P. balsamifera L., P. laurifolia Ledeb. Mine (fig.
25) on underside, rarely on upperside of leaf, folded
and situated between veins.
Material examined: 1d. Kazakhstan: 1d, env. Alma-
Ata, Zailiyskiy Alatau ridge, 22.ix.1957, M. Falkovitsh (-
ZIAS).
Phyllonorycter populi (Filipjev, 1926)
(figs. 5, 15-16, 26)
Lithocolletis populiella Filipjev, 1926: 284 (preoccupied by
L. populiella Chambers, 1878).
Lithocolletis populi Filipjev, 1931: 236 (replacement name
for L. populiella Filipjev), Kuznetzov 1981: 284.
Diagnosis. — Most similar to Ph. sagitella
(Bjerkander), but differs by distal part of valva not be-
ing narrowed and by presence of small tooth on apex.
External characters (fig. 5). — Wingspan 7.4-9.5
mm. Face and labial palpi white. Tuft on head con-
sists of whitish and brownish-grey piliform scales.
Scapus white. Flagellum greyish, weakly annulated
with grey. Thorax irrorate with white, brown or
brownish scales. Forewing with slightly diffuse pat-
tern, consisting of five white, diffuse costal strigulae
(irrorate with pale brownish scales) and four weakly
distinct, white dorsal strigulae. All strigulae are proxi-
NOREIKA & PupLesis: Gracillariidae
mally edged dark brown. Apical region of forewing
with two dark brown streaks. Cilia line at distal mar-
gin of forewing dark brown. Cilia light grey.
Hindwing dark creamy greyish, cilia light creamy
greyish. Abdomen whitish creamy.
Male genitalia (figs. 15-16). — Symmetrical. Uncus
extending slightly beyond valva. Vinculum triangu-
lar, more than twice as short as valva. Valva simple,
slightly broadened medially, distally blunt, and with
small denticle on apex between setae. Apical third of
valva proximally strongly sclerotized, and with bifur-
cate denticle. Transtilla symmetrical, with two proxi-
mal processes. Sternite 8 broad, almost reaching apex
of valva, with small invagination distally. However,
smaller specimens without invagination of 8th ster-
nite have been noted by us (fig. 16). Aedeagus with
small hook at apex.
Biology. — Host plants: Populus alba L., P. nigra L.
Mine (fig. 26) or underside of leaf, weakly folded,
oval in shape, and usually situated between midrib
and two lateral veins. Pupation in mine.
Distribution. — Southern Kazakhstan, Turkme-
niya, Uzbekistan, Tadzhikistan.
Material examined: 2d, 29. — Turkmeniya: 2d, 29,
env. Svintsovyy Rudnik (loc. Sayat), 28.viii.1990, larvae on
Populus alba, R. Noreika (ver).
Phyllonorycter populicola (Kuznetzov, 1975)
(figs. 6, 17, 27)
Lithocolletis populicola Kuznetzov, 1975: 419-420, 1981:
287-288.
Diagnosis: — Externally most similar to Ph. populi-
albae (Kuznetzov) and Ph. comparella (Duponchel),
but distinguished by absence of dark apical spot on
forewing. The male genitalia of this species are
unique in Phyllonorycter.
External characters (fig. 6). — Wingspan 6.5-7.5
mm. Face covered by white scales, tipped grey-
brown. Labial palpi white. Tuft on head consists of
white and grey-brownish scales. Scapus white with
few brown scales. Flagellum grey-brown, annulated
with fuscous. Thorax white with some grey-brown
and orangeish scales, tipped dark brown. Forewing
with slightly diffuse pattern, and covered by scales of
various colours: orangeish, grey, dark grey and white,
tipped dark brown to blackish. More or less distinct
pattern consists of five costal and four dorsal white
strigulae. All strigulae slightly irrorate with few
orangeish and greyish scales, tipped, blackish. White
strigular with brown to blackish lines at their proxi-
mal margin. Apical spot absent. Cilia grey, only at fo-
rewing apex orange. Hindwing and cilia grey.
Abdomen grey.
35
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
<=
Figs. 19-21. Male genitalia of Phyllonorycter. - 19, Ph. pruinosella; 20, Ph. asiatica; 21, Ph. dentifera, holotype. Scale 0.2 mm.
36
Male genitalia (fig. 17). — Symmetrical. Uncus not
reaching apex of valva. Vinculum broad, long, apical-
ly rounded. Valva complex, medially strongly broad-
ened and bulged; bulge with some large, strongly scle-
rotized spines; three to four smaller spines on distal
margin of bulge. Apical third of valva abruptly nar-
rowed, with dentate dorsal margin and cornutus-like
spine at apex. Small setae covering apical third of val-
va. Transtilla symmetrical and broad. Sternite 8 not
reaching apex of valva, broad and dentate distally.
Aedeagus longer than valva, smooth and gradually
narrowed toward apex, and with dorsal hook at apex.
Biology. — Host plant: Populus afghanica (Aitch. et
Hemsl.) Schneid. Mine (fig. 27) on underside of leaf,
occasionally on upperside, folded, situated in various
parts of leaf and strongly distorting it.
Distribution. — Tadzhikistan only.
Material examined: 86, 32. — Tadzhikistan: 24 (holo-
type and paratype), 30 km N Dushanbe, env. Varzob (loc.
Kondara), 3-22.vi.1971 larva on Populus sp., ex p.
7.vii.1971, R. Sherniyazova (zis); 16, same locality, 8.-
viii.1986, R. Puplesis (vei); 1d, 22, same locality, 15.20.-
viii.1990, R. Noreika (vei); 2d, 19, same locality, 15.-
viii.1990, larva on Populus afghanica, N. 5060, ex p.
23.vili-14.ix.1990, R. Noreika (ver); 16, 49, same locality,
20.vii-10.viii.1990, V. Sruoga (vei); 1d, same locality, 8.-
viii. 1990, J. Bajarunas (ver).
Phyllonorycter pastorella (Zeller, 1846)
(figs. 7, 18, 28)
Lithocolletis pastorella Zeller, 1846: 250; Kumata 1963: 54-
DI:
Lithocolletis fainae Gerasimov, 1931: 128/130.
Lithocolletis pastorella fainae, Gerasimov 1932: 240-241,
Kuznetzov, 1981: 281.
Diagnosis. — Externally closely resembling Ph. ap-
parella (Herrich-Schäffer) and Ph. chiclanella
(Staudinger), but distinguished by distinct, bright,
white strigulae of forewing. In male genitalia, resem-
bling Caucasian Ph. armeniella (Kuznetzov), but dif-
fering by valva being broader, not bilobed apically,
and with horn-like spine, and by longer aedeagus. In
male genitalia, Ph. pastorella also closely resembles the
Japanese Ph. pulchra (Kumata), which feeds on
Rosaceae, but pastorella is easily separeted from this
species by its long aedeagus.
External characters (fig. 7). — Wingspan 6.7-9.00
mm. Face and labial palpi white. Tuft on head white.
Scapus white with tuft of white scales at base.
Flagellum shining white. Thorax bright white, some-
times with pale ferruginous-brown spot at its centre.
Forewing with pale ferruginous-brown and white pat-
tern. Basal streak and dorsal streak touching forming
together one white spot, which does not reach costa
of forewing, with some ferruginous-brown scales al-
most at its centre. In medial and apical part of fore-
Noreika & PupLesis: Gracillariidae
wing, five costal and four dorsal white strigulae; these
proximally edged brown or black. First pair of strigu-
lae oblique and situated almost at middle of forewing.
First costal strigula reaching dorsal strigula, frequent-
ly joining each other. Second costal strigula almost
equal in width as first costal streak, but shorter.
Second costal strigula situated proximal to the tip of
the dorsal strigula. Apical three costal and dorsal strig-
ulae slightly oblique. Second dorsal is widest among
all other strigulae of distal region. Brown-blackish ap-
ical spot present. Cilia line at apex of forewing black-
ish-brown. Cilia whitish. Hindwing grey cilia whit-
ish. Abdomen ferruginous grey, but in distal part
yellowish white and laterally whitish.
Male genitalia (fig. 18). — Symmetrical. Uncus
reaching apex of valva. Vinculum triangular, and
shorter than width of valva. Valva simple, broad, and
in apical third narrowed, with horn-like spine on
apex. Near this spine small cluster of strongly sclero-
tized small spines present. Transtilla symmetrical and
U-shaped. Sternite 8 broad, slightly rounded and not
reaching apex of valva. Aedeagus tube-shaped, twice
as long as valva, and with hook-like process.
Biology. — Host plants: Salix spp. (in Central Asia
S. triandra L. and S. babylonica L.). Mine (fig. 28) on
underside of leaf, rarely in upperside, folded, and sit-
uated in various parts of leaf, but usually near midrib
vein; occasionally midrib also damaged.
Distribution. — Throughout Europe, Turkmeniya,
Tadzhikistan, Kirgiziya, Korea, Japan and possibly in
fareastern territories of Russia.
Material examined: 55d, 162 — Turkmeniya: 40 km E
Kara-Kala, Kopet Dag, 28.v.1988, R. Noreika (vei); 14d,
92, env. Svintsovyy Rudnik, 11.viii.1989, larvae on Salix
triandra, R. Noreika (ver); 19, same locality, 28.viii.1990,
R. Noreika (ver); 4d, Ashkhabad, botanic garden, 17.viii-
14.x.1991, larvae on Salix sp., E. Kokanova (coll.
Kokanova); 1¢, env. Ashkhabad, 24.vi.1991 larva on
Populus sp., E. Kokanova (coll. Kokanova). — Tadzhikistan:
408,29, env. Regar (loc. Tursunzade), 24.viii.1990, larvae
on Salix sp., R. Noreika (ver).
Phyllonorycter pruinosella (Gerasimov, 1931)
(figs. 8, 19, 29)
Lithocolletis pruinosella Gerasimov, 1931: 130-131, 1932:
127-129, Kuznetzov 1981: 283, 288, 289.
Diagnosis. — Externally and in male genitalia very
characteristic species; among Salicaceae feeding spe-
cies easily recognisable by strong asymmetry of valvae
and by long spine on left valva.
External characters (fig. 8). — Wingspan 5.0-6.1
mm. Face, labial palpi and tuft on head white. Scapus
and flagellum white without distinct annulation.
Thorax bright white. Forewing snow-white with
three pale yellow, perpendicular fasciae, more or less
3
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 22-31. Leaf-mines. - 22, Caloptilia stigmatella on Salix blakii, 23, Cameraria saliciphaga on Salix triandra, 24, C. obli-
quifascia on Populus afghanica; 25, Phyllonorycter populifoliella on Populus balsamifera, 26, Ph. populi on Populus alba, 27, Ph.
populicola on Populus afghanica; 28, Ph. pastorella on Salix triandra, 29, Ph. pruinosella on Populus pruinosa ; 30, Ph. asiatica
on Salix blakii ; 31, Ph. dentifera on Populus pruinosa. Scale 2 cm.
38
equal in width. All fasciae edged dark brown. First
and second fasciae situated in basal area of forewing,
and having characteristic shape (see fig. 8). Apical ar-
ea of forewing pale yellow with short costal strigula
just before apex and small dark-brown apical spot.
Distinct blackish cilia-line present. Hindwing and cil-
ia of both wings white. Abdomen bright white later-
ally, and pale golden dorsally.
Male genitalia (fig. 19). — Distinctly asymmetrical.
Uncus reaching beyond apex of valva. Vinculum
broad, slightly trapezium-shaped. Left valva twice as
wide as right one, tapering in apical part, and with
long, strongly angled spine. Right valva with small
tooth before apex. Transtilla symmetrical, and U-
shaped. Sternite 8 triangular, broad at base, and dis-
tally pointed. Aedeagus tube-like with small hook at
apex.
Biology. — Host plants: Populus pruinosa Schrenk,
P. euphratica Olivier, occasionally Salix sp. Mine (fig.
29) small, smooth, oval, situated on upperside or
underside of leaf, but on Salix only recorded from leaf
underside. Pupation in mine.
Distribution. — Kazakhstan, Turkmeniya, Uzbeki-
stan and Tadzhikistan.
Material examined: 105, 29.- Turkmeniya: 1d Tedzhen,
8.viii.1988, R. Puplesis (ver). — Uzbekistan: 7d (holotype and
paratypes), env. Bukhara, 5.viii.1928, larvae on Populus prui-
nosa, A. Gerasimov (zis); 19, Khiva (Novyi Urgench),
24.vii.1927, larva on Salix sp., A. Gerasimov (ZIAS). —
Tadzhikistan: 1d, 30 km N Dushanbe, env. Varzob (loc.
Kondara), 20.viii.1990, R. Noreika (vri); 12, 20 km S
Dzhilikul’, 13.vii.1990, R. Puplesis (ver).
Phyllonorycter asiatica (Gerasimov, 1931)
(figs. 9, 20, 30)
Lithocolletis connexalla asiatica Gerasimov, 1931: 130;
1932: 134-136.
Diagnosis. — Most similar to Ph. connexella
(Zeller), but differs externally by paler background of
forewing and colouration of flagellum, which has no
annulation. In male genitalia, Ph. asiatica is distin-
guished from Ph. connexella by longer proximal pro-
cess of vinculum.
External characters (fig. 9). — Wingspan 6.0-10.0
mm. Face white with some brownish-ferruginous
scales at its centre. Palpi yellowish brown. Tuft pale
yellowish at base and brownish-ferruginous at apex.
Scapus white, with tuft of white, piliform scales at
base. Flagellum white with annulation. Thorax yel-
lowish brown with whitish lateral and anterior mar-
gins. Forewing yellowish ferruginous with white
strigulae and streaks. Medio-basal streak short. White
costal streak additionally present, slightly shorter than
medio-basal streak. Three dorsal and three costal
strigulae present, proximally edged with black. First
Norerka & Purresis: Gracillariidae
costal strigula situated in middle of forewing. Second
costal strigula narrower but twice as long as third co-
stal strigula. Dorsal strigulae equally spaced. First and
second of these with equal width, and connected by
medio-basal streak at their apexes, third dorsal strigu-
la with wider base than other dorsals and curved dis-
tally. Distal part of forewing dark, irrorate with
brownish scales. Small, black apical spot present.
Apex of forewing with brownish-ferruginous line.
Cilia whitish. Hindwing and cilia greyish.
Male genitalia (fig. 20). — Asymmetrical. Uncus al-
most reaching apex of valva. Vinculum triangular, ta-
pering distally. Valva simple and with long comb.
Left valva shorter than right one, and with long,
slightly arcuate apical process. Transtilla slightly
asymmetrical; left process longer than right one.
Abdominal sternite 8 gradually narrowed towards
apex and with small distal invagination. Aedeagus
tube-like and with small hook at apex.
Biology. — Host plants: Salix alba, S. blakii Goerz,
S. fragilis L., Populus nigra L., and possibly also other
Salicaceae. Mine (fig. 30) on underside of leaf, occa-
sionally on upperside, folded and situated in various
parts of leaf.
Distribution. — Kazakhstan, Turkmeniya, Uzbeki-
stan, Tadzhikistan.
Material examined: 106, 282.—Turkmeniya: 2, © ; env.
Ashkabad, Firyusa, 1.vi.1988, larvae on Salix sp. R.
Noreika (vei); 1d, Kugitangtau, env. Svintsovyy Rudnik,
11.viii.1989, R. Noreika (vei); 1d, 19, env. Ashkhabad,
Firyusa, 22-27.v.1991, larvae on Salix sp., E. Kokanova (-
coll. Kokanova). — Tadzhikistan: 16, 30 km N Dushanbe,
env. Varzob (loc. Kondara), 9.viii.1986, R. Puplesis (vri);
23,102, same locality, 12.viii-6.1x.1990. 1990, R. Noreika
(ver); 22, same locality, 6.ix.1990, larvae on Salix blakii, N
5067, ex p. 15.ix.1990, R. Noreika (vei); 38, 99, same lo-
cality, 21.vii-10.viii.1990. V. Sruoga (vei); 1d, same local-
ity, 8.viii.1990, J. Bajarunas (vei).
Phyllonorycter dentifera Noreika sp. n.
(figs. 10, 21, 31)
Type material. — Holotype d : southern Tadzhikistan, 20
km S Dzhilikul’, 18.viii.1990, R. Noreika (ver). Paratypes:
134,99, same locality, 26.vii--18.viii.1990, R. Noreika, R.
Puplesis, V. Sruoga (vri); 1d, same locality, 17.viii.1990,
R. Noreika (RMNH). 1d, same locality, 13.vii.1990, larvae
on Populus pruinosa, N 4201, ex 1.1.17.viii.1990, R.
Puplesis (vei); 16, Turkmenistan, Tedzhen, 6.vii.1991, lar-
va on Populus sp., ex 1.13.vii.1991, R. Puplesis (ver).
Diagnosis. — Externally very distinct species.
Recognizable by connected costal and dorsal strigulae
in apical part of forewing, and by very large tuft of
white, piliform scales on scapus. In male genitalia,
slightly resembling some other Phyllonorycter species
with asymmetrical valvae, such as Ph. quinqueguttella
(Stainton), but easily distinguished from all these spe-
39
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
cies by shape of apical process of valva, which is short
and weakly sclerotized.
External characters (fig. 10). — Wingspan 5.2-5.8
mm. Face and palpi white. Tuft on head pale brown
with some white, piliform scales. Scapus white and
with large tuft of white piliform scales. Flagellum
white on upperside and slightly brownish on under-
side. Thorax ferruginous-brown with three longitudi-
nal pale lines. Forewing ferruginous-brown, usually
distally with green-grey tinge and with narrow white
strigulae. Costal-basal streak reaching middle of fo-
rewing and joining with first costal strigula. Medio-
basal streak slightly arcuate, and slightly shorter than
costal-basal streak. Dorsal-middle streak twice as
short as costal-basal streak. Four costal and four
oblique dorsal (except two apical ones, which are
straight) strigulae present, all white. First costal stri-
gula situated slightly beyond middle of forewing, and
connected basally with costal-basal streak. First dorsal
strigula wider than all other ones, situated before
middle of forewing, and reaching half width of fore-
wing. Second costal and second dorsal strigulae as
well as third costal and third dorsal connected at their
apexes and forming two narrow parallel fasciae.
Fourth costal and fourth dorsal streaks short and
straight. All strigulae proximally edged by blackish
scales. Apical spot small and elongate, distally bor-
dered by blackish scales. Apex of forewing edged
black. Cilia white, costally slightly fuscous. Hindwing
and cilia grey-creamy.
Male genitalia (fig. 21). — Asymmetrical. Uncus
reaching apex of valva. Vinculum triangular. Each
valva with small tooth-like ventral process near apex.
Transtilla symmetrical, U-shaped. Sternite 8 rather
narrow, and slightly arcuate in anterior view. Tube-
like aedeagus slightly shorther than valva and distally
somewhat narrowed. Small hook present before apex
of aedeagus.
Biology. — Host plant: Populus pruinosa Schrenk.
Mine (Fig. 31) on upperside of leaf, slightly folded,
relatively small, usually near leaf margin.
Distribution. — Turkmeniya and
Tadzhikistan.
southern
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Kumata, T., 1982. A taxanomic revision of the Gracillaria
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Kuznetzov, V. I., 1956. Novye listovertki (Tortricidae) mo-
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Kuznetzov, V. I., 1975. Novye vidy nizshikh cheshuekry-
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Kuznetzov, V. I., 1978. Novye vidy nizshikh cheshuekry-
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Kuznetzov, V. 1, 1979. Novye miniruyushchie moli-pes-
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SSSR. - Trudy Zoologicheskogo instituta Akademii
Nauk SSSR 88: 77-84.
Kuznetzov, V. I, 1981. Semeistvo Gracillariidae
(Lithocolletidae) - moli-pestryanki. - Opredelitel naseko-
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Kuznetzov, V. I. & N. I. Tristan, 1985. Obzor miniruyush-
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Gracillariidae) Palearkticheskoi fauny. +
Entomologicheskoe obozrenie 64: 177-199.
Mastumura, S., 1931. 6000 Illustrated insects of Japan-
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Noreika, R. V., 1991. Obzor fauny molei-pestryanok
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Received: 1 October 1991
Accepted: 27 February 1992
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
BOOK REVIEWS
Block, W., 1992. An annotated bibliography of antarctic in-
vertebrates (terrestrial and freshwater). — British Antarctic
Survey, Natural Environment Research Council,
Cambridge U. K.: 1-263, figs [ISBN 0 85665 148 6].
Available from British Antarctic Survey, NERC, High
Cross, Madingley Road, Cambridge CB3 OET, United
Kingdom.
In total 1430 references presented in alphabetical
order per author, are given. Each reference has key
words for the taxonomic groups. Abstracts provided
have been arranged per aspect (systematics, ecology).
The many indices (classification, taxonomic, species,
author, subject, geographical) make this publication a
welcome resource for anyone starting a study on ant-
arctic invertebrates. The work seems to be compiled
carefully and certainly gives many references which
cannot be traced easily from abstracting services. Just
one remark. The publication is spaciously printed in
a unnecessarily large typeface on luxurious paper. |
would suggest that an organisation working in area
where raw material for the production of paper is vir-
tually non-existent, adheres to a more reserved use of
paper.
[J. van Tol]
Heie, O. E., 1992. The Aphidoidea (Hemiptera) of
Fennoscandia and Denmark. IV. Family Aphididae: Part
1 of tribe Macrosiphini of subfamily Aphidinae. — Fauna
Entomologica Scandinavica 25: 1-189, figs. 1-585,
colour plates I-II. [ISBN 90 04 09514 4]. [ISSN 0106-
8377]. Published by E. J. Brill, Leiden. Price Hfl. 90.-.
From volume 25 onwards the series Fauna
Entomologica Scandinavica is published in the size of
Tijdschrift voor Entomologie, also using a two-col-
umn format. Although this is certainly an advantage
from many points of view, the author will regret that
the design of his work, published in at least six parts,
has changed for a second time. Descriptions and illus-
trations have the same high standard as the previous
works of the author. Keys are provided for the genera
and the species alike, with separate keys for the apter-
ous viviparous females, the alate viviparous females
and the oviparous females, whenever needed.
Printing as well as contents make this book a valu-
able addition to every biological library.
[J. van Tol]
Hansen, M., 1991. The Hydrophilid Beetles. Phylogeny,
classification and a revision of the genera (Coleoptera,
Hydrophiloidea). — Biologiske Skrifter Kongelige Danske
Videnskabernes Selskab 40: 1-367, figs 1-340. [ISSN
0366-3612, ISBN 87-7304-226-9. Price 700 Danish
Crowns].
42
Publication of “The Hydrophilid Beetles’ addresses
several long standing problems in the study of this
group of insects. Although many papers about
Hydrophiloidea were published by authors as
d'Orchymont and Balfour-Browne, an overview of
the group (146 genera, ca 2400 species) was not avail-
able up to now. Furthermore there was no agreement
as to the definition of the group as a whole.
The paper consists of three major parts: (1) a phy-
logenetic analysis of the Hydrophiloidea leading to a
reclassification of the group, (2) descriptions of and
keys to all genera and suprageneric taxa, (3) a cata-
logue of genus- and family-group names of
Hydrophiloidea with nomenclatorial details.
A detailed comparison of all known hydrophiloid
genera has not been published since the description of
tropical genera started in the 19th century and was
therefore long overdue. Most of the proposed classifi-
cations were rather subjective and usually without
any argumentation. Hansen's attempt towards a clas-
sification based on arguments is a large step forward.
The last supraspecific classification for the whole
group on a worldwide basis was published in 1924 by
Knisch in an issue of the Coleopterorum Catalogus.
From that time on numerous new genera were de-
scribed and articles treating supraspecific taxa or re-
gional faunas published. Newcomers attempting to
identify a tropical hydrophilid to at least generic level
were convicted to wade through decennia of the
‘Zoological Record’, usually ending with the conclu-
sion that the particular group one was looking at was
badly in need of revision. The keys, descriptions and
figures in Hansen's new book will in most cases en-
able an unambiguous identification to genus. For
each genus a description, a habitus illustration, the
number of species included and the global distribu-
tion is given. It is unfortunate that a summary of the
distributional data in a biogeographic analysis has not
been tried.
The extensive catalogue of family and genus group
names is especially important. It will most certainly
contribute to the nomenclatorial stability of the
group.
The book is well printed and bound. The quality of
the figures is excellent. Separate indexes are provided
on species, generic and suprageneric level. This book
is a must for everybody who's interest in
Hydrophiloidea goes beyond a regional fauna.
Besides I recommend it to anybody interested in the
general classification of Coleoptera. It may also be
useful for ecologists who need to sort samples from
more exotic locations.
J. Huijbregts)
THOMAS PAPE
Zoological Museum, Copenhagen, Denmark
PHYLOGENY OF THE TACHINIDAE FAMILY-GROUP
(DIPHRERANCAEMPIIRAMAE)
Pape, T., 1992. Phylogeny of the Tachinidae family-group (Diptera: Calyptratae). — Tijdschrift
voor Entomologie 135: 43-86, figs. 1-11, tabs. 1-5. [1ssN 0040-7496]. Published 15 July 1992.
Family-level phylogeny within the Tachinidae family-group (Oestroidea) is analyzed using the
computer package ‘Hennig86’. All families except the Rhinophoridae are split into a number of
groups, most of which are given subfamilial status, and terminal taxa are scored for a total of 50
characters. Outgroups are Anthomyiidae and Muscidae. The favoured cladogram lists the
Sarcophagidae as the sister group of the Tachinidae, the Oestridae as the sister group of the
Calliphoridae, and the Rhinophoridae as the sister group of the clade Calliphoridae +
Oestridae. The monophyly of all taxa of family rank and above within the Tachinidae family-
group is corroborated by explicit listing of apomorphic character states. Implications of various
interpretations of character states and transformation series are discussed.
Mystacinobia zelandica Holloway, 1976 is accepted as a member of the Tachinidae family-
group, but excluded from the analysis because of difficulties in interpreting character state ho-
mologies.
Dr. Thomas Pape, Zoological Museum, Universitetsparken 15, DK - 2100 Copenhagen,
Denmark
Key words. — Phylogeny, Tachinidae family-group, Oestroidea, Calyptratae, Diptera,
Hennig86.
CONTENTS Crosskey 1980a, Palearctic: Soós 1984-?, Australasian
/ Oceanian: Evenhuis 1989) more or less implicitly list
Introduction Mau ARE INTERNE 43 or issue the Diptera families in some kind of phyloge-
Terminology, methods and choice of terminal netic sequence. With respect to what is here called the
CAR At UE PRES HELL. OL AUREA 44 Tachinidae family-group, the Calliphoridae are usual-
@haracterstjand/charaeteristatese. nee ee 47 ly treated first, followed by the Sarcophagidae,
Unscored characters of potential phylogenetic Rhinophoridae and Tachinidae respectively (the latter
Valuese sante ee RENE ECM Er ARE be ino, 63 two sometimes united). The Oestridae take up various
Clades of the Tachinidae family-group ................ 69 positions: issued first (Palearctic catalogue), situated
@Oestridae ren REEL 71 between the Calliphoridae and the Sarcophagidae
Rhinophonidae m. MENO NEGER 72 (Australasian/Oceanian catalogue), or following the
Sarcophagidae VERAN TEE en 74 Tachinidae (other catalogues). Other, more explicit
Ia chinida West ente PV 75. phylogenetic hypotheses have proposed a sister group
Calliphoridae VALERA, IS UN relationship between the Oestridae (in part) and the
@alliphoridacen @estridact men 78 Tachinidae (Roback 1951), between the Rhino-
SarcophagidactHWliachinida ce rennen 78 phoridae and Sarcophagidae (Rohdendorf 1967),
Rhinophoridae + (Calliphoridae + Oestridae) ..78 between the Rhinophoridae and Calliphoridae
dachinidaesfarnilyzoroup EAP Re 78 (Tschorsnig 1985a), between the Rhinophoridae and
IMIVSIACINO IME 78 Tachinidae (Wood 1987b), between the Oestridae
Discussions ren FSB 79 and all other members of the Tachinidae family-group
Necknowledgements pester tee RER neo ees 81 (Hennig 1976, Tschorsnig 1985a), or between the
Referencestae Me MERIEN tone i). LER DTL ADG 81 Calliphoridae and all other members of the
Tachinidae family-group (Griffiths 1982). Shewell
I (1987b: 1162) was of the opinion that ‘biological and
NTRODUCTION
All recent regional Diptera catalogues (Nearctic:
Stone et al. 1965, Neotropical: Papavero 1966-?,
Oriental: Delfinado & Hardy 1977, Afrotropical:
distributional data suggest [that the Sarcophagidae]
are younger than both the Calliphoridae and the
Tachinidae’. In strict cladistic terms, this hypothesis
implies that Shewell considers the Sarcophagidae to be
43
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
the sister group of either the Oestridae, the
Rhinophoridae, or both — although this actually may
not have been the intention. Only McAlpine (1989)
has made an attempt of fully resolving the phylogeny
at the family level and he considered the clade
[Calliphoridae + Mystacinobiidae] + Sarcophagidae as
the sister group of the clade Oestridae + [Rhino-
phoridae + Tachinidae].
The primary objective of the present paper is to test
these hypotheses through reconsideration of available
data and incorporation of additional data. In order to
be as objective as possible and to facilitate and en-
courage future retesting, I have applied a numerical
cladistic approach including close to all (non-redun-
dant) characters for which I have been able to obtain
clearcut scorings. Several characters of potential phy-
logenetic value are discussed but not included in the
analyses because of my uncertainty with regard to
their proper scoring. It is hoped that the detailed an-
notations for most characters will serve as the basis for
extensive discussions on character (state) interpreta-
tion as well as promote a search for additional mor-
phological, behavioural and biochemical evidence.
The Tachinidae family-group contains the well
known blowflies, botflies, flesh flies, tachinid flies,
and woodlouse flies. Whether this entire group is giv-
en rank of family (Griffiths 1972), family-group
(Griffiths 1982) or superfamily (Hennig 1958,
McAlpine et al. 1981, Pape 1986a) is of minor con-
cern for the present discussion, and I have simply fol-
lowed current use of family rank for the subgroups in-
cluded. It should be mentioned that the Tachinidae
family-group in the present sense equals the
Calliphoroidea of Hennig (1958), the Tachinoidea of
Rohdendorf (1977) and Pape (1986a), and the
Oestroidea of McAlpine et al. (1981). The latter is
formally correct in the sense of the International
Code on Zoological Nomenclature as the oldest fam-
ily-group name within this taxon is based on Oestrus
Linnaeus (C.W. Sabrosky, in prep.). Griffiths (1972)
used Girschner's (1893) broad definition of the
‘Tachinidae’ for this taxon but later modified the
name to the “Tachinidae family-group’ (Griffiths
1982). I have adopted the latter suggestion rather
than applying a formal superfamily as I prefer to ap-
ply superfamily level much further down the clado-
gram of Diptera, as has been done by Griffiths
(1972), Crosskey (1980a), Woodley (1989), and
Thompson (1990). Furthermore, I prefer
“Tachinidae family-group to ‘Oestridae family-
group’ as the former is already in use (Griffiths 1982,
Thompson 1990), and the latter may easily be mis-
interpreted as a group consisting of what is here con-
sidered subfamilies of an Oestridae sensu lato, like the
Oestroidea of Hackman (1980) or the even more re-
stricted Oestroidea of Papavero (1977).
44
Paraphyletic assemblages of families arising when
excluding the Tachinidae family-group from more
inclusive clades, e. g. from the Calyptratae, are here
described as ‘non-tachinoid’, e.g. non-tachinoid cal-
yptrates.
‘TERMINOLOGY, METHODS AND CHOICE OF TERMINAL
TAXA
Terminology is straightforward and follows in ge-
neral McAlpine (1981) and Teskey (1981). The only
important exception is in the terminology of the
structures of the male terminalia, where I have fol-
lowed Michelsen (1988) and Wood (1990, 1991).
The character matrix (table 2) was analysed with
the computer package Hennig86 (version 1.5, copy-
right J.S. Farris 1988) on an IBM PS/2-30 (8086 mi-
croprocessor). Cladograms (trees) were generated
with ‘mh*;bb*;’ rather than any combination of im-
plicit enumeration (ie) and branch-swapping (bb). As
noted by both Farris (1988: Hennig86 documenta-
tion) and Fitzhugh (1989), this includes the possibil-
ity of not finding all minimum-length cladograms,
but it may be a convenient trade-off as it is consider-
ably faster and therefore will allow many more trials.
The procedure chosen was judged to have no family-
level impact on the resulting equally most parsimoni-
ous cladograms, although this hypothesis needs cor-
roboration.
All character states were treated as unordered when
producing a first set of cladograms (changing the
Hennig86 default by ‘cc-.;’), and a successive weight-
ing procedure (‘xsteps w; in combination with
‘mh*;bb*; and ‘cc; until weights no longer change)
was applied to any initial set of equally most parsimo-
nious cladograms. The latter procedure, which
weights characters according to their best fit to the
cladograms concerned through multiplying consis-
tency and retention indices and scaling these in the
range 0-10, means that clades based on more reliable
characters are given higher priority (Fitzhugh 1989).
Thus, the resulting cladogram(s) may be different
from any of the initial ones. Where successive weight-
ing could not reduce the number of equally most par-
simonious cladograms to one, a Nelson consensus
tree was produced by the ‘nelsen;’ command. As dis-
cussed by Carpenter (1988), any of the equally most
parsimonious cladograms will represent a stronger
hypothesis than the Nelson consensus tree, but I have
chosen the latter due to lack of good arguments for
selecting amongst these (often numerous) resulting
cladograms.
Transformation series for characters involving
more than two states were deduced from the weight-
ed Nelson consensus tree obtained through the proce-
dure outlined above. With this new input of phyloge-
netic information, data were reanalyzed.
The number of characters (here defined as transfor-
mation series, see for example Mickevich (1982) and
Platnick (1979)) in biological taxa is usually very
large, and any phylogenetic analysis can only take a
fraction of these into consideration. Exclusion of
those characters that would be scored identically for
all terminal taxa analysed and thus carry no phyloge-
netic information is straightforward. Similarly, those
unique (autapomorphic) character states in a given
analysis that define terminal taxa may be excluded as
well, as they have no impact on the relationships
between the taxa. In between is a potentially large
number of characters that possess phylogenetic infor-
mation in the given context. Of these, some are left
unscored simply because they remain unknown to the
observer, or because severe problems with regard to
interpretation of homology or proper delimitation
into states defer clearcut decisions. Therefore, the
present data matrix (table 2) contains only those non-
redundant characters for which I have been able to
obtain ‘reasonably unambiguous’ scorings. In the
present analysis, these characters are mostly well
known and generally used in descriptive works, but
the scorings — which are the heart of the analysis —
represent hypotheses that should be tested through
more information (morphological, biological, phylo-
genetic, etc.) on the terminal taxa considered.
All scorings are meant as features of the hypotheti-
cal ancestor of the group in question, and I have used
‘groundplan state unknown’ (scored simply as —')
whenever a character is inapplicable or when more
than one state occurs in a pattern that does not sug-
gest one to be a more probable groundplan feature
than the other(s).
When scoring larger groups one almost always has
to deal with exceptions, i.e. one or more species with-
in a group showing a character state differing from
that of the hypothetical groundplan chosen. Rather
than scoring every deviating member separately,
which would create an excessive number of terminal
taxa; or using ‘state unknown’ for all instances where
exceptions were found, which would greatly reduce
the amount of information actually at hand, general-
izations have sometimes been made. These are dis-
cussed in the annotations. Concurrently with the
emergence of explicit hypotheses on the basal splits
within each of the terminal taxa, these generalizations
— and thereby the hypotheses emerging from them —
become open to rigorous testing.
Two non-tachinoid members of the Calyptratae
have been included in the analysis: the Anthomyiidae
and the Muscidae. Both families were designated as
outgroup relative to the Tachinidae family-group
(outgroup = 0.1;’). The Fanniidae are sometimes in-
cluded within the Muscidae or considered their sister
Pare: Phylogeny of the Tachinidae family-group
group, but they are here excluded from the analysis.
All Fanniidae have the first anal vein (A, + CuA, or
just A,) shortened and may for this reason be consid-
ered as members of a group also containing the
Muscidae and the Tachinidae family-group. The re-
duction of A,, however, is associated with a strongly
curved second anal vein (A,), and this feature puts se-
rious doubt on the interpretation of the shortening of
the first anal vein as homologous to the condition
seen in the Muscidae. Pont (1986a: 41) even men-
tioned that the Fanniidae ‘may be the primitive sister-
group of the rest of the Muscoidea, or even of the
Calyptrata’, which finds interesting support in the
analysis of ribosomal RNA phylogeny provided by
Vossbrinck & Friedman (1989). McAlpine (1989)
treated the Fanniidae as the sister group of the
Muscidae, but he did not apply a numerical cladistic
approach and his hypothesis needs testing by inclu-
sion of more data. Non-tachinoid calyptrate phyloge-
ny is beyond the purpose of the present paper and I
have simply accepted Pont's statement cited above.
Anyway, the in- or exclusion of this family in the phy-
logenetic analysis presented below would probably
have no impact on groundplan estimates for the
Tachinidae family-group nor on hypotheses of phylo-
genetic relationships between its members.
Except for the Rhinophoridae, which have been
scored as a terminal taxon, I have split the tachinoid
families into a number of subgroups to enhance the
corroboration of family groundplan estimates. Note
that my use of subfamilial (or tribal) endings should
not necessarily be taken as an endorsement of subfa-
milial (or tribal) rank for these groups; they are sim-
ply more easily referred to as such for the present pur-
pose. These subgroups generally equal named and
well-known subfamilies or tribes, but a special case is
made for the Tachinidae, the subfamilial classifica-
tion of which is highly premature in so far as few cur-
rently recognized subfamilies emerge as well corrobo-
rated monophyletic groups. Crosskey (1980b:
822-823), for example, was of the opinion that the ‘-
traditional’ Tachininae ‘almost certainly [are]
polyphyletic’. According to Wood (1987c: 1201) ‘the
Dexiinae is the only subfamily that can be defined on
the basis of a synapomorphy, and Tschorsnig
(1985b: 120) stated that the monophyly of the
Tachininae as well as the Exoristinae is unsupported
(Die Monophylie sowohl der Tachininae als auch
der Exoristinae kann nicht begriindet werden’). Still,
however, recent suggestions on how to divide the
Tachinidae into major subgroups and hypotheses of
the phylogenetic relationships between these are al-
most exclusively restricted to the traditional concept
of the four subfamilies Phasiinae, Exoristinae,
Tachininae and Dexiinae (e.g. Shima 1989, Richter
1991)
45
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
For the present analysis I have applied a much wid-
er concept of the Tachininae than the one referred to
above, namely by uniting all species which embryo-
nate (or are suspected to embryonate) their eggs in the
uterus for at least some time after fertilization. This
concept was originally proposed by Wood (1985: 8),
who regarded the ‘possession of a distensible ovisac
for embryonating eggs .... as a synapomorphy of those
tachinids that possess it.” I have recognized the
Phasiinae as applied by Wood (1987c), and a ‘rest-
group’, here named the ‘Exorista genus-group’, in-
cluding the non-embryonating, non-phasiine
Tachinidae (mainly genera like Aplomya Robineau-
Desvoidy, Exorista Meigen, Medina Robineau-
Desvoidy, Meigenia Robineau-Desvoidy, Phorocera
Robineau-Desvoidy, and Winthemia Robineau-
Desvoidy, see Wood (1972, 1985)). This latter group
is thus possibly para- or even polyphyletic, but the
classification is considered superior to the more tradi-
tional division of the Tachinidae into four subfami-
lies as the suspected non-group is much less inclusive
in terms of genera and species. The Dexiinae will thus
be subordinate within the present Tachininae and are
not scored separately.
The Rhinophoridae need some minor comments.
The genus Mimodexia Rohdendorf was described in
the Sarcophagidae (Rohdendorf 1935) and later con-
sidered a distinct subfamily, Mimodexiinae, of the
Calliphoridae with a sister group relationship to the
remaining members of this family (Lehrer 1970).
Tschorsnig (1985a: 16) proposed Mimodexia as a sen-
ior synonym of Callidesia Kugler (Rhinophoridae).
The phallus is highly rhinophorid-like and possesses
the apical bend of the median sclerotization listed by
Pape (1986a) as a synapomorphy for Tromodesia
Rondani and Callidesia, and features like wing pat-
tern, wing venation, and narrowing of frontal vitta al-
so agree with this proposal (Rohdendorf 1935: figs.
79-82, Kugler 1978: figs. 10-11). Thus, I accept the
synonymy and thereby the assignment to the
Rhinophoridae.
Also, I prefer to include the New World Bezzimyia
Townsend and the Oriental Malayia Malloch in the
Rhinophoridae. Crosskey (1977) discussed the famil-
ial assignment of Bezzimyia and considered this genus
in the Tachinidae (exclusive of the Rhinophoridae),
because ‘the head facies … is conspicuously tachinid,
and the enlarged fore tarsi of the female are of the
minthoine type’ (p. 13). This was accepted without
further comments by Pape (1986a). However, still
more evidence is accumulating in favour of an inclu-
sion in the Rhinophoridae (Pape, unpubl.). The phal-
lus of what appears to be an undescribed species of
Bezzimyia (Mexico: Chiapas, El Triunfo, 1 male, 13-
15.v.1985, A. Freidberg, deposited in the Depart-
ment of Zoology, University of Tel Aviv) is strikingly
46
similar to the phallus found in the Rhinophoridae
(fig. 3), and as subscutellum, lower calypteres and ab-
dominal sternite 2 agree more with the
Rhinophoridae than with the Tachinidae, I prefer to
include Bezzimyia in the former. The rhinophorid ap-
pearance may have been the reason why Sabrosky &
Arnaud (1965) put Bezzimyia in the Rhinophorinae
(as a subfamily of the Tachinidae). Crosskey (1976:
163, 209) listed Malayia as a genus incertae sedis
within his possibly polyphyletic Tachininae (i.e. the
traditional concept), but also Malayia possesses the
rhinophorid-like features here given for Bezzimyia.
Admittedly, they do not fit easily into the cladogram
of rhinophorid genera produced by Pape (1986a), but
lack of evidence does not corroborate anything. Thus,
the only argument for treating these rhinophorid-like
genera as Tachinidae is that they possess characters
not found in any (? other) rhinophorid but which are
encountered scattered in the Tachinidae, e.g. a later-
ally compressed fore tarsus (many Bezzimyia, some
undescribed Malayia), heavily setose facial ridges that
are elongated ventrally (Bezzimyia busckii Town-
send), or a lobate first flagellomere (some undescribed
Bezzimyia). But as long as they cannot be shown to
share but a few possibly derived character states with
any specific group of Tachinidae, the many apomor-
phic character states which are shared with the
Rhinophoridae make an affiliation with this family
most corroborated. Most probably, the description of
either mature eggs or the first instar larvae of Malayia
and Bezzimyia — still unknown for both genera — will
provide definite clues to their systematic position.
Apart from Bezzimyia and Malayia, some other unde-
scribed species of Neotropical and Australasian
Rhinophoridae have turned up lately (Pape in prep.),
and as these do not fit easily in the cladogram of Pape
(1986a), the family groundplan remains uncertain for
some characters.
The lack of a well corroborated infrafamilial clas-
sification of the Calliphoridae urges some explanation
of the subgroups used. I have treated the Toxotars-
inae as including the Sarconesiinae, Kuschelomyinae
and Netinae of Lopes & Albuquerque (1982), but
distinct from the Chrysomyinae and Rhiniinae, with
which they share a row of setae on the posterodorsal
surface of the stem vein (remigium). This is not be-
cause I endorse subfamily rank for these taxa, but
simply because a splitting will bring more detailed
scorings and therefore result in better corroborated
hypotheses of groundplans and therefore of phyloge-
netic relationships. The Auchmeromyiinae of Patton
(1935a) [originally spelt Auchmeromyinae] is here
widened and synonymized with the Bengaliinae as
used by James (1966) and the Bengaliinae and
Tricycleinae used by Lehrer (1970). This taxon will
probably, in addition to Auchmeromyia Brauer &
Bergenstamm and Bengalia Robineau-Desvoidy, con-
tain the genera Booponus Aldrich, Cordylobia
Grünberg, Hemigymnochaeta Corti, Neocordylobia
Villeneuve, Pachychoeromyia Villeneuve, Termi-
toloemus Baranov and Tricyclea Wulp, the species of
which all are exclusively yellowish or brownish
(Rognes 1991, pers. obs.). Moreover, all members
may have the spermathecal ducts arranged in the ‘all
separate’ configuration (see discussion under charac-
ter 25 below), and all genera of this taxon examined
by me (Auchmeromyia, Bengalia, Cordylobia,
Termitoloemus, Tricyclea) have an elongated first anal
vein (A, + CuA,), reaching, or almost reaching the
wing margin (see also Villeneuve (1924)). Rognes
(1991) mentions that the complete first anal vein is a
possible autapomorphy for the Auchmeromyiinae, al-
though he simultaneously proposes to maintain the
equally equipped genus Bengalia in a subfamily of its
own. I know of no character states shared by the non-
bengaliine Auchmeromyiinae, which may corrobo-
rate their monophyly. A similarity between
Auchmeromyia, Cordylobia and Bengalia was, in fact,
already noted by Patton (1935a: 229), who men-
tioned that ‘practically all the chaetotactic characters
lof Auchmeromyia) are common to Bengalia and
Cordylobia as well’.
The monophyly of the Ameniinae and Mesem-
brinellinae, respectively, seems corroborated beyond
any doubt (Crosskey 1965, Guimaräes 1977), and
the Phumosiinae, including Euphumosia Malloch and
Phumosia Robineau-Desvoidy, may be defined by the
character states macro- (or uni-) larviparous repro-
duction (Ferrar 1978), elongated spermathecae, and
the ‘all separate’ configuration of spermathecal ducts.
The Helicoboscinae were dealt with by Rognes
(1986), with whom I fully agree. Note that already
Patton (1939: 107) suggested a calliphorid affiliation
of this taxon: ‘It is not possible at present to be certain
of the systematic position of Helicobosca muscaria and
Helicobosca distinguenda, one point, however, is cer-
tain they do not belong to the subfamily
Sarcophaginae; the terminalia suggest relationships
with the Calliphorinae’. Verves (1990) transferred the
genus back into the Paramacronychiinae, but without
a detailed discussion on how to explain the absence of
the diagnostic sarcophagid groundplan features in
this taxon.
The subfamilies Calliphorinae, Luciliinae, Mela-
nomyinae, and Polleniinae have received a detailed
and precise treatment by Rognes (1991) and need no
further comment in this context. The subfamilial and
tribal phylogeny proposed by Lehrer (1970) will not
be thoroughly discussed in the present paper as this
author did not explicitly list the (derived) character
states defining clades as well as terminal taxa.
The Prosthetosomatinae, which were transferred to
Pape: Phylogeny of the Tachinidae family-group
the Calliphoridae by Pont (1980a), have not been in-
cluded as a separate (calliphorid) taxon as they in my
opinion represent a group of convenience. The con-
stitutive feature is the habit of all members infesting
termite nest-mounds (in itself hardly a clearcut char-
acter state), and they are known from the larval stag-
es only. They will probably fit into one or more of the
groups applied in the present paper when adults be-
come available.
The division of the Sarcophagidae into only three
subfamilies has been dealt with by Pape (1987a).
Oestridae in the present sense equals that of Wood
(1987a), and they have been divided in four subfam-
ilies: Cuterebrinae, Gasterophilinae, Hypoder-
matinae and Oestrinae. The Gasterophilinae have
been further split into two subgroups: Cobboldiini
with the genus Cobboldia Brauer on the one hand,
and Gasterophilini with Gasterophilus Leach and
Gyrostigma Brauer on the other. The systematic posi-
tion of the monotypic genera Neocuterebra Grünberg
and Ruttenia Rodhain within the Oestridae is uncer-
tain (Zumpt 1965; Wood 1986, 1987a), and I have
followed the tentative suggestion of Wood (1987a),
that they belong to the Cuterebrinae.
The monotypic Mystacinobia Holloway from New
Zealand has not been included in the analysis, neither
as a separate taxon nor by inclusion in one of the ac-
cepted taxa. No shared character states have been
found which will convincingly indicate inclusion
within any of the currently applied families, but I do
not consider this justification for a separate family.
The strongly autapomorphic morphology renders ho-
mologies of most character states highly tentative or
even inapplicable, and many scorings are open to se-
rious doubt. I prefer to accept Mystacinobia as a mem-
ber of the Tachinidae family-group and to treat it as a
genus incertae sedis. Possible phylogenetic affinities
are discussed further in the section dealing with clades
of the Tachinidae family-group.
CHARACTERS AND CHARACTER STATES
Characters discussed below are given numbers ac-
cording to their scorings in the data matrix (table 2),
starting with 0 rather than 1 following the procedure
of numbering characters in Hennig86 (version 1.5).
All characters have been split into two or more states
denoted by consecutive numbers (0-n). For the
convenience of the later discussion, states have — whe-
re possible — been arranged in the transformation se-
ries deduced from the initial analysis and with the
more plesiomorphic state having the lowest number.
Character 0. — Postocular setae; size
0 - equal size, 1 - alternating size, 2 - redu-
ced.
47
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Apart from Pape (1991), this character has not pre-
viously been employed for diagnostic purposes or
otherwise explicitly mentioned in connection with
the Tachinidae family-group. In the non-tachinoid
Calyptratae, the row of postocular setae consists of
equal-sized setae or these may increase gradually in
size towards the vertex. In the Tachinidae family-
group, this row consists of setae distinctly alternating
in size, and only two exceptions are known to me:
The Oestridae and the Miltogrammatinae
(Sarcophagidae). All Oestridae have their general se-
tosity greatly modified, with most setae soft and hair-
like. The postocular setae are bristly in the sense that
they are stiff and tapering, but obviously reduced in
length, and they show no alternation in size. As this
reduced condition could easily have evolved from any
of the other two states, the oestrid condition has been
scored as a separate state: reduced (2). In the
Miltogrammatinae, species of Macronychia Rondani
sometimes have postocular setae slightly alternating
in size, but the subfamily has nevertheless been scored
for equal sized postocular setae.
Character 1. — Postcranium; shape
0 - convex, 1 - concave.
The concave condition is found in all Oestridae
and in the subfamily Miltogrammatinae of the
Sarcophagidae. The groundplan of all other terminal
taxa has been scored as convex.
Character 2. — Imaginal mouthparts; functionality
0 - fully developed and actively used, 1 - re-
duced and never used for feeding purposes.
Within the Oestridae, all species possess reduced
mouthparts, and only species of Cuterebra Clark and
some Hypodermatinae have been seen imbibing wa-
ter from wet surfaces (Bennett 1955, Catts & Garcia
1963, Grunin 1965). Solutions of sugar or proteina-
ceous substances are never exploited. The non-oestrid
taxa have all been scored for fully functional mouth-
parts (0) as the very few instances of much reduced
mouthparts (e.g. Dexia Meigen and Phasiops
Coquillett of the Tachinidae, Chauliooestrus
Villeneuve of the Sarcophagidae [not Gasterophilidae
as listed) "in: »Pont-"@980b); "see Paper (1991)],
Villeneuviella Austen of the Calliphoridae) seem to
involve genera far from the basal splits in their respec-
tive subfamilies.
Character 3. — General integument; coloration
0 - non-metallic, 1 - metallic.
The Anthomyiidae and Rhinophoridae possess no
members with metallic colours, and the few examples
in the Muscidae (eg. Neomyia Walker),
Sarcophagidae (only Sarcophaginae, e.g. Sarcophaga
metallescens Bezzi, Chlorosarcophaga Townsend), and
48
Tachinidae (e.g. Gymnocheta Robineau-Desvoidy of
the Tachininae) may convincingly be characterised as
isolated homoplasies by reference to what is known of
infrafamilial phylogeny in these groups. In the sub-
family Cuterebrinae, metallic colours occur in
Neocuterebra squamosa Griinberg and Dermatobia
hominis (Linnaeus, Jr). Members of the genus
Cuterebra sensu lato generally have a shining integu-
ment and a few species present a distinct metallic
tinge, e.g. Cuterebra atrox Clark. Ruttenia loxodontis
Rodhain has no metallic coloration. The groundplan
of the Cuterebrinae has been scored as unknown with
regard to this character.
Metallic colours are not present in the
Gasterophilini. In Cobboldia, the Afrotropical species
C. loxodontis Brauer and C. roverei Gedoelst have tho-
rax and abdomen metallic blue and green respective-
ly, but the Oriental C. elephantis (Cobbold) is non-
metallic.
In the Calliphoridae, no member of the
Helicoboscinae and the Bengaliinae possess a metallic
coloured integument, and these have accordingly
been scored as non-metallic (0). In the Polleniinae,
metallic colours are generally absent but at least some
New Zealand Pollenia Robineau-Desvoidy are metal-
lic green (Dear 1986). The ‘Australasian’ concept of
this genus, however, is much broader than that con-
taining only those Palearctic species with soft, yellow-
ish, crinkled hairs amongst the thoracic setae, and as I
have been unable to differentiate Pollenia (s.1.) from
the remaining genera of Polleniinae, the groundplan
of the subfamily has been scored as unknown for this
character. Also, several species of Rhiniinae and
Melanomya Rondani of Melanomyinae are non-me-
tallic and the groundplan of these subfamilies has
been scored as unknown for this character.
Remaining calliphorid subfamilies have been scored
as metallic (1).
Character 4. — Imago; clothing setae
0 - unmodified setae, 1 - most setae hairli-
ke.
The hairy condition of all members of the oestrid
subfamilies is well known and needs no further com-
ment. Rhyncomya Robineau-Desvoidy (Rhiniinae)
and Palearctic Pollenia (Polleniinae) of the
Calliphoridae possess whitish or yellowish hairs
among the setae, but never to a degree comparable
with any botfly, and the hairs are crinkled and unlike
those found in the Oestridae.
Character 5. — Prosternum; setosity
0 - bare, 1 - setose.
In the Sarcophagidae, most members of subfamily
Sarcophaginae have the prosternum setose and the
groundplan has been scored as such. No
Paramacronychiinae and in the Miltogrammatinae
only Chrysogramma Rohdendorf (preocc., no valid
name at present) have setae on this sclerite. No
Rhiniinae and Polleniinae have a setose prosternum,
while the setose condition is widespread in all other
terminal taxa of Oestridae and Calliphoridae. In the
Tachinidae, I have not seen the setose condition in
the Phasiinae nor in the Exorista genus-group, but
both conditions occur in the Tachininae, and the
groundplan of the latter has been scored as unknown
for this character.
Character 6. — Proanepisternum; setosity
0 - bare, 1 - setose.
A setose proanepisternum (upper, depressed part of
propleuron) may be found scattered throughout the
Sarcophagidae, but always in distinct clades that are
not engaged in the basal dichotomies of their respec-
tive subfamilies. Therefore, I feel confident by scoring
the groundplan of the contained subfamilies as bare
(0). In the Calliphoridae, I have found no instances of
a setose proanepisternum in the Mesembrinellinae,
Polleniinae and Rhiniinae — all scored as bare (0) —
while both states occur in the Bengaliinae (setose in
Tricyclea, bare in others) and Melanomyinae (bare in
Melanomya, setose in Melinda Robineau-Desvoidy),
which have been scored as unknown. Remaining cal-
liphorid subfamilies have been scored as setose (1) as
I know of no exceptions. No Rhinophoridae possess a
setose proanepisternum, and in the Tachinidae no
members of the Phasiinae and the Exorista genus-
group seen by me have this surface setose. Both con-
ditions occur in the Tachininae, the groundplan of
which accordingly has been scored as unknown. The
Oestridae probably have the setose condition as a
groundplan feature. In the Cuterebrinae the proane-
pisternum is bare in Neocuterebraand Ruttenia, setose
in Dermatobia Brauer and Cuterebra. Gasterophilus,
Gyrostigma and Cobboldia of the Gasterophilinae all
possess a setose proanepisternum. In the
Hypodermatinae, the setose condition is found in
Oestromyia Brauer, but only occasionally in the
Hypodermatini. No Oestrinae possess a setose proan-
episternum.
Character 7. — Postalar wall; setosity
0 - bare, 1 - setose.
The typical pattern of postalar setae is an isolated
tuft of setae centrally on the postalar wall. Several
genera of the sarcophagid subfamily Sarcophaginae
possess postalar setae in this configuration, but no
postalar setae are found in the genus Tricharaea
Thomson, which is often considered close to the ba-
sal split(s) of this taxon, and the sarcophagine
groundplan has been scored as unknown. No mem-
bers of the Paramacronychiinae possess postalar setae,
Pape: Phylogeny of the Tachinidae family-group
and where a few postalar setae occur in the Milto-
grammatinae, e.g. Metopia brasiliana Townsend
(Pape 1987), these are probably homoplasic as prob-
ably basal groups like Macronychia and
Eumacronychia Townsend do not possess any postalar
setae. Both conditions are found in the
Melanomyinae (bare in Melanomya, setose in
Melinda), but of general occurrence in the other calli-
phorid subfamilies, which have been scored as setose
(1). The oestrid taxa have been scored as setose (1), al-
though apparently bare conditions occur in the
Oestrinae, e.g. in Tracheomyia macropi (Frogatt) and
in some Gasterophilus spp. These exceptions all pos-
sess setae higher up on the postalar wall which are
confluent with the general setosity of notum, but I
have not been able to decide whether this setosity is
homologous to true postalar setae.
I have not found any Tachinidae with a setose
postalar wall and the three subgroups here recognized
have been scored as bare (0).
Character 8. — Metasternum; setosity
0 - bare, 1 - setose.
Note that what is here called metasternum (i.e. the
convex sclerite situated in front of the hind trochant-
ers) probably includes pleural elements, and the
metasternum proper is largely invaginated. I have
found no Anthomyiidae, Muscidae, Rhinophoridae
or Tachinidae with a setose metasternum. In the
Sarcophagidae metasternal setosity is present in all
Sarcophaginae, absent in Paramacronychiinae, and in
the Miltogrammatinae only present in Chryso-
gramma. All Oestridae seen by me have the metaster-
num setose. The calliphorid subfamilies have been
scored as setose (1) as the few exceptions, e.g. some
but not all Melanomya, are considered as derived from
the setose condition.
Character 9. — Meron; setosity
0 - bare, 1 - with a vertical row of bristles, 2
- with a patch or tuft of hairs.
Usually, the row of meral bristles runs vertically at
the posterior margin of the sclerite and may follow
this where it curves forwards along the anteroventral
margin of the metathoracic spiracle. Smaller species
may have the number of bristles reduced to 2-3, e.g.
many Rhinophoridae and small Tachinidae, while
larger species often have numerous additional hair-
like setae, especially anterior to the bristles. Very few
Tachininae, e.g. the genera Tarassus Aldrich
(Neotropical) and Lophosiosoma Mesnil (Oriental),
lack meral bristles altogether, but these are considered
reversals and the groundplan of the Tachininae has
been scored as 1. Rognes (1986) considered the mer-
al hairs found in the Oestridae as non-homologous to
the meral bristles of the remaining Tachinidae family-
49
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
group, possibly because the hairs usually form a patch
rather than a vertical row. In my opinion, the identi-
cal position compared to that of the bristly row of the
non-oestrid members of the Tachinidae family-group
should be taken as a priori evidence of homology. The
hairlike appearance of the meral setae is most parsi-
moniously regarded as part of the general hairiness,
and it should be noted that the meral hairs sometimes
are developed as a row of fine bristles, e.g. in
Dermatobia hominis (Cuterebrinae) (Wood 1987a,
pers. observation). Sabrosky (1986: 40) noted that
Cuterebra has the meron equipped with ‘a tuft of stiff,
bristly, black hairs in most species’.
Meral setae are also found in non-tachinoid
Calyptratae, especially well developed in the genus
Eginia Robineau-Desvoidy (Muscidae) but also in
more typical Muscidae (Pont 1986b, McAlpine
1989). A few Anthomyiidae possess meral setae, e.g.
Eutrichota Kowarz (Huckett 1987) and ‘McAlpine’s
fly’ (pers. obs.), which tentatively has been assigned to
the Anthomyiidae (Ferrar 1979, 1987). Only in
Eginia, however, is the configuration reminiscent ofa
(very short) vertical row of long and bristly setae.
Pont (1977) placed Eginia with some related genera
in their own family, Eginiidae, but later transferred
the group to the subfamily Phaoniinae of the
Muscidae (Pont 1986b, as Eginiini). Skidmore
(1985) was convinced that Eginiini did not belong in
the Muscidae and he suggested affinities to be with
the Tachinidae or the Calliphoridae. This decision,
however, was primarily based on the presence of well
developed parastomal bars in the cephalopharyngeal
skeleton of what was thought to be a puparium of
Eginia ocypterata (Meigen). As this puparium seems
to belong to the Sciomyzidae (V. Michelsen, pers.
comm.), the evidence for excluding Eginiini from the
Muscidae is strongly reduced. A discussion of the
systematic position of Eginiini is given by McAlpine
(1989: 1498).
Character 10. — Subscutellum, shape
0 - concave, 1 - slightly swollen, 2 - strong-
ly swollen.
The concave condition found in all Anthomyiidae
and Muscidae examined by me is easily distinguished
from the swollen conditions (1, 2). A similar concave
condition is found in the three sarcophagid subfami-
lies except in Nyctia Robineau-Desvoidy of the
Paramacronychiinae where the slightly swollen con-
dition obviously has evolved independently. The ta-
chinid taxa are here scored for the strongly swollen
condition (2), although some exceptions occur, e.g.
Cinochira Zetterstedt and Cylindromyiella Malloch.
Crosskey (1976: 36), however, noted that it ‘is not
completely certain that Cylindromyiella is a tachinid’.
Downes (1986) considered the groundplan of the
50
Tachinidae to be characterised by a medium-sized
subscutellum, which led him to assume that ‘the sub-
scutellum has become enlarged independently in
more than one lineage’ (p. 15) within the family. In
my opinion, it is more parsimonious to consider the
strongly swollen subscutellum of the Tachinidae as a
groundplan feature.
Within the Oestridae, the subscutellum of the
Cuterebrinae (Wood 1987a: fig. 9) and
Gasterophilinae is considered as state 1, while the
subscutellum of the Hypodermatinae (Wood 1987a:
fig. 10) is distinctly larger and is scored as such (2). In
the Oestrinae, Oestrus has a strongly swollen subscu-
tellum but other states occur, like the almost concave
subscutellum of Cephenemyia Latreille and
Pharyngomyia Schiner (Zumpt 1965). The subscutel-
lar groundplan of the Oestrinae has accordingly been
scored as unknown.
All Rhinophoridae have a slightly swollen subscu-
tellum (Wood 1987b: figs. 2,3), and so have the cal-
liphorid subfamilies Ameniinae, Bengaliinae,
Helicoboscinae, Mesembrinellinae and Polleniinae,
which all possess a subscutellum that definitely is con-
vex, but still much less swollen than the condition in
the Tachinidae. As both the concave and the slightly
swollen conditions occur in the Melanomyinae, this
taxon has been scored as unknown for this character.
As an intermediate condition, the slightly swollen
condition is poorly defined. More precise morpho-
logical descriptive terms will probably increase the
number of states and add important phylogenetic in-
formation.
Character 11. — Anatergite; setosity
0 - bare, 1 - setulose.
If present, the laterotergal ‘hairs or setae of
McAlpine (1989: 1498) [= infrasquamal setulae of
Shewell (1987b: fig. 108.23)] are arranged in a small
patch or cluster on the anatergite just below the low-
er calypter. I have not found infrasquamal setulae in
any Anthomyiidae, and in the Muscidae only in
Neomyia, and I have scored the groundplan of both
families as bare (0). All Calliphoridae, Rhinophoridae
and Tachinidae seem to possess infrasquamal setulae
(I have, however, seen some specimens of the rhino-
phorid Paykullia kugleri (Herting) with few and even
no setulae). Among the Sarcophagidae, all species of
Paramacronychiinae and Sarcophaginae have infra-
squamal setulae, but the large majority of
Miltogrammatinae have the anatergite completely
bare. Some exceptions, however, occur, e.g.
Eumacronychia and Macronychia, and occasional
specimens of Xiphidiella Villeneuve, and I have
scored the miltogrammatine groundplan as unknown
for this character. No Oestridae possess anatergal set-
ulae.
Character 12. — Metathoracic spiracle; configuration
of fringe
0 - continuous along posterior, ventral and
anterior rim, 1 - divided into an anterior
and a posterior fringe.
The Muscidae and the Anthomyiidae are scored for
a continuous fringe, although this fringe often is dis-
tinctly thinned ventrally. Within the Tachinidae fam-
ily-group, I know of only the following exceptions
from the divided fringe: Mystacinobia, which is dis-
cussed separately below, has no fringe at all. The
Mesembrinellinae were given family rank by
Guimaräes (1977), probably prompted by Crosskey's
(1965: 43) statement that they ‘may not be
Tachinoidea at all’. Their metathoracic spiracle devi-
ates strongly from that of other Calliphoridae in be-
ing equipped with a large continuous fringe, much
like some Muscidae (e.g. Phaonia Robineau-
Desvoidy). Bequaertiana Curran of the
Rhinophoridae has almost no fringe, which added to
Zumpt's (1956) conviction that this genus was an
acalyptrate. The generic cladogram of Pape (1986a),
however, suggests that the spiracular configuration of
Bequaertiana is apomorphic. Groundplan estimates
of the Oestridae are difficult as much variation oc-
curs. In the Cuterebrinae, Neocuterebra and Ruttenia
have a continuous fringe, all Cuterebra a divided
fringe. In the Gasterophilinae species of Cobboldia
have a divided fringe, those of Gasterophilus and
Gyrostigma have a continuous fringe. All
Hypodermatinae have a continuous fringe while all
Oestrinae have a divided fringe.
Several small Tachinidae and Calliphoridae (e.g.
Cinochira, some Melanomya), and all of
Rhinophoridae have small metathoracic spiracles, but
they still provide good examples of the divided condi-
tion. From the SEM figures given by Crosskey (1977:
figs. 41-44), it may be observed that Melanophora
roralis (Linnaeus), Phyto discrepans Pandellé and
Stevenia atramentaria (Meigen) possess distinctly di-
vided metathoracic fringes, and only the fringe seen
in Rhinophora lepida (Meigen) could perhaps be tak-
en for continuous, although even here a distinct break
of the fringe can be observed ventrally (which is al-
most to the right in Crosskey's figure).
Character 13. - Metathoracic spiracle; relative size of
anterior and posterior lappets
0 - both lappets small, 1 - narrow anterior
lappet and large posterior lappet, 2 - both
lappets large.
This attempt of fitting the configurational multi-
tude of metathoracic spiracular lappets into a few
states based on relative size alone may seem too sim-
ple as ‘a wide range of “opercular” metathoracic spir-
acles occurs in Tachinidae-Calliphoridae-Sarco-
Pare: Phylogeny of the Tachinidae family-group
phagidae’ (Crosskey 1977: 8). On the other hand,
scoring most types of fringe configurations different-
ly would cause an extreme splitting of taxa that would
be difficult (and very time-consuming) to score at the
present state of knowledge — and actually carry less
phylogenetic information as far as interfamilial rela-
tionships are concerned. Therefore, I have tried to re-
duce the number of groundplan types, and the
present three-state character seems in agreement with
Crosskey (1977: 8), as his ‘representative[s] of two
very common types’ fit my states 1 and 2 perfectly.
The Helicoboscinae, all Oestridae except those
with continuous fringe, and the sarcophagid and ta-
chinid taxa are scored for state 1 where the spiracular
fringe is divided into a narrow, crescent-shaped ante-
rior fringe (or lappet) and a broad, operculum-like
posterior lappet that is often distinctly narrowed at
base (Crosskey 1977: fig. 45; Rognes 1986: fig. 3).
Rarely, the two lappets are more equal to each other,
as in the genera Macronychia (Miltogrammatinae) and
Chrysotachina Brauer & Bergenstamm (Tachininae)
(Wood 1987c: fig. 169), a configuration which I con-
sider as apomorphic for these genera. Anthomyiidae,
Muscidae, Mesembrinellinae, Gasterophilus + Gyros-
tigma in the Gasterophilinae, and Hypodermatinae
have been scored as inapplicable for this character as
they possess a continuous fringe.
In many Calliphoridae, the anterior lappet is en-
larged. Rarely it is very much larger than the opercu-
lum, as in the genus Phumosia (Calliphoridae) depict-
ed by Crosskey (1977: fig. 46), but note that this
figure is of the right metathoracic spiracle, contrary to
all other spiracles figured. I have scored all non-helic-
oboscine calliphorid subfamilies for an enlarged ante-
rior lappet (2) although exceptions occur, e.g. some
Melanomya in the Melanomyinae. Species with a
small metathoracic spiracle, e.g. all Rhinophoridae,
often have a reduced and less operculum-like posteri-
or lappet, and this condition is here considered a dis-
tinct state (0).
Downes (1986), Pape (1986a) and Rognes (1986)
all mention more or less explicitly that a large meta-
thoracic spiracle is characteristic for (part of) the
Calliphoridae, and Downes (1986: 19) stated that he
considered the enlarged metathoracic spiracle to have
developed several times in that family: “The
Calliphoridae have repeatedly developed very large
posterior thoracic spiracles that intrude forward into
the hind margin of the meron.’ Apparently, the spi-
racular fringe has enlarged as well, and the large ma-
jority of species in all non-helicoboscine calliphorid
subfamilies show this state with unambiguous clarity.
Rognes (1991) provides many line-drawings of calli-
phorid metathoracic spiracles. The Helicoboscinae,
on the other hand, is a perfect representative of the
‘narrow/large’ condition (1), see Rognes (1986: fig.
51
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
3). The subfamily Ameniinae was for long considered
in the Tachinidae, but Crosskey (1965) convincingly
transferred them to the Calliphoridae, although not-
ing that ‘the Ameniinae [may be] as closely related to
the Sarcophagidae as to the Calliphoridae’ and that ‘it
may be better to treat [them] as a full family’ (p. 43).
The metathoracic spiracular fringe of most species is
perfectly ‘large/large’ (2) and I have all reasons to con-
sider this as homologous to the condition found in
other non-helicoboscine calliphorid subfamilies. The
few calliphorid species with a rhinophorid-like meta-
thoracic spiracle fall nicely within their respective taxa
through their close affinity with more typical calli-
phorids, like the close relationship between the genus
Melinda and the Melanomya genus-group (Rognes
1986, 1991).
All species of Mesembrinellinae possess a metatho-
racic spiracle with one continuous, large fringe
(Guimaraes 1977: fig. 8). This non-opercular condi-
tion brings some problems of homology, i.e. whether
all of the fringe or only the anterior part is homolo-
gous to the anterior lappet of other Calliphoridae.
Accordingly, this character has been scored as inappli-
cable for the Mesembrinellinae.
The shape of the rhinophorid metathoracic spirac-
ular fringe needs special mention as this could be con-
sidered intermediate between the operculate fringe
found in Calliphoridae, Oestridae, Tachinidae and
Sarcophagidae on the one side and the type with a
continuous fringe found in the non-tachinoid calyp-
trates on the other. Note, however, that an operculate
metathoracic spiracle occurs in the rhinophorid genus
Baniassa Kugler, where it may be considered a rever-
sal, i.e. apomorphic (Pape 1986a). Crosskey (1977: 8)
made the following description of the rhinophorid
spiracle: ‘In Rhinophoridae .... the external opening
of the spiracle lacks a definite occluding flap or pair of
flaps and instead is margined by complex short fring-
es that stand out from the spiracular rim (there being
normally an anterior and a posterior fringe that meet
ventrally but that are widely separated dorsally),’ and
he considered ‘the nature of the metathoracic spiracle
[as providing] the most important single character for
rhinophorid recognition’. McAlpine (1989: 1501)
characterised the rhinophorid spiracle as ‘subcircular,
without distinct operculum, and margined with erect
fringe of hairs’ and considered this an ‘important
plesiomorphic character’. I have no 4 priori reason to
do this, and McAlpine was, in fact, contradicted by
his own cladogram (fig. 116.8).
Both outgroups, i.e. Muscidae and Anthomyiidae,
have been scored as unknown/inapplicable for this
character. I have insufficient knowledge of the meta-
thoracic spiracular groundplan for these two families,
and the continuous nature of the fringe would make
any scoring rather tentative.
52
Character 14. — Anepimeron; setosity
0 - bare, 1 - setose.
Anepimeral setae occur in only a few
Anthomyiidae (Huckett 1987) and Muscidae
(Huckett & Vockeroth 1987), but in all members of
the Tachinidae family-group. Also, while bristly ane-
pimeral setae are virtually absent in the Anthomyiidae
and Muscidae, they are widespread and common in
the Tachinidae family-group. Therefore, the present
scoring of the anthomyiid and muscid anepimeral
groundplan as bare (0) and that of all other taxa as se-
tose (1), seems well founded.
Character 15. — Wing vein M; course distal to cross-
vein dm-cu
0 - bent anteriorly at junction with dm-cu,
1 - a straight (or at least non-bent) conti-
nuation of proximal part, 2 - bent anterior-
ly at a more or less right angle, with bend
distinctly removed from dm-cu, 3 - bent
anteriorly at an obtuse angle, with bend
distinctly removed from dm-cu.
Gasterophilus + Gyrostigma (Gasterophilini) show
the straight condition (1), while species of Cobboldia
as well as all Cuterebrinae and Oestrinae show the
sharply bent condition (2). The Hypodermatinae are
unique in having the bend occurring almost exactly at
junction between M and crossvein dm-cu. The ob-
tuse angle or gentle curve of M (state 3) is found in
the Mesembrinellinae (Calliphoridae) and
Rhinophoridae. This state is also found in many
smaller species of Calliphoridae (e.g. some
Melanomya), while some Tachinidae (e.g. Freraea
Robineau-Desvoidy, Cinochira, Cylindromyiella) have
the bend almost absent. These instances are here con-
sidered independent acquisitions derived from the
bent conditions. The total absence of the distal part
in various Tachinidae, and in Bezzimyia,
Bequaertiana, Melanophora asetosa Kugler and Oplisa
aterrima (Strobl) of the Rhinophoridae is obviously a
distinct state. No evidence, however, suggests that
this state may characterise the groundplan of any of
the taxa scored in the present analysis, and I have not
made a special entry. Note that some species of Phyto
Robineau-Desvoidy, e.g. P. armadillonis Kugler, and
Mimodexia spp. show a vein M with a bend that is
very reminiscent of the condition found in most
Calliphoridae, i.e. bent in an almost right angle
(Rohdendorf 1935: fig. 82; Kugler 1978: figs. 11,
26). These species, however, do not seem to be close
to the basal splits within the family and their venation
is accordingly more parsimoniously regarded as ho-
moplasic.
No Anthomyiidae possess a bent vein M, but in the
Muscidae many Muscinae have this vein distinctly
bent. However, as no other Muscidae to my know-
ledge share this condition, it is not considered a
groundplan state of the family.
Character 16. — Wing vein Sc; course
0 - with a convexity at level of humeral
break, 1 - straight at level of humeral break.
As noted by Herting (1955), vein Sc of all
Oestridae takes a parallel course relative to the costa
until it bends towards it, ultimately reaching the wing
margin (Wood 1987a: figs. 5-8). In the Muscidae and
the Anthomyiidae and the non-oestrid members of
the Tachinidae family-group, this vein has a distinct-
ly convex part (relative to the anterior wing margin)
at level of, or just distal to the humeral break
(McAlpine 1981: fig. 69).
Character 17. — Wing vein R,; configuration at level
of subcostal break
0 - with a knob, 1 - without a knob.
In most Calyptratae, and certainly in the non-oes-
trid taxa included in the present analysis, vein R, has
a small knob on the anterior surface at the level where
subcosta bends towards costa. This has been taken as
vestiges of a crossvein (sc-r) connecting subcostal and
radial veins (McAlpine 1981). All botflies examined
by me lack this knob of R,.
Character 18. — Posterodorsal surface of stem vein
(remigium); setosity
0 - bare, 1 - setulose.
To my knowledge, all members of Chrysomyinae,
Rhiniinae and Toxotarsinae possess the setose condi-
tion (1). Only two other occurrences of a setose remi-
gium are known within the Tachinidae family-group:
in two species of Mesembrinellinae, Mesembrinella
bellardiana Aldrich and M. peregrina Aldrich, and in
Pollenia atramentaria (Meigen) of the Polleniinae. As
P. atramentaria does not seem to be the sister group
of all remaining Polleniinae, I have scored the
groundplan of this subfamily for a bare remigium (0).
The groundplan of the Mesembrinellinae, however,
has been scored as unknown for this character as the
genus Mesembrinella Giglio-Tos seems to be defined
exclusively on the possession of three (instead of two)
humeral bristles (Guimaräes 1977), which probably is
plesiomorphic at this level. Thus, I cannot exclude
that M. bellardiana and M. peregrina are included in
the basal dichotomy of this subfamily.
Character 19. — First anal vein; length relative to
wing margin
0 - extending to or almost to wing margin,
1 - reduced and ending in wing membrane
distinctly before wing margin.
I have not seen any Muscidae, except for members
of the Eginiini, with an extended A,, and Huckett &
Pape: Phylogeny of the Tachinidae family-group
Vockeroth (1987: 1116) stated that for the Nearctic
Muscidae this vein is ‘always incomplete’. As already
mentioned, I have accepted the inclusion of Eginiini
within the subfamily Phaoniinae of the Muscidae,
and the groundplan of the family has been scored as
reduced (1) for the present character. In the
Anthomyiidae very few genera, Acridomyia
Stackelberg and Coenosopsia Malloch, have a reduced
first anal vein, but as there is no reason to consider
these genera close to each other and neither as sister
group of the remaining Anthomyiidae (V. Michelsen,
pers. comm.), the reduction is probably derived inde-
pendently. The family has accordingly been scored
for an extended first anal vein (0). In the Tachinidae
family-group all members of Bengaliinae and
Gasterophilini possess an extended first anal vein. In
the Cuterebrinae Neocuterebra possess an extended
first anal vein, but as all other species have a distinct-
ly shortened vein the groundplan of this subfamily
has been scored as unknown. A few genera in the
Tachininae (e.g. Siphona Meigen) and
Rhinophoridae (Malayia) show the extended condi-
tion. These instances are all considered as derived.
Character 20. — Base of vein R,.; setosity
0 - bare, 1 - setulose.
The large majority of species within the Tachinidae
family-group possess some setulae on the upper sur-
face of the base of vein R,.., either as a few setulae at
the very base (actually the common base of R,., and
R,,,) or as a distinct row along the vein. Usually, set-
ulae are present on the lower surface as well. In the
Cuterebrinae, the base of R,.. is setose in Dermatobia
hominisand the Cuterebra examined by me, while it is
bare in Neocuterebra and Ruttenia. Gyrostigma +
Gasterophilus of the Gasterophilini show the bare
condition, Cobboldia the setose. All Oestrinae seen
possess the setose condition, while both conditions
occur in the Hypodermatinae. In the Polleniinae, all
species of Morinia Robineau-Desvoidy and
Melanodexia Williston have this vein bare while it is
setulose in Pollenia. The Polleniinae has therefore
been scored as unknown for this character, although
the absence most probably is apomorphic for Morinia
+ Melanodexia, as already noted by Rognes (1991:
209). The only other instance of a bare base of vein
R,,, within the Tachinidae family-group known to
me is in the Phyto carinata species-group of the
Rhinophoridae (Pape 1987b), but several others may
occur. À setulose base of vein R,,, occurs sporadically
in the Muscidae, e.g. species of Graphomya
Robineau-Desvoidy, Musca Linnaeus, Neomyia, and
Stomoxys Geoffroy (Huckett & Vockeroth 1987: figs.
21-30, Zumpt 1973: fig. 94), but not in
Achanthiptera rohrelliformis (Robineau-Desvoidy),
the sole member of subfamily Achanthipterinae.
53
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Achanthiptera Rondani has abdominal spiracle 6 re-
tained in the female terminalia and has for this reason
been considered as one branch of the basal dichotomy
of the family (Carvalho 1989). Huckett & Vockeroth
(1987: 1117), however, found the female abdominal
spiracle 6 retained ‘in a few exotic species of several
subfamilies’, which weakens the argument that
Achanthiptera should be considered a basal lineage.
Awaiting more knowledge of muscid phylogeny, I
have scored the groundplan as unknown for this char-
acter. I have not seen any Anthomyiidae showing the
setose condition.
Character 21. — Lower calypter; shape
0 - narrow, 1 - oval, 2 - broad, 3 - tongue-
shaped.
With the convincing exclusion of Melanomya (here
including Angioneura Brauer & Bergenstamm) from
the Rhinophoridae presented by Downes (1986),
Rognes (1986, 1991) and Pape (1986a), all
Rhinophoridae possess a lower calypter, which is
bigger than the upper calypter, diverging from the
scutellum, and either slightly roundish or parallel-sid-
ed for part of its length (3). Species of the Rhiniinae
possess a very rhinophorid-like lower calypter and
have been scored similarly (3), while both the broad
and tongue-shaped conditions occur in the
Chrysomyinae (broad in Chrysomya Robineau-
Desvoidy, tongue-shaped in all others), and the
groundplan of this subfamily has been scored as un-
known.
The Anthomyiidae are scored for the widespread
narrow state (0), where the lower calypter does not
exceed the upper calypter in size. However, examples
occur where the lower calypter is subequal to or even
exceed the upper calypter in length (Huckett 1987),
and the character needs reconsideration. The
Muscidae are difficult to score as numerous states oc-
cur, ranging from the rather short type found in
Achanthiptera rohrelliformis with a posterior margin
running almost perpendicular to the median plane, to
lower calypteres fully as large and broad as those of
the Tachinidae family-group. The oval type of the
Muscidae (Huckett & Vockeroth 1987: fig. 15),
which is smaller and slightly more tapering than the
condition found in the Rhinophoridae, seems to be
most widespread (and incidentally also the most com-
mon) and is here postulated as groundplan. Further
knowledge of muscid infrafamilial phylogeny is need-
ed.
The groundplan of all other terminal taxa seems to
be characterised by the broad type with the median
margin following scutellum for a considerable dis-
tance before turning outwards almost perpendicular
to the median plane. The several non-rhinophorid
and non-rhiniine taxa of the Tachinidae family-group
54
with diverging lower calypteres, e.g. Cinochira and
Cylindromyiella (Tachinidae: Phasiinae), some species
of Johnsonia Coquillett and Neophyto Townsend (Sar-
cophagidae: Sarcophaginae), Morinia (Calliphoridae:
Polleniinae), and . Melanomya (Calliphoridae:
Melanomyinae) are easily classified as derived from
the broad condition by simple outgroup comparison
within their respective subfamilies. Morinia, for ex-
ample, may be the sister group of Melanodexia (cor-
roborated by black body colour and absence of setae
on wing vein R,..), and the tongue-shaped condition
in Morinia is therefore most parsimoniously consid-
ered apomorphic at this level as Melanodexia and all
other Polleniinae have broad lower calypteres.
Likewise, if it is accepted that the Miltogrammatinae
are the sister group of a monophyletic Para-
macronychiinae + Sarcophaginae (Kurahashi 1975;
Pape 1986b, 1987a; present analysis), the presence of
broad lower calypteres in all species of both non-sar-
cophagine subfamilies is convincing evidence that the
possession of narrow lower calypteres in Johnsonia
and Neophyto is apomorhic. Downes (1986), howev-
er, takes the opposite standpoint and argues from a
functional viewpoint. Broad lower calypteres are seen
as independent adaptations to increase flight speed
and agility, and the ‘primitive-looking groups are
considered as having been ‘shunted off from the
mainstream of calyptrate evolution’ (p. 17). Species
of the Tachinidae family-group are generally very ag-
ile flyers compared to most Muscidae and
Anthomyiidae, and this may indeed be correlated
with the size of the lower calypteres. Flight patterns,
however, are the functional manifestations of mor-
phological and physiological character states and are
thus obviously subject to reversals and homoplasy. I
see no reason or argument for postulating a main-
stream calyptrate evolution. It is more parsimonious,
i.e. requires less ad hoc assumptions, to consider the
shape of calypteres in Neophyto, Johnsonia and the
Rhinophoridae as independent evolutionary events.
Character 22. — Abdominal sternite 2; degree of
exposition
0 - freely exposed but only separated from
margins of tergite 1+2 by a narrow strip of
unfolded membrane, 1 - contiguous with
or slightly overlapped by margins of tergite
1+2, 2 - completely overlapped by margins
of tergite 1+2, 3 - overlapping margins of
tergite 1+2, 4 - freely exposed and widely
separated from tergal margins by more or
less folded membrane.
The degree of exposition of the non-terminal ab-
dominal sternites is a character of diagnostic value,
but I have focused on sternite 2 rather than all of ster-
nites 2-5 as this may be the easiest to score. The
Sarcophagidae are very homogenous, with abdominal
sternite 2 distinctly overlapping margins of corre-
sponding (syn)tergite in all but the aberrant and obvi-
ously apomorphic Asilodexia Townsend (Sar-
cophaginae). The Calliphoridae are more varied,
from a perfectly muscid-like condition in the
Mesembrinellinae to a rhinophorid-like condition in
some Melanomya, the Helicoboscinae, and the
Ameniinae. The configuration in the Rhinophoridae
is here scored like the Ameniinae and Helicoboscinae
(1) as I have no a priori reason to consider them non-
homologous. The Phasiinae have been scored like the
Rhinophoridae, although exceptions occur.
Character 23. - Alpha setae; presence/absence
0 - present, 1 - absent.
Alpha setae (sensilla trichodia) are found wide-
spread in most Rhinophoridae and scattered in the
Tachinidae (Lehrer 1973, Tschorsnig 1985b). They
are absent from the Ameniinae (Rognes 1986), but
are found in at least some species of all other sub-
groups of the Calliphoridae here recognized. No spe-
cies of Sarcophagidae are yet known to possess alpha
setae. The few botflies examined by me for this char-
acter (Cuterebra atrox Clark, Gasterophilus intestinalis
(De Geer), Hypoderma tarandi (Linnaeus), Oestrus
ovis Linnaeus) had no alpha setae, but what appears to
be typical alpha setae is shown for Dermatobia homi-
nis (Cuterebrinae) and Cobboldia chrysidiformis
Rodhain & Bequaert by Patton (1935b, 1937).
Character 24. — Female abdominal sternite 8;
median division
0 - entire, 1 - divided.
Depending on the degree of sclerotization of the
median part of the female abdominal sternite 8, this
may appear complete (0) or divided (1). The charac-
ter was extensively discussed by Herting (1957) and
needs only few additional comments. I have scored all
taxa of the Tachinidae family-group for the entire (or
undivided) condition although a few exceptions oc-
cur. In the Hypodermatinae, Hypoderma Latreille
provides examples of a completely divided female
sternite 8 (Wood 1987a: fig. 14). This may, however,
be shown to be a derived condition by reference to
the undivided condition found in the genera
Oestromyia and Portschinskia Semenov, which are
considered as basal lineages of the subfamily (Grunin
1965). Calliphora stelviana (Brauer & Bergenstamm)
of the Calliphorinae is another example (Rognes
1991: fig. 182), but here considered derived within
its subfamily.
Character 25. — Spermathecal ducts; configuration
relative to uterus
0 - two ducts fused just before entering the
Auteur: Verkorte titel van het artikel
uterus, 1 - all ducts enter the uterus sepa-
rately.
Within the Schizophora and possibly within all of
the Cyclorrhapha, it seems to be a groundplan condi-
tion that two of the three spermathecal ducts join
each other shortly before ending in the uterus, thus
leaving the latter with only two openings leading to
the spermathecal ducts (Griffiths 1982, Wiegmann
1989). Note that McAlpine (1989: 1429) considered
the groundplan of the Calyptratae to have all three
spermathecal ducts reaching the uterus separately,
relying on Sturtevant (1925-26). The latter author,
however, never made extensive dissections of calyp-
trate taxa.
Griffiths (1982), referring to unpublished data
from B. A. Holloway, delimits a clade of the
Tachinidae family-group consisting of the
Sarcophagidae, Tachinidae and Oestridae based on
the complete separation of all three spermathecal
ducts at junction with the uterus. My own dissec-
tions, however, have revealed another pattern (table
1). I have found the ‘fused’ condition with only 2
openings in the Muscidae (other non-tachinoid cal-
yptrate families not examined by me), in the
Cuterebrinae and Hypodermatinae, which are the
only botfly subfamilies examined (figs. 4, 5), and in
all calliphorid subfamilies except Helicoboscinae,
Bengaliinae and Phumosiinae (table 1). All
Rhinophoridae and Tachinidae examined possess
three separate openings. It should be noted that fig-
ures of female reproductive organs drawn for other
purposes may be unreliable with respect to this char-
acter, e.g. that of Mesembrinella peregrina illustrated
by Guimaraes (1977: figs. 39, 40). It is shown to pos-
sess the ‘all-separate’ condition. In the Sarcophaginae
only Sarcodexia lambens (Wiedemann) is known to
have two openings, but this is apparently an autapo-
morphic condition with the common spermathecal
duct very long (Lopes 1941: figs. 8, 10). Among the
species of Miltogrammatinae and Paramacro-
nychiinae dissected by me, none possess the ‘fused’
condition with two spermathecal ducts joining close
to the uterus (table 1). All possess three separate
openings, but the openings may be so close to each
other that they superficially appear as one. In
Sarcotachina subcylindrica Portschinsky and
Sarcophila latifrons (Fallen) (figs. 6, 7), both of the
Paramacronychiinae, and in Metopia argyrocephala
(Meigen) of the Miltogrammatinae, the three sper-
mathecal ducts open into a more or less tube-like or
stalk-like pouch of the uterus. More dissections of
species close to the basal splits within the
Miltogrammatinae and the Paramacronychiinae are
needed to settle the groundplan condition for these
two subfamilies — and thereby probably for all of the
Sarcophagidae. This is important as it could be
DD
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 1. Species dissected for examination of configuration
of spermathecal ducts relative to uterus (character 25). 1:1:1
indicates that all spermathecal ducts enter the uterus or a
common uterine pouch separately. 1:2 indicates that two
spermathecal ducts are fused just before entering the uterus
or a common uterine pouch. Subfamilial assignment indi-
cated where possible.
MUSCIDAE
Achanthipterinae
Achanthiptera rohrelliformis 1:2
RHINOPHORIDAE
Melanophora roralıs lei
Phyto melanocephala 1:1:1
Rhinomorinia sarcophagina 1:1:1
SARCOPHAGIDAE
Miltogrammatinae
Metopia argyrocephala LATE
Xiphidiella anorubra neilail
Paramacronychiinae
Brachicoma devia igileil
Nyctia halterata enen
Sarcophila sp. 1:1:1
Sarcotachina aegyptiaca 1:1:1
Sarcophaginae
Blaesoxipha batilligera leilaıl
Ravinia pernix TETE
Tricharaea sp. leılanı
TACHINIDAE
Exorista genus-group
Exorista larvarum hels
Winthemia quadripustulata 1:1:1
CALLIPHORIDAE
Helicoboscinae
Eurychaeta palpalis eile
Bengaliinae
Bengalia lorosa lateralis lenten
Tricyclea sp. 1:1:1
Polleniinae
Morinia melanoptera 2:1
Ameniinae
Silbomyia sp. 2:1
Mesembrinellinae
Mesembrinella bicolor 2:1
Phumosiinae
Phumosia abdominalis pallida leileil
Calliphorinae
Calliphora vicina 2:1
Luciliinae
Lucilia sericata Deil
Phormiinae
Protophormia terraenovae Dil
Toxotarsinae
Neta chilensis 2:1
OESTRIDAE
Cuterebrinae
Dermatobia hominis
Cuterebra atrox
Hypodermatinae
Hypoderma tarandi DE]
56
argued that the stalk-like pouch represents the fused
bases of all three spermathecal ducts, which then
should be given a separate scoring.
The configuration of the spermathecal ducts is
interesting as the recent exclusion of some presumed
Calliphoridae from the Rhinophoridae may be fur-
ther corroborated. An examination of Morinia mela-
noptera (Fallen), for example, reveals the fused condi-
tion, and according to Downes (1986: fig. 26), this
condition is found in Melanomya as well. Both these
genera have been treated as Rhinophoridae by
Herting (1961), Crosskey (1977), and Tschorsnig
(1985a), while they were put in the Calliphoridae by
Downes (1965, 1986), Rognes (1986, 1991) and
Pape (1986a, 1988). The separate spermathecal ducts
of the Sarcophagidae, Rhinophoridae and Tachinidae
may have triggered the evolution of a tripartite acro-
phallus several times. A tripartite acrophallus is thus a
groundplan feature of the subfamily Sarcophaginae
(Pape, unpubl.), and tripartition was recently discov-
ered in a single species of Miltogrammatinae (Pape
1989). Tripartition has probably evolved once in the
Rhinophoridae (Tschorsnig 1985a, Pape 1986a) and
several times in the Tachinidae (Andersen 1988). On
the other hand, tripartition apparently never evolved
in the Calliphoridae, Oestridae nor any of the non-
tachinoid calyptrate families. However, many genera
of the Bengaliinae show a complicated acrophallus
with flaps and membranes that may guide sperm
and/or accessory gland material into the openings of
the spermathecal ducts and thus effectively be a tri-
partition. The lateral acrophallic ducts guiding the
accessory gland material from the phallotreme to the
female lateral sacs in species of Lucilia Robineau-
Desvoidy are obviously not homologous to any of the
examples of acrophallic tripartition discussed above as
they are invaginations of the external wall of the phal-
lic tube (Merritt 1989, Rognes 1991). It should be
kept in mind that separate spermathecal openings
most probably is a groundplan feature of all of Dip-
tera (McAlpine 1981), while this groundplan prob-
ably had but a single male gonopore (Wood 1990).
Character 26. — Spermathecae; size
0 - equal sized, 1 - one reduced.
In the Oestridae, all Hypodermatinae and
Oestrinae have one of the three spermathecae reduced
(see figures in Grunin 1965, 1966). The
Gasterophilini have also been scored as ‘one reduced’
(1) as the total absence of one spermatheca in
Gasterophilus obviously has to be considered a reduc-
tion. I have no information on size or number of sper-
mathecae in the genus Gyrostigma. In the genus
Cobboldia, spermathecae of C. chrysidiformis and C.
elephantis are figured in Patton (1937), showing three
oval or slightly elongated spermathecae of which one
is clearly reduced in the former species, slightly re-
duced in the latter. All Cuterebrinae may have spher-
ical, equal-sized spermathecae, although I have exam-
ined only Dermatobia hominis (fig. 4) and Cuterebra
atrox, but see also the figure of C. fontinella Clark in
Patton (1935b, fig. 6). The Mesembrinellinae and
Phumosiinae have elongated spermathecae, which
may vary somewhat in size, but both taxa have been
scored for equal sized spermathecae (0).
I have scored the Tachininae as having spermathe-
cae of equal size, thereby considering the few instan-
ces of loss of one spermatheca, e.g. in Siphona
(Andersen 1983), as secondary.
Character 27. — Male sternite 5; configuration of
posterior margin
0 - deeply emarginated, 1 - simple or with
a shallow emargination.
Although the posterior outline of the male sternite
5 shows much variation, the shallow posterior emar-
gination in all Oestridae and in the subfamily
Paramacronychiinae of the Sarcophagidae is very dis-
tinct. In the Miltogrammatinae Chrysogramma and
Eumacronychia show a typical deep emargination,
while the remaining members of the subfamily have
no or a shallow emargination. The groundplan for
the Miltogrammatinae has been scored as unknown.
Character 28. — Male tergite 6; connection to
syntergosternite 7+8
0 - free, 1 - fused.
The fused condition is found in all of
Paramacronychiinae and apparently in all Oestridae.
The fused condition occurs scattered elsewhere in the
Tachinidae family-group but cannot be ascribed to
the groundplan of any other terminal taxon consid-
ered here.
Character 29. — Male bacilliform sclerites;
configuration
0 - well developed and parallel, 1 - reduced
and more or less perpendicular to the me-
dian plane.
All Sarcophagidae possess reduced bacilliform
sclerites (processus longi, divided sternite 10).
Usually only very short sclerites remain, and if dis-
tinct these are always more or less perpendicular to
the median plane. The groundplan of all non-sarco-
phagid terminal taxa scored here is characterized by
distinctly developed bacilliform sclerites, which are
parallel and directed posteriorly.
Character 30. — Gonostylus; presence/absence of
basal apodeme
0 - apodeme not present, 1 - apodeme
present.
Pape: Phylogeny of the Tachinidae family-group
It has been known for decades, that some or most
tachinoid calyptrates possess a small sclerite at the
base of the gonostylus (paramere of McAlpine
(1981)). This sclerite is figured but unnamed in
Rohdendorf (1937), named ‘Gelenkfortsatz’ by Salzer
(1968), ‘additional forceps’ by Lopes (1975), and ‘ba-
sal parameral apodeme’ by Pape (1990). The impor-
tance, in a phylogenetic context, of the presence/ab-
sence of this sclerite, was first recognized by
Tschorsnig (1985a, b). The latter author found this
structure in all Tachinidae, Sarcophagidae,
Rhinophoridae, and Calliphoridae examined, but not
in the few members of the Gasterophilinae, Oestrinae
and non-tachinoid Calyptratae examined.
Accordingly, he favoured the hypothesis of Hennig
(1976) that the Oestridae are the sister group of all
other members of the Tachinidae family-group
(Tschorsnig 1985a: 10). Rognes (1991) found no go-
nostylar apodeme in Cephenemyia trompe (Modeer)
(Oestrinae) and Hypoderma tarandi
(Hypodermatinae). I have examined males of
Cobboldia chrysidiformis, several Gasterophilus,
Hypoderma bovis (Linnaeus) and Oestrus ovis
Linnaeus, which all had no apodeme. However,
males of Cuterebra neomexicana Sabrosky, Oestromyia
leporina (Pallas) and Cephenemyia trompe examined
by me show an unambiguous, although small and eas-
ily overlooked apodeme at the base of the gonostylus
(fig. 1). In a single specimen of Ruttenia loxodontis ex-
amined, it appears as if the gonostylar apodeme is
partly fused to the gonostylus. Therefore, the ground-
plan of the Cuterebrinae is scored as present (1), that
of Gasterophilini and Cobboldiini as absent (0), and
that of Hypodermatinae and Oestrinae as unknown
(both states occur). A more careful search for this eas-
ily overlooked apodeme in the Oestridae is needed.
The sclerite may be homologous to the gonostylar
apodeme found in other Diptera, part of the gonosty-
lus proper, or a unique sclerite arisen de novo in the
Tachinidae family-group. Contrary to the internal
gonostylar apodeme found in other Diptera it forms
part of the external cuticle, and the first possibility
seems uncorroborated or even falsified. As nothing
seems to indicate that the apodeme should have split
from the base of the gonostylus, I am in favour ofa de
novo origin, but note that the use of the presence/ab-
sence of this apodeme in a phylogenetic context does
not require an explicit interpretation of its potential
homology with structures found in groups not in-
cluded in the analysis.
Character 31. — Phallus; ventral surface of distal
section
0 - smooth, 1 - with sclerotized denticles.
The denticles (Körnchen oder Dörnchen’ of
Tschorsnig (1985b: 61)) are found throughout the
57
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Tachinidae family-group, and they usually cover a
large part of the ventral surface. They are absent from
the acrophallus and part of the ventral surface in the
calliphorid subfamilies Melanomyinae and
Polleniinae (Rognes 1991). In the Sarcophagidae
they are absent in several members of the sarcophagid
subfamily Paramacronychiinae, e.g. Goniophyto
Townsend, Wohlfahrtia Brauer & Bergenstamm and
Sarcophila Rondani. They are restricted to the lateral
styli in many Sarcophaginae. In the Tachinidae,
Tschorsnig (1985b) recorded an absence of phallic
denticles in part of the subfamily Phasiinae. All taxa
of both the Sarcophagidae and the Tachinidae, how-
ever, have been scored as denticled (1) because no ev-
idence is present that favour a basal position of the
‘non-denticled’ taxa. In the Gasterophilinae, denticles
are present in Cobboldia (Patton 1937), absent in
Gasterophilus. | have not examined Gyrostigma for this
character. Similar phallic denticles are absent from
the Muscidae (personal observations, few samples),
and in non-tachinoid Calyptratae they are known on-
ly from a subgroup of Delia Robineau-Desvoidy
(Anthomyiidae), where the condition most probably
has developed independently (Griffiths 1982).
Character 32. — Dorsolateral phallic processes; apical
configuration
0 - confluent with distiphallic wall throug-
hout, 1 - separate from wall of distiphallus,
Le. with a free tip.
Rognes (1986, 1991) stressed the phylogenetic im-
portance of the free-tipped condition and considered
this as synapomorphic for Calliphoridae and
Rhinophoridae.
Subfamilies of the Sarcophagidae and Tachinidae
have been scored as inapplicable for this character as
their dorsolateral processes are fused along the dorso-
median line. Some Sarcophaginae possess what ap-
pears to be free dorsolateral processes, most notably
Sarcodexia lambens (fig. 2) but also some species of
Blaesoxipha Loew subgenus Acridiophaga Townsend.
These processes, however, although derived from the
dorsal plate, are probably not homologous to the pro-
cesses seen in the Calliphoridae. A very large clade in
Pape: Phylogeny of the Tachinidae family-group
the Sarcophaginae, containing both Sarcodexia
Townsend and Blaesoxipha, is defined by the presence
of a transverse, desclerotized strip distally on the dor-
sal side of the phallus, separating what is generally
called a juxta from the more proximal part of the dor-
sal plate. The juxta may well contain elements of the
fused dorsolateral processes, which means that any
free sclerotized prongs in juxtate Sarcophaginae can-
not be considered homologous to the free tips of the
dorsolateral processes in Calliphoridae and
Rhinophoridae.
All calliphorid subfamilies have been scored for free
tips except the Bengaliinae and the Rhiniinae. For the
Bengaliinae, my knowledge of phallic structures is
rather limited, and the groundplan of dorsolateral
processes has been scored as unknown/inapplicable.
As dorsolateral processes in the Rhiniinae seem to
have fused in the dorsomedian line, the character has
been scored as unknown/inapplicable as in the
Sarcophagidae and Tachinidae (see discussion under
character 34 below).
All Hypodermatinae possess sclerotized processes
laterally or dorsolaterally on the phallus and with tips
just free of the wall (see figures in Grunin 1965). No
other member of the Oestridae possesses similar pro-
cesses. These processes are strikingly similar to the
dorsolateral phallic processes of the Calliphoridae,
and apical microserration may suggest a similar func-
tion. They differ, however, by apparently originating
from the lateral plates rather than from the dorsal
plate. Rudiments of the dorsolateral processes may be
seen at the base of the dorsal plate (Grunin 1965: fig.
123). This puts serious doubt on the homology to the
dorsolateral processes seen elsewhere in the
Tachinidae family-group and I have scored the
Hypodermatinae for the confluent condition.
Both Rognes (1986) and Pape (1986a) found no
examples of free dorsolateral processes in the
Tachinidae, and checking the numerous illustrations,
eg. in Wood (1972), Shima (1986, 1988),
Tschorsnig (1985b) and Cantrell (1988), does not re-
veal any. It should be noted, however, that Malayia
and some undescribed Bezzimyia (Pape unpubl.) pos-
sess free tips of dorsolateral processes, but I regard
Figs. 1-3. Details of male terminalia. — 1. Cuterebra neomexicana Sabrosky (Oestridae: Cuterebrinae), phallus, gonocoxite and
gonostylus, posterior view; 2. Sarcodexia lambens (Wiedemann) (Sarcophagidae: Sarcophaginae), phallus, dorsal view; 3.
Bezzimyia sp. (?Rhinophoridae), phallus; a = distiphallus, dorsal view, b = phallus, lateral view. Abbreviations: DP = dorsolat-
eral process, G = gonocoxite, P = gonostylus, PA = gonostylar apodeme.
Figs. 4-7. Female internal genitalia, ovaries omitted. — 4. Dermatobia hominis (Linnaeus, Jr.) (Oestridae: Cuterebrinae), only
one accessory gland shown; 5. Hypoderma tarandi (Linnaeus) (Oestridae: Hypodermatinae), with enlarged subset showing
junction of spermathecal bases with uterus; 6. Sarcophila sp. (Sarcophagidae: Paramacronychiinae), with only one spermath-
eca shown and enlarged subset showing junction of ducts from spermathecae and accessory glands with uterus; 7. Nyctia hal-
terata (Scopoli) (Sarcophagidae: Paramacronychiinae) with only one spermatheca shown. Abbreviations: AG = accessory
gland, IP = remnants of incubatory pouch, O = uterus, S = spermatheca. Scale bar = 0.1 mm.
59
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
these to be Rhinophoridae as already discussed.
The Rhinophoridae have been scored as unknown
for the present character as both conditions are of
widespread occurrence. Processes free from distiphal-
lic wall are found in Mimodexia, Malayia, Phyto and
some Bezzimyia; processes confluent with distiphallic
wall are present in remaining genera (see figures in
Tschorsnig (1985a) and Pape (1986a)). The present
knowledge of generic phylogeny is largely that of
Tschorsnig (1985a) and Pape (1986a), with addition-
al comments in Pape (1987d), but both authors did
not consider Bezzimyia and Malayia. A proper evalu-
ation of the most probable groundplan state is there-
fore not possible at present.
As I have never encountered distinctly free tips of
dorsolateral processes within Muscidae and
Anthomyiidae, these have been scored as confluent
(0). This is in agreement with Hennig (1976: 690),
who pointed out the free tips of the calliphorid dorso-
lateral processes and mentioned that dorsolateral pro-
cesses in the Anthomyiidae were confluent with the
distiphallic wall (‘Bei den Anthomyiidae scheinen
[die dorsolaterale Fortsätze] nur die Seitenteile einer
geschlossenen in der Mitte membranösen oder eben-
falls sklerotisierten Dorsalwand zu bilden’).
Character 33. — Acrophallus; position
0 - strictly terminal, 1 - on ventral surface
of phallus.
All terminal taxa except the sarcophagid subfami-
lies Paramacronychiinae and Sarcophaginae possess a
groundplan phallus with the spermexit almost per-
fectly terminal and situated on the longitudinal axis
of the phallus, although this may be curved in the me-
dian plane. A very characteristic modification of both
the paramacronychiine and sarcophagine phallus is
the position of a very well-defined acrophallus ven-
trally on the distal part of the phallus.
Character 34. - Dorsolateral phallic processes;
dorsomedian configuration
0 - separate, 1 - fused.
Phallic dorsolateral processes are here considered
homologous throughout the Tachinidae family-
group with the exception of the juxtate
Sarcophaginae discussed under character 32 above.
Tschorsnig (1985b: 119) suggested that the dorsolat-
eral processes of the Calliphoridae and Rhino-
phoridae were homologous to the dorsal extension
found in the Tachinidae, which I fully accept. To my
knowledge, separate dorsolateral processes are un-
known in the Tachinidae (if the exclusion of Malayia
and Bezzimyia is accepted), and the tachinid taxa have
been scored as fused (1), although some species may
lack dorsolateral processes altogether. In the
Sarcophagidae, all species possess fused dorsolateral
60
processes (see for example Pape 1986c: figs. 1-4, Pape
1987a: figs. 29-33). Although some members of the
subfamily Miltogrammatinae may have the dorsal
plate partly desclerotized medially, they never present
fully separate dorsolateral processes. The Hypo-
dermatinae have been scored as unknown for this
character as I consider the dorsolateral plates to be re-
duced.
I have not been able to fully evaluate the phallic
groundplan of the Anthomyiidae and Muscidae,
which have been scored as unknown.
Character 35. — Uterus; presence of ventral bilobed
pouch
0 - not present, 1 - present.
A bilobed uterine pouch for embryonating eggs is
found in the Sarcophagidae and seems here to be
present without exceptions. In the botflies, all
Oestrinae have a bilobed sac or pouch extending from
the uterus, and although somewhat different in shape
and tracheation, the Oestrinae have been scored like
the Sarcophagidae (1). In the Tachininae (as here de-
fined) eggs are retained (incubated or embryonated)
in the uterus, which often becomes greatly distended,
but no pouch is formed. Eurychaeta palpalis
(Robineau-Desvoidy) of the Helicoboscinae has a
pouch-like extension of the uterus, but this is never
bilobed (Rognes 1986). Large incubatory pouches are
known from Bellardia Robineau-Desvoidy and
Onesia Robineau-Desvoidy (Calliphorinae), but these
are lateral (Rognes 1991).
Character 36. — Oviposition behaviour; deposition
relative to food source
0 - eggs (or larvae) deposited directly on
the larval food source, 1 - eggs (or larvae)
deposited away from the food source.
As correctly stated by Wood (1987b: 1189), the
deposition of eggs away from the host in the
Rhinophoridae is ‘unusual among calyptrates’.
McAlpine's (1989: 1501) claim that this is a plesio-
morphic groundplan character state of this family is
simply unsupported and may have come from
Crosskey's (1977: 6) statement that the Rhino-
phoridae possess an ‘oviposition habit of great sim-
plicity and apparent primitiveness'.
Another instance of oviposition away from the host
is found in the genus Pollenia. As the biology of oth-
er genera of the Polleniinae is largely unknown, this
subfamily has been scored as 1. I know of no other
calliphorid subfamily (as here defined), where ovipo-
sition away from the food source is a groundplan
character state. Within the Sarcophagidae,
Sarcophaga Meigen (s.str.) and at least some species of
Miltogrammatinae (Phrosinella Robineau-Desvoidy
and Eumacronychia) possess state 1 (Eberhardt 1955,
Downes 1955, Lopes 1982a). As the groundplan of
all other sarcophagid genera, for which extensive bio-
logical knowledge exists, seems to be characterised by
state 0, the sarcophagid subfamilies have been scored
as such. The gluing of eggs to the host-wasp carrying
prey to the nest within some Miltogrammatinae (Day
& Smith 1980) is here considered derived. Within
the Oestridae, all Cuterebra oviposit in places fre-
quented by the host, but never directly on a host spec-
imen. Dermatobia hominis uses a carrier for its eggs,
while the habit of oviposition for Neocuterebra and
Ruttenia is unknown. Almost all non-cuterebrine bot-
flies ovi- or larviposit directly on their host. One ex-
ception is Gasterophilus inermis (Brauer), which de-
posits eggs on grassblades and the like. This is here
considered a derived condition. Within the
Hypodermatinae, species of Portschinskia seem to
produce eggs without any attachment apparatus
(Grunin 1965), but no observation has been made of
the oviposition, nor have any naturally deposited eggs
been found. Grunin (1965) suggested an oviposition
behaviour similar to that of Cuterebra, and I have
scored the groundplan of the Hypodermatinae as un-
known for this character.
Character 37. — Egg, ventral surface
0 - not glued to substrate, 1 - glued to
substrate.
Many members of the Calyptratae will deposit eggs
that are sticky and therefore adhere to the surface on-
to which they are positioned. This, however, is easily
distinguished from the condition of all oviparous
members of the Oestridae and Tachinidae, where
eggs are glued firmly to the substrate. The
Sarcophagidae are mainly ovi-larviparous but some
will deposit embryonated eggs ready to hatch, espe-
cially when an ample larval food source induces pro-
lific larviposition. In the subfamily Miltogram-
matinae, some species may glue eggs to the host wasp
(Day & Smith 1980), while others deposit eggs freely
(Maneval 1929), and the groundplan of this subfam-
ily has been scored as unknown with regard to this
character. Nothing is known on egg structure and
deposition in Neocuterebra and Ruttenia.
Character 38. — Egg; hatching mechanism
0 - by peeling off a long flap, 1 - by discar-
ding an anterodorsal cap.
In the Tachinidae, eggs of the non-embryonating
groups Phasiinae and the Exorista genus-group pos-
sess anterior cleavage lines flanking the aeropyles and
delineating a somewhat operculum- or caplike struc-
ture through which the larva emerges (Herting 1960,
Wood 1987c: fig. 110.264). This is very similar to
the condition in plastronate groups, although with
shorter hatching sutures. I have scored both groups as
Pare: Phylogeny of the Tachinidae family-group
0. Many different hatching mechanisms are known
from the Tachininae, and they have been scored as
unknown with regard to the groundplan for this char-
acter. Very little is known about egg structure and
function in the Sarcophagidae (scored as inappli-
cable/unknown), but eggs of at least one species of
Miltogrammatinae hatch through pushing off an an-
terior egg cap or operculum (Day & Smith 1980). In
the genus Sarcophaga (s.l.), no preformed lines or
weaknesses seem to exist, and the larva emerges by
bursting the chorion (Baudet 1980). Eggs of all ovip-
arous Oestridae hatch by discarding an anterodorsal
cap (Wood 1987a: fig. 15, 17).
Within the Calliphoridae, Booponus of the
Bengaliinae is to my knowledge the only known tax-
on with a somewhat oestrid-like hatching mecha-
nism.
Character 39. — Progeny at deposition; stage of
development
0 - unembryonated eggs, 1 - embryonated
eggs or prehatched first instar larvae (ovo-
larviparous), 2 - larvae that for an extended
period have received nourishment within
the uterus (true larviparous).
In the Sarcophagidae, eggs do not necessarily hatch
immediately upon deposition. They are always em-
bryonated and except for Sarcophaga nigriventris
Meigen, which is stated to deposit second instar lar-
vae (Séguy 1965), I know of no cases of sarcophagid
larvae being nourished in utero. In a species of
Oebalia Robineau-Desvoidy, which oviposits directly
on the ‘host’ carrying the prey, the first instar larva
within the egg is fully mature and postpones hatching
simply to await being transported to the host nest by
its carrier (Day & Smith 1980). No embryonation
takes place in the Rhinophoridae, and in the
Oestridae only in the larviparous subfamily
Oestrinae. All Phasiinae are stated by Wood (1987c)
to deposit unembryonated eggs, and the Tachininae
are here defined as including all embryonating
Tachinidae.
Within the Calliphoridae the macro- or unilarvip-
arous habit of the Helicoboscinae, Ameniinae,
Phumosiinae and Mesembrinellinae is well known,
but only the Mesembrinellinae will nourish the larvae
during intra-uterine life (Rognes 1986, Crosskey
1965, Guimaraes 1977, Ferrar 1978).
Character 40. — Mandible of first instar larva;
relative development
0 - present as a distinctly sclerotized plate,
1 - reduced to an ill-defined sclerotization
on side of mouth opening.
In all Tachinidae, what is probably a reduced man-
dible is present as ‘a vaguely defined sclerotized area
61
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
on side of mouth opening, without hook-like
extension’ (Wood 1987c: 1196). None of the non-ta-
chinid terminal taxa applied here has a similar man-
dibular reduction as a groundplan feature. Note that
McAlpine's (1989) statement that calliphorid first in-
star larvae lack paired mandibles is erroneous and was
taken uncritically from Hennig (1973). All
Calliphoridae possess easily recognizable paired man-
dibles in the first instar larva, although sometimes
quite small (Rognes 1991).
Character 41. — Labrum of first instar larva;
connection to remaining cephalopharyngeal skeleton
0 - labrum more or less connected to, but
always distinct from remaining cephalop-
haryngeal skeleton, 1 - labrum firmly fused
to, and contiguous with remaining cephal-
opharyngeal skeleton.
The unique cephalopharyngeal skeleton of the first
instar tachinid larva is discussed and described suffi-
ciently in the literature (e.g. Ferrar 1987). With no
exceptions within the Tachinidae, and no homopla-
sies in the possible groundplan of any of the other ter-
minal taxa, this character needs no further explana-
tion.
Character 42. — Segments of first instar larva;
development of spines
0 - without bands of strong spines, 1 - with
bands of strong spines.
First instar larvae of all Oestridae are well-known
for their bands of usually strong spines (Grunin 1965,
1966, 1969; Wood 1987a). These spines are always
present in the first instar but may be modified or re-
duced in later instars.
Character 43. — Spiracular plates of third instar
larva; development of peritreme and ecdysial scar
0 - complete, 1 - incomplete.
All larvae of Sarcophagidae have the peritreme in-
complete ventro-medially and the ecdysial scar from
the second instar posterior spiracles is absent or indis-
tinct. A similar configuration belongs to the ground-
plan of the calliphorid subfamily Chrysomyinae
(Rognes 1991). In the Cuterebrinae both Dermatobia
and Neocuterebra show an incomplete peritreme
while that of Cuterebra is complete. The cuterebrine
groundplan has been scored as unknown for this
character.
Character 44. — Posterior spiracles of second and
third instar larva; position relative to surface
0 - more or less level with surrounding sur-
face, 1 - set in a recession or cavity.
In the Sarcophagidae, close to all species for which
larval morphology is known have the posterior spira-
62
cles concealed in a distinct recession. The only excep-
tions known to me are species of Amobia Robineau-
Desvoidy sensu stricto (Miltogrammatinae),
Macronychia, and at least some Tricharaea
(Sarcophaginae), but these are here considered homo-
plasies. Somewhat similar conditions in the
Gasterophilinae, where upper and lower lips can be
pulled together, enclosing the spiracles, and in the
Cuterebrinae, where the posterior spiracles can be
pulled into the preceding segment, are considered
non-homologous.
Character 45. — Parastomal bars of third instar larva;
presence/absence
0 - parastomal bars present, 1 - parastomal
bars absent.
Parastomal bars are easily recognized when present
and need no further comments (see figures in Ferrar
(1987)).
Character 46. — Posterior spiracle of second and
third instar larva; vertical position of ecdysial scar
0 - low (or ventrally) on the spiracular
plate, 1 - near the horizontal diameter of
the spiracular plate.
This character has not previously been discussed in
the literature, but illustrations of spiracular plates, e.g.
in Ferrar (1987), show the difference in position of
the ‘low’ ecdysial scar of non-botfly calyptrates com-
pared to the much higher ecdysial scar of the botflies.
Character 47. — Posterior spiracles of second and
third instar larva; configuration of openings
0 - three slits, 1 - a porous plate.
The porous condition is found in the Hypo-
dermatinae and Oestrinae of the Oestridae (Zumpt
1965, Howard 1980, Ferrar 1987). Cuterebra de-
serves mention as the configuration with highly tor-
tuous slits could be taken as resembling that found in
Oestrinae and Hypodermatinae (see, e.g., Baird &
Graham (1973: fig. 7D). However, the spiracular
openings are still conspicuous slits, not porous plates,
and with a clearly discernible tripartition. Typical tri-
partite slits occur in Dermatobia, Ruttenia and
Neocuterebra (Ferrar 1987).
Character 48. — Posterior spiracular plates of second
and third instar larva; configuration
0 - two isolated plates, 1 - plates fused in
the median line.
Second and third instar larvae of all members of
both Cobboldia and Gasterophilus + Gyrostigma have
the spiracular plates fused (Zumpt 1965). Ferrar
(1987: 137) mentioned that in Dermatobia hominis
‘the two plates [of the posterior spiracles] are appar-
ently joined, and have a single median button
between the slits’. Figures in Kremer et al. (1978),
however, show neither peritreme nor button. The
fused character state has not been found in any other
taxon of the Tachinidae family-group.
Character 49. — Puparium; hatching mechanism
0 - anterior end splits into a dorsal and a
ventral piece, 1 - anterior end splits off on-
ly one, dorsal piece.
The groundplan condition of puparial hatching in
the Cyclorrhapha (Ferrar 1987), seems to be a split-
ting off of an anterior cap, and this typically has a lat-
eral suture causing further splitting into a ventral and
a dorsal half. In the botflies, the ventral half is more or
less fused to the remainder of the puparium and the
dorsal half is flattened to a lid-like structure.
UNSCORED CHARACTERS OF POTENTIAL
PHYLOGENETIC VALUE
As mentioned previously, character matrices are
never complete, and some characters have not been
included in the analysis because of uncertainty with
regard to proper scoring. These are dealt with below
in the hope that future studies will allow inclusion in
the matrix.
Character a. — Eyes; holoptic/dichoptic condition
The typical holoptic condition with frontal vitta
completely obliterated between eyes is widespread
and common in the male sex of Calliphoridae,
Muscidae and Anthomyiidae (Huckett 1987: fig. 4,
Huckett & Vockeroth 1987: fig. 5, Shewell 1987a:
fig. 6). The holoptic condition occurs scattered in the
Tachinidae and is present in a few Rhinophoridae
(some Phyto and Stevenia Robineau-Desvoidy), Sar-
cophagidae (Agria Robineau-Desvoidy, Blaesoxiphella
Villeneuve) and Oestridae (some Cuterebra,
Pharyngobolus Brauer). Much variation exists between
and even within genera and groundplan estimates are
difficult to assess at the subfamily and family levels.
Character b. — Intrapostocular setulae;
presence/absence
Presence of intrapostocular setulae have been used
in the definition of the Calliphoridae in spite of sever-
al exceptions: absences within the Calliphoridae as
well as occurrences outside this family (Rognes
1986). I have not made a sufficiently extensive search
for intrapostocular setulae to give a reliable scoring.
They appear to be widespread in the non-rhiniine
and non-ameniine Calliphoridae, including rare oc-
currences in the possibly basal lineage
Helicoboscinae. I have not found any intrapostocular
setulae in the Oestridae, Sarcophagidae and
Rhinophoridae. Rognes (1986:86) found such setulae
Pape: Phylogeny of the Tachinidae family-group
in Dexiosoma caninum (Fabricius) and Tachina fera
Linnaeus of the Tachinidae. An extensive search in
the Muscidae and Anthomyiidae is needed.
Character c. — Antennal arista; setosity
Whether the arista is bare or plumose or some in-
termediate condition is often used for diagnostic pur-
poses at various levels within the Tachinidae family-
group. The character is, however, quite variable, and
I have failed in breaking it up into only a few clearcut
states. The setose condition covers a whole array of
types, from the single row of dorsal and ventral rays in
Musca (Muscidae) to the bottle-rinser-type of
Melanophora roralis (Rhinophoridae). However, the
type with rays arranged dorsally as well as ventrally,
with at least the dorsal row being double in the proxi-
mal part, and no hairs or pubescence on the sides is
very characteristic. This ‘calliphorid type’ arista is
found widespread in all non-helicoboscine
Calliphoridae. Eurychaeta Brauer & Bergenstamm,
which is the sole genus of the Helicoboscinae, has a
unique arista with medium long rays on the whole
surface, although with the dorsal and ventral rays lon-
gest. Members of the Rhiniinae have either a bare or
a pectinate arista, with the latter type being identical
to the typical calliphorid type except for the lack of
ventral rays.
In the Oestridae, Dermatobia and most Cuterebra
of the Cuterebrinae possess a plumose or pectinate
calliphorid type arista. All other botflies have a com-
pletely bare arista.
No members of the Rhinophoridae and
Tachinidae possess a calliphorid type arista as here de-
fined. When a similar arista occurs in the
Sarcophagidae, like in Agria and Sarcophila of the
Paramacronychiinae, the arista often has distinct pu-
bescence on the sides. In the Sarcophaginae, a plu-
mose arista may be a groundplan feature and at least
Ravinia Robineau-Desvoidy and Oxysarcodexia
Townsend have a calliphorid type arista. In the large
Sarcophaga sensu lato, however, the arista has pubes-
cence on the sides. The apparently bare arista of the
Miltogrammatinae and several Tachinidae is covered
with short or minute pubescence.
The Rhinophoridae present several examples of a
plumose arista, but never of the calliphorid type de-
scribed above. McAlpine's (1989: 1501) statement
that a plumose arista in the Rhinophoridae is an im-
portant plesiomorphic groundplan character state is
to some extent contradicted by himself as he simulta-
neously, although tentatively, used this state as autap-
omorphic for the Calliphoridae.
My knowledge of the Muscidae and Anthomyiidae
is too sparse to give a thorough discussion on varia-
tion in aristal setosity.
More accurate descriptions of different configura-
63
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
tions of aristal setosity, preferrably using SEM, and
further knowledge of their taxonomic distribution in
all of the Calyptratae are needed before this character
can effectively be used in interfamilial phylogenetic
analyses.
Character d. — Outer posthumeral thoracic seta;
position relative to the prealar seta
The position of the outer posthumeral seta outside
(i.e. lateral to) a sagittal line through the prealar seta
has often been used in the definition of the
Calliphoridae (Rognes 1986). This lateral position
seems to characterize the groundplan of all subfami-
lies except perhaps the Ameniinae, where both con-
ditions occur (Crosskey 1965), and the Polleniinae,
where the outer posthumeral seta is in line with the
prealar seta in Pollenia, absent in Morinia (Rognes
1991). The groundplan of both the Tachinidae and
Sarcophagidae has the outer posthumeral in a median
position relative to a sagittal line through the prealar
seta. Scoring the Rhinophoridae presents some pro-
blems. The outer posthumeral seta is generally absent,
but when present as a fine seta in some species of
Phyto, especially distinct in Phyto sordidisquama
Villeneuve, it is in the lateral position. A scoring as
absent would be equal to inapplicable and therefore
rather uninformative; a scoring as lateral would igno-
re all the absences. The character is inapplicable for
the Oestridae, as no bristly thoracic setae are differen-
tiated.
I have not been able to give a reliable estimate of
the groundplan for this character in the Antho-
myiidae and Muscidae.
Character e. — Ventral surface of costa between
junction with subcosta and R;; setosity
This character is diagnostically valuable for various
groups in the Calyptratae and clearly deserves more
attention (Rognes 1991: fig. 8, inset). Within both
the Calliphoridae and the Oestridae, this part of costa
is setose in the majority of species, while the bare con-
dition seems to be common and widespread in the
Tachinidae and Sarcophagidae. In the latter family
only a few genera of Sarcophaginae, e.g. Boettcheria
Parker, Emdenimyia Lopes and Notochaeta Aldrich,
possess the setose condition. Both conditions occur in
the Rhinophoridae, Muscidae and Anthomyiidae
with no obvious choice for the most probable
groundplan condition.
Character f. — Anterior malpighian tubules of adults;
length
This character state has not previously explicitly
been used as a tachinid groundplan feature, and it is
admittedly based on sparse evidence. Hori (1962)
depicts the alimentary canal of Sturmia sericariae
64
(Rondani), Prosena siberita (Fabricius) and Tri-
gonospila transvittatum (Pandellé), which all possess
reduced anterior malpighian tubules, which here are
defined as the pair of tubules arising from a common
malpighian duct in the right side of the gut and run-
ning anteriorly along the midgut. Three other tachin-
ids are listed in Hori's (1962) table 1 and are likewise
stated to have shortened anterior tubules. As in all ot-
her Calyptratae figured by Hori, the anterior tubules
of the Tachinidae run forwards along the midgut.
They do not, however, form a complete loop, with
the tubules running all the way back and terminating
at about the level of the rectal papillae. Instead, the
anterior tubules make a slight curve or a very short
loop, before ending well in advance of their starting
point. It should be noted that no rhinophorids were
dissected by Hori (1962), but specimens of Phyto me-
lanocephala (Meigen) dissected by me show the long
configuration of anterior malpighian tubules.
Character g. — Female terminalia; length
Long, retractile telescopic female terminalia are
common and widespread in both the Anthomyiidae
and Muscidae. In the Rhinophoridae, long telescopic
terminalia occur in all genera except Paykullia
Robineau-Desvoidy and Melanophora Meigen. In the
Calliphoridae, all genera which have retained a plesio-
morphic oviposition habit have telescopic female ter-
minalia. In the Tachinidae, with the present ex-
clusion of Malayia and Bezzimyia, all species possess
shortened female terminalia, if not secondary length-
ened, e.g. in some members of the phasiine tribe
Cylindromyini. In all of Sarcophagidae, female termi-
nalia are much shortened, and this holds for
Chilopodomyia Lopes & Tibana as well, even though
this genus was stated to have a telescopic ovipositor
(Lopes & Tibana 1984). The subfamilies of
Oestridae may also possess shortened female termina-
lia as a groundplan character state. Long female ter-
minalia are unknown in the Cuterebrinae and
Oestrinae, and the elongate terminalia of most fema-
le Gasterophilus are neither retractile nor telescopic. In
the Hypodermatinae females of Hypodermatini and
Oestromyini have long and telescopic ovipositors, but
female terminalia of the genus Portschinskia are short.
I have not been able to break the structural diver-
sity of female terminalia into unambiguous states ex-
cept as a simple, and therefore probably little phylo-
genetically informative division into a short versus a
telescopic condition. Also, the character is closely tied
to the mode of oviposition, and it may be directly
misleading to score larviparous groups and groups
with a derived oviposition behaviour.
Character h. — Female tergites 7 and 8; median
division
McAlpine (1989) considered the division of fema-
le abdominal tergites 7 and 8 as synapomorphic for
the Calliphoridae. I know of no Calliphoridae having
these tergites undivided, while the undivided con-
dition occurs in at least some species of all other fa-
milies of the Calyptratae (Herting 1957). However,
most non-calliphorid terminal taxa included in the
present analyses show examples of both the divided
and the undivided condition and would be scored as
unknown. Information is needed from genera consi-
dered to be close to the basal splits within each taxon.
Character i. — Spermathecal ducts; enclosion/non-
enclosion in acommon sheath
Hori (1961) distinguished between what he called
a 1:2 and a 1:1:1 configuration, where either two
spermathecal ducts were enclosed in a common she-
ath or all three were perfectly free of each other. This
should not be confused with character 25, which
deals exclusively with how the three ducts join the
uterus. Enclosion in a common sheath, therefore,
does not mean that they share a common duct or a
common opening into the uterus.
Apparently, all species of Sarcophagidae and
Tachinidae examined by Hori possess perfectly free
spermathecal ducts, while all Calliphoridae and al-
most all non-tachinoid calyptrates with 3 spermathe-
cal ducts examined showed the paired configuration.
Within the Rhinophoridae, freshiy killed female spec-
imens of Phyto melanocephala dissected by me showed
the paired condition with two spermathecal ducts
closely appressed, while a female of Melanophora ror-
alishad the two ducts only loosely connected. Among
the Sarcophagidae, the miltogrammatine species
Senotainia litoralis Allen figured by Allen (1926: fig.
25) seems to possess a paired configuration, while the
pairing of spermathecal ducts in S. trilineata (Wulp)
seems to be very loose (Allen 1926: fig. 20). I have
dissected freshly killed specimens of Brachicoma devia
(Fallen) of the Paramacronychiinae which showed ei-
ther a free condition or a condition with two loosely
connected ducts. In the Oestridae, the few figures
available do not allow a definite conclusion (Catts
1964; Grunin 1965, 1966), but in the Tachinidae no
examples of joined or paired spermathecal ducts are
known (S. Andersen, pers. comm.). While dissection
of dried females may give easy access to information
on how the spermathecal ducts enter the uterus, fresh
or properly fixed specimens are required to examine
whether or not the ducts are paired or free. I have not
been able to score the families consistently on the
sparse material examined.
Character j. — Female accessory glands; attachment
to oviducts
The detailed figures of Hori (1961) reveal another
Pape: Phylogeny of the Tachinidae family-group
character that may prove to contain phylogenetic in-
formation, the attachment/non-attachment of female
accessory glands to the oviducts through an extension
of their sheath of connective tissue. In all
Calliphoridae and non-tachinoid Calyptratae figured,
the female accessory glands are distinctly attached (see
also Kurahashi 1970: fig. 3), while they are free in all
Sarcophagidae and Tachinidae. Fresh females of
Phyto melanocephala and Melanophora roralis
(Rhinophoridae) examined by me both have the ac-
cessory glands unattached. A single female of
Brachicoma devia (Sarcophagidae: Paramacro-
nychiinae) seen had the accessory glands suspended in
a thin strand, and the female reproductive system of
Gonia sp. (Tachinidae: Tachininae) depicted in
Pantel (1910, identified as atra Cockerell and repro-
duced in Thompson (1963: fig. 2)) shows accessory
glands likewise loosely attached to the oviducts. The
only figures available of internal reproductive organs
of female botflies suggest that the accessory glands are
unattached (Catts 1964: figs. 3-7 (Oestrinae),
Grunin 1965: fig. 37 (Hypodermatinae), Grunin
1969: figs. 16-20 (Gasterophilinae)). These figures,
however, are hardly reliable as the dissections were
not made for the special purpose of examining this fe-
ature.
Character k. — Uterus; presence/absence of lateral
sacs
The uterus of many Calliphoridae, e.g. species of
the genera Calliphora Robineau-Desvoidy, Lucilia
and Pollenia, possesses distinct lateral sacs. These sacs
seem to receive the tips of the dorsolateral phallic pro-
cesses during copulation (Rognes 1991). Nothing
equivalent is known from the Sarcophagidae (Lopes
& Kano 1968), which do not possess lateral sacs
(Hori 1961), nor from the common house fly
(Degrugillier & Leopold 1973), which possesses dis-
tinct lateral sacs but no free dorsolateral processes. No
lateral sacs have been shown to occur in non-callip-
horid tachinoids (I have examined females of
Melanophora roralis and Phyto melanocephala of the
Rhinophoridae).
Character |. — Male terminalia; resting position
Wood (1987a: 1148) described the male terminalia
of the Oestridae as being ‘strongly retracted, and car-
ried rather far forward on underside of abdomen’.
This is distinct from the possible groundplan of the
Tachinidae family-group, where male terminalia are
almost apical. Many subfamilies within the
Calliphoridae, with the notable exception of the
Helicoboscinae, have the male terminalia slightly dis-
placed forwards on the underside of abdomen. I have
not, however, been able to produce an exact measure
for this character.
65
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Character m. — Testes; shape
Shape of testes may provide interesting phyloge-
netic information. In the Paramacronychiinae and
Sarcophagidae, testes are of a uniform shape with a
constriction proximal to middle. In the Tachinidae, a
pyriform shape of testes may belong to the ground-
plan (see numerous figures in Townsend (1942) and
Hori (1960)). Exceptions, however, exist and rigo-
rous use of this character depends on more infor-
mation from all terminal taxa.
Character n. — Testes; position relative to fat body
Based on data from Hori (1960), ‘testes enveloped
in fat body’ was used by McAlpine (1989) to corro-
borate the monophyly of the Calliphoridae.
However, data from many more calliphorid genera
and especially from the Oestridae and Rhino-
phoridae, are needed before it will find proper usage.
Character o. — Median phallic sclerotization; relative
development
This structure (= ‘Mesohypophallus’ of Salzer
(1968)) was given the poor term ‘spermduct scle-
rotization by Pape (1986a) and later renamed ‘me-
dian sclerotization’ (Pape (1987d) following
Tschorsnigs (1985a, b) ‘Medianleiste des
Ventralsklerits’). Although it may appear to be in-
ternal, as the ventral part of the phallus often is slight-
ly invaginated, it is part of the phallic tube, not the
sperm duct, as correctly stated by Tschorsnig
(1985a). A median sclerotization is often present and
well developed in the Tachinidae (see figures in
Tschorsnig (1985b)), less distinct and often shorte-
ned in the Oestridae. The groundplan of the
Rhinophoridae and Calliphoridae most probably is a
well developed median sclerotization running the full
length from the ventral plates to the acrophallus. In
the Sarcophagidae, the median sclerotization is never
complete. What could be a short median scle-
rotization occurs sporadically in the Miltogram-
matinae (e.g. Pape 1987a: fig. 133), but no trace of a
median sclerotization seems to be present in the
Paramacronychiinae. Within the Sarcophaginae, the
highly modified phallus makes homologies obscure,
but a median sclerotization is either absent or may
have given rise to the vesica and all or part of the me-
dian stylus.
Character p. — Dorsolateral phallic processes;
serration of tip
A point of interest relating to the calliphorid phal-
lus is that the dorsolateral processes often are equip-
ped with serrations along the free part of their tip. In
Lucilia cuprina (Wiedemann), these free, serrated tips
have been shown to tear slits in the cuticular lining
(intima) of the lateral sacs of the uterus. The curved
66
dorsolateral processes assist in guiding the accessory
gland material into intra-cuticular spaces (Merritt
1989). Eurychaeta palpalis (Helicoboscinae) and
several species of Calliphorinae, Chrysomyinae,
Melanomyinae, Mesembrinellinae and Polleniinae li-
kewise possess dorsolateral processes with some kind
of serration of the apical part (see figures in
Guimaräes (1977), Kurahashi (1970, 1971) and
Rognes (1986, 1987, 1988, 1991)). Rognes (1988:
342) considered the presence of ‘small to minute
tubercles’ on the tip of the dorsolateral processes as
apomorphic within Pollenia and as an underlying sy-
napomorphy corroborating a clade consisting of his
semicinerea-group and rudis-group. It may be,
however, that the synapomorphy should include all
of the Calliphoridae. I have not seen any serrations,
teeth or tubercles on the apical part of the very callip-
horid-like dorsolateral phallic processes of Phyto spp.
nor in the undescribed species of Bezzimyia shown in
fig. 3, although extensive SEM studies are needed. It
is tempting to postulate an ancestral calliphorid type
of copulation which, possibly repeatedly, led to an
elongation of the free tips of the dorsolateral process-
es with or without apical serration. It is interesting in
this context that many Calliphoridae with long, free
dorsolateral processes have the acrophallus equipped
with a pair of lateral ducts on the external wall of the
acrophallus. These ducts seem to facilitate the flow of
accessory gland secretion from the gonopore to the
slits made by the dorsolateral processes (Merritt
1989, Rognes 1991).
Character q. — ß-alanyl-L-tyrosine (‘sarcophagine’);
presence/absence in larval hemolymph
The accumulation of the dipeptide sarcophagine in
larval hemolymph of at least Agria housei Shewell
(Paramacronychiinae) and several species of Sarco-
phaga sensu lato (Sarcophaginae) is interesting as ‘8-
alanine and tyrosine have never been detected in
peptide linkage in proteins; nor has the free dipeptide
been observed in any other natural source’ (Bodnaryk
1970: 349). Later investigations by Bodnaryk (1972),
however, have revealed sarcophagine in one species of
Tachinidae. The Miltogrammatinae have not yet
been examined for this character.
Character r. — Larval pharynx; presence/absence of a
filter apparatus
McAlpine (1989) corroborated the monophyly of
his parasitic clade Oestridae + Rhinophoridae + Ta-
chinidae by the absence of a larval pharyngeal filter. I
have insufficient information on the distribution of
this character, e.g. in the Miltogrammatinae and
possibly basal calliphorid lineages like the Heli-
coboscinae and Bengaliinae, to be able to make a pro-
per scoring.
Character s. — Labrum of first instar larva; size
A labrum is completely absent in the Rhi-
nophoridae (Bedding 1973), it is small but still dis-
tinct in the Sarcophaginae (Lopes 1982b) and
Oestrinae (Grunin 1966). The labrum is short in the
Cuterebrinae and Hypodermatinae (Grunin 1965,
Ferrar 1987), long and tapering in the Gasterophilini
(Grunin 1969). In the Calliphoridae, a labrum is
usually well developed, but it is absent in the genera
Bellardia and Onesia (Calliphorinae) and in Proto-
calliphora Hough (Chrysomyinae) (Rognes 1991).
Character t. — Dorsal cornu in cephalopharyngeal
skeleton of third (and probably second) instar larva;
posterior incision
The large majority of species within the
Paramacronychiinae and Miltogrammatinae and all
of Sarcophaginae have a posterior incision in the
dorsal cornu. The few exceptions, however, make
groundplan estimates uncertain until generic phylo-
geny becomes better resolved. McAlpine (1989) listed
an unincised dorsal cornu as a tentative autapomor-
phic character state for the Calliphoridae, but he si-
multaneously gave this condition as autapomorphic
for his clade Oestridae + Rhinophoridae + Tach-
inidae.
Roback (1951) gave the incised condition as a
groundplan feature of the Anthomyiidae.
Character u. — Spiracular slits of posterior spiracles
in third instar larva; orientation
The pattern in the Sarcophagidae is generally that
of three more or less vertical slits, and in the
Calliphoridae the most widespread condition is three
slits radiating in a dorsolateral direction (see figures in
Ferrar 1987). In all Gasterophilinae, the three spira-
cular slits are parallel and vertical, with secondary
convolutions having occurred in Gyrostigma. In
Ruttenia the spiracular slits are horizontal or almost
so.
Character v. — Pupal prothoracic horns;
presence/absence
Prothoracic horns are present in the pupa of many
Calliphoridae and Oestridae and in at least some
Tachinidae, while they are seemingly absent in the
Rhinophoridae and Sarcophagidae. I have not inclu-
ded this character as the number of calliphorid and
tachinid subfamilies for which this character is dis-
cussed in available literature is strongly limited.
Character w. — Puparium; shape
The shape of the puparium was used by McAlpine
(1989) as a synapomorphy uniting his parasitic clade
Oestridae + Rhinophoridae + Tachinidae. These
three families were stated to possess a ‘puparium fully
Pape: Phylogeny of the Tachinidae family-group
inflated with both ends more or less equally hemi-
spherical’ (p. 1501). I have been unable to confirm
this.
Character x. — Egg; surface of area dorsal to
micropyle
The micropyle is here considered a morphological
landmark, which means that the chorion dorsal to
this may be considered homologous throughout the
Tachinidae family-group. In at least some and proba-
bly most Calliphoridae, the surface between the two
ridges or pleas demarcating the hatching sutures is
modified into a plastron with struts and crossbars. A
plastron seems likewise to be present in all
Rhinophoridae as well as most Muscidae and
Anthomyiidae (Hinton 1960a, b, 1963, 1965, 1967;
Bedding 1973; Kitching 1976; Leopold et al. 1978;
Richards & Morrison 1972; Erzinçlioglu 1988,
1989; Greenberg & Szyska 1984). The similarity in
position and ultrastructure suggest homology. No ta-
chinid egg is known to possess a plastron as here defi-
ned. Instead, one or more areas with perforations
(aeropyles) in the external covering, and therefore
with free access to the columnar middle layer, may be
found, as shown by figures and SEM-micrographs in
Wood (1972). Although the aeropylar areas someti-
mes are situated dorsal to the micropyle and between
the hatching pleas, I have no a priori evidence that
these are homologous to the plastron of the
Calliphoridae.
The sparse information available on the morpholo-
gy of oestrid eggs indicates ‘non-presence’ of a plas-
tronic network. It does, however, not allow a more
explicit interpretation of homology, e.g. relative to
the condition found in the Tachinidae. In
Dermatobia hominis (Cuterebrinae), the egg cap is
without any trace of a plastronic network and the ex-
ternal layer of chorionin, which in this species is sim-
ilar to the general egg surface, has no apparent open-
ing(s) (Leite 1988). Also in Cuterebra tenebrosa
Coquillett is the sculpturing of the egg cap hardly dif-
ferent from that of the general surface (Baird &
Graham 1973). In Cuterebra cuniculi (Clark) a mod-
ified sculpturing covers all of the egg cap (Baker
1986, as Cuterebra horripilum Clark). It does not look
like a plastron similar to that found in the
Calliphoridae, nor does it look like aeropyles as found
in the Tachinidae. The only SEM-studies of eggs of
Gasterophilini (Cogley & Anderson 1983) and
Hypodermatinae (Cogley et al. 1981) do not pay spe-
cial attention to the egg cap.
Too little is known on egg structure in the
Sarcophagidae to allow a reasonable scoring of the
groundplan. In Miltogramma Meigen, the egg seems
devoid of any plastron or aeropylar area (Thompson
1921), but SEM studies are needed to confirm this.
67
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
In Sarcodexia lambens, the egg is pitted with aeropyles
in the posterior third of what may be the dorsal sur-
face (Lopes & Leite 1989), while eggs of Sarcophaga,
to the extent they are known, only present a single
aeropyle, which is situated opposite the micropyle
(Baudet 1980, 1985). A true plastron is probably ab-
sent in the Sarcophagidae as this would be non-func-
tioning during intra-uterine incubation.
Character y. — 28s ribosomal RNA sequence data
Vossbrinck & Friedman (1989) used sequence data
from 28s ribosomal RNA to analyse relationships
between selected cyclorrhaphan species. Their data
have not been included in the present analysis for the
following reasons. Firstly, ribosomal RNA of far too
few taxa has been analysed, which would have given
this input of character states proportionally more
weight than it may deserve. Secondly, and perhaps
more important, I prefer to await a consistent metho-
dology for phylogenetic analysis of nucleotide se-
quence data. Of particular relevance before trans-
lating Vossbrinck & Friedman's (1989) table 1 into
character states is a discussion of homology between
the sequences analysed. However, it is obvious from
the cladograms produced by these authors that their
sequence data give results that show little congruence
with results obtained in the present analysis.
Character z. — Parasitism
Parasitism of a certain taxon has been used expli-
citly as an apomorphic character state for all major
parasitic clades of the Tachinidae family-group (Pape
1986a, Rognes 1986, McAlpine 1989). McAlpine
(1989: 1501) even stated that being ‘obligate pa-
rasites of other animals .... is presumed to be a pri-
mary, autapotypic adaption of [Oestridae +
Rhinophoridae + Tachinidae] (synapotypic character
of the component families)’. An important question,
however, is how parasitism fits into a character ana-
lysis, i.e. whether or not parasitism really is a char-
acter and in case how the states are defined. As I have
met numerous viewpoints on this issue in discussions
with fellow systematists, a rather detailed elaboration
of the problem seems justified.
Parasitism per se is here defined with the very broad
concept adopted by Price (1980: 4), who simply cites
the definition given in Webster's Third International
Dictionary: A parasite is ‘an organism living in or on
another living organism, obtaining from it part or all
of its organic nutriment, commonly exhibiting some
degree of adaptive structural modification, and caus-
ing some degree of real damage’.
Parasitism, then, could be defined as a character
with the states ‘developed’ and ‘not developed’. This
would imply that, say, the Agromyzidae (parasites of
angiosperms) would be scored exactly like the
68
Rhinophoridae (parasites of woodlice) in a family lev-
el phylogenetic analysis. This way of handling parasit-
ism seems falsified by the very definition of a charac-
ter accepted in the present paper, being an inheritable
feature that is independent of any other inheritable
feature(s). Parasitism per se, as here defined, is not
built in as a message in the genetic code. One could
elaborate the definition of the states so that states (of,
say, life habit) pertaining to parasites were defined
taxonomically as the most restricted (= smallest pos-
sible) recent taxon assumed to contain the hypotheti-
cal ancestral host or host-lineage. Put in a more infor-
mal wording, this means that the habit of parasitizing
a certain taxon is considered a derived character state
for the parasites of this taxon. In this way, parasitism
of true bugs (Heteroptera) would be a synapomorphy
of the Phasiinae, insect parasitism a synapomorphy of
the Tachinidae, mammal parasitism a synapomorphy
of the Oestridae, woodlice parasitism a synapomor-
phy of the Rhinophoridae, etc. But we still do not
have characters or character states in the sense of in-
heritable features — organisms simply do not parasi-
tize taxa, they parasitize other organisms. This may
seem pure epistemology, but parasitism in this taxo-
nomic definition can only be defined a posteriori, and
corroborating a group of species based on the com-
mon habit of parasitizing the same taxon (whether of
specific, generic, or any other rank) is unsound. The
Pieris -butterflies, for example, do not possess the
character state ‘phytophages of Capparales’. They will
readily eat any plant containing (or just tasting of)
mustard oil glucosides, e.g. members of the
Brassicaceae, Tropaeolaceae and some other
Capparales, and the real character state should rather
be ‘mustard oil glucosides promote larval feeding’
(David & Gardiner 1966) or perhaps even more spe-
cific ‘larval maxillary sensilla styloconica with chemo-
receptory cells sensitive to mustard oil glucosides’
(Schoonhoven 1967). Taxonomic host data is merely
information on the distribution of hosts within the
classification available. Modern classifications of host
organisms reflect their “descent with modification’,
which cannot trigger oviposition behaviour.
Thus, in my opinion the monophyly of a group
consisting of the Tachinidae + Rhinophoridae cannot
be argued on the shared occurrence of parasitism of
arthropods simply because arthropod parasitism has
no validity as a character state. And if arthropod par-
asitism cannot be considered a character state, insect
parasitism, wood-lice parasitism or animal parasitism
obviously cannot do either. When dealing with still
more restricted and/or specialized host taxa, however,
one will often increase the probability that its asso-
ciated parasites actually do belong to a monophyletic
group. This, however, is nothing but 4 posteriori sta-
tistics and reflects that many parasite clades in nature
really are restricted to one or a few host clades.
If parasitism (of a taxon) cannot be considered a
character, this implies that even if the Tachinidae and
Rhinophoridae constitute a clade one cannot make a
straightforward deduction that their common ances-
tor was a parasite of either insects or woodlice unless
we assume that the transition between woodlouse
parasitism and insect parasitism (or vice versa) is more
likely than the transition between scavenging and
woodlouse/insect parasitism. Only if synapomorphic
character states directly relating to the parasitic habit
are present can we convincingly argue for a single or-
igin of parasitism, and we may then consider wood-
louse parasitism to have developed from insect para-
sitism (or vice versa) simply because this is more
probable than the hypothesis of both habits arising si-
multaneously, i.e. in the same speciation event.
Wood (1987b: 1189) tentatively suggests one such
character: “The tendency to stimulate the formation
in the host of [a funnel-like integumental sheath] or
at least to use it to advantage for support and respira-
tion, may be a synapomorphy of the Rhinophoridae
and Tachinidae’. Two problems should be considered
in this context. Firstly, of course, one has to decide
whether or not to accept the character at all. In my
opinion, the formation of a sheath enveloping part of
the parasite is a typical immunological reaction that
would happen to any foreign body piercing the integ-
ument and penetrating into the haemocoel. Thus, the
formation of a sheath could be plesiomorphic at this
level. No sheath-formation has been reported from
hosts of parasitic Sarcophagidae, e.g. the grasshopper-
parasitizing Blaesoxipha (Léonide & Léonide 1986),
which seem to be independent of direct access to the
surrounding atmosphere in at least second and third
instar. More tricky is the question whether or not the
use of this sheath for support and respiration is a char-
acter state, but as the formation of the sheath prob-
ably is induced by the piercing of the integument, e.g.
a trachea, use of the sheath cannot be considered in
isolation. Numerous types of larval equipment for
piercing the host's integument may be found in the
Tachinidae, but all of these differ from what may be
analogous structures in the Rhinophoridae and ap-
parently without being related through transforma-
tion series. I have not accepted the use of a respirato-
ry funnel or any associated feature within
Rhinophoridae and Tachinidae as homologous.
Another matter is, that if the production within the
host of a sheath of host integument, or its being used
by the parasite, is considered a (derived) character
state, the character would be inapplicable with regard
to the Oestridae. Oestrid hosts, i.e. mammals, differ
radically from arthropods in cytology, physiology and
immunochemistry. A respiratory funnel, therefore,
cannot in itself provide evidence that the
Pape: Phylogeny of the Tachinidae family-group
Rhinophoridae are more closely related to the
Tachinidae than are the Oestridae. Note the few but
well documented cases of human myiasis produced
by tachinid flies that even have resulted in the emer-
gence of adults (Smith 1988). This shows that we
cannot a priori exclude the possibility that mammal
parasitism evolved from insect parasitism.
CLADES OF THE TACHINIDAE FAMILY-GROUP
Running the present character matrix (table 2)
through Hennig86 (version 1.5) with all states unor-
dered (outgroup = 0.1;cc-.;mh*;bb*;) results in an
initial set of 630 most parsimonious cladograms
(length = 111, consistency index = 55, retention in-
dex = 77). Subsequent successive weighting (xsteps
w;mh*;bb*;xsteps w;mh*;bb*; etc.) reduces these 630
cladograms to 15, although with no changes in fami-
ly level relationships. From the resulting Nelson
consensus tree (fig. 9), deductions of transformation
series between character states can be made for those
characters where more than two states are involved
(i.e. characters 0, 9, 10, 13, 15, 21, 22 and 39).
Characters 0, 9, 13 and 21 may thus be arranged in li-
near transformation series, which correspond to the
numeral sequences used when listing the states in the
discussion above. [Note that transformation series
need not be rooted at the lowest number, i.e. ‘0°.
Character 13 has the transformation series 0-1-2 but
is rooted at 1]. For characters 10, 15, 22 and 39, less
straightforward transformation series are involved,
and these characters were maintained as unordered
(cc-10,15,22,39;). Specifically for the subscutellum
(character 10), the cladogram in fig. 9 leads to the
assumption that the convex state (2) most probably
developed from the concave state (0) in the
Tachinidae and from the medium-sized state (1) in
the Hypodermatinae, and as the concave state also
may have given rise to the medium-sized state as well
as vice versa, all states are interconnected and should
be treated as unordered.
With this new input of phylogenetic information,
data were rerun (outgroup = 0: 1sce-
10,15,22,39;mh*;bb*;xsteps w; etc.). This resulted in
30 trees, the Nelson consensus tree of which did not
differ from the one resulting from all characters being
unordered. This cladogram is the one I consider as
the currently most corroborated hypothesis of family-
level phylogenetic relationships within the Tach-
inidae family-group (fig. 8).
Competing cladograms resulting from different
character or transformation series interpretations are
discussed below (figs. 10, 11), with character states
for non-terminal clades in tables 4, 5.
Explicit lists of apomorphic character states corrob-
orating the monophyly of families and supra-familial
69
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
SPUTSIEIOXOL =
SPUTTUTUA =~
eeuTAWUOSAIUI En
seutAuoue Ten
SBUTTTIONT
OPUTIOUATIIED
SEUTTSoUNnUd
eeur TUSWUY
SPEUTT TSOUTIAQUWISON
SEUTIUSTTOd 5
oeuttrefusg a:
SPUTDSOYODOTTSH
| | =0=0=0=0=0=0=0=0=0=0=0=0=0=0=0=0
SEUTIASISAND
TUTTPTOqqoD
TUTIIudors3se9
=:
SEUTJEWUISPOAÄH 2
dE
E
St
Gd AERO
SEUTIISSO
eeurbeydoozes Col Ark vanme Ne N ee Bi Er}
Se a = in © a NW oO
SEUTTYUDÄUOIDEUEIEG + =0=0=0=0=0=0=0 alal ot URS)
ee ||
SEUTTSEeUd
dnoxb-snueb PISTIOXH =
SEUTUTUDeL aq
oepIosnn
SEPTTÄUOUJUY RE)
Fig. 8. Cladogram of the Tachinidae family-group with apomorphies indicated for non-terminal clades. Numbers refer to
70
characters and states discussed in the text.
Pare: Phylogeny of the Tachinidae family-group
Table 2. Initial character matrix with scorings of 50 characters (0-49). See text for discussion of states.
Anthomyiidae 00000 00000 000-0 10000
Muscidae 00000 00000 000-0 10001
Rhinophoridae 10000 00001 11101 30001
Miltogrammatinae 01000 00001 0-111 20001
Paramacronychiinae 10000 00001 O1111 20001
Sarcophaginae 10000 10-11 01111 20001
Phasiinae 10000 00001 21111 20001
Exorista genus-group 10000 00001 21111 20001
Tachininae 10000 -0001 21111 20001
Cuterebrinae Dee De er Oe SOS
Gasterophilini 2101 2112127 1001 11100
Cobboldiini DNA VOM KO!
Hypodermatinae 21101 11112 200-1 01101
Oestrinae 21101 10-12 -Olll 21101
Helicoboscinae 10000 11011 11111 20001
Polleniinae 100-0 00101 11121 20001
Bengaliinae 10000 1-111 11121 20000
Mesembrinellinae 10010 10111 110-1 300-1
Ameniinae 10010 11111 11121 20001
Phumosiinae 10010 11111 01121 20001
Calliphorinae 10010 11111 01121 20001
Luciliinae 10010 11111 01121 20001
Melanomyinae 10020) = 12120001
Chrysomyinae 10010 11111 01121 20011
Rhiniinae 100-0 00111 01121 20011
Toxotarsinae 10010 11111 01121 20011
00001 00000 00000 00000 00000 00000
-1001 00000 00000 00000 00000 10000
13100 10000 11-00 01000 00000 10000
12310 10-01 11-01 10—1 00011 00000
12310 10111 11-11 100-1 00011 00000
12310 10001 11-11 100-1 00011 00000
12100 10000 11-01 00100 11000 10000
12200 10000 11-01 00100 11000 10000
12200 10000 11-01 001-1 11000 10000
-2400 00110 11000 01110 001-0 11001
02410 -1110 00-0— 00110 00100 11011
12400 -1110 01000 00110 00100 11011
—2410 01110 -100- 0-110 00100 11101
12410 -1110 -1000 10—1 00100 11101
12100 10000 11100 00—1 00000 00000
-2300 00000 11100 01000 00000 00000
123-0 10000 11-0- 0-000 00000 00000
12000 00000 11100 00—2 00000 —-0000
12110 00000 11100 0—-1 00000 0-0—
12300 10000 11100 0—-1 00000 0-0—
12300 00000 11100 00000 00000 00000
12300 00000 11100 00000 00000 00000
12300 00000 11100 00000 00000 00000
1-300 00000 11100 00000 00010 00000
13300 -0000 11-01 00000 00000 —0000
123— -0000 11100 00000 —0000 00000
clades within the Tachinidae family-group are listed
below, including additional character states that may
further corroborate the monophyly of the families,
but which were not scored for the analysis.
Apomorphic character states have been given a con-
cluding composite number (character: state) referring
to the discussion of characters and to the data matrix;
or a letter for reference to the discussion of unscored
characters.
Oestridae
As pointed out by Wood (1987a), the early con-
cept of the botflies as a ‘natural’ (i.e. monophyletic)
group has gained wider acceptance after a period with
attempts of splitting the group and assigning the
fractions to various other calyptrate families or even
to the acalyptrates (see Grunin (1969) for references).
Although mammal parasitism has evolved several ti-
mes in the Calliphoridae and Sarcophagidae, it has
never promoted the physiological coevolution shown
by the botflies and their hosts. Some species of
Wohlfahrtia (Sarcophagidae) and Booponus
(Calliphoridae) are subdermal parasites of mammals
and at least some species of the latter genus seem to be
quite host specific. But the warbles, if any, formed by
these species are simple swellings caused by the in-
flammatory response of the host. Contrary to the bot-
flies, these larval feeding sites often leave a considera-
ble scar.
NWW KRW D —=
Presently recognized botfly synapomorphies are:
. Postocular setae reduced (0:2);
. Postcranium concave (1:1);
. Mouthparts reduced (2:1);
. Clothing setae soft and hairlike (4:1);
. Meron with a patch of hairs (9:2);
. Anatergite bare (11:0);
. Subcostal vein straight at level of humeral break
(16:1);
. Vein R, without a knob at level where subcosta
joins with costa (17:1);
. Abdominal sternite 2 freely exposed and widely
separated from tergal margins by ample pleural
membrane (22:4);
. Two of the three spermathecal ducts fused just
before entering the uterus (25:0);
. Male sternite 5 simple or with shallow emargina-
tion (27:1);
. Male tergite 6 fused to syntergosternite 7+8
(28:1);
. Ventral surface of egg glued to substrate (37:1);
. Egg hatching through discarding an anterodorsal
cap or operculum (38:1);
. Segments of first instar larva with 1-2 bands of
strong spines (42:1);
. Second and third instar larva with ecdysial scar of
spiracular plate near horizontal diameter (46:1);
. Puparium hatching by splitting off only one, dor-
sal piece (49:1).
71
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Fig. 9. Nelson consensus cla-
dogram of 15 trees obtained
from running the matrix in
table 2 with all characters un-
ordered and successive
weighting. Weights and
character states of nodes are
given in table 3. For further
explanation see text.
Rhinophoridae
The Rhinophoridae have recently been subject to
some attention, but despite several attempts at a de-
finition based on apomorphic character states in the
adult morphology, only the morphology of the first
instar larva has provided what seems to be unambi-
guous and autapomorphic character states.
Crosskey (1977) reviewed the family and provided
a thorough discussion of the characters most often
used in the characterization of the family. He admit-
ted, however, that his recognition couplet for ima-
gines still fits “a very few aberrant tachinids’ (p. 7).
Tschorsnig (1985a), although including both
Melanomya and Morinia, both of which now general-
ly are accepted as Calliphoridae, described the struc-
ture of the male terminalia in detail and suggested
(tentatively) that broadened bacilliform sclerites and a
patch of setae anterobasally on the surstylus should be
considered synapomorphic for the Rhinophoridae
(see his figs. 7-10). Exceptions exist, and as some of
these represent undescribed genera (Pape, unpubl.) I
prefer to await a revised generic phylogeny with data
on this character incorporated before evaluating the
rhinophorid groundplan.
The phallic character state previously used by me,
the fusion of the ventral plates in the median line
(Pape 1986a), apparently does not have the diagnos-
tic or defining powers stated. Although it may be apo-
morphic at the level of the rhinophorid groundplan,
numerous homoplasies occur in the Calliphoridae
and Tachinidae, and even within the Rhinophoridae
reversals to the non-fused condition have to be as-
YA:
= Anthomyiidae
ah Muscidae
Bil
ie Tachininae
Exorista genus-group
Es
Phasiinae
3 ic Miltogrammatinae
3 6 È
IL TG Paramacronychiinae
32-— Sarcophaginae
f= Rhinophoridae
403 Oestrinae
| rok
Hypodermatinae
pre la Gasterophilini
27== Cobboldiini
L m: = Lo Cobb
mule ele Helicoboscinae
sl m age nee
ay) Te Polleniinae
29-— Mesembrinellinae
Ameniinae
Phumosiinae
Calliphorinae
Luciliinae
Melanomyinae
m Chrysomyinae
VE Rhiniinae
Toxotarsınae
Ar
[Tm
sumed (Paykullia, several species of Phyto). I do not
consider this character any further in this context.
Wood (1987b: 1189) clearly pinpointed the current
situation when he wrote: ‘no character or group of
characters has been found in the adult to render the
group recognizable or to suggest that it is
monophyletic’. But, on the other hand, the obviously
apomorphic larval character states bring substantial
evidence that a very well defined monophyletic group
exists, and only our sparse knowledge of this stage
prevents an easy delimitation of the group. Hence, I
cannot agree when Downes (1986: 17) ‘interpret[s
the Rhinophoridae] as an intentionally practical rath-
er than a natural taxon’ which ‘in practice....is analo-
gous to the Fungi Imperfecti’.
All species of Rhinophoridae are obligate parasites
(or parasitoids), but parasitism of woodlice will not
be included as a character state, as already discussed.
1. Metathoracic spiracle with both anterior and pos-
terior lappets small (13:0);
2. Vein M bend in an obtuse angle (15:3);
3. Lower calypter tongue-shaped (21:3);
4. Female ovipositing away from food source
(36:1);
5. First instar larva with completely reduced labrum
and closely appressed mandibles (s).
Some additional character states are found in the
larval morphology (see figures in Bedding 1973).
6. First instar larva with dorsal arc of mandibles
toothed or serrated;
7. First instar larva with anterior part of pharyngeal
sclerite elongated;
ál 42
2
40
GON MAGEN 39
Pape: Phylogeny of the Tachinidae family-group
35
BO 31 SAS” Sih Nor!
29
01
28
12
—
Node
D6 27
Weht
10
10
0
2
oo Ooo SooS OVO OO SO) OG)
-ooroooo- op _°>7o PO O _ _S SO _ SS _ _S _S _S _ _ SS 9
Hroo-©-oo005- ooo
Hm Oo Oo Hu o o oo o oo lr ono_IS xD SO OSO OSO OS OS _S __S _S _S © © © ©
-—oorooce o i o © © © OS O0 © © © SOS © © ©
OOS Ole Or OO OO LOOS OIS
MHOnmrOCCOF OO O0 © © mm O © © mm © © © © ©
Table 3. Character states and weights for clades 26-42 in fig. 9.
0
0
4
1
10
Il
0
4
4
0
3
2
0
0
4
3
0
4
10
0
10
10
10
3
10
2
10
10
10
10
Char
75
lobes can be considered homologous, as discussed
(and accepted) by Pape (1986a).]
whether or not the inflated vesicles and sac-like
lobes. [The use of this character state depends on
8. First instar larva with modified posterior end;
with inflated, ventral vesicles or terminal, sac-like
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 4. Character states and weights for clades 26-42 in fig. 10
40 ál 42
39
SOS ASS
920000997 11340 1351
Sil
290050
Node
DO AA AS
Weht
Char
So SoS err) SoS Soy) (=)
LOSS TOE
01
01
01
01
01
01
19
20
Di
N
4 014 0
1
0
0
N © ©
© © ©
© © ©
mm © © ©
© © ©
© © ©
i=)
>
== See ei
(©)
(LE ta CERCLE)
Sat eee © Sia
(©)
in) MENENS)
OO Om
OON NO Er
oO
SLR Oo O(a
i SS) SSS)
OO Sin
min NOM ER)
tm Om O0
SE SoCo a>
= em)
a iS SSS)
Sh AIS) SSS) SS)
|A
==
SITINIMNLIUANIIE
AAA DMM MDA M M
> PSP PO OS SO SO SO SS SS SO DOSSI
mm © © © © © © © ©
© © - © © © ©
© © © ©
© © © © ©
ODD
gite 1+2 (22:3);
1. Abdominal sternite 2 overlapping margins of ter-
2. Alpha setae absent (23:1);
3. Bacilliform sclerites (divided male sternite 10) re-
With the convincing exclusion of Eurychaeta provi-
ded by Rognes (1986), the Sarcophagidae have emer-
Sarcophagidae
duced and more or less perpendicular to the me-
dian plane (29:1);
ged as a very well-corroborated monophyletic group,
with the following probable apomorphies:
74
Pape: Phylogeny of the Tachinidae family-group
Anthomyiidae
ee Muscidae
Oestrinae
JT se Hypodermatinae
38 E Gasterophilini
n gio sm Cobboldini
Cuterebrinae
Phasiinae
40==
ata Rhinophoridae
Pel
Exorista genus-group
Tachininae
Helicoboscinae
zl
JE
Sarcophaginae
Paramacronychiinae
a
Mime 0== Miltogrammatinae
=i
Lif
Bengaliinae
m Phumosiinae
29 a Polleniinae
Lol Mesembrinellinae
Fr Ameniinae
r Calliphorinae
maus mes
Melanomyinae
È E Chrysomyinae
Are Rhiniinae
Toxotarsınae
Fig. 10. Nelson consensus cladogram of 72 trees obtained from running the matrix in table 2 with Rhinophoridae scored for
free tips of dorsolateral phallic processes. Weights and character states of nodes are given in table 4. For further explanation
see text.
12
. Uterus with a bilobed ventral pouch (35:1);
. Progeny deposited as embryonated eggs or pre-
hatched first instar larvae (39:1);
. Peritreme of posterior spiracles of second and
third instar larva incomplete and without a dis-
tinct ecdysial scar (43:1);
. Posterior spiracles of second and third instar larva
placed in a recession or cavity (44:1);
. Parastomal bars of third instar larva present
(45:0);
. Median sclerotization of phallus reduced (0);
. Dorsal cornu of third (and probably second) in-
star larval cephalopharyngeal skeleton with a pos-
terior incision (t);
. Prothoracic spiracular horn not protruding
through puparium (v);
ß-alanyl-L-tyrosine (‘sarcophagine’) present in
larval hemolymph (q).
[The use of the presence of larval sarcophagine as
an apomorphy corroborating the monophyly of
the Sarcophagidae is, of course, tentative as too
few records exist and especially as no
Miltogrammatinae have been tested. But the al-
most total absence of this dipeptide from the
non-sarcophagid calyptrates is strong evidence
that the presence in the Sarcophagidae is derived,
and testing one or more species of Milto-
grammatinae may indicate whether this character
state should be considered autapomorphic for the
clade Paramacronychiinae + Sarcophaginae or for
all of the Sarcophagidae.]
Some of the autapomorphic groundplan character
states used by McAlpine (1989) to define the
Sarcophagidae are not accepted here and should be
discussed briefly: Coiled male accessory glands, scle-
rotized plate (sigma) in dorsal wall of uterus, looping
of ovarian ducts and common oviduct, and reduced
labral sclerite in the first instar larva, are states that de-
fine all or part of the subfamily Sarcophaginae. These
states are not present in the Miltogrammatinae and
Paramacronychiinae and should not be considered
groundplan states for the family. Arista bare on at
least apical half needs reconsideration as already dis-
cussed. If the sister group of the Sarcophagidae is the
Tachinidae, a bare arista seems at least as probable for
the hypothetical groundplan.
Tachinidae
The Tachinidae form a huge family and the species
have diversified morphologically to an extent not seen
in the other families of the Tachinidae family-group.
However, apart from some species that are strikingly
rhinophorid-like in adult morphology, like
Litophasia Girschner, Tachinidae are easily recogni-
zed and they emerge as a well defined group of insect
parasites. The few non-insect parasites with centipe-
des, spiders and scorpions as hosts (Wood 1987c,
75
42
36 37 38 39 40 41
34 35
320033
BB SOS Sil
28
Node
DO 274
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Weht
Table 5. Character states and weights for clades 26-42 in fig. 11.
Char
NEKO FORSE Hoo-o0o000900000 SS Cee SEN)
01
O0 0000 mm ese se nm mm NN © © © mm nm © ©
iN © © © mm ON M © ©
000000 rm m = nsosor nam oo mm OO II OON ISIS: STE SS
One ee BN TOP NO NOOT 000001000000 0©
© len)
©
Om INNO NON HOOmTrOnrtoOooo OO
ei © =
E
Cent SS SENOS SS ON OC OS OS OS Oat Oe SEN SAMI SEI NOOC
= i i = u i vd vi Lon] Lun | =! — pd vl = = — | nl i en }
MAYHNOR WDHOCAAMNMNHORWDDADOANMMWMNHORDHDDOANHMTNHORADOCAAMNYTHVYDYD
OO SS SH CC NNNNNNNNNMMMMMMMMMMTTITITITTAT A
fined sclerotization on side of mouth opening
(40:1);
4. First instar larva with labrum firmly fused to, and
contiguous with remaining cephalopharyngeal
3. Mandible of first instar larva reduced to an ill-de-
skeleton (41:1);
76
10
Subscutellum greatly swollen (10:2);
2. Ventral surface of egg glued to substrate (37:1);
The following character states are here recognized
Williams et al. 1990), have most probably evolved
as tachinid synapomorphies:
from insect parasites.
IE
Anthomyiidae
Muscidae
met
Es
„em
42
Pape: Phylogeny of the Tachinidae family-group
Sarcophaginae
Paramacronychiinae
Miltogrammatinae
Pee
mp IoxeEarsinae
Rhiniinae
Chrysomyinae
Melanomyinae
Figa
TE
Luciliinae
Calliphorinae
Phumosiinae
Ameniinae
Mesembrinellinae
Bengaliinae
Polleniinae
Le bi Rhinophoridae
37
„Ie na Te JE
air
Tachininae
„JE Exorista genus-group
Phasiinae
Helicoboscinae
Cuterebrinae
30 Cobboldiini
À pet Gasterophilini
= [: fs
ly
Hypodermatinae
Oestrinae
Fig. 11. Nelson consensus cladogram of 14 trees obtained from running the matrix in table 2 without character 32. Weights
and character states of nodes are given in table 5. For further explanation see text.
5. Anterior malpighian tubules shortened in adults
(f);
6. Testes pyriform (m).
Calliphoridae
The blowflies have generally been recognized
among members of the Tachinidae family-group by
exclusion of the more easily defined remaining fami-
lies, and Griffiths (1982) concluded that no convin-
cing evidence corroborated the possible monophyly
of the Calliphoridae. Hennig (1973: 69) even
mentioned that the Calliphoridae could be paraphy-
letic (‘vielleicht eine paraphyletische Gruppe’). In the
present analysis the Calliphoridae emerge as mono-
phyletic, corroborated by the following character
states:
1. Phallus with free tips of dorsolateral processes
(32:1);
2. Parastomal bars present (45:0).
As already discussed, teeth or spines apically on the
free apices of dorsolateral processes may be another
synapomorphy (character p), but as long as both the
toothed and the non-toothed condition occur widely,
an assessment of the groundplan state within each of
the terminal taxa requires a better knowledge of the
distribution of both states as well as of generic phy-
logeny.
It is noteworthy that the metallic coloration of the
integument cannot be considered apomorphic at the
level of the Calliphoridae. Either this character state is
synapomorphic for Calliphoridae and Oestridae, or it
will define a subgroup of Calliphoridae, i.e. by exclu-
sion of at least the Helicoboscinae.
McAlpine (1989) mentioned several tentatively ap-
omorphic character states, but most of these need re-
consideration. A coprophagous larval food habit is
hardly ‘autapomorphic’ in the Calliphoridae, and
even McAlpine himself did not really believe this as
he used this state to corroborate the monophyly of his
clade Calliphoridae + Mystacinobiidae +
Sarcophagidae. It is worth noting that coprophagy in
the Calliphoridae may not be plesiomorphic either, as
none of the basal calliphorid lineages in the present
cladogram (fig. 8) are typical coprophages. Rognes
(1991: 27) is of the opinion that blowflies ‘primitive-
ly … lay their eggs on the exposed dead bodies of var-
ious animals, especially vertebrates, irrespective of
size’, but I find it very likely that the groundplan cal-
liphorid breeding strategy was one of small-carrion-
exploitation, e.g. dead invertebrates (especially snails)
and small vertebrates. This is in agreement with the
arthropod and mammal parasitism of the
Rhinophoridae, Tachinidae and Oestridae, which has
to have evolved from a non-parasitic life habit. The
hypothetical ancestors of these taxa could easily have
preferred small carrion: invertebrates for the
77
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Rhinophoridae and Tachinidae, rodents for the bot-
flies. "Thoracic spiracles large’ is closely connected to
the size of the fringes, which has been discussed in de-
tail above (character 13: state 2). Whether or not
large size of spiracle (or anterior fringe) belongs to the
groundplan of the Calliphoridae depends on the po-
sition of the Helicoboscinae. The ‘posterior thoracic
spiracle occluded by ... complex shutters’ is obviously
a groundplan feature of all of the Tachinidae family-
group, as was correctly stated by Crosskey (1977: 8),
although this author did not mention the Oestridae.
‘Arista plumose’ needs a more accurate scoring, but
may well be synapomorphic with the Oestridae.
Position of female abdominal spiracle 7 needs further
consideration through scoring of more species from
what may be considered basal lineages, and the pres-
ence of ‘anal lobes’ in the male perianal membrane
may belong to the groundplan of the Tachinidae fam-
ily-group. Other groundplan features have been dis-
cussed above.
Calliphoridae + Oestridae
1. Prosternum setose (5:1);
2. Proanepisternum setose (6:1);
[3. Postalar wall setose (7:1);]
4. Metasternum setose (8:1).
The third character state is given in brackets as its
status as a synapomorphy for Calliphoridae and
Oestridae is uncertain. The bare and the setose states
are equally likely for the groundplan of this clade. It
may, perhaps, be argued that an independent origin
of setae on this surface is less likely than independent
reductions.
Sarcophagidae + Tachinidae
1. Phallic dorsolateral processes fused (34:1).
Fused dorsolateral processes were used by Pape
(1987a: 11) to corroborate the monophyly of the
Sarcophagidae, but Tschorsnig (1985b) argued con-
vincingly that the dorsal (or posterior) phallic sclerot-
ization so common and widespread in the Tachinidae
is homologous to fused dorsolateral processes. The
character state will therefore indicate phylogenetic re-
lationship.
Rhinophoridae + (Calliphoridae + Oestridae)
1. Subscutellum slightly swollen (10:1).
This synapomorphy is, of course, highly dependent
on the scoring of subscutellar states, as will be dis-
cussed in more detail below.
78
Tachinidae family-group
Only two outgroups were used in the present stu-
dy, Anthomyiidae and Muscidae. As these may be
part of a separate clade, the Muscoidea of McAlpine
(1989), the present analysis cannot give a reliable esti-
mate of whether ground plan character states at the
level of the Tachinidae family-group are apomorphic
or plesiomorphic. Therefore, the states listed below as
probably synapomorphic for members of the
Tachinidae family-group may be tested when a more
detailed phylogeny of all of the Calyptratae becomes
available. However, in my opinion none of the states
listed below can be ascribed to the groundplan of any
non-tachinoid family of Calyptratae.
1. Postocular setae of alternating size (0:1);
2. Meron with row of setae (9:1);
3. Anatergite with setulae (11:1);
4. Metathoracic spiracle with divided (opercular)
fringe (12:1);
5. Anepimeron with setae (14:1);
6. VeinM with a distal angular bend (15:2);
7. Vein R,,; setose basally (20:1);
8. Lower calypter broad and following scutellum for
considerable distance (21:2);
9. Abdominal sternite 2 contiguous with or slightly
overlapped by margins of tergite 1+2 (22:1);
10. Spermathecal ducts join uterus separately (25:1);
11. Gonostylar apodeme present (30:1);
12. Phallus with denticles along the ventral surface of
distiphallus (31:1).
Mystacinobia
The single New Zealand species of this peculiar
taxon, M. zelandica Holloway, is highly adapted to
living in bat roosts and the morphology leaves few ap-
parent clues to phylogenetic relationships. Originally
described as a distinct family Mystacinobiidae belon-
ging to the Drosophiloidea (Holloway 1976),
Griffiths (1982) argued convincingly for an inclusion
in the Tachinidae family-group and proposed syno-
nymization with the Calliphoridae although no sy-
napomorphic character states with this family were
suggested. Kurahashi (1989) accepted an assignment
to the Calliphoridae and put Mystacinobia in a subfa-
mily of its own.
McAlpine (1989: 1500) recognized an affınity
between Mystacinobia and the Calliphoridae but en-
dorsed family rank for the genus because it possesses
‘many autapomorphies in all stages .... that set it apart
from all Calliphoridae’ — in my opinion a little con-
vincing argument. Furthermore, McAlpine's clado-
gram (Mcalpine 1989: fig. 116.8) corroborated the
monophyly of the non-mystacinobiine blowflies (his
Calliphoridae) by listing the synapomorphies 1) first
larval instar without paired mandibles, 2) arista plu-
mose, and 3) testes enveloped in fat body. The first
state was based on Hennig (1973) and is erroneous as
already mentioned. The second apomorphy is dubi-
ous and actually contradicted by McAlpine himself
on p. 1501, where he stated that a plumose arista is
present in the calliphorid + sarcophagid groundplan.
The third synapomorphy is hardly more convincing
as no mention is made of the actual distribution of
this character state, and Hori (1960), who first used
this character, examined only very few representatives
of Calliphoridae. Even if fresh or properly preserved
specimens of Mystacinobia were examined by
McAlpine, we are still in need of data from many oth-
er members, especially from the Oestridae and
Rhinophoridae, and from possibly basal calliphorid
lineages like the Helicoboscinae. McAlpine listed
more calliphorid autapomorphies in the text (p.
1499), but none of these will corroborate a separate
family status for Mystacinobia. The non-mystacino-
biine calliphorids cannot be defined by being co-
prophagous when their sister group is stated to be ‘liv-
ing in [bat] guano’ (p. 1500) and when coprophagy is
listed as a groundplan feature of Sarcophagidae +
Mystacinobiidae + Calliphoridae (fig. 116.8).
Indeed, Mystacinobia is highly aberrant, and as on-
ly a single species is involved, monophyly as such is ir-
relevant. A list of autapomorphic character states is
given by McAlpine (1989) and needs not be discussed
here. Griffiths (1982) gave three character states of
Mystacinobia that will corroborate its assignment to
the Tachinidae family-group: 1) 8th sternite of fe-
male entire, 2) phallus with denticles on ventral sur-
face of distal section, and 3) meron with row of setae
below metathoracic spiracle. To these could be added
the presence of anepimeral bristles (character 14 in
the discussion above). Many more characters are in-
volved in the definition of the Tachinidae family-
group, but several of these are wing-characters and
therefore inapplicable, which probably holds also for
the infrasquamal setulae (character 11). Thus, only
four characters need a comment. Postocular setae are
hardly recognizable in Mystacinobia, and the great re-
duction in eye-size and the deviating head-shape
makes this character inapplicable. Abdominal ster-
nites, in their degree of exposition, may be more like
the condition found in the Oestridae than that of any
other taxon here considered, and it is thus perfectly
possible — even probable — that it is derived relative to
the groundplan of the Tachinidae family-group. The
small metathoracic spiracle of Mystacinobia has no
fringe at all, and could thus easily be considered as de-
rived from any of the conditions found in the
Calyptratae. Moreover, it is almost to be expected
that a flightless species living in the constantly damp
atmosphere of a bat roost should show modified
Pare: Phylogeny of the Tachinidae family-group
metathoracic spiracles, as this is one of the major sites
for loss of water in a flying calyptrate fly. Finally, no
gonostylar apodeme is present, and this may in itself
be taken as evidence of a sister group relationship
with the remaining Tachinidae family-group.
However, other reductions are present in the male
copulatory apparatus, e.g. small gonocoxites and ab-
sence of epiphallus, and it seems somewhat premature
to assign familial status to Mystacinobia — not to men-
tion a sister group relation to the (remaining)
Tachinidae family-group — based exclusively on this
sparse evidence.
With regard to familial affinities, an inclusion
within any of the Oestridae, Rhinophoridae,
Sarcophagidae and Tachinidae seems highly improb-
able as this would imply that Mystacinobia should
have lost the numerous character states defining these
clades. Also, I have no convincing evidence that
Mystacinobia should belong to any subfamilial clade
within any of these families. Inclusion within the
Calliphoridae cannot be corroborated either.
Mystacinobia does not possess any of the two charac-
ter states here listed as synapomorphies for the
Calliphoridae, namely parastomal bars and free tips
of dorsolateral phallic processes, and there is no obvi-
ous calliphorid subfamily with which it could be affil-
iated. Still, however, the spermathecal ducts are of the
“two fused’ condition (see character 25 above), which
is absent from all of Rhinophoridae, Sarcophagidae
and Tachinidae. In the present analysis this character
defines a large subgroup of the Calliphoridae, namely
all subfamilies except Helicoboscinae and
Bengaliinae.
The dorsolateral processes are fused as in the clade
Sarcophagidae + Tachinidae and as in the Rhiniinae
of the Calliphoridae. However, no other evidence fa-
vours an inclusion in either of these groups, and no
member of the Rhiniinae has yet been reported from
New Zealand (Dear 1985).
I have not included Mystacinobia in the analysis at
the present state of knowledge. A strict scoring would
hardly give a reliable result and would rather interfere
with the not too stable phylogeny presented.
Therefore, I will simply consider Mystacinobia as a ge-
nus (and species) incertae sedis within the Tachinidae
family-group. Until a better corroborated hypothesis
for the inclusion within one of the existing taxa, or for
a sister group relationship to one or more of the oth-
er families is available, I prefer to avoid using a fami-
ly group name to accommodate the species.
Discussion
An interesting — although hardly surprising — result
of the present analysis is that the monophyly of all fa-
milies except the Calliphoridae is well corroborated.
79
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
The monophyly of the Calliphoridae is supported by
few, and perhaps not very convincing synapomor-
phies, and the family may even come out as polyphy-
letic with regard to the Helicoboscinae under certain
circumstances, which are discussed below. Also inte-
resting is the surprisingly few derived character states
that define suprafamilial clades, and it is noteworthy
that none of these states have been used previously in
corroborating inter-familial phylogenetic relation-
ships. We simply have no conclusive evidence on any
sister group relationship at the family level within the
Tachinidae family-group. As a consequence of this li-
mited evidence, of course, the cladogram favoured
here is far from stable. Reinterpretations of characters
and/or the addition of new characters by future stu-
dents are highly encouraged to test the present hypo-
thesis.
In the Calliphoridae, presence of parastomal bars is
a very consistent character state in the sense that ho-
moplasic occurrences are virtually non-existent. I
know of no calliphorid larva without parastomal bars
in the third instar, except for Mystacinobia if evidence
can be found that corroborates an inclusion in the
Calliphoridae. Parastomal bars do not occur at all in
the Tachinidae, Rhinophoridae and Oestridae. It is
interesting, however, that presence of parastomal bars
generally has been considered plesiomorphic in the
Calyptratae, e.g. by McAlpine (1989).
The free tips of dorsolateral phallic processes was
used by Rognes (1986, 1991) to unite the
Rhinophoridae and the Calliphoridae; with the
Rhinophoridae as a subfamily, but not necessarily as a
sister group of the remaining Calliphoridae. As al-
ready discussed, I have been unable to score the
groundplan of the Rhinophoridae for this character,
which in combination with its inapplicability with re-
gard to the Sarcophagidae and Tachinidae (see below)
really leaves it with very limited diagnostic power in
the present analysis. Thus, it is interesting to note
that if the Rhinophoridae are scored for free tips of
phallic dorsolateral processes in the matrix (i.e.
Rhinophoridae are scored for 1 in character 32 of ta-
ble 2), and if this is run as discussed above (outgroup
= 0.1;cc-10,15,22,39;mh*;bb*;xsteps w; etc.), the re-
sulting consensus tree will depict the Oestridae as the
sister group to the remaining Tachinidae family-
group and the Helicoboscinae as the sister group of a
clade Sarcophagidae + non-helicoboscine Callipho-
ridae (fig. 10). Scoring the Rhinophoridae for the al-
ternative state; namely dorsolateral processes conflu-
ent with phallic wall (0 in character 32 of table 2), has
no impact on the configuration of the resulting
Nelson consensus tree.
The clade Calliphoridae + Oestridae is corroborat-
ed by three character states with the possible addition
of a fourth. These states are presence of setae on well-
80
defined thoracic surfaces, which have been used wide-
ly in the diagnosis of the Calliphoridae. They have
probably been overlooked or neglected in the
Oestridae because of the long, dense hairs, which
tend to conceal the thoracic sclerites.
Only one character state, the fusion of phallic dor-
solateral processes, corroborates the monophyly of
the clade Tachinidae + Sarcophagidae. This fusion
complicates the assessment of at which level in the
cladogram the free-tipped dorsolateral processes are
apomorphic. I have argued that the dorsomedian fu-
sion of these phallic sclerotisations makes a scoring
for presence/absence of free tips inapplicable. With
the condition in the Rhinophoridae being uncertain,
this will imply that the possibility of free tips being a
groundplan state of the Tachinidae family-group can-
not be excluded. In the analysis, this would be equal
to omitting this character (outgroup = 0.1;cc-
10,15,22,39;cc]32;mh*;bb*;xsteps w; etc.). This re-
sults in the Sarcophagidae being the sister group to
the remaining Tachinidae family-group and the Cal-
liphoridae become polyphyletic as the Helicoboscinae
move to a sister group position of Oestridae (fig. 11).
The monophyly of the clade Rhinophoridae +
Calliphoridae + Oestridae is corroborated by a single
character state: presence of a slightly swollen subscu-
tellum. This is indeed weak evidence as the present
division of subscutellar shape in three states may be
too simplified. Differences between the various types,
however, are not easy to put on formal descriptive
terms. If the slightly swollen subscutellum seen in the
Rhinophoridae, the botfly subfamilies, and some of
the calliphorid subfamilies are considered as derived
independently in each terminal taxon, this character
will have no effect on interfamilial phylogeny and
could as well be excluded from the matrix. This exclu-
sion (outgroup = 0.1;cc] 10;cc-15,22,39;mh*;bb*;-
xsteps w; etc.) results in a cladogram identical to that
of fig. 10, with the Oestridae being the sister group of
the remaining Tachinidae family-group. This will
happen irrespective of how the Rhinophoridae are
scored with regard to free or confluent tips of dorso-
lateral processes.
Even the ordering of the subscutellar states is open
to discussion. As already discussed, I prefer to avoid
ordering these states as I have no reason to consider
the swollen condition of the Tachinidae to have aris-
en from a less swollen condition rather than from the
concave condition. If, however, these states are put in
a linear transformation series with increasing convex-
ity (i.e. 0-1-2), and if all characters but 15, 22 and 39
still are treated as ordered (outgroup = 0.1;cc-
15,22,39;mh*;bb*;xsteps w; etc.), the resulting clado-
gram will again put the Oestridae as the sister group
to the remaining Tachinidae family-group (fig. 10).
These examples are excellent illustrations of the
importance of character and transformation series
interpretation.
It should be obvious from the discussion above that
inter-familial phylogenetic relationships within the
Tachinidae family-group still are open to consider-
able discussion. While the monophyly of the non-cal-
liphorid tachinoids are becoming better and better
corroborated we are still short of a satisfactory defini-
tion of the Calliphoridae. However, the main prob-
lem relates to the position of the subfamily
Helicoboscinae (currently with the single genus
Eurychaeta), as the large size of the metathoracic spir-
acle (or its fringe) provides a character state consist-
ingly uniting all non-helicoboscine blowflies. It
should also be emphasized that the present study is
insufficient in the sense that several characters have
been excluded from the analyses because of uncertain-
ty with regard to their proper scoring. Presence in
Eurychaeta of intrapostocular setulae and divided fe-
male abdominal tergites 7-8 (characters b, h) may
provide important additional support for a mono-
phyletic Calliphoridae when included.
What should be given high priority in future stud-
ies is a better definition of the Calliphoridae.
Important information may be obtained through bet-
ter definitions of subfamilies as well as through pat-
terns of subfamilial relationships. Also the
Rhinophoridae need a closer examination. Their def-
inition relies heavily on larval characters, which are
known for less than half the included genera, and our
present ignorance of generic phylogenetic relation-
ships for genera like Bezzimyia and Malayia makes
groundplan estimates of certain characters ambiguous
within this family.
Inclusion of additional outgroups will be a proper
test of the hypothetical groundplan of the Tachinidae
family-group outlined in this study. Thus, studies on
the phylogenetic relationships of the non-tachinoid
calyptrate families are needed, and if McAlpine's
(1989) hypothesis of a monophyletic Muscoidea
finds additional support, outgroup information
should be obtained from the Hippoboscidae family-
group and/or from the Acalyptratae.
ACKNOWLEDGEMENTS
I extend my most sincerest thanks to Knut Rognes,
Stavanger, who spent much time reading and
critizising previous drafts of the manuscript. His most
valuable comments and suggestions are highly appre-
ciated, and although the final decisions and con-
clusions regarding interpretations of characters and
character states are entirely mine, there is no doubt
that the paper has improved considerably through his
effort. Monty Wood, Ottawa, and Stig Andersen,
Pape: Phylogeny of the Tachinidae family-group
Copenhagen, contributed with significant infor-
mation on various aspects of tachinid phylogeny and
morphology, and Verner Michelsen, Copenhagen,
gave important information on the Anthomyiidae
and Muscidae. Amnon Freidberg, Tel Aviv, kindly
arranged a loan of the undescribed species of Mexican
Bezzimyia, and Norman E. Woodley, Washington,
D.C. and Beverly A. Holloway, Auckland, generous-
ly provided some specimens of Mesembrinellinae and
Mystacinobia zelandica respectively for examination.
Thanks also to Brian M. Wiegmann, Maryland
University, for taking time to discuss the use of pa-
rasitism in phylogenetic analyses.
The present study was supported in part through
grant No. 11-6152 from the Danish Natural Science
Research Council.
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TILA M. PÉREZ
Laboratorio de Acaralogia, Universidad Nacional Autónoma de México
FEATHER MITES (ACARINA, XOLALGIDAE) IN THE
UROPYGIAL GLAND TUFT OF ARATINGA
HOLOGHLEORAK(SCLATER AVES, ESIMTACIDAE)
Pérez, T. M. Feather mites (Acaria, Xolgidae) in the uropygial gland tuft of Aratinga holochlo-
ra (Sclater) (Aves, Psittacidae). — Tijdschrift voor Entomologie, 135: 87-90, figs. 1-6. [Issn
0040-7496]. Published 15 July 1992.
The microhabitats for all stages of a new species of feather mite near Fainalges longissimus Mejia-
Gonzalez and Pérez, 1988 (Acarina, Analgoidea, Xolalgidae) on Aratinga holochlora (Sclater,
1859) (Aves, Psittacidae) are reported. Eggs are laid on the barbs of the eight modified down
feathers of the uropygial gland tuft, moulting takes place within the quills of these feathers, and
active instars inhabit the downy barbs of the tail coverts. The uropygial gland tuft for the host
bird is briefly described.
Correspondence: Dra. Tila M. Pérez, Laboratorio de Acarologia, Departamento de Biologia,
Facultad de Ciencias, Universidad Nacional Autönoma de México, 04510 México, D.F.,
México.
Key words. — Feather mite, Xolalgidae, parrot, Psittacidae, uropygial gland tuft.
Ontogenetic studies of astigmatid mites are limit-
ed. To add to the meager information on this subject,
I am associating all instars of seven species of the
feather mite genus Fainalges Gaud & Berla, 1964
(Xolalgidae) from field collected specimens of the
green conure, Aratinga holochlora (Sclater), taken in
Tamaulipas, Mexico.
The most demanding part of this and similar onto-
genetic studies is to unequivocally associate feather
mite larvae with their eggs. To do this, it is necessary
to dissect fully developed larvae from their eggs for
microscopic examination. In the Fainalges investiga-
tion, the larvae of six species had been identified and
associated with their eggs, but the egg of a seventh lar-
va, a new species near Fainalges longissimus Mejia-
Gonzälez & Perez (1988) remained undetected until
a small group of specialized feathers were examined,
namely, the uropygial gland tuft. Herein, this special-
ized microhabitat and their acarine occupants will be
described
MATERIALS AND METHODS
Uropygial gland papillae were taken from two
frozen, field collected Aratinga holochlora. The tuft
feathers were excised, the quills split lengthwise and
the quill contents either mounted on SEM stubs or
microslides. To obtain mature larvae for identifi-
cation, eggs were taken from the barbs and opened,
either on an SEM stub or in a small drop of Hoyers
mounting medium. Thus, correlation of eggs and
larvae were established. Additional specimens for on-
togenetic studies were obtained from samples taken
from approximately 30 museum study skins.
Dissections and measurements of the feathers were
done with a dissecting microscope equipped with an
ocular micrometer. Identification of the mite life
stages was with a Wild-Heerbrugg M-20 phase con-
trast microscope and specimens for electron scanning
microscopy were gold-coated and examined with an
Amray 1000 SEM.
Uropygial gland papilla and associated feathers
The uropygial gland (oil gland, preen gland, rump
gland) is positioned dorsomedially in the tail region
of a bird. When present and functional, the bilobed
gland is usually indicated by an elevation of the skin,
the uropygial eminence, from which a terminal papil-
la arises; the papilla usually contains two ducts ope-
ning to the exterior at its tip. The papilla is variously
shaped and may have a cluster of modified down or
modified semiplumes, termed the uropygial gland
tuft or circulus uropygialis, surrounding the openings
of the uropygial gland ducts; the tuft feathers are in
different numbers and arrangements according to
group (Lucas & Stettenheim 1972, Jacob & Ziswiler
1982: table 1, Johnston 1988). Tuft feathers have un-
usually large quills and relatively short barbs when
compared to typical down feathers.
In Aratinga holochlora the papilla is cylindrical (3.6
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mm in length, 2.2 mm in diameter, N=1) and the
uropygial gland tuft (barbs only, 7.9-9.4 mm, N=2) is
arranged as a single oval of eight modified down
feathers around the duct orifices. The tuft is saturated
with uropygial gland secretions, but when degreased
it resembles an airgun dart as all diverging barbs end
at the same level (Type I of Johnston 1988). Each
uropygial gland feather lacks a rachis and hyporachis
(aftershaft), and the barbs arise directly from the
upper rim of the calamus at the skin level (figs. 1, 2).
The calamus (approximately 3.0 mm, N=12), com-
pletely embedded in the skin, is a tubular structure
(fig. 1) filled with pulp caps. The eight quills form a
rosette around the apex of the papilla. The distal end
of the quill may represent the superior umbilicus, and
as mites can enter, the umbilicus must be open. By
being open, each quill probably contains uropygial
gland secretions (sebum) and can provide a refuge for
organisms unaffected by these secretions.
Fainalges and the uropygial gland tuft
When studying the biology of the seventh
Fainalges species mentioned above, all active instars
were observed on the down barbs of the tail coverts,
but neither eggs nor exuviae were noted. In previous
studies of the feather mites of parrots, it had been de-
termined that for some mite species, different instars
occur in different microhabitats (e.g. Perez & Atyeo
1984, Atyeo & Perez 1988). Therefore, the feathers
above and below the tail coverts were examined, but
again, neither eggs nor exuviae were discovered. The
only feathers never examined were those of the uro-
pygial gland tuft, a microhabitat that was never consi-
dered a viable candidate for mite habitation as it has
been thought that the uropygial gland secretions inhi-
bit parasites (e.g. Jacob & Ziswiler 1982). So, a tuft
was excised and examined under a dissecting micro-
scope. At the bases of the barbs and surrounding pa-
pillary skin, detritus could be seen, but under higher
magnification, the ‘detritus’ consisted of masses of
eggs, egg shells and exuviae. Eggs (fig. 2) and imma-
tures were observed on the barbs. Furthermore, the
eight quills of the tuft, examined through the translu-
cent papillary skin, appeared dark brown, a color con-
dition of quills containing feather mites.
Individual feathers and surrounding papillary tis-
sue were removed and quills split longitudinally (fig.
1). Within each quill there was a compact plug of ma-
terial which, when removed, revealed densely packed
and haphazardly arranged exuviae, pharates and ac-
tive stages for all immature instars partially covered
with uropygial gland secretions (fig. 3).
The density of mites within a quill varied. When
few mites were observed, limited numbers occupied
the more distal interspaces between the pulp caps, up
to 4/5 of the distal quill. Some quills were so packed
88
with mite material that the pulp caps were no longer
evident. These compact masses (plugs) always over-
flowed the quill interior, to form a mound of
detritus’ external to the quill proper. The lengths of
the plugs varied from quill to quill in the same bird.
From 30 museum study skins and eight field col-
lected Aratinga holochlora, the active stages of this
new species have been known for four to five years,
but neither oviposition nor moulting sites had not
been described until now. On a related host species,
the orange-fronted conure, A. canicularis (Linnaeus,
1758), the active instars of Fainalges longissimus were
described from a single microhabitat, the downy
barbs of the tail coverts (Mejia-Gonzälez & Pérez
1988), but oviposition sites were not studied.
From the information obtained from mites of
Aratinga holochlora, it was hypothesized that the uro-
pygial gland tuft is the oviposition and ecdysial site
for Fainalges longissimus and related species on other
parrot taxa. For a preliminary test of the hypothesis, a
tuft was removed from one specimen each of
Aratinga canicularis and a related taxon, the Carolina
parakeet, Conuropsis carolinensis (Linnaeus, 1758),
which has been extinct since the 1920s.
RESULTS
Each quill of the uropygial gland tufts examined
from Aratinga holochlora, A. canicularis and
Conuropsis carolinensis had at least small populations
of Fainalges species in all tuft feather quills and many
barbules had Fainalges eggs cemented to them. Each
host species supported a different species of the
Fainalges longissimus morphotype. This morphotype,
as characterized by Mejia-Gonzälez & Perez (1988),
has all instars with leaflike ventral setae on tarsi I and
II (fig. 6), and in immatures and females, tarsi III and
IV have many spinelike setae, minute ambulacral
discs, and ambulacral stalks longer than the corres-
ponding tibiae (fig. 4). Fainalges longissimus is asso-
ciated with Aratinga canicularis, whereas A. holochlo-
ra and Conuropsis carolinensis each support a new
species of the F. longissimus species complex.
Discussion
The probable scenario for these species of Fainalges
is: females oviposit on bases of the barbs, the barbules
of the tuft feathers (Fig. 2) and on the papillary skin;
the emerging larvae move to the downy barbs of the
tail coverts for feeding; and eventually go to a uropy-
gial gland quill for moulting. Each successive instar
has the same activity, that is, emerging from the quill,
feeding, and then returning to the quill for ecdysis.
Because of the out-of-quill sites for the active instars,
it is assumed that the uropygial gland secretions with-
PÉREZ: Feather mites of Aratinga holochlora
esky ToeU 120 10260 MCZ
Figs. 1-6. Scanning electron micrographs. — 1, One uropygial gland feather embedded in papillary tissue cut along longitudi-
nal axis (approximately 1/3 of quill cut off) to show interior of most of the calamus: two left arrows=area of quill from which
plug (fig. 3) was removed (constriction of quill is artifact due to pressure while cutting), right arrow = Fainalages egg; 2, de-
tail of uropygial gland feather with truncated barbs; arrow = Fainalages egg; 3, plug’ of mites and exuviae removed from quill
of fig. 1: arrows=legs III and IV of nymph; 4, enlargement of fig. 3: arrows=same legs in fig. 3; 5, exuviae emphasizing pro-
terosomata; 6, enlargement of exuvial leg I of fig. 5 showing tarsal structures needed for morphotype identification. Scale bars:
figs. 1, 2: 1000 um; fig. 3: 200 Um; figs. 4, 5: 50 um; fig. 6: 10 um.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
in the quill provide little or no nourishment, and also
that the secretions have no deleterious effect(s) on the
mites or their eggs.
Three questions need to be addressed: Is there a re-
lationship between the moult of the uropygial gland
tuft feathers and populations of feather mites? Does
preening reduce the mite populations of these mites?
Do undiscovered species of the Fainalges longissimus
morphotype have the same modus operandi vis-a-vis
oviposition and moulting sites?
The literature does not specifically answer the
question of whether or not the old gland feathers
moult. Even so, it is assumed that these feathers do
moult as populations of mites within the quills vary,
which also suggests that these feathers moult at differ-
ent times.
The previously reported microhabitats of feather
mites on the external feather surfaces relate to the
channels created by adjacent barbs, usually on the
ventral feather surfaces (e.g., Dubinin 1951, Atyeo &
Pérez 1988). Thus, these mites live in a three-dimen-
sional space bounded by ramal walls laterally and bar-
bules dorsally or ventrally; this space affords protec-
tion from the bills of preening birds.
Depending on the type of uropygial papilla, sebum
for preening is obtained by various methods, ‘either
directly from the drops passing out of the [gland] ori-
fices or by drawing away the fluid from the tuft circ-
let feathers’ (Jacob & Ziswiler 1982: 252). As regards
the uropygial gland tuft in Aratinga species, these par-
rots probably ’… take the sebum by brushing its beak
along the tuft whenever it needs it’ (Jacob & Ziswiler
1982: 252). With this suggested type of preening, it is
doubtful that many mite eggs or active instars would
be dislodged or destroyed.
Do species related to Fainalges longissimus oviposit
and ecdyse in the uropygial gland feather quills? The
genus Fainalges has never been revised, however, dis-
tinct morphotypes are known (Mejia-Gonzalez &
Pérez 1988). The F. longissimus type, restricted to
New World parrots, was characterized as having all
instars with leaflike ventral setae of tarsi I and II (fig.
6), and in immatures and females, pretarsi III and IV
(fig. 4) with many spinelike setae, minute ambulacral
discs, and ambulacra longer than the corresponding
tarsi (Mejia-Gonzälez & Pérez 1988). The F. longissi-
mus morphotype is now known from three species of
90
the Aratinginae (sensu Wolters 1975), therefore, it is
probable that this morphotype will be found on oth-
er New World Aratinginae, and possibly on other
parrots with tufted uropygial gland feathers.
ACKNOWLEDGEMENTS
The research, supported in part by the National
Science Foundation (BSR 89-08301), was conducted
in the Department of Invertebrates, Museum of
Comparative Zoology, Harvard University. I am in-
debted to Warren T. Atyeo, University of Georgia,
for reviewing this manuscript, and Mrs. Robin Pinto,
SEM Laboratory, Museum of Comparative Zoology,
for her assistance in taking the micrographs for this
publication.
REFERENCES
Atyeo, W. T. & T. M. Pérez, 1988. Species in the genus
Rhytidelasma Gaud (Acarina: Pterolichidae) from the
Green Conure, Aratinga holochlora (Sclater) (Aves:
Psittacidae). — Systematic Parasitology 11: 85-96.
Dubinin, V. B., 1951. Feather mites (Analgesoidea). Part I.
Introduction to their study. — Fauna USSR,
Paukoobraznye 6 (5): 1-363. [In Russian].
Jacob, J. & V. Ziswiler, 1982. The uropygial gland. — In
Farner, D. S., J. R. King & K. C. Parkes (eds.), Avian
Biology 6: 199-324. Academic Press, New York.
Johnston, D. W., 1988. A morphological atlas of the avian
uropygial gland. — Bulletin of the British Museum of
Natural History (Zoology) 54: 199-259.
Lucas, A. M. & P. R. Stettenheim, 1972. Avian Anatomy.
Integument. Part I and II. — U.S. Department of
Agriculture Handbook, Washington, D. C., 721 p.
Mejia-Gonzälez, E. & T. M. Pérez, 1988. Three new species
of Fainalges Gaud and Berla (Analgoidea : Xolalgidae)
with descriptions of their developmental series. —
Acarologia 29: 73-86.
Pérez, T. M. & W. T. Atyeo, 1984. Site selection of the
feather and quill mites of mexican parrots. — In D. A.
Griffiths & C. E. Bowman (eds.), Acarology 6 (1): 563-
570. Ellis Horwood Ltd., Chichester, England.
Wolters, H. E. 1975 (1982). Die Vogelarten der Erde. —
Paul Parey, Berlin, 745 p. (Lieferung 1, pp. 1-80, orginal-
ly published September, 1975, included the Psittacidae).
Received: 12 July 1991
Accepted: 1 October 1991
HARRY SMIT
Alkmaar, The Netherlands
WATER MITES FROM NEW SOUTH WALES AND
QUEENSLAND, AUSTRALIA
(ACARI, HYDRACHNELLAE)
Smit, H., 1992. Water mites from New South Wales and Queensland, Australia (Acari,
Hydrachnellae). — Tijdschrift voor Entomologie 135(1): 91-112, figs. 1-74. [ISSN 0040-7496].
Published 15 July 1992.
One new subgenus of the genus Unionicola, viz., Australionicola, and ten new species, viz.,
Hydrachna palustris, Aspidiobates unidorsalis, Kallimobates cooki, Unionicola hammeni,
Unionicola davidsi, Recifella pectinatus, Koenikea purpurea, Albia brokenensis, Arrenurus roobee-
ki, Arrenurus vanderpalae are described from New South Wales and Queensland, Australia.
Limnesia trituberculata is considered a junior synonym of Physolimnesia australis. In addition,
descriptions are provided of the opposite sex of various species known from one sex only. An
annotated list is given for all species recorded during this study.
H. Smit, Emmastraat 43-a, 1814 DM Alkmaar, The Netherlands
Key words. — Water mites, Australia, new species.
Cook (1986) is the most comprehensive work on
Australian water mites published so far. Of 263 spe-
cies treated by him, 203 were described as new. Since
then various studies of Australian water mites were
published by Harvey (1987, 1988a-c, 1989a-b,
1990a-d), Harvey & Cook (1988), and Smith &
Harvey (1989), all dealing with a limited number of
genera or families.
During a trip to New South Wales and Queensland
in the austral winter of 1989, the present author was
able to collect water mites at 46 locations, including
lentic as well as lotic habitats. The material was col-
lected by using a dip net and preserved in Koenike's
solution. All material, including the holotypes, is de-
posited in the collection of the Institute of Taxonomic
Zoology of the University of Amsterdam (Zoological
Museum). Localities are summarized in appendix 1.
SYSTEMATIC PART
The terminology of Cook (1974) is followed for
the description of the species. For a number of species
(especially the genera Flabellifrontipoda, Frontipoda
and Oxus), without external sexual characters, the sex
has not been determined when not necessary for
identification.
Hydrachna Müller
Hydrachna palustris sp. n.
(figs. 1-5)
Type material. — Holotype male: Hasties Swamp,
Queensland (loc. 28). — Paratype: Hasties Swamp,
Queensland, 1 ® (loc. 28).
Description
Male: Body 1800 u in length and 1704 u in width.
Integument papillate. Dorsum with one large plate,
the anterior portions extending laterally. Pre- and
post-ocularia and one pair of glandularia are incorpo-
rated in dorsal plate; eyes outside dorsal plate.
Posteromedial portions of fourth coxae long and nar-
row, apodemes of fourth coxae short. Dorsal lengths
of palp segments: PI 204 u, PIT 173 u, PIII 238 u,
PIV 82 u, PV 41 u.
Female: Body 1824 u in length and 1584 u in
width. Integument papillate. Dorsum with two large,
broad plates, extending anteriorly of eyes; posterior
portion of dorsal plates triangular. Pre-ocularia and
one pair of glandularia incorporated in dorsal plate.
Posteromedial portions of fourth coxae long and nar-
row, apodemes of fourth coxae short. Dorsal lengths
of last four palp segments: PII 184 u, PIII 252 u, PIV
84 u, PV 48 u; PIII stockier than that of male.
Etymology. — The species is named after its occur-
rence in a swamp.
Discussion. — The long and narrow posteromedial
portion of the fourth coxae and the form of the dorsal
plates are diagnostic. As stated by several authors (e.g.
Cook 1974), subgeneric classification of the genus
Hydrachna is somewhat arbitrary. The H. palustris
male could be assigned to the subgenus Hydrachna,
but the female shows the characters of Diplo-
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-5. Hydrachna palus-
tris sp. n. — 1, dorsal plate
6; 2, ventral view d; 3,
palp 6; 4, ventral view ®;
5, dorsal plates © (figs. 1-3
holotype).
hydrachna. For this reason no subgeneric assignment
is given here for H. palustris.
Hydrachna (Rhabdohydrachna) approximata Halik,
1940
The species has been reported before from West
Australia and Victoria (Halik 1941).
Material examined. — New South Wales: 1 d, loc. 43.
Hydrachna sp.
Lundblad (1941) described Hydrachna lateriscuta,
but later (1948) placed this species into synonymy of
Hydrachna bilobata Halik. Cook (1986) found his
specimens of H. bilobata to be in good agreement
with Halik's type material, but discovered differences
between the material described by Lundblad (1948)
as H. bilobata and his own (and thus Halik's) materi-
al of that species. Cook (op. cit.) thus suggested that
H. lateriscuta Lundblad is a distinct species. Three fe-
male Hydrachna specimens collected in the present
study agree with the description of the female of H.
bilobata presented by Lundblad (1948) and Cook
(1986) and tentatively considered as H. lateriscuta by
Cook. As no males were found, the three females are
left unnamed until the taxonomy of the species con-
cerned is better understood. The three specimens are
larger than the female collected by Cook. Body length
varies from 2280 to 2424 u. Measurements of dorsal
plates (length/width) are also larger, 816/1080 u;
768/1008 u and 840/1008 u respectively.
Material examined. — Queensland: 1 9, loc. 19; 1 9, loc.
DD EP locw2 8.
92
Eylais Latreille
Australian Zylais species were described by
Rainbow (1906), Lundblad (1948), Szalay (1953)
and Viets (1980). As the variation in morphological
characters is rather extensive, taxonomical problems
within the genus Eylais remain. Lundblad (1948)
stated that (in translation) ‘the Australian Eylais-spe-
cies seem to vary highly like everywhere in the world,
and it is difficult to find two specimens which are
completely alike’. Nothing is known about the va-
riation of the Australian species, and it seems to me
that a number of species are based on highly variable
characters, e.g. the eye-bridge. A revision is required
and therefore no effort is made here to describe the
material taken during this study.
Material examined. — New South Wales: 1, loc. 1; 1, loc.
10; Queensland: 7, loc. 39; 15, loc. 37; 1, loc. 36.
Hydryphantes Koch
Hydryphantes (Polyhydryphantes) haliki Cook,
1986
Cook (1986) described this species from a pond in
Queensland.
Material examined. — New South Wales: 1 ®, loc. 46.
Pseudohydryphantes Viets
Pseudohydryphantes aroona Harvey, 1988
(figs. 6-10)
The species is known from the holotype only, a
male collected in a stream in Victoria. The specimen
in this study is a female, and a description is given be-
low.
Female: Body 902 u in length and 630 u in width,
lateral side of body undulating. Integument papillate.
Dorsally with 6 pairs of dorsoglandularia and 5 pairs
of lateroglandularia; sclerites associated with glandu-
laria crescent shaped. Lateral eyes on ocular tubercles.
Median eye between dorsoglandularia 2 on a small
sclerite. Chelicera of normal size, each with a number
of short teeth. Ventrally with 5 pairs of ventroglandu-
laria. Coxae in four groups, tips of first, second and
third with thickened setae. Three pairs of acetabula,
two anterior pairs elongate and third pair of acetabu-
la rounded. Genital flaps with long setae on mesal
edge; genital field 184 u in length. Dorsal lengths of
palp segments: PI 46 u, PIT 91 u, PHI 62 u, PIV 139
u, PV 34 u; PIII with convex dorsal and ventral mar-
gins. Dorsal length of leg segments: I-leg-1 60 u, I-
leg-2 79 u, I-leg-3 89 u, I-leg-4 127 u, I-leg-5 151 u,
I-leg-6 156 u; IV-leg-1 120 u, IV-leg-2 98 u, IV-leg-
3 130 u, IV-leg-4 211, IV-leg-5 214 u, IV-leg-6 175
U. Legs III and IV with swimming setae. Claws with-
out serrations, but with a short dorsal tooth.
Discussion. — The enlarged sclerotisized rings sur-
rounding the median eye, preocularia and postocular-
ia, the crescent shaped dorsoglandularia sclerites and
the position of the median eye between dorsoglandu-
laria 2 are diagnostic for the species. The palp of the
female is stockier compared with the male.
Material examined. — Queensland: 1 9, loc. 33
Diplodontus Duges
Diplodontus haliki Lundblad, 1948
This species probably has a widespread distribution
in Australia; it has been reported from Tasmania,
Victoria and New South Wales (Lundblad 1948,
Cook 1986).
Material examined. — New South Wales: 1 ®, loc. 43.
Mamersa Koenike
Mamersa corndorl Harvey, 1988
(figs. 11-13)
Previously reported from the Northern Territory.
Material examined. — Queensland: 1 d, 1 @, loc. 28.
Hydrodroma Koch
Hydrodroma spec A Cook, 1986
Cook (1986) suggested that H. despiciens could be
SMIT: Water mites from Australia
divided into morphologically similar but distinct spe-
cies. I have compared the Australian material with
European specimens, and it is indeed difficult to find
any marked differences. As with Cook, the species
was found in ponds in Queensland.
Material examined. — Queensland: 1, loc. 17; 3, loc. 21;
3, loc. 25; 18, loc. 32; 1, loc. 36; 3, loc. 39; 1, loc. 46.
Hydrodroma spec B Cook, 1986
This species is characterized by the reduced num-
ber of swimming setae. This species occurred only in
lotic habitats, but Cook (1986) also found it in lentic
habitats.
Material examined. — Queensland: 3, loc. 18; 1, loc. 23;
5, loc. 33; 2, loc. 35.
Flabellifrontipoda Lundblad
Flabellifrontipoda pectinata Lundblad, 1948
This species has a widespread distribution in
Australia; it is reported from Tasmania to
Queensland.
Material examined. — Queensland: 4, loc. 18.
Flabellifrontipoda carteza Cook, 1986
The species is known from the holotype only, a fe-
male collected in a stream in Queensland. The speci-
men collected in this study is also a female. It is larg-
er than the holotype, 776 u in length and 630 in
height. Dorsal lengths of palp segments: PII 57 u,
PIII 60 u, PIV 84 u, PV 14 u.
Material examined. — Queensland: 1 9, loc. 18.
Frontipoda Koenike
Frontipoda spinosa Viets, 1977
Viets (1977) described this species from Lake
Eacham. It was recollected there, and also from the
nearby Lake Barrine.
Material examined. — Queensland: 1, loc. 29; 2, loc. 31.
Frontipoda tasmanica Viets, 1978
A species previously reported from Tasmania and
New South Wales.
Material examined. — Queensland: 4, loc. 16.
Oxus Kramer
Oxus meridianus (Lundblad, 1948)
Previously reported from Tasmania.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 6-10. Pseudohydryphantes aroona Harvey. — 6, dorsal view 9; 7, ventral view 9; 8, palp 9; 9, first leg 2; 10, lateral view
capitulum and chelicera 2. — Figs. 11-13. Mamersa corndorl Harvey.— 11, dorsal view 9; 12, ventral view 2; 13, palp ©.
94
Material examined. — New South Wales: 1, loc. 1; 4, loc.
6; 4, loc. 10. Queensland: 1, loc. 18; 2, loc. 15.
Oxus orientalis Walter, 1915
With the exception of being larger, the specimens
collected agree with the description by Cook (1986).
Males vary from 795-863 u (Cook: 684 u), females
from 863-1392 u (Cook: 851-972 u). The male from
New South Wales is 688 u in length. The specimens
of Walter (1915) measure 1500 and 1050 u. Cook
(op. cit.) collected only one male and two females, ap-
parently small specimens. Males and females differ in
length of the long seta on tip of IV-leg (up to 120 u
in length), and in the form of the coxae near the gen-
ital field.
The species was described from New Caledonia.
Material examined. — New South Wales: 1 d, loc. 10.
Queensland: 5 6, 14 9, loc. 21; 2 2, loc. 25; 1 d, loc. 32;
1 d, loc. 31.
Physolimnesia Halik
Physolimnesia australis Halik, 1940
Limnesia trituberculata Viets, 1955. Syn. n.
Cook (1986) could not distinguish females of
Physolimnesia australis and Limnesia trituberculata
Viets. I compared the females of P. australis with the
illustrations of Viets and concluded that the two spe-
cies are identical.
Material examined. — New South Wales: 17 4, 50 9,
loc. 3; 1 8,7 9, loc. 6. Queensland: 3 @, loc. 29;4 5,8 9,
loc. 34; 3 3,6 9, loc. 15.
Limnesia Koch
Limnesia longigenitalis Lundblad, 1941
A species previously reported from Victoria.
Material examined. — New South Wales: 2 4,5 9, loc. 1;
1 ®, loc. 46.
Limnesia dentifera Viets, 1980
(figs. 14-15)
Viets (1980) described only the female of this spe-
cies from the Myall Lakes area, New South Wales; a
description of the male is given below.
Male: Dorsum not sclerotisized, with only one pair
of small platelets. Dorsum 679 u long and 495 u
wide. Capitular bay V-shaped. Genital field triangu-
lar, 213 u in length and 136 u in width. Glandula
Limnesiae (see Cook 1986, fig. 414) shifted to poste-
rior margin of third coxae. Dorsal lengths of palp seg-
ments: PI 22 u, PII 108 u, PIII 89 u, PIV 151 u, PV
Smit: Water mites from Australia
77 u; PIL medially with one large seta. Peg-like seta
on ventral side of PII not on a tubercle. III-leg-4 and
5, and IV-leg-4 and 5 with 6-7 swimming setae.
Material examined. — New South Wales: 1 4,2 9, loc. 1;
No WAS 1l0c#;310; 3.2, Iinymph, loc. 8:
Limnesia parasolida Viets, 1984
The species is reported from Queensland, where it
is probably widespread. Viets (1984) described it
from the Alice Springs area, Northern Territory.
Material examined. — Queensland: 7 3,5 9, loc. 23; 30
1782103837303 BP loves TUG.
Limnesia solida Lundblad, 1948
(fig. 16)
Lundblad (1948) described the female from
Victoria. The male was hitherto unknown; a descrip-
tion is given below. Viets (1975) reports the species
without location, but probably in Queensland.
Male: Body soft, 650 u in length and 524 u in
width. Dorsum without platelets. Coxae in three
groups. The Glandula Limnesiae shifted to posterior
margin of third coxae. Three pairs of genital acetabu-
la, the first anterior pair noticeably separated from
second pair. Gonopore relatively small, 78 u in
length. Dorsal lengths of palp segments: PI 24 u, PII
91 u, PI 70 u, PIV 131 u, PV 36 u. Palp as illustrat-
ed for the female by Cook (1986). Dorsal lengths of
distal three segments of fourth leg: IV-leg-4 178 u,
IV-leg-5 178 u, IV-leg-6 156 u; IV-leg-5 with 1 and
IV-leg-4 with 3 swimming setae.
Material examined. — Queensland: 1 2, loc. 33; 2 2, loc.
ZIA oc 2262010085;
Limnesia lembangensis Piersig, 1906
A species with a wide distribution, known from
India through to Australia. Gledhill (pers. comm.)
has seen material from Sri Lanka. Halik (1941) re-
ported it from New South Wales.
Material examined. — Queensland: 7 &, 41 ©, loc. 21; 2
Doc 222
Hygrobates Koch
Hygrobates hamatus Viets, 1935
A widespread species. Hygrobates hamatus hamatus
is known from Indonesia and Queensland, the sub-
species H. hamatus bharatensis is recorded from India
(Cook 1967).
Material examined. — Queensland: 1 d, 3 ©, loc. 23; 1
Ds loes Biles
95
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 14 - 15. Limnesia dentifera Viets. — 14, ventral view d; 15, palp d. - Fig. 16. Limnesia solida Lundblad. — 16, ventral
view d.
Australiobates Lundblad
Australiobates mutatus Viets, 1978
Based on the very long palp segments one speci-
men is assigned to A.mutatus, the most common of
the Australian Australiobates species and known from
Tasmania to Queensland.
Material examined. — Queensland: 1 9, loc. 27.
Australiobates linderi Lundblad, 1941
(figs. 17-19)
Specimens from three locations are provisionally
assigned to A.linderi. Specimens from loc. 7, loc. 42
and some males from loc. 40, however, lack setae on
the ventral side of PII. This segment has a ventral
margin which is almost straight, slightly convex or
convex, similar to fig. 33B of Lundblad (1948). Palp
segments and leg segments are rather stocky, like A.
emalus Cook, which is known from one male only.
However, the ventral margin of PIII in A. emalus is
straight. The peg-like setae of PIV are bluntly and/or
sharply pointed (this character being variable within a
single specimen), so this is not a good character.
Some measurements are provided here of specimens
from loc. 7.
Male: Dorsal lengths of palp segments: PII 82 u,
PIII 120 u, PIV 108 u, PV 36 u. Dorsal lengths of
first leg: I-leg-4 182-209 u, I-leg-5 187-211 u, I-leg-
6 158-177 u.
96
Female: Dorsal lengths of palp segments: PI 43-46
u, PIT 106-108 u, PIII 149-151 u, PIV 134-139 u,
PV 43-48 u. Dorsal lengths of first leg: I-leg-4 272, I-
leg-5 281 and I-leg-6 213 u.
Material examined. — New South Wales: 4 d, 4 @, loc. 7.
Queensland: 6 d, 6 9, loc. 40; 2 4,28 9, loc. 42.
Coaustraliobates Cook
Coaustraliobates minor (Lundblad, 1948)
(fig. 20)
In addition to material from lotic habitats I also
found this species at lentic sites. Cook (1986), howev-
er, only collected this species from streams.
Specimens from Lake Eacham, Queensland have a
rather long anterior acetabulum and more setae on
the anterior margin of the genital field (see fig. 20).
Material examined. — New South Wales: 3 2, loc. 2; 1 6,
loc. 3; 2 ©, loc. 4; 1 6, loc. 6; 3 3b, 22 9, loc. 9; 1! 6, loc.
10. Queensland: 3 9, loc. 23; 8 d, 5 2, 1 nymph, loc. 30;
Doll Bo lores Bile i Le Mere, Sieh
Aspidiobates Lundblad
Aspidiobates geometricus Cook, 1986
Previously reported from Victoria, New South
Wales and Queensland.
Material examined. — Queensland: 1 2, loc. 20.
SMIT: Water mites from Australia
Figs. 17-19. Australiobates linderi Lundblad. — 17, palp 3; 18, palp 2; 19, palp 9. - Fig. 20. Coaustraliobates minor
(Lundblad), genital field 8. — Figs. 21-23. Aspidtobates unidorsalis sp..n. — 21, ventral view d , 22, dorsal view d ; 23, palp d
(figs. 21-23 holotype). — Figs. 24-26. Kallimobates cooki sp. n. — 24, ventral view 9; 25, palp 2; 26, lateral view capitulum
and chelicera 9 (figs. 24-26 holotype).
97
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 27-37. Unionicola. — Figs. 27-34. Unionicola hammeni sp. n. — 27, ventral view d ; 28, palp 3; 29, dorsal view palp d;
30, 4-leg-6 4 ; 31, 1-leg-6 d; 32, ventral view 9; 33, palp 2; 34, 4-leg-6 ® (figs. 27-31 holotype). — Fig. 35. Unionicola fla-
belliseta Cook. — 35, ventral view 2. — Figs. 36-37. Unionicola davidsi sp. n. — 36, ventral view 95,37, palpr 2.
98
Aspidiobates scutatus Lundblad, 1941
Previously reported from Tasmania, Victoria and
New South Wales.
Material examined. — Queensland: 1 à, loc. 11; 1 d, 2
®, loc. 13.
Aspidiobates unidorsalis sp. n.
(figs. 21-23)
Type material. — Holotype male: Darraboola Creek,
Lamington NP, Queensland (loc.13).
Description
Male: Dorsal and ventral shields present. Dorsal
shield complete, 591 u in length, 543 u in width and
with six pairs of glandularia each lying on a small
platelet, together with the associated seta. Genital
field 145 u in length and 78 u in width. Three pairs
of acetabula, anterior pair separated from the others.
Fourth coxae with small posteromedial projections.
Glandularia of the fourth coxae shifted onto the third
coxae. Dorsal lengths of palp segments: PI 26 u, PII
94 u, PHI 77 u, PIV 120 u, PV 38 u.
Female: Unknown
Etymology. — The species is named for its complete
dorsal shield.
Discussion. — This is the fifth known Aspidiobates
species with a complete dorsal shield. Viets (1969)
described two species from New Caledonia and
Hopkins (1975) described one species from New
Zealand. The only known Australian species with a
complete dorsal shield, A. bidewel Harvey, exhibits
sexual dimorphism in IV-leg-4. The form of the dor-
sal shield and the genital field distinguish the new
species easily from the other Australian Aspidiobates
species.
Caenobates Viets
Caenobates acheronius Viets, 1978
The species was previously known from Tasmania,
Victoria and New South Wales.
Material examined. — Queensland: 1 9, loc. 12.
Kallimobates Viets
Kallimobates cooki sp. n.
(figs. 24-26)
Type material. — Holotype female: Creek near Elabama
Falls, Lamington NP, Qld (loc.11). — paratype: Same loca-
tion as holotype, 19.
Description
Female: Dorsal and ventral shields present. Dorsal
Smit: Water mites from Australia
shield 563 u in length and 514 u in width and with
three pairs of glandularia, one pair lying in the dorsal
furrow posterior to dorsal shield. Three pairs of acet-
abula; gonopore 111 u in length and 97 u in width.
Capitular rostrum tapers gradually anteriorly. Dorsal
lengths of palp segments: PI 29 u, PI 89 u, PIII 117
u, PIV 74 u, PV 43 u. PII with a large projection; PI-
II with a smaller projection, which does not extend
distoventrally. Legs without swimming setae.
Male. — Unknown.
Etymology. — The species is named after Dr. D. R.
Cook.
Discussion. — Females of the new species differ
from those of the two known species of the genus
Kallimobates in the form of the palp. The projection
on PIII is intermediate between K. australicus Viets
(almost without a projection on PIII) and K. vietsi
Cook (with a large projection on PIII). The capitu-
lum is similar to that of K. vietsi. PIV is rather short,
as in K. australicus.
Encentridophorus Piersig
Encentridophorus sarasini Walter, 1915
A widespread species, known from New Caledonia
to India; from Australia it has been reported from
Victoria and Queensland.
Material examined. — Queensland: 1 9, loc. 39; 1 d, 13
9 Inymphy loc.,22; 6495 loc. 26; luda 2h ploc 37:2 d,
2319 Moc) 253-3) 2 loc. 36:
Unionicola Haldeman
Australionicola subgen. n.
Diagnosis
Characters of the subfamily Unionicolinae. Genital
field with 5 pairs of acetabula; male with one acet-
abular plate, female with two acetabular plates.
Pronounced sexual dimorphism in male palp and
fourth leg. Fifth palp segment of male relatively long,
with a large claw-like structure. Fifth palp segment of
female relatively long, with 3 short, thickened setae
on posterior margin. Legs with large claws with 2-4
clawlets; fourth leg of male with a small claw and
without clawlets.
Type species. — Unionicola (Australionicola) ham-
meni sp. n.
Discussion. — The sexual dimorphism found in the
male palp is not found in any other subgenus of
Unionicola (see Vidrine 1986, 1987). Also the large
claws with 3-4 clawlets are unique for the subgenus,
although subgenera (e.g. Anodontinatax) with large,
bifid claws do occur.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Unionicola (Australionicola) hammeni sp. n.
(figs. 27-34)
Type material. — Holotype male: Cattle pond Upper
Daintree Road, Daintree, Queensland (loc. 32). —
Paratypes: One male and one female from same location
as holotype.
Description
Male: Body soft, 592 u in length and 359 u in
width; eyes large. Apodemes of anterior coxae exten-
ding to third coxae. Epimeroglandularia 2 small. Two
acetabular plates (which are possibly fused anteriorly,
but this is difficult to determine) each with 5 aceta-
bula, the anterior two acetabula of each side and the
posterior three acetabula arranged one behind the
other; each acetabular plate with two setae. (The slide
of the type specimen is of poor quality, so the illust-
ration of the ventral side (fig. 27) is of the paratype).
Gonopore 107 u in length. Dorsal lengths of palp
segments: PI 17 4, PII 103 p, PIII 60 p, PIV 70 u,
PV 72 u. Fourth palp segment with two small tuber-
cles on posterior margin. Fifth palp segment with a
large, stiff lateral seta and a large claw-like structure.
Dorsal lengths of fourth leg: IV-leg-2 94 u, IV-leg-3
101 u, IV-leg-4 125 u, IV-leg-5 163 u and IV-leg-6
134 u; IV-leg-6 relatively short, with a small claw
without clawlets. All other legs with large claws and
2-3 clawlets. There are two types of long setae
present: broad setae which probably act as swimming
setae (and here referred to as swimming setae) and
long, hair-like setae. I-leg-4 with 4 hair-like setae, I-
leg-5 with 5 swimming setae; II-leg-3 with 1 hair-like
seta, II-leg-4 with 5 swimming setae, II-leg-5 with 3
swimming and 1 hair-like setae; III-leg-3 with 1 hair-
like seta, III-leg-4 with 5 swimming setae and III-leg-
5 with 3 swimming and 2 hair-like setae; [V-leg-4
with 4 hair-like setae and IV-leg-5 with 5 swimming
setae.
Female: Body soft, 747 u in length and 553 u in
width. Anterior coxae with apodemes, but it is not
possible to determine how far they extend posteriorly
due to the poor condition of the specimen.
Epimeroglandularia 2 small. Two pairs of acetabular
plates, lying close two each other; the anterior plates
with two pairs of acetabula and two setae, the posteri-
or plates with three pairs of acetabula (arranged one
behind the other) and each plate with one seta.
Dorsal lengths of palp segments: PI 17 u, PII 89 u,
PIII 38 u, PIV 84 u and PV 60 u; PIV with two small
tubercles, PV with three short setae on dorsal margin.
Dorsal lengths of fourth leg: IV-leg-2 142 u, IV-leg-3
154 u, IV-leg-4 178 u, IV-leg-5 221 u and IV-leg-6
187 u. Legs 1-3 with large claws and 3 clawlets (2
large and one small), leg 4 with 4 clawlets. Similarly
the female has long, hair-like setae and broad swim-
ming setae. I-leg-3 with 1 hair-like seta, I-leg-4 with 5
swimming setae, I-leg-5 with 5 swimming and 1 hair-
100
like setae; II-leg-3 with 1 hair-like setae, II-leg-4 with
6 swimming setae, II-leg-5 with 5 swimming setae;
III-leg-3 with 1 swimming and 1 hair-like seta, III-
leg-4 with 5 swimming and 1 hair-like setae, III-leg-5
with 3 swimming and 1 hair-like setae; IV-leg-5 with
3 swimming and 1 hair-like setae, IV-leg-4 with 2
swimming and | hair-like setae.
Etymology. — The species is named after Mr. Henk
van der Hammen.
Discussion. — The unique structure of the male
palp separates this species from any other member of
the genus. Other diagnostic characters for the male
include the number of clawlets of the claws. The
structure of the palp of the female is also diagnostic.
Unionicola ( Unionicola) flabelliseta Cook, 1986
(fig. 35)
Cook (1986) described only the male. Only fe-
males were found in the present study and a descrip-
tion is given below.
Female: Body soft, 999 u in length and 805 u in
width. Ventrally with a pair of raised glandularia tu-
bercles at posterolateral corners of body. Fourth cox-
ae with small posteromedial projections but with very
long setae. Epimeroglandularia 2 small. Two pairs of
acetabular plates, each plate with three acetabula.
Dorsal lengths of palp segments: PI 17 u, PII 214 u,
PHI 62 u, PIV 178 u, PV 82 u. The female, like the
male, has an expanded fan-shaped peg-like setae on
PIV. Dorsal lengths of distal segments of first leg: I-
leg-4 262 u, I-leg-5 223 u, I-leg-6 194 u. Dorsal
lengths of distal segments of fourth leg: IV-leg-4 252
u, IV-leg-5 369 u, IV-leg-6 349 u.
Discussion. — The fan-shaped peg-like seta of PIV
is diagnostic for the species. Cook described the spe-
cies from a river in Queensland, the females described
above were found in a lake in Queensland.
Material examined. — Queensland: 3 @, loc. 19.
Unionicola (Pentatax) longiseta Walter, 1915
A widespread species previously known from New
Caledonia and Australia (Tasmania and Victoria).
Material examined. — Queensland: 2 9, loc. 19; 1 2, loc.
36.
Unionicola (Pentatax) minutissima Lundblad, 1948
Reported before from Tasmania and Queensland.
Material examined: Queensland: 2 9, loc. 19; 3 2, loc.
18.
Unionicola (Pentatax) davidsi sp. n.
(figs. 36-37)
Type material. — Holotype female: Cattle pond north of
Bundjalung NP, New South Wales (loc. 6).
Description
Female: Body soft, 553 u in length and 398 u in
width. A pair of raised glandularia tubercles at poste-
rolateral corners of body. Tips of second and third
coxae rounded. Five pairs of genital acetabula, these
placed on two pairs of acetabular plates. Posterior pair
of acetabular plates with three pairs of acetabula and
one pair of heavy setae; anterior plates with two pairs
of acetabula and two pairs of heavy setae. Lengths of
palp segments: PI 14 u, PII 103 u, PIII 48 u, PIV
106 u, PV 55 u. Peg-like seta of PIV at extreme dis-
toventral end not on a tubercle; one of the hair-like
setae on PIV on a short tubercle. PII with a long
lateral seta; PII with two short lateral setae. Dorsal
lengths of first leg: I-leg-4 226 u, I-leg-5 218 u, I-leg-
6 151 u. Dorsal lengths of fourth leg: IV-leg-4 262 u,
IV-leg-5 320 u, IV-leg-6 272 u.
Male: Unknown.
Etymology. — The species is named after Dr. C.
Davids.
Discussion. — This species is close to U. minutissi-
ma Lundblad and U. alpa Cook. It differs from both
species in the structure of the palp (PV of U. davidsi
is relatively short and PIV has a different shape), the
larger size and the very long leg segments.
Unionicola (Pentatax) sp.
Two Unionicola-females from location 19 have
shorter palps and shorter leg segments than U. david-
si. They resemble the latter species in most details,
but may belong to a different species. For one of these
female the dorsal lengths of the palp and leg segments
are: PII 84 u, PIII 34 u, PIV 86 u, PV 53 u; I-leg-4
175 u, I-leg-5 168 u, I-leg-6 125 u; IV-leg-4 194 u,
IV-leg-5 261 u, IV-leg-6 240 u.
Unionicola (Giselatax) lundbladi Viets, 1975
Viets (1975) described the species from Queens-
land.
Material examined. — Queensland: 1 9, loc. 36.
Neumania Lebert
Neumania (Neumania) ambigua Piersig, 1906
A widespread species, known from Japan to
Indonesia, Philippines and eastern Siberia.
Material examined. — Queensland: 1 d, loc. 19; 1 d, loc.
DE Slee 2220245020 loc ERI LE
30; 1 6, loc. 32.
Neumania (Neumania) nodosa (Daday, 1898)
N. nodosa is a widespread species, reported from
India, Indonesia, Japan and also from the Northern
Smit: Water mites from Australia
Territory, Australia (Viets 1981).
Measurements and number of acetabula are vari-
able. The Australian specimens are larger than the
specimens taken by Cook (1967) in India. Australian
females measured 960-1248 u, females from India
653-714 u. Females from Indonesia are intermediate
i.e. 800-1000 u (Piersig 1906). There are also differ-
ences in the lengths of the leg and palp segments.
Dorsal lengths of first leg of Australian females: I-leg-
3 257 u; I-leg-4 378 u; I-leg-5 378 u; I-leg-6 262 u;
dorsal lengths of fourth leg: IV-leg-3 281 u; IV-leg-4
310-349 u; IV-leg-5 340-378 u; IV-leg-6 310-349 u;
dorsal lengths of palp segments: PII 137 u; PIII 89 u;
PIV 132 u; PV 50 u. Indian specimens have smaller
leg- and palp segments. Additionally, Australian fe-
males bear 20-22 acetabula, specimens from India
15-18 and specimens from Indonesia 28-34.
Material examined. — Queensland: 1 2,1 d, loc. 21; 1
Proc 22-59 loc 25; 21271028!
Neumania (Lemienia) gila Viets, 1975
Reported before from several localities in
Queensland (Viets 1975).
Material examined. — New South Wales: 5 2, loc. 6.
Queensland: 2 d, 1 9, loc. 17; 8 d, 5 2, loc. 36.
Neumania (Lemienia) falcipes Koenike, 1906
The first record of this species from Australia. The
species is known from Sri Lanka, Indonesia and
Africa.
Material examined. — Queensland: 2 d, loc. 38.
Recifella Viets
Recifella (Recifellida) tinka (Viets, 1975)
(figs. 38-39)
Viets (1975) described the male of this species
from a location in Queensland; the female was hither-
to unknown. At one location both males and females
were found and the female is described below.
Female: Dorsal shield 640 u in length and 553 u in
width. Dorsal shield bearing three pairs of glandular-
ia, the most posterior lying on small tubercles; a
fourth pair of glandularia lying free in the dorsal fur-
row posterior to the dorsal shield; these glandularia
on large tubercles. Glandularia of ventral shield on
small tubercles. Dorsal lengths of palp segments: PI
29 u, PII 98 u, PIII 46 u, PIV 96 u, PV 25 u. Palp
stocky, as illustrated for the male by Viets (1975).
Dorsal lengths of fourth leg: IV-leg-6 194 u, IV-leg-5
209 u,IV-leg-4 158 u, IV-leg-3 122 u, IV-leg-2 110
u; IV-leg-5 and IV-leg-4 each with three swimming
setae.
101
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
44 @ d
Figs. 38-45. Recifella. — Figs. 38-39. Recifella tinka (Viets). — 38, dorsal shield © ; 39, ventral view 2. — Figs. 40-45. Recifella
pectinata sp. n. — 40, dorsal view d ; 41, ventral view d ; 42, palp d ; 43, fourth leg d ; 44, dorsal shield ®; 45, ventral view
9 (figs. 40 - 43 holotype).
Discussion. — See under À. pectinatus sp. n. Reported before from New South Wales and
Material examined. — Queensland: 7 2, 1 à, loc. 30. Queensland.
Recifella (Recifellida) doomba Cook, 1986 Material examined. — Queensland: 1 d, 1 9, loc. 38
102
Recifella (Recifellida) pectinatus sp. n.
(figs. 40-45)
Type material. — Holotype male: Lake at Eungella Dam,
Queensland (loc. 19). — Paratypes: 5 d and 8 9 from same
location as holotype; 2 2, loc. 17.
Description
Male: Dorsal shield fused with ventral shield poste-
riorly thus making the dorsal furrow incomplete.
Dorsal shield 446 u in length and 393 u in width and
with 4 pairs of glandularia, the three most posterior
pairs on tubercles. Tips of first coxae rounded and
projecting beyond the body proper. Genital field with
15-20 acetabula. Dorsal lengths of palp segments: PI
24 u, PIT 96 u, PIII 46 u, PIV 84 u, PV 26 u. Dorsal
heavy terminal seta of PV diverging from the others;
ventral side of PII straight, PII stocky. Dorsal lengths
of fourth leg: IV-leg-2 107 u, IV-leg-3 116 u, IV-leg-
4 145 u, IV-leg-5 242 u, IV-leg-6 184 u. IV-leg-5
bowed, the three unmodified specialized setae in the
proximal half of segment. Proximal two of these setae
closer to each other than the two distally sited setae.
The distal end of this segment with a prominently
pectinate seta. [V-leg-4 and IV-leg-5 with 3 swim-
ming setae; IV-leg-4 with 2 large pectinate setae and
1 small pectinate seta, IV-leg-3 with 1 pectinate seta.
Female: Dorsal and ventral shield present; dorsal
shield 611 u in length and 495 u in width. Dorsal
shield with three pairs of glandularia on small tuber-
cles, a fourth pair of glandularia lying free in the dor-
sal furrow on large tubercles. Tubercles on ventral
shield large, especially on lateral side of body.
Posterior apodemes of anterior coxal groups extend-
ing to middle of third coxae. Acetabular plates with
numerous acetabula. Dorsal lengths of palp segments:
PI 24 u, PIT 96 u, PIII 46, PIV 89 u, PV 29 u; palp
as illustrated for the male. Dorsal lengths of distal
three segments of first and fourth leg: I-leg-4 136 u,
I-leg-5 155 u, I-leg-6 155 u; IV-leg-4 155 u, IV-leg-
5 208 u, IV-leg-6 194 u; Ill-and IV-leg-5, III- and
IV-leg-4 with three swimming setae, III- and IV-leg-
3 with two swimming setae.
Etymology. — The species is named for its large,
pectinate setae.
Discussion. — The fourth leg of the male of this
species resembles that of À. baltoona and R. tinka; it
differs from the first species in the absence of a bulge
on IV-leg-4 and the heavy pectinate seta on the distal
end. À. tinka has the three setae of IV-leg-4 close to
each other and on a small but clear bulge. Males and
females of R. tinka and R.pectinatus have a stocky sec-
ond palp segment; females of the two species differ in
the form of acetabular plates and tubercles.
Smit: Water mites from Australia
Koenikea Wolcott
Koenikea (Koenikea) sorpresa Cook, 1986
Known from a few specimens from Queensland
and New South Wales.
Material examined. — Queensland: 1 2, loc. 19.
Koenikea (Notomideopsis) australica Lundblad,
1941
Reported by Lundblad (1948) from Tasmania and
Victoria.
Discussion. — See K. verrucosa.
Material examined. — New South Wales: 2 8,4 9, loc. 8.
Queensland: 1 9, loc. 16.
Koenikea ( Notomideopsis) verrucosa Lundblad,
1948
Lundblad (1948) described the species from a new-
ly metamorphosed male from Tasmania. This male
differed from K. australica in having much larger
glandularia tubercles. Cook (1986) suggested that
older specimens have smaller tubercles, and therefore
questioned the validity of K. verrucosa. At one loca-
tion in this study males and females were found with
large glandularia tubercles and these were compared
with males and females with small tubercles assigned
to K. australica. It is clear that K. verrucosa should
stand as a distinct species. The glandularia tubercles
of K. verrucosa measure 24-29 U and those of K. aus-
tralica 12-16 U. Further, K. verrucosa has more and
larger papillae on the body than X. australica. The fe-
male of K. verrucosa is briefly characterized below.
Female: Body 630 u in length and 601 u in width;
dorsal shield 563 u in length and 495 u in width.
Dorsal shield with five pairs of large glandularia tu-
bercles. Body with large papillae. Dorsal lengths of
palp segments: PI 24 u, PI 74 u, PHI 48 u, PIV 72
iS ENYA Sho} (Uy
Material examined. — Queensland: 3 6, 2 9, loc. 14.
Koenikea ( Notomideopsis) branacha Cook, 1986
Previously reported from Queensland and New
South Wales.
One of the two females from loc. 18 is much larger
than other specimens found (and also larger than the
measurements given by Cook 1986), but is otherwise
similar. Dorsal shield of this female is 824 u in length
and 616 u in width; dorsal lengths of palp segments:
PI 25 u, PIL 96 u, PHI 67 u, PIV 106 u, PV 46 u;
dorsal lengths of distal three segments of first leg: I-
leg-4 213 u, I-leg-5 189 u, I-leg-6 199 u; ventral
shield is 858 u in length; gonopore 136 u in width. It
is provisionally assigned to K. branacha, but may be-
long to a different species.
103
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 46-47. Koenikea voloma Cook. — 46, dorsal shield 9 ; 47, ventral view 9. — Figs. 48-51. Koenikea purpurea sp. n. — 48,
ventral view d ; 49, dorsal shield 5; 50, palp 4; 51, ventral view ® (figs. 48-50 holotype).
Material examined: Queensland: 2 @, loc. 16; 2 9, loc.
18; 1 9, loc. 40.
Koenikea (Notomideopsis) voloma Cook, 1986
(figs. 46-47)
Previously reported from Tasmania and New
South Wales.
Cook (1986) provided a good description of this
species. Only females were found in the present
study, which, although not entirely fitting the de-
scription of K. voloma by Cook (1986), have all been
assigned to that species. A short description of these
specimens is given below (the measurements given by
Cook for the allotype and paratype are shown in pa-
rentheses).
Female: Dorsal shield 650-718 u ( 684-729) in
length and 504-548 u (562-638) in width. Dorsal
104
shield with 6 pairs of glandularia; glandularia 3, 4 and
5 more or less in a line. Tips of first coxae rounded
and not projecting to anterior end of body.
Epimeroglandularia 2 located near posterior suture
lines of fourth coxae. Genital field close to posterior
of body. Gonopore 116-126 u (120-122) in width;
acetabular region 281-310 u (318-347) in width.
Dorsal lengths of palp segments: PI 26 u (26-27), PII
77-84 u (85-86), PIII 48-53 u (48-51), PIV 77-84 u
(88-90), PV 38 u (40-43); peg-like seta of PIV on a
short tubercle. The location of this seta varies (even
within a specimen), from near the distal end of this
segment to somewhat more proximal. First leg with
long ‘rillborsten’ (fluted or grooved setae).
Discussion. — The females of this study have a
more elliptical dorsal shield compared with the origi-
nal description presented by Cook (1986). The loca-
tion of the peg-like seta of PIV varies, so this charac-
ter does not separate this species from other species,
e.g. K. jacunda.
Material examined. — Queensland: 5 ®, loc. 18; 1 9,
loc.40.
Koenikea (Notomideopsis) purpurea sp. n.
(figs. 48-51)
Type material. — Holotype male: Teewah Creek,
Cooloola NP, Queensland (loc.16). — Paratype: Same loca-
tion as holotype, 1 ©.
Description
Male: Body purple, 669 u in length and 640 u in
width. Dorsal furrow complete; dorsal shield 655 u in
length and 601 u in width and with 6 pairs of glan-
dularia on small tubercles. Tips of first coxae not ex-
tending to anterior end of body. Second coxae each
with one very large, stiff seta. Posterior apodemes of
anterior coxal groups extending posteriorly beyond
suture lines between third and fourth coxae.
Epimeroglandularia 2 lying in an indentation of the
posterior margin of the fourth coxae. Acetabular
plates broad, 116 U in length with one pair of glan-
dularia and 10-15 small acetabula. Gonopore 87 U in
length. Excretory pore terminal. Dorsal lengths of
palp segments: PI 22 u, PII 70 u, PIII 48 u, PIV 70
u, PV 36 u; peg-like seta of PIV not inserted on a tu-
bercle. Dorsal lengths of last three segments of fourth
leg: IV-leg-4 126 u, IV-leg-5 155 u and IV-leg-6 184
u. One swimming seta on II-leg-4, two on I-leg-5, II-
leg-5, III-leg-4, IIl-leg-5, and three on IV-leg-4 and
IV-leg-5. First two legs with ‘rillborsten’.
Female: Body purple, 718 u in length and 679 u in
width. Dorsal furrow complete; dorsal shield 708 u in
length and 621 u in width. Tips of first coxae not ex-
tending to anterior end of body. Posterior apodemes
of anterior coxal groups extending posteriorly beyond
suture lines between third and fourth coxae. Second
coxae each with one very long, stiff seta. Acetabular
fields with one pair of glandularia and 10-15 small
acetabula. Gonopore 107 u in length. Dorsal length
of palp segments: PI 21 u, PII 74 u, PIII 48 u, PIV
72 u, PV 43 u; setae on PIII larger than in male, PV
relatively more slender; peg-like seta on PIV not on a
tubercle. First two legs with ‘rillborsten’. Dorsal
lengths of last three segments of fourth leg: [V-leg-4
136 u, IV-leg-5 155 u and IV-leg-6 165 u. One
swimming seta on Il-leg-3 and IV-leg-4, two swim-
ming seta on I-leg-5, three swimming setae on I]-leg-
5, Hl-leg-5, IV-leg-4 and IV-leg-5; legs 3 and 4 with
rather large setae.
Etymology. — The species is named for its conspic-
uous colour.
Discussion. — The colour of the body (purple, even
after preservation in Koenike's solution) and the very
long stiff seta on the second coxa are diagnostic for
this species.
Smit: Water mites from Australia
Albia Thon
Albia (Albiella) brokenensis sp. n.
(figs.52-54)
Type material. — Holotype female: Broken River,
Eungella NP, Queensland (loc. 18).
Description
Female: Dorsal and ventral shields present. Body
1047 u in length and 737 u in width. Dorsal shield
with 5 pairs of glandularia, one pair of glandularia
lying in the dorsal furrow. Dorsoglandularia 3 and 4
lying very close to each other, dorsoglandularia 4 mo-
re medial. Glandularia of the fourth coxae extending
forward causing a slight bowing of the suture lines
between third and fourth coxae. None of the coxal
setae thickened. Acetabular plates with numerous
acetabula, inner margin free of acetabula. Pre-genital
shield present, triangular in form. Dorsal lengths of
palp segments: PI 35 u, PII 120 u, PIII 72 u, PIV
139 u, PV 53 u. Dorsal lengths of last three segments
of first leg: I-leg-4 110 u, I-leg-5 125 u, I-leg-6 132
u. Fifth segment of second, third and fourth legs with
numerous swimming setae; claws of legs with one
clawlet.
Male: Unknown.
Etymology. — The species is named after the type
locality.
Discussion. — This species is very close to A. austra-
lica Cook. It differs in the position of dorsoglandular-
ia 3 and 4, and in the form of the acetabular plates.
The anterior margin of the body of A. brokenensis is
almost straight, whilst that of A. australica is rounded.
Momoniella Viets
Momoniella australica Cook, 1986
Widespread in Australia, and reported from
Tasmania, New South Wales and Queensland.
Previously only collected in streams, but location 31
is Lake Eacham in the Atherton Tablelands area.
Material examined. — Queensland: 1 à , loc. 20; 1 9, loc.
DIANO
Gretacarus Viets
Gretacarus sp.
Males of Gretacarus exhibit the most useful taxo-
nomic characters and a number of species are known
only from males. Since only a female was found dur-
ing this study, no name can be applied to this speci-
men.
Material examined. — Queensland: 1 9, loc. 40.
105
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 52-54. Albia broke-
nensis sp. n. — 52, dorsal
view ®; 53, ventral view 9;
54, palp 2 (figs. 52-54 hol-
otype).
Arrenurus Dugès
Arrenurus (Arrenurus) balladoniensis Halik, 1940
The type locality of this species is Farm Balladonia,
southwestern Australia. Also reported from Victoria
(Lundblad 1948) and Queensland (Cook 1986).
Material examined. — Queensland: 1 d, loc. 39.
Arrenurus (Arrenurus) pseudoaffinis Piersig, 1906
A widespread species, also reported from Thailand,
Singapore, India and Indonesia.
Material examined. — Queensland: 1 2, loc. 21.
Arrenurus (Megaluracarus) tricornutus Viets, 1955
comb. n.
(figs. 55-58)
Viets (1955) described the female from a pool near
Mossman, Queensland. The species is also men-
tioned by Viets (1975), without giving the precise lo-
106
cation (but possibly Queensland). The male was
hitherto unknown. At one location both females and
males were collected. The male characters indicate
that the species belong to the subgenus
Megaluracarus. A description is given below.
Male: Body 1536 u in length , 1248 u in width
and 1416 u in height. Dorsum with three large, well
separated humps (as in the female) and four smaller
humps in anterior portion of dorsum. Eyes located on
the most lateral, rather pointed humps; the more me-
dial pair of these humps less pointed. Dorsal furrow
complete. Ventral side with a small epimeral field.
Acetabular plates extending to lateral margins of
body, the posterior margin slightly undulating.
Gonopore small, 68 jt in length. Dorsal lengths of
palp segments: PI 52 u, PIT 122 u, PIII 72 u, PIV
134 u, PV 62 u; PII with five setae on medial side.
Legs II, III and IV with swimming setae; IV-leg-4
without a distal projection.
Material examined: 1 d,2 9, loc. 22; 1 d, 1 ?, loc. 25;
le 261 loc 32 CPR lees 57
Smit: Water mites from Australia
Figs. 55-58. Arrenurus tricornutus Viets. — 55, dorsal view d ; 56, ventral view d; 57, lateral view d ; 58, palp d.
Arrenurus (Megaluracarus) rostratus degeneratus
Viets, 1984
À. rostratus is a widespread species, known from
China, Singapore, India, Shri Lanka and Indonesia.
Viets (1984) described the subspecies À. rostratus de-
generatus from the Northern Territory.
Material examined. — Queensland: 1 9, loc. 29.
Arrenurus (Megaluracarus) spec À
(figs. 70-72)
Only one female has been collected, and therefore
no name is assigned. It is possibly a subspecies of A.
rostratus (see discussion), and may therefore be placed
in the subgenus Megaluracarus.
Female: Length of body 844 u, width 630 u and
anteriorly pointed. Dorsal shield incomplete; lateral
sides with small indentations. Body posterior with a
short, truncate cauda. First coxae anteriorly pointed.
Gonopore 136 p in length. Dorsal length of palp seg-
ments: PI 24 u, PII 60 u, PIII 41 pi, PIV 62 u, PV 41
u; PII and PIV short, PII with two setae on medial
side. (Note: antagonistic bristle of normal length, but
depicted rather short in fig. 72). Legs II, III and IV
with swimming setae; I-leg-4 and 5 with only 1-2
swimming setae.
Discussion. — This species is possibly a subspecies
of A. rostratus Daday. Viets (1984) described A. ros-
tratus degeneratus from the Northern Territory,
Australia. Similar characters with this and other sub-
species are the pointed first coxae, the anteriorly
107
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 59-65. Arrenurus roobeeki sp. n. — 59, dorsal view d ; 60, ventral view d ; 61, lateral view 3; 62, palp d ; 63, dorsal view
? ; 64 ventral view 9; 65, lateral view 2. (figs. 59 - 62 holotype).
pointed body and the short PII and PIV. This female
differs in the form of the cauda and a very short PII
and PIV.
Material examined. — One @ from a pond near
Normanton, Queensland (loc. 37).
108
Arrenurus (Megaluracarus) roobeeki sp. n.
(figs. 59-65)
Type material. — Holotype male: Old river branch, south
of Mackay, Queensland (loc. 21). — Paratypes: Same loca-
tion as the holotype, 4 2,1 d.
Figs. 66-69. Arrenurus vanderpalae sp. n. — 66, dorsal view
3; 67, ventral view d ; 68, lateral view d ; 69, palp d (figs.
66-69 holotype).
Description
Male: Body 1392 u in length, 1140 u in width and
1416 u in height (including the humps). Dorsal fur-
row complete. Dorsum with three large humps, two
anterior bifid humps and one posterior. Epimeral
field small; fourth coxae with a medial side.
Acetabular plates extending to the lateral body mar-
gins; acetabular plates not well sclerotisized, and the-
refore difficult to observe. Gonopore small, 82 u in
length. Epimeroglandularia 2 on tubercles. Dorsal
lengths of palp segments: PI 53 u, PIL 110 u, PIII 62
u, PIV 125 u, PV 62 u; PII with three setae on me-
dial side. Second, third and fourth leg with many
swimming setae; IV-leg-4 without a distal projection.
Female: Body 1584 u in length, 1440 u in width
and 1584 u in height (including the humps). Dorsal
furrow complete. Dorsum with four large, bifid
humps. Third and fourth coxae almost without a me-
Smit: Water mites from Australia
dial side. Acetabular plates extending and expanding
towards to the lateral sides of body and not well scler-
otisized. Gonopore 136 u in length and 145 u in
width. Epimeroglandularia 2 on tubercles. Dorsal
lengths of palp segments: PI 53 u, PIL 118 u, PIII 67
u, PIV 127 u, PV 62 u. Legs II, IT and IV with many
swimming setae.
Etymology. — The species is named after Mr. C. F.
Roobeek, who spend many hours drawing water
mites.
Discussion. — The new species belongs to a group
of water mites mainly distributed on the Pacific is-
lands, e. g. A. lohmanni Piersig, 1898, A. bicornutus
Piersig, 1898, and A. multicornutus Walter, 1915.
Arrenurus (Megaluracarus) vanderpalae sp. n.
(figs. 66-69)
Type material. — Holotype male: Cloncurry River,
Cloncurry, Queensland (loc. 38).
Description
Male: Body 1176 u in length and 572 u in width.
Dorsal furrow incomplete, with two pairs of glandu-
laria. Dorsum with two small humps, on which glan-
dularia are located. Cauda 398 u in length, broade-
ning posteriorly. Posterior margin with two small
hyaline appendages and a small hyaline area. Dorsal
side of posterior end of cauda with a rudimentary
petiole consisting of two small spines. Glandularia of
the cauda on tubercles. Gonopore 78 u in length.
Acetabular plates short and broad, 24 u in length.
First coxae pointed anteriorly; medial space between
third and fourth coxae narrowing posteriorly; fourth
coxae large. Dorsal lengths of palp segments: PI 26 u,
PII 74 u, PIII 48 u, PIV 72 u, PV 41 u; PII with a
patch of setae. Leg IV with swimming setae; [V-leg-4
with a short distal projection. Legs I, II and III with
long spines; legs II and III with a few short, rudi-
mentary swimming setae.
Female: Unknown.
Etymology. — The species is named after Mrs. G.
M. van der Pal, who assisted me with the field work.
Discussion. — The shape of the cauda and the struc-
ture of the hyaline appendages are diagnostic for the
species.
Arrenurus (Micruracarus) forcipatoides Lundblad,
1941
Previously reported from Victoria and South
Australia (Lundblad 1948), this species is widespread
in ponds and small lakes in Queensland, including in-
land localities.
Material examined. — Queensland: 2 9, loc. 39; 4 d, 2
Le Zi Sole ZR UO O5 15 loc Bd, oe
IPO Cloer
109
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
74
Figs. 70-72. Arrenurus spec A. — 70, ventral view 9; 71, dorsal view 9; 72, palp 9. — Figs. 73-74. Arrenurus spec B . — 73,
ventral view 9; 74, palp 9.
Arrenurus (Truncaturus) tasmanicus Lundblad,
1941
The species has been reported by Lundblad (1948)
from Tasmania.
Material examined. — New South Wales: 1 4,8 ®, loc. 1;
lag) 2Hloe28.
Arrenurus spec B
(figs. 73-74)
Description
Female: Body 892 u in length and 752 u in width;
dorsal furrow incomplete. Gonopore 155 u in length.
Acetabular plates short and broad; genital field 436 u
in width. Ventral shield posterior with two small
humps. Dorsal length of palp segments: PI 34 u, PII
86 u, PIII 58 u, PIV 82 u, PV 46 u; PII medially
with three long setae. Third and fourth leg with ma-
ny swimming setae, second leg with only a few swim-
ming setae.
110
Male: Unknown.
Discussion. — As only the female is known, no
name is assigned.
Material examined. — Cloncurry River, Cloncurry,
Queensland, 1 2 (loc. 38)
ACKNOWLEDGEMENTS
I am indebted to G. M. van der Pal for her assistan-
ce during the field work and Dr. T. Gledhill for criti-
cally reading the manuscript. C. F. Roobeek made a
number of illustrations.
REFERENCES
Cook, D. R., 1967. Water mites from India. — Memoirs of
the American Entomological Institute 9: 1-411.
Cook, D. R., 1974. Water mite genera and subgenera. —
Memoirs of the American Entomological Institute 21: 1-
860.
Cook, D. R., 1986. Water mites from Australia. — Memoirs
of the American Entomological Institute 40: 1-568.
Halik, L. 1941. Beitrag zur Kenntnis der
Wassermilbenfauna Australiens. — Sbornik entomolo-
gickelo oddeleni Zemského Musea v Praze 19: 103-118.
Harvey, M. S., 1987. New and little-known species of the
water mite genera Tartarothyas, Pseudohydryphantes and
Cyclohydryphantes from Australia (Chelicerata:
Actinedida: Hydryphantidae). — Memoirs of the
Museum of Victoria 48 (2): 107-122.
Harvey, M. S., 1988a. Three new unusual water mites from
Australia (Chelicerata: Acarina:
Hydryphantidae, Hygrobatidae, and Athienemanniidae). —
Memoirs of the Museum of Victoria 49 (2): 355-361.
Harvey, M. S., 1988b. Two new species of the water mite
genus Aspidiobates Lundblad from Western Australia
(Acarina: Hygrobatidae). — Records of the Western
Australian Museum 14 (2): 199-209.
Harvey, M. S., 1988c. New species of the water mite family
Hydryphantidae (Acarina) from the Northern Territory,
Australia. — The Beagle, Records of the Northern
Territory Museum of Arts and Sciences 5 (1): 17-26.
Harvey, M. S., 1989a. A review of the water mite genus
Australorivacarus K. O. Viets (Chelicerata: Actinedida:
Hygrobatidae). — Invertebrate Taxonomy 3: 155-162.
Harvey, M. S., 1989b. A new species of Wuria K. Viets from
Northern Australia (Acarina: Arrenuridae). — The Beagle,
Records of the Northern Territory Museum of Arts and
Sciences 6 (1): 85-88.
Harvey, M. S., 1990a. A review of the water mite family
Limnocharidae in Australia (Acarina). — Invertebrate
Taxonomy 3: 483-493.
Harvey, M.S., 1990b. A review of the water mite family
Anisitsiellidae in Australia (Acarina). — Invertebrate
Taxonomy 3: 629-646.
Harvey, M. S., 1990c. Two new species of Partidomomonia
Cook from south-eastern Australia (Acarina:
Momoniidae). — Memoirs of the Museum of Victoria 50
(2): 337-340.
Harvey, M. S., 1990d. Two new water mite genera from
south-western Australia (Acarina: Aturidae:
Mideopsidae). — Memoirs of the Museum of Victoria 50
(2): 341-346.
Harvey, M. S. & D. R. Cook, 1988. Water mites of the ge-
nus Aspidiobates from Victoria, Australia, with the de-
scription of two new species (Chelicerata: Acarina:
Hygrobatidae. — Memoirs of the Museum of Victoria 49
(1): 51-57.
Hopkins, C. L., 1975. New species of Hygrobatidae and
Lebertiidae (Acari: Hydrachnellae) from New Zealand. —
Journal of the Royal Society of New Zealand 5 (1): 5-11.
Lundblad, O., 1941. Neue Wassermilben. Vorlaufige
Mitteilung. — Entmologisk Tidskrift 62: 97-121.
Lundblad, O., 1948. Zur Kenntnis australischer
Wassermilben. — Arkiv for Zoologi 40A (2): 1-82.
Piersig, R., 1906. Uber Siisswasser-Acarinen von Hinter-
' Most authors (including Lundblad himself) cite this paper
from Lundblad incorrectly as published in 1947.
However, it is printed in 1947, but published in 1948. In
the Zoological Record it is cited as Lundblad 1948.
Smit: Water mites from Australia
indien, Sumatra, Java und den Sandwich-Inseln (Reise
von Dr. Walter Volz.). — Zoologische Jahrbiicher,
Abteilung fiir Systematik 23: 321 - 394.
Rainbow, W. J., 1906. A synopsis of Australian Acarina. —
Records of the Australian Museum, Sydney 6: 145-193.
Smith, I. M. & M. S. Harvey, 1989. Description of adults of
Arrenurus (? Micruracarus) kitchingi sp. nov. (Acarina:
Arrenuridae) from water filled tree holes in Australia. —
Canadian Entomologist 121: 283-289.
Szalay, L., 1953. New data on Tasmanian water-mites
(Hydrachnellae) with a list of recorded species. — Papers
and Proceedings of the Royal Society of Tasmania 87: 73-
80.
Vidrine, M. F., 1986. Revision of the Unionicolinae (Acari:
Unionicolidae). — International Journal of Acarology 12
(4): 233-243.
Vidrine, M. F., 1987. Comments on the taxonomy of
Unionicola (Unionicolidae). — International Journal of
Acarology 13 (2): 157-158.
Viets, K., 1955. Kleine Sammlungen europäischer und
aussereuropäischer Wassermilben (Hydrachnellae,
Acari). — Abhandlungen naturwissenschaflichen Verein
zu Bremen 34: 1-26.
Viets, K. ©., 1969. Études Hydrobiologiques en Nouvelle-
Calédonie (Mission 1965 du Premier Institut de
Zoologie de l'Université de Vienne). VIII. Wassermilben
(Hydrachnellae, Acari). — Cahier Office de la Recherche
Scientifique et Technique Outre-Mer, série
Hydrobiologie 2 (3-4): 35-77.
Viets, K. O. 1975. Neue Wassermilben (Acari,
Hydrachnellae) aus Australien. — Zoologica Scripta 4: 93-
100.
Viets, K. O., 1976. Über drei neue Wassermilben
(Hydrachnellae, Acari) aus Australien. — Gewässer und
Abwässer 60/61: 71-84.
Viets, K. O., 1980. Weitere neue Wassermilben
(Hydrachnellae, Acari) aus Australien. — Gewässer und
Abwässer 66/67: 143-169.
Viets, K. O., 1981. Wassermilben (Acari, Hydrachnellae)
aus dem Northern Territory, Australia. — Entomologica
Scandinavica Supplement 15: 333 - 336.
Viets, K. O. 1984. Über Wassermilben (Acari,
Hydrachnellae) aus Australien. — Archiv für
Hydrobiologie 101 (3): 413-436.
Walter, C, 1915. Les Hydracariens de la Nouvelle-
Calédonie. — In: F. Sarasin & J. Roux, Nova Calédonia.
Forschungen in Neu-Caledonien und auf den Loyalty-
Inseln., Zool. 2, 7: 97-122.
Received: 9 July 1991
Revised version accepted: April 1992
111
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
APPENDIX I
Localities
New South Wales
1. Pond near Bombah Point, Myall Lakes NP
(National Park). July 11, 1989.
2. Muirs Creek, near Myal! Lakes NP. July 11,
1989.
3. Pond near Boolambay Creek, east of Bulahdelah.
July 12, 1989.
4. Boolambay Creek, east of Bulahdelah. July 12,
1989.
5. Pond near Iluka, 50 km NE of Grafton. July 14,
1989
6. Cattle pond, north of Bundjalung NP, south of
Woodburn. July 15, 1989.
7. Stream near Gap Road, Booroora Rest Area, near
Bundjalung NP. July 16, 1989.
8. Heath pond, northern part Bundjalung NP. July
16, 1989.
9. Stream near Gap Road, north of Bundjalung NP.
July 17, 1989.
10. Pond near Gap Road, Bundjalung NP. July 17,
1989.
42. Stream near Blackman campsite, Warrumbungle
NP, 25 km east of Coonabarabran. August 21,
1989.
43. Spring, near Blackman campsite, Warrumbungle
NP. August 2, 1989.
44. Cattle pond nr. 1 near Blackman campsite,
Warrumbungle NP. August 21, 1989
45. Cattle pond nr. 2 near Blackman campsite,
Warrumbungle NP. August 21, 1989
46. Small lake near Bundeena, Royal NP, 25 km
south of Sydney. August 25, 1989.
Queensland
11. Creek at Elabama Falls, Lamington NP, 100 km
south of Brisbane. July 18, 1989.
12. Blue Pool, Lamington NP. July 18, 1989.
13. Darraboola Creek, Lamington NP. July 19,
1989.
14. Poona Lake, Cooloola NP, 50 km NE of
Gympie. July 21, 1989.
15. Pond along road to Rainbow Beach, Cooloola
NP. July 21, 1989.
16. Teewah Creek, Cooloola NP. July 22, 1989.
112
117%
18.
19:
20.
Ds
22.
23:
24.
29)
26.
DIE
28.
29:
30.
gE
32:
33.
34.
35.
36.
OÙ
38.
39;
40.
41.
Ululah Lagoon, Maryborough. July 22, 1989.
Broken River, Eungella NP, 80 km west of
Mackay. July 27, 1989.
Lake near Eungella Dam, 10 km west of
Eungella. July 28, 1989.
Small stream, tributary of Broken River, Eungella
NP. July 28, 1989.
Old river branch, south of Mackay. July 29,
1989.
Pond near Crocodile Rd, Bowling Green Bay
NP, 40 km south of Townsville. July 31, 1989.
Alligator Creek, Mnt. Elliot NP, 30 km south of
Townsville. July 31, 1989.
Small lake, Townsville Common NP, north of
Townsville. August 1, 1989.
Pond Townsville Common NP. August 1, 1989.
Freshwater Lagoon, Horseshoe Bay, Magnetic
Island. August 3, 1989.
Douglas Creek, Palmerston NP, 30 km SE of
Millaa Millaa. August 5, 1989.
Hasties Swamp NP, 4 km south of Atherton,
Atherton Tablelands. August 6, 1989.
Lake Barrine, 15 km east of Atherton, Atherton
Tablelands. August 7, 1989.
Lake near Tinaroo Falls Dam, Tinaroo, 15 km
NE of Atherton, Atherton Tablelands. August 7,
1989.
Lake Eacham, 15 km east of Atherton, Atherton
Tablelands. August 8, 1989.
Cattle pond, Upper Daintree Road, Daintree.
August 9, 1989.
Elizabeth Creek, east of Mnt. Surprise. August
11, 1989.
Creek near Eubanangee Swamp NP, 15 km
north of Innisfail. August 11, 1989.
Creek Innot Hot Springs, 30 km west of
Ravenshoe. August 11, 1989.
Cattle pond, 21 km west of Georgetown. August
12, 1989.
Pond north of Normanton. August 14, 1989.
Cloncurry River, Cloncurry. Dry river bed with
some ponds. August 14, 1989.
Pond north of Longreach. August 15, 1989.
Carnarvon Creek, Carnarvon NP, 100 km SE of
Rolleston. August 17, 1989.
Pond between Carnarvon NP and Injune, ap-
proximately 38 km east of Carnarvon. August 19,
1989.
JAN H. STOCK
Institute of Taxonomic Zoology, University of. Amsterdam
PYCNOGONIDA FROM SOUTHERN BRAZIL
Stock, Jan H., 1991. Pycnogonida from southern Brazil. — Tijdschrift voor Entomologie, 135:
113-139, figs. 1-80. [issn 0040-7496]. Published 15 July 1992.
Records of Pycnogonida collected at 85 different stations in southern Brazil, between 18°34' S
and 34°27' S. Four of these were sampled by hand or by skin-diving in the littoral and upper
infralittoral zones, six stations were located on the continental slope between 250 and 1100 m,
the remaining 75 were on the continental shelf. Forty-four species are represented, dispersed
over 16 genera in seven families. Seven species are new to science, in the genera Eurycyde (E.
platyspina sp. n.), Achelia (A. columnaris sp. n.), Nymphon (N. bullatum, N. inaequipes, N. vul-
canellum spp. n.), Anoplodactylus (A. spurius sp. n.), and Pycnogonum (P. ornans sp. n.). New to
the fauna of Brazil are eight species. New synonymies are proposed in Pallenopsis (P. boehmi
Schimkewitsch, 1930 = P. longicoxa Stock, 1966) and Anoplodactylus (A. petiolatus (Kroyer,
1844) = A. guyanensis Child, 1977).
J.H. Stock, Institute of Taxonomic Zoology, P. O. Box 4766, 1009 AT Amsterdam, The
Netherlands.
Key words. — Pycnogonida, taxonomy, distribution, Brazil, littoral, shelf, continental slope.
The pycnogonid fauna of southern Brazil is rela-
tively well-studied, mainly through the monographic
treatment of Marcus (1940b) and a number of satel-
lite papers in the following decades by Corrêa (1948),
Sawaya (1945, 1947, 1950, 1951), Zilberberg
(1963), Zago (1970), de Mello-Leitao (1945, 1946,
1949 a, b, 1955), Marcus & Marcus (1963) and
Marcus-du Bois-Reymond (1952). Occasionally,
large expeditions collected material on the Brazilian
coast, e.g. the ‘Challenger’ (Hoek 1881), the ‘Vettor
Pisani’ (Schimkewitsch 1890), the ‘Calypso’ (Stock
1966), and some American cruises (Child 1982a).
Nevertheless the present collections from southern
Brazil form a fine addition to our previous know-
ledge. The material was brought together by the late
Dr. Plinio Soares Moreira of the Instituto
Oceanographico of the University of Sao Paulo (usp),
mainly on board of the Oceanographic Research
Vessel ‘Prof. W. Besnard’.
The amount of previous work done explains why
the number of new species in the present collection is
relatively low (7 out of 44, or 16%). However, taken
into consideration that another eight species are new
to Brazil, a total of 15 species is added to the Brazilian
fauna, which is not too bad a score (more than one-
third of the species represented in this collection).
Biogeographically, endemics and warm-water spe-
cies are predominant in the Brazilian fauna, not sur-
prising in the light of the South Atlantic Drift which
distributes tropical waters along major parts of the
coastline of the country.
As table 1 shows, a good number of species (viz.
21) is endemic to Brazil (and neighbouring territo-
ries, the Mar del Plata and the Guyanas). Another
group of eleven species are mainly warm-water ele-
ments which have their distribution centre in the
Caribbean. One of these species, Ammothella spinife-
ra, occurs on the Pacific coasts of the Americas as
well. Interesting is a small group of seven species oc-
curring on both sides of the Atlantic Ocean (so-called
amphi-Atlantic species). Finally, some Magellanean
cold-water species reach their northern distribution
limits on the shelf of southern Brazil, but this group
contains only three species in all.
METHODS
Entire specimens were studied and measured in
glycerin or lactophenol, dissected appendages on per-
manent slides in Reyne's modification of Faure's me-
dium. All specimens have been deposited in the
Zoölogisch Museum, Amsterdam (ZMA).
DESCRIPTIVE PART
Family Ammotheidae
Eurycyde platyspina sp. n.
(figs. 1-8)
Type material. - 1 9 (holotype), 1 d (paratype), Project
Isotan-Draga I, Stn. MBT 155: 22°22' S 41°32' W, 34 m, 5
Sep 1970 (zma Pa. 3443).
Description. — Unlike certain other species of the
113
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-8. Eurycyde platyspina sp. n. (3 holotype and © allotype). 1, trunk. 2, dorsal (scale A); 2, trunk, ©, from the left (A);
3, chelifore 4 (B); 4, palp, & (B); 5, oviger, 4 (C); 6, compound spine 5 of oviger segment 7, d (D); 7, leg 3, d (A); 8, dis-
tal segments of leg 3, & (B). Scales after fig. 80.
114
genus, the two sexes are very similar in spination, etc.,
even in the morphology of the oviger. Secondary sex-
ual differences (presence of femoral cement gland
tube in male, large genital pores on ventral surface of
coxa 2 of all legs in female) may serve to discriminate
between the sexes.
Trunk completely segmented; mid-dorsal line
without tubercles. Lateral processes separated by nar-
row intervals, distally with low tubercle. Oviger im-
plantation free of first lateral process. Proboscis with
articulated stalk, distal part directed downward.
Abdomen straight in basal part; at about 80% of its
length 6 setae are implanted at bend of abdomen.
Ocular tubercle cylindrical with pointed tip; eyes
small, well-pigmented, touching; no setae on ocular
tubercle.
Chelifore scape 2-segmented, segments subequal in
length; segm. 1 with 3 long spines, segm. 2 with 7.
Chela not retracted in distal end of scape; hand glob-
ular, with 1 rudimentary finger and 2 setae.
All spines (on chelifores, coxae 1 and 2, femur and
tibiae 1 and 2) are of rather peculiar morphology (fig.
1, detail), viz. flattened, with more or less obtuse tip
and with distal spinule, both margins with some lat-
eral setules.
Palp segm. 1 with very strong lateral spur; segm. 2
minute; segm. 3 longest; segm. 6 shorter than 7;
segms. 7 to 9 subequal, segm. 10 shorter.
Oviger segm. 4 longest; segms. 7 to 10 with 2 rows
of compound spines, in longest row according to for-
mula 6:4:4:5. Compound spines with 3 small proxi-
mal marginal teeth, then 1 large tooth, and 4 to 5
smaller distal teeth. Terminal claw short, slightly
curved, smooth.
Coxa 1 with 2 long disto-dorsal spurs and 2 more
proximal dorsal spines. Coxa 2 with 2 + 2 dorsal
spines. Coxa 3 with minute spinules only. Femur of
male with swelling at about 40% of posterior surface,
bearing tube of cement gland; femur of female with-
out such swelling; distal end of femur with 4 long
spines. Tibiae 1 and 2 with several (5-7) long dorsal
spines, plus some long lateral spines. Tarsus short.
Propodus slightly curved, distally widening, no heel;
sole with 12-14 spinules of uniform size. Claw short,
curved; no auxiliary claws.
Female genital pores on distoventral surface of coxa
2 of all legs; male cement glands on femur of all legs.
Measurements (mm) of holotype: Length trunk
(frontal margin cephalic segment to tip 4th lateral
process) 1.71; diameter across 2nd lateral processes
1.17; length abdomen 0.58; length proboscis 1.36.
Paratype, leg 3: first coxa 0.31; second coxa 0.49;
third coxa 0.27; femur 1.01; first tibia 1.17; second
tibia 1.17; tarsus 0.10; propodus 0.70; claw 0.23.
Etymology. - The specific name, platyspina, refers
to the curious shape of the spines on chelifores, abdo-
Jan H. Stock: Brazilian Pycnogonida
men and legs.
Remarks. — Child (1988b) has presented a key to
the species of Eurycyde. Since the construction of this
key, three more species became known: E. antarctica
Child, 1987 (which can be traced down to couplet 7
of the key), E. longioculata Müller, 1990 and E. dia-
cantha Stock, 1990 (both ‘strand’ in couplet 3 of the
key). Child (1988b: 7) remarks that £. acanthopus
Stock, 1979 (described from a female) is probably the
opposite sex of E. curvata Child, 1979 (described
from a male). If this supposition is true, it might be
useful to know which name has priority. It turned out
to be almost a photofinish: E. acanthopus was pub-
lished Sep 10, 1979, and E. curvata Sep 27, 1979, so
the former name has priority.
Eurycyde platyspina sp. n. can be followed in
Child's key to couplet 7, along with the couple acan-
thopus | curvata, hispida (Kroyer, 1844) and antarcti-
ca. It differs from antarctica in having long spines on
the abdomen (abdomen practically glabrous in ant-
arctica), and from acanthopus/curvata in having the
abdominal spines arranged in a single dorsal group
(versus scattered over the abdomen). Finally it differs
from hispida in the presence of a tubercle on the later-
al processes and 2 spurs on coxa 1 (none in hispida).
Nymphopsis melidae Sawaya, 1947
N. melidae Sawaya, 1947: 83-87, pls. II.
Material. - 1 chelate juvenile, Project Isotan-Draga I,
Stn. MBT 157: 21°46'S 40°58' W, 15 m, 5 Sep 1970; 1 9,2
chelate juveniles, Project Isotan-Draga I, Stn. MBT 165:
21°20' S 40°39' W, 28 m, 6 Sep 1970; 1 6, Project
Petrobas/usp, Stn RD 54: 18°54.8'S 39°15.4 W, 41 m, 12
Aug 1972.
Remarks. - Up to now this species has been record-
ed only once: Sawaya (1947) described a single male
from the upper littoral zone of Ilha das Palmas, Baia
de Santos (ca. 24° S). The present records are more
north, and from deeper shelf waters (15-41 m).
The juveniles in this collection have an unsegment-
ed scape, the adults a 2-segmented scape. This fea-
ture, often used as a key character, should be used
with some care.
Cilunculus sp.
Material. - 1 juvenile, Project Isotan-Draga I, Stn MBT
143: 24°08'S 42°28'W, 1020 m, 2 Sep 1970.
Remarks. - This juvenile has still retained its che-
lae, and the ovigers are an undeveloped, sausage-like
structure. In most of its characters, it resembles close-
ly €. europaeus Stock, 1978, from deep waters of the
Bay of Biscay and the entrance of the English
Channel (absence of an ocular tubercle, long setae on
the legs, long auxiliary claws, absence of mid-dorsal
115
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
trunk tubercles), but it differs from C. europaeus in
the naked lateral processes (setose in europaeus) and in
the greater space between the anterior lateral process-
es (interval 3 times the diameter of the lateral pro-
cess). The propodal heel bears 1 larger spine (2 in eu-
ropaeus) and the sole bears only 6 spines, which are
longer than those of europaeus.
I presume this specimen represents an undescribed
species, but in absence of adults I refrain from nam-
ing it. The genus Cilunculus is new to Brazil.
Ammothella appendiculata (Dohrn, 1881)
A. appendiculata, Child 1979: 9 [refs.].
Material. - 1 d, usp Stn. F 23: Enseada do Flamengo,
Ubatuba, State of Sao Paulo; intertidal of rocky shore; 13
Feb 1965; 1 ®, Project Isotan-Draga I, Stn. MBT 165:
21°20'S 40°39'W, 28 m, 6 Sep 1970.
Remarks. - The female of Stn. MBT 165 is in agree-
ment with the general diagnosis of this species. The
male of Stn. F 23 is intermediate between A. appendi-
culata and A. rugulosa (vide infra), in that it has the
short first scape segment of the former, but it possess-
es a hollow spine on the scape and spines on the later-
al processes, that characterize the latter. Presumably
the two species are identical.
A. appendiculata is an amphi-Atlantic species, al-
ready known from Brazil (Marcus 1940b).
Ammothella rugulosa (Verrill, 1900)
A. rugulosa; Child 1979: 11 [refs.].
Material. — 1 d, 1 juv., Project Isotan-Draga I, Stn. MBT
155: 22°22'S 41°32'W, 34 m, 5 Sep 1970.
Remarks. — The adult male agrees well with the
characteristics presumed to characterize this species.
However, in view of the large range of variation ob-
served in A. appendiculata (vide supra), and in agree-
ment with the opinion of Stock (1975a) and Child
(1979), A. rugulosa is probably synonymous with that
species.
A. rugulosa is a West Indian species, recorded be-
fore from Brazil by Marcus (1940b).
Ammothella spinifera Cole, 1904
A. spinifera Cole, 1904: 275-277, pl. 12 fig. 8, pl. 20 figs. 7-
9, pl. 21 figs. 1-6; Hilton 1915: 204; Child 1979: 11-12,
fig. 3f-h [older refs.]; Wicksten 1980: 198; Austin 1985:
428; Salazar-Vallejo & Stock 1987: 269; Müller & Roth
1990: fig. 4; Müller 1990b: 278.
Material. - 1 d (ovig.), 1 4, 4 2, 3 juvs., usp Stn. F 10:
Ponta de Santa Rita, Enseada do Flamengo, State of Säo
Paulo; amongst ascidians, bryozoans and sponges in infralit-
toral zone on rocks; 13 Feb 1965.
116
Remarks. — The present material falls perfectly
within the range of variation described by Child
(1979). The species is mainly known from the Pacific
(from southern California to the coast of Panamá and
the Galapagos Islands). Child (1979) was the first to
record it from the Caribbean coast of Panamá, and re-
cently Müller (1990b) recorded it from Martinique
and Barbados. Its range is considerably extended
southward, to the coast of southern Brazil, by the
present record.
Achelia besnardi Sawaya, 1951
A. besnardi; Stock 1975b: 70-71, figs. 1-9 [older refs.].
Material. - 1 d, Project Petrobas/usp, Stn. RD 40:
19°15.8'S 38°56.3'W, 59 m, 11 July 1972.
Remarks. — This specimen resembles closely
Sawaya's (1951) good illustrations. It is a relatively
rare species, having been found off Rio de Janeiro, in
the Virgin Islands, and the Bahamas. There is no ma-
terial available that substantiates Fry & Hedgpeth's
claim (1969: 102) that the species occurs in the for-
mer Belgian Congo or (on their distribution map, fig.
155) on the South American coast as far south as 45°.
Probably, these authors have confused A. besnardi
with A. sawayai (vide infra) which was recorded by
Fage (1949) from Zaire.
Contrary to Fry & Hedgpeth's (1969: 105) opin-
ion, I consider this species closely related to A. saway-
ai, with which it agrees in two essential characters: (1)
all spines on the propodal sole are of equal size, and
(2) absence of a male genital spur.
Achelia columnaris sp. n.
(figs. 9-18)
Type material. - 1 d (ovig.), holotype and 3 ® para-
types, Project Isotan-Draga I, Stn. MBT 155: 22°22'S
41°32'W, 34 m, 5 Sep 1970 (zma Pa. 3444).
Description. - Male: Trunk compact, hardly any
space between lateral processes. Trunk segments 1, 2,
and 3 separated by articulation lines, segments 3 and
4 fused. Trunk segments 2 and 3 with bifid mid-dor-
sal spur (anterior branch longer than posterior one),
tipped with spinules. Trunk segment 4 and proximal
end of abdomen with lower, simple mid-dorsal spur.
Anterior margin of cephalic segment with pointed tu-
bercle on either corner. Ocular tubercle with 2 min-
ute lateral knobs and small distal knob; eyes small,
pigmented. Lateral processes with spiniferous spurs,
some bifid or trifid at tip.
Proboscis of type C (greatest width at less than
mid-length, distal part tapering). Abdomen reaching
to distal end of coxa 1 of leg 4, slightly upcurved;
apart from dorsal spur in basal part, it bears 2 lateral
Jan H. Stock: Brazilian Pycnogonida
Figs. 9-14. Achelia columnaris sp. n. (6 holotype and © paratype). 9, trunk, à , dorsal (scale A); 10, contour of dorsum, à,
from the right (A); 11, proboscis, d, ventral (A); 12, palp, © (B); 13, leg 3, d (A), 14, leg 3, 2 (A). Scales after fig. 80.
WL,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
spiniferous tubercles near distal end.
Chelifore scape with 1 tall mediolateral spur, and 2
distolateral spurs of unequal length. Chela with 2
pointed processes (rudiments of fingers).
Palp 8-segmented; basal segments covered with
short spinules; segm. 2 with distolateral spiniferous
process; a much taller, pillar-shaped process present
on distolateral corner of segm. 3. Segm. 2 > 4; segm.
5 < 6;6>7; 7 < 8. Distal segnıs. slender, with ventral
brush of setae.
Oviger segms. 4 and 5 subequal, with recurved row
of spinules. Segm. 6 with strong, reversed spine in ba-
sal part, and several other, smaller, reversed spinules.
Compound spines elongate, with some 8 pairs of
marginal teeth; formula 2:2:1:2.
Legs: Coxa 1 with 1 or 2 tall spiniferous spurs on
anterior margin, 2 on posterior margin, and 2 or 3 on
distal margin. Coxa 2 with 3 or 4 tall spurs on anteri-
or and posterior margins. Coxa 3 with 1 or 2 spinifer-
ous tubercles. Femur with numerous spiniferous pro-
cesses on posterior and dorsal surface, taller towards
distal end. Tibia 1 with many very tall, pillar-shaped,
spiniferous spurs on dorsal and posterior surface, tib-
ia 2 with still more numerous, but slightly less tall
such spurs. Tarsus short, with 4 or 5 ventral spines.
Propodus slightly curved, heel with 3 or 4 large
spines; sole with some 7 smaller spines. Auxiliary
claws more than half as long as main claw. Genital
spur long, thump-shaped, on ventrodistal corner of
coxa 2 of legs 3 and 4.
Female: Mid-dorsal trunk tubercles simple (not bi-
fid); tubercle over base of abdomen lacking. Spurs on
lateral processes and legs as tall as in male, but coxa 1
without marginal spurs (only with distal spurs, occa-
sionally with 1 small posterior spur). Coxa 2 with 1
pair of marginal spurs and 1 pair of distal spurs.
Femur more swollen than in male. Genital apertures
on ventral surface of coxa 2 of all legs.
Measurements (mm) of male (those of female in
parentheses). — Length proboscis (ventral) 1.10
(1.15); greatest diameter proboscis 0.44 (0.43);
length trunk (to tip 4th lateral process) 1.30 (1.26);
width across 2nd lateral processes 1.16 (0.94); length
abdomen 0.59 (0.59); length scape 0.57 (0.67); di-
ameter egg on male oviger 0.15.
Leg 3: first coxa 0.46 (0.34); second coxa 0.50
(0.57); third coxa 0.34 (0.38); femur 1.177 (1.37);
first tibia 1.27 (1.39); second tibia 1.31 (1.46); tarsus
0.13 (0.11); propodus 0.71 (0.74); claw 0.39 (0.40);
auxiliary claw 0.23 (0.28).
Etymology. - The proposed specific name, colum-
naris (Latin, pillar-shaped) alludes to the shape of the
spurs on femur and tibiae.
Remarks. — The new species shows, by the presence
of a row of mid-dorsal trunk tubercles, a strong simi-
larity to the Achelia assimilis/wilsoni/variabilis com-
118
plex (see Hedgpeth 1961), frequently recorded from
the major oceans of the southern hemisphere.
However, none of the recorded ‘variations’ has tall,
pillar-shaped spurs on femur and tibiae. Similar tall
spurs are present on palp segments 2 and 3 of the new
species, whereas in the A. assimilis group just a tuber-
cle is present.
The very strong armature of the legs is not unlike
that found in À. setulosa (Loman, 1912), but that spe-
cies lacks mid-dorsal trunk and palp spurs, and has a
less slender distal palp segment.
Other very spinose Achelia species from the central
Atlantic Ocean are A. turba Stock, 1990 (which has
however no mid-dorsal trunk spurs, shorter spurs on
the long leg segments, a shorter male genital spur, and
a less pointed proboscis), and A. armata (Bouvier,
1916) (which lacks spurs on the long leg segments or
in the mid-dorsal line and has less elongate femur,
tibiae 1 and 2, and propodus).
Achelia gracilis Verrill, 1900
A. gracilis, Hedgpeth 1948: 244, fig. 38 f-g [older refs.];
Stock 1954: 117; Stock 1979: 10; Stock 1986: 416;
Zambrana et al. 1985: 30 [refs.]; Markham 1986: 276,
fig. 87.
Material. — 1 d, Project Petrobas/usp, Stn. RD 50:
19°02.5'S 39°30.6'W, 28 m, 12 July 1972.
Remarks. — This species was known from Bermuda
to the West Indies. It is new to the fauna of Brazil.
Achelia sawayai Marcus, 1940
A. sawayai, Krapp & Kraeuter 1976: 342-343 [older refs.];
Stock 1986: 414-416 [older refs.]; Stock 1989: 87-89;
Müller 1989: 124; Müller 1990a: 105-106; Müller
1990b: 277; Stock 1990: 218-219.
Material. - 64 spms., usp Stn. P.E.: Ponta do Espia,
Ubatuba, State of Sao Paulo, on seaweeds of rocky shore, in-
fralittoral, 20 Nov 1964; 3 6,6 2,25 juvs., usp Stn. F 23:
Enseada do Flamengo, Ubatuba, State of Sao Paulo; infralit-
toral, on large boulders, amongst ascidians, bryozoans,
sponges, 13 Feb 1965; 1 2, 1 juv., usp Stn. F 10: Ponta de
Santa Rita, Enseada do Flamengo, Ubatuba, State of Sao
Paulo; intertidal, rocky shore, on Dyctiota sp., 12 Dec 1965;
1 d ovig., 1 d,1 ©, Project Isotan-Draga I (PID) Stn. MBT
152: 23°03'S 41°10'W, 98 m, 4 Sep 1970; 1 juv., PID Stn.
MBT 159: 22°00'S 40°06'W, 90 m, 5 Sep 1970; 1 juv., PID
Stn. MBT 161: 21°42'S 40°15'W, 56 m, 6 Sep 1970; 2 d, 1
2,5 juv., Pip Stn. MBT 164: 21°15'S 40°50'W, 19 m, 6 Sep
1970; 1 d ovig., Project Petrobas/usp, Stn. RD 54:
18°54.8'S 39°15.4'W, 41 m, 12 July 1972; 1 6,1 2, 1 juv.,
Project Petrobas/usp, Stn. RD 56: 18°54.7'S 38°53.5'W, 38
m, 12 July 1972.
Remarks. — Originally described from Brazil, the
species is now recorded from all over the West Indies,
the Gulf of Mexico, the American east coast as far
north as Georgia, the Cape Verde Islands, Mauritania
Jan H. Stock: Brazilian Pycnogonida
Figs. 15-18. Achelia columnaris sp. n. (d holotype and © paratype). 15, chelifore, 2 (scale B); 16, oviger, 2 (B); 17, com-
pound spine of oviger segment 10 (free-hand sketch); 18, oviger, d (B). Scales after fig. 80.
(unpubl. record based on material in the Museum Gr.
Antipa, Bucarest), Zaire, Madagascar, and the Society
Islands.
Tanystylum acuminatum Stock, 1954
(figs. 19-21)
T. acuminatum Stock, 1954: 125-127, fig. 29; Stock 1979:
11; Müller 1990b: 278.
Material. — 15 spms. (incl. ovigerous males), usp Stn.
P.E.: Ponta do Espia, Ubatuba, State of Sao Paulo, on sea-
weeds growing in the infralittoral of rocky shore, 20 Nov
1964; 1 2, usp Stn F 27: Rocky shore between Praia do
Dionisio and Praia do Flamengo, Enseada do Flamengo,
Ubatuba, State of Sao Paulo, on algae at low tide, 15 Feb
1965; 1 2, usp Stn F 10: Ponta Santa Rita, Enseada do
Flamengo, Ubatuba, State of Sao Paulo, intertidal of rocky
shore, on Dyctiota sp., 12 Dec 1965; 1 9, Project Isotan-
Draga I, Stn. MBT 157: 21°46'S 40°58'W, 15 m, 5 Sep 1970.
Remarks. - The male has never been described be-
fore. It resembles the female in almost every charac-
ter, except for the tuberculation of coxa 1, which is
slightly stronger in the male, and of course the struc-
ture of the oviger. The male oviger has an ecto-distal
apophysis on segment 7, which places 7° acuminatum
in a group of species comprising 7. tubirostrum Stock,
1954, 7. dowi Child, 1979, 7. oculospinosum Hilton,
1942, T. isthmiacum Stock, 1955, and 7. isabellae
Marcus, 1940.
T. acuminatum is new to Brazil; it was previously
known from the Lesser Antilles only (St. Barts,
Anguilla, Martinique).
Tanystylum evelinae Marcus, 1940
T. evelinae Marcus, 1940b: 99-102, fig. 13.
Material. - 1 d ovig., usp Stn. P.E.: Ponta do Espia,
Ubatuba, State of Sao Paulo, on seaweeds growing on rocky
shore, infralittoral, 20 Nov 1964.
Remarks. - The single specimen is badly damaged,
but is probably this species. 7° evelinae was already
found in the same general area by Marcus (1940b),
but was never recorded again. As point of fact it re-
sembles strongly 7. haswelli Child, 1990, an
Australian form. The palps of haswelli are 4-segment-
ed, those of the types of evelinae 6-segmented, but in
the present specimen, articulation lines in the basal
part of the palp are obscure, resulting in a 4-segment-
ed appearance of the appendage.
Tanystylum isabellae Marcus, 1940
(figs. 22-26)
T. isabellae Marcus, 1940b: 95, 102-105, fig. 14; Fage 1949:
29, figs:
Material. - 1 d, 3 juvs., usp Stn. P.E.: Ponta do Espia,
Ubatuba, State of Sao Paulo, on seaweeds growing on rocky
shore, infralittoral, 20 Nov 1964.
Remarks. - The types come from the same area and
the same depth. Fage (1949) records it moreover
from Zaïre.
119
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 19-21. Tanystylum acuminatum Stock, 1954 (& from Stn. usp/P.E.). 19, palp (scale E); 20, oviger (E); 21, spine 1 of
oviger segment 10 (free-hand sketch).
Figs. 22-26. Tanystylum isabellae Marcus, 1940 (3 from Stn. usr/P.E.). 22, ocular tubercle from the left (free-hand sketch);
23, proboscis and chelifores (A); 24, palp (E); 25 distal part of oviger (E); 26, leg 2 (B). Scales after fig. 80.
120
Some new illustrations are incorporated here,
mainly to show the shape of the proboscis. The pe-
nultimate palp segment is slightly less lobate than
Marcus’ figure 14E suggests, but it is in good agree-
ment with Fage's fig. 5.
This species is strikingly similar to 7° oculospino-
sum Hilton, 1942 (vide Child & Hedgpeth 1971:
619, fig. 5, and Child 1979: 34), a Pacific species
ranging from Baja California to the Galapagos
Islands. The only difference seems to be the second
tibia which looks shorter in Child & Hedgpeth's fig.
5c. I suppose, however, that both species are synony-
mous, in which case Marcus’ name has priority. Child
(1979) records another Tanystylum, T. tubirostrum
Stock, 1954 and several other species of Pycnogonida
from both the Atlantic and the Pacific side of the
Americas, so from a biogeographic point of view it
would not be too surprising if the Pacific 7. oculospi-
nosum and the Atlantic 7: isabellae were synonymous.
Tanystylum isthmiacum difficile Stock, 1966
T. isthmiacum difficile Stock, 1966: 389-390, fig. 2 a-h; Fage
& Stock 1966: 318; Stock 1975a: 984; Child 1979: 29-
30; Stock 1979: 11-13, fig. 3 b-f; Zambrana et al. 1985:
31 [refs.].
Material. — 1 juv., Project Petrobas/usp, Stn. RD 50:
19°02.5'S 39°30.6'W, 28 m, 12 July 1972.
Remarks. - This young specimen, with still poorly
developed ovigers, agrees well with the published de-
scriptions of this (sub)species. It was already known
from the same general depth in the same area (Stock
1966).
Tanystylum orbiculare Wilson, 1878
T. orbiculare, Krapp 1973: 69, fig. 4; Child 1990: 319, fig.
3 [holotype re-illustrated].
Material. - 5 spms., usp Stn. P.E.: Ponta do Espia,
Ubatuba, State of Sao Paulo, on seaweeds of intertidal zone
of rocky shore, 20 Nov 1964; 1 2, usp Stn. F 27: Rocky
shore between Praia do Dionisio and Praia do Flamengo,
Enseada do Flamengo, Ubatuba, State of Sao Paulo, on al-
gae slightly exposed at low tide, 15 Feb 1965.
Remarks. — The proboscis shape resembles Marcus
(1940b) figure rather than Krapp's (1973) illustra-
tion. The female oviger is 10-segmented (versus 9-
segmented in the closely similar 7. conirostre) and the
propodal armature is as Krapp describes for 7. orbic-
ulare. The abdomen too, resembles Marcus fig. 15A,
instead of Child's (1990) fig. 3 A-B. Perhaps this is
not orbiculare at all, but another look-alike in this dif
ficult genus.
Family Nymphonidae
Jan H. Stock: Brazilian Pycnogonida
Nymphon bullatum sp. n.
(figs. 27-34, 47-51)
Type material. - 1 d (holotype), 8 d (of which 1 oviger-
ous), 6 9, 25 juvs. and larvae (paratypes), Project Isotan-
Draga I (pip), Stn. MBT 149: 23°41'S 41°55'W, 250 m, 4
Sep 1970 (zma Pa. 3446).
Other material. - 1 juv., Pip, Stn. MBT 153: 22°48'S
41°14'W, 65 m, 4 Sep 1970; 1 (fragm.), rip, Stn. MBT
181: 25°35'S 46°03'W, 136 m, 29 May 1971.
Description. - Trunk completely segmented, un-
adorned. Lateral processes narrow, separated by inter-
vals distinctly larger than their own diameter. Neck
strongly widened in anterior end; narrow, parallel-
sided portion just in front of oviger implantation; lat-
ter practically in contact with first lateral process.
Ocular tubercle truncate in frontal view, with 2 min-
ute distolateral tubercles; in side view as illustrated;
eyes well-pigmented. Proboscis almost cylindrical.
Abdomen reaching to distal end of coxa 1 of leg 4.
Chelifore scape thin, longer than proboscis, armed
with some setules in middle and at distal end. Chela
much shorter than scape; palm much shorter than
fingers; both fingers curved; movable finger with 19
regular teeth, immovable finger with 14.
Palp segm. 2 longest; segm. 3 ca. two-thirds of
segm. 2; segm. 4 distinctly shorter than segm. 5; long
setae on segms. 3 to 5.
Oviger of male highly characteristic: segm. 5 long-
est, somewhat S-shaped, with strong apophysis at
two-thirds of its endal margin; segm. 6 curved, setif-
erous. Compound spine formula 7:4:3:4; compound
spines with 2 or 3 pairs of marginal teeth. Distal claw
with 7 endal teeth.
Oviger of female: segm. 5 hardly longer than segm.
4, straight. Compound spine formula 6:5:4:4. Distal
claw with 6 endal teeth.
Legs thin, slender. Coxa 2 shortest in leg 1, longest
in legs 3 and 4; very slender in posterior legs. Femur
< tibia 1 < tibia 2. Femur of male with 3 ventral ce-
ment gland apertures, shaped as truncate cones.
Femur of female: basal two-thirds swollen (contain-
ing eggs). All long leg segments with many setae,
mostly about as long as segment diameter. Tarsus
slightly shorter than propodus; both segments with
row of small spinules of uniform size on ventral mar-
gin; no heel or heel spines differentiated. Claw
curved, about 60% of length of propodus; auxiliary
claws feeble, about 30% of length of claw.
Measurements of & holotype (mm). - Length pro-
boscis (dorsal) 1.04; greatest diameter proboscis 0.45;
greatest diameter crop 0.91; length first trunk segm.
1.61; length second trunk segm. 0.80; length third
trunk segm. 0.78; length fourth trunk segm. (to tip
abdomen) 1.24; width across second lateral processes
1.57; length scape 1.41; length chela 0.91.
Leg 3: first coxa 0.63; second coxa 1.92; third coxa
121
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 27-34. Nymphon bullatum sp. n. (from Stn. MBT 149). 27, trunk, d (scale F); 28, ocular tubercle, ® (free-hand sketch);
29, ocular tubercle, 2, from the left (free-hand sketch); 30, oviger, ? (B); 31, leg 3, d (F); 32, leg 3, ® (F); 33, cement gland
apertures of leg 3, 4 (B); 34, distal segments of leg 3, $ (B). Scales after fig. 80.
122
Jan H. Srock: Brazilian Pycnogonida
Figs. 35-41. Nymphon inaequipes sp. n. (from Stn. MBT 142). 35, trunk, ©, dorsal (scale F); 36 ocular tubercle, 2, from the
left (free-hand sketch); 37 ocular tubercle, 2, frontal (free-hand sketch); 38, leg 1, 5 (F); 39, leg 3, d (F); 40, leg 4, 2 (B;
41, cement gland apertures of leg 1, & (free-hand sketch). Scales after fig. 80.
0.43; femur 2.74; first tibia 3.59; second tibia 4.05;
tarsus 0.95; propodus 1.07; claw 0.58; auxiliary claws
0.15.
Etymology. - The specific name is derived from
bulla (Latin = knob, bump) and alludes to the apoph-
ysis on oviger segment 5 in the male.
Remarks. - Two outstanding features of the male
differentiate this species from the 200-odd others in
the genus: (1) the cement glands discharging through
volcano-shaped prominences; and above all (2) the
apophysis on oviger segment 5 which is not apical (as
in many other Nymphon species) but localized at two-
thirds of the segment.
Cement gland apertures similar to those of the new
species, are known from two other species only: N.
discorsicoxae Child, 1982 and N. caldarium Stock,
1988. The former, however, is a brevitarsal species
(whereas N. bullatum is longitarsal) and the latter is
uniunguiculate (whereas N. bullatum possesses auxil-
iary claws).
Morphologically closest to N. bullatum are perhaps
N. sandersi Child, 1982 (from off Rio de la Plata,
Argentina, 3815-4435 m) and N. longicollum Hoek,
1881 (off Valparaiso, Chile, 2225 fms = 4072 m).
Since some of the most useful diagnostic characters in
the genus Nymphon are sex-linked (male oviger and
male cement glands), it is hard to judge about the re-
lationships of these two species, of which the (adult)
male is unknown. Both differ from N. bullatum in
that the auxiliary claws are tiny or lacking, and the oc-
ular tubercle is slender and conical, without eyes.
Some additional differences berween N. sandersi and
N. bullatum are: (1) palp segm. 2 = 3 in sandersi, 2 >
3 in bullatum; (2) oviger claw with 3 teeth in sander-
si, 6-7 in bullatum; (3) compound oviger spines with
5 pairs of teeth in sandersi, 2-3 in bullatum. In N. lon-
gicollum (vide Gordon 1932a) the chela is straighter
and the palm is longer than in N. bullatum, and each
finger bears 11-12 teeth only (14-19 in bullatum).
123
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Nymphon inaequipes sp. n.
(figs. 35-46)
Type material. - 1 & holotype, 1 9 allotype, 23 para-
types, Project Isotan-Draga I (PID), Stn. MBT 142: 23°46'S
43°00'W, 150 m, 2 Sep 1970 (zma Pa. 3445).
Other material. — 1 9, rip, Stn. MBT 94: 25°56'S
46°42'W, 124 m, 18 June 1970; 2 ©, rip, Stn. MBT 139:
30°52'S 49°51'W, 126 m, 27 June 1970; 1 9, pip, Stn. MBT
148: 23°19'S 41°57'W, 136 m, 3 Sep 1970; 7 spms., PID,
Stn. MBT 151: 23°36'S 41°23'W, 475 m, 4 Sep 1970; 1 9,
PID, Stn. MBT 152: 23°03'S 41°10'W, 98 m, 4 Sep 1970; 2
3,1 9, pip, Stn. MBT 194: 24°27'S 44°57'W, 26 m, 1 June
1971; 1 d, 2 juvs., PID, Stn. MBT 195: 24°03'S 44°59'W, 73
m, 1 June 1971; 1 6, 1 9, rip, Stn. MBT 196: 23°31'S
44°57'W, 38 m, 1 June 1971; 208 spms., PıD, Stn. MBT 197:
23°47'S 44°44'W, 65 m, 1 June 1971; 5 spms., Pip, Stn.
MBT 198: 24°03'S 44°30'W, 124 m, 2 June 1971; 47 spms.,
PID, Stn. MBT 206: 23°14'S 44°03'W, 52 m, 3 June 1971; 4
spms., PID, Stn. MBT 208: 23°20'S 43°26'W, 78 m, 4 June
1971; 34 spms., PID, Stn. MBT 211: 23°59'S 43°01'W, 220
m, 4 June 1971; 1 ®, R. V. ‘Prof. W. Besnard’, Stn. RIG
591: 24°27'S 44°00'W, 500 m, 8 Feb 1969; 2 juv., R. V.
‘Prof. W. Besnard’, Stn. rıG 1330: 23°13'S 44°06'W, 40 m,
16 Feb 1968.
Description. - Trunk completely segmented,
naked. Lateral processes long, slender, separated by
intervals more than twice as wide as diameter of each
lateral process. Neck widened at anterior end, fol-
lowed by long, narrow, parallel-sided part. Oviger
implantation in contact with anterior margin of first
lateral process; on dorsal side of neck, right and left
implantations fused (looking like segmentation line,
but such line absent on ventral surface of neck).
Ocular tubercle mainly cylindrical, rather low;
rounded point at distal end; eyes small, well-pigment-
ed. Abdomen short (in dorsal view not extending be-
yond 4th lateral processes), directed obliquely up-
ward and backward (angle with horizontal ca. 75°).
Proboscis swollen at basal third, and again at distal
third, shorter than neck.
Chelifore scapes strongly diverging, practically un-
armed. Chela much longer than scape. Fingers longer
than palm, curved near tip. Each finger with numer-
ous (ca. 55) unequal teeth: a larger tooth is alternating
with 2 to 4 smaller teeth (some of the smaller teeth bi-
fid); larger teeth of movable finger shorter and more
obtuse, on immovable finger larger and pointed. In
juveniles, number of teeth on fingers lower.
Palp segms. 2 to 5 very slender, in proportions
9:7:8:6. Long setae in particular on segms. 4 and 5.
Oviger of male: Segm. 5 almost straight, by far
longer than segm. 4, with row of recurved spinules on
ventral margin and with small distal apophysis. Segm.
6 curved, about one-third of length of segm. 5, with
inconspicuous tubercle on dorsal margin. Compound
spines, according to formula 8:6:6:6, with 3 large ba-
sal teeth and 3 much smaller distal teeth on either
margin. Distal claw long, slender, with 7 teeth.
124
Oviger of female: Much weaker than that of male,
segm. 5 not so much longer than segm. 4, straight,
without recurved spinules. Segm. 6 straight. Spine
formula 10:7:6:8::10.
Legs long and thin. Coxae almost naked; coxa 2 of
leg 1 less elongate than that of P2-P4. Femur < tibia
1; tibia 2 slightly longer than tibia 1. Several rather
short setae, in particular on tibiae. Leg 1 having distal
segms. different from those of other legs (heteropo-
dy); in leg 1 tarsus > propodus, both densely set with
small spinules of uniform size on ventral surface;7
claw of leg 1 short (ca. 25% of length of propodus);
auxiliary claws almost 90% of length of main claw. In
remaining legs, tarsus < propodus, sparsely armed
with small spinules; claw long (about 50% of length
of propodus); auxiliary claws about 80% of length of
main claw. Heteropody already visible in juveniles.
Femur of female strongly swollen in basal half, for
conception of eggs. Femoral gland pores in male in-
conspicuous; a bulbous, subcutaneous vesicle opens
through a longish, narrow, subcutaneous duct into a
minute pore; 8 or 9 glands present on each leg. Sexual
openings of female on distoventral end of coxa 2 of all
legs.
Measurements (mm) of female from Stn. MBT 142.
— Length proboscis (dorsal) 0.93; greatest diameter
proboscis 0.45; length first trunk segm. 1.47; length
second trunk segm. 0.51; length third trunk segm.
0.53; length fourth trunk segm. (to tip fourth lateral
process) 0.59; width across second lateral processes
1.40; length scape 0.72; length chela 1.63. First leg:
tarsus 0.71; propodus 0.64; claw 0.16; auxiliary claws
0.14. Third leg: first coxa 0.33; second coxa 1.40;
third coxa 0.39; femur 2.03; first tibia 2.77; second
tibia 2.85; tarsus 0.64; propodus 0.96; claw 0.49;
auxiliary claws 0.40.
Etymology. — The specific name, inaequipes (Latin,
with unequal legs) alludes to the morphological dif-
ferences between leg 1 and the remaining legs.
Distribution. — This is a rather common, some-
times even abundant species in our samples. It has
been found in depths between 26 and 500 m.
Remarks. - The present species is rather similar to
the next, N. vulcanellum. Look-alikes must be sought
amongst such taxa as N. macrum Wilson, 1880, N.
gracillimum Calman, 1915, and N. hiemale Hodgson,
1907.
N. inaequipes differs from these as follows:
— From N. vulcanellum by the cement gland aper-
tures which are flush with the femoral surface (not
raised), the fingers of the chela which bears >50 teeth
of unequal size (versus <26 teeth of uniform size), the
longer auxiliary claws, and the heteropodous nature
of the legs. In mixed collections of N. vulcanellum
and N. inaequipes, which are common, the latter can
be distinguished at first glance by its smaller size,
JAN H. Stock: Brazilian Pycnogonida
Figs. 42-46. Nymphon inaequipes sp. n. (from Stn. MBT 142). 42, chela, (scale B); 43, palp, 3 (B); 44, oviger, d (A); 45,
distal part of oviger, d (G); 46, oviger, 9 (A).
Figs. 47-51. Nymphon bullatum sp. n., 3, from Stn. MBT 149. 47, chela (scale B); 48, palp (B); 49, oviger (B); 50, oviger claw
(free-hand sketch); 51, compound spines of oviger segment 10 (free-hand sketch). Scales after fig. 80.
125
TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
longer neck, and shorter, erect abdomen.
— From North Atlantic N. macrum by the second
tibia which is hardly longer than the first (versus 1.3
to 1.5 times as long as tibia 1), the oviger implanta-
tion which is in contact with the first lateral process
(well in front of it in N. macrum), the number of
compound oviger spines, viz. a total of 25-31 (versus
> 60 in macrum), and the number of teeth on the ovi-
ger claw ( <11, versus ca. 19 in macrum).
— From N. gracillimum and N. hiemale (which
might be synonymous, see Gordon, 1932b: 42), from
cold to polar waters of the Southern Hemisphere, by
the presence of a distal apophysis (though small) on
oviger segment 5 of the male in inaequipes, the propo-
dal armature (sole with uniform spinules in inae-
quipes, versus unequal in the other two), the subequal
tibiae of inaequipes (versus second tibia 1.4-1.5 times
as long as the first), the higher number of teeth on the
fingers of the chela ( >50 versus <36), and the longer
auxiliary claws.
In none of these species heteropody is recorded. In
fact, this is a rather rare phenomenon, more usually
found in unrelated genera, such as Ascorhynchus.
Nymphon vulcanellum sp. n.
(figs. 52-61)
Nymphon spec. Stock 1966: 392-393, fig. 3a-d.
Type material. - 1 d (ovig.) holotype, 1 9 allotype, 9 4
(partly ovig.), 2 9, 39 juvs and larvae (paratypes), Project
Isotan Draga I (Pip), Stn. MBT 148: 23°19'S 41°57'W, 136
m, 3 Sep 1070 (zma Pa. 3447).
Other material. - 14 larvae (probably this species), PID,
Stn. MBT 111: 27°22'S 47°27'W, 140 m, 21 June 1970; 1
®, pip, Stn. MBT 116: 27°59'S 47°31'W, 150 m, 22 June
1970; 1 juv., 2 larvae, Pip, Stn. MBT 141: 23°25'S 43°00'W,
113 m, 2 Sep 1970; 3 2, 19 juvs. & larvae, PID, Stn. MBT
142: 23°46'S 43°00'W, 150 m, 2 Sep 1970; 1 d (ovig.), 2
QE) juvs., PID, Stn. MBT 145: 23°25'S 42°27'W, 130 m, 3
Sep 1970; 1 d, 9 larvae, Pip, Stn. MBT 149: 23°41'S
41°55'W, 250 m, 4 Sep 1970; 1 juv., rip, Stn. MBT 152:
23°03'S 41°10'W, 98 m, 4 Sep 1970; 5 larvae, pip, Stn. MBT
156: 22°06'S 40°53'W, 32 m, 5 Sep 1970; 30 juvs. & lar-
vae, PID, Stn. MBT 161: 21°42'S 40°15'W, 56 m, 6 Sep
1970; 1 spm. (fragm.), rip, Stn. MBT 173: 25°32'S
47°11'W, 55 m, 27 May 1971; 1 juv., Pip, Stn. MBT 195:
24°03'S 44°59'W, 73 m, 1 June 1971; 2 ®, Pip, Stn. MBT
197: 23°47'S 44°44'W, 65 m, 1 June 1971; 1 larva, PID,
Stn. MBT 210: 23°46'S 43°10'W, 128 m, 4 June 1971; 4
juvs., PID, Stn. MBT 211: 23°59'S 43°01'W, 220 m, 4 June
1971; 1 2, R. V. ‘Prof. W. Besnard’ Stn. 1490: 22°08'S
41°00'W, 21 m, 12 Mar 1971.
Description. - Trunk completely segmented, prac-
tically unarmed. Lateral processes > diameter trunk,
intervals slightly larger than diameter of lateral pro-
cess. Anterior part of neck strongly widened; short
parallel-sided basal part. Ocular tubercle truncate,
with 2 minute lateral, and 1 terminal tubercle; eyes
126
present. Oviger implantation in contact with anterior
margin of first lateral process. Abdomen long, reach-
ing beyond middle of coxa 1 of leg 4, almost horizon-
tal. Proboscis barrel-shaped, short.
Chelifore scapes strongly divergent, poorly armed.
Chela longer than scape. Fingers not in same plane as
palm, longer than palm. Movable finger curved at tip,
immovable finger regularly curved; each finger with
20-25 teeth, those of immovable finger longer than
those of movable finger.
Palp segm. 2 > 3 > 4 > 5; segms. 3 and 4 unusually
slender.
Oviger of male: Segm. 5 >> 4, somewhat S-shaped,
distally swollen. Segm. 6 most remarkable, because
showing a subdivision into 2 segments (segmentation
more or less distinct, sometimes looking functional,
see arrow in fig. 58). Compound spines with 3, rarely
4, marginal teeth; formula 12:8:6:6. Claw with 6
teeth.
Oviger of female: Segm. 5 straight, slightly longer
than 4; segm. 6 not subdivided. Spine formula
5:6:6: 72:5).
Legs: Femur of female slightly swollen in proximal
part. Femur of male with some 16 cement gland aper-
tures on ventral margin of all legs. Apertures shaped
like low, truncate volcanoes. Coxa 2 and both tibiae
with some long setae. Femur < tibia 1; tibia 1 slightly
longer than tibia 2. Tarsus slightly longer than propo-
dus; ventral margin of tarsus and propodus densely
packed with minute spinules of uniform size (no heel,
no heel spines). Claw short; auxiliary claws about half
length of main claw. Legs homoiopodous.
Measurements of 9 from Stn. MBT 142 (mm). —
Length proboscis (dorsal) 1.24; greatest diameter pro-
boscis 0.64; length first trunk segm. 1.59; length sec-
ond trunk segm. 0.87; length third trunk segment
0.82; length fourth trunk segment (to tip fourth lat-
eral process) 1.05; width across second lateral process-
es 2.15; length abdomen 0.91.
Leg 3: first coxa 0.90; second coxa 1.64; third coxa
0.63; femur 3.78; first tibia 5.29; second tibia 5.12;
tarsus 1.40; propodus 1.11; claw 0.40; auxiliary claws
0.20.
Etymology. — The specific name, vulcanellum
(Latin = little volcano) alludes to the low, volcano-
shaped cement gland apertures on the femur of the
male.
Remarks. - This is no doubt the same species re-
corded as “Nymphon spec.’ by Stock (1966), from one
of the ‘Calypso’ stations off the South American
Atlantic coast. In 1966, though it was recognized that
probably a new species was concerned, it was not
named because only a female was available. Now, the
male being discovered, I have no hesitation any more
to attribute this highly characteristic animal to a new
species.
Jan H. Srock: Brazilian Pycnogonida
55:
53
&
Figs. 52-56. Nymphon vulcanellum sp. n. (3 from Stn. MBT 149, 2 from Stn. MBT 142). 52, trunk, 9, dorsal (scale F); 53
ocular tubercle, 3, frontal (free-hand sketch); 54 chela, 9 (A); 55, leg 3, & (F); 56, femur, 9 (F). Scales after fig. 80.
127
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
By the large chela, the absence of propodal heel
spines, and the slender palp segments, the new species
looks like the North Atlantic N. macrum Wilson,
1880. Obvious differences are, apart from the volca-
no-shaped cement gland apertures, the ratio tibia
1/tibia 2 (subequal in the new species, tibia 2 by far
the longest in N. macrum), the low number of denti-
cles on the oviger claw, and the low number of margi-
nal teeth on the compound oviger spines of the new
species. Finally, the oviger implantation is in contact
with the first lateral process in the new species, im-
planted well in front of it in N. macrum.
For the differences from N. inaequipes, consult the
remarks under the latter.
Distribution. - Found often together with N. inae-
quipes (vide supra), but is less abundant. Its bathy-
metrical range lies between 21 and 250 m.
Family Callipallenidae
Callipallene evelinae Marcus, 1940
C. evelinae Marcus, 1940b: 31-33, fig. 2 A-D.
Material. — 1 juv., usp, Stn. F 10: Rocky intertidal of
Ponta de Santa Rita, Enseada do Flamengo, Ubatuba, State
of Sao Paulo, on Dyctiota sp., 12 Dec 1965; 2 ®, Project
Isotan-Draga I (PID), Stn. MBT 145: 23°25'S 42°27 W, 130
m, 3 Sep 1970; 1 9, PID, Stn. MBT 155: 22°22'S 41°32'W,
34 m, 5 Sep 1970.
Remarks. - C. evelinae is mainly characterized by
male characters (presence of plumose setae on distal
leg segments, absence of lobe on 5th oviger segment).
Thus, the identification of the present material (lack-
ing males) must be taken with some reservation.
However, nothing in the morphology seems to pre-
vent identification with C. evelinae, previously known
only from Ilha das Palmas, Bahia de Santos, 0-5 m.
The present records extend the range to east of Rio de
Janeiro, and to a depth of 130 m.
Callipallene gabriellae Corrêa, 1948
C. gabriellae Corrêa, 1948: 1-4, figs. 1-6.
Material. - 76 spms. (incl. ovigerous males and juve-
niles), usp, Stn. P.E.: Collected by hand by divers in the in-
fralittoral zone on seaweeds growing on rock, Ponta do
Espia, Ubatuba, State of Säo Paulo, 20 Sep 1964.
Remarks. - This species is extremely close to C.
emaciata (Dohrn, 1881), known from the
Mediterranean and from New England to the West
Indies. Child (1979: 41) examined specimens from
the entire range of C. emaciata and made some obser-
vations on its morphology. In the configuration of
the propodal spines and the auxiliary claws, the
present material agrees with Child's observations.
128
Evidently, the shape of the trunk, oviger spines, pro-
portions of the leg segments, curvature of the propo-
dus, and length of the auxiliary claws, C. gabriellae
and C. emaciata are practically identical. Corrêa used
the presence or absence of a suture between trunk seg-
ments 3 and 4 as a key character to separate the two
species, but in the large sample from Brazil at hand,
this feature proved to be too variable to be reliable.
The only characters that made me decide to retain
C. gabriellae as a distinct species, are its more swollen
male(!) femur and the number of teeth on the fingers
of the chela. Child (1979, in key) records about 15
teeth on each finger of C. emaciata, whereas in the
present material of C. gabriellae there are 6-9 teeth.
The number of propodal heel spines, considered to be
diagnostic character by Child, varies between 4 and 5
in C. gabriellae.
C. gabriellae has been described from the upper lit-
toral zone of Ilha das Palmas, Bahia de Santos
(Brazil). As far as I know, no further specimens have
ever been recorded.
Callipallene margarita (Gordon, 1932)
Pallene margarita Gordon, 1932b: 82-85, figs. 40-41.
Callipallene margarita; Marcus 1940b: 30; Corrêa 1948: 6;
Child 1982a: 26.
? Callipallene margarita, Hedgpeth 1961: 3, fig. 1.
Material. - 1 d, 3 juvs., Project Isotan-Draga I, Stn. MBT
149: 23°41'S 41°55'W, 250 m, 4 Sep 1970.
Remarks. - The formula of the compound oviger
spines in the present male is 8:6:8:8, which is lower
than the number recorded by Gordon (1932b), but
about similar to that found by Hedgpeth (1961).
The species is known from South Georgia, N. of
the Falkland Islands, 120-320 m (Gordon 1932b);
southern Chile, 70 m (Hedgpeth 1961), and from the
Argentine slope, between 36°53'S and 38°13'S, 454-
2323 m (Child 1982a). The present record extends
the range of the species further north to 23°41'S, but
falls within the bathymetrical range.
Callipallene producta (Sars, 1888)
C. producta; Stock 1990: 227 [refs.].
Material. - 1 d, Project Isotan-Draga I, Stn. MBT 145:
23°25'S 42°27'W, 130 m, 3 Sep 1970.
Remarks. - This specimen resembles closely Sars’
(1891) figures based on material from northern
Europe, except that the second tibia is provided with
some long setae. This species is mainly distributed in
the northern and central Atlantic, from Norway to
the Cape Verde Islands. It is new to Brazil.
Jan H. Stock: Brazilian Pycnogonida
Figs. 57-61. Nymphon vulcanellum sp. n. (6 from Stn. MBT 149, © from Stn. MBT 142). 57, palp, 9 (scale A); 58, oviger, d
(A); 59, proximal part of oviger, 2 (A); 60, compound spine of oviger segment 10, & (free-hand sketch); 61, terminal oviger
claw, d (free-hand sketch). Scale after fig. 80.
Pallenoides amazonica Stock, 1975
P. amazonica Stock, 1975a: 1012-1015, figs. 23-24; see also
Arnaud & Child 1988: 142-143.
Material. — 1 juv., Project Isotan-Draga I (PID), Stn. MBT
148: 23°19'S 41°57'W, 136 m, 3 Sep 1970; 1 9, 2 juvs.,
PID, Stn. MBT 157: 21°46'S 40°58'W, 15 m, 5 Sep 1970; 2
9, 2 juvs., rip, Stn. MBT 163: 21°29'S 40°56'W, 17 m, 6
Sep 1970; 1 d ovig., 3 2, 2 juvs., PID, Stn. MBT 165:
21°20'S 40°39'W, 28 m, 6 Sep 1970.
Remarks. — The eggs carried by the ovigerous male
from Stn. MBT 165 are, in preserved state, of a deep-
brown colour.
The species was described from Brazilian shelf wa-
ters, but much further north (between 00°31 N and
02°15'S). Arnaud & Child's South African record
(1988) based on a larva, is ‘certainly questionable’
(1988: 143).
Family Phoxichilidiidae
Pallenopsis (Pallenopsis) boehmi Schimkewitsch,
1930
P. boehmi Schimkewitsch, 1930: 236-237 [new name for P.
fluminensis, sensu Böhm 1879, nec Kroyer 1844]. —
Marcus 1940a: 190-191; Marcus 1940b: 22; Stock 1973:
349-350, figs. 1-10 [redescr. of types]; Stock 1975a: 1018
[in key].
P. longicoxa Stock, 1966: 394-396, fig. 4 [syn. n.].
Material. — 1 d,R. V. ‘Prof. W. Besnard’, Stn. RIG 588:
23°50'S 44°16'W, 100 m, 7 Feb 1968; 2 6, 1 9, R. V.
‘Prof. W. Besnard’, Stn. RG 1333: 23°13' 44°18'W, 35 m,
17 Feb 1968; 2 4, 3 9, R. V. Prof. W. Besnard’, Stn. 1477:
23°29'S 42°54'W, 105 m, 9 March 1971; 1 ®, Project
Isotan-Draga I, Stn. MBT 153: 22°48'S 41°14'W, 65 m, 4
Sep 1970.
Remarks. — P. boehmi was based on females (see
Stock 1973), whereas P. longicoxa was based on
males. Some of the present samples contain both sex-
es, and prove boehmi to be a senior synonym of longi-
coxa, as already suspected by Stock (1973). The
length of the auxiliary claws is variable in these sam-
ples: in some specimens they are some 40% of the
length of the main claw, in others they are fully half as
long.
P. boehmi is predominantly a cold-water species,
distributed from Magellan Strait to southern Brazil.
One of the above stations, Stn. 1477, is located very
close to the type-locality of P. longicoxa.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Pallenopsis (Pallenopsis) candidoi de Mello-Leitäo,
1949
P. (P.) candidoi, Stock, 1975a: 1030, fig. 31 a-b [refs.].
Material. - 1 d, 1 2, R. V. ‘Prof. W. Besnard’, Stn. RD
54: 18°54.8'S 39°15.4'W, 41 m, 12 July 1972.
Remarks. - This species is known from southern
Brazil to Surinam, between 15 and 65 m.
Pallenopsis (Pallenopsis) fluminensis (Kroyer, 1844)
P. (P.) fluminensis; Stock 1975a: 1020-1021, fig. 25 [refs.].
Material.- 1 d,R.V. Prof. W. Besnard’, Stn. RG 1325:
23°14'S 43°56.5 W, 49 m, 16 Feb 1968.
Remarks. - A Brazilian species, known from the
State of Säo Paulo to N. of Pernambuco. Depth range
0-49 m.
Pallenopsis (Pallenopsis) kempfi Stock, 1975
P. (P.) kempfi Stock, 1975a: 1025-1028, figs. 28-29; Stock
1986: 427-429, fig. 8 a-f.
Material. - 1 36, R. V. Prof. W. Besnard’, Stn. 1848:
30°22'S 48°41'W, 178 m, 3 Aug 1972.
Remarks. — Previously recorded from Florida (ca.
24°36'N) to Brazil (ca. 04°13'S). The present
records forms a significant southward extension of
the range.
Pallenopsis (Pallenopsis) tumidula Loman, 1923
P. (P.) tumidula Loman, 1923: 32-34, fig. G; Marcus
1940a: 191; Marcus 1940b: 24 [in key]; Stock 1957:
103-105, fig. 18; Stock 1966: 393, fig. 3e; Stock 1975a:
1018 [in key].
Material. — 1 d ovig., 2 2, R. V. ‘Prof. W. Besnard’, Stn.
RG 1332: 23°14.3'S 44°13.3'W, 42 m, 17 Feb 1968; 1 &
ovig., 2 2, R. V. ‘Prof. W. Besnard’, Stn. 1883: 30°22'S
48°41'W, 178 m, 3 Aug 1972.
Remarks. - Previously recorded from off La Plata
(ca. 37°30'S). The present records extend the range to
southern Brazil (23°14'S).
Anoplodactylus aragaoi Sawaya, 1950
(figs. 63-64)
A. aragaoi Sawaya, 1950: 63-67, 77-80, pls. I-II.
Material. - 1 d ovig., R. V. ‘Prof. W. Besnard’, Stn. RD
56: 18°54.7'S 38°53.5'W, 38 m, 12 July 1972.
Remarks. - The chelifore and chela were not illus-
trated in great detail before; therefore some figures of
this appendage are incorporated in the present paper.
The single specimen collected agrees in every detail
130
with Sawaya's description. Previously recorded from
20°33'S 40°14'W, 35 m.
Anoplodactylus brasiliensis Hedgpeth, 1948
A. brasiliensis Hedgpeth, 1948: 224 [pro A.pygmaeus sensu
Marcus 1940b: 63-65, Pl. VI fig. A-D, nec Hodge,
1864].
Material. — 2 2, 6 juvs. & larvae, usp, Stn. P.E.: Ponta do
Espia, Ubatuba, State of Sao Paulo, on seaweeds growing on
rocky shore, infralittoral, 20 Nov 1964.
Remarks. — I agree with Lebour (1945) and
Hedgpeth (1948) in considering this south Brazilian
shallow water species different from A. pygmaeus,
under which name it was recorded by Marcus
(1940b). Hedgpeth coined the name A. brasiliensis
for it, which is adopted here. The species differs from
A. pygmaeus by the presence of small auxiliary claws,
and the absence of a tubercle on the lateral processes
(a single spinule is present, however). The propodal
sole has a rather short lamina, preceded by 3 or 4 sole
spines. Unfortunately, the present collection contains
no males, so I can add nothing to Marcus’ descrip-
tion.
Anoplodactylus evelinae Marcus, 1940
A. evelinae, Child 1982b: 368-369 [refs.].
Material. - 24 spms., usp, Stn. P.E.: Ponta do Espia,
Ubatuba, State of Sao Paulo, on seaweeds growing on rocky
shore, infralittoral, 20 Nov 1964; 1 2, usp, Stn. F 10: Ponta
Santa Rita, Enseada do Flamengo, Ubatuba, State of Sao
Paulo, on intertidal algae (Dyctiota sp.) on rocky shore, 12
Dec 1965; 1 ©, usp, Stn F 27: Rocky shore between Praia
do Dionisio and Praia do Flamengo, Enseada do Flamengo,
Ubatuba, State of Sao Paulo, on algae slightly exposed at low
tide, 15 Feb 1965.
Remarks. - The species was described from the
same general area; later records extended the range to
shallow waters of the West Indies.
Anoplodactylus insignis (Hoek, 1881)
A. insignis, Stock 1975a: 1056-1058, fig. 44 [refs.]; Stock,
1986: 437-438, fig. 15 a-b.
Material. - 1 d (fragm.), Project Isotan-Draga I, Stn.
MBT 165: 21°20'S 40°39'W, 28 m, 6 Sep 1970; 1 9, R. V.
‘Prof. W. Besnard’, Stn. RD 54: 18°54.8'S 39°15.4' W, 41
m, 12 July 1972.
Remarks. - Originally described by Hoek from
Brazil (Bahia, 13-37 m). Later records show it to have
a vast range in the Caribbean and on the American
east coast up to North Carolina. The female from
Stn. RD 54 belongs to var. calcaratus Stock, 1986, pre-
viously found in the West Indies, Venezuela and the
Guyanas.
Jan H. Stock: Brazilian Pycnogonida
Fig. 62. Anoplodactylus stictus Marcus, 1940 (® from Stn. MBT 165), chela (scale E). Figs. 63-64. Anoplodactylus aragaoi
Sawaya, 1950 (4 from Stn. rp 56). 63, chelifore (scale B); 64, chela (E). Scales after fig. 80.
Anoplodactylus maritimus Hodgson, 19 14
A. maritimus, Stock 1975a: 1072, 1074, fig. 54 [syn. &
distr.].
Material. - 1 ©, R. V. ‘Prof. W. Besnard’, Stn. RG 1331:
23°14.7'S 44°09.4'W, 45 m, 16 Feb 1968; 2 6, Project
Isotan-Draga I (pip), Stn. MBT 153: 22°48'S 41°14'W, 65
m, 4 Sep 1970; 2 6, Pip, Stn. MBT 164: 21°15'S 40°50'W,
19 m, 6 Sep 1970; 2 5, 4 9, Pip, Stn. MBT. 184: 24°33'S
46°50'W, 30 m, 30 May 1971; 4 &,2 9, 2 juvs., Pip, Stn.
MBT 195: 24°03'S 44°59'W, 73 m, 1 June 1971; 15 gd, 12
2, pip, Stn. MBT 206: 23°14'S 44°03'W, 52 m, 3 Jun
1971"
Remarks. - This species, often associated with
floating Sargassum, is widely distributed in the (sub)-
tropical Atlantic. It is recorded here from the first
time from Brazil.
Anoplodactylus monotrema Stock, 1979
A. monotrema; Child 1982b: 372 [refs.].
Material. - 1 5,1 9,11 juv., usp, Stn. F 10: Ponta
de Santa Rita, Enseada do Flamengo, Ubatuba, State
of Sao Paulo, on intertidal algae (Dyctiota sp.) on
rocky shore, 12 Dec 1965; 1 6, R. V. ‘Prof. W.
Besnard’, Stn. RD 54: 18°54.8'S 39°15.4'W, 41 m, 12
July 1972.
Remarks. - The males in this collection have one
slit-like cement gland aperture on the legs, and agree
also otherwise with the original description. The fe-
male from Stn. F 10 has more ‘angular’ corners of the
proboscis, and thus resembles A. robustus (Dohrn,
1881), but has well-spaced lateral processes 1 and 2 as
in A. monotrema.
These records confirm the presence of A. monotre-
ma on the coasts of Brazil, already anticipated by
Stock (1979).
Anoplodactylus petiolatus (Kroyer, 1844)
A. petiolatus, Stock 1975a: 1072-1075, fig. 53 [incl. A. lon-
gicollis (Dohrn, 1881)].
A. guyanensis Child, 1977 [syn. n.].
Material. - 8 d (of which 1 ovig.), 2 2, Project Isotan-
Draga I (erp), Stn. MBT 50: 23°39'S 45°06'W, 28 m, 8 May
1970; 1 2, ip, Stn. MBT 77: 23°39'S 43°09'W, 128 m, 12
May 1970; 4 d, 1 @, PID, Stn. MBT 101: 25°51'S 47°54 W,
41 m, 19 June 1970; 1 d, rip, Stn. MBT 106: 26°45'S
47°16'W, 130 m, 20 June 1970; 9 spms., PID, Stn. MBT 113:
27°19'S 48°15'W 54 m, 21 June 1970; 3 d, 2 9, Pip, Stn.
MBT 116: 27°59'S 47°31'W, 150 m, 22 June 1970; 1 9,
PID, Stn. MBT 127: 29°33'S 48°46'W, 118 m, 25 June 1970;
1 2, Pip, Stn. MBT 128: 29°23'S 49°10'W, 54 m, 25 June
1970; 1 6, rip, Stn. MBT 141: 23°25'S 43°00'W, 113 m, 2
Sep 1970; 5 8,1 ?, PID, Stn. MBT 145: 23°25'S 42°27'W,
130 m, 3 Sep 1970; 3 dg, Pip, Stn. MBT 147: 23°01'S
41°59'W, 66 m, 3 Sep 1970; 1 4,3 9, 1 juv., PID, Stn. MBT
148: 23°19'S 41°57'W, 136 m, 3 Sep 1971; 1 d, 3 juvs.,
PID, Stn. MBT 152: 23°03'S 41°10'W, 98 m, 4 Sep 1970; 5
3, 2 2, 8 juvs., 1 larva, PID, Stn. MBT 161: 21°42'S
40°15'W, 56 m, 6 Sep 1970; 1 d, pip, Stn. MBT 164:
21°15'S 40°15'W, 19 m, 6 Sep 1970; 6 ®, pip, Stn. MBT
171: 25°05'S 47°40'W, 20 m, 27 May 1971; 16 spms., PID,
Stn. MBT 173: 25°32'S 47°11'W, 55 m, 27 May 1971; 1 dg,
3 ®, pip, Stn. MBT 185: 24°12'S 46°09'W, 38 m, 30 May
1971; 4 d, 3 2, 2 juvs., Pip, Stn. MBT 186: 24°28'S
45°54'W, 59 m, 30 May 1971; 1 d, pip, Stn. MBT 195:
24°03'S 44°59'W, 73 m, 1 June 1971; 5 3, 3 Q, pip, Stn.
MBT 197: 23°47'S 44°44'W, 65 m, 1 June 1971; 2 d,1 9,
PID, Stn. MBT 210: 23°46'S 43°10'W 128 m, 4 June 1971;
1 d, R. V. ‘Prof. W. Besnard’ (RvwB), Stn. RIG 588: 23°50'S
44°16'W, 100 m, 7 Feb 1969; 2 2, RvwB, Stn. RD 4:
19°59'S 39°56'W, 41 m, 6 July 1972; 1 4, RvwB, Stn. RD
5: 19°54'S 39°50'W, 51 m, 6 July 1972; 1 à ovig., RVWB,
Stn. RD 50: 19°02.5'S 39°30.6'W, 28 m, 12 July 1972; 1 d,
1 2, RvwB, Stn. RD 54: 18°54.8'S 39°15.4'W, 41 m, 12 July
1972; 1 2, RvwB, Stn. RD 71: 18°34'S 39°14.6'W. 35 m, 14
July 1972; 1 d, rvws, Stn. RD 74: 18°34'S 38°38'W, 59 m,
14 July 1972; 1 d, RvwB, Stn. RD 82: 19°55'S 39°54'W, 46
m, 16 July 1972; 1 d ovig., Project Petrobas/usp, Stn. RD
36: 19°18.1'S 39°39.4'W, 16 m, 10 July 1972.
131
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Remarks. - The present material gave me consider-
able trouble. After re-examination of many samples
from various origins (North Sea, Bay of Biscay, Israel,
the Guyanas), I came to the conclusion that A. petio-
latus is a variable species and that the variation occurs
in a geographically unordered way. North Sea speci-
mens have the lowest number of spines on the propo-
dal sole in front of the lamina (viz. 3 to 6), medium-
sized tubercles on the lateral processes, and no
tubercle on coxa 1. Guyana specimens have 4 to 8
spines on the propodal sole, very tall tubercles on the
lateral processes, and sometimes a distinct tubercle on
coxa 1. Finally Brazilian specimens have the highest
number (4 to 10) of spines on the propodal sole, me-
dium-sized tubercles on the lateral processes, and a
vestigial tubercle on coxa 1.
The Bay of Biscay specimens look very much like
A. guyanensis Child, 1977, but the abdomen is slight-
ly longer and less erect. Israel specimens look strong-
ly like A. vemae Child, 1982, but oviger segment 2 is
shorter. The length of the cement gland tube and the
trunk segmentation can vary as well. In our Brazilian
collection, the tube is usually rather short, and speci-
mens with an unsegmented trunk are more common,
but other specimens show a segmentation line
between trunk segments 1 and 2, or between 1, 2 and
3. Comments on the variability of À. petiolatus are al-
ready briefly provided in the papers by Stock (1966,
1975a). For these reasons, I consider A. petiolatus a
variable species, and has united A. longicollis (Dohrn,
1881) from the Mediterranean and A. guyanensis
Child, 1977 with A. petiolatus (Kroyer, 1844).
A. petiolatus was recorded before from Brazil
(Marcus, 1940b) and northern Argentina (Stock,
1966).
Anoplodactylus spurius sp. n.
(figs. 65-75)
Type material. - 1 d holotype, 1 juv., Project Isotan-
Draga I (erp), Stn. MBT 211: 23°59'S 43°01'W, 220 m, 4
June 1971 (zma Pa. 3448); 1 ®, 3 juvs. (paratypes), PID,
Stn. MBT 142: 23°46'S 43°00'W, 150 m, 2 Sep 1971 (zma
Pa. 3449); 3 & (paratypes), Pip, Stn. MBT 145: 23°25'S
42°27'W, 130 m, 3 Sep 1970 (zma Pa. 3450); 1 9, 1 juv.
(paratypes), PID, Stn. 159: 22°00'S 40°06'W, 90 m, 5 Sep
1970 (zma Pa. 3451); 1 2 (paratype), Pip, Stn. MBT 164:
21°15'S 40°50'W, 19 m, 6 Sep 1970 (zMa Pa. 3452).
Description. - Trunk segments 1 and 2, and 2 and
3 well-articulated, intersegmental line between seg-
ments 3 and 4 vestigial. One tall spur and 1 or 2 spi-
nules on distodorsal end of each lateral process.
Lateral processes separated by interval greater than
own diameter. Ocular tubercle with 2 minute lateral
tubercles, tip conical; eyes pigmented. Abdomen
more or less strongly directed upward, as long as 4th
lateral process.
152
Proboscis of adult female with 2 ventral alar pro-
cesses, free part of each lobe pointing forward. In ven-
tral view, lobes are heart-shaped. Male proboscis
without alar processes. Proboscis (4, ) surrounded
by collar-like expansion with on either side a rounded
swelling which might represent a rudiment of palp.
Chelifore scape just overreaching tip of proboscis,
with low distal spur. Chela with slender palm; curved
fingers slightly longer than palm. Both palm and ectal
margin of movable finger with several long setae.
Each finger with 2 proximal setae on endal margin.
Oviger 6-segmented; segm. 2 slightly less than half
as long as segm. 3; segm. 3 very elongate; segms. 5
and 6 with several long setae; segm. 6 longer than
wide, pointed.
Legs setose, especially femur and tibiae. Tubercles
and spurs variously developed (strongest develop-
ment in one of the females!). Coxa 1 with 2 to 10 tu-
bercles, most tubercles spine-tipped. Coxa 2 with
ventrodistal genital spur only (male spur thump-
shaped, present of legs 3 and 4; female spur broadly
rounded, on all legs). Coxa 3 sometimes with ventro-
distal, rounded process. Femur slightly longer than
tibiae 1 and 2, with distodorsal spur; setae implanted
on low or vestigial tubercles. Tibiae 1 and 2: setae im-
planted on tubercles, which may assume columnar
shape. Tarsus short. Propodus with strong heel form-
ing right angle with sole, armed with 1 or 2 basal
spines and row of thin spinules; propodal lamina very
short. Rudimentary auxiliary claws present. Cement
gland slightly proximad of middle of dorsal surface of
femur of all legs in male, discharging through strong,
heavy, tapering, thump-shaped tube directed
obliquely distad; length of tube variable: usually as il-
lustrated in fig. 70, but in male from Stn. MBT 211
more slender (fig. 71).
Measurements (d from Stn. MBT 211) in mm. -
Length proboscis (ventral) 1.14; greatest diameter
proboscis 0.34; length trunk (frontal margin cephalic
segment to tip 4th lateral process) 1.76; width across
2nd lateral processes 1.08. Leg 3: first coxa 0.30; sec-
ond coxa 0.96; third coxa 0.46; femur 1.85; first tib-
ia 1.78; second tibia 1.82; tarsus 0.15; propodus
0.64; claw 0.49.
Etymology. - The specific name, spurius (Latin =
illegitimate child) alludes to the dubious taxonomic
position of the new species.
Remarks. — By the presence of two ventral out-
growths on the female proboscis, as well as by almost
all points of the general morphology (structure of
chela, oviger, shape of propodus, short propodal lam-
ina, genital spurs in both sexes...) the present materi-
al shows a close similarity to A. stictus Marcus, 1940.
The differences with A. stictus pertain to the degree of
setosity and tuberculation of legs and lateral processes
and to a longer cement gland duct. These characters
Jan H. Stock: Brazilian Pycnogonida
Figs. 65-75. Anoplodactylus spurius sp. n. (65, 68, 69, 70 and 74 from Stn. MBT 145; 66, from Stn. MBT 165; 67, 73 and 75
from Stn. MBT 159; 71 and 72 from Stn. MBT 211). 65, trunk, d, dorsal (scale F); 66, cephalon, 9, from the right (free-hand
sketch); 67, proboscis, 9, ventral (B); 68, oviger, d (B); 69 leg 3, & (F); 70, 71, cement gland of leg 3, 5 (B); 72, lateral pro-
cess, coxae 1 and 2 of leg 2, d, dorsal (A); 73, leg 3, 2 (A); 74, distal segments of leg 3, & (B); 75, chela, 2 (E). [c= cephal-
ic segment; Lp. = lateral process; pa. = palp.] Scales after fig. 80.
133
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
may be subject to, variation in Anoplodactylus. The
length and number of tubercles, processes and spurs
fluctuate for instance in À. insignis (Hoek, 1881) (see
Stock 1986: 437), whilst the length the cement gland
duct is variable in A. petiolatus (vide supra). To com-
plicate matters, the material attributed here to A. spu-
rius sp. n. is not uniform throughout, but shows
marked variability in degree of development of the leg
armature and length of the gland tube. Moreover,
Bremec et al. (1986: figs. 7-8) illustrate material from
Argentina with a distinct tubercle on the lateral pro-
cesses (as in A. spurius), but with a low cement gland
duct and non-tuberculate legs (as in A. stictus).
Therefore, I find it hard to decide whether the
present material represents a new species or just ‘a
more strongly armed’ variety of A. stictus. I have de-
cided to describe it as a new species, mainly to draw
attention to the plasticity of the morphological char-
acters in this genus. More material will be necessary
to decide if the chosen solution was correct.
The following expression of character states separ-
ates for the moment A. spurius from A. stictus (1) lat-
eral processes with tall distodorsal tubercle in spurius,
tubercle lower or absent in stictus, (2) coxa 1 with 2 to
10 tubercles in spurius, none in stictus (3) cement
gland rather long to long in spurius, short or rather
short in stictus, (4) femur and tibiae with numerous
long setae, placed on (tall) basal tubercles, in spurius
few setae and no basal tubercles in stictus.
Comparison with other species. - Child (1979) has
described A. reimerae from the Pacific coast of
Panama that looks very similar to A. spurius (lateral
processes and coxa 1 with tubercles). But À. reimerae
has lateral (not ventral) alar processes on the female
proboscis, its oviger segments have different propor-
tional lengths, and a different terminal oviger seg-
ment setation. A. spurius is also very similar to A. sim-
ulator Stock, 1975 from Florida, but this species lacks
ventral alar processes on the female proboscis (Stock
1986: 440).
Anoplodactylus stictus Marcus, 1940
(fig. 62)
A. stictus Marcus, 1940b: 65-68, pl. VI fig. 7E, pl. VII fig.
7A-D; Sawaya 1945: 231-234, pl. XXIII; de Castellanos
1965: 16-17; Stock 1966: 405; Bremec et al. 1986: 36-
37, figs. 7-8.
Material. - 1 ©, usp, Stn. P.E.: Ponta do Espia, State of
Sao Paulo, on seaweeds growing in the infralittoral of rocky
shore, 20 Nov 1964; 1 d, 5 9, Project Isotan-Draga I (PID),
Stn. MBT 50: 23°39'S 45°06'W, 28 m, 8 May 1970; 1 9,
PID, Stn. MBT 149: 23°41'S 41°55'W, 250 m, 4 Sep 1970; 1
8,2 9,3 juvs., PID, Stn. MBT 155: 27°22'S 41°32'W, 34 m,
5 Sep 1970; 1 ®, rip, Stn. MBT 156: 22°06'S 40°53'W, 32
m, 5 Sep 1970; 1 ®, 16 juvs., PID, Stn. MBT 157: 21°46'S
40°58'W, 15 m, 15 Sep 1970; 1 2, Pip, Stn. MBT 171:
134
25°05'S 47°40'W, 20 m, 27 May 1971; 2 ®, pip, Stn. MBT
173: 25°32'S 47°11'W, 55 m, 27 May 1971;3 6,1 Q, pip,
Stn. MBT 195: 24°03'S 44°59'W, 73 m, 1 June 1971; 5 d,
6 ®, rip, Stn. MBT 197: 23°47'S 44°44'W, 65 m, 1 June
1971; 1 6, rip, Stn. MBT 206: 23°14'S 44°03'W, 52 m, 3
June 1971; 1 2,R. V. ‘Prof. W. Besnard’ Stn. RD 4: 19°59'S
39°56'W, 41 m, 6 July 1972.
Remarks. - This species is known shallow waters,
mostly in the upper part of the shelf, of the coast of
Brazil (Marcus 1940b, Sawaya 1945) and northern
Argentina (de Castellanos 1965; Stock 1966; Bremec
et al. 1986). The bathymetrical range of the species is
extended by the present records to 250 m. An illustra-
tion of the chela is presented here (fig. 62), to allow
for proper comparison with A. spurius (see above).
Endeis biseriata Stock, 1968
Endeis biseriata Stock, 1968: 57-60, fig. 21; Stock 1970b: 1;
Stock 1974: 17; Stock 1979: 28-30, fig. 9.
Endeis biserata [lapsus calami]; Child 1988b: 20; Child
1990: 332-333.
Phoxichilus meridionalis Loman 1908 [nec Böhm, 1879]:
78-79, pl. XI figs. 153-155.
Material. - 1 d, usp, Stn. P.E.: Ponta do Espia, Ubatuba,
State of Sao Paulo, on seaweeds in the infralittoral of rocky
shore, 20 Nov 1964; 1 ©, 2 juvs., usp, Stn. F 23: Enseada
do Flamengo, Ubatuba, State of Sao Paulo, amongst ascid-
ians, bryozoans and sponges growing of large boulders in in-
fralittoral, 13 Feb 1965.
Remarks. - This species is known from almost the
entire tropical Indo-Pacific, from Hawaii to the Great
Barrier Reef in the east, to the Gulf of Eilat and
Madagascar in the west. The only published Atlantic
record (Stock 1979) is, like the present records, from
Brazil, but in zma there is also a sample from the me-
diolittoral zone of Antigua (West Indies). The single
available male of the present collection has the cement
gland pores arranged in 2 rows on the posterior sur-
face of the tibiae. A count of the longest row on leg 4
amounts to about 29 pores. The 3 distal femoral spurs
are rather short (as in Stock's 1979, fig. 9c). This is a
shallow-water species, found between 0 and 46 m.
Family Colossendeidae
Colossendeis geoffroyi Mafie-Garzón, 1944
C. geoffroyi Mane-Garzón, 1944: 1-6, figs. 1-5; Stock 1966:
396-397, fig. 3f; Larramendy 1974: 149-152, figs. 1-3.
Material. - 1 specimen (fragm.), probably this species,
Project Isotan-Draga I, Stn. MBT 141: 23°25'S 43°00 W,
113 m, 2 Sep 1970; 4 spms., R. V. ‘Prof. W. Besnard', Stn.
1743: 33°50'S 51°51'W, 65 m, 19 Apr 1972.
Remarks. — The record from Stn. 1743 is rather
close to the type-locality, 36°05'S 53°03'W, 130 m.
If the fragmentary specimen from Stn. MBT 141 (dis-
Jan H. Stock: Brazilian Pycnogonida
E 1 mm F
C 0.5 mm D
0.05 mm
Figs. 76-80. Pycnogonum ornans sp. n. (4 holotype). 76, trunk, dorsal (scale A); 77, trunk, from the left (A); 78, oviger (E);
79, leg 3 (A); 80, distal segments of leg 3 (B).
tal leg segments and palps lacking) is really this spe-
cies, the range would be considerably extended in
northward direction. Minnaard & Zamponi (1984:
272-275, fig. 11) describe a Colossendeis identified as
C. geoffroyi from the “Walther Hertwig 1966 cruise,
Stn. 340, 200 m. This station is off Mar del Plata,
Argentina (ca. 40°S). Both the rough figures and the
measurements provided by the Argentinean authors
seem to indicate that a different species is concerned,
because the tarsus is almost twice as long as the pro-
podus, whereas in all other specimens of C. geoffroyi
recorded, there is no great length difference between
tarsus and propodus.
Family Rhynchothoracidae
Rhynchothorax mediterraneus Costa, 1861
R. mediterraneus, Child 1988a: 56 [refs.]; Stock 1988: 507;
Arnaud & Krapp 1990: 4.
Material. - 14 spms., Project Isotan-Draga (PID), Stn.
MBT 155: 22°22'S 41°32'W, 34 m, 5 Sep 1970; 1 spm., PID,
Stn. MBT 156: 22°06'S 40°53'W, 32 m, 5 Sep 1970; 4
spms., PID, Stn. MBT 159: 22°00'S 40°06'W, 90 m, 5 Sep
1970; 1 spm., rip, Stn. MBT 169: 21°50'S 39°52'W, 1100
m, 8 Sep 1970; 1 spm., Project Petrobas/usp (pp), Stn. RD
31: 19°27.5'S 39°03.4'W, 58 m, 10 July 1972; 1 spm., pp,
Stn. RD 53: 18°56'S 39°26'W, 27 m, 12 July 1972.
Remarks. - The present specimens agree pretty
well with the illustrations of R. mediterraneus (cf.
Dohrn 1881; Stock 1974), but for a few details that I
do not consider of great importance. These differenc-
es concern (1) the anterior and posterior spurs on
coxa 1 which in some specimens of the present collec-
tion are taller; (2) the presence in some specimens of
a well-marked tubercle on palp segment 2; (3) a nar-
row, unarmed 2nd segment in the palp, vaguely indi-
cated in Mediterranean material, is more distinctly
present in Brazilian specimens; (4) oviger segments 5
and 6 are subequal (versus 6>5); (5) the posteroven-
tral spur on coxa 3 of leg 3 d is less pronounced than
Dohrn's figures suggests.
The species was recorded before from Brazil by
Zago (1970) and Zilberberg (1963). Child (1988a)
re-examined some of Zilberberg s specimens and con-
cluded that they consisted of a mixture of R. mediter-
raneus and À. monnioti. Apart from Brazil, R. mediter-
raneus has been recorded from the Mediterranean
basin (see Arnaud & Krapp 1990, for references),
Madagascar (Stock 1974), off Gibraltar (Stock 1988),
135
‘TijDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
and the Seychelles (Child 1988a). The present
records fall within the known bathymetrical range of
the species (1-200 m), except for Stn. MBT 169, which
extends this range to 1100 m.
Family Pycnogonidae
Pycnogonum (Retroviger) cessaci Bouvier, 1911
P. (R) cessaci; Stock 1975a: 1085, fig. 58d [syn., refs.];
Child 1979: 72 [refs.]; Child 1980: 325; Stock 1990:
231-232.
Material. - 37 spms., usp, Stn. P.E.: Ponta do Espia,
Ubatuba, State of Säo Paulo, on seaweeds, rocky infralitto-
ral, 20 Nov 1964.
Remarks. - The species was recorded before from
Brazil, under the name of Pycnogonum pamphorum
Marcus, 1940, a junior synonym.
Pycnogonum (Retroviger) gibberum Marcus &
Marcus, 1963
P. gibberum Marcus & Marcus, 1963: 3-7, pl. I.
Material. - 1 4,1 ®, R. V. ‘Prof. W. Besnard’ (ws), Stn.
RG 1325: 23°14'S 43°56.5'W, 49 m, 16 Feb 1968; 1 ®,
wB, Stn. RIG 587: 23°22'S 44°26'W, 50 m, 7 Feb 1969; 1
juv., Project Isotan-Draga I (PID), Stn. MBT 50: 23°39'S
45°06'W, 28 m, 8 May 1970; 1 larva, Pip, Stn. MBT 161:
DIRADIS 402152 56M 65 Sep1970: 01 32, Project
Petrobas/usp (pp) Stn. RD 4: 19°59'S 39°56'W, 41 m, 6 July
1972; 1 juv., pp, Stn RD 25: 19°38'S 39°27'W, 9 Aug 1972.
Remarks. — Pycnogonum elephas Stock, 1966, from
the same general area (between Rio de Janeiro and
Rio de la Plata) is very similar to P. gibberum, but is
almost twice as large. Moreover, the tubercles on the
lateral processes and on coxa 1 are much less conspic-
uous than in P. gibberum, and the proboscis is, espe-
cially in lateral view more slender. It is possible that
the male of P. elephas lacks ovigers; the male of P. gib-
berum has 7- (or 8-2) segmented ovigers.
Apart from the three specimens on which the orig-
inal description is based, P. gibberum has never been
recorded again. The original material came from Ilha
Lage de Santos, 32 m (Marcus & Marcus 1963: 6)
and Alcatrazes Island, 36 m (Marcus & Marcus 1963:
3), both in Brazil. The present records extend both
the geographical and the bathymetrical ranges slight-
ly, to 19°38'S and 56 m.
Pycnogonum (Retroviger) ornans sp. n.
(figs. 76-80)
Type material. - 1 d holotype, R. V. ‘Prof. W. Besnard’,
Stn. RD 49: 19°05.5'S 39°14.3'W, 47 m, 12 Aug 1972 (ZMA
Pa. 3441); 1 ® paratype, Project Isotan-Draga I, Stn. MBT
196: 23°31'S 44°57'W, 38 m, 1 June 1971 (zMa Pa. 3442).
Description. - A small-sized Pycnogonum.
136
Male: Trunk long with respect to width, complete-
ly segmented. Lateral processes widely spaced. Ocular
tubercle low, truncate; eyes well-pigmented. Three
pointed tubercles behind ocular tubercle. Three mid-
dorsal bosses near posterior margin of trunk segments
1, 2, and 3; bosses as tall as ocular tubercle; bosses 1
and 2 broadly conical, boss 3 narrower; each boss
coarsely tuberculate. Similar bosses on dorsal surface
of all lateral processes, that on lateral process 4 tallest.
Anterior and posterior margins of lateral processes 2
and 3, posterior margin of lateral process 1, and ante-
rior margin of lateral process 4 with 1 or more large
tubercles. Abdomen reaching beyond middle of coxa
2 of leg 4, with small tubercles; distally truncate.
Proboscis widest near proximal end, slightly taper-
ing, distally truncate. Dorsal surface with numerous
small tubercles. Oviger 7-segmented; 1 spine on seg-
ments | and 2. Distal claw long, sharp, curved.
Legs short, thin. Coxa 1 of legs 2 to 4 with large,
tuberculate or branched, somewhat upcurved, pro-
cesses on dorso-anterior and dorso-posterior surface;
coxa 1 of leg 1 with low tubercles only. Coxa 2 not
strongly ornamented. Coxa 3 smooth. Femur not
widened; with ventroproximal, triangular hump and
several large tubercles; dorsodistal end marked by tu-
berculate bump and 1 strong seta. Tibia 1 slightly
longer than femur, with smaller tubercles and shorter
distal seta. Tibia 2 much shorter than tibia 1, about 3
times as long as wide, with some tubercles and rather
short distal seta. Tarsus with ventral ‘brush’ of some 7
spinules. Propodus curved, longer than tibia 2; sole
evenly armed with some 10 spinules, some of these
bifid. Claw curved, less than half as long as propodus;
no auxiliary claws.
Female: No ovigers. Ornamentation of dorsum
and legs slightly less pronounced than in male.
Measurements of holotype (mm). - Length pro-
boscis (dorsal) 0.71; greatest diameter proboscis 0.34;
distal diameter proboscis 0.18; length first trunk
segm. 0.73; length second trunk segm. 0.42; length
third trunk segm. 0.33; length fourth trunk segm. (to
tip lateral process) 0.25; width across second lateral
processes 0.74; length abdomen 0.36. Leg 3: first
coxa 0.17; second coxa 0.19; third coxa 0.15; femur
0.35; first tibia 0.36; second tibia 0.30; tarsus 0.06;
propodus 0.35; claw 0.16.
Etymology. - The specific name ornans (Latin =
ornamented) alludes to the ornamentation of trunk,
lateral processes, proboscis, and certain leg segments
with bosses and tubercles.
Remarks. - The narrow body shape is reminiscent
of P. cessaci Bouvier, 1911 and P. guyanae Stock,
1975. However, P. cessaci possesses small auxiliary
claws (absent in the new species), a more barrel-
shaped proboscis, whereas the propodal armature
tends to concentrate in the distal half of the sole
(evenly spread over sole in the species), tibia 1 shorter
than the femur and post-ocular tubercles are absent.
Both in P. cessaci and in P. guyanae, the coxae and the
proboscis are devoid strong tubercles and bosses, so
characteristic for the habit of the new species.
Although P. letictae de Mello-Leitäo, 1945 is gen-
erally considered synonymous with P. cessaci, its pro-
podal armature resembles that of the new species.
However, by the absence of strong tubercles on the
legs, it is clear that the present species cannot be iden-
tified with P. leticiae.
Pentapycnon geayi Bouvier, 1911
P. geayi, Stock, 1975a: 1088 [refs.].
Material. — 1 juv., Project Isotan-Draga I, Stn. MBT 164:
21°15'S 40°50'W, 19 m, 6 Sep 1970; 1 juv., Project
Petrobas/usp (pp), Stn. RD 51: 19°05.1'S 39°40.4'W, 18 m,
12 July 1972; 1 9, pp, Stn RD 67: 18°34'S 39°40', 15 m, 13
July 1972.
Remarks. - This extra-legged form is known from
Puerto Rico, the Guyanas, and the northern coast of
Brazil (southernmost record hitherto 06°04.7'S). The
present records extend the range to 19°05.1'S.
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de Zoologie expérimentale et générale 86, Notes et Revue
1: 20-31.
Fage, L. & J. H. Stock, 1966. Pycnogonides des campagnes
de la Calypso aux îles du Cap Vert. - Annales de l'Institut
océnographique 44: 315-327.
Fry, W. G. & J. W. Hedgpeth, 1969. The fauna of the Ross
Sea, 7: Pycnogonida, 1. - New Zealand Oc-anoprphic
Institute, Memoir 49: 1-139.
Gordon, I., 1932a. Re-description of some type-specimens
of Pycnogonida of the genus Nymphon. — Annals and
Magazine of natural History (10) 9: 97-120.
Gordon, I., 1932b. Pycnogonida. - Discovery Reports 6: 1-
138.
Hedgpeth, J. W., 1948. The Pycnogonida of the western
North Atlantic and the Caribbean. — Proceedings of the
United States national Museum 97 (3216): 157-342.
Hedgpeth, J. W., 1961. Reports of the Lund University
Chile Expedition 1948-49, 40: Pycnogonida. — Lunds
Universitets Ärssrift n.F. (2) 57 (3): 3-18.
Hilton, W. A., 1915. Pycnogonids collected during the
summer of 1915, at Laguna Beach. - Journal of
Entomology and Zoology, Pomona College 7 (3): 201-
206.
Hoek, P. P. C., 1881. Report on the Pycnogonida. — Report
on the scientific Results of the Voyage of H.M.S.
Challenger (Zoology) 3 (10): 1-167, Pls. I-XXI.
Krapp, F., 1973. Pycnogonida from Pantellaria and
Catania, Sicily. - Beaufortia 21 (277): 55-74.
Krapp, F. & J. N. Kraeuter, 1976. Additions to the
Pycnogonida of Georgia. - Bonner zoologische Beiträge
27: 336-346.
Larramendy, M. L., 1974. Anotaciones sobre Colossendeis
geoffroyi (Mañe-Garzon, 1944). - Neotropica 20 (63):
149-152.
Lebour, M. V., 1945. Notes on the Pycnogonida of
Plymouth. - Journal of the marine biological Association
of the United Kingdom 26: 139-165.
Loman, J. C. C., 1908. Die Pantopoden der Siboga
Expedition. - Siboga Expeditie 40: 1-88, pls. I-XV.
Loman, J. C. C., 1923. The Pycnogonida. - Further zoolog-
157
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
ical Results of the Swedish Antarctic Expedition 1901-
1903, 1 (2): 3-41.
Mafié-Garzén, F., 1944. Notas sobre Pantöpodos, 1:
Colossendeis geoffroyi nov. sp., de la plataforma continen-
tal frente al Rio de la Plata. - Communicaciones zoologi-
cas del Museo de Historia natural de Montevideo 1 (15):
1-7.
Marcus, E., 1940a. Pallenopsis fluminensis (Kröyer) e as
Pallenopsis sul-americanas restantas (Arthropoda,
Pantopoda). — Revista entomologica 11 (1-2): 180-199.
Marcus, E., 1940b. Os Pantopoda brasileiros e os demais
sul-americanos. — Boletim da Faculdade de Filosofia,
Cienciàs e Letras, Universidad de Sao Paulo 4: 3-179, Pls.
I-XVII.
Marcus, E. du Bois-Reymond, 1952. A hermaphrodite pan-
topod. - Anais da Academia brasileira de Ciências, Rio de
Janeiro 24 (1): 23-30.
Marcus, E. & E. Marcus, 1963. A new Pycnogonum from
Brazil. - Boletim do Instituto oceanogräfico, Sao Paulo
1228):55>10:
Markham, J., 1986. Class Pycnogonida (= Pantopoda) (Sea
Spiders): 275-277. In: W. Sterrer, ed. Marine fauna and
flora of Bermuda. (John Wiley & Sons, New York).
Mello-Leitäo, A. de, 1945. Uma espécie nova do gênero
Pycnogonum Brünnich, 1764 (Pycnogonidae,
Pantopoda). — Boletim do Museu nacional, Rio de
Janeiro (n.S.) (Zool.) 42: 1-4.
Mello-Leitäo, A. de, 1946. Novo gênero de Pantopodes de
Baia de Guanabara. — Anais da Academia brasileira de
Ciéncias 18 (4): 291-296.
Mello-Leitäo, A. de, 1949a. Uma nova especie de Pallenopsis
do Atläntico Sul (Pantopoda, Pallenidae). - Arquivos do
Museu Paranaense, Curitiba 7 (7): 299-307, Pls. IX-X.
Mello-Leitäo, A. de, 1949b. Nova especie de Halosoma
Cole, 1904 (Pantopoda, Phoxichlidiidae (sic) Sars, 1981.
— Anais da Academia braileira de Ciéncias 21 (2): 167-
173:
Mello-Leitäo, A. de, 1955. Novo Pantopoda da Baia de
Guanabara (Melloleitanianus candidoi, Phoxhichi-
lidiidae). - Revista de Biologia marina, Valparaiso 5
(1952): 119-124.
Minnaard, V. & M.O. Zamponi, 1984. Estudios sistemati-
cos de algunos Pantöpodos de la región subantärtica. —
Historia natural, Corrientes 4 (28): 257-279.
Müller, H.-G., 1989. Shallow-water Pycnogonida from co-
ral reefs at Moorea, Society Islands, with description of
Rhynchothorax tiahurensis sp. n. - Bonner zoologische
Beiträge 40 (2): 123-139.
Müller, H.-G., 1990a. On some Indo-West-Pacific
Pycnogonida from the Zoological Museum,
Copenhagen. — Zoologische Abhandlungen Staatliches
Museum für Tierkunde in Dresden 45 (10): 103-110.
Müller, H.-G., 1990b. Shallow-water Pycnogonida from
Martinique, French Antilles, with description of
Nymphon macabou sp. n. - Bonner zoologische Beiträge
41 (3/4): 277-285.
Müller, H.-G. & M. Roth, 1990. Asselspinnen.
Morphologie, Lebensweise, Präparation. - Microcosmos
79 (6): 162-174.
Salazar-Vallejo, S. I. & J. H. Stock, 1987. Apparant parasit-
ism of Sabella melanostigma (Polychaeta) and Ammothella
spinifera (Pycnogonida) from the Gulf of California. -
Revista de Biologia tropicale 35 (2): 269-275.
Sars, G. O., 1891. Pycnogonida. - Den Norske Nordhavs-
Expedition (Zoology) 20: 1-163, Pls. I-XV, 1 chart.
138
Sawaya, M. P., 1945. Anoplodactylus stictus Marc.
(Pantopoda) em Caiobá, Estado do Paraná. - Aquivos do
Museu Paranaense, Curitiba 4 (10): 231-236, PI. XXIII.
Sawaya, M. P., 1947. Nymphopsis melidae sp. n., e
observagöes sôbre e respiracäo nos Pantopoda. — Boletim
de Faculdade de Filosofia, Ciencias e Letras, Univerdidad
de Sao Paulo (Zoologia) 12: 83-91, Pls. II.
Sawaya, M. P., 1950. Anoplodactylus aragäoi, sp. n. and oth-
er (sic) collected by the hydrographic research-ship ‘Rio
Branco’. - Memorias do Instituto Oswaldo Cruz 47 (1/2)
(‘1949’): 63-86, Pls. I-II.
Sawaya, M. P., 1951. Achelia besnardi, sp. n. (Pantopoda -
Ammotheidae). — Boletim de Faculdade de Filosofia,
Ciéncias e Letras, Universidad de Sao Paulo (Zoologia)
16: 271-280.
Schimkewitsch, W., 1890. Sur les Pantopodes, recueillis par
M. le lieutenant G. Chierchia pendant le voyage de la cor-
vette ‘Vettor Pisani’ en 1882-1885. — Atti della reale
Accademia dei Lincei 1889 (4), Memorie della Classe di
Scienze, Fisiche, Matematichi e Nataturali 6: 329-347,
PI. I.
Schimkewitsch, W., 1930. Pantopoda 2. - Fauna de l'URSS
et des pays limitrophes: 225-555, pls. V-X.
Stock, J. H., 1954. Four new Tanystylum species, and other
Pycnogonida from the West Indies. - Studies on the
Fauna of Curagao 5: 115-129.
Stock, J. H., 1957. Pantopoden aus dem Zoologischen
Museum Hamburg, 2. - Mitteilungen aus den
Hamburgischen Zoologischen Museum und Institut 55:
81-106.
Stock, J. H., 1966. Campagne de la Calypso au large des
côtes Atlantiques del'Amérique du Sud (1961-1962), I,
4. Pycnogonida. - Annales de l'Institut océanographique
44: 385-406.
Stock, J. H., 1968. Pycnogonida collected by the Galathea
and Anton Bruun in the Indian and Pacific oceans. —
Videnskabelige Meddelelser fra Dansk naturhistorisk
Forening i Kjobenhavn 131: 7-65.
Stock, J. H., 1970a. Endeis flaccida Calman, 1923, in
Florida: a pycnogonid new to the Atlantic Ocean. —
Entomologische Berichten, Amsterdam 30 (1): 3-4.
Stock, J. H., 1970b. A new species of Endeis and other pyc-
nogonid records from the Gulf of Aqaba. - Bulletin
Zoölogisch Museum Universiteit van Amsterdam 2 (1):
1-4.
Stock, J. H., 1973. A re-description of the pycnogonid
Pallenopsis boehmi Schimkewitsch, 1930 based on the
original material from Strait Magellan. — Netherlands
Journal of Zoology 23 (3): 347-352.
Stock, J. H., 1974. Medio- and infralittoral Pycnogonida
collected during the 1.1.O.E. near the landbase on Nossi-
Bé, Madagascar. — Bulletin Zoölogisch Museum
Universiteit van Amsterdam 4 (3): 11-22.
Stock, J. H., 1975a. Pycnogonida from the continental
shelf, slope, and deep sea of the tropical Atlantic and east
Pacific. - Bulletin of marine Science 24 (4): 957-1092.
Stock, J. H., 1975b. Pycnogonida found on fouling panels
from the east and west coast of America. -
Entomologische Berichten, Amsterdam 35: 70-77.
Stock, J. H., 1979. Pycnogonida from the mediolittoral and
infrakittoral zones in the tropical western Atlantic. -
Studies on the Fauna of Curagao 59 (184): 1-32.
Stock, J. H., 1986. Pycnogonida from the Caribbean and
the Straits of Florida. - Bulletin of marine Science 38 (3):
399-441.
Stock, J. H., 1988. Faunistic transit between the Atlantic
and the Mediterranean: the deep-water Pycnogonida. —
Cahiers de Biologie marine 28: 505-519.
Stock, J. H., 1989. Pycnogonida collected in the interstitia
of coral sand and rubble in the Netherlands Antilles. —
Bijdragen tot de Dierkunde 59 (2): 87-92.
Stock, J. H., 1990. Macaronesian Pycnogonida. -
Zoologische Mededelingen (Leiden) 63 (16): 205-233.
Wicksten, M. K., 1980. Crustacea and Pycnogonida. In:
Southern California marine invertebrates. — Allan
Hancock Foundation technical Report 3: 196-223.
Zago, M. S. Arcifa, 1970. Söbre o Pantópodo
Rhynchothorax mediterraneus Costa, 1861. -
Contribuigöes do Instituto oceanografico, Univerdad de
Säo Paulo (Oceanografia y Biologia) 21: 1-5.
Zambrana Diaz, M., M. I. Garcia Palmer, O. Rivero Bostos
& T. Eligio Castillo, 1985. Lista de especies y bibliografia
de los picnogönidos (Arthropoda) del Mediterräneo
Americana. — Revista de Investigacién marina, La
Habana 6 (1): 29-38.
Zilberberg, F., 1963. Notes on Pantopoda. - Boletim del
Instituto oceanografico, Universidad de Sao Paulo 13 (2):
21-32.
Received: 1 October 1991
Accepted: 2 March 1992
Table 1. Taxa represented in the usp collections and their
biogeographic characterization.
In brackets after the species name, the following abbrevia-
tions are used:
AA amphi-Atlantic elements (on the warm and warm-
temperate Atlantic coasts of America and Europe,
the Mediterranean basin, and the northern and
central Atlantic coasts of Africa)
CT circum-tropical elements
E endemics of Brazil, the Mar del Plata region and
the Guyanas
M Magellanean elements (cold-water elements of
South America, from Argentina south of the Rio de
la Plata to the Falkland Islands and South Georgia,
southern Chile, sometimes extending northward to
southern Brazil).
P also on the warm and warm-temperate Pacific
coasts of the Americas
WAW western Atlantic warm-water elements (from
southern Florida and the West Indies, the
Guyanas, to Brazil and the Mar del Plata in north-
ern Argentina).
Family Ammotheidae
Genus Eurycyde Schiödte, 1857
E.platyspina sp. n. [E]
Genus Nymphopsis Haswell, 1884
N. melidae Sawaya, 1947 [E]
Genus Cilunculus Loman, 1908
C. sp. [E?]
Genus Ammothella Verrill, 1900
A. appendiculata (Dohrn, 1881) [AA]
A. rugulosa (Verrill, 1900) [WAW]
A. spinifera Cole, 1904 [WAW + P]
Genus Achelia Hodge, 1864
JAN H. Stock: Brazilian Pycnogonida
A. besnardi Sawaya, 1951 [WAW]
A. columnaris sp. n. [E]
A. gracilis Verrill, 1900 [WAW]
A. sawayai Marcus, 1940 [mainly AA, possibly
CT]
Genus 7anystylum Miers, 1879
T. acuminatum Stock, 1954 [WAW]
T. evelinae Marcus, 1940 [E]
T. isabellae Marcus, 1940 [E]
T. isthmiacum difficile Stock, 1966 [WAW]
T. orbiculare Wilson, 1878 [AA, possibly CT]
Family Nymphonidae
Genus Nymphon J.C. Fabricius, 1794
N. bullatum sp. n. [E]
N. inaequipes sp. n. [E]
N. vulcanellum sp. n. [E]
Family Callipallenidae
Genus Callipallene Flynn, 1929
C. evelinae Marcus, 1940 [E]
C. gabriellae Corrêa, 1948 [E]
C. margarita (Gordon, 1932) [M]
C. producta (Sars, 1888) [AA]
Genus Pallenoides Stock, 1951
P. amazonica Stock, 1975 [E]
Family Phoxichilidiidae
Genus Pallenopsis (subgenus Pallenopsis) Wilson, 1881
P. (P.) boehmi Schimkewitsch, 1930 [M]
P. (P.) candidoi de Mello-Leitäo, 1949 [E]
P. (P.) fluminensis (Kroyer, 1844) [E]
P. (P.) kempfi Stock, 1975 [WAW]
P. (P.) tumidula Loman, 1923 [M]
Genus Anoplodactylus Wilson, 1878
. aragaoi Sawaya, 1950 [E]
. brasiliensis Hedgpeth, 1948 [E]
. evelinae Marcus, 1940 [WAW]
. insignis (Hoek, 1881) [WAW]
. maritimus Hodgson, 1914 [AA]
. monotrema Stock, 1979 [WAW]
. petiolatus (Kroyer, 1844) [AA]
. spurius sp. n. [E]
stictus Marcus, 1940 [E]
Genus Endeis Philippi, 1843
E. biseriata Stock, 1968 [CT]
SR RS
Family Colossendeidae
Genus Colossendeis Jarzynski, 1870
C. geoffroyi Mané-Garzón, 1944 [E]
Family Rhynchothoracidae
Genus Rhynchothorax Costa, 1861
Rh. mediterraneus Costa, 1861 [AA, probably CT]
Family Pycnogonidae
Genus Pycnogonum Brünnich, 1764, subgenus Retroviger
Stock, 1968
P. (R.) cessaci Bouvier, 1911 [AA]
P. (R.) gibberum Marcus & Marcus, 1963 [E]
P. (R) ornans sp. n. [E]
Genus Pentapycnon Bouvier, 1910
P. geayi Bouvier, 1911 [WAW]
139
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
BOOK ANNOUNCEMENTS AND
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Skou, Peder, 1991. Nordens Ugler. Handbog over de i
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Graphic design
Ontwerpers B.V., Aad Derwort, ’s-Gravenhage
STEPHEN J. BROOKS & STEPHEN J. ICES
CZ
RARY
‘The Natural History Museum, London
"James Cook University of North Queensland
A NEW SPECIES OF OREAGRION (ODONATA:
JAN 2 7 1993
HARVARD
UNIVERSITY
COENAGRIONIDAE): MONTANE DAMSELFLIES
FROM NEW GUINEA
Brooks, S. J. & S. J. Richards, 1992. A new species of Oreagrion (Odonata: Coenagrionidae):
montane damselflies from New Guinea. — Tijdschrift voor Entomologie 135: 141-144, figs. 1-
8. [1ssN 0040-7496]. Published 1 December 1992.
Oreagrion pectingi sp. n. (Coenagrionidae) is described from specimens collected above 3000 m
on the slopes of Mt Wilhelm and Mt Capella, Papua New Guinea. The discovery of this new
species sheds new light on the relationship of Oreagrion with other ischnurines.
Correspondence: S. J. Brooks, Department of Entomology, The Natural History Museum,
Cromwell Road, London SW7 5BD, U. K.
Key words. — Odonata, /schnura, Oreagrion, New Guinea, phylogeny.
The ischnurine genus Oreagrion Ris was originally
based on a single female collected at 3000 m in the
Snow Mountains (now called the Maoke Range) of
central New Guinea (Irian Jaya) (Ris 1913). Thirty
years later, from the same general area, Lieftinck
(1949) described a further three species collected ne-
ar Lake Habbema and Mt Wilhelmina (now called
Mt Trikora) between 3000-3500 m. Since then no
further species of the genus have been described, alt-
hough Lieftinck predicted that others would probably
be found residing throughout the sub-alpine high-
lands of New Guinea. About 700 km to the east of
Lake Habbema, in Papua New Guinea, similar tro-
pic-alpine conditions occur on Mt Wilhelm and it
was here that a fifth species was discovered by George
Beccaloni in August 1990. Subsequently, in
November, 1991 one of us (SJR) discovered another
population on the southern slopes of Mt Capella, in
the Star Mountains, central New Guinea.
At both localities, no other insects were flying on
the cool, exposed slopes, so it is of note that this spe-
cies of damselfly can tolerate such harsh conditions.
However, Oreagrion species are stout and robust, and
the head, thorax, and basal segments of the abdomen
have a sparse covering of long, fine hairs which may
play a role in thermoregulation. Similar hairs occur in
Protallagma, a neotropical genus of ischnurines from
the high Andes.
Because of the absence of a ventral spine on the 8th
abdominal segment in females of Oreagrion, Lieftinck
(1949) regarded the genus as the primitive progenitor
of Ischnura which it otherwise resembles. However,
Oreagrion pectingi sp. n. does possess a ventral spine
which throws into doubt Lieftinck’s interpretation of
the phylogenetic relationships of the genus.
‘TAXONOMY
Oreagrion pectingi sp. n
(figs. 1-8)
Type material. — Holotype d: Papua New Guinea,
Chimbu Province, Mt. Wilhelm, near waterfall entering
west end Lake Piunde, 3600 m, 6 August 1990, G.
Beccaloni (BMNH). — Paratypes: 1 2, same data as holotype;
28 1@ (incl. one pair), Papua New Guinea, Star
Mountains, southern slopes of Mt Capella, Dokfuma, pond
in subalpine meadow, 5°1'S 141°7’E, 3000-3200 m, 16/19
Nov 1991, S. J. Richards (all paratypes in BMNH).
Description
Male holotype. Note that the markings described
below are based on living specimens; after the death
the pale areas become orange-brown.
Head (fig. 1). Width across eyes 4.1 mm. Eyes (in
life) dark brown in dorsal half, pale blue ventrally.
Labrum blue-green with narrow black basal stripe
and black median triangular spot. Base of mandible
blue-green. Gena blue-green. Clypeus black. Frons
blue-green. Vertex black with small, rounded, pale
blue post-ocular spots and narrow, pale blue occipital
stripe. Long fine hairs sparse on labrum, clypeus,
vertex (especially in post-ocular region), occiput;
dense on frons.
141
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-2. Oreagrion pectingi sp. n. — 1, male, right lateral view; 2, female, left lateral view;
Prothorax (fig. 1). Anterior lobe black with small
lateral blue-green spot; propleuron blue-green; me-
dian lobe black with small lateral blue-green spot;
posterior lobe black. Hairs on posterior lobe sparse,
dense on posterior margin of propleuron.
Mesostigmal lamina (fig. 3): black medially, perip-
herally blue-green, with small, rounded submedian
tubercles on posterior margin; tubercles black on
anterior face, blue-green dorsally and posteriorly.
Pterothorax (fig. 1). Bronze-black on dorsum, pale
antehumeral stripe absent; mesepimeron black in
dorsal half, blue-green below; metepisternum almost
entirely blue-green but with narrow black stripe ad-
jacent to second lateral suture; metepimeron with
narrow black stripe adjacent to second lateral suture,
the rest blue-green. Fine long hairs covering most of
synthorax but absent antero-laterally. Legs: black
with narrow blue-green stripe on flexor surface of fe-
mora; coxa blue-green. Wings: hyaline with faint am-
ber suffusion. Venation black. Pterostigma (fig. 4)
medially dark brown in fore wing, pale brown in hind
wing, with narrow yellow-brown border in all wings;
2.5 cells below pterostigma in fore wing, 1.25-1.50
cells in hind wing. Fore wing with 12 postnodals, 10-
11 in hind wing. R3 arises between 4th and 5th post-
nodals in fore wing, 3rd and 4th in hind wing.
Measurements. Hind wing 17.5 mm, 4.5 mm at gre-
atest width, pterostigma (fore wing) 0.80-0.84 mm
long, 0.5-0.6 mm wide, (hind wing) 0.56-0.62 mm
long, 0.36-0.38 mm wide.
Abdomen (fig. 1). SI with black dorsal spot in
anterior half, apple green posteriorly; S2-S7 black
dorsally, apple green ventro-laterally; S8-S9 entirely
sky-blue dorsally; S10 entirely black dorsally. Fine
hairs in ventro-lateral region of S1 and S2, but hairs
confined to anterior half of S3. Appendages: superior
(figs. 5-6) apple green at extreme base and extero-late-
rally, black dorsally and interno-laterally; about 3 ti-
mes as long as inferior, about same length as S10; co-
142
nical and diverging in dorsal view with broad, re-
curved median ventral tooth which narrows towards
its apex. Hairs short, sparse. Inferior with apex of
dorsal branch black, rest apple green; bifurcate with
broad, blunt ventral branch and narrow, cylindrical
dorsal branch. Total length 30.5 mm, abdomen 24
mm (including appendages 0.58 mm).
Female paratype. — As male except the following.
Head (fig. 2). Eyes dark brown in dorsal half, apple
green ventrally. Labrum apple green with small me-
dio-basal black spot. Base of mandible apple green.
Gena apple green. Clypeus black with narrow apple
green border. Frons apple green. Vertex black; post-
ocular spots apple green, large and confluent with oc-
cipital stripe.
Prothorax (fig. 2). Anterior lobe apple green; prop-
leuron apple green; median lobe black dorsally with
large apple green lateral spot; posterior lobe black me-
dially, apple green laterally. Mesostigmal lamina (fig.
7). Black medially with anterior margin and lateral lo-
bes apple green; lateral lobes reniform.
Pterothorax (fig. 2). Blonze-black on dorsum with
broad apple green antehumeral stripe; mesepimeron
with narrow black stripe adjacent to first lateral sutu-
re, the rest apple green; metepisternum apple green
with short black stripe near posterior margin; metepi-
meron apple green with black posterior spot on se-
cond lateral suture. Fine, long hairs over most of synt-
horax but absent from antero-lateral region. Legs:
femur and tibia apple green with lateral black stripe
on external face; tarsus black. Wings: hyaline with pa-
le amber suffusion. Venation black. Pterostigma (fig.
8) pale brown in fore and hind wing; 1.5 cells below
pterostigma. Measurements: hind wing 19.5 mm, 4.0
mm at greatest width; pterostigma 0.72 mm long,
0.32 mm wide in fore wing; 0.62 mm long, 0.32 mm
wide in hind wing.
Abdomen (fig. 2). S1 apple green with small black
Brooks & RICHARDS: À new species of Oreagrion
So con
CA ga
© sen
>
TX,
Figs. 3-8. Oreagrion pectingi sp. n. — 3, male mesostigmal lamina, dorsal view; 4, male apex of fore and hind wing; 5, male an-
al appendages, right lateral view; 6, male superior appendages, dorsal view; 7, female mesostigmal lamina, dorsal view; 8, fe-
male apex of fore and hind wings.
lateral spot connected by fine black line to black
dorsal spot in anterior half; S2 apple green with small
black antero-dorsal spot and black sagittal spot in
posterior half; S3-S10 with black dorsal stripe, nar-
row anteriorly, broadening posteriorly, apple green
laterally. S8 with ventral spine. Ovipositor short, ex-
tending 0.2 mm beyond apex of abdomen. Anal ap-
pendage (fig. 2) short, conical, blunt apically and
slightly downturned. Measurements: total length
30.5 mm, abdominal length 22.5 mm (including an-
al appendages 0.38 mm).
143
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Habits. — The type locality is a steep east-facing
bank, supporting alpine vegetation, about 100 m
above Lake Piunde on the slopes of Mt Wilhelm. A
full description of the terrain is given by Smith
(1982). The lake is oligotrophic and appeared to be
devoid of higher plants and invertebrates. During
August rain may fall every afternoon but up to 22
consecutive dry days have been recorded. Night tem-
peratures fall below freezing and day temperatures
may exceed 20°C. The holotype and paratype were
collected over a shallow seepage trickling through
grass, low herbage and bare rocks. The seepage was
unshaded and exposed to the wind. Several addition-
al zygopterans, probably of the same species, were
observed, but not collected, flying in sunshine at the
locality. No other odonates were seen. The holotype
and paratype were photographed ovipositing in tan-
dem into rotting grass stems in about one cm of
water. Both male and female had their legs in contact
with emerging grass stems during oviposition.
The locality at Mt Capella is similar in character.
The meadow floor is covered with a thick mat of ferns
with clumps of large tree-ferns and stunted rhodo-
dendrons. The entire meadow is very boggy and with
scattered small areas of open water. O. pectingi occur-
red around these pools. The ponds were devoid of ve-
getation and supported a depauperate fauna. No
Odonata larvae were discovered in the ponds.
Temperatures in November fell to 1°C at night and
got no higher than 15°C during the day. Individuals
flew during brief periods of sunshine. They ceased
activity during cloudy or rainy weather. At one pond,
about 4 m in diameter, five males rigorously defended
territories around the perimeter, chasing off intruding
males. Two of these males successfully mated at least
once during the three days they were observed.
Remarks. — Females of Oreagrion pectingi may be
distinguished from other members of the genus by
the presence of a ventral spine on the 8th abdominal
segment. Otherwise pectingi has similar markings to
O. oreadum Lieftinck. However, the lateral lobes of
the mesostigmal lamina are rounded in pectingi but
narrow and taper apically in oreadum.
The abdominal markings of the male pectingi also
most closely resemble male oreadum but the pale co-
louration is yellow rather than blue-green in the latter
species. In oreadum males there is a broad antehumer-
al stripe which is absent in pectingi and the pterostig-
ma of the fore wing is uniformly brown and lacks the
pale apical outer angle of oreadum. The anal appen-
dages most readily differentiate pectingi from other
Oreagrion species and are not as densely pubescent.
However, they are closest in appearance to oreadum.
Like this species the superior appendages taper apical-
ly but are more acutely pointed in pectingi. The large
144
ventral tooth arises subapically in pectingi, but is api-
cal in oreadum. The inferior appendages in oreadum
are narrow, directed posteriorly and are considerably
longer than the superiors. The inferiors of pectingi are
short and bifurcate, although the narrow dorsally di-
rected branch is again reminescent of oreadum. Male
and female of O. pectingi are considerably larger than
oreadum. The similarities of the characters noted
above suggest that O. pectingi is most closely related
to O. oreadum, which Lieftinck (1949) thought was
the most primitive of the Oreagrion species.
Lieftinck (1949) considered Oreagrion to be an
ischnurine because the males have dissimilar pte-
rostigmata in the fore and hind wing and because of
certain penile characters, synapomorphies that sugge-
st a close relationship with /schnura. However, unlike
Ischnura, the females of the four Oreagrion species
known to Lieftinck lacked a ventral spine, which
implied that they were less derived than /schnura.
Nevertheless, in O. pectingi, a fully developed ventral
spine is present. If Lieftinck’s assertion is correct, and
Oreagrion is a primitive ischnurine, then the ventral
spine must have arisen independently once in O.
pectingi and again in the /schnura-related ischnurines.
Another interpretation of the phylogeny of Oreagrion
is that the ventral spine has been secondarily lost in
the other four species of Oreagrion, but plesiomorphi-
cally retained in O. pectingi. This would imply that,
rather than being a stem-group ischnurine, Oreagrion
is more closely related to /schnura than Lieftinck sug-
gested. However, until more species of Oreagrion are
discovered and more work is done on ischnurine phy-
logeny this problem will be difficult to resolve.
ACKNOWLEDGEMENT
We are very grateful to George Beccaloni for al-
lowing us to study the dragonflies he collected while
on the Imperial College Expedition to Papua New
Guinea.
REFERENCES
Lieftinck, M. A., 1949. The dragonflies (Odonata) of New
Guinea and neighbouring islands. Part VII. Results of the
third Archbold Expedition 1938-1939 and of the Le
Roux Expedition 1939 to Netherlands New Guinea (II.
Zygoptera). — Nova Guinea (New Series) 5: 1-271, figs.
Ris, F., 1913. Die Odonata von Dr. H. A. Lorentz’
Expedition nach Siidwest-Neu-Guinea 1909 und einige
Odonata von Waigeu. — Nova Guinea 9: 471-512.
Smith, J. M. B., 1982. Origins of the tropicalpine flora. p.
287-308. — In: J. L. Gressitt (ed), Biogeography and eco-
logy of New Guinea. Junk, The Hague.
PINGPING CHEN' & NICO NIESER?
! Beijing Academy of Agriculture and Forestry Sciences, Institute of Plant and Environmental
Protection, Beijing, China
2 Tiel, The Netherlands.
GERRIDAE, MAINLY FROM SULAWESI AND PULAU
BUTON (INDONESIA)
Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), III.
Chen, P. & N. Nieser, 1992. Gerridae, mainly from Sulawesi and Pulau Buton (Indonesia).
Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), III. — Tijdschrift voor
Entomologie 135: 145-162, figs. 1-41, table 1. [rss 0040-7496]. Published 1 December 1992.
Gerridae mainly from Sulawesi, Pulau Buton and Sabah (E. Malaysia) are revised and keys to
Indonesian genera and Sulawesi species are provided, with records and distributional notes.
Nine new species are described: Cylindrostethus akanthinos, Metrocoris breviculus,
Potamometropsis anomalis, Rheumatometroides drepanephoros, R. makraitos and Ventidius xyele
from Sulawesi, Ventidius xiphibion from Sulawesi and Buton, Tenagogonus akanthinos and
Ventidius kurtokalami from Sabah.
Correspondence: Dr. N. Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.
Key words. — Sulawesi; Buton; Sabah; Gerridae; keys; new species.
As our contribution to the Fauna Malesiana pro-
gram, we are working on the Nepomorpha and
Gerromorpha mainly collected by staff members of
the Zoological Museum Amsterdam and the
National Museum of Natural History Leiden. The
first part of the Gerridae, viz. a revision of the genus
Limnometra in the Malay archipelago, was published
as the second part of our series (Nieser & Chen
1992a), while Naucoridae, Nepidae and Notonec-
tidae appeared in the first part (Nieser & Chen
1991).
The specimens studied in this paper were mainly
collected by J. P. Duffels, N. Nieser, J. van Tol and
G. Zimmermann on several expeditions to Sulawesi,
the nearby island of Buton, and to Borneo. For more
details (e. g. a list of localities) the reader is referred to
Nieser & Chen (1991, 1992a). In addition, unidenti-
fied Gerridae from Indonesia kept in RMNH and ZMA
have been studied. Dr. D. A. Polhemus (presently at
Bishop Museum, Honolulu, Hawaii) is working on a
revision of the genus Ptilomera Amyot & Serville.
Most of the specimens of this genus taken by the col-
lectors mentioned above are at his disposal.
A key to the genera of Gerridae occurring in
Indonesia and E Malaysia is included. When rele-
vant, keys to species of Sulawesi and Buton are given
as well. These should be used with caution as the fau-
na of the region is still poorly known.
Measurements are in millimetres and present the
range or the (arithmetic) mean based on five random-
ly chosen specimens, or in case there are less than fi-
ve, on all specimens available. The width of the head
is measured across the eyes, the width of the thorax is
measured across the metaacetabula.
Most of the females in the samples carried eggs; on-
ly where such was not the case this has been indicated.
MATERIAL
Localities are in Indonesia, unless stated otherwise.
The areas in Sulawesi (Celebes) used with the locali-
ties agree with the Indonesian provinces, viz. Sulawesi
Utara, Tengah and Tenggara for respectively for
Northern, Central and Southeastern Sulawesi (see
Whitten et al. 1988). Pulau Buton is mentioned se-
parately although it belongs administratively to
Sulawesi Tenggara.
Collections from which material has been studied
or in which material has been deposited (with abbre-
viations in brackets) are: Museum Zoologi Bogor,
Bogor, Jawa (MBBJ); Bagian Pertanian, Universitas
Haluoleo, Kendari, Sulawesi (BPUH); J. T. Polhemus
Collection (University of Colorado Museum),
Englewood, Co. U. S. A. (yrec); Snow Entomological
Museum, University of Kansas, Lawrence, Kansas, U.
S. A. (semc); N. Nieser collection, Tiel, The
Netherlands (Nc); P. Chen collection, Beijing, P. R.
China (pcHc); National Museum of Natural History
(Rijksmuseum van Natuurlijke Historie), Leiden,
The Netherlands (RMNH); G. Zimmermann collec-
145
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
tion, Marburg, B. R. D. (zc); Zoölogisch Museum,
Afdeling Entomologie, Amsterdam, The Netherlands
(zma); Zoological Museum of the University,
Copenhagen (zMC).
Specimens collected by J. P. Duffels are deposited
in ZMA, those by J. van Tol in RMNH, those by G.
Zimmermann in zc and those by N. Nieser in his
own collection, unless otherwise stated. Deposition of
material is usually specified only with rare or new spe-
cies.
ACKNOWLEDGEMENTS
Thanks are due to Dr. J. P. Duffels (ZMA), Mr. J.
van Tol (RMNH) and Dr. G. Zimmermann (Marburg,
B. R. D. ) for putting specimens in their care to our
disposal; to Dr. N. Moller Andersen (zmc) for discus-
sing several problems in identification and Dr. I
Made Ebeneser (BPUH) for showing the senior author
some interesting localities.
SYSTEMATIC PART
Key to Indonesian and East Malaysian genera of
Gerridae
(Adapted from Andersen 1982)
1. Ventral lobes of head produced (fig. 1), first ab-
dominal sternum present. Female ovipositor well
developed, serrate (Rhagadotarsinae) ..................
VERA CUES aterm iad ae eo Rhagadotarsus
— Ventral lobes of head not produced, first abdomi-
nal sternum absent or fused with metasternum.
Female ovipositor short and nonserrate ........... 2
2. Middle femora usually stout, shorter than middle
tibia and hind femora (Trepobatinae)
— Middle femora slender, usually distinctly longer
thantmiddletibiaer ea 5
3. Metasternal scent gland opening absent .............
ER ren fia OM AT ET IN EN Metrobatopsis
— Metasternal scent gland opening small but dis-
Une REN e OME ia LE el 4
4. Metasternum not produced anteriorly, scent
gland opening in the centre of metasternum ......
EL. RAR SEEN Rp Rheumatometroides
— Metasternum produced anteriorly, scent gland
opening near anterior border ............. Stenobates
5. Metasternum extremely reduced, usually only re-
presented by a very short subtriangular plate en-
closing the scent gland opening (Fig. 2). Claws of
hind tarsi modified, straight or S-shaped
(Halobatinae) ieee ee eee Ar nee 6
— Metasternum well developed, laterally clearly re-
aching the metaacetabular region. Claws of hind
tarsi, ifipresent, aleatee yes eee 10
6. Clypeus with basal margin well defined, anterior
146
10.
12:
13:
14.
margin of head not smoothly rounded.
Metasternum reduced but reaching metaacetabu-
laslaterally marine Halobates
Clypeus with basal margin obliterated or lost, an-
terior margin of head broadly and smoothly
rounded. Metasternum strongly reduced, not re-
aching metaacetabula laterally … … … … … … … 7
. Metanotum with lateral longitudinal elevation
reaching intersegmental suture between meso-
and metanotum. Male third antennal segment
with stiff hairs on margin ........................ Esakia
Metanotum with lateral longitudinal elevation
indistinct or ending somewhere on metanotum,
not reaching intersegmental suture between me-
so- and metanotum. Pilosity of third antennal
segment variable, but no stiff hairs on margin . 8
. Eyes overlapping anterolateral angles of mesono-
WD E N. OR 9
Eyes not overlapping anterolateral angles of mes-
ONO tlm EAN MR EN Metrocoris
. Metaacetabula and hind coxa of female produced
medially Penn Ventidiopsis
Metaacetabula and hind coxae of female not pro-
dueedmediallya een Ventidius
Fore tarsi long, at least one half the length of fore
tibia. Metaacetabular groove dorsally reaching
anterior end of first abdominal tergum
(Btilomerimae)h. Mn. N 11
Fore tarsı shorter. Metaacetabular groove not re-
aching anterior end of first abdominal tergum …
AM RACE RN, 14
. Hind coxae with a small spine on apical margin,
hind femora much longer than middle femora ...
M BARS. een) | Ates N er Ptilomera
Hind coxae without spines, hind femora sube-
qual to or shorter than middle femora ........... 12
Anterior margin of head rounded, first antennal
segment shorter than three following segments
together GE Rheumatogonus
Anterior margin of head not rounded, first anten-
nal segment about as long as or longer than three
following segments together .......................... 13
First antennal segment about as long as three fol-
lowing segments together, middle and hind tarsi
with distinct Claws … Potamometropsis
First antennal segment longer than three follo-
wing segments together, middle and hind tarsi
without distinct Claws ...................... Pleciobates
Rostrum short, not surpassing the prosternal
margin, forth antennal segment short and curved.
First segment of fore tarsi less than one-half the
length of second (Cylindrostethinae)
Etre ear Ser, Bateau AE LORO Cylindrostethus
Rostrum longer, clearly surpassing the prosternal
margin, forth antennal segment straight. First
segment of fore tarsi usually more than one-half
the length of second segment … 15
15. Wingless form with posterior lobe of pronotum
absent, small species (length up to 5 mm), anteri-
or lobe of pronotum with a central and a pair of
sublateral longitudinal light stripes, tarsal claws of
middle and hind legs present (Eotrechinae) … …
bbl Seer ais ita oh AA Amemboa
— Wingless form with posterior lobe of pronotum
present, usually larger species, if length about 5
mm or less, then tarsal claws of middle tarsi ab-
sent or anterior lobe of pronotum light with a
dark longitudinal median stripe (Gerrinae) ... 16
16. Dorsal surface of head and abdominal venter ge-
nerally dark. Metathoracic spiracle located only a
little more than its own length from base of fore
WIND Ren ee nine ester ne rig Aquarius
— Dorsal surface of head light or dark with longitu-
dinal dark stripes or elongate spots, venter gene-
rally light. Metathoracic spiracle located much
more than its own length from base of fore wing
17. Anterior lobe of pronotum dark with a large me-
dian light spot. Claws absent on middle tarsi .....
FREE EERE nn Neogerris
— Anterior lobe of pronotum light or dark with a
pair of small light spots. Claws present on middle
(TAI a aes O I OR 18
18. Posterior lobe of pronotum dark, usually with a
median longitudinal white stripe ..... Limnogonus
— Pronotum basically light with a median longitu-
dinal dark stripe, sometimes with quite extensive
Othermdarksmankinesnecse ct I 19
19. Connexival spines normal. Male abdomen not re-
duced. Last ventral abdominal segment plus geni-
tal segments shorter than preceding four seg-
ments. Hind coxae of male rarely extending
beyond middle of second ventral abdominal seg-
EN EHER RE eee tee ene ALT AS A EE Limnometra
— Connexival spines reduced. Male abdomen redu-
ced. Ventral abdominal segments short, last seg-
ment plus the genitalia at least as long as prece-
ding four segments, usually longer. Hind coxae of
male nearly reaching or surpassing posterior mar-
gin of second ventral abdominal segment … … …
VORO o Ra bd Tenagogonus
GERRINAE Bianchi, 1896
Limnogonus Stäl, 1868
A tropicopolitan genus with a relatively small num-
ber of, often widespread, species. Andersen (1975)
provides an excellent revision of the Old World spe-
cies.
Most Limnogonus live on small stagnant waters in-
cluding cattle ponds and pools and ditches in sawahs.
CHEN & Nieser: Gerridae from Sulawesi
Some species have good tolerance for brackish water.
Three species, one with two subspecies, are known
from Indonesia; all three occur in Sulawesi.
Key to Limnogonus of Sulawesi
1. Connexival spines well developed (fig. 3), prono-
tal lobe usually without a mesial pale stripe ........
Rl NN L. nitidus
— Connexival spines reduced, pronotal lobe with a
mesialipalefsEripep ennn 0 enne 2
2. Pale stripe on upper part of mesopleuron tapering
posteriorly, ending in front and below second
spiracle (Ho ARRET L. fossarum skusei
— Pale mesopleural stripe widened posteriorly, en-
ding above the second spiracle (fig. 5) ................
RE E L. hungerfordi
Limnogonus fossarum fossarum (Fabricius)
Cimex fossarum Fabricius, 1775: 727.
Limnogonus fossarum; Cheng 1966: 121-127 (biology).
Limnogonus fossarum fossarum; Andersen 1975: 30-36, figs.
30-36, 49-53, 56-61, 64-67, 69, 70, 72, 73, 78-85 (rede-
scription, synonymy); Selvanayagam & Rao 1989: 474-
484 (biology); Rubia & Heong 1990: 34-35 (life history).
Material. — INDONESIA. N. Sumatera: Alas valley,
Kutacane, Tanah merah, 3°31’N 97°47’E, ca. 200m, 8. VI.
1972, 39, leg. J. Krikken No 9; NE Sumatra: Kuala
Simpang, lowland cultivated area, at light, XI. 1953, 24
59; same II. 1964, 1d, leg. A. Sollaert. — MALAYSIA, Sabah:
16 km NE Tenom Agr. Res. Station, 115°5Y’E, 5°11’N,
TL, 180m, 11. VI. 1986, leg. J. Huisman, 28 29 (18
brach. ); same 23. XI. 1987, leg. J. Huisman & R. de Jong,
1d; Kinabalu Park HQ, roadside, ML, 116°32’E, 6°00°N,
1500m, 14. V. 1987A, 1 9 ; leg. J. Huisman (all RMNH).
Distribution. — Widespread: India (except the
NE), Indochina, extreme S of China including Hong
Kong and Taiwan, Ryu Kyu Is., Philippines,
Sumatera and N Borneo (Andersen 1975).
Remarks. — One of the very few species of tropical
Asian Gerridae of which the biology has been studied
(Cheng 1966, Selvanayagam & Rao 1989, Rubia &
Heong 1990). Chinese texts report that it is useful as
a component of a cure against malaria (Anonymous
1982).
Limnogonus fossarum skusei (Bueno)
(figs. 3-4)
Limnometra skusei Bueno, 1926: 129 (new name for
Hydrometra australis Skuse, preoccupied by Hydrometra
australis Say ).
Limnogonus fossarum; Lundblad 1933: 374-380, fig. 119
(redescription based on specimens from Java).
Limnogonus fossarum skuset, Andersen 1975: 36-39, figs. 54,
71, 74-77, 86-89, 91-92 (subspecific status).
147
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-16. — 1, diagram of head of Gerridae, left Rhagadotarsus right Cylindrostethus, stippled area is the ventral lobe; 2, dia-
gram of thoracic sternites in Gerridae, left Halobatinae, right Gerrinae, sclerite with stippled margin is metasternum with
scent gland orifice; 3, apex of abdomen in female Limnogonus, left L. nitidus, right L. fossarum: 4, lateral view of thoracical
pattern in Limnogonus fossarum skusei; 5, idem, L. hungerfordi 6, Tenagogonus akanthinos sp. n., holotype à , apex of abdo-
men ventral view, scale 0.5 mm; 7, idem, allotype 9, apex of abdomen ventral view, scale 1 mm; 8, idem, lateral view of en-
dosomal sclerites of paratype, scale 0. 25 mm; 9-12, Cylindrostethus, left C. brachyakanthinos sp. n., right C. persephone Kirk;
9, apex of abdomen of 2, scale 1 mm; 10, suranal plate of d, scale 1 mm; 11, paramere, scale 0.2 mm; 12, lateral sclerite of
endosoma, scale 0.2 mm; 13, left half of suranal plate of Prilomera oribasis Bred. 3, scale Imm; 14, idem, P. laelaps Bred.;
15, idem, P. dorceus Bred.; 16, idem, unnamed new species.
148
Material. — Sulawesi Utara: Dumoga Bone National
Park, Toraut, Fischteich südl. Base Camp, 18. X. 1985, 14
12; Danau Alea, Bergsee bei Kotamobagu, 1500m, X.
1985, 19; S. Coast, Malibagu, mangrove, 8. XI. 1985, 2d
32 (no eggs). Sulawesi Tenggara: Desa Kagunyala, N8908,
19; Aopa Marsh, N8913, 4d 59 (34 59 apterous); Jalan
Asera, N8918 1d 19; Sungai Kolaka N8921B, 2d 29;
Tamborasi, N8925, 48 69 (on the sea, without eggs);
Mowewe, N8924, 19; Pomalaa, N8930 12, N8931 39 (2
apt. ); Teluk Kendari, N8948, 48 6%. Buton: Desa
Gareng-Gareng, N8937, 15 29 (19 apt. ). West Flores:
19, leg. F. A. F. Verheijen (RMNH). Macropterous unless
otherwise indicated.
Distribution. - Widespread: Indonesia (except
Sumatera), Mariana Is, New Guinea, N and E
Australia to Cook Is.
Limnogonus hungerfordi Andersen
(fig. 5)
Limnogonus hungerfordi Andersen, 1975: 46-47, figs. 115,
116, 120-122.
Material. — INDONESIA. Sulawesi Utara: 50 km N
Kotamobagu, Thermalquellen, 26. X. 1985, 19, leg. G.
Zimmermann. Sulawesi Tengah: Palu 60 km SE of: Lore
Lindu N. P., Danau Tambing and brooklets, UTM SJ95,
1600m asl, Pandanus, 7. XII. 1985, 1d, leg. J. van Tol.
Sulawesi Tenggara: Wawonggole, N8901, 6d 72 (36 29
apterous), N8902, 19 apt. ; Kendari, N8903A, 1? apt.,
12 macr. ; Desa Kagunyala, N8905, 18 19; N8907, 16;
Sungai Sampara, N8910, 38 29 (18 apt. ), N8911, 1d
19, N8912, 2d 49; Jalan Asera, N8916, 28 32, N8917,
5d 89, N8918, 236 169 (26 apt. ); Pomalaa, N8928 56
29 (48 29 apt. ), N8929 5d 49 (apt. ); Mowewe,
N8932, 4d 59 (36 19 apt.), Teluk Kendari, N8945, 16.
P. Buton: Desa Gareng-Gareng, N8937 12; Mangrove
swamp, N8939, 1d 19 apt. ; N8943, 18 19. Irian Jaya:
Neth. New Guinea Exp., Star Range, Bibil, 1290m, 2?
(RMNH). — MALAYSIA: Sabah: 60 km W Lahad Datu, DVFC,
building site staff quarters, ML-light, 117 48E 4°58’N,
150m, ML-light, 28. X. 1987, 16; 23 km W Sandakan,
Sepilok tree tower, 118°06’E 5°49’N, 0-100m, ML, 1. XI.
1987, 1 ; 20 km W Sandakan, Sepilok-laut, nr. Resthouse,
118°06’E 5°49’N, Om, ML-light, 4. XI. 1987, 19; 23 km
W Sandakan, Sepilok, 118°06’E 5°49’N, 0-100m, ML-
light, 31. IX. 1987, 16; 12.5 KM S Nabawan, Kg.
Pamuntaria, nr river, 116°27’E 5°02’N, 400m, ML-light,
16. XI. 1987, 1d, leg. J. Huisman & R. de Jong (RMNH).
Macropterous unless otherwise indicated.
Distribution. - Widespread: Taiwan, Malay
Peninsula, Philippines, Indonesia to New Guinea and
NE Australia (Andersen 1975).
Limnogonus nitidus (Mayr)
(fig. 3)
Hydrometra nitidus Mayr, 1865: 443.
Limnogonus nitidus, Lundblad 1933: 387-388, fig. 122 (re-
description); Andersen 1975: 62-65, figs. 191, 192, 195-
203, 205-208; Selvanayagam & Rao 1989: 474-484 (bio-
CHEN & Nigser: Gerridae from Sulawesi
logy); Nieser & Chen 1992b: (in print).
Material. — INDONESIA. Sulawesi Utara: Dumoga Bone
N. P., Toraut alt. 210m, sec. growth, at light, 28. VII.
1985, 18 19, leg. J. Huijbregts; S coast, Malibagu,
Mangrove, 8. XI. 1985, 1d 29 (no eggs). Sulawesi Tengah:
Palu, 65 km SSE of: Lore Lindu N. P., Marena shelter,
UTM-SJ62, 600m, 15. XII. 1985, 29, leg. van Tol &
Krikken (RMNH). Sulawesi Tenggara: Wawonggole, N8901,
1d; Sungai Sampara, N8912, 1 2; Jalan Asera, N8916 16;
N8917, 2d; N8918, 1d 49; Sungai Kolaka, N8921B, 24
32; Tamborasi, N8925 (on the sea), 38 49 (39 without
eggs); Wolo, N8926 (pools on banks), 1d 19; Pomalaa,
N8927, 16 (apterous) 19; N8930, 19; N8931, 19. P.
Buton: Desa Gareng-gareng, N8936, 19. — MALAYSIA:
Sabah, 16 km NE Tenom, Agr. Res. Station, 115°59’E
SEEN, 180m; DE eV 8 GRA STE uisrman:
Macropterous unless otherwise indicated.
Distribution. — Widespread: Maldive Is., India,
Indochina, Indonesia to Sulawesi, Sumbawa and
Flores.
Neogerris Matsumura, 1913
A tropicopolitan genus which until recently was
usually treated as a subgenus of Zimnogonus, with
which it deals habitat preference. A revision of Old
World species is given in Andersen (1975). Only one
species occurs in Sulawesi.
Neogerris parvula (Stàl)
Gerris parvula Stàl, 1859: 265.
Limnogonus parvulus, Lundblad 1933: 384-387, fig. 121, pl.
11 (redescription).
Neogerris parvula; Andersen 1975: 86-89, figs. 288, 289,
292, 297, 298, 300, 302-306, 309-312.
Material. — Sulawesi Tenggara: Wawonggole, N8901,
19, N8902, 38 19 (24 apterous); Desa Kagunyala,
N8905, 19; N8906, 28 19 (18 apt. ), N8907, DG DY
apt., N8908, 12; Sungai Sampara, N8912, 1d; Aopa
marsh, N8913, 5d 29 (5d 19 apt. ); Jalan Asera, 18 19,
N8918, 1435 149 (28 apt. ); Tamborasi, N8925, 19 (on
the sea); Pomalaa, N8927 1% apt., N8929, 18 19 (apt. ),
N8931, 18 39 (29 apt. ), 1 lv V. — Maluku: Bacan,
Wayauna, alt. ca. 50m, sec. growth, eutrophic pool, 06-07.
VII. 1985, 12. Macropterous unless otherwise indicated.
Distribution. — Widespread: SE tip of Arabian
Peninsula, SE Iran, India Indochina, Macao, Amoy,
Taiwan, Ryu Kyu Is., Philippines, Indonesia, New
Guinea and Solomon Is.
Aquarius Schellenberg, 1800
This genus is not represented in Sulawesi. The
nearest localities where species have been encountered
are Sumatera and West Australia (Andersen 1990).
149
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Limnometra Mayr, 1865
The Indo-Australian species of this genus have
been reviewed recently (Nieser & Chen 1992a). This
is essentially a SE Asian and Australian genus with
thirty known species of which seven occur in Sulawesi
and one on Buton. Sulawesi is the region richest in
species for this genus. For a key to the species and
other details readers are referred to Nieser & Chen
(1992a).
Tenagogonus Stàl, 1853
This genus is represented in Africa, Myanma and
Sumatera through Indonesia and the Philippines to
New Guinea. Not recorded from Sulawesi.
Tenagogonus akanthinos sp. n.
(figs. 6-8)
Type material. - Holotype à, allotype 9, 4 paratype d:
E. Malaysia, Sabah, Danum Valley, 70 km W Lahad Datu,
Main Trail West 5 South 3, 150 m, narrow creeks in rainfo-
rest, 16. XII. 1989, sample Sab. 70, M. J. & J. P. Duffels, all
apterous (ZMA, 2 paratypes NC).
Description
Apterous specimens. - Dimensions. Length d 6.0-
7.6, 2 6.8; width of head 6 1.23-1.50, $ 1.41;
width of thorax d 2.22-2.95, £ 2.80. Colour. Pale
brown, legs medium brown, antennae dark brown.
Eyes and apical segment of rostrum shiny dark casta-
neous. Interoculus with dark brown V-shaped me-
dian mark and margins along eyes. Pronotal lobe with
blackish margin all around and a dark median longi-
tudinal stripe. Pleura with a dorsal blackish band
which is very broad and anteriorly dived in an upper
and lower part by a narrow light line on mesopleuron.
Ocellated spot dorsally in front of mesoacetabula,
dorsal edge of connexiva and lateral margins of abdo-
minal tergites dark brown to blackish. Venter with
mesoacetabular suture and patch anteriorly black.
Some variable and often ill defined brownish marks
on meso- and meta- sternum and first abdominal
sternum. Sometimes somewhat similar to the pattern
in 7. maai Hungerford & Matsuda.
Length of antennal segments I : II : III : IV male
2.1 : 1.7 : 2.4 : 2.8; female 1.9 : 1.3 : 1.9 : 2.5.
Rostrum moderately slender, reaching slightly over
one third of mesosternum. Posterior lobe of prono-
tum rather narrow and exposing small parts of meso-
notum around its tip. Connexiva rather broad, no
connexival spines. Last abdominal segment, however,
with a pair of dorsocaudolateral projections, better
developed in male than in female, which look like
connexival spines. Hind coxae reaching halfway
fourth abdominal sternite in male, reaching third in
female. Last abdominal sternite as ling as three prece-
150
ding ones. In male last sternite with genital segments
as long as preceding part of abdominal venter. Leg
measurements, see table 1.
Male first genital sternite with a mediocaudal
groove bordered by a pair of broad caudally slightly
diverging carinae (fig. 6). Ninth segment without ob-
vious hair tufts. Endosomal sclerites as in fig 8.
Female, caudal margin of last abdominal sternite
with small lateral spines and a short, broad median lo-
be (fig. 7).
Comparative notes. — At first sight very similar to
T° pravipes Bergroth (= Limnometra brevis Lundblad,
= T. robustus Hungerford & Matsuda, see Polhemus
1991), which lacks, however, the spine like projec-
tions caudally on segment 7 an differs in details of ge-
nital segments. 7. kampaspe Kirkaldy which has cau-
dal projections on 7th segment in male, has these
placed more ventrally, has a different colour pattern,
and different genital segments. 7. maai Hungerford
& Matsuda (= T. quinquemaculata Miyamoto) (see
Polhemus 1991), which may look similar and occurs
in the same region, is smaller (6 length 5.5 mm,
width of thorax 1.64), more slender (over three times
as long as wide) and has more pronounced spine-like
projections of the last abdominal segment.
Etymology. — Akanthinos (Greek adjective mea-
ning spinose) refers to the spine like projections of
seventh abdominal segment, which differentiate this
species from 7. robustus.
CYLINDROSTETHINAE Andersen, 1975
Cylindrostethus Mayr, 1865
A tropicopolitan genus, the species live mostly on
shaded, quiet parts of streams, like most representati-
ves of the genus Aquarius. At first sight, species of
both genera are quite similar in size and general sha-
pe. Two species have been found in Sulawesi. The
comparative notes under C. brachyakanthinos sp. n.
will serve to separate them.
Cylindrostethus persephone Kirkaldy
Cylindrostethus persephone Kirkaldy, 1899: 508; Hungerford
& Matsuda 1962: 97-99, pl. 3 figs. E, F, pl. 5 figs. h, i
(redescription).
Material. — Sulawesi Utara: Dumoga Bone National
Park, Tumpah river nr. confl. Toraut, UTM XL0063, c.
210 m als., 18. V. 1985-A, 25 19 (1 copula); same 23. V.
1985-A, 18 29 (1 copula); same 3. VI. 1985-A 26 39 3
lvv, leg. J. van Tol. Dumoga Bone N. P. Tumpah R.
Staustufe, 13. X. 1985, 38 32; Tumpah R., Sandinsel u
Stromschnelle, 19. X. 1985, 48 39; Dumoga R. 22. X.
1985, 2d 29, leg. G. Zimmermann; Str. nach Malibogu,
Bach, 18. XI. 1985, 36 39 leg. G. Zimmermann. —
Sulawesi Tengah: Palu, gardens and agricultural fields, 28.
XI. 1985-A, 1d 19, leg. J. van Tol. — Sulawesi Tenggara:
Sungai Sampara, N8911, 12d 139. All apterous.
Distribution. — Sulawesi.
Cylindrostethus brachyakanthinos sp. n.
(figs. 9-12)
Type material. — Holotype d, allotype 2, 14 paratypes
3, 15 paratypes 9, Sulawesi Tengah, Luwuk area, Sungai
Batui, 15 Nov 1989, J. van Tol (RMNH). Additional paraty-
pes (larvae excluded): Luwuk area, S. Batui, 17 Nov. 1989,
364 189; Luwuk area, Sungai Biak, 21. X. 1989, 95 89;
NE Luwuk, 24. I. 1989, 9d 39, J. van Tol (RMNH). NE
Luwuk, Sungai Bantayan, near road Kayutanyo-Siuna,
(0°47’S 123°00’E), 50-100m, medium sized river in slightly
disturbed lowland rainforest, sta. Sul 2, 7. Oct 1989, 184
129 1 lv; SW Luwuk, between Desa Seseba and Singsing
Camp (ca 1°09’S 122°31’E), 80m, narrow stream in low-
land rainforest, sta. Sull2, 4 Oct 1989, 58 59 1 lv; SW
Luwuk, Sungai Batui at Singsing Camp (ca 1°09’S
122°31’E), 90m, middle sized stream in lowland rainforest,
sta. Sull4, 14 Oct 1989, 48 59 13 lv; SW Luwuk, Totop
Camp along Sungai Batui (1°09’S 122°31’30”E), 120m,
lowland rainforest, sta. Sull8, 19/21 Oct 1989, 136 209
(12 macropterous) 25 lv, leg. J. P. Duffels (zmA). Apterous
unless otherwise indicated.
Description
Apterous specimens. - Dimensions. Length d
10.9-11.7, 2 13.9-15.2; width of head d 1.60-1. 64,
® 1.80-1.89; width of thorax d 2.50-2.54, ® 3.24-
3.48. Colour and pilosity. Dorsally generally black
with conspicuous stripes of silvery hairs at dorsal mar-
gins of pleura and ditto dots on acetabula. Median
band dorsally on head, thorax and abdomen glabrous
in spite of short golden pubescence especially on ab-
dominal tergites. Apex of head with dirty yellowish
pilosity at sides, becoming blackish with sparse lighter
hairs posteriorly. Fulvous spot on base of head and
narrow median fulvous stripe on pronotum rather in-
distinct. Lateral margins of mesonotum broadly with
short thick velvety black pilosity. Outer (upper) mar-
gin of connexiva fulvous. Anterior femur black with
yellowish base and variable yellowish stripes on inner
and outer faces, tibia and tarsus blackish. Middle and
hind legs blackish except for coxae and trochanters
mainly yellowish. Venter with silvery pilosity on dark
base in males; background except for dark spots ante-
riorly on thorax, yellowish in females.
Length of antennal segments I : II : III : IV, male
1.8: 1.1 : 0.6 : 0.95, female 2.1 : 1.15: 0.6 : 1.0. In
dorsal view head longer than pronotum (1.2 in males,
1.4 in females). Pronotum medially depressed.
Mesonotum not depressed but with a very thin me-
dian groove over its length in female, only in posteri-
or part in males. Caudal part of mesonotum in fema-
le without oblique ridge. Metanotum flat, its rear
margin sinuate. Connexiva slightly less than half the
CHEN & Nieser: Gerridae from Sulawesi
width of tergites, flaring laterally in males, more or
less erect in females (except when abdomen swollen
with eggs). Connexival spines short, reaching halfway
first genital segment in both sexes, tips not or hardly
divergent caudally. Metasternum with lateral groove
leading from omphalium, deep and well marked,
much more prominent than caudal margin of metas-
ternum, which is subequal to or slightly longer than
median length of first two abdominal sternites com-
bined. Leg measurements, see table 1.
Male. Suranal plate in male as shown in fig. 10;
wings convex lateroproximally. Last two abdominal
sternites of subequal median length, posterior margin
of last abdominal sternite evenly concave, posterior
margin of first genital sternite slightly produced to
nearly straight medially, their median length sub-
equal. Parameres and lateral sclerite of endosoma as in
fies 11,410,
Female. Last abdominal sternite medially slightly
longer than penultimate, its hind margin slightly and
obtusely produced medially. Macropterous female
(apex of hemielytra broken off) as apterous except for
development of pronotum and wings. Length 14.5,
humeral width of pronotum 2.35, thoracic width
3.25. Hemielytra dark brown with blackish veins.
Comparative notes. — Very similar to Cylindro-
stethus persephone Kirk., also from Sulawesi. There is a
series of small differences: C. brachyakanthinos is
slightly larger on average, the fulvous marks on head
and pronotum are less distinct, the male abdomen is
darker, the posterior part of mesonotum in female is
not differentiated. The connexival spines in C. bra-
chyakanthinos are shorter (fig. 9), the wings of the su-
ranal plate of male are convex basally (straight in C.
persephone, fig. 10) and the paramere is broader at ba-
se and consequently more tapering in C. brachyakant-
hinos (fig. 11). The endosomal sclerites are nearly
identical, only the laterals show very small differences
in shape (fig. 12).
Etymology. — Brachyakanthinos (Greek adjective
meaning ‘with short spines’) refers to the short con-
nexival spines.
EOTRECHINAE Matsuda, 1980
No species of this subfamily have been reported
from Sulawesi. Members of this subfamily live rather
cryptically on wet places along waters. If any are to be
found they will most likely belong to the widespread
genus Amemboa Esaki, of which species have been re-
corded from Java and Mindanao (Polhemus &
Andersen 1984).
RHAGADOTARSINAE Lundblad, 1933
151
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
al 30
Figs. 17-31. — 17, lateral view (left) of seventh abdominal segment of Ptilomera doreceus Bred., 2, scale 2 mm; 18, idem, P.
oribasus Bred.; 19, idem, P. laelaps Bred.; 20, idem, unnamed new species; 21, idem, P. sumizone Esk. (redrawn after
Hungerford & Matsuda 1965); 22, Metrocoris breviculus sp. n., holotype, scale 0.5 mm, paramere; 23, idem, endosomal scle-
rites, lateral view; 24, forelegs of male Ventidius kurtokalami sp. n. paratype (Halobatinae), scale 1 mm, 25, idem, Metrocoris
breviculus sp. n., holotype; 26, Parameres of Ventidius xiphibion sp. n., scale 0. 25 mm; 27, idem, V. xyele sp. n.; 28, idem, V.
kurtakalami sp. n.; 29, lateral view of endosomal sclerites of Ventidius xiphibion sp. n., scale 0. 25 mm; 30, idem, V. xyele sp.
n.; 31, idem, V. kurtakalami sp. n.
Rhagadotarsus Breddin, 1905 Rhagadotarsus kraepelini Breddin
IX small tropical genus with one species in Asia, Rhagadotarsus kraepelini Breddin, 1905: 136-139, figs. 12-
which is also known from Sulawesi, and one (attribu- I ee Le pl. 11; Cheng &
ted to a different subgenus) in Africa. Re NE SN Go
152
Sulawesi Tenggara: Wawonggole, N8901, Id 19;
Sungai Sampara, N8909 4d 79 (29 macr. ). Apterous un-
less otherwise indicated.
A very widespread species, from Sri Lanka,
Myanma and SW China to Philippines, Sulawesi and
Sumbawa.
PTILOMERINAE Esaki, 1927
Ptilomera Amyot & Serville, 1843
Widely distributed in tropical Asia. Large and
conspicuous inhabitants of streams usually near
places with rather strong current on which they ven-
ture when disturbed. Regularly on very narrow
streams. In spite of their size and obvious morpholo-
gical specialization identification is difficult as there
are several species groups consisting of very similar
species. At present the genus is under revision by D.
A. Polhemus (Bishop Museum, Honolulu, Hawaii);
material of our expeditions will be included in
Polhemus’ study. We present a preliminary key to
Sulawesi species (Polhemus & Polhemus 1986).
Key to Prilomera of Sulawesi
1. Specimens with posterior sclerite in dorsal view
(suranal plate) with a pair of lateral wings, a pair
of caudally directed more or less curved styli
(which are actually the parameres) and in ventral
view a large boat-shaped posterior segment (the
pygophore) with a pair of lateral extensions, ma-
VEREEN Ba ie Kuhn ER ade 2
— Apex of abdomen in lateral view with a broad la-
teral plate on last segment which hides the apex of
abdomenytemaleste tect EN RE ee 7
2. Lateral wings of suranal plate with rostral and
caudal margins nearly parallel in apical part (fig.
13) [Sulawesi Selatan & Tengah] .......................
RO ty. LOTTA RER P. oribasus Breddin, 1901
— Rostral and caudal margins of lateral wings of su-
ranal plate convergent lateroapically ................ 3
3. Lateral wings of pygofer long and slender, apical-
ly acutely pointed and laterally surpassing the
wings of suranal plate [Sulawesi Tengah] … … …
ERSTE a nn P. pamphagus Breddin, 1901
— _ Lateral wings of pygofer shorter, laterally at most
as long as the wings of suranal plate or with bro-
ddlvatnuncaterapexape eee a ete eee 4
4. Caudal margins of lateral wings of suranal plate
perpendicular on longitudinal body axis (fig. 14)
— Caudal margins of lateral wings of suranal plate
pointing obliquely caudad compared to longitu-
dinalbodyiaxiss m. int ae ee eee 6
5. First genital segment strongly swollen dorsally in
caudal half [Sulawesttlienpeara| ren.
CHEN & NIESER: Gerridae from Sulawesi
a as At OI Bel ARR mu P. sumizone Esaki, 1925
— First genital segment not swollen dorsally
(Sulawesi Selatan] … … P. laelaps Breddin, 1901
6. Caudal margins of lateral wings of suranal plate
nearly straight (fig. 14) [Sulawesi Utara] … … …
hs RIO Pees AL LD P. dorceus Breddin, 1901
— Caudal margins of lateral wings of suranal plate
convex (fig. 15) [Sulawesi Tenggara] ..................
IC SER Meee ee A oe fai Le ai Ptilomera sp. n.
7. Dorsal lobe of seventh abdominal segment (later-
al view) long and spine-like, reaching beyond well
developed connexival spines (fig. 17) . P. dorceus
— Dorsal lobe relatively shorter ........................... 8
8. Dorsal lobe well developed and narrow, reaching
to apex of connexival spines (fig. 18) P. oribasus
=" Dorsalllobe/broader eee 9
9. Incision between dorsal and ventral lobe of seven-
th abdominal segment in lateral view wide,
ventral lobe reduced (fig. 19) .............. P. laelaps
— Incision narrow, ventral lobe more distinct ... 10
10. Caudoapical margin of dorsal lobe of seventh ab-
dominal'seementivert ale 11
— Caudoapical margin of dorsal lobe of seventh ab-
dominal segment pointing obliquely caudad (fig.
LT, Ptilomera sp. n.
11. Lateral lobe of seventh abdominal segment vent-
robasally with an incision (fig. 21) .. P. sumizone
— Lateral lobe of seventh abdominal segment vent-
robasally without an incision ......... P. pamphagus
Pleciobates Esaki, 1930
One species known from the Malay Peninsula, P.
tuberculatus Esaki, 1930.
Rheumatogonus Kirkaldy, 1909
Three species, of which À intermedius Hungerford,
was described from Sumatera and occurs in the Malay
Peninsula too (Cheng & Fernando 1969). The other
two occur in the Philippines, one extending into N.
Borneo.
Rheumatogonus borneensis Esaki
Rheumatogonus borneensis Esaki, 1927: 267-268, fig. 3d;
Andersen 1967: 267-272, figs. 12-14, 16-17 (redescrip-
tion).
Material. - MALAYSIA, Sabah, Danum Valley, 70 km W
Lahad Datu, Main Trail W 12, 180 m, 2. XII. 1989, narrow
stream, sample 52, 76 29 ; same, 4 km S main trail W 5, nr.
Sungai Segama, 150 m, 3. XII. 1989, middle sized stream
and water fall, sample 54, 1d 19; same, Main Trail West 5
South 3, 150 m, 18. XII. 1989, narrow creeks in rainforest,
sample 70, 2d 59, leg. M. J. & J. P. Duffels (zma), all ap-
terous.
153
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 1 Measurements of leg segments of Gerridae described in this paper. In species marked with an *, the tarsal segments
have not been measured separately, the entire length is given under tars1.
femur tibia tarsl tars2
Cylindrostethus brachyakanthinos
d fore leg 3.1 2.9 0.2 0.6
d middle leg 11.1 7.4 3.5 0.8
d hind leg 11.4 6.0 0.5 0.3
? fore leg 3.6 555) 0.3 0.65
9 middle leg 12.1 8.5 5.1 1.0
9 hind leg 122 8.9 0.7 0.4
Metrocoris breviculus*
d fore leg 2.03 1.68 0.70
3 middle leg 5.65 3.93 1.82
d hind leg 4.79 2.50 0.72
9 fore leg 1.74 1.43 0.70
? middle leg 4.80 3.48 1.70
9 hind leg 4.20 2.42 0.71
Potamometropsis anomalis
d fore leg Sell 4.0 19) 0.85
3 middle leg 19.2 Poll 3.0 0.47
3 hind leg 13.2 4.6 0.10 0.12
? foreleg 5A 3) 2.1 0.98
? middle leg 129 72 34 0.50
? hind leg 13.2 5.0 0.10 0.13
Rheumatometroides drepanephoros
d fore leg 0.85 0.7 0.09 0.36
3 middle leg 2112 2:59, 0.79 0.77
G hind leg 2.24 0.80 0.11 0.27
9 fore leg 0.85 0.74 0.09 0.38
? middle leg 2.18 2.59 0.81 0.79
2 hind leg 2.30 0.88 0.12 0.31
Rheumatometroides makraitos
d fore leg IR 0.96 0.1 0.5
d middle leg 2.82 3.26 1.18 1.08
d hind leg 3.01 1.16 0.19 0.38
Potamometropsis Lundblad, 1933
Three species, one from Sumatera and two from
the Philippines are known (Hungerford 1957). We
add a new species from North Sulawesi.
Potamometropsis anomalis sp. n. (figs. 38-41)
Type material. - Holotype d allotype 9 (zma), 1d 89
paratypes (1? macropterous): Sulawesi Utara, Dumoga
Bone N. P., Tumpah R. Staustufe (barrage), 23 Oct. 1985,
leg. G. Zimmermann (19 MBBJ, 1d 3® Nc, 19 RMNH, 42
zc). — Additional paratypes: Sulawesi Utara, Dumoga Bone
N. P., Base Camp, bridge, 16. X. 1985, leg. G.
Zimmermann, 3d 49 (1d 29 macropterous) (Nc, zo);
Mumipahy River beach, 198 xo 19855) 012) lec G
Zimmermann same, Edwards Camp, Tumpah River, 900m,
23.11.1985, Stat. 28 (Project Wallace), 16 leg. J. P. Duffels
(ZMA), apterous unless otherwise indicated.
Description
Apterous specimens. — Dimensions. Length à 7.8-
8.4, 2 9.4-10.3; width of head d 1.41-1.45, © 1.41-
1.48; thoracic width & 2.46-2.50, 2 2.90-3.05.
Colour black. Head yellowish brown with a large
154
femur tibia tars] tars2
9 fore leg 1217. 0.92 0.12 0.60
? middle leg 3.17 3.59 1.30 1.25
? hind leg 3.05 1.25 0.21 0.37
Tenagogonus akanthinos
d fore leg Sal 255 0.35 0.64
3 middle leg 8.1 5.6 2.0 0.58
3 hind leg 7.6 3.2 0.64 0.62
? fore leg 2.9 2.3 0.3 0.57
? middle leg 6.5 4.8 2.0 0.60
? hind leg 6.4 2.9 0.55 0.51
Ventidius kurtokalami*
3 fore leg 1.2 1.0 0.40
3 middle leg 3.3 1.8 181
d hind leg 3.6 1.5 0.5
? fore leg 1.6 1.0 0.5
? middle leg 3.6 1.8 el
9 hind leg 4.0 17 0.6
Ventidius xiphibion*
3 fore leg 1.1 0.9 0.35
3 middle leg 32 1.9 1.1
d hind leg 39 1.5 0.4
? fore leg 1.2 0.9 0.3
? middle leg 33 169 Le
9 hind leg 3.7 1.6 0.4
Ventidius xyele*
d fore leg 12 1.0 0.4
3 middle leg 3.4 1.9 1.1
3 hind leg 3.9 127 0.5
? fore leg 12 1.0 0.4
? middle leg 3.5 1.9 1.1
9 hind leg 3.9 1.8 0.5
median blackish spot anteriorly and a pair of smaller
blackish spots in posterior part of interoculus. Venter
of head, rostrum except for apex, prosternum, poste-
rior half of mesosternum, metasternum, abdominal
venter, acetabula, coxae and ventral and dorsal stripe
apically and proximally widened on fore femur, yel-
low. Antennae and legs dark castaneous.
Mesosternum, metasternum and abdominal venter
with dense short silvery pilosity. A broad band of sil-
very pilosity on upper part of mesothoracic pleuron
extending on dorsum of metaacetabulum. A pair of
submedian bands of silvery pilosity on pro- and mes-
onotum, in some specimens hardly developed. A
band of silvery pilosity just ventrally and just dorsally
of base of connexiva.
Length of antennal segments I : II : HI: IV d 3.51
: 1.43 : 1.23 : 1.09, © 3.67 : 1.33 : 1.22: 1.11. So se-
cond segment longer than third. Pronotum slightly
wider than head, median length 0.8. Median length
of mesonotum three times as long as pronotum and
metanotum. Fore tibia laterally compressed, with a
slight constriction subapically, accentuated by tufts of
setae proximally and distally of constriction. Inner
apical process bluntly triangular, equally developed in
both sexes. First segment of fore tarsus slightly over
two times as long as second (table 1). Middle and
hind femur of subequal length in both sexes someti-
mes middle > hind, sometimes the reverse. Leg meas-
urements, see tabel 1.
Male. Tergite 7 two times as long as tergite 6, ster-
nite 7 caudally broadly and shallowly emarginate.
Tergite 8 twice as long as sternite, caudally broadly
rounded. Paramere, well developed, rather broad
compared to other species in the genus (fig. 40), in
rest hidden in the prominent pygophore. Endosoma,
fig. 41.
Female. Tergite 7 nearly two times as long as 6.
Sternite 7 twice as long as 6, with a broad mediocau-
dal lobe, in ventral view covering most of the genital
segments (fig. 39).
Macropterous form as apterous except for develop-
ment of thorax and pronotum. Length (including he-
mielytra) 10.0-10.2, humeral width of pronotum
2.4-2.5. Pronotum mostly shining black with rough
impressed punctures, median length 2.9. Anterior lo-
be divided from posterior lobe by a medially inter-
rupted ridge caudally accentuated by a groove with
some silvery pilosity, posterior part of anterior lobe
and anterior part of posterior lobe with a median lon-
gitudinal carina. Hemielytra rather membraneous,
medium brown with swollen, slightly darker, veins.
Etymology. — Anomalos, a greek adjective meaning
aberrant.
Comparative notes. — P. anomalis differs from
other species in the genus by having second antennal
segment longer than third and first segment of fore
tarsus over twice as long as second. The male has a
shorter and broader paramere than other species, the
abdominal sternite 7 of the female is quite different
from the other species.
Remarks. — The macropterous female from
‘Staustufe’ has the hemielytra broken off which seems
to be the normal condition in this genus (Hungerford
1957). The macropterous specimens from ‘bridge’
have the hemielytra complete, these are recently
moulted specimens, with belly still soft (bending in-
ward under slight pressure). Its general form, absence
of specialization of the venter in female and presence
of claws on middle and hind tarsi place this species in
Potamometropsis. This species differs in several charac-
ters from all its congeners (see comparative notes).
Pending a cladistic analysis of the entire subfamily we
refrained from describing a new genus for it.
We have seen a teneral female from Sabah, Kg.
Sapulut, 116°36’E 4°38’N, leg. Huisman in RMNH,
which might belong to another species of this genus
but as the abdomen is shrivelled, this material is in-
sufficient for a description.
CHEN & NIESER: Gerridae from Sulawesi
HALOBATINAE Bianchi 1896
Halobates Eschscholtz, 1822
Exclusively marine. Several species live on the open
ocean, have wide distributions and are only found
near or even on the shore after high winds. Halobates
micans Eschscholtz is tropicopolitan and the only spe-
cies found in the open Atlantic. It has been recorded
from Laut Sulawesi. A second species has been recor-
ded from the same area : Halobates germanus White
(Herring 1958). These species are easily separated by
size, H. micans is over 4-4.5 mm long, H. germanus
3.5-3.9 mm. We have seen two specimens of the lar-
gest species in the genus H. princeps which measures
6-7 mm. So far this is the only species recorded from
the coast of Sulawesi (Andersen 1991b). The genus
was revised by Herring (1961); a survey of phylogeny
and evolution is given by Andersen (1991a).
Halobates princeps White
Halobates princeps White, 1883: 44, pl. 1 fig. 3; Herring
1961: 267-270, fig. 70 (redescription); Andersen 1991b:
158, fig. 8.
Material. — Sulawesi Utara, N. Coast, Molosso, reef, rock
pools, 16. X. 1985 2 9, leg. G. Zimmermann.
Distribution. — Range included Solomon Isles, N
New Guinea, NW Australia, Nusa Tenggara, Malay
Peninsula, extreme S of Philippines and Palau
(Andersen 1991b). This species has apparently been
found on more or less open sea (the type is from
‘Celebes Sea’) and on coastal habitats.
Metrocoris Mayr, 1865
Widespread, from the Arabian Peninsula to SE
Asia. The species from Malay Archipelago and
Philippines have recently been revised by Polhemus
(1990). The genus as a whole will be revised by Chen
& Nieser (in prep.). Two species are known from
Sulawesi.
Key to Metrocoris from Sulawesi
1. Transverse black band on pronotum covering an-
terior third to nearly half of pronotum, body
length = CS mn rer ee M. celebensis
— _ Transverse black band on pronotum covering less
than anterior third of pronotum, body length 4-
RS) SOV TY, Bone pace aceericoe EE M. breviculus
Metrocoris celebensis D. Polhemus, 1990
Metrocoris celebensis D. Polhemus, 1990: 8-9, figs. 3, 19-22,
47).
155
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Only known by the type series from Lore Lindu
National park (Sulawesi Tengah), we have seen 14
1? apterous paratypes in ZMA.
Metrocoris breviculus sp. n.
(figs. 22-23, 25)
Type material. — Holotype à, allotype 9, 19 paratype:
Sulawesi Tengah, between Desa Seseba and Singsing Camp,
SW of Luwuk, c 1°09’S 122°31’E, 90m, narrow stream in
lowland forest, 14. X. 1989, Sta. Sull2, J. P. Duffels. Batui
River at Singsing Camp, 90m, middle sized stream in low-
land rainforest, 14-17. X. 1989, Sul. Sta. 14, J. Duffels 1 4
1 paratypes, all apterous (ZMA, NC).
Description
Apterous specimens. - Dimensions. Length d 4.3,
? 4.0-4.2; width across metaacetabula & 2.2-2.3, 9
2.3-2.4; width of head d 1.32-1.33, 2 1.29-1.31.
Colour. Dark pattern variable, somewhat reduced
compared to other species. Interoculus without dark
mark or anterior half dark with caudal arms along in-
ner margins of eyes. Dark mark of pronotum T-sha-
ped, dark mark of mesonotum T-shaped with lateral
edges of T produced caudally. Metanotum with a
broad anterior and mesal band. First abdominal tergi-
te black with a pair of yellowish spots, connexiva an-
teriorly blackish, posteriorly yellowish, remaining ter-
gites yellowish with blackish margins. Mesopleura
yellow without longitudinal dark stripe. Fore femur
with an apical dark ring, which may extend over the
apical quarter of femur, ventral longitudinal stripe
present, other indistinct or absent. Venter yellowish,
area around metasternal scent gland opening dark to
black in both sexes.
Width of an eye slightly less than its posterior
width, 0.42 and 0.46 in 6, 0.50 and 0.53 in ® re-
spectively, width of interoculus 0.48. In lateral view
posterior half of eye covering anterior three fifths of
propleura. Length of antenna d 4.6, 2 3.7. Length
of antennal segments 1: II : IN : IV d 1.92: 1.11:
0.91 : 0.60, 9 1.51 : 0.79 : 0.81 : 0.60. Antenna in
male reaching well beyond the apex of abdomen, in
female reaching the apex of abdomen, antenna in ma-
le distinctly thicker than in female. Antennal seg-
ments II and III in males with curved woolly pilosity,
especially developed on ventral face, its thickness over
half the width of the segments. Pronotum not bul-
bous, its width 1.4-1.5, slightly more than width of
head. Meso- and metapleura with blackish pilosity
which tends to become bristle-like at rear end of me-
tapleura. Male fore femur (fig. 25) not incrassate, ra-
tio length: width about 6, without stiff dark spinules,
long hair fringe on inner face, constriction, indenta-
tion or tooth; fore tibia without tooth like elevation.
Female fore femur, ratio length : width 6-6.5. Length
of hind trochanter 0.5 in both sexes, in female with-
156
out a tuft of long curved hairs apically. Middle and
hind femur without long hair fringe. Leg measure-
ments, see table 1.
Male. Pregenital tergite and genital segments rather
small, pregenital segment, length 0.2, width 0.5.
Parameres, fig. 22, endosomal sclerites fig. 23.
Female. Subgenital sternite large, its median length
subequal to the median length of preceding abdomi-
nal sternites together, caudal margin sinuate without
tongue-like caudal process, apex of abdomen visible
in ventral view.
Macropterous form unknown.
Etymology. — Breviculus (latin adjective meaning
rather small) refers to the small size of the species.
Comparative notes. — Recognized at once by its
small size in combination with yellowish venter, ab-
sence of mesopleural stripes and the paramere fig. 22.
M. breviculus is 1 mm or more shorter than the only
other species occurring in Sulawesi: M. celebensis D.
Polhemus.
Esakia Lundblad, 1933
Species of Esakia are very similar to Ventidius and
Ventidiopsis. The genus is considered by Matsuda
(1960) to have evolved from Ventidius by acquiring
some specialized characters such as the strongly modi-
fied third antenna of the male. Three of the six species
known occur in the Malay Peninsula, E. ventidioides
Lundblad (1933) was described from Sumatera.
Ventidiopsis Miyamoto, 1967
Monotypic; V. imadatei Miyamoto (1967) was de-
scribed from N. Borneo (Brunei).
Ventidius Distant, 1910
The largest and most widespread genus within this
group of small Halobatinae, 16 species have been de-
scribed. Greatest species density seems to be in the
Malay Peninsula (Cheng 1965, Lansbury 1990) but
various regions of its range have been poorly sampled.
Ventidius xiphibion sp. n.
(figs. 26, 29)
Type material. - Holotype d, allotype 2 (RMNH), 42d
(18 macropterous) 642 paratypes: Sulawesi Tenggara,
Small stream 8 km E of Sungai Sampara along road
Kendari-Wawotobi, N8911, 22 Feb 1989. — Additional pa-
ratypes: P. Buton: Stream just N of Bau-bau, N8935, 8 Mar
1989, 358 402 (12 macr. ); stream about 15 km E of Bau-
bau, N8942, 10 Mar 1989, 406 182 (19 macr. teneral).
Apterous, unless indicated otherwise. Samples N8935 and
N8942 contained also many larvae.
Description
Apterous specimens. Dimensions. Length d 2.7-
3.0, 2 3.0-3.3, width of thorax d 1.95-2.05, 9
2.20-2.32. Colour. Eyes and anterior half of intero-
culus blackish, posterior half of interoculus yellow.
Pronotum black with 5 small yellowish spots: antero-
median, one pair halfway laterally at posterior border
and a pair laterally at margins of eyes. These spots are
variable, specifically the three central ons may be
confluent. meso- and meta- notum with a large centr-
al bilobed yellow area (general pattern as in V. henryi
and V. werneri Hungerford & Matsuda, but with
light area larger although quite variable between se-
ries). A broad black longitudinal band, which may be
interrupted in lightly coloured specimens, on border
of notum and pleuron. Posterior part of metanotum,
first abdominal tergite black, with caudally tapering
lateral black bands on remaining abdominal tergites,
connexiva from second segment on yellow. Genital
segments black in male, laterally and caudally black in
female. Variable black dots anteriorly of acetabula.
Venter yellow with a pair of small black dots at ante-
rior point of mesoacetabular suture. Apex of rostrum,
antennae except for base of first segment, apex of fore
femur and rest of fore leg, middle and hind leg blac-
kish. Body dorsally with rather sparse short black pi-
losity giving the specimens a dull appearance.
Antennae when folded back over venter as long as
body in male, slightly shorter in female. Length of an-
tennal segments I : II : III : IV, male 1.22 : 0.48 : 0.34
: 0.46, female 1.23 : 0.48 : 0.44 : 0.50. Eyes twice as
long as head, overlapping lateral margin of prothorax
and anterolateral angles of mesothorax. Anterior and
posterior margins of pronotum medially concave. In
lateral view, mesonotum softly convex dorsally, me-
tanotum declivent. Intersegmental suture between
meso- and meta- notum as well as anterior margin of
first tergite indistinct or lost. Lateral longitudinal su-
ture of metanotum not reaching the level of
meso/metanotal intersegmental suture. Posterolateral
angle of metaacetabula bilobate. Ventral margin of
front femora nearly straight in female; convex, but
without tubercle in the middle, in male; ratio length:
width 5.7 in male, 6.4 in female. Leg measurements,
see table 1.
Male. Parameres asymmetrical, fig. 26. In the cent-
re of mesosternum is a small patch of minute black
bristles which are not on a tubercle.
Macropterous form. — As apterous except for deve-
lopment of pronotum and hemielytra. Length à 2.8-
3.3, ® 3.4-3.6, humeral width of pronotum d 1.65-
1.70, 2 1.80-1.95. Pronotum black, lateral margin
and apex brownish, anterior margin variable from
black to a predominantly yellow transverse band; a
pair of large pale yellow pear-shaped patches on cent-
re of disc. Hemielytra extending beyond apex of ab-
CHEN & NIESER: Gerridae from Sulawesi
domen, black with costal area and inner margin dark
brown.
Comparative notes. — The bilobed posterolateral
angle of metaacetabula, the sexual dimorphism of fo-
re femur, asymmetrical parameres and small patch of
black bristles on mesosternum of male place this spe-
cies in the subgenus Ventidoides Hungerford &
Matsuda. V. xiphibion differs from V. pulai Cheng,
which has a very similar colour pattern, apart from
being slightly larger on average, by lacking a tubercle
on fore femur of male, having the patch of minute
black bristles on male mesosternum developed, al-
though not on a tubercle, and different parameres.
The macropterous form of V. pulai has the light mes-
onotal spots fused so there is one large central spot.
Other Malay species differ in head coloration.
Etymology. — Xiphibion (greek: small sword) a
noun in apposition refers to the shape of left parame-
re.
Remarks. — This and the next are the first species of
this genus to be reported from Sulawesi. The locali
ties where V. xiphibion was taken are small to medi-
um sized streams, all three had turbid water due to
suspended loam. The species tends to aggregate at
places with little current.
Ventidius xyele sp. n.
(figs. 27, 30)
Type material. - Holotype d, allotype 2 (RMNH),
Sulawesi Utara, Dumoga Bone National Park, R. Toraut,
Maze, 16 Nov. 1985, leg. G. Zimmermann. — Paratypes
(the adults only) same data as holotype, 38 99; Dumoga
Bone N. P. Base Camp, 16. X. 1986, 18 19; Dumoga
Bone River, downstream of bridge, 22. X. 1985, 19;
Tumpah River, Beach, 19. X. 1985 8d 99, 28 lv III-V;
Tumpah River, Staustufe (low barrier), 23. X. 7d 79:
Malibagu Z. 8. XI. 1985, 54 19 (2d 1% macropterous);
Südküste, Strasse, Bach (S coast, road, rivulet), 18. XI.
1985,48 59 5 lv IV-V, apterous unless otherwise indicated
(zc, 88 89 Ne, 16 19 each in BPUH, JTPC, MBBJ, PCHC,
ZMA, ZMC).
Description
Apterous specimens. - Dimensions. Length d 2.9-
3.0, 2 3.0-3. 2, width of thorax d 2.0-2. 12, 9 2.20-
2.33. Colour, as in V. xiphibion except: pronotum
without yellow spots, meso/metanotal yellow mark,
smaller (but essentially of the same shape), black area
on border of notum and pleuron broader and not in-
terrupted.
Antennae when folded back over venter as long as
body in male, slightly shorter in female. Length of an-
tennal segments I: II: III: IV, male 1.29: 0.50: 0.41:
0.46, female 1.22: 0.50: 0.52: 0.48. Eyes twice as
long as head, overlapping lateral margin of prothorax
and anterolateral angles of mesothorax. Anterior and
posterior margins of pronotum medially concave. In
157
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
37
36
h_—_—_—_—_—_—_____—_—_—_—_____ +
Figs. 32-41. — 32, Rheumatometroides makraitos sp. n. paratypes, dorsal view of body of ®, scale 2 mm; 33, idem, apical ab-
dominal segments of & in lateral view; 34, idem, proctiger in dorsal view, scale 0.5 mm; 35, idem, endosomal sclerites in la-
teral view, scale 0. 2 mm; 36, Rheumatometroides drepanephoros sp. n., paratype à , scale 0.2 mm, proctiger in dorsal view; 37,
idem, endosomal sclerite in lateral view; 38, Potamometropsis anomalis sp. n., apex of abdomen ¢ paratype in lateral view; 39,
idem, apex of abdomen ? paratype in ventral view; 40, idem, d paratype, paramere; 41, idem, 3 paratype, endosomal scle-
rites, scales: 38, 39 1 mm; 41, 42 0.25 mm.
lateral view, mesonotum softly convex dorsally, me-
tanotum declivent. Intersegmental suture between
meso- and meta- notum as well as anterior margin of
first tergite indistinct or lost. Lateral longitudinal su-
ture of metanotum not reaching the level of
meso/metanotal intersegmental suture. Posterolateral
angle of metaacetabula bilobate. Ventral margin of
front femora nearly straight and without ventral
tooth in male, slightly concave in female; ratio length:
width 5.5 in male, 6.7 in female. Leg measurements,
see table 1.
Male. Parameres asymmetrical, fig. 27. In the cent-
re of mesosternum is a small patch of minute black
bristles which are not on a tubercle. Endosomal scle-
rites fig. 30.
Macropterous form as apterous except for develop-
ment of pronotum and hemielytra. Length d 3.1-
3.2, 2 3.33; humeral width of pronotum d 1.6-1.7,
9 1.74. Pronotum dark brown to black with a pair of
clavate yellowish spots on the centre constricted pos-
teriorly, in one male and the female with the light
158
brown posterior margin of pronotum. Hemielytra ex-
tending beyond apex of abdomen, proximally blac-
kish, apically dark brown with black veins, costal
margin medium brown.
Comparative notes. — Very similar to V. xiphibion
which has, however, a more extensive yellow pattern
dorsally and different parameres. The colour pattern
is intermediate between V. pulai and V. kuiteri
Hungerford & Matsuda, which both have, however,
a medioventral tubercle on fore femur in male and so-
mewhat different parameres.
Etymology. — Xyele (greek: knife, falchion) a noun
in apposition refers to the shape of left paramere.
Ventidius kurtokalami sp. n.
(figs. 24, 28, 31)
Type material. - Holotype à, allotype 9 E. Malaysia,
Sabah, Danum Valley, 70 km W Lahad Datu, 4 km S main
trail W5 nr. Sungai Segama, 150m, middle sized stream and
waterfall, 3 Dec. 1989, sample Sab. 54, M. J. & J. P. Duffels
(ZMA); paratype d Sabah, Danum Valley, 70 km W Lahad
Datu, main trail W 12, 180m, narrow creek, 2. XII. 1989,
sample Sab. 52, M. J. & J. P. Duffels (Nc). All apterous.
Description
Apterous specimens. - Dimensions. Length d 2.8-
3.0, 2 3.45; width of thorax d 2.0, ® 2.60. Colour.
Eyes and anterior half of interoculus blackish, poste-
rior half of interoculus yellow. Pronotum black, me-
so- and metanotum with a large central bilobed yel-
low area. An irregular broad black longitudinal band
on border of notum and pleuron. Posterior part of
metanotum, basal half of abdominal dorsum black,
caudal half largely yellow. Basal third of metapleuron
black, apical part largely yellow. Venter yellow with a
pair of very small black dots at anterior point of me-
soacetabular suture. Apex of rostrum, antennae ex-
cept for base of first segment, apex of fore femur and
rest of fore leg, middle and hind leg blackish.
Antennae when folded back over venter as long as
body in male, slightly shorter in female. Length of an-
tennal segments I : II : III : IV, male 1.22 : 0.57 : 0.55
: 0.56, female unknown. Eyes twice as long as head,
overlapping lateral margin of prothorax and anterola-
teral angles of mesothorax. Anterior and posterior
margins of pronotum medially concave. In lateral
view, mesonotum softly convex dorsally, metanotum
declivent. Intersegmental suture between meso- and
metanotum as well as anterior margin of first tergite
not visible. Lateral longitudinal suture of metanotum
not reaching the level of meso/metanotal interseg-
mental suture. Posterolateral angle of metaacetabula
bilobate. Ventral margin of fore femora slightly con-
cave, ventral margin of fore tibia nearly straight in fe-
male; fore femur with a low hump, fore tibia ventral-
ly irregular in male (fig. 24). Leg measurements, see
table 1.
Male. Parameres nearly symmetrical, fig. 28. In the
centre of mesosternum is a small patch of minute
black bristles which are not on a tubercle. Endosomal
sclerites fig. 31.
Comparative notes. — Although the parameres are
nearly symmetrical, this species fits best in subgenus
Venditioides. V. kurtakalami differs from similar spe-
cies by the form of fore tibia in male.
Etymology. — Kurtakalami is a Greek noun mea-
ning ‘curved shin’ and refers to the structure of male
fore tibia.
TREPOBATINAE Matsuda, 1960
Metrobatopsis Esaki, 1926
Representatives of this genus have so far only been
found in Papua New Guinea and Solomon Isles.
CHEN & NIESER: Gerridae from Sulawesi
Stenobates Esaki, 1927
Representatives of this genus have been reported fr-
om Singapore and Australia (an undescribed species
Andersen 1982 fig. 366). Stenobates is very similar to
Rheumatometroides, the main difference is in the loca-
tion of the metasternal scent gland opening which in
Stenobates lies near the anterior margin in the anteri-
or extension, whereas in Rheumatometroides the ope-
ning lies in the centre of the metasternum.
Rheumatometroides Hungerford & Matsuda, 1958
Two species are known from Papua New Guinea
and Solomon Isles (Lansbury 1992). Like Stenobates
they live on the sea and in estuaries in mangrove and si-
milar habitats. We add two new species from Sulawesi:
Rheumatometroides makraitos sp. n.
(figs. 32-35)
Type material. - Holotype d, allotype 2 (RMNH) and
5d 29 paratypes: Sulawesi Tenggara, Teluk Kendari, 13
Mar. 1989, N8947 (Nc, 1d 19 zMA 1d zmc), all apterous.
Description
Apterous specimens. — Dimensions. Length d 3.7-
4.3, 2 4.0-4.2; width of head d 0.90-0.98, 2 0.93-
0.98; width of thorax d 1.32-1.48, 2 1.66-1.70.
Colour, interoculus yellow with dark markings, eyes
dark, antennae dark, ventral side of first antennal seg-
ment, yellowish. Thorax and dorsum of abdomen
yellowish with dark markings (fig. 32). Venter yel-
low, legs light brown with dark stripes.
Eyes large, about two thirds as wide as interoculus.
Antennae shorter (0.6) than length of body, first seg-
ment with about 10 suff black setae. Length of anten-
nal segments I : II : HI: IV male 1.19 : 0.57 : 0.47:
0.43; female 0.95 : 0.55 : 0.49 : 0.47. Rostrum relati-
vely stout, reaching anterior quarter of mesosternum.
Pronotum distinctly (0.8) narrower than width of
head. Meso and metanotum in females with a very
broad mesal, weakly chitinized, sulcus (fig. 32), this is
not apparent in males. Fore femur of male only very
slightly thicker than in female, ventral margin straight
without modifications. Fore tibia flattened and so-
mewhat widened apically in both sexes.
Mesosternum 6-7 times as long as metasternum,
which is not produced anteriorly. Scent gland ope-
ning surrounded by a circular structure situated more
or less in the centre of metasternum.
Male. Pregenital sternite three quarters as long as
preceding abdominal sternites together. Sternite 8
slightly shorter than sternite 7, sternite 9 slightly
shorter than 8. Proctiger with a pair of long thin ro-
stroventrally directed horns (figs. 33, 34). Endosomal
sclerites fig. 35.
159
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Comparative notes. — There are two further species
in the genus: À browni Hungerford & Matsuda from
Solomon Isles and New Britain and A. serena
Lansbury from Papua New Guinea and New Britain
(Lansbury 1992). R. makraitos differs from both by
having the yellowish mark on pronotum larger, the
female in spite of a broader mesonotal sulcus being
relatively narrower and the horns of proctiger of male
being relatively shorter.
Etymology. — Makraitos, a Greek adjective, mea-
ning ‘extended’ and referring to the broad mesonotal
sulcus.
Remarks. — The species was found on very small
and shallow puddles between mangrove air roots. If
disturbed they jumped over the wet mud to the next
puddle. Despite their small size they are rather quick
and, as the net is hampered by the air roots, difficult
to catch.
Rheumatometroides drepanephoros sp. n.
(figs. 36-37)
Type material. — Holotype à, allotype 2 (RMNH), and
93 109 paratypes: Sulawesi Tenggara, Teluk Kendari,
Pulau Nambo, 13 Mar 1989, N8945 (Nc, 18 12 MBBj 19
BPUH 1d 19 yrec 1d 19 zmald 19 zmc), all apterous.
Description
Apterous specimens. — Length d 3.0-3.5, 2 3.2-
3.5; width of head d 0.87-0.90, 2 0.89-0.91; width
of thorax 6 1.15-1.20, 2 1.32-1.40. Colour, intero-
culus yellow with a broad median brown stripe, wide-
ned posteriorly with a small yellowish dot in the wide
part. A pair of brown lines at inner margins of eyes
also widened posteriorly and two sublateral brown li-
nes converging anteriorly and fused there with the
median stripe. Eyes dark, first antennal segment yel-
lowish with a dorsal brown line, segments 2-4 vent-
rally light, dorsally medium brown. Pronotum brown
with a pair of more or less circular yellow spots cover-
ed with pruinose silvery grey pubescence.
Mesonotum yellowish with a pair of submesal and a
pair of lateral longitudinal brown stripes, not fused
with the broadly brown hind margin, posterior yello-
wish part covered with silvery grey pubescence. In fe-
males this pattern is split by the broad yellowish cuta-
neous part. Abdominal terga largely brown, in
females alternating with yellow transverse stripes,con-
nexiva dark yellowish. First genital tergite yellowish, .
second basally yellowish, apically brownish. Venter
yellow, legs light brown with dark stripes.
Eyes large, about two thirds as wide as interoculus.
Antennae shorter (0.7) than length of body, first seg-
ment with 7- 12 stiff black setae. Length of antennal
segments I : II : III : IV male 0.71 : 0.51 : 0.48 : 0.49;
female 0.67 : 0.49 : 0.48 : 0.51. Rostrum relatively
160
stout, reaching anterior quarter of mesosternum.
Pronotum distinctly (0.7) narrower than width of
head. Meso and metanotum in females with a very
broad mesal, weakly chitinized, sulcus, this is not ap-
parent in males.
Fore femur of male not noticeably thicker than in
female, ventral margin straight without modifica-
tions. Fore tibia flattened and somewhat widened api-
cally in both sexes. Leg measurements, see tabel 1.
Mesosternum 6-7 times as long as metasternum,
which is not produced anteriorly. Scent gland ope-
ning surrounded by a circular structure which is mo-
re distinct in males than in females, situated near the
anterior margin of metasternum. Leg measurements,
see table 1.
Male. Pregenital sternite one third as long as prece-
ding abdominal sternites together. Sternites 8 and 9
of subequal length, about three quarters as long as
sternite 7. Proctiger with a pair of relatively short
crescent shaped rostroventrally directed horns (fig.
36). Endosomal sclerites fig. 37.
Comparative notes. — The appendages of the proc-
tiger of male are similar in form, though somewhat
shorter, as those in R. serena Lansbury and À. browni
Hungerford & Matsuda, which both have a different
pronotal pattern with a single median yellowish spot.
R. makraitos sp. n. is larger has also a different colour
pattern and the proctiger horns are more or less
straight.
Etymology. — Drepanephoros (Greek adjective mea-
ning bearing a sickle or scythe) refers to the form of
the appendages of the proctiger in the male.
Remarks. — This species was found on a small stre-
am with a sand/clay bottom flowing through a coco-
nut planation into the bay of Kendari. The plantation
is on a very small island, Pulau Nambo, in the bay.
This habitat is structurally quite different from the
one in which the preceding species was found.
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Gerridae) in Malaya, with a description of four new spe-
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Cheng, L., 1966. Studies on the biology of the Gerridae
(Hemiptera: Heteroptera) I: Observations on the feeding
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the Malayan Gerridae (Hemiptera: Heteroptera) with no-
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Distant, W. L., 1910. Some undescribed Gerrinae. — Annals
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Esaki, T., 1926. The water-striders of the subfamily
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Esaki, T., 1927. Revision of the Ptilomera-group of the
Gerridae, with descriptions of three new species
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Esaki, T., 1930. New or little known Gerridae from the
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162
KLAUS-GERHARD HELLER & KLAUS REINHOLD
Institut für Zoologie, Erlangen
A NEIWNBUSEI@GRICKEITOFESTETE GENUS
POECILIMON FROM THE GREEK ISLANDS
(ORTHOPTERA: PHANEROPTERINAE)
Heller, K.-G. & K. Reinhold, 1992. A new bushcricket of the genus Poecilimon from the Greek
Islands (Orthoptera: Phaneropterinae). — Tijdschrift voor Entomologie 135: 163-168, figs 1-
24. (1ssN 0040-7496). Published 1 December 1992.
Poecilimon paros sp. n. from the islands of Paros and Naxos, Greece, is described and illustra-
ted. It differs distinctly in calling song and structure of male cercus from its closest relative,
Poecilimon hamatus.
Dr. K.-G. Heller, K. Reinhold, Institut ftir Zoologie der Universitat Erlangen, Staudtstrasse 5,
D-8520 Erlangen, Germany.
Key words. — Orthoptera, Phaneropterinae, Poecilimon, taxonomy, bioacoustics, Greece.
The genus Poecilimon is the largest bushcricket ge-
nus in Europe, containing more than hundred species
(including species in the East Mediterranean and
around the Caspian Sea). Many species have restric-
ted ranges in the mountains of the Balkan peninsula
and Asia Minor, but only few are confined to islands
in the Mediterranean: P. cretensis Werner, 1903 to
Crete and Naxos, ikariensis Willemse, 1982 to Ikaria,
aegaeus Werner, 1932 to some Aegean islands, mytile-
nensis Werner, 1932 to Lesvos and Limnos and de-
planatus Brunner von Wattenwyl, 1891 to some is-
lands of the Southern Sporadhes (Willemse 1984).
Here we describe a new species, Poecilimon paros,
known only from the islands of Paros and Naxos
(Kikladhes, Greece) in the Aegean sea. Though close-
ly related to P. hamatus Brunner von Wattenwyl,
1878, known from Rhodes and Asia Minor, the new
species may be distinguished from the latter by slight
differences in morphology, but is readily recognizable
by its distinct song.
METHODS
Calling song and wing movement were registered
as described in Heller (1988). For song terminology
see Willemse & Heller (1992).
Poecilimon paros sp. n.
(figs. 1-5, 9, 11, 13, 15-19)
Type material. — Holotype d : Greece, Kikladhes, island
of Paros, Ag. Ilias-Pantes, 500-600 m, above the village of
Lefkes, 17.v.1991, collected as subadults, leg. K. Reinhold
(collection Heller, University of Erlangen). — Paratypes: sa-
me data, 84 79, including ® allotype (collection Heller,
except 1d 1% paratypes in collection Willemse,
Eygelshoven); Naxos: Sifones, 26.v.1976 (19)[ & 2d 39
subadults]; Moutsouna, 10.vi.1982, 2d 19), both leg. A.
Malicky (collection Willemse, Eygelshoven).
Description
Male (fig. 1) medium sized, integument shiny.
Head: fastigium of vertex produced anteriorly,
nearly twice as long as broad, narrow, as broad as
third antennal segment or a little bit narrower, lateral
margins parallel or slightly tapering anteriorly, from
above longitudinally impressed.
Thorax: pronotum with V-shaped transverse sulcus
cutting median line just behind middle of pronotal
length; metazona not widening and very slightly in-
flated posteriorly; hind margin straight and ventral
margin slightly convex in the prozona, sloping up-
wards in the metazona. Auditory spiracle large, more
than half pronotal height. Elytra with basal two third
covered by pronotum, apical margins reaching mid-
dle of first tergite; stridulatory file (fig. 11) of left elyt-
ron as seen from below distally not reaching hind
margin of elytron, in profile slightly bulging distally
of middle of length, shortest distance between proxi-
mal and distal most tooth 1.6 mm; in proximal two
thirds about 55 well sclerotised teeth and in distal
third about the same number, gradually decreasing in
size to the margin of the elytron; in the middle of the
file about 65 teeth/mm. Hind femur unarmed.
Abdomen: hind margins of tergites straight. Cercus
strongly bifurcate; inner branch terminating into
several small tubercles and distinctly longer than api-
cal one; apical branch slightly curved outwards and
upwards, terminating into a strong black tooth poin-
ting upwards in a 45° angle relative to the basal part
of the cercus (figs. 3-5, 9).
Subgenital plate reaching from mid to distal third
163
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-2. Poecilimon paros. - 1, 3, dorsal view (holotype); 2, 2, lateral view (allotype). - Figs. 3-10. Poecilimon, à , left cer-
cus (scale 1 mm). - 3-5, P. paros, dorsal view, three different specimens (paratypes); 6, P. hamatus, dorsal view (Dimilia,
Rhodes); 7-8, P. hamatus, dorsal view, two different specimens (Lindos, Rhodes); 9, P. paros, lateral view (paratype); 10, P.
hamatus, lateral view (Lindos, Rhodes).
of cercus, hind margin weakly incised, postero-lateral
edges slightly protruding, apical part of ventral side
with an obtuse median and on either side a more dis-
tinct lateral keel, extending into postero-lateral edges.
Coloration: in general appearance very similar to P.
hamatus (see Heller 1988, jacket photograph); varico-
164
loured, general colour yellowish red. Vertex, lateral
and dorsal sides of prozona and pronotum, tergites
and legs finely speckled rusty brown or blackish.
Antennae black, finely annulated yellow-white.
Dorsum of pronotum with short yellowish median
line frontally and black markings laterally and in the
HELLER & rEINOLD: New Poecilimon
Figs. 11-12. Poecilimon, ventral view of stridulatory file of left d elytron, distal end to the left (scale 300 um). — 11, P. paros
(paratype); 12, P. hamatus, (Lindos, Rhodes). - Figs. 13-14. Poecilimon, base of ovipositor, lateral view (scale 1 mm). - 13, P.
paros (allotype); 14, P. hamatus (Lindos, Rhodes).
centre, metazona reddish brown. Elytra white lateral-
ly up to the apical parts contrasting with the black
central areas. Abdomen from above yellow with bro-
ad, black midline and broad, black lateral lines; first
and last two tergites nearly completely black; sternites
yellowish, each with a reddish brown spot of various
intensity. Cerci reddish brown, tips black. Subgenital
plate yellowish to reddish brown. All femora with
black stripes and between the yellow lower keels dark
brown to black; auditory tympana of fore tibia each
with a large black spot; hind tibiae dorsal with black
spines; all tarsi dark brown to black.
Female (fig. 2). — Auditory spiracle large, but less
than half pronotal height. Elytra: left overlapping
right, clearly visible, strikingly coloured black and
white as in male; stridulatory apparatus present, simi-
lar to that in females of P. hamatus (see fig. 1 in Heller
& von Helversen 1986). Basal fold of ovipositor (fig.
13) with two pits and two lobes forming an oblique
T-shaped structure as seen from above: one pit situ-
ated above the horizontal line of the T, the other in
front of the vertical line of the T and opening ventro-
anteriorly. Coloration as male.
Measurements (length in mm). — Body ¢ 20-22,
165
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
19 24
Hd
Figs. 15-24. Poecilimon, 3, synchronous registration of left elytron movement (above, upward = opening, downward = clo-
sing) and calling song (below) (scale 0.5 sec.). — 15-19, P. paros (body temperature ca. 32°C.). - 15, holotype; 16-18, three
different paratypes; 19, specimen from fig. 18, variation. - 20-24, P. hamatus (body temperature ca. 26°C.). - 20-22, three dif-
ferent specimens (Lindos, Rhodes); 23, specimen from Dimilia, Rhodes, normal pattern; 24, specimen from fig. 23, rarely
observed variation.
166
® no data (all females preserved in absolute ethanol);
pronotum d 4.7-5.5, ® 5.7-6.2; elytron (visible
part) d 1.0-2.2, 2 1.0-1.2; hind femur d 14.0-14.7,
3 14.7-15.6; ovipositor 7.0-7.5.
Bioacoustics. — Male calling song consists of short
echemes repeated at intervals of several seconds
(mostly 3-5 s). Each echeme (figs. 15-19) consists of a
series of wing movements (opening and closing) with
increasing amplitude producing a series of syllables
with increasing intensity; wing movements repeated
regularly except the last movement cycle (and sylla-
ble) which is produced after a longer interval than be-
tween the previous cycles. Frequency of stridulatory
movements within the central part of each echeme
between 23 and 28 Hz (body temperature of the ani-
mals about 32°C). Number of movement cycles dif-
ferent between both individual males (most frequent
numbers in different males: 9-10, 11, 11-13, 14; ran-
ge 6-15) and circumstances (short echemes during
acoustic interactions). The power spectrum of the
song showed its peak at about 28 kHz (20 dB below
peak 21-42 kHz).
Distribution. — Only known from the islands of
Paros (500-600 m altitude) and Naxos.
Differential diagnosis
The new species can easily be distinguished from
all other species of the genus except for Poecilimon
cervus Karabag, 1950 and P. hamatus, which show
also a strongly bifurcate shaped male cercus. In P. cer-
vus, known from Central and North Anatolia, the tip
of the apical branch of the male cercus has a series of
strong spines instead of a single and smaller one in the
new species and the inner branch is wider and apical-
ly widened instead gradually tapering as in paros
(compare figs. 3-5 with fig. 25 in Karabag 1950).
However, morphological distinction between the
new species and P. hamatus, known from West
Anatolia and some adjacent Aegean islands (Willemse
1984), is less obvious. Apparently they represent sis-
ter species. Their distinction is mainly restricted to
the male cercus: the inner branch in P. hamatus being
longer and somewhat more slender (figs. 6-8;
Karabag 1950: fig. 26, Willemse 1988: fig. 127,
Heller 1984: fig. 2) and the apical branch pointing
more outwards with the terminal tooth being smaller
and pointing straightly upwards (fig. 10) instead of
upwards and at the same time obliquely inwards in
the new species (fig. 9). In the female of P.paros the
pit of the basal fold of the ovipositor that opens vent-
ro-anteriorly is broader and the bordering lobe more
prominent than in hamatus. Another difference be-
tween both species, present in both sexes, refers to the
elytra which in the new species are less covered by the
HELLER & rEINOLD: New Poecilimon
pronotum and to a greater extent visible than in P.
hamatus. However, most obvious and reliable diffe-
rences between the two species are found in the struc-
ture and duration of the calling songs, defined by the
number of movement cycles (opening and closing).
In P. hamatus two cycles at most have been registra-
ted, the first one often producing only a very faint
sound if detectable at all (figs. 20-24; Heller & von
Helversen 1986: fig. 3, Heller 1988: fig. 36K). Rarely
even a single opening and closing movement was re-
corded (fig. 24). These short echemes of P. hamatus
differ strongly from those of paros in which at least 6
and commonly 9 to 14 successive opening and clo-
sing movements per echeme are produced (figs. 15-
19).
Remarks
The type locality is the slope of the mountain Agios
Ilias-Pantes and can be reached by car using the road
from the village of Lefkes to the O. T. E. station
(Greek Telegraph Station) on one of the summits.
The subadult animals of P. paros were found mainly
on Cistus. Their distribution was clustered with so-
metimes high population densities of up to 25 ani-
mals per m’. Up to now P. paros is the only member
of the genus known from Paros.
Etymology
A noun in apposition, named after the island of
Paros.
ACKNOWLEDGEMENT
We thank Dr. Fer Willemse for his valuable com-
ments.
REFERENCES
Heller, K.-G., 1984. Die ‘Zsophya-Arten von Rhodos. —
Articulata 2: 74-77.
Heller, K.-G., 1988. Bioakustik der europäischen
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sensory time window. — Behavioural Ecology and
Sociobiology 18: 189-198.
Karabag, T., 1950. Five new species of Poecilimon Fischer
(Orthoptera: Tettigoniidae) from Turkey. — Proceedings
of the Royal Entomological Society of London (B) 9-10:
150-155.
Willemse, F., 1982. A survey of the Greek species of
Poecilimon Fischer (Orthoptera, Ensifera, Phanerop-
terinae). — Tijdschrift voor Entomologie 125: 155-203.
Willemse, F. , 1984. Catalogue of the Orthoptera of Greece.
— Fauna Graeciae 1: i-xii, 1-275.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Willemse, F. & K.-G. Heller, 1992. Notes on systematics of N
Greek species of Poecilimon Fischer, 1853 (Orthoptera: 1 ne 1991
Phaneropterinae). — Tijdschrift voor Entomologie 135: Accepted: February 1992
DD):
168
P. J. van HELSDINGEN & A. E. DECAE
National Museum of Natural History, Leiden, The Netherlands
ECOLOGY, DISTRIBUTION AND VULNERABILITY
OF MACROTHELE CALPEIANA (WALCKENAER)
(ARANEAE, HEXATHELIDAE)
Helsdingen, P. J. van & A. E. Decae, 1992. Ecology, distribution and vulnerability of
Macrothele calpeiana (Walckenaer) (Araneae, Hexathelidae). — Tijdschrift voor Entomologie
135: 169-178, figs. 1-2, tabs. 1-4. [Issn 0040-7496]. Published 1 December 1992.
Data on distribution, habitats, local densities, and observations on the biology of Macrothele
calpeiana (Walckenaer) are presented. A distribution-map (5 X 5 km UTM-grid) is provided.
Our results confirm the earlier published distribution in Spain. Sites where the species was not
found are also indicated. The occurrence in North Africa is discussed. The inclusion of this spe-
cies in Appendix II of the Convention on the Conservation of European Wildlife and Natural
Habitats (Bern Convention) is criticized.
Correspondence: P. J. van Helsdingen, National Museum of Natural History, Raamsteeg 2,
Leiden, NL-2311 PL Netherlands.
Keywords. — Araneae, Hexathelidae, Macrothele calpeiana, ecology, distribution, Spain, Bern
Convention.
Macrothele calpeiana (Walckenaer, 1805) was de-
scribed early in the 19th century as Mygale calpeiana
after a male specimen from the Gibraltar region.
Hardly any new records have been published since
and Gibraltar (type locality of M. calpeiana),
Algeciras (type locality of the junior synonym Mygale
luctuosa Lucas, 1855), Malaga and Ronda in Spain,
and El-Arouch in Algeria remained the recorded lo-
calities of this species until very recently. Then sud-
denly and for reasons unknown the species received
new attention from two sides independently.
Blasco & Ferrandez (1986) produced a substantial
morphological and taxonomical account on M. cal-
peiana, and settled the discussion on the synonymy of
M. luctuosa with M. calpeiana. Blasco & Ferrandez
(1986) also presented an updated distribution based
largely on recently collected material.
Snazell & Allison (1989) published an important
paper on the species in which they repeated the dis-
tributional and ecological data published earlier by
Snazell (1986). They also included and discussed the
distribution data of Blasco & Ferrandez (1986) and
of others which were not available in 1986.
Furthermore they, elegantly, corroborated the synon-
ymy of M. calpeiana with M. luctuosa. It appeared
that a subadult male of M. calpeiana had not been
recognized as such and described as the female of a
new species (M. luctuosa), an understandable mistake
for any one who knows these animals. Snazell &
Allison (1989) presented additional morphological
details and information on the prey, a description of
the nest and web, observations on courtship and mat-
ing in captivity, and finally a description of the sec-
ond European representative of the genus, M. cretica
Kulczynski 1903.
As to its ecology, Snazell (1986) emphasized the re-
lationship of M. calpeiana with undisturbed Cork
Oak (Quercus suber L.) woodland in the southern-
most part of Spain, which he refers to as the optimum
habitat for this species. He suggested an apparent in-
ability of the spiders to survive in situations more
than a few metres away from the woodland, although
populations are said to survive on bare banks of roads
and streams for some time even though the surround-
ing woodland has almost entirely degenerated.
Extensive areas of Cork Oak woodland are still
present in the region and, he says, there seems to be
no immediate problem for M. calpeiana in Spain.
However, according to him, intensive cultivation of
the woodland and the development of housing pro-
jects currently undertaken in the area might become a
serious threat. Collins & Wells (1987) picked up this
message and, treating M. calpeiana as a bio-indicator
for Cork Oak woodland, included the species in their
selection of endangered invertebrate species to be list-
ed in the Bern Convention. In the same year the spe-
cies was placed on Appendix II of the Convention.
Not being convinced by Snazell’s (1986) argu-
ments for the strict habitat requirements of M. cal-
peiana, we decided to carry out an independent sur-
vey. The fieldwork was carried out in 1989 by A. E.
Decae and A. Hallensleben during two trips to
169
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 1. Synopsis of localities in Spain and their main characteristics where M. calpeiana was found. Density refers to the
counted number of nests of adult spiders within an area of approximately 20 m’ around the first nest observed; (number of
collected specimens in brackets); no number means webs and/or spiders observed but not counted.
Locality UTM grid- date altitude site vegetation type density
reference inm (number collected)
El Captan UF 1846 30.1 200 hillside cork oak cult. 1 (1)
El Captan UF 1847 30.ili 250 hillside cork oak cult. 1 (0)
Guadiario TE 9419 05.iv 20 coastal zone olive cult. 10 (4)
Gaucin TF 9445 17.ix 600 highlands cork oak 6-7 (1)
Crestellina TF 9242 17.ix 500 hilltop oak/olive 3 (0)
Nuevo Castellar TF 8220 18.ix 70 lowlands dense oak 1 (1)
Almoriama TF 8020 18.ix 80 lowlands oak/eucal./pine 13 (1)
Castellar de la Frontera 57.923 19.ix 260 hilltop castle wall 10-15 (2)
Sotogrande TF 9218 20.ix 20 lowlands cork oak cult. 0-5 (0)
Jimena de la Frontera TF 8036 20.ix 200 hilltop castle wall
La Tienera Vieja TE 7839 20.ix 200 hillside shrubs/oak/pine 1-2 (0)
La Tienera Vieja TE 7740 20.ix 300 highlands oak/shrubs 6-8 (0)
Sambana TF 8832 20.ix 100 hillside shrubs 7 (1)
Ronda-Grazalema TF 9773 23.1x 720 highlands oak cult. 1-0 (1)
Grazalema TF 8770 23.ix 920 mountainside grass/herbs 1 (0)
Puerto de la Boya TF 8570 23.ix 1100 mountain pass pine cult. 4-6 (1)
Sierra Grazalema TF 8371 23.ix 960 highlands oak/shrubs
Benamahona DEVIS IX 900 highlands pine
El Bosque TF 6972 23.ix 200 highlands grass/olive 5 (1)
Arcos de la Frontera TF 5470 23.ix 200 highlands oak/pine 8 (1)
El Santiscal TF 5172 23.ix 100 highlands acacia cult. 1 (0)
San José del Valle TF 5556 23.ix 120 hillside olive cult.
Tempul TF 6058 23.ix 100 river valley shrubs/gallery 1-2 (0)
Cortes de la Frontera TF 7346 23.ix 500 highlands oak forest 2-3 (0)
Puerto del Bujeo MES ICH 26:x 150 roadside cork oak
Jubrique MISI 2 300 hilltop oak/pine/olive 10-14 (0)
Santuaria de la Luz TF 6301 27 ix 30 lowlands oak forest 8 (2)
Almodovar/Sierra del Niño TF 6106 27.1x 50 roadside oak/grass 1-3 (1)
Sierra del Nino TF 6408 27.ix 150 hillside oak forest 1 (1)
Sierra del Niño TE 6504 27.ix 250 hillside oak forest 1 (0)
Sierra del Niño TF 6503 27.1x 250 hillside oak forest 1-2 (0)
Puerto de Ojen TF 6707 27.ix 350 hillside oakforest 4-5 (1)
Los Barros TF 7008 27 ix 200 hilltop shrubs 4 (0)
Los Barros TE 7109 27.ix 200 hillside oak forest 7 (1)
Casares TE 9935 28.ix 180 hillside oak forest 7-9 (5)
Casares TF 9636 29.ix 500 hilltop shrubs 1-2 (1)
Casares TE 9737 29.ix 400 hillside olive cult. 11 (6)
Estepona UF 0836 18.ix 80 hillside eucalyptus cult. 1 (1)
Sierra Bermeja UF 0738 18.ix 450 hilltop oak/pine 1-3 (3)
Sierra Bermeja UF 0240 18.ix 500 mountainside pine/oak 1-2 (0)
Sierra Bermeja UF 0642 18.ix 600 mountainside chestnut 4 (0)
Sierra de Palmetera UF 1749 25.ix 450 mountainside shrubs 1 (1)
Sierra de Palmetera UE 1755 25.ix 900 mountainside roadside/pine 1-3 (0)
Acueducto UF 1170 25.ix 700 highlands grass/roadside 1-2 (0)
Puerto del Viento UF 2073 25.ix 1180 mountain pass shrubs/roadside 3-4 (2)
Genalguacin UF 0048 26.ix 400 hillside olive cult. 3-5 (0)
Buenas Noches WHOS ie 23: 20 lowlands eucalyptus 15 (3)
Casares UF 0033 28.ix 100 hillside oak/olive 1-2 (0)
Puerto de Piñas Blancas UE 0643 26.ix 700 hillside oak/pine 6 (1)
Sierra Bermeja UF 0642 21.ix 450 hilltop oak/pine 4 (0)
Puerto de Piñas Blancas UF 0643 21.ix 700 hillside pine cult. 5 (2)
Jubrique UF 0848 21.ix 500 hilltop pine cult.
Jubrique UF 0448 2l.ix 350 hilltop oak/pine
Jubrique UF 0150 21.ix 300 highlands cork oak cult. 6-8 (0)
Ronda UF 0769 21.ix 750 canyon/ruins almond cult. 12 (4)
Ronda UF 0769 22.ix 750 city park palmtrees 1 (0)
Estación de la Indiana UF 0372 23.ix 750 roadside cork oak
170
Andalusia. They collected both ecological and distrib-
utional data.
METHODS
A first trip was made in the period between 25
March and 8 April 1989. The coastal region, roughly
bordered by the lines Gibraltar-Ronda in the West,
Ronda-Lanjarön in the North and Lanjarön-Albunol
in the East, was surveyed for the presence of M. cal-
peiana. Twenty localities were investigated. In only
three localities, all of them West of the city of Mälaga,
M. calpeiana was found (fig. 1). Approximately ten
specimens were collected (females and juveniles) and
taken home for further study.
A second trip was made in the period between 17
and 29 September 1989. During this trip attention
was focussed on the ecology and natural history of M.
calpeiana rather than on the geographical distribu-
tion. The region surveyed was largely the same as
Snazell’s (1986) study area. Eighty localities were sur-
veyed, in 58 of which M. calpeiana was found.
In table 1 we have listed all localities where
Macrothele calpeina has been found. We have includ-
ed information on the relative population densities
(number of nests within approximately 20 m’). Fifty
adult specimens were collected, among which three
adult males. All specimens are deposited in the collec-
tion of the Rijksmuseum van Natuurlijke Historie
(National Museum of Natural History) at Leiden,
The Netherlands. Table 2 lists all the localities where
the species could not be found, an equally important
observation. In the tables we have listed our localities
with a 1X1 km UTM-grid, but visualized the distri-
bution on the map with a 5x5 km accuracy.
RESULTS
Habitat
The most western localities visited during the first
trip lie within the region where Snazell (1986) found
M. calpeiana most commonly and predominantly in
association with Cork Oak. The reported frequent
occurrence of M. calpeiana in Cork Oak vegetations
in that region (mainly cultivated and often fenced in
forestland) was confirmed, although dense popula-
tions of M. calpeiana were also found in very different
situations, such as along roadsides, in picnic areas, in
fruit orchards, in pine- and Eucalyptus plantations, on
recently burnt grassy slopes, on rubbish dumps, in
old stone walls, on cliff faces and even in the Palm-
trees of the city parks in Ronda. Further to the East,
in the coastal region east of the city of Malaga, an area
not surveyed by Snazell, M. calpeiana was not found.
This seems to support Snazell’s (1986) remark on the
VAN HELSDINGEN & DECAE: Macrothele calpeiana
very restricted distribution of M. calpeiana in Spain.
During the second trip large and dense populations
of M. calpeiana were found, in association with Cork
Oak as well as in very different situations. It appeared
that M. calpeiana, rather than being dependent on
undisturbed Oak-wood, is a culture following species
that profits from human modifications of the land-
scape.
Counts of nests showed that in some of these local-
ities population densities reach higher levels than in
Cork Oak forests (see tables 1 and 3). Population
densities in the relatively undisturbed forests of the
‘Parque Natural Sierra de Grazalema’, the ‘Reserva
Nacional de Cortez de la Frontera’ and the ‘Reserva
Torrecilla de Serrania’ were found to be lower than in
typically man-made habitats such as stone walls and
rubbish dumps (see table 3).
Table 4 illustrates the relationship of the occur-
rence of M. calpeianawith more or less disturbed hab-
itats. The following classification was designed for the
purpose:
Category 1. — Non-cultivated/non-forested: in-
cludes the typical mediteranean Garigue with Cistus
sp., Thymus sp., Lavandula sp., Trifolium stellatum L.,
Quercus coccifera L., Chamaerops humilis L., etc., bar-
ren mountain passes and exposed river beds;
Category 2. — Relatively undisturbed mixed forest:
Maquis with Pinus pinea L., Quercus ilex L., Q. suber
L., Laurus nobilis L., Pistacia lentiscus L., Ruscus acu-
leatus L., Myrtus communis L., etc., with a dense
understorey;
Category 3. — Cultivated Cork Oak stands: a forest
type of almost entirely Cork Oak (Q. suber) from
which the understorey is cleared; most of these woods
are apparently private property and all are in use for
the production of cork;
Category 4. — Unmanaged culture land: includes
road sides, old walls, rubbish dumps and generally
polluted picnic grounds;
Category 5. — Managed culture land: includes city
parks, gardens, parking lots and development areas.
The first two categories contain habitat types
which have developed as a result of historical human
activities and possible changes in climate but which
are currently relatively free of human influences.
Reproducing females of M. calpeiana were found in
the typical ‘garigue’ (cat. 1) as well as in the dense, al-
most inpenetrable ‘maquis’ (cat. 2) type of habitat,
but population densities in both these ‘undisturbed’
habitat types were relatively low. Population densities
were even lower in the fifth recognized category
(managed culture land), although nests of adult spi-
ders were found even in such thoroughly maintained
places as private gardens, city parks and parking lots.
The optimum habitat for M. calpeiana in Southern
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Spain obviously lies in terrains that are easily access-
ible to man. These habitats, here placed in categories
3 and 4, are either man-made origin or to some extent
exploited by man. As far as road sides, old walls and
rubbish dumps are concerned this needs no further
explanation. Cork Oak woodland is very common
and regarded as a natural resource in the region. This
leads to a careful management that, as far as our ob-
servations go, rarely has adverse effects on the M. cal-
peiana populations. In 20 out of the 22 Cork Oak fo-
rests thus managed M. calpetana populations were
found. In one case, where clearing went as far as bull-
dozing all the understorey, M. calpeiana apparently
still managed to retain a foodhold by becoming arbo-
real. Here numerous webs of adult spiders were found
more than 2.5 meters above the ground on the trunks
of the Oak trees.
Summarizing we may conclude that M. calpeiana
thrives on sites with open and accessible soil with
some form of shade and cover.
Biology
Males
Adult males have a body length (excluding the
spinnerets) of approximately 25 mm. In contrast to
most mediterranean Mygalomorphae, males of M.
calpeiana are present virtually all year round. Males
were collected in the period February - June (Blasco
& Ferrandez 1986 and Snazell 1986) and in
September (this study).
It is not clear if there is a particular mating season.
During September fully adult males were found to
hunt actively from their own nests. Males of mediter-
ranean mygalomorphs more usually abolish feeding
to become reproductively active.
Females
Adult females of M. calpeiana range in body length
(excluding the length of the spinnerets) from approxi-
mately 25 mm to 37 mm. The relatively large varia-
tion in body size of adult females indicates the exis-
tence of several year-classes in this group. This in turn
would mean that, as is usual in Mygalomorphae, M.
calpeiana females are reproductively active for several
years after reaching adulthood.
Juveniles
In September several females were found to have
young with a body length of approximately 3 mm in
their nests. These juveniles were moving freely in
their mothers’ nests and presumably represented the
generation which would disperse. Juvenile spiders of
intermediate sizes were observed to inhabit individu-
al nests.
172
Activity rythms
M. calpeiana, unlike most other mediterranean
mygalomorphs, is not strictly nocturnal in its activi-
ties. Prey-capture was observed both at night and du-
ring daylight hours. At night some, but not all spiders
in a colony took up positions in the entrance of their
nests near the web. Occasionally spiders were obser-
ved in this position during daylight. Other activities
such as dispersal, nestbuilding, or mating were not
observed.
The nest
The nest of M. calpeiana is composed of an expo-
sed ‘sheetweb’ and a set of underground silk tubes.
Sometimes the sheetweb is absent and the exposed
part of the nest is reduced to a densely woven closed
silk cell. The web extends from the nest entrance and
may cover an irregularly shaped area up to 1600 cm’.
It may be directly attached to the substrate or be sus-
pended between low vegetation around the nest en-
trance. The web consists of a more or less dense mesh
of very fine silk strands spun in an irregular criss-cross
pattern. Some aerial strands may extend to some 20
cm above the nest entrance. In general smaller spiders
spin smaller webs, although the size and density of the
web may vary considerably within all spider size clas-
ses.
M. calpeiana spins silk continually as it moves.
When the spider re-enters the nest after a short sally
on the web, it covers the entrance behind itself with a
few flimsy strands of silk.
The nest entrance may consist of one, two or more
silk funnels converging towards the inside of the nest
in one main nest tube. The main tube may have one
or more flimsy side tubes and is suspended in a pre-
existing cavity. This cavity may be a deserted rodent
burrow, a rock crevice or washout, a hollow tree etc.
Depending on the conditions the main nest tube may
extend more than fifty centimetres underground.
Indigestible remains of prey are attached to the side
tubes or to the outside of the main tube.
Prey-capture
Natural prey-capture behaviour was not observed.
Prey-capture could only be induced in spiders having
a sheetweb extending from the entrance of their nests.
Those having a closed silkcell at the entrance were not
actively hunting and were probably moulting.
Skincasts were found both in sheetweb nests and in
‘cellnests’.
A whole range of arthropods placed in the web of
M. calpeiana induced prey-capture behaviour. The
capture behaviour is rather straightforward and ster-
eotyped, regardless of the type of prey. Movements of
the prey in the web cause the spider to come to the
entrance of the nest where it pauses for some time,
VAN HELSDINGEN & DECAE: Macrothele calpeiana
Table 2. Synopsis of localities and their characteristics where M. calpeiana could not be found in 1989.
Locality UTM grid-reference date
Santuaria de la Luz TE 6395 27.ix
Manilva TF 9928 17.ix
Manilva TF 9632 72x
Casares TF 9536 17.ix
Crestellina TF 9239 17.ix
Crestellina TF 9341 17.ix
San Martin de Tesserillo TE 9224 19.ix
San Martin de Tesserillo TF 8624 19.ix
Castelar/Ferro Caril TF 8324 19.ix
Sotogrande TF 8619 19.ix
Charco les Hurrones TF 6757 DIX
Puerto de Galiz TF 6750 23.1x
Gibraltar TF 8802 05.iv
Sotogrande TF 9218 05.iv
Sierra Bermeja UF 0738 21.ix
Puerto del Viento UF 1773 25.ix
Sierra de Palmetera UF 1355 25.ix
Paruta UF 1260 NS
Sierra de Palmetera UF 1655 25.ix
Estepona UF 0534 24.ix
Puerto de Alija UF 1647 25.ix
Yunquea UF 3169 30.ii
Tolox UF 3161 30.11
Rio Grande UF 3661 30.11
Puerto de Abejas UF 2969 30.1
Mirador del Guarda Forestal UF 2471 30.iii
El Burgo UF 2872 30.11
Moclinejo UF 8869 03.1v
Benamargosa UF 9377 03.iv
Puente Tablata UF 9876 02.iv
Estepona UF 0837 18.ix
Frigliana VE 1772 27.
Cueva de Nerja VF 2369 29.1
Motril VE 5570 28. iii
Pénon del Fraile VF 6181 28.111
Haza de Lino VE 7275 28.111
Popolos VF 7674 28.iii
Puerto Camacho VF 6878 18.11
Frigliana VF 1870 27.111
Cueva de Nerja VF 2471 7.iv
apparently to locate the source of disturbance. After a
short interval it dashes straight at the prey, grabs it
with the fangs and drags it in reverse back to the nest
entrance.
Prey
Captures of the following arthropods were obser-
ved: Oniscoidea (two species), Orthoptera, Ceram-
bycidae, Staphylinidae, Carabidae, Diplopoda, Scor-
pionida. Escaping from the web very quickly was a
Scolopendra spec.
Snazell & Allison (1989) reported a very broad
range of different arthropod prey being captured and
consumed by M. calpetana. This strongly indicates
general or opportunistic feeding habits of this species
and suggests that the availability of food is unlikely to
limit the natural distribution.
altitude in m. site vegetation type
30 lowlands grass
50 lowlands vine cult.
200 hilltop shrubs
400 highlands shrubs
400 highlands olive/shrubs
450 hilltop olivebush
20 river bed gallery forest
30 creek bed shrub/gallery
40 river bed gallery forest
50 undulating cork oak cult.
160 highlands grass
540 highlands oak forest dense
40 hillside maquis
10 coastal zone cork oak cult.
200 hillside pine
1190 mountain pass shrubs
1050 mountainside barren
1000 highlands pine forest
800 mountainside pine forest
10 lowlands shrubs
400 mountainside barren
700 highlands maquis
300 highlands olive cult.
250 highlands garigue
820 highlands olive cult.
1000 highlands pine forest
800 highlands garigue
100 hillside maquis
200 hillside maquis
100 highlands olive cult.
150 hilltop shrubs/pine
300 highlands olive cult.
20 coastal zone garigue
20 coastal zon road side
900 highlands garigue
1250 highlands cork oak cult.
600 hillside olive cult.
1220 highlands olive cult.
250 highlands olive cult.
30 coastal zone garigue
Our investigations of prey remains found in ten
webs support this idea. Oniscoidea, Coleoptera and
Formicidae constitute the bulk of the prey both in bi-
omass and number of individual prey. Interesting is
the large discrepancy in the sizes of individual prey
animals found in single webs. Remains of beetles with
body lengths of approximately 20 mm were found to-
gether with the remains of ants of hardly 2 mm. It has
still to be investigated if adult spiders catch this ex-
traordinary range of prey sizes or that maybe the small
prey is captured by juveniles actively hunting in their
mother’s web.
Very few mygalomorphs are known to capture fly-
ing prey. The presence of a bee (family Apidae) re-
ported by Snazell & Allison (1989) and a
Hymenopteron in one of our samples indicates that
M. calpeiana is an exception.
173
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Ectoparasites
Virtually all specimens collected were heavily infes-
ted with Acari which clung in large numbers to the
carapace, usually concentrating in the fovea, and
sometimes to the abdomen of the spiders. This is in
agreement with what Snazell & Allison (1989) found.
Numerous mites may be present on one individual
without the spider seeming to suffer very much.
Spiders kept in captivity were all very quickly free of
mites. It is not known what caused the disappearance
of the mites. The mite was described by Baker (1991)
as a new species of parasitic mite (Androlaelaps pilosus)
after specimens collected by Snazell.
Distribution in Spain
The recent work by others (Snazell 1986, Blasco &
Ferrandez 1986, Snazell & Allison 1989) and oursel-
=
|
f
=
D
(
SS
O Literature records
+ Own records
A Negative results
Fig. 1. Distribution map of Macrothele calpeiana (Walckenaer) with 5x5 km accuracy on UTM-grid; early record from Al-
geria indicated.
174
ves certainly has broadened our knowledge of
Macrothele calpeiana. Our own investigations affirm
and reinforce the distribution pattern which emerges
from the literature. But science is exigent and never
satisfied. Many questions remain yet unanswered.
In Europe M. calpeiana remains restricted to the
South of Spain, the region of Andalusia, mainly in the
provinces of Cadiz and Malaga. The occurrence in the
province of Huelva, based on old and recent records
(Blasco & Ferrandez 1986) demonstrates a disjunct
distribution with the valley of the Guadalquivir as a
possible lowland ecological barrier (open, unshaded,
lowland area). The three Huelva records represents the
northernmost as well as the westernmost finds so far.
The population to the southeast of the
Guadalquivir reaches from Malaga in the East to the
City of Cadiz in the West and from 37° North lati-
tude down to Tarifa, the very southern tip of the pe-
ninsula. In this region we may call M. calpeiana a
common species. Our attempts to find the species
East of Malaga remained without success. However,
Barbara York Main from Nedlands, Western
Australia, has found a small population (a juvenile
and some cast skins were collected) at Las Nimbres,
1.5 km E of Puento de la Mora, at an elevation of
1390 m, in the Sierra Harana north of Granada
(Snazell & Allison 1989: 69; also pers. comm.),
which means a considerable eastward extension of the
known range. The presently known northern limit of
the distribution in the province of Malaga at 37° N
does not seem a very natural one at first sight and we
may expect new records more to the north.
The three Huelva records indicate the occurrence
on at least the southern slopes of the Sierra Morena.
This region, the northern halves of the provinces of
Huelva, Sevilla and Cordoba probably is undercol-
lected.
Which factors limit the distribution of M. calpeia-
na is not known. The species seems to prefer cover
and shade over exposed open soil. From this we may
infer that the animal is sensitive to low and high tem-
perature extremes. Humidity may also be a regulating
factor. The discovery of populations at higher alti-
tudes, where the temperature will be lower on the av-
erage, indicates a fair rate of tolerance. We have found
a population at 1100 m in a pine forest, on a south-
facing slope on a mountain pass at Puerto de la Boya
(36°46’N, 5°24’W; province of Cadiz), and another
one at 1180 m on a roadside bank on a north-facing
slope on a mountain pass near Puerto del Viento
(36°48’N, 5°01’W; province of Malaga). The popu-
lation established by B. Y. Main at the Sierra Harana
at ca 1400 m also proves the ability of M. calpeiana to
survive under the climatic conditions at high alti-
tudes.
Snazell & Allison (1989) stress the correlation of
VAN HELSDINGEN & DECAE: Macrothele calpeiana
the distribution of M. calpeiana with the area of rela-
tively high rainfall (80-200 cm), warm winters and
high summer temperatures. They also link the species
firmly to oakwoods with the Cork Oak (Quercus
suber L.) as dominant species. The Cork Oak has a
much wider distribution in Spain and certainly is not
restricted to Andalusia. Other parts of Spain than the
presently known distribution area of M. calpeiana, in
particular the southern slopes of the Sierra Morena,
have precipitations about as high as the Cadiz-Malaga
mountainous region. This supports the hypothesis of
a more extensive distribution along these slopes.
Distribution outside Spain
Two records from North Africa indicate a wider
distribution South of the Mediterranean Sea. The
first is an old record from Algeria, where it was found
somewhere in the surroundings of El-Arouch in the
province of Constantine (Lucas 1846). Our attempts
to locate the original material have failed. According
to Lucas the species was not common in that locality,
where it was found under stones. The second record
is of more recent date. A. Blasco collected a single
male in 1981 at Ceuta in Spanish Morocco (Blasco &
Ferrandez 1986). Even though these two records are
spaced out geographically and in time they are sugge-
stive of a much wider distribution in the West
Mediterranean. The mediterranean zone of North
Africa is grossly undercollected and hardly anything is
known of the Mygalomorphae of that region.
If M. calpeiana would be more wide-spread in
North Africa, the geographically restricted occurrence
in southern Spain would be easier to understand. The
restricted presence in Spain would only be a bridge-
head on the European continent from a much larger
population in North Africa. The Atlas mountains
might offer suitable habitats. Further investigation of
this region will throw more light on the distribution.
It would not at all be surprising if North Africa
harboured a number of Macrothele species. The genus
now comprises five species in West Africa, calpeiana
and cretica in Europe, and seven more species in
India, South-east Asia, China and Taiwan. Most spe-
cies have been described after one sex only, usually
the female, which shows the paucity of available ma-
terial.
DISCUSSION
M. calpeiana, according to our observations, is not
restricted to oak woodland. No undisturbed oak
woodlands persist in southern Spain today. All oak-
woods, with the Cork Oak (Quercus suber L.) as do-
minant tree species, or mixed woods are the result of
human interference through mild exploitation during
175
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Table 3. The ten highest densities of Macrothele calpeiana
nests counted in plots of approximately 20 m’ within diffe-
rent types of habitat in Southern Spain. Note the absence of
relatively undisturbed Oak forest (maquis) in this ranking.
type of habitat location number of
nests per 20 m’
1 castle wall Castelar de la Frontera 15
2 rubbish dump Buenas Noches 15
3 mixed forest Jubrique 14
4 ruin wall Ronda 12
5 Cork Oak (cult.) Casares 9
6 picnic place Santuaria de la Luz 8
7 picnic place Arcos de la Frontera 8
8 Cork Oak (cult.) Jubrique 8
9 Cork Oak (cult.) La Tienera Vieja 8
10 castle wall Jimena de la Frontera 8
the long Spanish history. In these semi-natural forests
M. calpeiana occurs at low densities if compared to
other (unnatural) habitats, such as semi-cultivated
Cork Oak woodland with the understorey cleared
away, rubbish dumps, roadsides and old walls (table
3).
The general impression is that the species benefits
from the milder activities of man and makes use of
the opportunities thus offered. These observations
contradict the opinion expressed by Snazell & Allison
(1989: 69), who describe the optimum habitat as un-
disturbed Quercus suber woodland with a light under-
storey and many large to medium-size stones. Snazell
& Allison maintain this view despite the several con-
trasting opinions and observations by others and re-
ported by themselves in the same publication. We
have shown that the population densities outside the
Cork Oak woodland are often higher than within.
The suggestion of Snazell that M. calpeiana is charac-
teristic of undisturbed Cork Oak woodland, and that
of Collins & Wells (1987) that it could serve as bio-
indicator for that type of habitat has herewith become
most unlikely.
Apparently the ecological spectrum of this species
is very broad. It has been found in a wide variety of
habitats. A species which invades new habitats and
Table 4. Average densities of Macrothele calpeiana nests
counted in plots of appproximately 20 m? within five distin-
guished categories of habitat in Southern Spain.
distinguished categoriess number average max
of habitat of counts density density
1 non-cultivated/non-forested 22 1.4 5
2 relatively undisturbed forest 20 3.0 7
3 cultivated Cork Oak 21 5.8 14
4 unmanaged culture land 33 SE) 15
5 managed culture land 14 0.4 3
176
settles, temporarily or permanently, in man-made
habitats is to some extent a culture follower. It is jus-
tified to call this species aggressive as to its ecological
strategy. Biological strategies, such as the number of
offspring, the dispersal of the young, the time and
amount of food needed to reach adulthood, and the
life-span are unknown so far. It is difficult, if not im-
possible, to analyse a possible extension of the distri-
bution area in the past. The sequence of the earlier
records may be suggestive of a recent colonization of
the Iberian Peninsula. However, this is no proof of its
speed of dispersal.
Nature conservation aspects
Snazell’s earlier report (Snazell 1986) on the distri-
bution, habitat and status of M. calpeiana in Spain ca-
me under the notice of the IUCN Conservation
Monitoring Centre at Cambridge (UK) when compi-
ling a list of invertebrates to be used in the Bern
Convention. They placed the species on their provi-
sional list (Collins & Wells 1987) and classified it as
‘vulnerable’, following Snazell’s (1986) description of
the species status as ‘fairly secure’ and ‘in no imme-
diate danger’ but forseeing as major threat ‘the im-
provement of the woodlands or the spread of housing
developments’. In the same year (1987) the species
was officially placed on Appendix II. This second ap-
pendix lists the endangered species the habitat of
which should be protected. Most species listed come
from the better known groups, such as Lepidoptera,
Coleoptera, Odonata, Mollusca, etc. Macrothele cal-
peiana is the only spider.
In 1991 the Habitat Directive of the European
Community has been accepted by the member states.
When a country ratifies the Directive it compels itself
to take relevant measures for the protection of the list-
ed species and their habitats. Macrothele calpeiana is,
again, in the appendix to this Directive because the
Bern Convention list of Invertebrates served as an ex-
ample and was copied without any criticism. Spain
thus is obliged to protect the habitat of M. calpeiana
as well as the spider itself. The Habitat Directive, in
contrast with the Bern Convention, is obligatory: if a
country does not satisfy the set rules it can be sum-
moned before the European Court.
Collins & Wells developed a set of seven criteria for
the selection of the species for their list (Collins &
Wells 1987). First of all (1) the species should be
under serious threat (endangered or vulnerable) or of
widespread conservation concern. Since the list is to
be used as an appendix to the Convention on the
Conservation of European Wildlife and Natural
Habitats, generally known as the Bern Convention,
(2) the species’ range in Europe should not be margi-
nal to a much wider range outside Europe. The Bern
Convention has political significance in the first place
VAN HELSDINGEN & DECAE: Macrothele calpeiana
Fig. 2. Macrothele calpeiana (Walckenaer) on its web
and this has led to the suggestion that (3) the species
must be reasonable easy to identify, and preferably fa-
miliar to members of the general public. Even though
challengeable on evolutionary grounds one can
understand that (4) in line with other listings in the
Appendices to the Convention only full species
should be considered. This is not a criterion, but a
practical rule. There are three more criteria which in
fact are recommendations of the compilers of the lists
for the different appendices: the final selection of spe-
cies (5) should come from a wide range of habitats,
particularly threatened ones, and (6) from a wide va-
riety of phyla and classes, while it should also (7) em-
brace a wide geographical range.
When judging a single species on the list one has to
consider criteria 1 to 3. If we do this with M. calpeia-
na, we easily arrive at the conclusion that the species
should not at all be listed in the Bern Convention.
We have pointed out that this spider is not under
threat, endangered or vulnerable in its relatively small
Spanish range of distribution. It is not strictly tied to
any specific habitat, but can be found in a wide varie-
ty of, often man-made, habitats. We have presented
arguments in support of the hypothesis that the spe-
cies has also a wider distribution outside Europe.
Thus we find the first two criteria not applicable to
M. calpeiana.
Remains the third criterion. M. calpeiana is a large,
black spider, upto to 7 cm long (legs included), with
two conspicuous ‘tails’, the spinnerets, at the end of
its abdomen (fig. 2). The general public at the most
will know its large web. The animal itself leads a cryp-
tic life in its burrow. It is the only spider of this size,
shape and habits that lives in the area and is quite
conspicuous when seen (e.g. by the people who man-
age the oak forests). We already indicated above that
this criterion is a political rather than a scientific one.
It has practical implications for the enforcement of
any directives in relation to the Bern Convention. For
a matter so serious as the protection of Invertebrates
and their natural habitats, however, this criterion has
no meaning.
A species becomes threatened with extinction usu-
ally through the activities of man. The threat is aimed
either directly at the animal, e.g. by hunting, over-
fishing or the use of selective biocides; or indirectly
through the many drastic changes in the environ-
ment, such as extensive drainage schemes, the bring-
ing into cultivation of natural habitats, the use of un-
selective biocides, the cutting down of forests, the
construction of motorways and town-development.
In the case of Invertebrates the indirect factors are
usually the cause of threat. Stenoecous species with
very strict habitat or food requirements are more vul-
WHY,
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
nerable than euryoecous species which can live nearly
anywhere. When a habitat is scarce and deteriorates
by one of the above influences all species narrowly
adapted to that habitat are threatened. If one wishes
to protect such a species one has to protect or restore
the habitat in general or a site in particular by ward-
ing off the threatening forces. It is the general practice
in nature conservation legislation to protect a habitat
through the species, i.e. the species is named and the
law compels that the habitat or site be protected or
managed in such a way that the species can survive.
It is a good policy to select a species of which the
biology and ecology are well known, the distribution
is properly mapped, and the densities measured. It
must be feasible to monitor the species and follow the
developments and the results of the measures eventu-
ally taken. The selection of the species should be
based on such (scientific) arguments, not on the pos-
sible recognition by the general public. The general
public should be interested in the lasting biological
diversity in general and support politicians to develop
legislation and give priority to nature conservation.
The general public must be kept informed in general
terms of the progress made. In our view the third cri-
terium should read as follows: “The species must be
taxonomically unambiguous and sufficiently defined
as to its biology and ecology. At the same time the
reasons for its decline should be fully understood’.
Species such as M. calpeiana then would not be listed
in the Bern Convention.
Future research
As we have pointed out the distribution in Spain is
as yet not clearly established, especially in the East
and Northwest. Further investigation should focus on
these regions, in particular the southern slopes of the
Sierra Morena and the Sierra Harana north of
Granada, and possibly the Sierra Nevada as well. It
would also be an enticing project to investigate the
northern countries of Africa, from Morocco to
Algeria, or even further to the East. Investigation in
this region will throw more light on the distribution
of M. calpeiana.
178
REFERENCES
Audouin, J. V., 1826. Explication sommaire des planches d’
arachnides de l'Egypte et de la Syrie, publiées par Jules-
César Savigny. Description de l’Egypte. — Atlas. Histoire
Naturelle 1(4): 99-186.
Bacelar, A., 1932. Sur quelques Araignées Theraphoses de la
faune Iberique. — Bulletin de la Société portugaise des
Sciences naturelles 11(15): 169-174.
Baker, A., 1991. A new species of the mite genus
Androlaelaps Berlese (Parasitiformes: Laelapidae) found in
association with the spider Macrothele calpeiana
(Walckenaer) (Mygalomorphae: Hexathelidae). — Bul-
letin of the British Arachnological Society 8(7): 219-223.
Blasco, A. & M. A. Ferrandez, 1986. El Genero Macrothele
Ausserer 1871 (Araneae, Dipluridae) en peninsula
Iberica. — Actas X Congresso Internacional de Arac-
nologia, Jaca/Espana 1: 311-320.
Collins, N. M. & S. M. Wells, 1987. Invertebrates in need
of special protection in Europe. — Council of Europe,
Nature and Environment series 35: 1-100.
Lucas, H., 1846. Histoire naturelle des animaux articulés. —
In: Exploration scientifique de l’Algérie pendant les
années 1840, 1841 et 1842, 1: 89-271. Paris.
Lucas, H., 1855. Sur une nouvelle espèce d’aranéide (Mygale
luctuosa) qui habite Espagne méridionale. — Annales de
la Société entomologique de France 3(3): 15-20.
Snazell, R. G., 1986. The spider genus Macrothele Ausserer
in Spain (Araneae; Dipluridae). — Bulletin of the British
Ecological Society 17(2): 80-83.
Snazell, R. G. & R. Allison, 1989. The genus Macrothele
Ausserer (Araneae, Hexathelidae) in Europe. — Bulletin of
the British Arachnological Society 8(3): 65-72.
Walckenaer, C. A., 1805. Tableau des aranéides: 1-88. —
Paris.
Received: 1 November 1991
Revised version accepted: September 1992
BREIDCHRATCEIEEE
University of Nebraska State Museum
NEW SREGIES AND COUNTRY RECORDS OF
BRAZIONINIGYELO GEREAKAKEOLEOPRTERA:
SCARABAEIDAE: DYNASTINAE)
Ratcliffe, B. C., 1992. New species and country records of Brazilian Cyclocephala (Coleoptera:
Scarabaeidae: Dynastinae). — Tijdschrift voor Entomologie 136: 179-190, figs. 1-34. [issN
0040-7496]. Published 1 December 1992.
Nine new species of Cyclocephala Latreille are described from Brazil: C. amplitarsis, C. rondon-
iana, C. sarpedon, and C. schmitzorum from Rondonia; C. deltoides from Para; and C. iani, C.
nodanotherwon, C. pseudoconfusa, and C. sarahae from Amazonas. New country records for
Brazil are given for C. boulardi Dechambre, C. guianae Endrödi, C. longa Endrödi, C. munda
Kirsch, C. ocellata Burmeister, and C. picipes (Olivier). Finally, the female of C. panthera
Dechambre is reported for the first time and briefly characterized.
Correspondence: Dr. B. C. Ratcliffe, Systematics Research Collections, W436 Nebraska Hall,
University of Nebraska State Museum, Lincoln, NE, 68588-0514, U.S.A.
Key words. — Brazil, Neotropics, Cyclocephala, new species, new country records.
In this paper are described new species and records
of Cyclocephala that I collected during my stay in
Brazil from 1976-1978, from collecting trips taken in
1980 and 1991, and from specimens either given or
loaned to me in the interim. Each new species has
been correlated with the most recent key to the genus
provided by Endrédi (1985) in order that the reader
might know where they fall out in the key.
Appropriate notations can then be made for future re-
ference. The parameres of the males, diagnostic in vir-
tually all cases, are illustrated and accompanied by
photo habitus pictures or line drawings to clarify the
descriptions of body morphology.
Several new country records are also listed.
Although I and a few others have known of these oc-
currences for several years, they have never been for-
mally placed into the literature. The purpose of lis-
ting these records is to provide an enhanced level of
reliability concerning identification of taxa that fall
outside of the previously published ranges.
Specimens have either been borrowed from or de-
posited in several collections. These collections are re-
ferred to by their four-letter acronyms as published in
Arnett and Samuelson (1986) or as indicated here if
not in Arnett and Samuelson:
BCRC - Brett C. Ratcliffe, Lincoln, NE, USA; CASC
- California Academy of Sciences, San Francisco, CA,
USA; DERC - David E. Russell, Davis, CA, USA;
HAHC - Henry and Anne Howden, Ottawa, Canada;
INPA - Instituto Nagional de Pesquisas da Amazonia,
Manaus, Brazil; LGBC - Larry G. Bezark, Sacramento,
CA, USA; MAMC - Miguel A. Morón, Xalapa,
Mexico; MZSP - Museo de Zoologia da Universidade
de Säo Paulo, Brazil; PKLC - Paul K. Lago, University,
MI, USA; vcpc - University of California, Davis,
CA, USA; UNSM - University of Nebraska State
Museum, Lincoln, NE, USA; USNM - U.S. National
Museum, Washington, D. C., USA.
Cyclocephala amplitarsis sp. n.
(figs. 1, 9-12)
Type material. — Holotype à, allotype ?, and eight pa-
ratypes labelled: ‘BRAZIL: Rondonia, 62 km S. Ariquemes,
Faz. Rancho Grande, 10° 32’ S, 62° 48’ W, XI-11-22-1991,
B. C. Ratcliffe, taken at light;’ 12 paratypes with same data
except collected by L. G. Bezark and D. E. Russell; 4 para-
types with same data except 25.xi.1991, S. L. Heydon, Hg-
vapor light; 5 additional paratypes with same data except da-
tes of 27-xi-1991 (2), 29-xi-1991 (1), 4-xii-1991 (1),
5-xii-1991 (1); 1 paratype labelled BRASIL: Rondonia,
Porto Velho, BR319, km 866, 28-1-1980, Jorge Arias,
C.D.C., 6-M (eters); 1 paratype labelled BRASIL: Mato
Grosso, Reserva Humboldt, 10° 11’ S, 59° 48’ W, 13-19-
VII-1977, B. C. Ratcliffe, taken at light. Holotype and allo-
type deposited at UNSM. Paratypes deposited at MZSP, CASC,
USNM, UNSM, UCDC, DERC, LGBC, and BCRC.
Description
Male holotype (fig. 1). — Length 12.9 mm; width
across humeri 5.5 mm. Colour of head, pronotum,
scutellum, pygidium, venter, and legs black; elytra
testaceous with all margins, spots on humeral and
apical umbones, and triangular area behind scutellum
black.
179
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-8. Cyclocephala spp., habitus of males. — 1, C. amplitarsis 2, C. deltoides, 3, C. iant; 4, C. nodanotherwon; 5, C. rondo-
niana, 6, C. sarahae, 7, C. sarpedon; 8, C. schmitzorum.
Head: Surface moderately densely punctate; punc-
tures on frons and basal half of clypeus small, punc-
tures on apical half of clypeus minute and denser.
Clypeus (fig. 9) with sides weakly converging, apex
broadly truncate, narrowly reflexed. Interocular
width 3.3 transverse eye diameters. Antenna with 10
segments, club subequal to segments 2-7.
Pronotum: Surface with punctation similar to that
of frons. Base lacking marginal line. Prosternal pro-
cess only moderate in length, obtusely rounded at ap-
ex.
Elytra: Surface minutely alutaceous, moderately
densely punctate; punctures moderately large, shal-
low, with double rows on disc. Sparse, minute, pale
setae present, especially in posterior half.
Pygidium: Surface completely, finely rugulose with
sparse, minute, pale setae. In lateral view, surface we-
akly convex.
Legs: Foretibia bidentate, both teeth close to each
other at apex of tibia. Foretarsus (fig. 10) with seg-
ment 4 expanded ventrally as a strong tooth; segment
5 greatly expanded for entire length, wider at base
180
than at apex. Claw immensely expanded, subtriangu-
lar, apex entire. Posterior tarsus almost twice as long
as posterior tibia.
Parameres: figs. 11-12.
Female allotype. — Length 12.6 mm; width across
humeri 5.6 mm. As holotype except in the following
respects: Head: Clypeus in apical half with punctures
not smaller, instead subequal in size to those on rest
of head. Elytra: Lateral margin slightly diluted at
about middle and with strong, protuberant knob at
about level of apical umbone. Pygidium nearly flat in
lateral view. Legs: Foretibia tridentate, teeth equally
spaced from one another. Foretarsi and claw not en-
larged.
Variation. — Males (18 paratypes). Length 12.3-
14.0 mm; width across humeri 5.5-5.9 mm. The pa-
ratypes do not differ significantly from the holotype.
Females (14 paratypes). — Length 12.1-13.5 mm;
width across humeri 5.4-6.1 mm. The females are
also remarkably similar to the allotype with no signi-
ficant differences noted.
Distribution
Cyclocephala amplitarsis is known from primary
lowland rainforests in central Rondonia and north-
western Mato Grosso in Brazil. All specimens were ta-
ken at lights, and one (collected by Jorge Arias) was
taken at a CDC light trap suspended at a height of 6
m.
Remarks
The enlarged foretarsi and claws of C. amplitarsis
bring to mind C. longimana Dechambre, 1980 and C.
tarsalis Dechambre, 1979, but those species have a
bordered pronotum as well as many other different
character states. Cyclocephala amplitarsis will key only
so far as couplet 239 (242) in Endrödi (1985) (if the
surface is considered setose, couplet 221/272) where
neither choice concerning foretibial teeth fits the cle-
arly bidentate state of this species.
If the elytra are considered to have short setae (lat-
ter half of couplet 221/272), then C. amplitarsis will
go as far as couplet 328/337 where neither choice
(concerning tridentate foretibia) is acceptable.
Etymology
From the Latin amplus, meaning large and the
Greek tarsor, meaning flat of the foot; large foot, a
noun in apposition in reference to the greatly swollen
foretarsi of the male.
Cyclocephala boulardi Dechambre
This species was described from Surinam, and the
following new country record from Brazil is not surpri-
sing in view of the relative affınities between the fau-
na of northern Brazil and the Guianas.
Brazil: Amazonas, Reserva Ducke, 26 km NE
Manaus, 1-xii-1976, B. C. Ratcliffe, at UV light (1
male in BCRC ).
Cyclocephala deltoides sp. n.
(figs. 2, 13-15)
Type material. — Holotype d labelled ‘BRASIL: Pará,
Urua, 65 km SW Itaituba on BR230, X-12-15-1977, B. C.
Ratcliffe, habitat: tropical evergreen forest, collected at
light.’ (UNSM).
Description
Male holotype (fig. 2). — Length 18.0 mm; width
across humeri 8.0 mm. Colour of clypeus, pronotum,
scutellum, pygidium, legs, and venter piceous; frons
black as well as all elytral margins (expanded into bro-
ad triangle behind scutellum), elongate spots on hu-
meral and apical umbones, and punctures on elytra.
Remainder of elytra testaceous.
Head: Frons and base of clypeus moderately dense-
RATCLIFFE: New Brazilian Cyclocephala
ly punctate; punctures moderate in size. Fron-
toclypeal suture a distinctly impressed, biarcuate line.
Clypeus in apical 2/3 finely and completely roughe-
ned; sides barely converging to very broad, truncate
apex, apex reflexed. Interocular width 3.0 transverse
eye diameters. Antenna 10-segmented, club subequal
in length to segments 2-7.
Pronotum: Surface moderately punctate on disc,
becoming a little denser on sides; punctures small on
disc, becoming moderate in size on sides. Base lacking
marginal line. Prosternal process short, mostly obscu-
red by brush of long setae.
Elytra: Surface with moderate to large punctures in
striate rows, double rows of punctures distinct; sur-
face, especially posterior half, with minute, moderate-
ly dense, reddish brown setae. Pygidium: Surface co-
arsely shagreened, setigerous; setae reddish brown,
minute at base, short on disc and moderately long ne-
ar apex. In lateral view, surface evenly convex.
Legs: Foretibia bidentate, both teeth close to one
another near apex of tibia. Foretarsus with segment 5
(claw segment) greatly expanded for its entire length
(fig. 13), wider at base than at apex. Claw of foretar-
sus enlarged, apex entire. Posterior tarsus about twice
as long as posterior tibia.
Parameres: figs. 14-15.
Distribution
Cyclocephala deltoides is known only from the Urua
locality in Brazil’s Amazon National Park, which was
just being established in 1977. It was taken at lights
on a road cut in primary lowland rainforest near the
end of dry season.
Remarks
Like Cyclocephala amplitarsis described elsewhere in
this paper, the enlarged foretarsi also resemble C.
longimana Dechambre and C. tarsalis Dechambre,
but those species have a margined base of the prono-
tum as well as other different character states.
Cyclocephala deltoides most closely resembles C. amp-
litarsis but differs from it in greater body size, smaller
size and different form of the foretarsal claw in males,
different form to the parameres, tricoloured body
(black, piceous, testaceous), larger eyes, longer and
denser pygidial setae, and with larger and more dis-
tinctive elytral punctures.
If the elytra are considered setose (Endrédi’s cou-
plet 221/272), then C. deltoides will key only to cou-
plet 239/242 where neither choice of foretibial teeth
works. If the choice is made for the second part of
couplet 221/272 (species with very short elytral
setae), then this species keys to couplet 328/337 whe-
re neither choice (concerning tridentate foretibia)
works.
181
TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
12
13
14
19
16 18
Figs. 9-12. Cyclocephala amplitarsis. — 9, dorsal view of head; 10, dorsal view of left foretibia and tarsus; 11, 12, caudal and la-
teral views of parameres; Figs. 13-15. C. deltoides. — 13, dorsal view of left foretibia and tarsus; 14, 15, caudal and lateral views
of parameres; Figs. 16-17. C. iani, caudal and lateral views of parameres; Figs. 18-19. C. nodanotherwon, caudal and lateral
views of parameres.
182
Etymology
From the Greek delta meaning shaped like a delta
or triangle; so named in reference to the large, black,
postscutellar mark on the elytra.
Cyclocephala guianae Endrödi
Endrödi described Cyclocephala guianae from
French Guiana. The records listed below are a new
country record for Brazil.
Brazil: Amazonas, Reserva Ducke, 26 km NE Manaus,
14-11-1977, B. C. Ratcliffe, taken at black light (3 males and
2 females in BCRC , 1 female at UNSM); same data but 14-i-
1978 and A. Soares collector (1 male at UNSM); same data
but 18-vi-1976 and A. Luna Dias collector (1 male at INPA,
1 male at UNSM); same data but 14-i-1978 and L. P.
Albuquerque collector (1 female at INPA). Amazonas, 60 km
N. Manaus, ZF3, km 23, 9-12-vi-1980, B. C. Ratcliffe (1
male and 2 females in BCRC ).
Cyclocephala iani sp. n.
(figs. 3, 16-17)
Type material. — Holotype d and allotype 9 labelled
‘BRASIL: Amazonas, Reserva Campinas, 60 mi N Manaus,
11-10-1977, W. W. Benson, ex: Annona nitida” Types de-
posited at UNSM.
Description
Male holotype (fig. 3). — Length 16.1 mm; width
across humeri 8.0 mm. Colour testaceous with black
frons and elytral markings; elytral marks consist of
large postscutellar spot, small spot just mesad of hu-
meral umbone, a smaller spot behind humerus, and a
short, transverse zigzag band just behind middle; cly-
peus, pygidium, legs, and venter piceous.
Head: Surface of frons moderately densely puncta-
te, punctures moderately large; clypeus densely punc-
tate, punctures mostly moderate in size, surface beco-
ming rugose in narrow band at apex. Clypeus with
apex nearly semicircular, margins narrowly reflexed.
Interocular width 2.4 transverse eye diameters.
Antenna with 10 segments, club subequal in length to
segments 2-7.
Pronotum: Surface with disc sparsely punctate,
punctures small; sides with punctures moderately
dense and moderately large, subequal in size to those
of frons. Base without marginal line. Prosternal pro-
cess long, posterior face at apex flattened.
Elytra: Surface finely alutaceous, moderately den-
sely punctate; punctures large, shallow, double rows
weak. Minute, testaceous setae present; setae modera-
te in density, most apparent on posterior half.
Pygidium: Surface sparsely punctate in centre, be-
coming denser on sides and rugose in lateral angles;
punctures on centre small, becoming large on sides,
setigerous; setae minute, testaceous. In lateral view,
RATCLIFFE: New Brazilian Cyclocephala
convexity moderate.
Legs: Foretibia tridentate, basal tooth small and
greatly removed from other teeth. Foretarsus enlar-
ged, larger claw with apex finely cleft. Posterior tarsus
slightly longer than posterior tibia.
Parameres: figs. 16-17.
Female allotype. — Length 16.5 mm; width across
humeri 8.1 mm. As holotype except in the following
respects: Head: Frons with punctures sparser, instead
only moderate in density; clypeus with surface more
generally roughened so punctures not as distinct.
Elytra: Lateral margin just behind middle weakly ex-
panded. Pygidium: Surface slightly more flattened,
small setae greatly reduced in size and number, barely
discernible. Legs: Foretarsi not enlarged.
Distribution
Cyclocephala iani is known only from Reserva
Campinas (Instituto Nacional de Pesquisas da
Amazonia) north of Manaus. This region is characte-
rized by lowland rainforest mixed with islands and
peninsulas of white sand soils (campinas) and associa-
ted low vegetation (campinarana). This soil/ vegeta-
tion type, known as campinas amazonicas (Anderson
et al. 1975), is where the reserve takes its name.
Remarks
In Endrödi (1985), both sexes of C. ani will key to
C. undata (Olivier) in their respective parts of the
Cyclocephala key (329/330 for males, 297/298 for fe-
males). The significant differences between them,
however, clearly indicate they are not conspecific. The
differences are primarily in the sculpturing of the cly-
peus, frons, and pronotum and especially in the form
of the parameres of the male. The parameres, in fact,
most closely resemble those of C. picipes (Olivier), but
other characters of clypeal shape, surface sculpturing,
and colour patterns separate the two.
Etymology
I am pleased to name this species after my youngest
son, Ian, who has so faithfully assisted me in the field
while collecting beetles.
Cyclocephala longa Endrödi
Endrédi (1963) described this species from Bolivia.
The locality data listed below, actually just across the
border from Bolivia, is a new country record.
Brazil, Rio Branco, 25-vi-1962, Hawarth collector
(1 male in BCRC ).
Cyclocephala munda Kirsch
Although widely scattered over northern South
America (Surinam, Colombia, Peru; Endrödi 1985),
183
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
C. munda has not been recorded from Brazil. The fol-
lowing is a new country record.
Brazil: Amazonas, Reserva Campinas, 60 mi N
Manaus, 14-1-1977, B. C. Ratcliffe (1 male in BCRC );
Amazonas, 15 km SE Barcelos, 14-1-1978, N.D.
Penny (1 male in BCRC ).
Cyclocephala nodanotherwon sp. n.
(figs. 4, 18-19)
Type material. — Holotype d and allotype © labelled
‘BRAZIL: Amazonas, Lago Anama, 160 km WSW Manaus,
XI-1981, R. Best.’ Single paratype with same data but date
of 1-v-1980. Holotype and allotype deposited at UNSM; pa-
ratype deposited in BCRC .
Description
Male holotype (fig. 4). — Length 15.6 mm; width
across humeri 7.9 mm. Colour reddish brown except
for black, triangular area mesad of each eye on frons.
Head: Surface finely alutaceous, moderately punc-
tate; punctures moderate in size, shallow.
Frontoclypeal suture distinct, weakly arcuate.
Clypeus with apex broadly and deeply emarginate,
narrowly reflexed. Interocular width 4.6 transverse
eye diameters. Antenna 10-segmented, club subequal
to segments 2-7.
Pronotum: Surface finely alutaceous, moderately
punctate, punctures becoming denser on sides; punc-
tures moderate in size, shallow, becoming a little lar-
ger and deeper on sides. Base without marginal line.
Prosternal process long, apex flattened in posterior
half.
Elytra: Surface finely alutaceous, moderately den-
sely punctate; punctures moderately large, shallow,
with 2 pair of double rows on disc; punctures in pos-
terior fourth with minute setae.
Pygidium: Surface moderately densely punctate;
punctures mostly large, shallow, setigerous; setae pa-
le, minute. In lateral view, surface evenly convex.
Legs: Foretibia tridentate, basal tooth very small,
well separated from other teeth. Foretarsus with 1
claw strongly enlarged, apex finely cleft, smaller ra-
mus not widely diverging. Posterior tarsus about 1/3
longer than posterior tibia.
Parameres: figs. 18-19.
Female allotype. — Length 15.8 mm; width across
humeri 7.9 mm. As holotype except in the following
respects: Colour of elytra light reddish brown. Head:
Clypeus with apex a little less emarginate. Pronotum:
Punctures sparse on disc to moderate in density on
sides. Elytra: Discal rows of punctures not distinct.
Sparse setae near apices long. Lateral margin slightly
expanded just behind middle with strongly expanded
lateral flange just behind this. Pygidium: Sparse setae
long. Convexity, in lateral view, weak. Legs: Claw of
184
foretarsus not enlarged.
Variation. — Male (1 paratype). Length 16.2 mm;
width across humeri 8.2 mm. Head: Surface a little
more densely punctate. Clypeus with apex a little less
emarginate. Elytra: Setae near apices short, distinct.
Pygidium: Setae moderately dense, short to modera-
tely-long.
Distribution
Cyclocephala nodanotherwon is known only from
Lake Anama WSW of Manaus.
Remarks
This species will key to either couplet 298 (C. gra-
vis Bates) or 306 (C. munda Kirsch) in Endrôdi
(1985) depending on the interpretation of the degree
of punctation on the pronotum (couplet 299). In eit-
her case, however, neither the parameres, vestiture, or
surface sculpturing conforms to that of C. nodan-
otherwon.
The examples of this new species were collected by
Robin Best (deceased), a colleague of mine at Brazil’s
National Institute of Amazonian Research (INPA),
while conducting research on manatees at Lago
Anama.
Etymology
This new species name is the result of an arbitrary
combination of letters (hence, treated as an indeclina-
ble noun) and results in a species name not inapprop-
riate in such a large genus.
Cyclocephala ocellata Burmeister
Known previously from Ecuador and Peru
(Endrédi 1985) and French Guiana (Dechambre,
1979), the following represents a new country record.
Brazil: Amazonas, 46 km N Manaus, 1-29-viii-
1979, H. M. Savage (1 female in BCRC ).
Cyclocephala panthera Dechambre
Dechambre (1979) described this distinctively
marked species from a single male specimen taken in
Para State, Brazil. I report here ten additional speci-
mens from the Canadian National Collection of
Insects that verify the valid status of Dechambre’s
species. The specimens (six males, three females) are
labelled BRAZIL: Distrito Federal, Estacion Florestal
Cabeca do Veado, 1100 m, 13-27 Oct. 1971, E. G.,
I. & E. A. Munroe.
The previously unknown females have the elytral
margin in the posterior third slightly (but distinctly)
thickened. In Endrédi’s (1985) key to female
Cyclocephala, it will key as far as couplet 261. Couplet
262 should be modified as follows:
262 (263) Surface bare.
262a. Head and pronotum piceous, elytra with
variable pattern (Figs. 270, 273). Guyana,
Peru, Brazil (Amazonas), Bolivia … … … …
ET N pugnax Arrow
262 b. Head with frons only black, pronotum
and elytra with distinct pattern. Central
Brazile ses panthera Dechambre
Cyclocephala picipes (Olivier)
Previously known from French Guiana (Endrödi
1985), the following constitute a new country record.
Brazil: Amazonas, Reserva Campinas, 60 km N
Manaus, 15-1-1977 (1 male) and 18-1-1978 (1 fema-
le), B. C. Ratcliffe (both in BCRC ).
Cyclocephala pseudoconfusa sp. n.
(figs. 20-24)
Type material. — Holotype à , allotype ®, and two para-
types labelled “BRASIL: Amazonas, Reserva Campinas, 60
km N Manaus, XI-22-1976, B. C. Ratcliffe.’ Holotype and
allotype deposited at UNSM. Paratypes deposited in BCRC .
Description
Male holotype. — Length 9.4 mm; width across hu-
meri 4.8 mm. Body form short, oval. Colour testace-
ous except for black frons.
Head: Surface of frons coarsely alutaceous with
moderately dense and moderately large punctures.
Clypeus with similar punctation mostly obscured by
surface roughness; sides subparallel, apex broadly
truncate, narrowly reflexed (fig. 20). Interocular
width 2.15 transverse eye diameters. Antenna 10-seg-
mented, club subequal to segments 2-7.
Pronotum: Surface finely alutaceous with punctu-
res moderate in density; punctures moderate in size
along midline, becoming moderately large on disc
and sides. Base without marginal bead. Prosternal
process long, posterior face at apex flattened.
Elytra: Surface finely alutaceous with moderate to
large, shallow punctures, double rows of punctures al-
most indistinct.
Pygidium: Surface densely punctate, punctures
small. In lateral view, pygidium evenly convex.
Legs: Foretibia (fig. 21) tridentate, basal tooth
slightly removed from other teeth. Claw of foretarsus
with apex deeply split (fig. 22). Posterior tarsus sub-
equal in length to posterior tibia.
Parameres: figs. 23-24.
Female allotype. — Length 10.6 mm; width across
humeri 5.2 mm. As holotype except in the following
respects: Elytra: Lateral margin slightly expanded just
behind middle. Pygidium: Surface weakly convex, ne-
arly flat. Legs: Foretarsus with claw not enlarged.
RATCLIFFE: New Brazilian Cyclocephala
Variation. — Males (2 paratypes). Length 11.1-12.0
mm; width across humeri 5.5-5.8 mm. The paratypes
do not differ appreciably from the holotype.
Distribution
Cyclocephala pseudoconfusa is known only from the
type locality north of Manaus. This is an area of whi-
te sand soils (campinas) that occur in peninsulas or
patches. This particular soil/vegetation type was char-
acterized by Anderson et al. (1975).
Remarks
This small, oval species will key as far as couplet
405 (404) in Endrödi (1985) which is C. confusa
Endrödi. While fitting in all respects to the characters
listed in the couplet, the parameres are vastly diffe-
rent. The same can also be said for the similarity in
external characters with C. perconfusa Dechambre,
1992. These three species should cluster together in
any key and be separable only by the form of the pa-
rameres in the males and the presence or absence of
an expanded elytral margin in females (margin simp-
le in C. perconfusa, with slight expansion in C. confu-
sa and C. pseudoconfusa).
Etymology
From the Greek pseudes, meaning false, i.e., the fal-
se C. confusa.
Cyclocephala rondoniana sp. n.
(figs. 5, 25-26)
Type material. — Holotype à , allotype ?, and one para-
type labelled "BRASIL: Rondonia, 62 km S. Ariquemes, Faz.
Rancho Grande, 10° 32’ S, 62° 48’ W, XI-11-22-1991, B.
C. Ratcliffe, collected at light.’ Additional paratype with sa-
me data and date of 12-22-XI-1991 and collected by L. G.
Bezark and D. E. Russell, Hg vapor light. Holotype and al-
lotype deposited at UNSM. Paratypes deposited in LGBC and
BCRC.
Description
Male holotype (fig. 5). — Length 9.1 mm; width
across humeri 4.0 mm. Colour testaceous except for
black frons, small spot on side of pronotum at middle,
oblique elytral vitta, sutural margin of elytra, and ba-
se of elytra for short distance either side of scutellum.
Head: Surface finely alutaceous, moderately punc-
tate; punctures small, becoming slightly less distinct
toward apex of clypeus. Frontoclypeal suture impres-
sed, arcuate. Clypeus with anterior angles rounded,
apex truncate, margins thickened and reflexed.
Interocular width 3.0 transverse eye diameters.
Antenna with 10 segments, club subequal in length to
segments 2-7.
Pronotum: Surface finely and weakly alutaceous,
sparsely punctate on disc, punctures small; punctures
185
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
2
AR
0
Sa 21
22
2: :
(\
25
31
JP
4 34
29 30 33
Figs. 20-24. Cyclocephala pseudoconfusa. — 20, dorsal view of head; 21, dorsal view of left foretibia and tarsus; 22, large ramus
of left foreclaw; 23, 24, caudal and lateral views of parameres; Figs. 25-26, C. rondoniana, caudal and lateral views of para-
meres; Figs. 27-28, C. sarahae, caudal and lateral views of parameres; Figs. 29-31, C. sarpedon. — 29, dorsal view of left fore-
tarsus; 30, 31, caudal and lateral views of parameres; Figs. 32-34. C. schmitzorum. — 32, dorsal view of left foretarsus; 33, 34,
caudal and lateral views of parameres.
186
becoming slightly larger and denser on sides. Base
with complete marginal line. Prosternal process short,
apex simply rounded.
Elytra: Surface finely alutaceous, punctate; punctu-
res moderate in density, small, shallow (often indis-
tinct), some setigerous; setae sparse, small, stout, red-
dish brown. Double rows not evident.
Pygidium: Surface minutely shagreened, moderate-
ly punctate; punctures small, shallow to indistinct.
Sparse setae present at base. In lateral view, surface re-
gularly convex.
Legs: Foretibia tridentate, basal tooth simply a we-
ak angulation, teeth subequally spaced. Foretarsus en-
larged, large claw with apex entire. Posterior tarsus
twice as long as posterior tibia.
Parameres: figs. 25-26.
Female allotype. — Length 8.7 mm; width across
humeri 3.8 mm. As holotype except in the following
respects: The black, elytral vitta reduced to 2 weakly
connected, elongate spots. Head: Punctures strong
and distinct over entire surface. Elytra: Punctures dis-
tinct. Margin slightly expanded at middle. Pygidium:
In lateral view, surface nearly flat. Legs: Foretibia dis-
tinctly tridentate, teeth subequally spaced. Foretarsus
and claw not enlarged.
Variation. — Females (2). Length 8.8 mm each;
width across humeri 3.9 mm each. Except for the
elytral markings being reduced to 2 small, separate
spots, the females do not vary significantly from the
allotype.
Distribution
Cyclocephala rondoniana is known only from the
lowland rainforests on the Schmitz farm in
Rondonia, Brazil.
Remarks
This species will key only so far as couplet 203
(202) in Endrödi (1985), after which nothing fits.
The parameres appear closest to C. quadripunctata
Höhne and C. peruana Endrödi, but those species
lack a marginal line on the base of the pronotum. The
oblique line on the elytra of the holotype is distincti-
ve although this is reduced to two spots in the three
females.
Etymology
The proper name of the territory of Rondonia has
been converted into a noun in the genitive to indica-
te ‘of Rondonia.’
Cyclocephala sarahae sp. n.
(figs. 6, 27-28)
Type material. — Holotype à, allotype ® and single pa-
ratype labelled ‘BRASIL: Amazonas, Rio Demeni nr.
RATCLIFFE: New Brazilian Cyclocephala
equator, IV-10-1978, L. R. Lacey.’ Holotype and allotype
deposited at UNSM; paratype deposited in BCRC .
Description
Male holotype (fig. 6). — Length 11.4 mm; width
across humeri 6.2 mm. Colour reddish brown except
for black frons and testaceous elytra. Form short,
oval.
Head: Surface of frons moderately punctate (more
so in anterior half); punctures moderate in size, deep.
Frontoclypeal suture distinct, sinuate. Clypeus with
surface transversely rugopunctate; sides convergent to
broad, subtruncate apex, apex reflexed. Interocular
width 2.25 transverse eye diameters. Antenna 10-seg-
mented, club a little longer than segments 2-7.
Pronotum: Surface moderately punctate, beco-
ming densely punctate on sides; punctures deep, mo-
derately large on disc, becoming large on sides.
Posterior angles broadly rounded. Base without mar-
ginal line. Prosternal process long, flattened at apex
on posterior half.
Elytra: Surface with large, shallow, umbilicate
punctures; punctures not in distinct rows, surface be-
tween punctures minutely alutaceous and shining.
Pygidium: Surface similar to that of pronotum
with moderately dense and moderately large punctu-
res on disc, punctures becoming denser and larger on
sides and base. In lateral view, pygidium strongly con-
vex.
Legs: Foretibia tridentate, basal tooth distinctly re-
moved from others. Foretarsus with 1 claw strongly
enlarged, apex finely cleft; smaller ramus of claw not
widely divergent from larger ramus. Posterior tarsus
short, subequal in length to posterior tibia.
Parameres: figs. 27-28.
Female allotype. — Length 13.5 mm; width across
humeri 7.4 mm. As holotype except in the following
respects: Head: Frontoclypeal suture broader, deeper.
Clypeus with surface more coarsely rugopunctate.
Elytra: Lateral margin barely expanded at about mid-
dle. Pygidium: Surface densely punctate on disc, be-
coming rugopunctate on base and sides. Convexity
weak. Foretarsus with claw not enlarged.
Variation. — Male (1 paratype). Length 10.3 mm;
width across humeri 5.6 mm. The single paratype
does not differ appreciably from the holotype.
Distribution
Cyclocephala sarahae is known only from the ‘type
locality’ on the Rio Demini north of Barcelos (a tri-
butary of the Rio Negro above Manaus).
Remarks
This species is externally similar to C. bicolor
Castelnau (couplet 408/409 in Endrôdi, 1985) ex-
cept that the large foretarsal claw is finely split in C.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
sarahae and broadly split in C. bicolor. The parameres
are also vastly different; in fact, no known species of
Cyclocephala has parameres even remotely resembling
those of C. sarahae. Cyclocephala sarahae will key as far
as couplet 404/405 in Endrödi (1985) which is C. te-
stacea ab. ovulum Bates. Cyclocephala testacea ab. ovul-
um has much simpler parameres, has the basal tooth
of the foretibia barely separated from the others, and
is not distinctly bicoloured as in C. sarahae.
This new species was collected by a colleague, Larry
Lacey, while we were both working for Brazil’s
National Institute for Amazonian Research (INPA).
During a boat trip upriver by Lacey, periodic stops
were made to spend the night, hence the imprecise lo-
cality information.
Etymology
I am pleased to name this species after my daugh-
ter, Sarah, who, even though she was not quite three
years old at the time, nevertheless helped me to col-
lect Amazonian insects during our stay in Brazil.
Cyclocephala sarpedon sp. n.
(figs. 7, 29-31)
Type material. — Holotype à , allotype 2, and 13 paraty-
pes labelled “BRASIL: Rondonia, 62 km S. Ariquemes, Faz.
Rancho Grande, 10° 32’ S, 62° 48’ W, XI-11-22-1991, B.
C. Ratcliffe, collected at light.’ Additional paratypes with sa-
me data (18) except collected by L. G. Bezark and D. E.
Russell and BRASIL: Mato Grosso, Reserva Humboldt, 10°
11° S, 59° 48’ W, 16-22-III-1977, B. C. Ratcliffe, collected
at light (4); Venezuela: Aragua, Rancho Grande, 20-VII-
1986, R. S. Miller colr. (1); Suriname, Raleigh Falls, 25-27-
VII-75, Coll: D. Engleman, collected at lights (1). Holotype
and allotype deposited at UNSM. Paratypes at CASC, MZSP,
UCDC, UNSM, USNM, PKLC, DERC, LGBC, HAHC, MAMC, and
BCRC .
Description
Male holotype (fig. 7). — Length 11.7 mm; width
across humeri 5.0 mm. Colour testaceous with pice-
ous tarsi and 3 small, black elytral spots (1 behind hu-
merus, a larger, oblique, elongate spot mesad of that,
and 1 in centre of elytron behind middle).
Head: Surface minutely alutaceous, moderately
densely punctate; punctures small, becoming smaller
and denser at clypeal apex. Clypeus with apex nar-
rowly parabolic, reflexed. Interocular width 3.0 trans-
verse eye diameters. Antenna with 10 segments, club
subequal in length to segments 2-7.
Pronotum: Surface similar to that of frons except
punctures slightly larger and sparser on disc. Base
with marginal line. Prosternal process short, conical.
Elytra: Surface with moderately dense, large, very
shallow punctures, double rows on disc barely evi-
dent; a few, short, pale setae present on apical half in
barely discernible rows.
188
Pygidium: Surface roughened with moderately
dense, short, reddish brown setae. In lateral view, sur-
face convex.
Legs: Foretibia bidentate. Foretarsus expanded in
size, claw bearing segment greatly enlarged at base on
medial surface (fig. 29), larger claw with apex entire
but bearing small tubercle subapically. Posterior tarsi
about 1.8 times longer than posterior tibia. Meso-
and metatibia with stout (not bristle-like) setae.
Parameres: figs. 30-31.
Female allotype. — Length 11.0 mm; width across
humerus 4.8 mm. As holotype except in the following
respects: Flared region of elytral margin black. Head:
Punctures slightly larger and denser. Elytra: Lateral
margin expanded just behind middle. Pygidium:
Surface glabrous, less convex. Legs: Foretibia triden-
tate, basal tooth only a weak angulation, all teeth sub-
equally spaced from one another.
Variation. — Males (13). Length 10.7-12.3 mm;
width across humeri 4.8-5.2 mm. The paratypes are
all similar to the holotype except that the size of the
elytral spots varies slightly (1 example with spots ne-
arly obsolete), and the Surinam specimen has the cly-
peal apex a little less narrowly rounded.
Females. (23). — Length 11.0-12.8 mm; width
across humeri 4.8-5.5 mm. The female paratypes do
not differ significantly from the allotype.
Distribution
Cyclocephala sarpedon is known from the lowland
rain-forests of central Rondonia and northwestern
Mato Grosso states in Brazil and from north-central
Surinam and north-central Venezuela. All specimens
were collected at lights.
Remarks
The specimens of the type series are remarkably
consistent in expression of their character states. The
narrowly parabolic clypeal apex, bidentate tibia, and
the form of the male parameres are diagnostic for this
species. It is very close in appearance to C. schmitzo-
rum, but the size is generally smaller, the parameres
differ in shape (usually difficult to see due to dense
setae on the parameres), and the males have the fifth
tarsomere greatly expanded at the base medially (fig.
29). Like C. schmitzorum, this species will key only so
far as couplet 193 (196) in Endrödi (1985) where, af-
ter that, character states no longer fit.
Etymology
Named after Sarpedon, a son of Zeus and Europa
and king of Lycia who was killed in the Trojan War
… and also because it was the very first epithet in a
scientific name (Graphium sarpedon) that | learned as
a boy. I’ve liked the sound of that word ever since.
Cyclocephala schmitzorum sp. n.
(figs. 8, 32-34)
Type material. — Holotype d, allotype 2, and 5 para-
types labelled ‘BRASIL: Rondonia, 62 km S. Ariquemes,
Faz. Rancho Grande, 10° 32’ S, 62° 48’ W, XI-11-22-1991,
B. C. Ratcliffe, collected at light.’ Five additional paratypes
with same data except collected by L. G. Bezark and D. E.
Russell, and Brasil: Amazonas, Reserva Ducke, 26 km NE
Manaus, 11-xi-1976, B. C. Ratcliffe (1); Suriname, Raleigh
Falls, 25-27-vii-1975, Coll: D. Engleman (1). Holotype and
allotype deposited at UNSM. Paratypes at USNM, CASC, DERC,
and BCRC .
Description
Male holotype (fig. 8). — Length 13.1 mm; width
across humeri 5.8 mm. Colour testaceous with pice-
ous tarsi and small, black elytral spots (1 behind hu-
merus, a smaller one mesad of that, and 1 in centre of
elytron just behind middle).
Head: Surface minutely alutaceous, moderately
densely punctate; punctures small, becoming smaller
and denser at clypeal apex. Clypeus with apex nar-
rowly parabolic, reflexed. Interocular width 3.0 trans-
verse eye diameters. Antenna with 10 segments, club
subequal in length to segments 2-7.
Pronotum: Surface similar in sculpturing to frons,
punctures a little sparser on disc, becoming a little lar-
ger and denser in posterior angles. Base with comp-
lete marginal line. Prosternal process short, conical.
Elytra: Surface with moderately dense, large, shal-
low punctures, double rows on disc evident; 6 rows of
setae present (1 sutural row, 2 rows on disc, 2 on side,
1 on lateral margin); setae short, stout (almost bristle-
like), reddish brown, sparse.
Pygidium: Surface completely roughened, with
moderately dense punctures; punctures small, setige-
rous; setae dense, long, reddish brown. In lateral view,
surface weakly convex.
Legs: Foretibia bidentate. Foretarsus expanded in
size (fig. 32), larger claw not split at apex. Posterior
tarsi about 1.6 times longer than posterior tibia.
Meso- and metatibia with bristle-like setae.
Parameres: figs. 33-34.
Female allotype. — Length 13.2 mm; width across
humeri 6.0 mm. As holotype except in the following
respects: Expanded region of elytral margin is black.
Elytra: Only a few setae present in apical half, rows
absent. Lateral margin dilated at level of first sternite
with swollen tumosity on side just above margin.
Pygidium: Surface glabrous. Legs: Foretibia tridenta-
te, basal tooth only a weak angulation, all teeth sube-
qually spaced from one another.
Variation. — Males (5). Length 11.2-13.2 mm;
width across humeri 5.4-6.2 mm. The male paratypes
do not differ significantly from the holotype. One
specimen has the elytral setae reduced or abraded
away on the anterior half.
RATCLIFFE: New Brazilian Cyclocephala
Females (2). — Length 12.0-12.9 mm; width across
humeri 5.8-6.0 mm. The female paratypes are similar
to the allotype.
Distribution
Cyclocephala schmitzorum is known from primary
lowland rainforests in central Rondonia and by a
single specimen each from near Manaus, Brazil and
north central Surinam. All specimens were taken at
lights.
Remarks
Although apparently subject to some abrasion, the
presence of almost bristle-like setae in rows on the
elytra in combination with the narrowly parabolic
clypeal apex, bidentate tibia in the male, and form of
the parameres will distinguish this species. It is very
close in overall appearance to C. sarpedon, but the si-
ze is slightly larger, the parameres differ (often diffi-
cult to see due to the dense pilosity on the parameres),
and the enlarged claw joint in the males is not flared
at the base (fig. 32). If the elytral surface is considered
setose (couplet 183/174 in Endrédi 1985), then it
will key to couplet 193 (196) and no further.
Etymology
I take great pleasure in naming this species after
Harald, Barbara, Tomas, Uta, and Aike Schmitz of
the Fazenda Rancho Grande in Rondonia, Brazil.
Their efforts to preserve rainforest habitat and to en-
courage entomological explorations on their ranch
not only brought to light this new species but also ser-
ves as a shining example of benevolent stewardship of
tropical biota.
ACKNOWLEDGEMENTS
For the loan and/or gift of specimens I thank Jorge
Arias (Brasilia, Brazil), Dodge Engleman (Margarita,
Panama), Steve Heydon (University of California at
Davis), Larry Bezark (Sacramento, CA), David
Russell (Davis, CA), and Paul Lago (University of
Mississippi). David Myers (Weeping Water, NE) and
the University of Nebraska Foundation are gratefully
acknowledged for making available travel funds to
Brazil in 1991. I am deeply appreciative of the efforts
of Thomas Emmel (University of Florida) for coordi-
nating the 1991 Brazil trip. For their warm hospitali-
ty and fine efforts to promote rainforest conservation
in Rondonia, I thank Harald, Barbara, Tomas, and
Uta Schmitz of the Fazenda Rancho Grande. Laura
Williams, Gail Littrell, and Charles Messenger (all
University of Nebraska) are acknowledged for their
line drawings, word processing, and photographic as-
sistance, respectively.
189
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
RESUMO
Nove novas especies de Cyclocephala do Brasil sao descri-
tas. Estas sao C. amplitarsis, C. rondoniana, C. sarpedon, e C.
schmitzorum da Rondônia; C. deltoides do Para; e C. iani, C.
nodanotherwon, C. pseudoconfusa, e C. sarahae do Amazonas.
Novos registros no Brasil sao C. boulardi Dechambre, C.
guianae Endrödi, C. longa Endrödi, C. munda Kirsch, C.
ocellata Burmeister, e C. picipes (Olivier). Finalmente, a fê-
mea de C. panthera Dechambre é relatada pela primeira vez
e caracterizada brevemente.
REFERENCES
Anderson, A. B., G. T. Prance, & B. W. P. de Albuquerque,
1975. Estudos sobre a vegetaçäo das Campinas
Amazonicas-III. — Acta Amazonica 5: 225-246.
Arnett, R. H. Jr. & G. A. Samuelson, 1986. The insect and
spider collections of the world. — E. J. Brill/Flora and
Fauna Pubs., Gainesville. 220 pp.
190
Dechambre, R.-P., 1979. Missiones entomologiques en
Guyane et au Brésil (*) [Coleoptera Dynastidae]. — Revue
Frangaise d’Entomologie (N. S.) 1: 160-168.
Dechambre, R.-P., 1980. Six nouvelles espéces de
Cyclocephala (Coleoptera, Dynastidae]. — Revue Française
d’Entomologie (N.S. ) 2: 42-49.
Dechambre, R.-P., 1992. Nouveaux Cyclocephalini des
genres Cyclocephala et Aspidolea (Col. Dynastidae). — In:
Lachaume, G., Les coléoptéres du monde, vol. 14,
Dynastidae américains. Sciences Nat., Venette, France.
Endrödi, S., 1963. Neue Cyclocephala-Arten. — Annales
Historico-Naturales Musei Nationalis Hungarici 55:
323-333.
Endrödi, S., 1985. The Dynastinae of the world. — Dr. W.
Junk Publishers, Dordrecht. 800 pp.
Received: 7 May 1992
Accepted: 10 August 1992
RAI ASCHOUIMEN
Museon, Department of Biology, Den Haag
REVISION OF THE GENERA EUCHROMIUS GUENEE
AND MIYAKEA MARUMO (LEPIDOPTERA:
CRAMBIDAE: CRAMBINAE)
Schouten, R. T. A., 1992. Revision of the genera Euchromius Guenée and Miyakea Marumo
(Lepidoptera: Crambidae: Crambinae). — Tijdschrift voor Entomologie 135: 191-274, figs. 1-
198. [issN 0040-7496]. Published 1 December 1992.
The Palaearctic, Nearctic, Neotropical, Oriental and Australian species of the genus
Euchromius Guenée, 1845 are revised. Euchromius is a senior synonym of Pseudoancylolomia
Ahmad, Zaidi & Kamaluddin, 1982. Four new species are described: £. circulus (Burma), £.
confusus (Afghanistan, Iran, Tadzhikistan), E. minutus (Brazil) and E. ornatus (India). The fol-
lowing synonymies are established: £. pulverosus (Christoph, 1887) is a senior synonym of E. co-
chlearellus (Amsel, 1949). Pseudoancylolomia qadrii Ahmad, Zaidi & Kamaluddin, 1982 is a
junior synonym of E. ocelleus (Haworth, 1811). E. viette: Bleszynski, 1961 is a senior synonym
of E. karsholti Ganev, 1987. The genus Miyakea Marumo, 1933 is revised, M. sinevi sp. n.
(Mongolia) is described and M. raddeellus (Caradja, 1910) is reinstated as a valid species. E. lu-
shanus Inoue, 1989 is transferred to Miyakea. E. delicatalis (Hampson, 1919) is transferred to
the genus Aurotalis Bleszynski, 1970. A key is provided together with full (re)descriptions of all
species, with notes on distribution and biology.
R. T. A. Schouten, Museon, Dept. of Biology, Stadhouderslaan 41, 2517 HV Den Haag, The
Netherlands.
Keywords. — Pyralidae; Crambinae; Euchromius; Miyakea; key; new species; checklist; tympanal
organs.
The classification of the Pyralidae has been subjec-
ted to several changes during the last two decades. In
the present paper the genera Euchromius Guenée,
1845 and Miyakea Marumo, 1933 are treated as be-
longing to the subfamily Crambinae of the
Crambidae, regarding all ‘pyralids’ at superfamily le-
vel as proposed by Minet (1982, 1991) and confirm-
ed by Solis & Mitter (1992).
The taxonomic position of Euchromius and
Miyakea has never been the subject of much discus-
sion. These taxa have always been placed in the
Crambinae. Zeller (1863) subdivided the Crambinae
as ‘Crambos’ and ‘Chilones’. Bleszynski (1965)
treated Euchromius and Miyakea close to
Metaeuchromius Bleszynski, 1960, Pseudargyria
Okano, 1962 and Eschata Walker, 1856. Gaskin
(1975) redefined the tribus Crambini and erected the
subtribus Crambina and Corynophorina with
Euchromius included in the subtribus Crambina.
Most of these subdivisions within the Crambinae are,
however, ill-defined and still little understood.
This paper provides the revision of all species of the
genera Euchromius and Miyakea are easily recognized
within the Crambinae by their yellow medial fascia
and the terminal black dots on the forewing.
Euchromius has a worldwide distribution with the ex-
ception of Indonesia, the Philippines and the islands
in the Pacific, while Miyakea is restricted to the east-
ern Palaeartic. Most species of Miyakea are known
from a few specimens and a few localities only, pro-
bably since very few collectors have visited China and
surrounding countries.
Euchromius and Miyakea are very similar in exter-
nal characters. As a consequence, several species of
Miyakea have been included erroneously in
Euchromius. In this paper a new set of distinguishing
characters is given.
For more information on the taxonomic history,
distribution and biology of the genera I refer to the
generic description.
MATERIAL AND METHODS
In the species descriptions the statement, ‘labial
palp two’ stands for: the length of the labial palp is
twice the diameter of the eye. The formula for the
black terminal dots mentioned in the species descrip-
tions is read as follows; 2-2-3-2 stands for: the black
terminal dots closest to the apex start with a group of
two, followed by a second group of two dots, a group
of three dots and finally a group of two dots near the
tornus (fig. 7).
Holotypes and lectotypes are referred to in both
191
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
con.
f. ty.
I. ty.
Spl
pro. ty. tym
ven. p po. ty
b. ty. ra. ty
con. sac. ty
sac. ty
ven. s
Figs. 1-6. Morphological characters. — 1, venation in Euchromius 2, venation in Aurotalis delicatalis 3, tympanal organs in
Euchromius. Abbreviations, b. ty. = bulla tympani; con. = conjunctivum; con. sac. ty. = conjunctivum saccus tympani; f. ty. =
fornix tympani; |. ty. = ligna tympani; po. ty. = pons tympani; pro. ty. = processus tympani; ra. ty. = ramus tympani; sac. ty.
= saccus tympani; spi. = spinula; tym. = tympanum; ven. p. = venula prima; ven. s. = venula secunda; 4, spinula in Euchromius,
5, tegula, evenly mottled, no dark patch; 6, tegula with dark patch in the middle.
192
the synonymy list and the paragraph ‘material’. The
spelling of the localities is taken literally from the la-
bels to make tracing of the material easier. For
Chinese material the Pinyin transcription is given.
Distribution maps are based on personally exam-
ined material only, except for Æ. ocelleus, where the
North-European localities are mostly based on reli-
able literature references.
The sequence of the species followed here is based
on overall similarity. No phylogenetic conclusions
should be drawn from this sequence. The species of
Euchromius which were revised and (re)described re-
cently (Schouten 1988, 1990) are only briefly treated,
but when available, new information has been added.
Dissections
Dissections and mounting of the genitalia, abdo-
mina and tympanal organs were carried out in the
way described by Robinson (1976) with some adjust-
ments. The abdomen is completely separated from
the thorax, taking care not to disrupt the tympanal
organs. Then, the abdomen is macerated for three mi-
nutes ina KOH 10% solution at 100° Celsius and
transferred to ethanol 70% for cleaning. Snipe feath-
ers are used to descale the abdomen and micro-pins
mounted into a match are used to remove remaining
tissue. Genitalia are separated from the abdomen by
cutting the posterior membrane of tergite and sterni-
te VIII. The aedeagus of the male genitalia is separa-
ted from the anellus and treated separately. After
staining with Chlorozol Black 1% (diluted in 70% et-
hanol), abdomen and genitalia are fixated in 96% et-
hanol. Male genitalia are fixated with the valvae spre-
ad. Abdomen and genitalia are preserved in Euparal
together on one slide, but under separate cover-slips.
The abdomen is placed under an elevated cover-slip
to avoid accidental flattening of the tympanal organs.
Since the Euparal essence will slowly evaporate, a va-
cuum underneath the coverslip may occur and air-
bubbles can arise, which can be avoided by putting
the slide in a stove at 40 °C for three days. The edges
of the coverslip are then sealed off with transparent
nail-polish.
Distortions caused by manipulations during the
preparation of the genitalia or by pressure of the
coverslip may easily arise. The coverslip can change
the orientation of the genitalia to a great extent. The
angle in which the valvae, aedeagus or any other part
of the genitalia, are bent upward or sideward can be
far outside the usual range of variation of the species
under consideration. Ignoring these artifacts easily
leads to incorrect interpretations, which may be ex-
emplified by the synonyms E. siuxellus Ganev &
Hacker, 1986 and E. gartheellus Derra, 1985 being a
result of a too narrow interpretation of differences ob-
served in genital slides (see Hacker 1986, Derra
SCHOUTEN: Euchromius and Miyakea
1987). Moreover, the differences observed are for
some part caused by distortions.
Depositories
Abbreviations (codens) for depositories follow
Arnett & Samuelson (1986), with the addition of:
DERR (collection G. Derra, Bamberg, Germany); GIEL
(collection C. Gielis, Lexmond, The Netherlands);
GORD (collection A. D. Gordon Agriculture and
Nature Study Institute, Degangya A, Israel); Huts
(collection K. J. Huisman, Wezep, The Netherlands);
HULL (collection E. Hull, Helsby, Great Britain); IN-
ou (collection H. Inoue, Tokyo, Japan); LAND (col-
lection B. Landry, Ottawa, Canada); Luca (collection
J. A. W. Lucas, Rotterdam, The Netherlands); MAES
(collection K. Maes, Gent, Belgium); PRIN (collection
W. de Prins, Berchem/Antwerpen, Belgium); ROBI
(collection R. Robineau, Tremblay-les-Gonesse,
France); RTAS (author’s collection); woLF (collection
H. W. van der Wolf, Nuenen, The Netherlands).
SYSTEMATIC PART
Euchromius Guenée, 1845
Euchromius Guenée, 1845: 324. Type species: Tinea bella
Hiibner, 1796: 29, designated by Desmarest (1857: 255).
Eromene Hiibner, [1825]: 366. Type species: Tinea bella
Hiibner, 1796: 29, by monotypy [nomen praeoccupa-
tum: Eromene Hübner, [1821]: 256., Noctuidae].
Araxes Stephens, 1834: 315, sensu Bleszynski (1963: 106),
(nec Araxes Walker, 1863). Type species sensu Bleszynski
(1963): Palparia ocellea Haworth, 1811: 486 [junior in-
valid designation].
Ommatopteryx Kirby, 1897: 274. Unnecessary replacement
for Euchromius Guenée, 1845 [mistaken homonymy with
Euchromia Hübner, 1819 and Euchromia Stephens,
1829].
eee Ahmad, Zaidi & Kamaluddin, 1982: 14.
Type species: Pseudoancylolomia gadrii Ahmad, Zaidi &
Kamaluddin, (1982: 15), by monotypy. Syn. n.
The taxonomic history at genus level has been
thoroughly explained by Bleszynski (1961, 1963) and
will not be repeated here. Ahmad, Zaidi &
Kamaluddin (1982) erected the new monotypic ge-
nus Pseudoancylolomia for the new species qadrii.
Pseudoancylolomia gadrii, however, is a junior syno-
nym of Euchromius ocelleus (Haworth, 1811) as is
pointed out under that species.
Pseudoancylolomia can be distinguished from other
Crambinae genera (Ahmad et al. 1982) by the well
developed proboscis and the anteriorly projecting
frons. These characters are, however, represented in
all species of Euchromius as well. Vein R5, mentioned
as stalked with the partly fused R3-R4, is identical to
that of all members of the genus Euchromius (see
A ina Met) alE1982 MANE et 2)" Therefore
Pseudoancylolomia Ahmad, Zaidi & Kamaluddin,
193
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
sl Pa d
Figs. 7-8. Forewings. — 7, fringes whitish at base, then even-
ly coloured; 8, forewing, fringes with three dark lines.
Abbreviations, f = fringes; pa = posterior area; df = double
fascia; aa = anterior area; td = terminal dots; sl = subterminal
line; d = dorsal edge; fl = fringes lines.
1982 is a junior synonym of Euchromius Guenée,
1845.
The firstly described species of the current genus
Euchromius is Tinea bella Hübner, 1796.
Subsequently, Haworth (1811) described the second
species now in Euchromius. Palparia ocellea. Costa
(1829), Treitschke (1832), Robinson (1870) and
Turati (1924) described species, later to be synonym-
ized with ocelleus. Until 1900 Zeller was the main au-
thor dealing with species now attributed to
Euchromius. He described anapiellus (Zeller, 1847),
vinculellus (Zeller, 1847), zonellus (Zeller, 1847) [=
ramburiellus (Duponchel, 1836)], superbellus (Zeller,
1849), wockeella (Zeller, 1863) [= superbellus (Zeller,
1849)] and cambridgei (Zeller, 1867). After 1900
mainly three authors were dealing with Euchromius
194
species. Caradja described four species in the period
1910-1937. Amsel described eight species in the peri-
od 1949-1961, of which five are now regarded as jun-
ior synonyms. Bleszynski in 1960-1967 made a seri-
ous effort to revise Euchromius, resulting in the
discovery of many synonyms and the description of
13 new species, of which only one turned later to have
a older synonym.
The Palaearctic species of Euchromius were revised
by Bleszynski (1965a), the Neotropical and South-
Nearctic ones (in part) by Capps (1966). Study of
material in museum collections not seen by previous
authors and material collected after the publication of
the above mentioned articles, has revealed many new
data. Study of material seen by Bleszynski and other
authors lead me several times to conclusions conflict-
ing with theirs. These conclusions and the new data
resulted in the updated revision of the genus
Euchromius presented in this paper.
The Oriental species are here revised for the first
time. The species from the Afrotropical and
Australian region were revised by Schouten (1988,
1990), but are included in the key. The present revi-
sion deals with 47 species, four of which are new to
science. The females of nivalis (Caradja, 1937) and
malekalis (Amsel, 1961) are described for the first
time. Of two species the female and of three the male
is unknown. Additional new species can be expected,
especially from Africa, China and the Oriental region
since large areas in these regions have never been vis-
ited by collectors of Lepidoptera.
Diagnosis
Dorsal insertion of ductus ejaculatorius is subter-
minal; gnathos is, in principle, armed with dorsal
thorns; tergite VIII of males with sclerotized pattern;
frons always projecting forward; M1 of hindwing lo-
cated in the upper angle of the open cell; base of M2
and M3 of hindwing without basal pointing stalk (fig.
1); spinula present (except in nivalis), simple hair-
like, without distinctly swollen base (fig. 4).
Morphology
The morphology is only discussed for features not
treated by Schouten (1988).
Intraspecific variation
The species of Euchromius show little intraspecific
variation. Only cambridgei (Zeller, 1867), gratiosellus
(Caradja, 1910), ramburiellus (Duponchel, 1836),
superbellus (Zeller, 1849) and vinculellus (Zeller,
1847) vary substantially in colouration. Variation in
structure of the genitalia is limited as well; some va-
riation is known in tanalis Schouten, 1988 and vin-
culellus.
SCHOUTEN: Euchromius and Miyakea
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195
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Tympanal organs
The tympanal organs (figs. 3, 4) have two func-
tions, detection of bat-sounds and locating the
sounds made the female (Minet 1983). The latter is
deduced from the sexual dimorphism of Catharia py-
renaelis (Dup.) (Pyraloidea: Crambidae: Odontiinae).
The male of this diurnal species has normally devel-
oped tympanal organs, whereas the female has atro-
phic tympanal organs. The dimorphism does not oc-
cur in night-flying species which are prone to be
attacked by bats. This suggests that the male uses his
tympanal organs to locate the female producing
sounds by vibrating her wings. Dahm et al. (1971)
demonstrated the use of the tympanal organs in the
localization of partners in Achrota grisella (Fabricius)
(Pyraloidea: Pyralidae: Galleriinae) in combination
with the use of pheromones. The diagnostic value at
subfamily and family level has been demonstrated by
Minet (1982, 1983, 1985, 1991) and Maes (1985).
The terminology of the tympanal organs follows
Maes (1985). In Euchromius the tympanum and con-
junctivum make an angle and the bulla tympani are
of the ‘open’ type. Consequently, Euchromius has
tympanal organs of the ‘Crambidae’ type, as defined
by Minet (1982) and Maes (1985). The tympanal or-
gans are located ventrally on the anterior part of the
abdomen.
Description of the tympanal organs in Euchromius
(figs. 3, 4). — The praecinctorium is simple sac-shaped
without a thorn-like sclerotization at the connection
with the pons tympani. The bulla tympani are bean-
shaped. The fornix tympani, supporting the conjunc-
tivum, are well developed and clearly visible. The ra-
mi tympani are connected, forming a well defined
semi-circle, enclosing the sacci tympani. At species
level the sacci tympani vary in size and shape.
Ventrally the two sacci tympani are connected by a
membrane, the conjunctivum saccus tympani. In
many cases, the width with which the two sacci tym-
pani are connected, in combination with the position
of the conjunctivum saccus tympani provide good di-
agnostic characters at the species level. The processi
tympani vary in size and are largest in ramburiellus,
occupying nearly half the breadth of the bulla tympa-
ni in that species. The processus tympani is orientat-
ed dorsal-ventrally and thus it is not always possible
to measure their size.
The processus tympani is connected to the tympa-
num by the scoloparium and spinula. The scolopar-
ium is a chordonotal nerve consisting of four scolo-
pale cells (Minet 1983). In mounted specimens, the
scoloparium is almost always lost, due to necessary
cleaning of the interior of the bulla tympani. The
scoloparium is connected to the spinula, a sclerite on
the inside of the tympanum. At a magnification of 40
times it usually can be seen as a small point or minute
196
Fig. 10. Venation in Miyakea.
line. In Crambinae spinulae have several distinct
shapes. The spinula in Euchromius (fig. 4) is a simple
hair-like structure, without a distinct swollen base,
implantated at an oblique angle on a minute elevation
and gradually tapering towards the distal end. The
distal end can be clean and thus pointed, or may still
have some remains of the scoloparium or tracheal tis-
sue connected to it, giving it a frayed tip. To estimate
the type of spinula of a genus or species usually many
slides should be checked. The spinula must be viewed
from several angles before a correct lateral drawing of
it can be made. The venula prima, situated lateral to
the fornix tympani, is always present. The venula se-
cunda, posterior of the fornix tympani, is more or less
an extension of the venula prima. In Euchromius the
venula secunda is well developed.
Distribution
The highest numbers of species occur in Africa
(19), the Mediterranean (16) and the Near and
Middle East (16). Australia has only one single spe-
cies, North America two and South America four. No
species are found on the islands of South-East Asia.
E. subcambridgei has a disjunct distribution in the
Sudan, Tunesia and the Cape Verdes. E. ocelleus with
an almost cosmopolitic distribution seems to be a mi-
gratory species which establishes temporar popula-
tions in localities far outside its main distribution
area from time to time.
Biology
E. ocelleus has been mentioned infesting food
(Hinton 1943). I am not aware of any recent confir-
mation of larvae found in food-products and, there-
fore I am sceptical about this claim. The caterpillars of
several species have been reared from dead leaves near
the base of plants, accidently larvae attack living plants
and flower-heads. Transportation of pupae with plant
remains seems likely (Meyrick 1895). Dead leaves of
mainly Compositae and one species of Dipsacaceae
are recorded as foodplants (Millière 1868).
Euchromius species are found in various habitats,
but most of them prefer dry and warm areas; no spe-
cies really enters the tropical rain forest. E. gnathosel-
lus and zephyrus have only been found in plantations
mixed with remains of secondary forest in West-
Africa, once covered with lowland tropical rain forest.
Several species are restricted to mountain areas and
plateaus e.g. tanalis and hampsoni in Africa, nivalis in
China and saltalis in South America. Many species are
found from sea-level up to more than 1000 m alti-
tude. £. ocelleus is recorded from the highest altitude,
4200 m (Afghanistan).
Excluded species
E. brunnealis (Hampson, 1919), micralis
(Hampson, 1919) and pygmea (Hering, 1903) do not
belong to the genus Euchromius in the present sense.
They should be classified in a separate genus to be de-
scribed in a forthcoming paper (Schouten, in prep.).
E. delicatalis (Hampson, 1919) is transferred to
Aurotalis Bleszynski, a genus not closely related to
Euchromius. Bleszynski (1970) regards Aurotalis a
close relative of Conotalis Hampson, 1919. The fore-
wing of E. delicatalis (fig. 2) has the Sc partly fused
with R1, whereas R2 is fused with R3+R4, a feature
never encountered in Euchromius (fig. 1). The male
genitalia of delicatalis lack any processus on the val-
vae, which are broad and of the same type as in
Aurotalis.
E. kuphitincta (Lucas, 1898) was transferred from
Diptychophora to Euchromius by Gaskin (1975a) .
The type material of this species has not been located
(Gaskin 1975a). The original description (Lucas
1898: 80), however, leaves no doubt that the species
is not a member of Euchromius. Several dentate lines
in the forewing are mentioned and the base of the
hindwing is said to have up to six or seven fine violet
fuscous transverse lines. Both features have never
been found in Euchromius. Especially the description
of the hindwing suggests attribution of this species to
the Diptychorini or the Pyraustinae. As long as the
holotype has not been located it will be impossible to
classify this species with certainty.
SCHOUTEN: Euchromius and Miyakea
Checklist of extant Euchromius Guenée
ZE
18.
39:
40.
SANADWNAKWN
ornatus sp. n.
. circulus sp. n.
. cornus Schouten, 1990
. californicalis (Packard, 1873)
. matador Bleszynski, 1966
. gnathosellus Schouten, 1988
. zephyrus Bleszynski, 1962
. tanalis Schouten, 1988
. mythus Bleszynski, 1970
. geminus Schouten, 1988
. galapagosalis Capps, 1966
. limaellus Bleszynski, 1967
. minutus sp. n.
. saltalis Capps, 1966
. ocelleus (Haworth, 1811)
cyrilli (Costa, 1829)
funiculella (Treitschke, 1832)
texana (Robinson, 1870)
gigantea (Turati, 1924)
qadrii (Ahmad et al., 1982) syn. n.
. pulverosus (Christoph, 1887)
cochlaearellus (Amsel, 1949) syn. n.
confusus sp. n.
vinculellus (Zeller, 1847)
corsicalis (Hampson, 1919)
asbenicola (Rothschild, 1921)
joiceyella (Schmidt, 1934)
bahrlutella (Amsel, 1949)
. anapiellus (Zeller, 1847)
. bellus (Hübner, 1796)
. bleszynskiellus Popescu-Gorj, 1964
roxanus Bleszynski, 1965
. bleszynskii Roesler, 1975
. scobiolae Bleszynski, 1965
. superbellus (Zeller, 1849)
wockeella (Zeller, 1863)
cypriusella (Amsel, 1958)
. keredjellus (Amsel, 1949)
. malekalis (Amsel, 1961)
. mouchai Bleszynski, 1961
. nivalis (Caradja, 1937)
. rayatellus (Amsel, 1949)
. gozmanyi Bleszynski, 1961
. gratiosellus (Caradja, 1910)
gartheellus Derra, 1985
siuxellus Ganev & Hacker, 1986
. jaxartellus (Erschoff, 1874)
. ramburiellus (Duponchel, 1836)
zonellus (Zeller, 1847)
luteella (Caradja, 1910)
islamella (Amsel, 1949)
. zagulajevi Bleszynski, 1965
. donum Schouten, 1988
. sudanellus Bleszynski, 1965
. subcambridgei Bleszynski, 1965
. cambridge: (Zeller, 1867)
luciella (Chrétien, 1907)
prototypa (Meyrick, 1935)
congruentella (Amsel, 1958)
szijjartoi (Gozmany, 1959)
ilkui (Gozmany, 1959)
viettei Bleszynski, 1961
karsholti Ganev, 1987 syn. n.
hampsoni (Rothschild, 1921)
197
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
41. klimeschi Bleszynski, 1961
42. discopis (Hampson, 1919)
43. labellum Schouten, 1988
44. aris Schouten, 1988
45. erum Schouten, 1988
46. locustus Schouten, 1988
47. nigrobasalis Schouten, 1988
Key to the species of Euchromius and Miyakea
The key can only be used when specimens are in
good condition. For a definite identification one
should check the genitalia.
ils
Eight, nine or ten black dots at termen of the
FOE wiley: ih seis hese ARR een nn tes 2
Four to seven black dots at termen of the fore-
wing (in E. rayatellus, E. superbellusand E. gozman-
yian eighth dot in apical position may occur) ..58
. Medial fascia double, divided by a silvery or white
line, inner fascia sometimes reduced … … … … …. 3
Medraltfasciatsing| NARE 28
. Black terminal dots arranged according to the
formula 2-2-2, 2-2-2-2-1 or 2-2-2-2-2, a group
offthreeldotsjisineyernpreseng eee tes ee
Black terminal dots arranged to the formula 1-2-
3-2, 2-2-3-2, or 2-3-1, a group of three dots is al-
WAYS! PECSENE sneren ese n tn 6
4. Frons rounded without a point ........... M. sinevi
— Frons pointed, with or without ridges ............. 5
5. Frons pointed, slender with several ridges (fig.
14). Forewings slender, tapering at the apex
Ra SAR RO TIE er o Ee NE er E. circulus
— Frons produced forward, pointed, ridges absent.
orewings NOFMAl eno E. matador
. Frons strongly produced forward, armed with
severaliridges a. on en E. zephyrus
Frons produced forward, one point, a ventral
ridge may be present or not (figs. 12, 21, 38) … 7
. Clypeus laterally lobbed, lobs produced forward
(Hig ID) eni aN A ROME E. ornatus
Clypeus not distinct laterally lobbed (figs. 21, 38)
. The outer medial fascia ending at circa one-third
of the dorsum; line between double fascia usually
whitish, seldom only silvery
The outer medial fascia ending at one-fourth to
one-sixth of the dorsum; line between double fas-
Cla silveryAs chien) Le nee 10
. The outer medial fascia nearly always bent out-
ward, especially near the inner margin. Forewing,
area adjacent to black terminal dots broad and
wihitisht., 8.2. ee EE ER E. bellus
The outer medial fascia straight; inner medial fas-
cia sometimes reduced; forewing, area adjacent to
black terminal dots narrow and usually yellow ...
RAT EERE E. anapiellus
198
10.
JRE
14.
157
16.
205
ZIE
DIA
Frons with clear ventral ridge, frons strongly con-
ical with corneous point ............... E. gnathosellus
Frons without ventral ridge (figs. 21, 38), frons
strongly conical or normally projecting, with cor-
NÉOUS POINTS. AN ee 11
Forewing brownish grey from base to inner medi-
al fascia, with no or little irroration by darker
scalesi rn ARE. E. galapagosalıs
Forewing, groundcolour creamy white, irroration
by darker scales from base to inner medial fascia
ee RTS TRE ME EL 12
Eemales Amel ati Aa en vets 20
. Gnathos with two basal projections (figs. 99,
KON SER nt a an RR 14
Gnathos without two basal projections (figs. 106,
MOD ti DER N Ere ENE a 15
Juxta without two large dorsal projections (fig.
(OO) Sr. Man E. californicalis
Juxta with two large dorsal projections (fig. 99)
RL ARE ER UE E. cornus
Aedeagus with one group of cornuti forming a
Packedirowi(tigs 21 06410) WENEN 16
Aedeagus with three groups of cornuti (figs. 109-
VNO dt A 117
Cucullus club-shaped. Processus basalis normal
sized, not blade-like at base. Two clear processi
interiorsvalyaci(tı 2100) E. tanalıs
Cucullus pointed. Processus basalis long, broad,
blade-like at base. One clear processus inferior
valva (ig MODE Re E. mythus
. The largest group of cornuti coiled, having a
woodrasp appearance (fig. 109) … … E. limaellus
The largest group of cornuti more or less straight,
forming a double row (figs. 110-112) ........... 18
. Dorsal spine of processus basalis arising from
middle of blade (fig. 112) ................... E. ocelleus
Dorsal spine of processus basalis arising from out-
ercomen (figlie 19
. Aedeagus with several cornuti at each side, point-
ing backwards giving the aedeagus a spear-like
appearance (fi gle E. saltalis
Aedeagus without backwards pointing cornuti at
the Sales (tte AURON E. minutus
Ostium tooth-shaped (figs. 156, 157)
Ostium toadstool or lip-shaped, not tooth-
shapeds(figs WIG 0) NE en etes DD
Ductus bursae with light sclerotizations near duc-
tus seminalis. Sclerotization under ostium most
prominent at the edges. Ostium longer than
broad, circa 1.5 times, (fig. 156) ......... E. tanalis
Ductus bursae without sclerotizations near duc-
tus seminalis. Sclerotization under ostium more
in centre, less at edges. Ostium about as high as
broad (figs 157) 0. ete ee E. mythus
Signa of equal size (figs. 151, 162, 163)
23:
24.
33.
34.
Signa unequal in size (figs. 158, 160, 161) .... 25
Ductus bursae short, not clearly sclerotized (fig.
ONERE eo ee ada de. 2 tti E. cornus
Ductus bursae long, clearly sclerotized (figs. 162,
163)
Hook-like projections lateral of the ostium.
Length ductus bursae circa two times that of bur-
sa copulatrix. Edges of tergite VIII connected
(6512281 C29 Henri ice au Lun E. saltalis
Hook-like projections lateral of ostium absent.
Length ductus bursae circa three times that of
bursa copulatrix. Edges of tergite VIII not con-
Nectedi(fig 163) en. E. ocelleus
. Edges of tergite VIII connected or not, wrinkled
(ipa) kent ne 26
Edges of tergite VIII connected, not wrinkled
(EUCH OSSEO 27
. Rim of ostium clearly dentate … … … E. limaellus
Rim of ostium not dentate ......... E. californicalis
. Larger signum three times as long as smaller sig-
UNI (NPA ee EE E. geminus
Larger signum less than two times as long as
smaller signum (fig. 161) .................. E. minutus
. Medial fascia broad, the entire length sprinkled
with black scales, reminding of a fingerprint.
Anterior area of the forewing for most part cov-
ered with black scales .................. E. nigrobasalis
Medial fascia normal, not or sprinkled with sil-
very scale. Anterior area of the forewing not cov-
ered with black scales, sometimes a few black
scales present, anterior area usually greyish or
brownish Ara N. een lee Bell, 29
costa
Medial fascia straight or gently arched, part near
inner margin may be lost or faint .................. 33
MIM ales mied ee a atin kbd de 31
Remalest anioni Annia leser IL voelen 32
. Uncus with a dorsal thorn (fig. 137) ..................
RR i leo ca nel E. sudanellus
Uncus without a dorsal thorn (fig. 139)
RR ritegno nin ast. Sala veu ed E. cambridgei
. One long and one shorter signum, a somewhat
triangular, broad sclerotized plate at the ventral
site of membrane of tergite VIII (fig. 187)
MERLINO Pennini ue REM cms im usi E. sudanellus
Signa of equal size, two very small slender sclero-
tized plates at the ventral site of membrane of ter-
re AUDI Gat ROSES SES E. cambridgei
Tegulae evenly mottled, no dark patch in the
middle (fig. 5)
Tegulae with dark patch in the middle (fig. 6) …
Ee rt FoF veh aria meet ee: 44
Fringes of forewing whitish at base, then evenly
ochreous brown (fig. 7)
Fringes of forewing with up to three dark lines
55)
36.
39:
40.
41.
42.
43.
44.
45.
SCHOUTEN: Euchromius and Miyakea
(LERS NEN diner dani sal, 39
Males (male of E. zagulajevi is unknown) … 36
Female ritieni mete 37
Cucullus with slight constriction about halfway.
Stalk of gnathos not terminally connected to ba-
sal part; stalk and basal part at right angle (fig.
11 SEE ica E. confusus
Cucullus without constriction. Stalk of gnathos
terminally connected to basal part; no clear angle
between stalk and basal part (fig. 114)
te A EREN Al ee E. pulverosus
. Ostium without prominent projection (fig. 184)
warn ee leet. alen en E. zagulajevi
Ostium with prominent projection (figs. 164,
165)
. Projection of ostium with parallel sides (fig. 164)
RR ee oi E. confusus
Projection of ostium without parallel sides, sides
convex: (Hp 1165) ETRE E. pulverosus
Malestein RAR AE RE RN 40
Females if. talia rate 42
Processus of sacculus slender, slightly bent up-
ward, not or slightly overlapping cucullus; pro-
cessus inferior valvae rounded, not elongated (fig.
SA) dent MARI E. gratiosellus
Processus of sacculus stout, clearly bent upward,
overlapping cucullus; processus inferior valvae
flat elongarediltie 155) rt 41
Processus of sacculus reaching almost halfway the
cucullus, at least one and a half times as long as
the breadth of cucullus (fig. 136)
ee ee E. ramburiellus
processus of sacculus at most reaching one-third
of the cucullus, at most as long as breadth of cu-
cullusi(fi 1135) Re ee E. jaxartellus
Ductus bursae under ostium only slightly en-
larged, with, only minor, second bulb-like projec-
tion (fie 2120) 2.2. ee E. gratiosellus
Ductus bursae under ostium enlarged, with clear-
ly separated sac-like projection (figs. 183, 185) …
RN dana ARTE 1 43
Projection of ostium long, extending over mem-
brane of tergite VIII (fig. 185) ... E. ramburiellus
Projection of ostium short, broad, circa equal-
sided, triangular, top just reaching membrane of
tergitenvlIN(fio A18 5) Penne E. jaxartellus
Frons conical with a small point (figs. 44, 70).
Fringes of the forewing with several dark lines
(fig. 8) or evenly brownish-grey at the end (fig. 7)
Frons bluntly produced forward, without point
(figs. 88, 96). Fringes of the forewing evenly
brownish-grey at the end (fig. 7)
Fringes of forewing with several dark lines (fig. 8)
Bea CORRE NANO PO ARENA PO 80
46.
47.
48.
. Tegumen with appendix angularis (fig. 148)
. Bursa copulatrix with two signa (fig. 195)
TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Labial palp two; subterminal fascia of hindwing
absent; fringes of forewing creamy white at base,
then two brown lines ......................... E. donum
Labial palp three; subterminal fascia of hindwing
present; fringes not clearly creamy white at base,
up to three brown lines ............ E. bleszynskiellus
Mal seca eee cen NN 48
Eemalesgart nafta Cao. 53
Gnathos without two dorsal thorns and terminal
part, processus basalis hardly visible or absent
(figs. 145, 147, 148)
Gnathos with two dorsal thorns and terminal
part, processus basalis clearly visible (figs. 140,
143, 144)
TEN AA HE AN E. locustus
147)
Valvae without heavy spines (fig. 145) .... £. aris
. Dorsal spike at base of processus of sacculus (fig.
DAAN ESP el ae E. discopis
Dorsal spike at base of processus of sacculus ab-
sents (figs 140) MAS) en 52
. Processus basalis narrowing abrupt, anterior part
of aedeagus normal (fig. 140) .............. E. viettei
Processus basalis narrowing soon, but gradual,
anterior part of aedeagus very slender (fig. 143) .
AA. MOREE Te LE NA LU LD A E. hampsoni
ae 54
Bursa copulatrix with one signum (figs. 190,
193)
. Projection formed by connected edges of tergite
VIII broadest at its base (fig. 195) ........... E. aris
Projection formed by connected edges of tergite
VIII not broadest at base, but more posterior (fig.
ISA) RR ERS IE NINE a) E. locustus
. Ductus bursae for most part strongly sclerotized
with large cornuti (fig. 196) .................. E. erum
Ductus bursae not strongly sclerotized (figs. 190,
IOT OI) er een ee: 56
. Ostium tooth-shaped (fig. 193) ......... E. discopis
Ostium not tooth-shaped (figs. 190, 191) ..... 57
. Lamella antevaginalis without anterior fold (fig.
OORDEEL fan ERE MEN Der E. viettei
Lamella antevaginalis with clear anterior fold (fig.
VON eren ee nd E. hampsoni
. Groundcolour of the forewing pure white, usual-
ly a clear dark brown to black spot in posterior
ACA He nne A PROS SA CUS E. nivalis
Groundcolour dirty to creamy white ............. 59
IMedialfascia double a ean eee: 60
Medialfscia she 65
. Yellow apical marking on forewing touching me-
AIRES Caen et ziale ine M. expansa
70.
The
>:
713:
. Frons produced forward with point (fig. 90)
Yellow apical marking clearly separated from me-
dial'fascian retreat ME ale cea REN 61
mi Aa eis) GIRONE VEEN RAS ann a RR 62
Female IE REIT aa te An RAM RUES 63
. Processus basalis absent (fig. 118) … M. lushanus
Processus basalis large, distinct (fig. 119) … … …
FILO di dn za he Lana tie fe M. raddeellus
. Under the ostium an elongated pouch, clearly
separated from the ductus bursae (fig. 199)
…. 64
Elongated pouch under the ostium absent
. Ostium without tongue-shaped projection (fig.
NGO) ER NES on M. lushanus
Ostium with tongue-shaped projection (fig. 167)
BEL EN ee dE RS RE ANA M. raddeellus
. Medial fascia clearly angled inward below costa
(fig. 9: E. labellum, E. cambridgei) … 66
Medial fascia straight or slightly arched ......... 68
. Medial fascia not sharply bent, angle in the mid-
dlefofithetwing. pete ene E. subcambridgei
Medial fascia more sharply bent, angle closely be-
lowithe eosta reet Anna EEE ER 67
wel A TE E. cambridgei
Frons produced forward without point
na RS ER E. labellum
. Frons rounded, without point; area adjacent to
black terminal dots white................. E. klimeschi
Frons rounded or not, with, a sometimes very
small, point, when absent then area to black ter-
mimalfdorshyello was ren me en 69
. Tegulae evenly mottled, no dark patch in the
middle (fig. 5)
Tegulae with dark patch in the middle (fig. 6), if
very faint then area adjacent to black terminal
dotsibroad era A ee eee 74
Area adjacent to black terminal dots narrow, usu-
ally yellow; subterminal line about midway
between terminal dots and termination of poste-
HOREN ARR Re PR Re E. vinculellus
Area adjacent to black terminal dots broad,
white; subterminal line closer to termination of
posterior area than to terminal dots … … … … 71
Malese Dre tee. bt sc ELEN 72
FEMALE Te LA TERA SE 0 78
Dorsal edge of sacculus with pointed projection,
stout spines at base of processus of sacculus ab-
SENI EMO) MEE E. bleszynskii
Dorsal edge of sacculus without pointed projec-
tion, stout spines at base of processus of sacculus
(ONU ZO RE RR Ae E. scobiolae
Ductus seminalis broad for some length, projec-
tion of ostium rectangular (fig. 174)
ELL ALE dl PERDO E. bleszynskii
Ductus seminalis broad for a short length, projec-
tion of ostium rounded (fig. 175) .... E. scobiolae
74. Fringes of forewing with one or two ochreous
brown lines, most clearly at the apex ............. 75
— Fringes of forewing highly shiny, evenly grey to
grey-brown, no ochreous brown lines ............ 80
dea Mall esa oord 76
SAMREMALES TE IRR nr rere 78
76. Aedeagus without cornuti (fig. 129) E. malekalıs
— Aedeagus with two or three groups of cornuti
(= Eee dI 77
77. Aedeagus with two groups of cornuti, processus
of sacculus short, but clearly free (fig. 128) … …
et i RR ra a E. keredjellus
— Aedeagus with three groups of cornuti, processus
of sacculus indistinct (fig. 130) … … E. mouchai
78. Ostium bean-shaped (fig. 179) … … E. malekalis
— Ostium lip-shaped (figs. 177, 178) ............... 79
79. Edges of tergite VIII connected, knot-shaped at
point of connection (fig. 177) … … E. keredjellus
— Edgesoftergite VIII not connected (fig. 178) ....
ER RER IZ ER AL Te Chio diri an Sd E. mouchai
SOMM alen EE N NP en 81
Eer al CSUN AREA QE PARA I 83
81. Dorsal thorns of gnathos elongated, terminal part
long (her) EEE NE E. superbellus
— Dorsal thorns of gnathos normal, short, terminal
partishore(fio nl 32)n..H nn 82
82. Aedeagus with three groups of cornuti; processus
of sacculus long, cucullus slender (fig. 132) ........
ORI RI DIA AI E. rayatellus
— Aedeagus with one large ridged cornutus; proces-
sus of sacculus short, cucullus broad (fig. 133) ...
OA ne zano E. gozmanyi
83. Ductus bursae with sleeve-like structure above
ductus seminalis (fig. 182) ............. E. gozmanyi
— Ductus bursae without sleeve-like structure (figs.
IGNIS DER E AO Re 84
84. Lamella antevaginalis large, shield-shaped, split
in the middle; membrane of tergite VIII without
thumb-like projection (fig. 180) ..... E. rayatellus
— Lamella antevaginalis small, not shield-shaped;
membrane of tergite VIII with thumb-like pro-
yectionm (figs Io) E. superbellus
1. Euchromius ornatus sp. n.
(figs. 11-13, 98)
Type material. — Holotype: 6, [India] ‘Dibidi, N. Coorg.
Newcome 5.12.06’, GS 17617 (BMNH).
Diagnosis. — Differs from all species in having the
clypeus laterally lobed in combination with the pres-
ence of a group of three dots in the terminal black
dots.
External characters male (fig. 11). — Wingspan 18
mm. Frons produced forward into very sharp point,
SCHOUTEN: Euchromius and Miyakea
creamy white to light brown, laterally lobed, ventral
ridge large; vertex creamy white; labial palp two, sides
creamy white at base, becoming brown, creamy white
from above and below; maxillary palp creamy white,
brown ringed at base of last segment; antenna creamy
white. Thorax brown; patagia brown; tegulae brown,
evenly mottled. Forewing, groundcolour white
densely suffused with ochreous to dark brown scales;
medial fascia double, slightly arched, running one-
fourth to one-fifth of the dorsum; subterminal line
dark brown, closer to termination of posterior area
than to terminal dots; area adjacent to terminal dots
white; eight black terminal dots, formula 2-2-3-1;
fringes shiny, at least one brownish line, too worn to
be described in more detail. Hindwing brown-grey,
subterminal fascia inconspicuous, termen darkly bor-
dered; fringes creamy white with brownish line.
Tergite VIII (fig. 12). — Sclerite normally sclero-
tized, only posteriormost part of posterior part vis-
ible.
Male genitalia (fig. 98). — Uncus basal-dorsal with
crown-like projection armed with heavy spines, more
posteriorly bearing tubercle; gnathos longer, two dor-
sal thorns large, terminal part long; tegumen without
appendix angularis; sacculus normal, processus of sac-
culus absent, processus basalis small, bent inward,
placed central on edge of valvae, processus inferior
valvae very prominent, forming tuberkel, cucullus
broad, sharply pointed upward; juxta triangular; vin-
culum normal; aedeagus small, two faint groups of
very small cornuti-like sclerotizations.
Biology. — Unknown. Holotype caught in
December.
Distribution (fig. 13). — Southern India.
Etymology. — Ornatus (Latin) refers to the graceful
ornamentation on the uncus.
2. Euchromius circulus sp. n.
(figs. 9, 13, 14, 150)
Type material. — Holotype: 9, [Burma] ‘Rangoon’, GS
17618. Paratype: 1 ?, [Burma] “Thyetmyo’, GS 17765
(holotype and paratype in BMNH).
Diagnosis. — Differs from all species with a double
fascia, except E. matador, in lacking a group of three
black terminal dots. Can be distinguished from E.
matador by the conspicuous ridges on the frons.
External characters female (figs. 9, 14). —
Wingspan 17 mm. Frons slender, sharply produced
forward, clear point and several ridges, light brown,
ventral ridge very small; vertex light brown; labial
palp two, sides creamy white becoming light brown,
creamy white from above and below; maxillary palp
creamy white; antenna creamy white. Thorax light
brown; patagia light brown to brown; tegulae light
201
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
circulus
ornatus
Figs. 11-20. Euchromius species. — 11, E. ornatus, head: lateral aspect, scales removed; 12, idem, sclerite of tergite VIII; 13,
distribution map of E. ornatusand E. circulus, 14. E. circulus, lateral aspect of head, scales removed, labial palp partly missing,
maxillary palp completely missing; 15, E. californicalis, head: lateral aspect, scales removed; 16, idem, sclerite of tergite VIII;
17, idem, distribution map (open circle: exact locality not found); 18, £. galapagosalis, head, lateral aspect, scales removed; 19,
idem, sclerite of tergite VIII; 20, idem, distribution map. Scale bar 0.5 mm to figs. 11-12, 14-16, 18-19.
brown to brown evenly mottled. Forewing, ground-
colour creamy white only posterior area densely suf-
fused with ochreous to dark brown scales; medial fas-
cia double, nearly straight, running to one-fourth of
the dorsum; subterminal line brown, about midway
between terminal dots and termination of posterior
area, area adjacent to terminal dots white, nine black
terminal dots, formula 2-2-2-2-1; fringes shiny
202
creamy white with two brown lines. Hindwing grey-
brown, subterminal fascia present; termen darkly bor-
dered; fringes creamy white with brownish line.
Female genitalia (fig. 150). — Papillae anales nor-
mal; membrane of tergite VIII with three narrow scle-
rotized bands, edges of tergite VIII around papillae
anales many folded; ostium complex heavily sclero-
tized; ductus bursae long, narrow; ductus seminalis
starting broad narrowing soon; bursa copulatrix ob-
long, two signa, club-shaped.
Male. — Unknown.
Biology. — Unknown.
Distribution (fig. 13). — Burma.
Etymology. — Circulus (Latin) refers to the circular,
ring-like shape of tergite VIII.
3. Euchromius cornus Schouten, 1990
(figs. 99, 151)
Euchromius cornus Schouten, 1990: 265. Holotype: à,
‘Sherlock R. W. Australia. E. Clements. 98-188’, GS
17616 (BMNH) [examined].
Material. — 21 G, 16 2.
Diagnosis. — Externally almost indistinguishable
from Euchromius species with a double medial fascia
and frons with one point. Differs in male genitalia
from all species in having the uncus armed with a
short, double, dorsal projection. Differs in female
genitalia from £. geminus, E. californicalis, E. limael-
lus, E. ocelleus, E. saltalis, E. mythus and E. tanalis by
its short ductus bursae without clear sclerotizations.
Distribution. — Australia.
4. Euchromius californicalis (Packard, 1873)
(figs. 15-17, 100, 152)
Eromene californicalis Packard, 1873: 264. Lectotype (desig-
nated by Capps 1966: 5): 4, ‘Type 14297’. (mczc) [ex-
amined].
Material. — 98 5,11 2. Canada: Alberta: Manyberries, 1
2 (BMNH). British Columbia: Nicola, 1 d (BMNH). USA:
California: 2 mi SW of Moraga, 2 ¢ (cısc); 4 mi E of
Petaluma, 1 d (RTAS), 2 & (ucpc); 8 km N of Tuckee, 1 4
(1ccm); 10 mi W of Simmler, 1 d (cısc); Arcata, 1 d
(vepe); Berkely, 1 d (casc), 1 d (cısc), 1 & (LAcM);
Bieber, 1 d (ucpc); Cambria, 4 d (AMNH), 1 d (RTAS);
Davis, 1 & (casc), 2 & (ucpc); El Cerrito, 1 d (casc), 1 &
(cisc); Half Moon Bay, 1 d (1ccm); Long Beach, 1 d
(AMNH); Moraga, 5 d (cısc); Napa, 3 d (BMNH), 3 d
(casc), 5 d (Mczc); Novata, 2 d (casc); Petaluma, 6 &
(LACM); Pinole, 1 d (cısc); Sacromento, 1 d (ucpc); San
Simeon, 3 d (AMNH), 1 d (RTAS); Santa Rosa, 1 d (LACM);
Sonoma, | d (cisc); Topaz, 1 & (Lacm); Ukiah, 1 d (cısc);
Vacaville, 1 4 (vepe). Colarado: Denver, 6 & (BMNH); Fort
Collins, 2 & (AMNH); Moffat, 2 &, 2 2 (1ccm), 1 d, 1 2
(RTAS). Idaho: East Fork Jarbidge, 1 & (Lacm); Moscow, 2
d (AMNH). Montana: Butte, 1 $ (MNHN). Nebraska: Grand
Island, 1 d (rccm). Nevada: 18 mi N of Paradise Valley, 1
3 (AMNH); 40 mi SW of Denio, 1 d (curc). New Mexico;
Chaca Canyon Nat. Mon., 1 & (LACM). South Dakota: 15
km W of Gattysbury, 1 2 (GIEL). North Dakota: Slope Co.
(locality not found), 1 d (AMNH). Oregon: 2 mi S of Ukiah,
1 d (cisc); Baker, 2 d (AMNH); Corvallis, 1 d (AMNH);
Klamath, 2 d (cısc); New Pine Creek, 1 d (ZMAN). Utah:
Hanksville, 1 6 (RTAS); Provo, 1 d, 1 2 (mczc); Vernal, 1
3,2 2 (teem), 1 à, 1 ® (RTAS). Washington: Pullman, 2
SCHOUTEN: Euchromius and Miyakea
d (ansp); 1 d (cuic). Wyoming: 5 mi N of Sundance, 1 4
(AMNH); 25 mi S of Bitterkreek, 1 d (iccm); Douglas, 3 4
(casc); Lone Tree, 1 9 (1ccm); Rock Springs, 1 2 (AMNH);
South Fork of Crazy Woman Creek, 1 9 (casc).
Diagnosis. — Externally difficult to separate from
species with a double fascia and frons with one clear
point. Differs from £. ocelleus, E. galapagosalis, E. sal-
talis, E. minutus, E. limaellus, E. mythus and E. tanal-
is in male genitalia by the basal projections on the
gnathos, from £. cornus in lacking the projections of
the juxta. E. californicalis can be distinguished in fe-
male genitalia by a mushroom-shaped ostium in com-
bination with signa unequal in size and a drop-shaped
sclerotization in the membrane of tergite VIII.
External characters male, female (fig. 15). —
Wingspan 20-24 mm. Frons sharply produced for-
ward into clear point, creamy white to light brown,
no ventral ridge; vertex creamy white; labial palp two
and a half to three, sides white at base, becoming
creamy white to light brown, creamy white from
above and below; maxillary palp white to creamy
white, brown ringed at base of last segment; antenna
creamy white, from about the middle very inconspic-
uous darkly ringed. Thorax creamy white; patagia
creamy white; tegulae creamy white evenly mottled,
sometimes with dark patch in the middle. Forewing,
groundcolour white, densely suffused with ochreous
to dark brown scales; medial fascia double, gently
arched to nearly straight, running to one-fifth to one-
sixth of the dorsum; subterminal line ochreous
brown, inconspicuous, closer to termination of poste-
rior area than to terminal dots; area adjacent to termi-
nal dots broad, white; nine or eight black terminal
dots, formula 2-2-3-2 or 2-2-3-1; fringes shiny
creamy white with two or three brown lines.
Hindwing creamy white to brown-grey, subterminal
fascia present or not, termen darkly bordered; fringes
white with brownish line.
Tergite VIII (fig. 16). — Sclerite normally sclero-
tized, posterior part rectangular, stalk broad ending
abruptly, anterior pattern invisible.
Male genitalia (fig. 100). — Uncus slightly bent,
with thin dorsal crest; gnathos longer, two dorsal-ba-
sal projections, two dorsal thorns, terminal part long;
tegumen without appendix angularis; sacculus nor-
mal, processus of sacculus absent, processus basalis
small, two processi inferiores valvae; cucullus bent
upward, with strong basal fold forming clear lob-like
projection; juxta small, triangular; vinculum long; ae-
deagus long, slender with three groups of cornuti,
sometimes only two clearly visible.
Female genitalia (fig. 152). — Papillae anales small;
membrane of tergite VIII with inconspicuous drop-
shaped sclerotization, edges of tergite VIII grooved,
not connected; ostium mushroom-shaped, not clear,
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
roughly granulate; ductus bursae long, more strongly
sclerotized in upper part; ductus seminalis narrow;
bursa copulatrix oblong, with two long signa, un-
equal in size.
Biology. — Unknown. Specimens caught from
March to September, with peaks in June, July and
August. Recorded from 300 m up to 2500 m altitude.
Distribution (fig. 17). — A West-Neartic distribu-
tion. Canada: British Columbia and Alberta. USA:
Washington, Idaho, Montana, North Dakota,
Oregon, Wyoming, California, Nevada, Utha,
Colorado, Nebraska and New Mexico.
Remarks. — The lectotype specimen in Mczc bears
only the label “Type 14297’. A second specimen with-
out any label is presumably a paralectotype. The third
specimen can almost certainly be excluded from the
syntype series since it has not been collected by
Edwards (Miller & Hodges 1990). The label reads:
C.H.F. March 95 Cal Behr 17, indicating that the
specimen was caught in March 1895 which is after
the date of publication of californicalis. The female
genitalia figured by Corbet and Tams (1943: 72, figs.
125 and 162) do not belong to £. californicalis.
5. Euchromius matador Bleszynski, 1966
(figs. 9, 101, 153)
Euchromius matador Bleszynski, 1966: 470. Holotype: 6,
‘Elisabethville Belgian Congo 4.V.1947 Ch. Seydel’, GS
4342, type 8921 (cncı) [examined]. Schouten (1988: 33)
[redescription].
Material. — 20 4, 13 9.
Diagnosis. — Differs from all species, except circu-
lus, in having a double medial fascia in combination
with the lack of a group of three black terminal dots.
Can be distinguished from circulus in lacking the
ridges on the frons.
Distribution. — Tanzania and Zaire.
6. Euchromius gnathosellus Schouten, 1988
(figs. 102, 154)
Euchromius gnathosellus Schouten, 1988: 35. Holotype: à,
‘Degbezere loc 7. 12 km E Bouafle 15-XII-1983 at light,
Cote D'Ivoire Bouafle R. T. A. Schouten & J. R. M.
Buijsen’, GS RS 101 (RMNH) [examined].
Material. — 14 6, 32 9.
Diagnosis. — Differs from all species, except orna-
tus, by its double medial fascia in combination with a
clear ventral ridge, a sharply pointed frons and the
presence of a group of three black terminal dots. Can
be distinguished from ornatus in lacking the lateral
lobes.
Distribution. — Senegal, Sierra Leone, Ivory Coast,
204
Ghana, Togo, Nigeria and the Central African
Republic.
7. Euchromius zephyrus Bleszynski, 1962
(figs. 103-105, 155)
Euchromius zephyrus Bleszynski, 1962: 129. Holotype: d,
‘Ilesha So. Nigeria (Capt. Humfrey)’, GS 7201 (BMNH)
[examined]. Schouten (1988: 36) [redescription].
Material. - 27 8,30 9.
Diagnosis. — Differs from all other species in hav-
ing a much ridged frons and vertex.
Distribution. — Senegal, Mali, Ivory Coast, Ghana
and Nigeria.
8. Euchromius tanalis Schouten, 1988
(figs. 106, 156)
Euchromius tanalis Schouten, 1988: 30. Holotype: d, ‘Tana
R. B.E. Africa. 3800 ft. 2.1.99. R. Crawshay 99-216’, GS
17482 (BMNH) [examined].
Material. — 13 6,2 2.
Diagnosis. — Difficult to distinguish externally
from other species with a double medial fascia.
Differs in male genitalia from all other species in hav-
ing a club-shaped cucullus in combination with a
packed row of cornuti in the aedeagus. In female gen-
italia it is characterized by a tooth-like ostium, the
edges directly under the ostium also sclerotized and
two signa of unequal length.
Distribution. — Ethiopia and Kenya.
9. Euchromius mythus Bleszynski, 1970
(figs. 107, 157)
Euchromius mythus Bleszynski, 1970: 2. Holotype: 6,
‘Diego Suarez’, GS 11310 (BMNH) [examined]. Schouten
(1988: 32) [redescription].
Material. — 32 6, 19 2.
Diagnosis. — Difficult to distinguish externally
from other species with a double medial fascia.
Differs in male genitalia in having the long pointed
processus basalis broad at base in combination with a
slender long packed row of cornuti in the aedeagus.
Differs in female genitalia in posessing a tooth-shaped
ostium, the edges directly under the ostium not scle-
rotized and two signa of unequal length.
Distribution. — Kenya, Tanzania, Zaire, Malawi,
Comoro Islands, Madagascar, Zimbabwe, Namibia
and South Africa.
10. Euchromius geminus Schouten, 1988
(fig. 158)
Euchromius geminus Schouten, 1988: 29. Holotype: 9,
‘Nairobi, BEA. may 1927 (D. M. Hopkins)’, GS 17483
(BMNH) [examined].
Material. — 1 9.
Diagnosis. — Male unknown. Differs from other
species with double medial fascia in female genitalia:
the two signa unequal in length in combination with
the ductus bursae sclerotized, only for a short part, di-
rectly under the ostium and having the edges of ter-
gite VIII connected.
Distribution. — Kenya.
11. Euchromius galapagosalis Capps, 1966
(figs. 18-20, 108, 159)
Euchromius galapagosalis Capps, 1966: 5. Holotype: 9,
‘South Seymour Galapagos April 23 1923’, GS 12,115
(USNM) [examined].
Material. — 24 6, 63 9. Ecuador: Pinzon, 7 d, 2 9
(casc), 2 d (RTAS). Santa Cruz: Academy Bay, 7 6, 4 9
(BMNH), 4 6, 51 ® (casc), 2 d, 4 2 (rras); Bella Vista
Trail, 1 & (casc); Conway Bay, 1 d (AMNH). South
Seymour, 1 2 (AMNH), 1 ® (USNM).
Diagnosis. — Difficult to distinguish externally
from E. californicalis, E. cornus, E. tanalis, E. mythus,
E. limaellus, E. ocelleus, E. saltalis, E. minutus. Can be
separated from these species in having the anterior
area of the forewing brownish grey with little or no ir-
roration by darker scales. In male genitalia it differs in
having a long slender aedeagus with three groups of
cornuti, no lateral cornuti, in combination with a
broad elongated processus basalis provided with a
small terminal spine which does not arise dorsally. £.
galapagosalis can be distinguished in female genitalia
by the edges of tergite VIII being connected in com-
bination with a toad-stool or lipshaped ostium with-
out lateral folds and signa unequal in size.
External characters male, female (fig. 18). —
Wingspan 15-21 mm. Frons produced forward with
point, creamy white to light brown, no ventral ridge;
vertex creamy white; labial palp two, sides white at
base, becoming light grey-brown, light grey-brown
from above and below; maxillary palp light brown,
dark ringed at base of last segment; antenna creamy
white to light grey. Thorax creamy white to light
brown; patagia creamy white to light brown, some-
times with two inconspicuous, broad, longitudinal
darker stripes; tegulae creamy white to light brown,
evenly mottled. Forewing, groundcolour creamy
white, posterior area densely suffused with dark
brown scales, anterior area evenly greyish coloured,
SCHOUTEN: Euchromius and Miyakea
without darker irroration or with but a few darker
scales; medial fascia double, gently arched or straight,
running to one-fifth of the dorsum; subterminal line
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white; eight, sometimes nine black terminal
dots, formula 2-2-3-1 or 2-2-3-2; fringes shiny, with
two, sometimes three brown lines. Hindwing grey,
subterminal fascia faint, termen darkly bordered;
fringes creamy white with brownish line.
Tergite VIII (fig. 19). — Sclerite normally sclero-
tized, posterior part rectangular, stalk ending abrupt-
ly, anterior part very faint, rounded.
Male genitalia (fig. 108). — Uncus normal, slightly
bent; gnathos longer, two dorsal thorns, terminal part
long; tegumen without appendix angularis; sacculus
narrow, processus of sacculus absent, processus basal-
is broad, elongated, provided with small terminal
spine, not arising dorsally, two processi inferiores val-
vae, cucullus very broad, bent upward, dorsal edge
minutely dentate; juxta triangular with two small
more strongly sclerotized dorsal projections; vincu-
lum large; aedeagus long, three groups of cornuti, one
of which consists of a double row of large cornuti.
Female genitalia (fig. 159). — Papillae anales nor-
mal; membrane of tergite VIII with inconspicuous
sclerotized line, edges of tergite VIII only connected
in middle part; ostium broad, lip-shaped, armed with
minute spines; ductus bursae long, for most part
strongly sclerotized; ductus seminalis narrow; bursa
copulatrix oblong, two signa, unequal in size.
Biology. — Unknown. Specimens caught in
January, February (top), April, May and July.
Distribution (fig. 20). — Endemic to the Galapagos
Islands.
12. Euchromius limaellus Bleszynski, 1967
(figs. 21, 22, 25, 109, 160)
Euchromius limaellus Bleszynski, 1967: 43. Holotype: à,
‘Lima Peru 20 May 1920 Cornell Univ. Expedition lot
607 sub 55’, GS 5161 (curc) [examined].
Material. - 4 d, 6 2. Columbia: Buena Ventura, 1 d
(ZMAC). Peru: Callao, 1 2 (MNHN); Lima, 1 d (BMNH), 1 d,
4 2 (cure), 1 d, 1 ® (RTAS).
Diagnosis. — Almost indistinguishable from species
with a double medial fascia and frons with one point.
Male genitalia differ from the other species in largest
group of cornuti being coiled, giving it a wood-rasp
appearance. Differs in female genitalia from £. gemi-
nus, E. minutus, E. cornus, E. ocelleus, E. tanalis, E.
mythus in having a mushroom-shaped ostium with-
out hooke-like projections, from E. californicalis in
having the rim of the ostium clearly dentate.
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
25
ocelleus
limaellus
saltalis
450%
minutus
Figs. 21-27. Euchromius species. — 21, E. limaellus, head, lateral aspect, scales removed; 22, idem, sclerite of tergite VIII; 23,
E. minutus, head, lateral aspect, scales removed; 24, idem, sclerite of tergite VIII; 25, distribution map of £. limaellus, E. min-
utus, E. saltalis and E. ocelleus, 26, E. saltalis, head, lateral aspect, scales removed; 27, idem, sclerite of tergite VIII. Scale bar
0.5 mm to figs. 21-24, 26-27.
External characters male, female (fig. 21). —
Wingspan 17-18 mm. Frons produced forward with
point, creamy white to brown, no ventral ridge; ver-
tex creamy white; labial palp two to two and a half,
sides white at base, becoming brown, creamy white
from above and below; maxillary palp creamy white
to light brown, dark ringed at base of last segment,
terminal part white; antenna creamy white. Thorax
creamy white to light brown; patagia creamy white to
light brown with two broad longitudinal light brown
stripes; tegulae creamy white, evenly mottled.
Forewing, groundcolour creamy white, densely suf-
fused with ochreous to dark brown scales; medial fas-
cia double, arched or nearly straight, running to one-
206
fifth to one-sixth of the dorsum; subterminal line
ochreous to dark brown, about midway between ter-
minal dots and termination of posterior area; area ad-
jacent to terminal dots white; nine black terminal
dots, formula 2-2-3-2; fringes shiny, creamy white
with two or three brown lines. Hindwing creamy
white, subterminal fascia present or not, termen dark-
ly bordered; fringes white to creamy white with
brownish line.
Tergite VIII (fig. 22). — Sclerite normally sclero-
tized, posterior part rounded to rectangular, stalk
quivered, anterior part faint.
Male genitalia (fig. 109). — Uncus tapering to sharp
pointed tip; gnathos longer, two dorsal thorns, termi-
nal part long; tegumen without appendix angularis,
membrane of tegumen armed with minute spines;
sacculus narrow, processus of sacculus absent, proces-
sus basalis small, sharply pointed, two processi inferi-
ores valvae, one at base of processus basalis, cucullus
very broad, bent upward, dorsal edge finely dentate in
basal half; juxta triangular with two small points; vin-
culum long; aedeagus long, three groups of cornuti,
largest group coiled, having a woodrasp appearance.
Female genitalia (fig. 160). — Papillae anales small;
membrane of tergite VIII without sclerotizations,
edges of tergite VIII strongly grooved, only connect-
ed in the middle forming a rectangular patch above
the ostium; ostium large mushroom-shaped, with
clear, roughly granulate lip; ductus bursae long,
strongly sclerotized, armed with many minute cornu-
ti; ductus seminalis narrow; bursa copulatrix oblong
to rounded with two long signa, unequal in size.
Biology. — Unknown. Caught in February and
May.
Distribution (fig. 25). — Columbia and Peru.
13. Euchromius minutus sp. n.
(figs. 23-25, 110, 161)
Type material. — Holotype: d, ‘19.3 Joazeiro Bras.
Exped. Penther ‘03’, GS 465 Naturhistorisches Museum,
Wien. Paratypes: 1 d, same data as holotype, 1 d, 1 9,
same data as holotype, but caught on 16 Mar and 17 Mar re-
spectively. All in nHMw. 1 d, 1 ®, same data as holotype,
but caught on 3.3 and 29.3 respectively, (RTAs). 1 ?
‘Brasilia Rio de Janeiro 23.X11.1961 (port) Gy. Topal’, GS
582 (zmuc).
Diagnosis. — Externally almost indistinguishable
from species with a double medial fascia and frons
with one point. Differs in male genitalia from E. cal-
ifornicalis, and E. cornus in lacking the basal projec-
tions of the gnathos, from E. tanalis, E. mythus, E. li-
maellus and E. ocelleus in having the dorsal edge of the
processus basalis minutely dentate, from £. saltalis in
lacking the lateral cornuti of the aedeagus. Differs in
female genitalia from £. tanalisand E. mythus in lack-
ing a tooth-shaped ostium, from E. cornus, E. saltalis
and £. ocelleus in having signa unequal in size, from £.
californicalis, E. galapagosalis, E. limaellus and E. gem-
inus in having the ostium simple, with two hook-like
projections.
External characters male, female (fig. 23). —
Wingspan 16-20 mm. Frons produced forward with
clear sharp point, creamy white to light brown, no
ventral ridge; vertex creamy white to light brown; la-
bial palp two, sides creamy white at base, becoming
light brown to brown, creamy white from above and
below; maxillary palp creamy white to light brown;
antenna creamy white. Thorax creamy white to light
SCHOUTEN: Euchromius and Miyakea
brown; patagia creamy white; tegulae creamy white to
light brown, evenly mottled. Forewing, groundcolour
creamy white densely suffused with ochreous to dark
brown scales; medial fascia double, gently arched to
nearly straight, running to one-fifth to one-fourth of
the dorsum; subterminal line ochreous brown, about
midway between terminal dots and termination of
posterior area; area adjacent to terminal dots white;
eight, sometimes nine black terminal dots, formula 2-
2-3-1 or 2-2-3-2; fringes shiny, creamy white with
one to two brown lines. Hindwing creamy white to
light brown, subterminal fascia absent, termen darkly
bordered; fringes white with sometimes inconspicu-
ous brownish line.
Tergite VIII (fig. 24). — Sclerite normally sclero-
tized, posterior part small, rounded, stalk long, ante-
rior part narrow, faint.
Male genitalia (fig. 110). — Uncus normal, slightly
bent, tapering to pointed tip; gnathos longer, two
dorsal thorns, terminal part long; tegumen without
appendix angularis; sacculus narrow, processus of sac-
culus absent, processus basalis large, broad, dorsal
edge minutely dentate, outer corner with bent spine,
two processi inferiores valvae, cucullus very broad,
dorsal edge dentate; juxta small, triangular with two
small dorsal projections; vinculum long; aedeagus
slender, three groups of cornuti, one group consisting
ofa double row.
Female genitalia (fig. 161). — Papillae anales nor-
mal; membrane of tergite VIII with one, half-circular
sclerotization, edges of tergite VIII connected, anteri-
or edge waved; ostium simple, two hook-like projec-
tions, minutely dentate; ductus bursae long heavily
sclerotized, armed with many cornuti in upper half;
ductus seminalis narrow; bursa copulatrix roundish,
signa unequal in size.
Biology. - Unknown. Specimens caught in March
and December.
Distribution (fig. 25). — Brazil.
Etymology. — Minutus (Latin) refers to the minute
dorsal dentation of the processus basalis.
14. Euchromius saltalis Capps, 1966
(figs. 25-27, 111, 162)
Euchromius saltalis Capps, 1966: 6. Holotype: d, ‘Salta
Argentina’, GS 12,110 (usnM) [examined].
Material. — 14 dg, 13 ©. Argentine: Alumine, 1 d
(zmuc); Arroyito, 1 d (RTAS), 7 d, 10 2 (zmuc); Chos
Malal, 1 2 (RTAS), 1 d, 1 2 (zmuc); Salta, 2 d, 1 2
(BMNH), 1 d (USNM); Tucuman, 2 d (ZMUN).
Diagnosis. — Externally almost indistinguishable
from species with a double medial fascia and frons
with one point. Differs in male genitalia from all spe-
cies in having a group of lateral cornuti posterior of
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
the anellus connection. Differs in female genitalia
from E. minutus, E. geminus, E. californicalis, E. li-
maellus, E. ocelleus, E. cornus, E. mythus and E. tanal-
is in having the edges of tergite VIII connected in
combination with signa equal in size.
External characters male, female (fig. 26). — 18-24
mm. Frons produced forward, with clear point,
creamy white, no ventral ridge; vertex creamy white;
labial palp two, sides creamy white to light brown,
creamy white to light brown from above and below;
maxillary palp light brown, terminal part lighter; an-
tenna creamy white, from about halfway in conspicu-
ous darkly ringed. Thorax light brown; patagia
creamy white to light brown; tegulae creamy white to
light brown, evenly mottled. Forewing, groundcolour
creamy white densely suffused with ochreous to
brown scales; medial fascia double, gently arched to
straight, running to one-fifth of the dorsum; subter-
minal line ochreous brown, about midway between
terminal dots and termination of posterior area; area
adjacent to terminal dots white; nine or eight black
terminal dots, formula 2-2-3-2 or 2-2-3-1; fringes
shiny with two ochreous brown lines. Hindwing
creamy white, subterminal fascia absent, termen
darkly bordered; fringes white with creamy white
line.
Tergite VIII (fig. 27). — Sclerite normally sclero-
tized, posterior part rectangular, stalk narrow, anteri-
or part faint.
Male genitalia (fig. 111). — Uncus normal, slightly
bent, tapering to pointed tip; gnathos longer, two
dorsal thorns, terminal part long; tegumen without
appendix angularis; sacculus narrow, processus of sac-
culus absent, processus basalis broad rectangular, dor-
sal edge smooth to very fine dentate, outer corner
provided with inwardly bent small spine, two proces-
si inferiores valvae, cucullus very broad, dorsal edge
minutely dentate; juxta normal sized, triangular; vin-
culum long; aedeagus long, armed with group of large
lateral cornuti posterior of anellus connection (these
may be lost due to copulation, the holotype only has
two lateral cornuti left), a double row of large cornu-
ti and two groups of small cornuti.
Female genitalia (fig. 162). — Papillae anales nor-
mal; membrane of tergite VIII with very inconspicu-
ous band-shaped sclerotization, edges of tergite VIII
connected, forming a rectangular patch above the os-
tium; ostium simple, lateral with hook-shaped projec-
tions, projections minutely dentate; ductus bursae
long, very strongly sclerotized with many cornuti in
upper half; ductus seminalis narrow; bursa copulatrix
oblong with two long equal-sized signa.
Biology. — Unknown. Caught in March,
November and December from 850 m (Chos Malal)
up to 2500 m (Salta) altitude.
Distribution (fig. 25). — Argentina.
208
15. Euchromius ocelleus (Haworth, 1811)
(figs. 25, 28-31, 112, 163)
Palparia ocellea Haworth, 1811: 486. Holotype: &, [United
Kingdom] ‘ocellea’, GS 17541 (BMHN) [examined].
Crambus cyrilli Costa, 1829: 11 [type material presumably
lost].
Phycis funiculella Treitschke, 1832: 200 [type material lost].
Eromene texana Robinson, 1870: 155 [type material pre-
sumably lost].
Eromene gigantea Turati, 1924: 63. Lectotype 9 (here desig-
nated): ‘Cyrenaica Bengasi 2.III.22 Geo. C. Kruger’, GS
17540 (BMNH) [examined].
Pseudoancylolomia gadrii Ahmad, Zaidi & Kamaluddin,
1982: 15. Holotype: d, ‘Kala Shan Kaku, on light.
(ESKK) [examined]. Syn. n.
Euchromius ocelleus (Haworth, 1811). Schouten (1988: 38)
[redescription].
Material. — 1955 6, 1733 ©.
Diagnosis. — Difficult to separate from other spe-
cies with a double medial fascia and a pointed frons.
Differs from these species in male genitalia by the
very broad cucullus, the dorsal edge being finely den-
tate in basal half in combination with processus basal-
is large, broad, provided with strong dorsally bent
spine, arising from middle of blade and the aedeagus
with three groups of cornuti, one of which consists of
a double row. Differs in female genitalia in having a
small mushroom or lipshaped ostium in combination
with the edges of tergite VIII separate and smooth
and signa of equal length.
Biology. — E. ocelleus has been found in various
habitats, viz. cultivated areas, macchia in the
Mediterranean, dry savanna in Africa, salt-steppe in
the Middle East and salt-areas in North-America. It
lives from sea leavel up to high altitudes, viz. up to
3100 m in the La Sal Mountains in Utah, at 1700 m
in the Greater Atlas, at 2600 m in Iran and 4200 m in
Afghanistan and at 2300 m in the north-eastern part
of Burma.
The larvae has been described by Hinton (1943:
202) from a single larva. It is said to be found on
stored food-products and owes it wide distribution to
human transportation with these products. Since
Hinton only had a single larva, it is not certain that it
was E. ocelleus, or it may be have been accidental that
it was found associated with stored food-products. I
agree with Hinton on the assumption that the wide
distribution of E. ocelleus is, at least partly, very likely
caused by human transportation. Lhomme (1935-
1946) gives dead and decaying leaves, etc. as the food-
source, Capps (1966) states that the larvae live on the
roots of corn and Milo maize (Sorghum). It seem very
plausible that ocelleus, as larvae or pupae, is transport-
ed with remains of plants and is not to be regarded as
a pest. It is also certainly migratory since several spec-
imens have been caught at sea as far as 50 miles out of
the coast.
Distribution (figs. 25, 28-31). — Africa: Ethiopia,
Somalia, Kenya, Tanzania, Central African Republic,
Zaire, Uganda, Malawi, Seychelles, Madagascar,
Mozambique, Zimbabwe, South Africa, Botswana,
Namibia, Angola, Equatorial Guinea, Nigeria, Niger,
Bourkina Fasso, Ghana, Senegal, Morocco, Algeria,
Tunisia, Libya, Egypt, Sudan. — Europe: Portugal,
Spain, France, Great Britain, Belgium, The
Netherlands, Denmark, Norway, Sweden, Finland,
Italy, Yugoslavia, Greece, Rumania, Bulgaria. — West
and Central Asia: Ukraine (Krim). — Near and
Middle East: Turkey, Israel, Jordan, Iraq, Saudi
Arabia, Yemen, Oman, Kuwait, Iran, Afghanistan,
Georgia, Turkmeniya, Tadzhikistan, Kirgiziya. —
Southern Asia: Pakistan, India, Nepal, Burma. —
Pacific: Hawaii. — Central and South America:
Mexico, Panama, Surinam, Netherlands Antilles. —
USA: California, Oregon, Washington, Nevada,
Utah, Arizona, New Mexico, Kansas, Colorado,
Texas, Oklahoma, Florida and Pennsylvania.
Remarks. — Ahmad et al. (1982: 15) published the
new species and genus Pseudoancylolomia qadrii.
Together with the description they figure the male
genitalia and wing-venation. Since Ahmad et al. were
clearly unaware of the existence of the genus
Euchromius and all the species belonging to it, they
make no reference as how to distinguish P. gadrii
from Euchromius ssp. The characters in which gadrii
differs from other species are therefore to be found
under the genus description. Arguments to synonym-
ize the genus Pseudoancylolomia with Euchromius have
been summarized before, while P. gadrii is evidently a
synonym of E. ocelleus (Haworth, 1811).
16. Euchromius pulverosus (Christoph, 1887)
(figs. 32-34, 114, 165)
Eromene pulverosus Christoph in Romanoff 1887: 47.
Lectotype 2 (designated by Bleszynski (1965a: 82)),
‘Ordubad’, GS 9538 (zmas) [examined].
Eromene cochlearella Amsel, 1949: 231. Holotype: 6, ‘Iran,
Elbursgebirge Keredj 1936 Leg. Brandt’, GS 317 (LNKD)
[examined]. Syn. n.
Material. — 13 &, 14 ®. Armenia: Ordubad, 1 9 (zmas).
Iran: Demavend, 1 d (INER); Derbend, 1 d (BMNH);
Keredj, 2 ? (BMNH), 1 d (LNKD); Nesa, 1 2 (NHRS). Iraq:
Rayat, 1 d (BMNH). Lebanon: Bcharre, 2 d (NHMW), 2 d
(zsMc); Beyrouth, 1 d (MNHN). Turkey: Aksehir, 1 d,1 9
(zsmc); Amanus, 1 d (zsmc); Artvin, 2 2 (Luca); Ergani, 1
3, 2 2 (mung); Kagizman, 1 @ (zmuc); Malatya, 1 ©
(BMNH); Marash, 1 9 (INER), 1 2 (NHMW), 1 d, 1 9
(ZSMC).
Diagnosis. — Externally indistinguishable from E.
SCHOUTEN: Euchromius and Miyakea
zagulajevi and E. confusus, but can be distinguished
from all other species by the eight terminal black dots,
tegulae evenly mottled, fringes of the forewing with-
out brown lines and a single medial fascia. Differs in
male genitalia from £. confusus in having the stalk of
the gnathos terminally connected to basal part. Basal
part of gnathos and stalk without clear angle. Differs
in female genitalia from E. confusus in having the pro-
jection of the ostium with convex sides. The projec-
tion of the ostium lacks in E. zagulajevi.
External characters male, female (fig. 32). —
Wingspan 20-23 mm. Frons produced forward, with
point, creamy white, no ventral ridge; vertex creamy
white; labial palp two, sides creamy white at base, be-
coming brown, creamy white from above and below;
maxillary palp brown, terminal part creamy white;
antenna creamy white to grey brown. Thorax light to
dark brown; patagia creamy white with two broad
longitudinal brown stripes; tegulae creamy white to
brown, evenly mottled. Forewing, groundcolour
creamy white, densely suffused with ochreous to dark
brown scales; medial fascia single, slightly arched,
running to one-third of the dorsum; subterminal line
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white; eight black terminal dots, formula 2-2-3-
1; fringes creamy white at base then evenly grey-
brown. Hindwing grey-brown, subterminal fascia ab-
sent, termen darkly bordered; fringes creamy white
with brownish line.
Tergite VIII (fig. 33). — Sclerite normally sclero-
tized, posterior part triangular, stalk short broad, an-
terior part faint.
Male genitalia (fig. 114). Uncus normal, tapering
to pointed tip; gnathos longer, at base with several
dorso-lateral spines, dorsal thorns absent, terminal
part short swollen; tegumen without appendix angu-
laris; sacculus broad, processus of sacculus not reach-
ing end of cucullus, tip slightly bent upward, strong-
ly sclerotized, processus basalis stout, nearly straight,
processus inferior valvae small, elongated, cucullus
normal sized, bent upward; juxta triangular; vincu-
lum normal; aedeagus long, slender one small row of
cornuti.
Female genitalia (fig. 165). — Papillae anales nor-
mal; membrane of tergite VIII with broad tie-like
sclerotization armed with node-like central projec-
tion, edges of tergite VIII connected; ostium complex
with rounded finger-like projection, no parallel sides;
ductus bursae long, slightly grooved in upper part;
ductus seminalis narrow; bursa copulatrix oblong, no
signa.
Biology. — Largely unknown. One long generation,
starting in May ending in first half of September.
Most abundant in May and June. This species has
been caught up to 2100 m.
209
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
pulverosus
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210
Distribution (fig. 34). — Found in Turkey, Iran,
Iraq, Armenia and Lebanon.
Remarks. — There has been much confusion about
the identity of this species. E. pulverosus (Christoph,
1887) was originally described from female material
only. Amsel (1949: 234) mentioned a male which he
found somewhat different from Æ. pulverosus, which
he conditionally named ‘cochlearella. Bleszynski &
Collins (1962: 305) first used the name Euchromius
cochlearellus (Amsel, 1949). Bleszynski (1965a) re-
corded five male specimens not conspecific with the
male of £. cochlearellus (Amsel, 1949), which he re-
garded the male of E. pulverosus. He also found two
females, difficult to separate from the females of E.
pulverosus, which he regarded the females of Æ
cochlearellus.
It appeared, however, that the males and females of
the two species had been mixed up by Bleszynski.
Due to the rearranging of the males and females, £.
cochlearellus has become a junior synonym of E. pul-
verosus, leaving the other species without a name. The
name for the latter is Euchromius confusus.
17. Euchromius confusus sp. n.
(figs. 34-36, 113, 164)
Euchromius cochlearellus auct., nec Amsel [misidentifica-
tion].
Type material. — Holotype: 9, ‘1-9.8.1962 Afghanistan.
Paghman 30 km NW v. Kabul, 2100m E. & A. Vartian
leg”, GS 17758 (BMNH). Paratypes: 2 6, ‘20.u.28.VI.1965
Afghanistan Paghman, 30 km NW v. Kabul, 2500m Kasy &
Vartian, (NHMW). 1 9, ‘20.-30.7.1962 Afghanistan
Paghman, 30 km NW v. Kabul, 2100m E. & A. Vartian
leg.’, (NHMW). 1 2, same data as holotype, (NHMw). 1 9, ‘J.
Klapperich Ghorbandtal 1900m 26.8.52 O-Afghanistan’,
(BMNH). 1 d, 1 ©, same data as previous, (RTAS) respective-
ly (LNKD). 3 d, Kondara Tadzhikistan Gissar-range 21.VII
946, (BMNH). 1 d, O-Afghanistan Gulbahar 1700m 2-9,
(LNKD). 1 ®, Zeitun, (BMNH). 2 9, ‘Iran Taug-Ab nr.
Firuzabad Fars 4000ft 6-IV-50 FF19 E.P. Wiltshire’, GS
17761 (BMNH). 1 d, no locality label, GS 16600 (BMNH). 1
3, no locality label, (BMNH). 1 d,1 2, Tadzhikistan Gissar
27.v.1985, (ZMAS).
Diagnosis. — Externally indistinguishable from £.
zagulajevi and E. pulverosus, but can be distinguished
from all other species by the eight terminal black dots,
SCHOUTEN: Euchromius and Miyakea
tegulae evenly mottled, fringes of the forewing with-
out brown lines and a single medial fascia. Differs in
male genitalia from £. pulverosus in having the stalk of
the gnathos not terminally connected to basal part.
Basal part of gnathos and stalk connected at right an-
gle. Differs in female genitalia from £. pulverosus in
having the projection of the ostium parallel-sided.
The projection of the ostium lacks in E. zagulajevi.
External characters male, female (fig. 35). —
Wingspan 20-23 mm. Frons produced forward, with
point, creamy white to brown, no ventral ridge; ver-
tex creamy white to brown; labial palp two, sides
creamy white at base, soon becoming brown, creamy
white from above and below; maxillary palp brown,
terminal part creamy white; antenna grey to grey-
brown. Thorax light to dark brown; patagia creamy
white with two broad longitudinal light to dark
brown stripes; tegulae light to dark brown, evenly
mottled. Forewing, groundcolour creamy white
densely suffused with ochreous to dark brown scales;
medial fascia single, slightly bent, running to one-
third of the dorsum; subterminal line brown, about
midway between terminal dots and termination of
posterior area; area adjacent to terminal dots white;
eight black terminal dots, formula 2-2-3-1; fringes
creamy white at base then evenly grey-brown.
Hindwing grey-brown, subterminal fascia absent, ter-
men darkly bordered; fringes creamy white with
brownish line.
Tergite VIII (fig. 36). — Sclerite normally sclero-
tized, posterior part triangular, stalk and anterior pat-
tern not visible.
Male genitalia (fig. 113). — Uncus normal, bent;
gnathos longer, basal part enlarged, basal edge den-
tate with strong spines, terminal part short, connect-
ed at right angle to stalk; sacculus normal, processus
of sacculus reaching up to half of cucullus, bent up-
ward, strongly sclerotized, processus basalis long,
slightly bent inward, dorsal edge inconspicuous
roughly dentate in posterior part, processus inferior
valvae elongated, cucullus long, slender, slightly con-
stricted about halfway; juxta slightly triangular; vin-
culum normal sized; aedeagus normal sized, one small
clear row of cornuti, posterior a second faint group of
cornuti.
Female genitalia (fig. 164). — Papillae anales nor-
mal; membrane of tergite VIII with tie-like sclerotiza-
Figs. 28-40. Euchromius species. — 28, distribution of E. ocelleus, European and North African distribution (distribution in
Mauritania, Mali, Niger, Tchad, the Central African Republic, Ethiopia and all other countries south of these are not depict-
ed, but see Schouten 1988); 29, distribution of E. ocelleus in North America; 30, distribution of E. ocelleus in the Oriental re-
gion; 31, distribution of E. ocelleus in Hawaii; 32, E. pulverosus, head, lateral aspect, scales removed; 33, idem, sclerite of ter-
gite VIII; 34, distribution of E. confusus and E. pulverosus, 35, E. confusus, head, lateral aspect, scales removed; 36, idem,
sclerite of tergite VIII; 37, distribution of E. vinculellus (distribution in Niger and Kenya are ommited); 38, E. anapiellus,
head, lateral aspect, scales removed; 39, idem, sclerite of tergite VIII; 40, distribution of E. anapiellus. Scale bar 0.5 mm to
figs. 32, 35-36, 38-39.
211]
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
tion, armed with small central projection, edges of
tergite VIII connected; ostium complex, large tooth-
shaped projection with parallel sides; ductus bursae
long, minutely grooved in upper part; ductus semi-
nalis narrow; bursa copulatrix oblong, no signa.
Biology. — Unknown. Collections from April,
May, June, July, August and September. Most speci-
mens in July and August, up to 2500 m altitude.
Distribution (fig. 34). — A Central-Asian species,
found in Tadzhikistan, Afghanistan and Iran.
Remarks. — See under E. pulverosus.
18. Euchromius vinculellus (Zeller, 1847)
(figs. 9, 37, 115-117, 166)
Crambus vinculellus Zeller, 1847: 760. Neotype: d (desig-
nated by Bleszynski (1960: 211)), GS 628 (zmHB) [not
examined].
Ommatopteryx corsicalis Hampson, 1919: 534. Holotype: 6,
‘Vizzavona Corsica 11.VI.1899 Wlsm 1910. 166’, GS
5647 (BMNH) [examined].
Ommatopteryx asbenicola Rothschild, 1920: 220. Holotype:
2, ‘Auderas, Asben 26 July 20 (A. Buchanan)’, GS 5675
(BMNH) [examined].
Eromene joiceyella Schmidt, 1934: 538. Holotype: 6,
‘42.27. Timmel S.E. Slopes Great Atlas Morocco
20.V.27 at light Talbot & Le Cerf. (BMNH) [examined].
Eromene bahrlutella Amsel, 1949: 236. Lectotype: d (desig-
nated by Bleszynski (1965a: 82)), ‘Sudende des Toten
Meeres 15.-27.3.33 Aigner leg. H. Amsel’, GS 320
(LNKD) [examined].
Euchromius vinculellus (Zeller, 1847) Schouten (1988: 26)
[redescription].
Material. — 57 4, 67 2. Afghanistan: Sarobi, 2 4,2 ?
(LNKD). Algeria: 70 km NE of Tamanrasset, 1 9 (BMNH); El
Kantara, 1 d, 1 2 (BMNH); Hammam-Meskoutine, 1 d
(BMNH); Lambese, 1 2 (BMNH); Marsa-ben-Mehidi, 1 à
(BMNH); Sebdou, 2 dd (BMNH); Sidi-bel-Abbes, 1
? (BMNH). Cyprus: Ayios Joannis, 1 9 (NHMW); Limasol, 1
® (nHMw). France: Ajaccio, 1 ® (NHMW); Vizzavona, 1 d
(BMNH). Georgia: Lagodekhi, 1 2 (zmas). Iran: 11 km NE
of Karevandar, 1 d, 1 2 (LNKD); 22 km N of Bandar-
Abbas, 3 6 (NHMw); 52 km S of Sirjan, 1 d, 1 2 (NHMW);
Ahwas, 1 d (BMNH); Bender Abbas-Sardabad, 1 d, 1 9
(NHRS); Bender Chahbahar, 1 ® (LNKD), 1 d (NHRS);
Bender Chahbahar-Iranshar, 1 9 (LNKD); Firouzabad, 1 &
(BMNH); Kazeroun-Bouchir, 1 ® (NHRS); Keredj, 1 2
(nHRs). Israel: Jericho, 1 & (LNKD), 1 2 (NHMw); South
end of Dead Sea, 1 5, 1 2 (LNKD). Italy: Arcu Neridu, 1 3
(LNKD); Bucherri, 1 d (RTAS); Can Gutluru Mannu, 1 9
(LNKD); Musei, 1 & (LNKD); Partenico, 1 9 (INER); Porto
Santoru, 1 9 (mer); Putifigari, 1 d, 1 2 (izu1); Sinecola, 1
d (LNKD); Teulada, 1 2 (LNKD). Jordan: Amman, 2 ®
(ZFMK); Zarga, 2 2 (LNKD). Kenya: South Horr, 1 ®
(LACM). Morocco: 15 km SW of Tazenakht, 1 2 (zMuc);
Barrage Cavagnac, 1 9? (MHNG); Chechaouen, 2 @ (NHMW),
1 d (zemk); Demnate, 1 2 (BMNH); Goundafa, 1 ©
(BMNH), 4 2 (NHMW); Ijoukak, 1 2 (MNHN); Ljourar, 1 9
(BMNH); Ketama, 1 ® (LNKD), 1 & (MNMs); Quirgane, 1 d
(LNKD), 1 d, 1 © (zrmk); Tangier, 1 9 (MNMs); Taroudant,
212
1 d (mMNHN); Timmel, 1 d (BMNH), 1 d (MNHN);
Tinerhir, 1 ® (zMuc); Zemmouro, 1 2 (BMNH). Niger:
Aouderas, 1 ® (BMNH). Nikhichevan: Ordubad, 1 @
(ZMAS). Oman: Ghubra, 1 2 (BMNH); Misfah, 1 d (BMNH);
Muscat, 1 d, 1 2 (BMNH); Wadi al Khawb, 1 2 (BMNH);
Wadi Bani Khalid, 2 &, 1 2 (BMNH); Wadi Fanjah, 1 9 (-
BMNH). Saoud Arabia: Buraiman, 1 2 (BMNH); Makakah, 1
? (NHMB); Sadiyah, 5 4,5 © (NHMB); Wadi Majarish, 2 &
(NHMB). Spain: Altea, 1 9 (zrMK); El Rompido, 1 ® (-
LNKD); Nerva, 2 ® (HULL); Periane, 1 9 (Grei), 1 6,2 9 (-
HULL); Ronquilla, 1 & (DERR), 3 d (GIEL), 1 (HULL), 5 d
(RTAS); San Pablo, 1 d (crer). Tunisia: 25 km SE of Ain
Draham, 1 d (zmuc); Bou-Hedma, 1 d (INER). Turkey:
Malatya, 1 d (BMNH). Yemen Arabic Republic: Suq as Sabi,
1 ® (RTAS).
Diagnosis. — Differs from all other species by its
straight single medial fascia in combination with six
or seven black terminal dots and having a yellow area
adjacent to the black terminal dots.
Biology. — Largely unknown. Flight-periods can be
split into three parts. Firstly south of the
Mediterranean Sea (Morocco, Algeria and Tunisia),
where this species has two generations and was caught
from March to September, with peaks in May-June
and August-September. Secondly the area north of
the Mediterranean Sea (Spain, France and Italy),
where there are two more or less continuous genera-
tions from April to October with peaks in May and
August-September. Thirdly, east of the
Mediterranean where specimens have been caught all
year round, with peaks in March-April-May and
September-October. Highest recorded altitude 1400
m in Morocco.
Distribution (fig. 37). — Spain, France, Italy,
Cyprus, Turkey, Georgia, Afghanistan, Iran, Oman,
Saudi Arabia, Jordan, Israel, Yemen Arabic Republic,
Kenya, Niger, Tunisia, Algeria and Morocco.
Remarks. — Bleszynski (1965a: 81) and Schouten
(1988: 26) incorrectly state the male with GS 628 Bl.
as the lectotype; this should read ‘neotype’.
19. Euchromius anapiellus (Zeller, 1847)
(figs. 38-40, 122, 171)
Crambus anapiellus Zeller, 1847: 757. Lectotype: 2 (desig-
nated by Bleszynski (1960: 211)), ‘Syracus Zell. coll.
1884’, GS 5655 (BMNH) [examined].
Eromene bellus var. minorella Chrétien: in Lhomme (1935-
1946: 84) [unavailable, manuscript name of Chrétien,
specimens in Paris Museum under minorella belong to E.
anapiellus).
Material. — 50 d, 57 ©. Algeria: Bone, 1 d, 1 ©?
(-MNHN); Djebel-Aurés, 1 d (BMNH); Guelt, 1 2 (MNHN);
Hamman R'irha, 2 d, 1 9 (BMNH); Lambese, 4 d, 1 9
(BMNH), 1 6 (NHMB); St. Charles, 2 d (MNHN); Qued
Hamidou, 1 d (BMNH); Tarfaia, 1 2 (MNHN). France:
Anduse, 1 2 (MNHN); Bize, 4 d, 4 2 (MNHN); Bonnieux, 1
® (rras), 1 2 (wozr); Durban-Sigean, 2 ® (zmuc); Feu-
du-Var, 1 & (RoBI); La Penne-s-L’Ouveze, 1 4,2 9 (RTAS),
4 2 (worr); Le Mans, 1 d (MHNG); Les Carbonnes de
Fleurs, 5km N. of Narbonne, 1 2 (Huis); Nimes, 1 dé
(MNHN); Pouzols, 1 d (Huts); Saignon, 1 9 (ROBI); Ste
Croix, 1 d (MNHN); Ste Guilhem, 1 9 (MNHN). Italy: Casel
Daccia, 1 d (iner); Sicilia, 1 ® (Mczc), 1 d,1 2 (NHMW);
Syracus, 2 d,6 $ (BMNH), 1 2 (LNKD), 4 d, 1 2 (NHMB),
76,5 2 (NHMW), 14,1 2 (NHRs), 16,1 2 (ZFMK), 1 9
(zsmc); Zappulla, 1 d (NHMw). Morocco: Azzou, 1 d
(BMNH); Meknes, 1 ¢ (BMNH); Rabat, 1 3 (BMNH); Sebou,
1 d (BMNH); Tangier, 3 6,8 ? (BMNH), 1 d,1 2 (ZFMK).
Spain: Jaca, 3 ® (BMNH); Korb, 1 2 (BMNH); San Antonia,
1 d (NHRs); San Roque, 1 9 (HULL); Tirade, 1 2 (HULL).
Tunisia: Ain Draham, 3 2 (BMNH); El Gounia 1 d (zsMc);
Sfax, 1 2 (BMNH).
Diagnosis. — Similar to E. bellus, but can be distin-
guished by the narrow yellow area adjacent to termi-
nal dots, which is broad and white in £. bellus. The
male of E. anapiellus differs from E. bellus in having a
cornutus which is curved less than 180° (figs. 122,
123). Differs in female genitalia from £. bellus in hav-
ing the sclerotization in ostium two times as long as
broad, three times as long as broad in E. bellus (figs.
IA)
External characters male, female (fig. 38). —
Wingspan 13-17 mm. Frons produced forward, with
point, creamy white, no ventral ridge; vertex creamy
white; labial palp two, sides white at base, becoming
brown, creamy white from above and below; maxil-
lary palp creamy white, brown ringed at base of last
segment, terminal part creamy white; antenna creamy
white to light grey, from about the middle inconspic-
uous darkly ringed. Thorax creamy white; patagia
creamy white with two broad longitudinal light
brown stripes; tegulae creamy white with dark patch
in the middle. Forewing, groundcolour white densely
suffused with ochreous to dark brown scales; medial
fascia double, inner one usually reduced, outer fascia
straight, running to one-third of the dorsum; subter-
minal line ochreous to dark brown, about midway
between terminal dots and termination of posterior
area; area adjacent to terminal dots narrow, yellow;
nine or eight black terminal dots, formula 2-2-3-2 or
2-2-3-1; fringes shiny, evenly grey-brown. Hindwing
light brown to dark grey, subterminal fascia present,
termen darkly bordered; fringes creamy white with
brownish line.
Tergite VIII (fig. 39). — Sclerite normally sclero-
tized, posterior part slightly larger than anterior part,
stalk stout.
Male genitalia (fig. 122). — Uncus normal, tapering
to sharp pointed tip; gnathos longer, with two dorsal
thorns, terminal part long; tegumen without appen-
dix angularis; sacculus swollen, swollen, partly armed
with thin elongated crest, processus of sacculus short,
blunt, dorsal processus of sacculus pointed, processus
basalis broad at base, tapering soon, bent inward, ven-
SCHOUTEN: Euchromius and Miyakea
tral terminal part minutely dentate, processus inferior
valvae at base of processus basalis, small, cucullus
broad, slightly bent upward at tip; juxta broad trian-
gular; vinculum rectangular; aedeagus small, one
large pointed cornutus.
Female genitalia (fig. 171). — Papillae anales nor-
mal; membrane of tergite VIII without clear sclerot-
izations; edges of tergite VIII not connected; ostium
simple, somewhat pointed, sclerotization two times as
long as broad, ductus bursae posterior part broad;
ductus seminalis narrow; bursa copulatrix elongated,
signa absent.
Biology. — Unknown. Flies after sunset (Zeller
1847). One generation, the first specimens were
caught in the first half of May. It continues through
June, July, August to the end of September.
Distribution (fig. 40). — A West-Mediterranean
species, occurring in Spain, France, Italy, Tunisia,
Algeria and Morocco.
Remarks. — Roesler (1983: 12) records Sardinia.
Records from Portugal (Mendes D’Azevedo 1904-
1905: 226, Vieilledent 1905: 201, Zerkowitz 1946:
145) and Egypt and Asia Minor (Bleszynski 1965a:
69) seem to be based on misidentified material.
Material under E. anapiellus in MzcP from Portugal
turned out to belong to E. gozmanyi.
20. Euchromius bellus (Hübner, 1796)
(figs. 9, 41-43, 123, 172)
Tinea bella Hübner, 1796: 29. Syntypes: ‘Hungary [lost].
Eromene bellalis Hübner, [1825]: 366 [lapsus calami].
Euchromius bellus (Hübner, 1796); Guenée 1845: 324.
Ommatopteryx bellus (Hübner, 1796); Kirby 1897: 274.
Ommatopteryx bellus var. minorella Chrétien in Lhomme
(1935-1946: 84) [name unavailable, manuscript name of
Chrétien, incorrect identification, the specimens belong
to E. anapiellus).
Material. — 253 6, 215 ®. Armenia, 1 d, 1 £ (teem).
Bulgaria: Bunkera, 1 d (RTAS); Ichtyman, 1 d (nHMw);
Kozuch, 2 ® (RTAS); Sistova, 2 d (INER), 1 d (sMTD), 9 6,
7 ® (zsmc). France: Abries, 1 d (MNHN); Aizenay, 1 9
(MNHN); Allauch La Clu, 1 2 (MNHN); Autun, 1 4,1 9
(MNHN); Berre, 1 d, 1 2 (MNHN); Borgo, 1 ® (MNHN);
Camargue, 1 ® (LNKD); Cannes, 1 d, 1 $ (BMNH);
Chatebaillon, 1 & (MNHN); Col de Homme Mort, 1 d (Ro-
BI); Corse, 1 2 (MNHN); Cravant, 2 2 (MNHN); Decoster, 1
? (MNHN); Lantosque, 1 d (MNHN); Digne, 1 d (BMNH), 2
d (INER), 1 2 (LNKD), 5 d, 5 2 (MNHN); Entrevaux, 2 d,
1 2 (BMNH); Florae, 1 2 (MNHN); Foret St. Baume, 9 d, 2
2 (woLF), 1 d (RTAS); Gemenos, 1 d (worr); La Banne, 1
3,1 2 (MNHN); La Baume, 1 6,2 ® (anse); La Voulte sur
Rhone, 6 d (MNHN); Le Mans, 1 d, 1 2 (mung); Le
Restugaud, 12 d, 5 ® (roBt); Le Rozier, 1 d (MNHN);
Montpellier, 1 d (MNHN); Nans, 1 d, 1 9 (ansp); Nievre,
2 4,2 2 (MNHN); Pelissanne, 2 d (MNHN); Ste André les
Alpes, 1 d (MNHN); St Barnabe, 1 2 (MNHN); Ste-Croix, 3
2 (MNHN); St. Guilhem le Desert, 1 2° (MNHN); Vaucluse,
213
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 41-51. Euchromius species. — 41, E. bellus, head, lateral aspect, scales removed; 42, idem, sclerite of tergite VIII; 43,
idem, distribution map (open circle: exact locality not found); 44, E. bleszynskiellus, head, lateral aspect, scales removed; 45,
idem, sclerite of tergite VIII; 46, idem, distribution map; 47, E. bleszynskii, head, lateral aspect, scales removed; 48, idem,
sclerite of tergite VIII; 49, idem, distribution map; 50, E. scobiolae, head, lateral aspect, scales removed; 51, idem, distribu-
tion map. Scale bar 0.5 mm to figs. 41-42, 44-45, 47-48, 50.
214
2 2 (MNHN); Volonne, 1 d, 1 2 (NHMB). Georgia: Gonia,
1 2 (zmas); Tbilisi, 1 d (zmas). Greece: Chalkidike, 4 9
(1zu1); Corfu, 1 d (NHMw); Kournas, 2 ® (LNKD);
Lithochoron, 1 d (LNKD); Niki, 1 & (Luca); Lindos, 2 9
(RMNH); Spili, 1 2 (LNKD); Vrises, 1 d (LNKD). Hungary:
no locality, 1 d (BMNH), 1 d (LNKD), 1 d (mezc), 1 d
(MHNG), 3 6,1 2 (NHMW), 6 6,2 2 (RMNH), 1 d (ZMUN),
2 3 (zsmc); Budafok, 2 d (BMNH). Iran: Keredj, 2 4, 1 9
(NHRS); Kermanshah, 2 d, 2 © (LNKD); Persepolis, 1 d, 1
® (LNKD); 100 km Mahabad-Sardascht, 2 2 (LNKD); 40 km
S. of Shahabad, 1 2 (LNKD); 90 km S. of Teheran, 1 3
(LNKD); 53 km S. of Zanjan, 1 d (LNKD). Iraq: Diana, 1 4
(BMNH); Mosul, 5 d, 2 2 (nHMw); Rowanduz, 1
(BMNH); Salahuddin, 1 ¢ (BMNH). Israel: Cesaree, 4
(MNHN); Haifa, 1 d (LNKD), 1 9 (zrmk); Tel Aviv, 1
(NHMW); Sedom, 1 d, 1 9 (BMNH). Italy: Albano, 1
(zsmc); Anagui, 2 d, 3 2 (MNHN); Bologna, 4 d, 2
(BMNH); Brummana, 1 6, 3 2 (BMNH); Campo Basso, 1
1 2 (BMNH); Fondi, 1 d, 1 9 (ner); Furbara, 5 6, 8
(INER); Maiella, 3 d, 1 2 (INER); Montagnana, 1
(MNsM); Monte Autore, 1 d (NHMB); Monte Vernita, 1 d
(casc), 1 2 (PRIN), 10 8,9 9 (RTAS), 1 6,1 2 (rmsa), 1
3, 1 2 (ZMAN); Mtgna Grande, 1 4, 3 2 (BMNH), 1 gd
(LNKD), 14 G, 8 ® (zsmc); Ovindoli, 1 &, 2 £ (BMNH);
Orvieto, 1 5,1 ® (INER); Prato, 1 d (NHMW); Ravone, 1 9
(zsmc); San Pietro, 4 6, 8 2 (MNsM); Sestola, 1 d, 1 9
(BMNH), 1 d (LNKD), 1 2 (NHMB); Spoleto, 1 4, 1 9
(NHMB); Syracus, 1 d (NHMw); Tivoli, 3 d, 4 2 (zsmc);
Toscane, 1 d (NHMB); Tradone, 1 d (zsmc). Jordan:
Amman, 1 d (zrMk); Dehbeen, 1 9 (LNKD); Zarga, 1 d
(LNKD). Lebanon: Beirut, 4 6, 1 2 (MNHN), 1 2 (NHMB), 1
6, 2 2 (NHMW), 1 d (ZMAN). Morocco: Ifrane, 1 9
(LNKD). Rumania: Bucuresti, 1 2 (BMNH), 1 2 (MNHN);
Mehadia, 1 9 (BMNH); Rimnicu Sarat, 1 d (MNHN);
Tecuci, 1 d (BMNH). Russia: Rubas, 1 d (zMas); Sarepta, 1
3, 1 2 (MNHN); Taganrog, 2 d, 2 2 (zMmas). Spain:
Andalusia, 1 4,1 © (curc); Burgos, 1 d (MNHN); Jaca, 12
d, 9 2 (BMNH); Murcia, 1 d (BMNH), 1 d (NHRS);
Tragacete, 1 & (Grrr). Syria: no locality, 1 2 (BMNH), 1 ©
(MHNG), 1 9 (RMNH). Tadzhikistan: Kurgan-Tjube, 1 ó
(NHMW). Turkey: Adana, 2 d (MNHN); Aksehir, 2 d, 1 9
(NHMW), 1 2 (zsmc); Alexandrette, 1 d (MNHN); Amassia,
1 4,3 $ (MNHN); Ankara, 2 d, 1 2 (MHNG), 2 6,19
(NHMW); Bursa, 1 d (BMNH), 2 6, 6 @ (iner), 1 ©
(NHMW); Diyarbakir, 1 d (BMNH), 1 d, 1 © (zEMk);
Gallipoli, 1 d, 1 2 (BMNH); Gümüshane, 1 & (zmuc); 7
km W. of Horasan, 1 d (Luca); Konya, 2 d, 6 9 (LUCA);
Malatya, 1 2 (BMNH); Marasch, 1 & (BMNH), 1 6,1 © (in-
ER), 1 4,3 2 (smtp), 1 6,5 ® (zsmc); Palandoken, 1 &
(Luca); Pontus, 1 d, 1 2 (BMNH); Smyrna, 1 2 (BMNH);
Tekirdech, 1 9 (Mars); Yenisehir, 1 d,2 2 (Mars); Yuksek
Dagh, 2 9 (zsmc); Zanapa, 5 d, 6 ® (mars). Ukraine:
Kara-dag, 1 d, 4 ® (zmas); Stevastopol, 1 d (zMas).
Yugoslavia: Drenovo, 2 2 (zsmc); Gruz, 1 2 (nHMw);
Istria, 1 6 (NHMw); Radobje, 1 d (nHMw); Ragusa, 1 d, 1
? (NHMW); Rovinj, 1 d (LNKD); Spalato, 2 d (NHMW).
Os +0 ® 40 +0 Os Os Os
Diagnosis. — Resembling E. anapiellus for differ-
ences see under diagnosis of that species.
External characters male, female (figs. 9, 41). —
Wingspan 14-19 mm. Frons produced forward with
small point or without, creamy white, usually with
darker centre, no ventral ridge; vertex creamy white
to light brown; labial palp two and a half, sides
SCHOUTEN: Euchromius and Miyakea
creamy white at base, becoming light brown to dark
grey-brown, creamy white from above and below;
maxillary palp creamy white to light brown, light
brown to dark grey at base of last segment; antenna
creamy white, inconspicuous greyish ringed. Thorax
creamy white to grey-brown; patagia creamy white
with two broad longitudinal light brown to grey-
brown stripes; tegulae creamy white with dark patch
in the middle. Forewing, groundcolour creamy white,
densely suffused with ochreous to dark brown scales,
anterior area yellow to brown-grey; medial fascia dou-
ble, arched to nearly straight, inner fascia reduced or
not, line between double fascia white, seldom silvery,
outer fascia running to one-third of the dorsum; sub-
terminal line ochreous to dark brown, closer to termi-
nation of posterior area than to terminal dots; area
adjacent to terminal dots broad, white; nine or eight
black terminal dots, formula 2-2-3-2 or 2-2-3-1;
fringes shiny, evenly greyish. Hindwing creamy white
to dark grey, subterminal fascia present, termen dark-
ly bordered; fringes creamy white with grey to brown-
ish line.
Tergite VIII (fig. 42). — Sclerite normally sclero-
tized, posterior part large, rectangular, anterior part
small.
Male genitalia (fig. 123). Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part long; tegumen without appen-
dix angularis; sacculus swollen, processus of sacculus
short, blunt, two dorsal processi, processus basalis
broad at base tapering soon, bent inward, ventral ter-
minal part minutely dentate, processus inferior valvae
small, at base of processus basalis, cucullus broad,
pointed tip bent upward; juxta broad triangular; vin-
culum normal; aedeagus large, one large curved cor-
nutus.
Female genitalia (fig. 172). — Papillae anales nor-
mal; membrane of tergite VIII without clear sclerot-
izations; edges of tergite VIII not clearly connected;
ostium simple, large rectangular sclerotization con-
tinued into ductus bursae, three times as long as
broad, ductus bursae posterior part very broad, sac-
shaped, partly wrinkled, anterior part more narrow,
ductus seminalis narrow, bursa copulatrix elongated,
very lightly sclerotized, signa absent.
Biology. — The single generation flies from May up
to the end of August. The flight reaches its peak in
July. In Israel a specimen has been caught on the 30th
of March. In Italy £. bellus has been caught up to
1400 m, in Turkey up to 2200 m and in Iran up to
2000 m. Hogenes (pers. comm.) caught the moths in
Italy at very hot and dry localities, with scarce herbal
vegetation with very few grasses. The larva is men-
tioned to live on dry leaves of Picris and Scabiosa, also
recorded to attack the roots and flower heads
(Milliere 1868: 219).
215
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Distribution (fig. 43). — This species is one of the
commonest of the genus. It occurs in Spain, France,
Italy, Yugoslavia, Hungary, Rumania, Bulgaria,
Greece, Turkey, Russia, Ukraine (Krim), Georgia,
Tadzhikistan, Iraq, Iran, Syria, Jordan, Lebanon,
Israel and Morocco. Bleszynski (1965A: 69) also
records Czechoslovakia.
Remarks. — Butler (1882: 42) mentions £. bellus
from Honolulu. This reference refers to Æ ocelleus.
21. Euchromius bleszynskiellus Popescu-Gorj, 1964
(figs. 44-46, 124, 173)
Euchromius bleszynskiellus Popescu-Gorj, 1964: 13.
Holotype: ®, ‘Periprava, pad. Letea, 29.VII.1963, leg.
Dr. A. Popescu-Gorj’, GS 12.331/637 (MGAB) [not ex-
amined]. Bleszynski (1969: 25) [synonymization of roxa-
nus].
Euchromius roxanus Bleszynski, 1965a: 73. Holotype: d,
‘Guberli’, GS 9432 (zmas) [examined].
Material.-6 d, 11 ?. Rumania: Periprava, 1 £ (LNKD);
Sulina, 1 2 (BMNH), 1 d (RMNH). Russia: Guberli, 1 d
(zmas); Sarepta, 1 d (zmas). Turkey: Dalyan, 2 2 (BMNH);
Kusadasi, 2 6, 4 2 (BMNH), 1 d, 2 2 (zemk). Ukraine:
Sem-Kolodezej, 1 2 (zsmc).
Diagnosis. — Differs from E. erum, E. locustus, E.
discopis, E. viettei, E. hampsoni and E. aris by the co-
nical frons with a small point. Can be distinguished
from E. donum in having labial palp three, two in E.
donum. Male genitalia of E. donum unknown. Female
genitalia of E. donum without signa, E. bleszynskiellus
with two signa.
External characters male, female (fig. 44). —
Wingspan 15-17 mm. Frons produced forward with
clear point, creamy white to light brown, no ventral
ridge; vertex creamy white to light brown; labial palp
three, sides creamy white at base, becoming brown,
creamy white from above and below; maxillary palp
creamy white to light brown; antenna creamy white,
greyish ringed. Thorax creamy white to brown; pata-
gia creamy white with two broad longitudinal light
brown stripes; tegulae creamy white with dark patch
in the middle. Forewing, groundcolour creamy white
densely suffused with ochreous to dark brown scales,
posterior area with up to three small dark brown
streaks; medial fascia single, straight or nearly so, run-
ning to one-third of the dorsum; subterminal line
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white to yellow; eight or nine black terminal
dots, formula 2-2-3-1 or 2-2-3-2; fringes shiny with
up to three brown lines. Hindwing creamy white to
light brown, subterminal fascia present, termen dark-
ly bordered; fringes creamy white with brownish line.
Tergite VIII (fig. 45). — Sclerite normally sclero-
tized, anterior part small, posterior part larger.
216
Male genitalia (fig. 124). — Uncus normal, tapering
to sharp pointed tip; gnathos longer, with two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus swollen, processus of sacculus
short, blunt, dorsal processus more or less straight,
processus basalis short, broad at base tapering soon,
processus inferior valvae small, cucullus relatively
stout, bent upward; juxta triangular; vinculum small;
aedeagus normal, one elongated group of small cor-
nuti.
Female genitalia (fig. 173). — Papillae anales nor-
mal; membrane of tergite VIII without sclerotiza-
tions, edges of tergite VIII connected; ostium small;
ductus bursae short, enlarged under ostium, halfway
with small sac-like projection; ductus seminalis nar-
row; bursa copulatrix oblong, two thorn-shaped sig-
na.
Biology. — Popescu-Gorj (1970) has caught bles-
zynskiellus in the dunes of the Donau river-delta. The
specimens were caught in May, June, July and
August.
Distribution (fig. 46). — Greece, Rumania, Turkey,
Russia and Ukraine.
Remarks. — The holotype of bleszynskiellus could
not be studied since the collections in Bucarest were
closed for research during my visit to the museum.
22. Euchromius bleszynskii Roesler, 1975
(figs. 47-49, 125, 174)
Euchromius bleszynskii Roesler, 1975: 233. Holotype: à,
‘Mongolia, Gobi Altaj aimak, Zachuj Gobi 10 km N von
Chatan chajrchan Gebirge, 1150 m, Exp. Dr. Z. Kaszab,
1966’. — ‘Nr.594, 27.VI.1966’, GS 6556 (HNHM) [exam-
ined].
Material. - 3 G, 13 2. Mongolia: Gobi Altaj aimak, 10
2 (BMNH), 2 d, 1 2 (HNHM), 2 2 (LNKD), 1 d (RTAS).
Diagnosis. — See under E. scobiolae.
External characters male, female (fig. 47). —
Wingspan 18-22 mm. Frons produced forward, with
clear point, brown, no ventral ridge; vertex creamy
white to light brown; labial palp two to two and a
half, sides white at base, becoming brown, creamy
white from above and below; maxillary palp brown,
terminal part creamy white; antenna light brown,
darker ringed. Thorax light to dark brown; patagia
light brown with two broad longitudinal dark brown
stripes; tegulae light brown, evenly mottled.
Forewing, groundcolour creamy white, densely suf-
fused with ochreous to dark brown scales, posterior
area usually with dark spot; medial fascia single,
slightly arched, running to one-third to one-fourth of
the dorsum; subterminal line brown, closer to termi-
nation of posterior area than to terminal dots; area
adjacent to terminal dots broad, white; six or seven
black terminal dots, very seldom an eighth vestigial
dot, formula 2-3-1 or 2-3-2, very seldom 1-2-3-1;
fringes shiny evenly grey-brown. Hindwing grey-
brown, subterminal fascia present, termen darkly bor-
dered; fringes white with brownish line.
Tergite VIII (fig. 48). — Sclerite normally sclero-
tized, anterior part small, rectangular, posterior part
larger, rounded, stalk normal.
Male genitalia (fig. 125). Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus very broad, rounded, dorsal
edge with pointed projection, processus of sacculus
short, processus basalis short, broad at base narrowing
soon, bent inward, processus inferior valvae incon-
spicuous, cucullus long, bent upward with anterior
fold; juxta broad triangular; vinculum normal sized,
pointed; aedeagus stout, most anterior part constrict-
ed.
Female genitalia (fig. 174). — Papillae anales large
rounded, hairs long, bent at top; membrane of tergite
VIII without sclerotizations, edges of tergite VIII
connected; ostium with rectangular projection; duc-
tus bursae short; ductus seminalis first part broad
then narrowing; bursa copulatrix rounded to oblong,
with inconspicuous more sclerotized part.
Biology. — Unknown. All specimens were caught in
June at 950-1200 m altitude.
Distribution (fig. 49). - Mongolia.
Remarks. — Roesler (1975) figures the female geni-
talia with a distinct signum. There is no real signum
present in the bursa copulatrix, only a faint more
strongly sclerotized area.
23. Euchromius scobiolae Bleszynski, 1965
(figs. 9, 50, 51, 126, 175)
Euchromius scobiolae Bleszynski, 1965a: 79. Holotype: à,
‘Kuschk’, (MGAB) [not examined].
Material.—1 8,4 2. Afghanistan: Kuschk, 1 d (BMNH).
Tadzhikistan: Kurgan-Tjube, 1 9 (BMNH). Turkmeniya:
Dort-Kuju, 1 2 (zmas); Peski, 1 ® (zmas); Sumbar, 1 9
(ZMAS).
Diagnosis. — Externally indistinguishable from £.
bleszynskii. Differs in male genitalia in lacking the
pointed projection of the dorsal edge of sacculus.
Differs in female genitalia in having the ductus semi-
nalis broad for a short length and the projection of
the ostium rounded, not rectangular.
External characters male, female (figs. 9, 50). —
Wingspan 16-21 mm. Frons produced forward with
clear point, creamy white to light brown, no ventral
ridge; vertex creamy white; labial palp two and a half,
sides creamy white at base, becoming brown, creamy
white from above and below; maxillary palp creamy
SCHOUTEN: Euchromius and Miyakea
white to brown, dark ringed at base of last segment;
antenna creamy white inconspicuous darkly ringed.
Thorax brown; patagia light brown with two broad
longitudinal brown stripes; tegulae brown, evenly
mottled. Forewing, groundcolour creamy white, very
densely suffused with ochreous to dark brown scales,
posterior area sometimes with dark spot; medial fascia
single, very faint, slightly arched, running to one-
third of the dorsum; subterminal line ochreous to
dark brown, closer to termination of posterior area
than to terminal dots; area adjacent to terminal dots
broad, white; seven or six terminal black dots, 2-3-2,
1-2-3-1 or 2-3-1; fringes shiny, grey-brown, no clear
lines. Hindwing grey-brown, subterminal fascia faint-
ly present, termen darkly bordered; fringes creamy
white with brownish line.
Tergite VIII not seen due to bad dissection.
Male genitalia (fig. 126). Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus very broad, rounded, processus
of sacculus stout, pointed upward, several stout spines
at base, processus basalis short, broad at base, narrow-
ing soon, bent inward, processus inferior valvae
prominent, cucullus slender; juxta triangular; vincu-
lum normal; aedeagus stout, most anterior part re-
stricted.
Female genitalia (fig. 175). — Papillae anales large
rounded, hairs long, bent at top; membrane of tergite
VIII without sclerotizations, edges of tergite VIII
connected; ostium with rounded projection; ductus
bursae short; ductus seminalis starting broad, narrow-
ing soon; bursa copulatrix oblong, roundish more
sclerotized patch in centre, no signa.
Biology. — Unknown. Specimens caught in April.
Distribution (fig. 51). — Turkmeniya,
Tadzhikistan and Afghanistan.
Remarks. — The holotype could not be studied
since the collections in Bucarest were closed for re-
search during my visit to the museum.
24. Euchromius superbellus (Zeller, 1849)
(figs. 52-54, 127, 176)
Crambus superbellus Zeller, 1849: 314. Holotype: à , ‘super-
bella m n 1/1 81°, GS 2835 (BMNH) [examined].
Eromene wockeella Zeller, 1863: 53. Lectotype: d (designat-
ed by Bleszynski (1961: 458)), ‘44, GS 5654 (BMNH)
[examined].
Ommatopteryx cypriusella Amsel, 1958: 51. Holotype: à,
‘6.4.47 Kyrenia’. (LNKD) [examined].
Material. — 62 d, 52 2. Albania: no locality, 1 9
(NHNW). Austria: no locality, 2 d (mczc). Armenia:
Yerewan, 2 2 (NHMW). Bulgaria: Blagoevaradski, 2 ®
(MNHN), 2 6,1 ® (RTAS); Kuzuch, 1 d (RTAS); Nessebar, 1
3 (BMNH). France: Corsica, 1 & (LNKD). Greece: 5 km S of
217
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Ge
==
Figs. 52-63. Euchromius species. — 52, E. superbellus, head, lateral aspect, scales removed; 53, idem, sclerite of tergite VIII; 54,
idem, distribution map (open circles: exact locality not found); 55, E. keredjellus, head, lateral aspect, scales removed; 56,
idem, sclerite of tergite VIII; 57, idem, distribution map; 58, E. malekalis, head, lateral aspect, scales removed; 59, idem, scler-
ite of tergite VIII; 60, idem, distribution map; 61, E. mouchai, head, lateral aspect, scales removed; 62, idem, sclerite of ter-
gite VIII; 63, idem, distribution map. Scale bar 0.5 mm to figs. 52-53, 55-56, 58-59, 61-62.
Monemvasia, 1 2 (RMNH); Heraklion, 1 d, 1 2 (Luca); d, (Luca); Fano, 3 6,5 2 (BMNH), 1 d (LNKD), 1 d,1 ®
Iria, 1 © (rras); Kallonis, 2 &, 2 2 (HULL); Kassandra, 1 d (zsmc); Ficuzza, 1 2 (BMNH); Livorno, 1 6, 1 2 (BMNH), 1
(LNKD); Katerini, 1 9 (ZFMK); Kyrenia, 1 2 (BMNH), 2 d 3 (NER), 16,1 2 (LNKD), 3 6,4 2 (NHMW), 1 2 (zsMC);
(LNKD); Meteora, 1 d, 1 @ (RTAS), 4 6, 4 @ (rzui). Pescara, 1 9 (mer); Sicily, 3 d (BMNH), 1 d (LNKD);
Hungary: no locality, 3 4, 1 9 (NHMw). Italy: Catania, 1 Toscane, 4 d, 1 ® (BMNH), 1 d (NHMW). Spain: Malaga, 1
218
d (BMNH), 1 d (NHMB). Turkey: Aksaray, 1 ® (zmuc);
Akshehir, 1 4 (zsmc); Alanya, 1 d (Luca); Aralik 10 km N,
2 2 (zmuc); Brusa, 1 (NHMW); Hazar Gölü, 1 ® (zmuc);
Kagizman 14 km E, 1 2 (zmuc); Konya, 1 d (Luca);
Kusadasi, 1 3 (BMNH). Ukraine: Jalta, 2 2 (zmas); Kara-
dag, 8 d, 1 2 (zmas). Yugoslavia: Rabac, 2 d (LNKD);
Saraj, 1 d (Luca); Skopje, 1 2 (rzur); Spalato, 4 d, 2 9
(NHMW); Split, 1 d,6 9 (mung), 1 2 (RTAS); Sucurac, 2 9
(NHMW).
Diagnosis. — E. superbellus externally resembles E.
gozmanyi, E. rayatellus, E. malekalis, E. keredjellus and
E. mouchai. Differs from the latter three in lacking
the one or two brown lines on the fringes of the fore-
wing. Differs in male genitalia from E. gozmanyi and
E. rayatellus by the dorsal thorns on the gnathos being
elongated. Differs in female genitalia in having the
ostium armed with a thumb-shaped projection.
External characters male, female (fig. 52). —
Wingspan 13-15 mm. Frons produced forward with
point, creamy white to brown, no ventral ridge; ver-
tex creamy white; labial palp two, sides white at base,
becoming brown to dark lead-grey, brown from
above, creamy white from below; maxillary palp
creamy white to dark brown-grey, terminal part light-
er; antenna creamy white to grey, from about halfway
darkly ringed. Thorax creamy white to brown-grey;
patagia creamy white with two broad longitudinal
light brown to lead-grey stripes; tegulae creamy white
with dark patch in the middle. Forewing, groundco-
lour creamy white densely suffused with ochreous
brown to dark grey scales; medial fascia single,
straight or nearly so, running to one-third of the dor-
sum; subterminal line ochreous brown, about mid-
way between terminal dots and termination of poste-
rior area, sometimes closer to termination of posterior
area than to terminal dots; area adjacent to terminal
dots white; seven or six, seldom eight, black terminal
dots, formula 2-3-2, 2-3-1 or seldom 1-2-3-2; fringes
highly shiny evenly silver-grey. Hindwing light to
dark grey, subterminal fascia present, termen darkly
bordered; fringes creamy white to dark grey with one
or two greyish lines. Specimens with very lightly
coloured wings do occur, mainly from Italy.
Tergite VIII (fig. 53). — Sclerite normally sclero-
tized, posterior part large, rectangular, stalk and ante-
rior part not distinguishable.
Male genitalia (fig. 127). Uncus bent, tapering to
sharp pointed tip, usually armed with thin dorsal
crest; gnathos longer, two elongated dorsal thorns,
terminal part long; tegumen without appendix angu-
laris; sacculus broad, dorsal edge armed with dentate
Projection, processus of sacculus absent, processus
basalis short, sharply bent inward, processus inferior
valvae small, cucullus normal, bent upward; juxta tri-
angular; vinculum normal; aedeagus normal, two
patches of small cornuti.
SCHOUTEN: Euchromius and Miyakea
Female genitalia (fig. 176). — Papillae anales nor-
mal; membrane of tergite VIII with rounded sclerot-
ization near papillae anales, second sclerotization
thumb-shaped, projecting near ostium, edges of ter-
gite VIII not connected; ostium simple; ductus bur-
sae normal; ductus seminalis starting broad, narrow-
ing soon; bursa copulatrix oblong, one inconspicuous
roundish area slightly more sclerotized.
Biology. — Found in gras- and weed-areas directly
bordering the beach, but specimens are also caught
up to 1700 m. There is one more or less continuous
generation from April to September. The flight
reaches its peak in June and July.
Distribution (fig. 54). — Confined to the northern
part of the Mediterranean, and recorded from Spain,
France, Italy, Austria, Yugoslavia, Hungary, Albania,
Greece, Bulgaria, Turkey, Ukraine and Armenia.
Possibly also in Portugal (Zerkowitz 1946, but see
Monteiro 1982). Records of Leech (1901), Caradja
(1925), Caradja & Meyrick (1933) from various
parts of China refer to Metaeuchromius species or E.
raddeellus. Records for Mozambique (Joannis 1927,
Vari & Kroon 1986)) refer to E. klimeschi.
Remarks. — The paralectotypes of wockeella Zeller,
1863 are lost, none have been found in BMNH or any
other museum.
25. Euchromius keredjellus (Amsel, 1949)
(figs. 55-57, 128, 177)
Eromene keredjellus Amsel, 1949: 233. Holotype: à , ‘Iran,
Elbursgebirge Keredj 4.6.1936 leg Brandt, GS 318
(LNKD) [examined].
Material. — 6 6, 6 9. Afghanistan: Herat, 2 ? (BMNH), 2
d (LNKD); Kasi, 1 & (BMNH). Iran: 90 km S. of Teheran, 1
d (LNKD), 1 (RTAS); Keredj, 2 d (LNKD), 2 2 (NHRS);
Shiraz, 1 2 (NHRS).
Diagnosis. — E. keredjellus externally resembles £.
malekalis, E. mouchai, E. superbellus, E. rayatellus, E.
gozmanyi. Differs from the latter three in having one
or two brown lines on the fringes of the forewing near
the apex. £. keredjellus differs in male genitalia in hav-
ing two cornuti in the aedeagus in combination with
a short terminal part of the gnathos and a clear pro-
cessus of the sacculus. In female genitalia £. keredjel-
lus can be separated in having the edge of tergite VIII
connected and forming a little knot-shaped sclerot-
ization at point of connection.
External characters male, female (fig. 55). —
Wingspan 13-21 mm. Frons produced forward, with
clear point, creamy white, no ventral ridge; vertex
creamy white; labial palp two and a half, sides white
to creamy white at base, becoming light brown to
brown, creamy white from above, light brown from
below; maxillary palp creamy white to light brown,
219
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
brown ringed at base of last segment; antenna creamy
white, in some grey-brown ringed. Thorax light
brown to brown; patagia creamy white with two
broad longitudinal light brown stripes; tegulae
creamy white with, sometimes very faint, dark patch
in the middle. Forewing, groundcolour white, dense-
ly suffused with ochreous to dark brown scales, ante-
rior area with faint streaks, posterior area sometimes
with up to three dark brown streaks; medial fascia
single slightly arched to nearly straight, running to
one-third of the dorsum; subterminal line ochreous
to dark brown, closer to termination of posterior area
than to terminal dots; area adjacent to terminal dots
broad, white; seven black terminal dots, formula 2-3-
2; fringes shiny, with up to three ochreous brown
lines, most clearly at apex. Hindwing creamy white to
light brown, subterminal fascia present or not, termen
darkly bordered: fringes pure white no brownish line.
Tergite VIII (fig. 56). — Sternite lightly sclerotized,
only posterior part partly visible.
Male genitalia (fig. 128). — Uncus normal, slightly
bent; gnathos longer, two dorsal thorns, terminal part
short; tegumen without appendix angularis; sacculus
broad, dorsal edge with strongly sclerotized ridge,
processus of sacculus stout, short, straight, processus
basalis small, bent inward, processus inferior valvae
normal, cucullus large, slightly bent upward; juxta tri-
angular; vinculum rectangular; aedeagus stout, anteri-
or one group of small cornuti, posterior second group
of large, circa five, cornuti.
Female genitalia (fig. 177). — Papillae anales large,
hairs long; membrane of tergite VIII with inconspic-
uous band-shaped sclerotization, edges of tergite VIII
connected, knot-like at point of connection; ostium
slightly toad-stool shaped; ductus bursae short, broad
under ostium; ductus seminalis starting broad nar-
rowing soon; bursa copulatrix oblong, inconspicuous
patch of stronger sclerotization visible.
Biology. — Collected in April, May and June.
Occurs up to 1600 m.
Distribution (fig. 57). — Only found in Iran and
Afghanistan so far.
Remarks. — Bleszynski (1965a) mentions NHRS as
the museum where the holotype is deposited, but it
was found in LNKD.
26. Euchromius malekalis (Amsel, 1961)
(figs. 58-60, 129, 179)
Eromene malekalis Amsel, 1961: 330. Holotype: à, ‘Iran,
Baloutchistan Bender Tchahbahar-Iranchar. Tahte
Malek 750 m leg. Brandt, 1938’, GS 3559 A (NHRs) [ex-
amined].
Material. - 2 d, 3 2. Iran: Tahte Malek, 1 d (NHRS).
Jordan: Azraq ed Druz, 1 2 (BMNH); Wadi er Retem, 1 d,
2 2 (BMNH).
220
Diagnosis. — E. malekalis externally resembles E.
mouchai, E. keredjellus, E. superbellus, E. rayatellus and
E. gozmanyi. Differs from the later three in having
fringes of the forewing with one or two brown lines,
most clearly at the apex. Differs in male genitalia
from E. mouchai and E. kereajellus in lacking a group
of cornuti in the aedeagus. Differs in female genitalia
by the bean-shaped ostium.
External characters male, female (fig. 58). —
Wingspan 20-21 mm. Frons produced forward, with
clear point, creamy white to light brown, no ventral
ridge; vertex creamy white; labial palp three, white at
base, becoming light brown, white from above and
below; maxillary palp brown, terminal part white; an-
tenna creamy white, from about the middle greyly
ringed. Thorax white to light brown; patagia white
with two broad longitudinal brown stripes; tegulae
with dark patch in the middle. Forewing, groundco-
lour white densely suffused with ochreous to brown
scales; medial fascia single, slightly arched to straight,
running to one-fourth of the dorsum; subterminal
line ochreous brown, about midway between termi-
nal dots and termination of posterior area; area adja-
cent to terminal dots white; seven black terminal
dots, formula 2-3-2; fringes with one to two brown
lines at apex. Hindwing creamy white to brown, sub-
terminal fascia present, termen darkly bordered;
fringes white with brownish line.
Tergite VIII (fig. 59). — Sclerite normally sclero-
tized, pattern only visible at posterior part.
Male genitalia (fig. 129). — Uncus normal, tapering
to pointed tip; gnathos longer, dorsal thorns normal,
terminal part short; tegumen without appendix angu-
laris; sacculus very broad, rounded, processus of sac-
culus short, bent upward, strongly sclerotized, proces-
sus basalis short, bent inward, processus inferior
valvae at base of processus basalis, cucullus normal
sized, slightly bent upward; juxta triangular, two
small dorsal projections; vinculum normal sized; ae-
deagus normal sized, no cornuti.
Female genitalia (fig. 179). — Papillae anales large
rounded, hairs long, bent at top; membrane of tergite
VIII without sclerotizations; ostium bean-shaped,
strongly sclerotized; ductus bursae lightly sclerotized;
ductus seminalis starting broad narrowing soon; bur-
sa copulatrix broad, with inconspicuous more sclero-
tized patch.
Biology. - Unknown. Caught in April and May. In
Iran at 750 m altitude.
Distribution (fig. 60). — Jordan and Iran.
27. Euchromius mouchai Bleszynski, 1961
(figs. 61-63, 130, 178)
Euchromius mouchai Bleszynski, 1961: 464. Holotype: à,
‘1865 Rossia m., Sarepta, Chr.’, GS 1142 [presumably
lost].
Material. — 4 d, 4 ®. France: Corsika, 1 2 (MNHN).
Georgia: Lagodechi, 1 ® (zmas). Italy: Syracus, 1 d
(BMNH). Russia: Sarepta, 3 d, 1 ® (BMNH). Turkey: Konya,
1 2 (MNHN).
Diagnosis. — Externally resembling £. keredjellus,
E. malekalis, E. superbellus, E. rayatellus and E. goz-
manyi, but differs from the latter three in having the
fringes of the forewing with one or two brown lines,
most clearly at the apex; male genitalia have three
groups of cornuti in the aedeagus. Differs in female
genitalia in having a lip-shaped ostium in combina-
tion with free edges of tergite VIII.
External characters male, female (fig. 61). —
Wingspan 14-16 mm. Frons sharply produced for-
ward with clear point, creamy white, brown in mid-
dle, no ventral ridge; vertex creamy white; labial palp
two and a half to three, sides white at base, becoming
brown, white from above, creamy white from below;
maxillary palp brown terminal part creamy white; an-
tenna creamy white to light brown-grey, from about
middle inconspicuously dark ringed. Thorax creamy
white; patagia creamy white with two broad longitu-
dinal brown stripes; tegulae creamy white with dark
patch in the middle. Forewing, groundcolour creamy
white densely suffused with ochreous to dark brown
scales, anterior area with inconspicuous brown spot
near medial fascia; medial fascia single, straight or
nearly so, running to one-third of the dorsum; subter-
minal line ochreous to dark brown, sometimes incon-
spicuous, closer to termination of posterior area than
to terminal dots; area adjacent to terminal dots broad,
white; seven black terminal dots, formula 2-3-2;
fringes shiny creamy white with two brown lines.
Hindwing grey-brown, subterminal fascia present,
termen darkly bordered; fringes white with brownish
line.
Tergite VIII (fig. 62). — Sclerite normally sclero-
tized, posterior part rectangular larger than anterior
part, stalk normal.
Male genitalia (fig. 130). — Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus broad, dorsal edge with trans-
parent narrow ridge, processus of sacculus very short,
processus basalis short, broad at base, narrowing
soon, processus inferior valvae small, cucullus very
long, slender; juxta broad triangular; vinculum rec-
tangular; aedeagus stout, three groups of cornuti.
Female genitalia (fig. 178). — Papillae anales nor-
mal; membrane of tergite VIII with two horizontal
sclerotizations enclosing a square part of membrane,
edges of tergite VIII not connected; ostium simple,
no clear projections; ductus bursae short, broad near
ostium; ductus seminalis starting broad at base, nar-
rowing soon, bursa copulatrix oblong, inconspicuous
SCHOUTEN: Euchromius and Miyakea
more strongly sclerotized area in lower part.
Biology. — Unknown. Very rare species.
Distribution (fig. 63). — France, Italy, Turkey,
Georgia and Russia.
Remarks. — The holotype seems to be lost. Dr.
Razowski informed me, that there is no Crambinae
material in ZMPA.
28. Euchromius nivalis (Caradja, 1937)
(figs. 64-66, 131, 181)
Eromene nivalıs Caradja in: Caradja & Meyrick, 1937: 151.
Lectotype: d (designated by Bleszynski (1965a: 83)), ‘Li-
Kiang Provinz Nord Yuennan 20.VII.1935 Hone’ GS
1741 (MGAB) [not examined].
Material. — 4 d, 1 9. China: Lijiang [Li-kiang],
(BMNH), 1 d (RTAS), 1 d, 1 2 (zFMKk); Yunnan,
(BMNH).
3
3
mi pa
Diagnosis. — E. nivalis can be distinguished from
all other species by it pure white groundcolour of the
wings.
External characters male, female (fig. 64). —
Wingspan 15-19 mm. Frons produced forward with
inconspicuous point, pure white, no ventral ridge;
vertex pure white; labial palp two and a half, sides yel-
low-white at base, becoming yellow, then yellow-
grey, white from above, yellow from below; maxillary
palp pure white, yellow at base; antenna creamy
white, darkly ringed. Thorax pure white; patagia pure
white with two broad longitudinal yellow stripes; teg-
ulae pure white. Forewing, groundcolour pure white,
anterior area not suffused with dark scales, posterior
area densely suffused with ochreous to dark brown
scales, posterior area with distinct dark brown spot
near medial fascia; medial fascia single, bent outward,
running to half to one-third of the dorsum; subtermi-
nal line ochreous to dark brown, inconspicuous to ab-
sent, closer to termination of posterior area than to
terminal dots; area adjacent to terminal dots broad,
white; seven or six black terminal dots, formula 2-3-2
or 2-3-1; fringes shiny, evenly yellow-grey. Hindwing
grey-white, subterminal fascia absent, termen darkly
bordered; fringes white, no clear line.
Tergite VIII (fig. 65). — Sclerite normally sclero-
tized, no pattern visible.
Male genitalia (fig. 131). — Uncus slender, tapering
to sharp pointed tip; gnathos longer, with two large
dorsal thorns, terminal part short; tegumen without
appendix angularis; sacculus normal, processus of sac-
culus absent, processus basalis more or less free from
back of valvae, swollen, ending in small point, proces-
sus inferior valvae elongated, cucullus slender, dorsal
edges slightly more sclerotized; juxta triangular with
two dorsal projections; vinculum very long; aedeagus
long, slender, with set of short strongly bent cornuti,
221
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 64-75. Euchromius species. — 64, E. nivalis, head, lateral aspect, scales removed; 65, idem, sclerite of tergite VIII; 66,
idem, distribution map; 67, £. rayatellus, head, lateral aspect, scales removed; 68, idem, sclerite of tergite VIII; 69, idem, dis-
tribution map (open circle: exact locality not found); 70, £. gozmanyi, head, lateral aspect, scales removed; 71, idem, sclerite
of tergite VIII; 72, idem, distribution map; 73, E. gratiosellus, head, lateral aspect, scales removed; 74, idem, sclerite of tergite
VIII; 75, distribution map £. gratiosellus in the West Palaearctic. Scale bar 0.5 mm to figs. 64-65, 67-68, 70-71, 73-74.
second, more anteriorly located group of long cornu-
ti.
Female genitalia (fig. 181). — Papillae anales small;
membrane of tergite VIII without sclerotization,
edges of tergite VIII faintly connected; ostium sim-
ple, grooved; ductus bursae long; ductus seminalis
narrow; bursa copulatrix roundish, no signa.
222
Biology. - Unknown. Specimens were caught in
July and September at an altitude of 2800 to 3200 m.
Distribution (fig. 66). - Only known from the type
locality, China, Province of Yunnan.
Remarks. — The lectotype of nivalis could not be
studied since the collections in Bucarest were closed
for research during my visit to the museum.
29. Euchromius rayatellus (Amsel, 1949)
(figs. 9, 67-69, 132, 180)
Eromene rayatellus Amsel, 1949: 278. Holotype: d , ‘13.7.35
Rayat’, GS 12384 (BMNH) [examined].
Material. — 42 d, 55 ®. Afghanistan: Balkh, 1 ® (LNKD);
Derweshan, 1 & (BMNH); Polichomri, 1 2 (BMNH), 1 d, 1
2 (LNKD). Bulgaria: Burgas, 1 d (NHMW), Nessebar, 6 d, 8
Q (BMNH), 1 6,1 ® (zut), 1 2 (LNKD), 1 d (RTAS), 1 9
(ZFMK). Cyprus: Kolossi, 2 d, 2 ® (BMNH); Lanarca, 1 9
(BMNH). Greece: Chalkidike, 1 4,3 2 (zur); Drepanon, 1
3 (PRIN); Ialyssos, 1 2 (zmuc); Lithokhoran, 1 d (LNKD);
Malia, 1 & (zmuc); Neapolis, 5 4, 1 (NHMW); Phaistos,
1 d (ZMAN); Pitsidia, 1 d (LNKD); Platamon, 1 9 (ZFMKk);
Rhodos, 1 ® (zrmk). Iran: Kendavan, 1 ® (INER). Iraq:
Rayat, 1 d (BMNH), 1 d (LNKD). Israel: Hulch Swamp, 2 ?
(BMNH); Daphne Settlement, 1 d (BMNH); Deganya, 1 d, 1
2 (corp). Italy: Fondi, 1 & (INER); Fano, 1 © (BMNH);
Livorno, 1 d (RMNH). Spain: Murcia, 1 9 (NHRS). Syria: no
locality, 1 2 (MNHN). Turkey: Adana, 1 9 (INER), 1 4,3 2
(MNHN); Aksehir, 1 9 (Luca); Amanus, 1 ® (zsmc);
Ankara, 1 £ (Luca); Diyadin 10 km NE, 1 4,1 9 (zmuc);
Diyarbakir, 1 9 (zrMmk); Ergani, 1 2 (MHNG); Karatas, 1 d,
1 © (LNKD), 1 d (MNHN); Kusadasi, 3 6, 1 2 (BMNH), 5 9
(zeMK); Mut, 1 d,2 2 (NHMw); Sarkikaagas, 1 9 (INER);
Tuz Gölü N., 1 9 (zmuc). Turkmeniya: Kara-Kala, 2 9
(isNB). Ukraine: Kara-dag, 4 d (zMas). Yugoslavia: Crna
Gora, 1 ® (LNKD).
Diagnosis. — £. rayatellus externally resembles £.
gozmanyi, E. superbellus, E. malekalis, E. keredjellus
and E. mouchai. Differs from the latter three in lack-
ing one or two brown lines in the fringes of the fore-
wing. E. rayatellus differs in male genitalia from E.
gozmanyi and E. superbellus by its aedeagus with three
groups of cornuti. Differs in female genitalia by the
large shield-like lamella antevaginalis which is split in
the middle.
External characters male, female (figs. 9, 67). —
Wingspan 13-16 mm. Frons produced forward, with
clear point, creamy white to brown, dorsal part dark-
er, no ventral ridge; vertex creamy white to light
brown; labial palp two and a half, sides creamy white
at base, becoming brown, creamy white from above
and below; maxillary palp creamy white, sometimes
brown ringed at base of last segment; antenna creamy
white. Thorax creamy white to brown; patagia
creamy white with two broad longitudinal light
brown stripes; tegulae creamy white with dark patch
in the middle. Forewing, groundcolour creamy white,
densely suffused with brown to dark brown scales;
medial fascia single, nearly straight, running to one-
third of the dorsum; subterminal line ochreous
brown, about midway between terminal dots and ter-
mination of posterior area, seldom closer to termina-
tion of posterior area; area adjacent to terminal dots
white, seldom broad; seven black terminal dots, for-
mula 2-3-2, in some specimens an additional eighth
dot occurs, formula 1-2-3-2; fringes shiny, evenly
ScHOUTEN: Euchromius and Miyakea
grey-brown. Hindwing light brown to grey-brown,
subterminal fascia present, termen darkly bordered;
fringes creamy white with brownish line.
Tergite VIII (fig. 68). — Sclerite normal sclerotized,
posterior part convex, stalk and anterior part very faint.
Male genitalia (fig. 132). — Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus normal, processus of sacculus
long, straight, processus basalis small, broad at base,
tapering soon, strongly bent inward, processus inferi-
or valvae slightly projecting, cucullus slender, bent
upward; juxta broad triangular; vinculum rectangu-
lar; aedeagus small, stout, three groups of cornuti.
Female genitalia (fig. 180). — Papillae anales nor-
mal; membrane of tergite VIII with rounded punc-
tate sclerotization, edges of tergite VIII connected; os-
tium small, lamella antevaginalis shield-like, split in
the middle; ductus bursae short; ductus seminalis
starting broad narrowing soon; bursa copulatrix ob-
long, with inconspicuous more sclerotized part in the
centre.
Biology. — There is one long flight-period, starting
in April, May and continuing until mid September,
peak in July, August. This species ranges in altitude
from sea-level up to 3000 m.
Distribution (fig. 69). — Spain, Italy, Yugoslavia,
Greece, Bulgaria, Turkey, Ukraine, Turkmeniya,
Afghanistan, Iran, Iraq, Syria and Israel.
30. Euchromius gozmanyi Bleszynski, 1961
(figs. 70-72, 133, 182)
Euchromius gozmanyi Bleszynski, 1961: 462. Holotype: d,
‘Hispania Korb Chiclana 1912. IV.V°, GS 1204 (HNHM)
[examined]. Glaser, 1975: 50 [description of female].
Material. — 36 8,46 2. Algeria: Aflou, 1 © (BMNH); Aön
Fezza, 1 d (BMNH); Biskra, 1 6 (BMNH); El Tarf, 1 ©
(MNHN); Foret de Tenira, 1 & (BMNH); Hassi-Babah, 1 d
(INER), 1 (MNHN); Marsa-Ben-Mehidi, 2 4, 2 2 (BMNH);
Sebdou, 1 9 (BMNH); Sidi-bel-Abbes, 2 2 (BMNH); St.
Charles, 1 2 (MNHN). France: Bormes, 1 2 (MNHN); Corse,
Tour de Vignate, 1 ® (1zur); Durban-Sigean, 1 d (zMuc);
Ile du Levant, 1 d (MNHN); L’Almanarres, 2 d (MNHN); Les
Carbannes de Fleury, 1 d (murs); Petit Bagnes, 3 d, 1 9
(MNHN); Port La Nouvelle, 2 d (MNHN). Italy: Licata, 6 d
(BMNH); Musei, 1 2 (LNKD); Sailletta, 1 d (IMMER).
Morocco: Tangier, 3 2 (BMNH), 1 ® (LNKD), 2 2 (MNHN),
5 ® (ZFMK); Taourirt, 1 £ (BMNH); Thami, 1 2 (MNHN);
no locality, 4 2 (BMNH). Portugal: Coimbra, 2 d (MNHN);
Setubal, 1 d (MNHN). Spain: Aljarque, 1 9 (zmuc); Cadiz,
2 4,29 (cieL), 1 ® (prın),2 6, 5 £ (RTAS); Chiclana, 1
3 (BMNH); El Rampido, 1 & (LNKD); La Escala, 1 2
(LNKD); Penalba, 1 4 (zmuc); Periane, 1 & (GIEL); Puerto
de Sta. Maria, 5 6, 9 2 (Hutt); Rosas, 2 2 (LNKD); San
Lucar, 2 2? (BMNH); Trebujena, 2 2 (BMNH). Tunisia: Ain
Draham, 1 d (BMNH); Cap Bon, 1 d (MNHN); Tozeur, 1 9
(INER).
223
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Diagnosis. — E. gozmanyi resembles E. superbellus,
E. rayatellus, E. mouchai, E. keredjellus and E. malek-
alis. Can be distinguished from the latter three species
in lacking brown lines on the fringes of the forewing.
Differs from above mentioned species in male geni-
talia in having one large cornutus. In female genitalia
E. gozmanyi is distinguished by a sleeve-like structure
above the ductus seminalis.
External characters male, female (fig. 70). —
Wingspan 14-21 mm. Frons sharply produced for-
ward, with clear point, creamy white to light brown,
no ventral ridge; vertex creamy white; labial palp two
and a half, sides white to creamy white at base, be-
coming light brown to brown from above and below;
maxillary palp light brown, terminal part creamy
white; antenna creamy white, from about the middle
inconspicuous darkly ringed. Thorax creamy white to
light brown; patagia creamy white to brown with two
broad longitudinal brown stripes; tegulae creamy
white to light brown with dark patch in middle.
Forewing, groundcolour white densely suffused with
ochreous to dark brown scales, anterior area some-
times with inconspicuous brown spot near medial fas-
cia; medial fascia single, straight or nearly so, running
to one-third to one-fourth of the dorsum; subtermi-
nal line ochreous brown, faint, about midway
between terminal dots and termination of posterior
area; area adjacent to terminal dots white; seven or six
black terminal dots, formula 2-3-2 or 2-3-1; fringes
highly shiny, evenly grey to brown-grey. Hindwing
creamy white to grey, subterminal fascia present or
not, termen darkly bordered; fringes creamy white
with brownish line.
Tergite VIII (fig. 71). — Sclerite normally sclero-
tized, posterior part rectangular, stalk broad, anterior
part inconspicuous.
Male genitalia (fig. 133). — Uncus normal, tapering
to sharply pointed tip; gnathos longer, two dorsal
thorns normal sized, terminal part short; tegumen
without appendix angularis; sacculus very broad, pro-
cessus of sacculus short but free, strongly sclerotized,
processus basalis short, sharply bent inward, proces-
sus inferior valvae small, at base of processus basalis,
cucullus relatively broad, bent upward; juxta normal;
aedeagus normal sized, slender, tapering anterior of
anellus connection, one ridged cornutus.
Female genitalia (fig. 182). — Papillae anales nor-
mal; membrane of tergite VIII with strongly sclero-
tized horizontal patch near papillae anales, second
patch less clear, rounded, more towards the ostium,
edges of tergite VIII connected; ostium simple,
rounded; ductus bursae long with clear sleeve-like
structure above ductus seminalis; ductus seminalis
starting broad, narrowing soon; bursa copulatrix
elongated with two, not clear, sclerotized patches in
the middle.
224
Biology. — This species seems to prefer the shores of
salt-lakes and is reported to live on tidal shores. The
caterpillar is said to survive temporary floods by the
sea and live on Salicornia spec. (Luquet, pers.
comm.). There are two, more or less continuous, gen-
erations from half April up to half October. The
peaks in the flight-periods are June-July and
September.
Distribution (fig. 72). — A West-Mediterranean
species, occurring in Portugal, Spain, France, Italy,
Tunisia, Algeria and Morocco.
Remarks. — From Biskra (Algeria) 1 d is known
with eight and nine terminal dots on its forewing, GS
BM 17612, BMNH. Upon checking, the genital slide
number of the holotype proved to be 1204 instead of
1075 (Bleszynski 1961).
31. Euchromius gratiosellus (Caradja, 1910)
(figs. 73-76, 134, 170)
Eromene ramburiella gratiosellus Caradja, 1910: 116.
Lectotype: d (designated by Bleszynski (1965a: 76)),
‘Lob-Noor, 1909, B.H.’, (MGAB) [not examined].
Euchromius ramburiellus, Bleszynski (nec Duponchel),
(1960: 204).
Euchromius gartheellus Derra, 1985: 237. Holotype: à,
“Türkei, Prov. Ankara Tuz Gölu 900 m, Leg. G.Derra
3.8.1984’, GS 2284 (DERR) [not examined]. Ganev &
Hacker (1986: 329) [hew localities]. Derra (1987: 31)
[synonymization with gratiosellus].
Euchromius siuxellus Ganev & Hacker, 1986a: 82.
Holotype: 4, Türkei, Prov. Ankara, 39°10’N/33°20’E,
Tuz Gölu N-Ufer, 900 m, 3.8.1984’, GS 911 Ganev,
Sofia [not examined]. Hacker (1986: 62) [synonymiza-
tion with gartheellus Derra, 1985].
Material. — 85 d, 67 ®. Algeria: El Mesrane, 1 9
(LNKD); Hassi Bahbah, 1 2 (BMNH); 3 9 (INER); 1 4,5 9
(nHMw). Armenia: Chiva Urgene, 1 ® (BMNH). France:
Corsika, 1 d (MNHN). Iran: 90 Km S of Teheran, 1 6, 1 9
(LNKD); Shaku, 1 ? (BMNH). Italy: Livorno, 1 d (NHMW).
Kazakhstan: Emba, 2 & (NHRS); Uralsk, 1 4,3 2 (MNHN);
Uzun-Agach, 2 ® (zmas). Kirgiziya: Naryn, 30 km E., 1 d
(zmas). Mongolia: 10 km NNE of Dalanzadgad, 3 4,6 9
(BMNH); Talaiin Bulag, 2 d (BMNH). Russia: Katunda, 10
km W., 1 d (zmas); Sarepta, 2 2 (BMNH), 1 (INER), 4 d
(NHMW), 1 9 (ZFMK); Troikoe, 2 2 (RTAS). Spain: Alcaniz,
1 ® (GtEL); Dolores, 1 &,1 2 (Gr), 1 3,1 £ (RTAS), 1 d
(rMsa); El Ronquillo, 1 ® (HuLL); Penalba, 1 d (zmuc);
Puerto de Sta. Maria, 1 9 (HULL); Puerto Lambreras, 1 &
(PRIN). Tadzhikistan: Dusti, 1 & (RTAS). Turkey: 20 km
NO of Konya, 2 ® (LNKD), 5 d (NHMw); 40 km SW of
Elazig; Aksehir, 1 d, 1 9 (NHMw); Ankara, 4 d, 1 ?
(BMNH), 1 6, 3 2 (mung), 1 (NHMw); Cay, 10 d,4 9
(MAES); Diyadin 10 km NE, 1 d,2 2 (zmuc); Eregli, 2 à,
1 2 (zsmc); Karapinar, 2 d (LNKD), 2 d,2 2 (NHMw);
Kelkit, 4 6,4 2 (BMNH); Konya, 1 d, 1 2 (LNKD); Tuz
Golu, 6 d,2 2 (BMNH), 2 gd (casc);3 6,3 $ (DERR), 2 d,
1 2 (RTAS), 1 d (rMsA); Tuz Golu N.shore, 2 d, (LNKD), 3
d (NHMW), 2 6,2 © (RTAS), 1 d, 2 2 (zmuc), 4 6,3 2
(zsmc). Ukraine: Karadag, 1 & (zmas).
Diagnosis. — Externally undistinguishable from £.
jaxartellus and E. ramburiellus. On the average E. gra-
tiosellus is slightly larger and lighter coloured. E. gra-
tiosellus differs from E. superbellus, E. rayatellus and E.
gozmanyi in having nine or eight black terminal dots
whereas the former species almost always have six or
seven dots. In male genitalia Æ. gratiosellus differs
from E. jaxartellus and E. ramburiellus in the having
sacculus less broad and processus of sacculus more
straight and slender. Processus inferior valvae of E.
gratiosellus is rounded, elongated in E. ramburiellus.
In female genitalia the ostium of E. gratiosellus is
broadly rounded with parallel sides, longer and point-
ed in £. ramburiellus, broader and dilated in £. jaxar-
tellus. Spined sack-like terminal part of ductus bursae
much smaller in £. gratiosellus than in E. jaxartellus
and £. ramburiellus.
External characters male, female (fig. 73). —
Wingspan 17-23 mm. Frons produced forward with
point, creamy white to light brown, no ventral ridge;
vertex creamy white to light brown; labial palp two
and a half, sides white to creamy white at base becom-
ing light brown to brown, white from above and be-
low; maxillary palp creamy white, dark ringed at base
of last segment, terminal part creamy white; antenna
creamy white, darkly ringed, sometimes with dorsal
grey line. Thorax creamy white to light brown; pata-
gia creamy white, no clear broad iongitudinal brown
stripes; tegulae creamy white to light brown, evenly
mottled. Forewing, groundcolour white, densely suf-
fused with grey to dark brown scales, anterior area
more or less evenly coloured, posterior area with a yel-
lowish to dark brown spot in the middle; medial fas-
cia single, gently arched to nearly straight, running to
one-third of the dorsum; subterminal line ochreous
brown, sometimes only faintly visible, about midway
between terminal dots and termination of posterior
area; area adjacent to terminal dots white; eight or
nine black terminal dots, formula 2-2-3-1 or 2-2-3-2;
fringes shiny creamy white, three or four brown lines.
Hindwing creamy white to grey, subterminal fascia
present; termen darkly bordered, fringes white with
or without a brownish line.
Tergite VIII (fig. 74). — Sclerite normally sclero-
tized, posterior part triangular, stalk short inconspic-
uous, anterior part narrow, very faint.
Male genitalia (fig. 134). — Uncus normal, tapering
to sharply pointed tip; gnathos longer, two dorsal
thorns normal sized, terminal part short; tegumen
without appendix angularis; sacculus narrow, proces-
sus of sacculus slender, slightly bent upward, not or
just slightly overlapping cucullus (watch out for dis-
tortions caused by the coverslip), strongly sclerotized,
processus basalis short, broad at base narrowing soon,
pointing inward, strongly sclerotized, processus infe-
rior valvae large, distinctly projecting, rounded, cu-
ScHOUTEN: Euchromius and Miyakea
cullus very slender more or less bent upward; juxta
normal; vinculum normal; aedeagus short, broad,
three groups of cornuti.
Female genitalia (fig. 170). — Papillae anales nor-
mal; membrane of tergite VIII without sclerotiza-
tions, edges of tergite VIII connected; ostium thumb-
shaped, broadly rounded, with parallel sides; ductus
bursae under ostium enlarged armed with many small
spines. very small sac-like pouch connected to en-
larged part of ductus bursae; ductus seminalis narrow;
bursa copulatrix oblong with two thorn-shaped signa.
Biology. — Several times recorded from salt deserts.
Caught in April up to October at an altitude of up to
1450 m.
Distribution (figs. 75, 76). — Algeria, Spain,
France, Italy, Turkey, Iran, Ukraine, Russia,
Armenia, Kazakhstan, Kirgiziya, Tadzhikistan and
Mongolia.
Remarks. — The holotype of gratiosellus could not
be studied since the collections in Bucarest were
closed for research during my visit to the museum.
32. Euchromius jaxartellus (Erschoff, 1874)
(figs. 76-79, 135, 183)
Eromene jaxartella Erschoff, 1874: 82. Holotype: 2, no la-
bel, GS 7602 (zmas) [examined].
Material. - 45 d, 60 ©. Afghanistan: 22 km E of Kabul,
1 2 (LNKD); Bashgul, 1 © (LNKD); Kabul, 1 2 (LNKD);
Kandahar, 1 ® (zsmc); Kuschk, 1 2 (BMNH); Pol-i-
Charchi, 1 9 (LNKD), 1 9 (RTAS); Sarobi, 1 6 (LNKD).
Armenia: Chiva Urgene, 1 d, 1 2 (BMNH). Iran: 95 km N
of Kermanschah, 1 9 (LNKD); Keredj, 1 d (LNKD), 1 @
(NHRS); Shaku, 1 2 (numw). Kazakhstan: Aulie Ata, 1 &
(BMNH); Dzambulskaya, 35 d, 36 ® (zmas). Kirgiziya:
Naryn, 1 2 (zmas). Mongolia: Bayan Hongor, oase Echin
gol [exact locality not found], 1 4 (BMNH). Pakistan: Gilgit,
4 2 (zsmc); Hunza-Nagar, 1 9 (zsmc). Tadzhikistan:
Kurgan-Tjube, 1 d (NHMw). Turkey: Aralik 10 km NW, 2
9 (zmuc); Diyadin 10 km NE, 2 d (zmuc); Kagizman 14
km E, 2 d, 1 2 (zmuc). Turkmeniya: Kara-Kala, 1 9
(BMNH), 1 6,2 ® (1sNB), 1 2 (RTAS).
Diagnosis. — £. jaxartellus externally resembles £.
ramburiellus, E. gratiosellus, E. confusus, E. pulverosus,
E. zagulajevi. Differs from latter three species in hav-
ing up to three dark lines on the fringes of the fore-
wing. E. jaxartellus differs in male genitalia from £.
confusus and E. pulverosus in having three cornuti in
the aedeagus, differs from £. gratiosellus and E. ram-
buriellus in the processus of sacculus, at the most,
reaching one-third of the cucullus and the processus
is not longer than the breadth of cucullus. Differs in
female genitalia in having the projection of the os-
tium broad triangular.
External characters male, female. (fig. 77). —
Wingspan 14-17 mm. Frons produced forward with
225
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
@ gratiosellus
B jaxartellus
Figs. 76-84. Euchromius species. — 76, distribution map of E. gratiosellus and E. jaxartellus, 77, E. jaxartellus, head, lateral as-
pect, scales removed; 78, sclerite of tergite VIII; 79, distribution map of E. jaxartellus in the West Palaearctic; 80, E. rambu-
riellus, head, lateral aspect, scales removed; 81, idem, sclerite of tergite VIII; 82, idem, distribution map; 83, £. zagulajevi,
head, lateral aspect, scales removed; 84, idem, distribution map. Scale bar 0.5 mm to figs. 77-78, 80-81, 83.
minute point, creamy white, dorsal slightly darker, no
ventral ridge; vertex creamy white; labial palp two
and a half, sides creamy white at base, becoming
brown, creamy white from above and below; maxil-
lary palp creamy white, brown at base of last segment;
antenna creamy white, from about the middle greyish
226
ringed. Thorax creamy white to light brown; patagia
creamy white with two broad longitudinal brown
stripes; tegulae creamy white, evenly mottled.
Forewing, groundcolour creamy white, densely suf-
fused with ochreous to dark brown scales, posterior
area sometimes with small dark brown streaks; medi-
al fascia single, arched to nearly straight, running to
one-third of the dorsum; subterminal line brown,
about midway between terminal dots and termina-
tion of posterior area; area adjacent to terminal dots
white to yellowish; nine or eight black terminal dots,
formula 2-2-3-2 or 2-2-3-1; fringes slightly shiny
with up to four brown lines. Hindwing creamy white
to brown-grey, subterminal fascia present, termen
darkly bordered; fringes creamy white with brownish
line.
Tergite VIII (fig. 78). — Sclerite normal sclerotized,
anterior part small, very faint, posterior part large,
rectangular.
Male genitalia (fig. 135). — Uncus normal, tapering
to sharp pointed tip; gnathos longer, two dorsal
thorns, terminal part short; tegumen without appen-
dix angularis; sacculus narrow, processus of sacculus
short, bent upward, processus basalis short bent
downward, processus inferior valvae large, elongated,
cucullus normal, bent upward; juxta triangular; vin-
culum rectangular; aedeagus short, swollen, three
groups of cornuti.
Female genitalia (fig. 183). — Papillae anales nor-
mal; membrane of tergite VIII without sclerotiza-
tions, edges of tergite VIII connected; ostium with
broad equal-sided triangular projection, top narrow,
not broad rounded; ductus bursae under ostium en-
larged, armed with many spines, small sac-like appen-
dix connected to enlarged part of ductus bursae; duc-
tus seminalis narrow; bursa copulatrix rounded,
armed with two thorn-shaped signa.
Biology. — Unknown. One flight-period from
April up to half September, peak in July and August.
Caught up to 2200 m altitude.
Distribution (figs. 76, 79). — Armenia, Turkme-
niya, Tadzhikistan, Kirgiziya, Kazakhstan, Iran,
Afghanistan, Pakistan and Mongolia.
Remarks. — Ganev & Hacker’s (1984: 238) record
from Turkey is very doubtful and may concern
Euchromius gratiosellus or Euchromius ramburiellus.
Mariani’s (1938: 160) records from Sicily may refer
to E. gratiosellus or E. ramburiellus.
33. Euchromius ramburiellus (Duponchel, 1836)
(figs. 80-82, 136, 185)
Crambus ramburiellus Duponchel, 1836: 83. Holotype: ?,
‘Crambus ramburiellus, GS 3778 (MNHN) [examined].
Crambus zonellus Zeller, 1847: 758. Holotype: 9, [Italien,
Sicilien, type lost].
Eromene ramburiellus var. luteella Caradja, 1910: 116.
Lectotype: d (designated by Bleszynski (1965a: 74)),
‘Biskra 8.VI.07, Chrétien’, (MGAB) [not examined].
Eromene islamella Amsel, 1949: 235. Lectotype: 4 (desig-
nated by Bleszynski (1965a: 74)), ‘Iran, Fars, 1937
Strasse Kazeroun-Bouchir Tchouroum 1000 m. leg.
Brandt 17.IIT, GS 319 (LNKD) [examined].
SCHOUTEN: Euchromius and Miyakea
Material. — 145 8, 150 9. Afghanistan: Polichomri, 1 d
(LNKD). Algeria: Aflou, 1 ? (BMNH); Aön Sefra, 1 4,3 9
(BMNH); Beni Ounif, 1 d, 2 2 (MHNG), 1 d (NHMB);
Biskra, 7 & (BMNH), 1 2 (zsmc); Bou-Saada Faroult, 1 9
(BMNH); El Golea, 1 d, 2 2 (INER), 3 d,2 2 (MNHN), 1 d
(zsmc); El Kantara, 6 d, 6 2 (BMNH); El Outaya, 1 9
(BMNH); Ghardaöa, 2 d (BMNH); Hammam-es-Salahin, 1 9
(BMNH); Hassi Babak, 1 d (nHMw); Hassi Bahbah, 1 9
(INER), 1 & (nHMw); Laghouat, 3 d, 1 ? (MNHN); Oued
Nssa, 1 d (BMNH); Sebdou, 1 d, 3 ® (BMNH); Sidi-bel-
Abes, 1 d,3 2 (BMNH); Sidi-Okba, 1 ? (BMNH). Cyprus:
Kolossi, 2 d, 1 2 (BMNH); Limasol, 2 2 (BMNH). Egypt:
Alexandria, 1 d (MNHN); Aswan, 1 d, 1 2 (BMNH); Caöro,
2 2 (NHMW); Khamissa, 2 d, 2 2 (BMNH); Siwa, 6 d, 12 ®
(BMNH). France: Camargue, 1 ® (LNKD), 1 2 (ZFMKk);
Digne, 2 d, 2 ® (BMNH); Ile du Levant, 1 d (MNHN);
Minervois, 1 d (Huis); Oraison, 2 (LNKD); Sorgues, 1 9
(LNKD); Volonne, 6 4, 2 ® (tsNB). Iran: Kazeroun-Bouchir
Tchouroum, 1 d (LNkD), 2 4,2 ® (NHRs); Shadegan, 1 d
(LNKD); Tchouroum, 1 £ (NHRS). Iraq: Amarah, 3 d, 5 9
(BMNH); Bagdad, 5 d, 4 9 (BMNH), 1 d, 1 2 (LNKD);
Karradah, 1 d (BMNH). Israel: Haöfa, 5 4,4 2 (numw); En
Gedi, 1 4,1 2 (BMNH). Italy: Livorno, 1 2 (BMNH), 2 d,
1 2 (NHMW). Jordan: Azraq ed Druz, 2 4, 5 © (BMNH);
Dead Sea, 1 2 (LNKD), 1 9 (zemk); Fuhes, 1 6, 1 9
(LNKD); Qa el Umari, 1 2 (BMNH); Wadi er Retem, 1 d, 2
? (BMNH). Libya: Sidi Mesri (locality not found), 1 4,1 9
(BMNH). Morocco: Asni, 1 ® (LNKD); Erfoud area, 9 4,7 9
(zmuc); Ksar es Souk, 1 & (LNKD); Tafilalt, 1 2 (MNHN);
Tansikht, 1 ® (LNKD). Russia: Sarepta, 3 d (BMNH). Saudi
Arabia: 150-600 km SSW of Riad, 1 2 (LNKD). Spain:
Albarracin, 1 & (INER); Alhama de Murcia, 1 d (ZFMK);
Huelva, 1 2 (BMNH); Jandia, 1 9 (zrMk); Orgiva, 1 9
(BMNH); Malaga, 1 © (Gier); Maspalomas, 1 d, 1 9
(LNKD), 1 6,5 2 (nHMw); Mazagon, 1 d (LNKD); Motril,
1 2 (zemx); Murcia, 1 & (GIEL); Periana, 1 & (DERR), 2 6,
2 2 (cieL), 2 6,1 & (RTAS); Rio de Baza, 1 d,2 2 (zmuc);
Ronquillo, 1 ® (GEL); San Roque, 1 ® (cIEL);
Valdeltormo, 1 6, 1 2 (GIEL); Zaragosa, 1 2 (MNHN);
Zujar, 3 d (rei), 1 2 (PRIN), 1 d (TMsa), 2 d (RTAS).
Tunisia: 65 km NW of Tozeur, 2 & (zmuc); Maknassy, 1 9
(INER); Nefta, 1 & (INER), 1 2 (zmuc); Sfax, 10 4,15 £
(BMNH), 16 d, 7 ® (HNHM); Tozeur, 1 2 (INER), 2 £
(MNHN). Turkey: Alanya, 1 ® (Luca); Kusadasi, 1 d
(Luca). Yugoslavia: Sucurac, 1 9 (BMNH), 1 d (NHMW).
Diagnosis. — See under £. gratiosellus.
External characters male, female (fig. 80). —
Wingspan 16-20 mm. Frons produced forward with
point, creamy white to light brown, no ventral ridge;
vertex creamy white; labial palp two, sides white at
base, becoming brown, white to creamy white from
above and below; maxillary palp creamy white, brown
ringed at base of last segment, terminal part white;
antenna creamy white, from about the middle darkly
ringed. Thorax creamy white to brown; patagia
creamy white to brown-grey; tegulae creamy white to
brown-grey, evenly mottled. Forewing, groundcolour
creamy white densely suffused with ochreous to dark
brown scales, usually with dark brown spot in middle
of posterior area; medial fascia single, gently arched or
nearly straight, running to one-third of the dorsum;
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
subterminal line ochreous to dark brown, about mid-
way between terminal dots and termination of poste-
rior area; area adjacent to terminal dots white; nine or
eight black terminal dots, formula 2-2-3-2 or 2-2-3-
l; fringes shiny, creamy white with two or three
brown lines. Hindwing creamy white to grey-brown,
subterminal fascia present or not, termen darkly bor-
dered; fringes creamy white with brownish line.
Tergite VIII (fig. 81). — Sternite normally sclero-
tized, posterior part rounded, stalk normal, anterior
part narrow.
Male genitalia (fig. 136). — Uncus normal, tapering
to sharply pointed tip; gnathos longer, two dorsal
thorns normal sized, terminal part short; tegumen
without appendix angularis; sacculus broad, proces-
sus of sacculus stout, clearly bent upward overlapping
cucullus, strongly sclerotized, processus basalis short,
sharply bent inward, gradually tapering, strongly scle-
rotized, processus inferior valvae large, flat, elongated,
cucullus slender, bent upward; juxta normal; vincu-
lum normal; aedeagus short, relatively broad, three or
four groups of cornuti.
Female genitalia (fig. 185). — Papillae anales small;
membrane of tergite VIII with very faint sclerotiza-
tion above ostium, edges of tergite VIII connected;
ostium thumb-shaped, more or less pointed; ductus
bursae enlarged and armed with many cornuti near
ostium, small sac-like pouch at middle of ductus bur-
sae; ductus seminalis narrow; bursa copulatrix round-
ed to oblong, two thorn-shaped signa.
Biology. — Lhomme (1935-1946) mentions the
dead leaves of Cirsium spec., Hieracium spec. and
Cichorium spec. (all Compositae) at the base of the
plants as the food for the larvae. The larvae can be
found in silk tunnels until October.
Flight-periods: south of the Mediterranean Sea,
Canary Islands eastward including Egypt from
February to early November (peaks April-May-June
and August); north of the Mediterranean Sea, Spain
eastward including Turkey from March till the end of
September (peak July to September); and the area east
of the Mediterranean Sea, Israel and eastwards the
specimens were caught from March up the first week
of July and in September up to the first week of
November, with peaks in March, April and May.
This species only occurs at lower altitudes (highest
record Morocco 1200 m).
Distribution (fig. 82). — Spain, France, Italy,
Yugoslavia, Turkey, Cyprus, Russia, Afghanistan,
Iran, Iraq, Jordan, Israel, Egypt, Libya, Tunisia,
Algeria and Morocco. Also recorded from Portugal
(Monteiro 1972: 13) and Bulgaria (Ganev 1985:
172)
Remarks. — The lectotype of Euchromius islamellus
(Amsel, 1949) is not in NHRS as stated by Bleszynski
(1965a: 74), but in LNKD.
228
34. Euchromius zagulajevi Bleszynski, 1965
(figs. 83, 84, 184)
Euchromius zagulajevi Bleszynski, 1965a: 80. Holotype, 9,
‘Kafirnigan, 31-VII.’, GS 3983 (zmas) [examined].
Material. — 1 9. Tadzhikistan: Kafirnigan, 1 ® (zmas).
Diagnosis. — See under Euchromius pulverosus.
External characters female (fig. 83). — Wingspan
17 mm. Frons slightly produced forward with very
minute point, creamy white, no ventral ridge; vertex
creamy white; labial palp two, sides creamy white at
base, becoming brown, creamy white from above and
below; maxillary palp creamy white, dark brown
ringed at base of last segment; antenna creamy white,
from about halfway inconspicuous dark ringed.
Thorax creamy white; patagia creamy white with two
broad longitudinal light brown stripes; tegulae
creamy white to light brown, evenly mottled.
Forewing, groundcolour creamy white densely suf-
fused with ochreous to dark brown scales; medial fas-
cia single, straight or nearly so, running to one-third
of the dorsum; subterminal line ochreous to dark
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white, eight black terminal dots, formula 2-2-3-
1, fringes white at base then one broad ochreous
brown line. Hindwing creamy white, subterminal fas-
cia faintly present, termen darkly bordered; fringes
creamy white with faint brownish line.
Female genitalia (fig. 184). — Papillae anales nor-
mal; membrane of tergite VIII without sclerotiza-
tions; tergite VIII with inconspicuous collar, edges
connected; ostium simple, ductus bursae strongly
sclerotized, especially at edge in middle part, ductus
seminalis narrow, bursa copulatrix oblong, drop-
shaped, no signa.
Biology. — Unknown. The unique holotype was
caught at the end of July.
Distribution (fig. 84). — Tadzhikistan.
Remarks. — This species is only known from the fe-
male holotype.
35. Euchromius donum Schouten, 1988
(fig. 186)
Euchromius donum Schouten, 1988: 28. Holotype: 9,
‘Haro-Ali, Gurra 6. April 01. (C. V. Erlanger).’ GS
12111 (BMNH) [examined].
Material. — 1 9.
Diagnosis. — Differs from all species by its single
medial fascia in combination with nine or eight black
terminal dots, the bursa copulatrix without a signum
and the ostium simple tongue-shaped.
Distribution. — Ethiopia.
36. Euchromius sudanellus Bleszynski, 1965
(figs. 85-87, 137, 187)
Euchromius sudanellus Bleszynski, 1965a: 79. Holotype: à,
‘Sudan sept. or. Port Sudan Rhoz Arbaat 23.V1.1962 leg
R. Remane’, GS 1308 (zsmc) [examined].
Material. — 1 d, 1 9. Sudan: Port Sudan, 1 d, 1 9
(ZSMC).
Diagnosis. — Externally almost indistinguishable
from E. cambridgei see under that species.
External characters male, female (fig. 85). —
Wingspan 13 mm. Frons produced forward with
small point, creamy white, no ventral ridge; vertex
creamy white; labial palp two and a half, sides white
to creamy white at base, becoming brown to dark
lead-grey, white from above and below; maxillary
palp white to creamy white, dark brown ringed at
base of last segment, terminal part white; antenna
creamy white, from about the middle inconspicuous
darkly ringed. Thorax creamy white to light brown;
patagia creamy white with two broad longitudinal
brown stripes; tegulae creamy white with dark patch
in the middle. Forewing, groundcolour white densely
suffused with ochreous to dark brown scales, anterior
area with dark brown spot of various size near medial
fascia, posterior area with dark brown spot in middle;
medial fascia single, clearly angled under costa, not
broadening at inner margin, running to one-third of
the dorsum; subterminal line ochreous brown, about
midway between terminal dots and termination of
posterior area; area adjacent to terminal dots white to
yellow; eight or nine black terminal dots, formula 2-
2-3-1 or 2-2-3-2; fringes shiny, creamy white with
three brown lines. Hindwing light brown, subtermi-
nal fascia present, termen darkly bordered; fringes
creamy white with brownish line.
Tergite VIII (fig. 86). — Sclerite strongly scleroti-
zed, posterior part small, rectangular, stalk long, slen-
der, anterior part clearly visible, narrow, rectangular.
Male genitalia (fig. 137). — Uncus rather broad,
clearly bent, big dorsal thorn at two-fifth of pointed
tip; gnathos longer, broad dorsal thorn elongated, ter-
minal part short; tegumen with very small appendix
angularis; sacculus narrow, processus of sacculus slen-
der reaching to about the end of cucullus, processus
basalis broad at base tapering soon, bent inward,
strongly sclerotized, processus inferior valvae slightly
pointing inward, cucullus very slender, nearly
straight; juxta broad; vinculum with small basal pro-
jections; aedeagus small, tapering ventral and dorsally
posterior of anellus connection, one patch of minute
cornuti.
SCHOUTEN: Euchromius and Miyakea
Female genitalia (fig. 187). — Papillae anales nor-
mal; membrane of tergite VIII with broad triangular
sclerotized patch; ostium rounded at top, heavily scle-
rotized; ductus bursae lightly sclerotized, widening
proximal; ductus seminalis narrow; bursa copulatrix
large, roundish, small signum without, larger signum
with median ridge.
Biology. — Unknown. The two specimens were
caught at the end of June.
Distribution (fig. 87). — One of the rarest species of
Euchromius, only known from North East Sudan.
Remarks. — In his original description Bleszynski
(1965a: 79) mingled the external characters of the
male and female of E. sudanellus and E. subcambrid-
gei concerning the medial fascia. Only in E. subcam-
bridgei the medial fascia is dilating at the inner mar-
gin and not in E. sudanellus as stated by Bleszynski.
Bleszynski (1965b: 1-4) depicts the female genitalia
of E. sudanellus with the legends of £. subcambrigei
and vice versa.
37. Euchromius subcambridgei Bleszynski, 1965
(figs. 87-89, 138, 188)
Euchromius subcambridgei Bleszynski, 1965a: 78. Holotype:
3, ‘Sudan Ed Damer Hudeiba 29.V.1962 leg
R.Remane’, GS 1315 (zsmc) [examined].
Material. — 14 8, 20 9. Cape Verde: St. Vincent, 1 d
(BMNH). Sudan: Hudeiba, 1 $ (BMNH), 2 G, 12 (zsMc).
Tunisia: Sfax, 8 d, 12 2 (BMNH), 1 ® (curc), 2 d,4 9
(RTAS), 1 d, 1 9 (ZEMK).
Diagnosis. — Externally very similar to E. cambrid-
gei and E. labellum. Differs from these species in hav-
ing the medial fascia not sharply bent, and the angle
in the middle of the forewing and the fascia broaden-
ing at the inner margin.
External characters male, female (fig. 88). —
Wingspan 14-15 mm. Frons produced forward,
rounded without point, creamy white, darker in cen-
tre, no ventral ridge; vertex creamy white; labial palp
two to two and a half, sides white at base, becoming
light brown, creamy white from above and below;
maxillary palp light brown, dark ringed at base of last
segment, terminal part white; antenna creamy white,
from about the middle inconspicuous darkly ringed.
Thorax creamy white to light brown; patagia creamy
white with two broad longitudinal brown stripes; teg-
ulae creamy white with dark patch in the middle.
Forewing, groundcolour white densely suffused with
ochreous to dark brown scales, anterior area with dark
brown basal patch; medial fascia single, clearly angled
in middle of wing, broadening at inner margin, run-
ning to one-third of the dorsum; subterminal line
ochreous brown, about midway between terminal
dots and termination of posterior area; area adjacent
229
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
subcambridgei
87
sudanellus
86
ey
= x
N nenn
DEE ——
in 89
LE
===
91
one = LS
@ FA
2
\
92
Figs. 85-92. Euchromius species. — 85, E. sudanellus, head, lateral aspect, scales removed; 86, idem, sclerite of tergite VIII; 87,
distribution map of E. sudanellus and E. subcambridgei, 88, E. subcambridgei, head, lateral aspect, scales removed; 89, idem,
sclerite of tergite VIII; 90, E. cambridgei, head, lateral aspect, scales removed; 91, idem, sclerite of tergite VIII; 92, idem, dis-
tribution map. Scale bar 0.5 mm to figs. 85-86, 88-89, 90-91.
to terminal dots white to yellow; six or seven black dered; fringes white with brownish line.
terminal dots, formula 2-3-1 or 2-3-2; fringes shiny, Tergite VII (fig. 89). — Sclerite clearly visible, pos-
evenly yellowish. Hindwing creamy white to brown- terior part more or less square, anterior part smaller,
grey, subterminal fascia present, termen darkly bor- with two rounded caudal pointing tips, stalk normal.
230
Male genitalia (fig. 138). — Uncus slender, slightly
bent, tapering to sharp pointed tip; gnathos longer,
slender, dorsal thorns normal sized, terminal part
long, lightly sclerotized; tegumen with clearly visible
appendix angularis; sacculus normal, processus of sac-
culus slender, bent, reaching to three-fifth of cucul-
lus, processus basalis broad at base tapering soon, tips
bent inward, strongly sclerotized, processus inferior
valvae slightly pointing inward, cucullus normal
sized, bent upward; juxta normal; vinculum angled;
aedeagus small, relatively broad, tapering ventral, tip
overhanging, one large patch of minute cornuti con-
taining one larger cornutus.
Female genitalia (fig. 188). — Papillae anales small;
membrane of tergite VIII with small triangular medi-
um sclerotized spot; ostium straight at top, strongly
sclerotized, lamella antevaginalis shield-like, split in
the middle; ductus bursae moderately broad; ductus
seminalis broad at base; bursa copulatrix large, elon-
gated, two round signa, one small other normal sized.
Biology. — In Tunisia this species flies from half
June to half October with a peak in August-
September. In Sudan it has been caught in the end of
May, August and November. On the Cape Verde
Islands it was caught in August.
Distribution (fig. 87). — A rather disjunct distribu-
tion, Cape Verde Islands, Tunisia and the Sudan.
Remarks. — See under £. sudanellus.
38. Euchromius cambridgei (Zeller, 1867)
(figs. 9, 90-92, 139, 189)
Eromene cambridgei Zeller, 1867: 370. Holotype: 6,
‘Egypte O.P.C. 1864’, GS 5653 (BMNH) [examined].
Eromene luciella Chrétien, 1907: 178. Lectotype: 3 (desig-
nated by Bleszynski (1965a: 77)), ‘Ste Lucie Aude France
8.7.86’, GS 3646 (MNHN) [examined].
Argyria prototypa Meyrick, 1935: 571. Holotype: à , ‘Ksar el
Souk Morocco R.12.5.33’, GS 5668 (BMNH) [examined].
Ommatopteryx congruentella Amsel, 1958a: 64. Holotype:
6, ‘Dhahran 24.8.57 AS. Talhouk coll.’, GS 3325
(LNKD) [examined].
Ommatopteryx szijjartoi Gozmany, 1959: 364. Holotype: 9,
‘Idfu, Egypt 28.x.1957 Exc. Egypt. Mus. Nat. Hung. leg.
Dr. Gozmany’, GS 1002 (HNHM) [examined].
Ommatopteryx ilkui Gozmany, 1959: 366. Holotype: ®,
‘Komosin, Egypt Distr. Fayum 9.x.1957 Exc. Egypt.
Mus. Nat. Hung. leg. Dr. Gozmany’, GS 1003 (HNHM)
[examined].
Material. — 207 5, 169 ©. Algeria: Biskra, 5 d, 4 9
(BMNH), 1 d (INER), 4 d, 1 2 (MNHN); Djanet, 1 d
(BMNH); El Golea, 1 9 (INER), 6 d, 8 2 (MNHN), 2 d
(isNB), 2 d (zsmc); El Kantara, 1 d (BMNH); Hammam-es-
Salahim, 1 d (BMNH); Laghouat, 2 d, 2 2 (MNHN); Le
Tarf, 1 6, 3 2 (MNHN); Sebdou, 1 d (BMNH); Sidi-bel-
Abbes, 1 d (BMNH), 1 d (MNHN). Bahrain: Al Jupair, 1 d
(zMuc); Manama, 5 d (BMNH). Egypt: Aswan, 3 d, 2 9
(BMNH); Cairo, 2 d,2 2 (NHMW), 1 © (MNHN); Idfu, 1 G
SCHOUTEN: Euchromius and Miyakea
(HNHM); Komosin, 1 9 (BMNH), 1 2 (HNHM), 1 d (ISNB);
Minyeh, 1 d (BMNH); Niltakarakt, 2 d, 2 2 (NHMw);
Maryut, 2 d (BMNH), 1 d (LNKD); Sids, 1 d (HNHM); Siwa,
4 2 (BMNH); Zeitoun, 1 d,4 ® (BMNH). France: Bastia, 2
3 (wnkD); Boulouris, 1 d (MNHN); Camarque, 1 d
(BMNH), 1 2 (murs), 1 4,1 2 (LNKD), 1 d (zMFK); Cannes,
1 d (MNHN); Le Canet, 1 2 (BMNH); Les Carbonnes de
Fleurs, 5 km N. of Narbonne, 4 4, 1 ® (nuts); Pinarello, 4
3,4 2 (1zu1); Sole Nzara, 1 d (zut); St. Lucie, 2 4, 1 2
(MNHN); Tour de Vignale, 1 d (1zu1). Iran: Bander Abbas,
5 6,2 ® (nHMw); 30 km E of Bander Abbas, 2 d, 1 9
(NHMW); Bender Tchahbahar, 1 ® (NHRs); Dalaki, 2 9
(LNKD); 35km N. of Minab, 1 & (LNKD); street Bander
Abbas-Sirjan, 1 d, 1 ® (LNKD). Israel: Dead Sea, 1 9
(BMNH); Deganya, 2 d (corp); Sedom, 6 d, 1 © (BMNH);
South end Dead Sea, 1 d (INER), 1 d,1 2 (LNKD), 1 46,1
? (MNHN), 1 2 (NHMW), 1 2 (MHNG), 3 2 (zsmc). Italy:
Siniscola, 2 d (LNKD). Jordan: Fühes, 1 2 (LNKD); Karak, 1
? (LNKD); Zarqa, 1 d, 2 ® (BMNH). Morocco: Aôt Melloul,
2 d (MNHN); Ksar el Souk, 1 d (BMNH); Mehedya, 1 d
(MNHN); Si Aebal Tazi, 1 2 (MNHN); Taourirt, 1 (BMNH).
Oman: Jiddat al Harasis, 1 d, 1 ® (BMNH). Pakistan:
Hydrarabad, 3 2 (BMNH); Karachi, 1 d (BMNH), 1 ©
(LNKD), 1 d, 1 2 (MNHN), 1 2 (NHMW). Saudi Arabia:
Dhahran, 1 d (LNKD); Riad, 3 ® (LNKD). Spain: Bajamar, 3
d (BMNH); Cruz, 1 d (nHMw); El Medano, 1 ® (LNKD);
Fuerteventura, 1 d, 1 ® (zmrk); Hildago, 1 ® (LNKD), 1 d,
1 2 (mnms); La Escala, 2 d (LNKD); Las Palma, 1 ©
(NHMW); Maspalomas, 1 9 (LNKD), 2 d (NHMw); Palma, 1
2 (nHMw); Port Orotava, 1 2 (BMNH), 1 d (RMNH); Rio de
Baza, 1 d (zmuc); Rosas, 2 d (zsmc); Sta. Cruz, 3 6,2 2
(BMNH); Tafira, 1 d (MNHN); Tejina, 1 & (MNMS);
Zaragosa, 1 ® (MNHN). South Yemen: Wadi Maseila, 1 2
(BMNH). Sudan: Ed Damer, 6 d, 11 2 (zsmc); Faras, 1 9
(NHMw); Wadi Halfa, 1 8,1 9 (BMNH), 16,1 2 (nHMw);
Toshka, 1 d, 1 2 (nHMw). Tunisia: Ain Draham, 1 d
(zmuc); Bou-Hedna, 1 & (inER); El Gouina, 2 d (zsmc);
Gafou, 1 4 (MNHN); Metlaoui, 1 & (MNHN); Sfax, 50 d, 55
2 (BMNH), 1 d (casc), 1 ® (curc), 5 6, 4 2 (HNHM), 5 d
(ISNB), 1 & (LNKD), 10 d, 7 2 (MNHN); Tozeur, 1 6
(MNHN). Yugoslavia: Spalato, 1 2 (BMNH), 1 2 (NHMW).
Diagnosis. — Externally very similar to £. sudanel-
lus, E. subcambridgei and E. labellum. Differs from the
last two species by a small point on the frons. In male
genitalia £. cambridgei can be distinguished from E.
sudanellus in lacking the dorsal thorn on the uncus.
The female genitalia have two equally sized signa and
two very small slender sclerotized plates at the ventral
site of the membrane of tergite VIII. The signa are
unequal in size in E. sudanellus and the single plate at
the ventral part of membrane of tergite VIII is trian-
gular.
External characters male, female (figs. 9, 90). —
Wingspan 12-17 mm. Frons produced forward with
small point, creamy white to light brown, more dark
in centre, no ventral ridge; vertex creamy white to
light brown; labial palp two and a half, sides white to
creamy white at base, becoming creamy white to
brown, white from above and below; maxillary palp
white to creamy white, darker ringed at base of last
segment; antenna creamy white, ringed grey. Thorax
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
light brown to brown; patagia light brown to brown
with two very inconspicuous longitudinal brown
stripes; tegulae light brown to brown with a not to
clear dark patch in the middle. Forewing, groundco-
lour creamy white densely suffused with ochreous to
dark brown scales, anterior area usually with dark
brown streak, in dark specimens anterior area entirely
dark brown, posterior area usually with dark brown
streak and a yellowish to dark brown spot, in dark
specimens posterior area entirely dark brown, also
very light specimens occur; medial fascia single, clear-
ly angled under costa, not broadening at inner mar-
gin, running to halfway of the dorsum; subterminal
line ochreous brown sometimes only faintly visible,
about midway between terminal dots and termina-
tion of posterior area; area adjacent to terminal dots
white to yellow; eight to four black terminal dots,
group three usually present; fringes shiny, creamy
white to yellow-brown, sometimes with, not clearly
separated, darker lines. Hindwing grey to light
brown, subterminal fascia present, termen darkly bor-
dered; fringes creamy white with brownish line.
Tergite VIII (fig. 91). — Sclerite normally sclero-
tized, posterior part broad, rounded, faint, stalk faint,
short, anterior part very faint and narrow.
Male genitalia (fig. 139). — Uncus tapering to
sharply pointed tip, inconspicuous dorsal crest
present; gnathos longer, two small dorsal thorns, ter-
minal part short; tegumen without appendix angular-
is; sacculus broad, processus of sacculus slender,
slightly bent upward, reaching to circa three-fifth of
cucullus, processus basalis slender, sharply bent, pro-
cessus inferior valvae small, flattened, cucullus nor-
mal, more or less bent upward; juxta triangular; vin-
culum normal; aedeagus small, one group of cornuti.
Female genitalia (fig. 189). — Papillae anales nor-
mal; membrane of tergite VIII with collar-like scle-
rotization, edges of tergite VIII not connected; os-
tium small; ductus bursae normal; ductus seminalis
narrow; bursa copulatrix roundish, two long signa.
Biology. — Recorded from inland-dune landscape
mixed with salt-areas (Amsel, 1958). Flies in Spain,
France, Italy, Yugoslavia, Tunisia, Algeria and
Morocco in two more or less continues generations
from April until November with peaks in April-May
and September-October. On the Canary Islands it is
also present in November up to April. In Pakistan,
Iran, Oman, South Yemen, Saudi Arabia, Jordan,
Israel, Egypt and Sudan it has been caught through-
out the year, with peaks in February-May and
October-November.
Distribution (fig. 92). — Spain, France, Italy,
Yugoslavia, Iran, Pakistan, Bahrain, Oman, South
Yemen, Saudi Arabia, Jordan, Israel, Egypt, Sudan,
Tunisia, Algeria and Morocco.
Remarks. — Most variable species in the genus, in
232
size as well as in colour. The darkest coloured speci-
mens occur on the Canary Islands and in the West-
Mediterranean countries. The palest and smallest
specimens are found in Iran.
39. Euchromius viettei Bleszynski, 1961
(figs. 93, 140, 141, 190)
Euchromius viettei Bleszynski, 1961: 455. Holotype: à,
“Arabia: Hejaz, Jidda. 22.X1.1926 H. St. J. B. Philby’, GS
5091 (BMNH) [examined]. Schouten (1988: 24) [rede-
scription].
Euchromius karsholti Ganev, 1987: 44. Holotype: à , ‘Saudi
Arabia FEB 1980 Djidda, Northern Creek U. Seneka
Nielsen leg’, GS 968 (zmuc) [examined]. syn. n.
Material. — 10 6, 4 2. Chad: Binni Erdi, 4 d, 2 9
(BMNH). Oman: Khubayt, 1 d (BMNH); Qurm, 1 6
(BMNH). Saudi Arabia: Burayman, 1 d, (BMNH); Djidda, 1
? (zmuc); Hejaz, 3 d, 1 2 (BMNH).
Diagnosis. — Difficult to distinguish externally
from other species with a single medial fascia. Differs
in male genitalia in having the processus of sacculus
forming a heavily spined lobe in combination with
the processus basalis broad at base, narrowing very
abruptly. Differs in female genitalia in a roundish
bursa copulatrix with one round signum in combina-
tion with edges of tergite VIII connected forming a
minute projection and having the lamella postvagin-
alis rectangular.
Biology. — Unknown. Specimens caught in
February, November and December in Saudi Arabia
and Oman, in August in Chad.
Distribution (fig. 93). — Occurs in the Saudian
Peninsula and the Tibesti Mountains of Chad.
Remarks. — Ganev (1987: 44) describes the species
Euchromius karsholti based on a single female from
Saudi Arabia. In the diagnosis he gives no distinct dif-
ferences between E. viettei Bleszynski, 1961 and kars-
holti. He only states: “VIII. tergite very complicated,
however differing from those of Euchromius hampso-
niand Euchromius viette?. The quality of the drawing
of the female genitalia is too poor to make any state-
ment on the degree of difference between the species
concerned. Study of the holotypes proved no differ-
ences between E. karsholti Ganev, 1987 and E. viettei
Bleszynski, 1961. E. karsholti Ganev, 1987 is there-
fore a junior synonym of E. viettei Bleszynski, 1961.
40. Euchromius hampsoni (Rothschild, 1921)
(figs. 143, 191)
Ommatopteryx hampsoni Rothschild, 1921: 220. Holotype:
3, ‘Azzal N. of Agades 13. July 20. (A. Buchanan)’, GS
5676 (BMNH) [examined]. Schouten (1988: 25) [rede-
scription].
Material. — 1 4,1 9.
Diagnosis. — Difficult to distinguish externally
from other species with a single medial fascia. Differs
in male genitalia in having the processus of sacculus
forming a heavily spined lobe in combination with
the processus basalis broad at base, narrowing gradu-
ally. Differs in female genitalia in a roundish bursa
copulatrix with one round signum in combination
with lamella postvaginalis square, lamella antevagin-
alis with clear anterior fold.
Distribution. — Niger.
41. Euchromius klimeschi Bleszynski, 1961
(figs. 142, 192)
Euchromius klimeschi Bleszynski, 1961: 467. Holotype: à,
‘Natal Weenen 1-iii-1927 H. P. Thomasset’, GS 5096
(BMNH) [examined]. Schouten (1988: 14) [redescrip-
tion].
Material. — 14 4, 30 9.
Diagnosis. — Differs from all other species by a
rounded frons without a point, a single medial fascia,
six or seven black terminal dots and a white area adja-
cent to terminal dots.
Distribution. — Ethiopia, Somalia, Kenya,
Tanzania, Zaire, Zambia, Zimbabwe, Madagascar,
Mozambique, South Africa, Namibia and Burkina
Faso [=Upper Volta]. B. Landry collected the first
specimen for West Africa in Burkina Faso:
Kompienbiga, 15 km W. of Pama, 5-13.VIII.1988
(LAND).
Remark. — Due to an incorrectly positioned female
genitalia the ductus seminalis was wrongly interpret-
ed and recorded as narrow (Schouten 1988). In con-
trast to this the ductus seminalis starts broad and nar-
rows soon, the sclerotized ring starts in the broad part
of the ductus seminalis.
42. Euchromius discopis (Hampson, 1919)
(figs. 144, 193)
Ommatopteryx discopis Hampson, 1919: 534. Holotype: 9,
“Transvaal 1907-122 Pretoria 21.10.06 A. J. T. Janse II’
GS 7049 (BMNH) [examined].
Euchromius discopis (Hampson, 1919). Schouten (1988: 16)
[redescription].
Material. - 23 6, 24 2.
Diagnosis. — Difficult to distinguish from other
species with eight or nine black terminal dots and a
single medial fascia. Differs from these species in male
genitalia in having the processus of sacculus long and
slender, nearly reaching the end of the cucullus, a
SCHOUTEN: Euchromius and Miyakea
dorsal spike at base of the processus. Differs in female
genitalia in having a tooth-shaped ostium in combi-
nation with a oblong bursa copulatrix with one faint
signum.
Distribution. —
Africa and Namibia.
Zimbabwe, South
Botswana,
43. Euchromius labellum Schouten, 1988
(figs. 9, 146, 196)
Euchromius labellum Schouten, 1988: 17. Holotype: 9,
‘Afrika Kenya Samburu Game Reserve, Lodge 15.2.1975
LF leg. D. Buckh.’, GS 393 (mung) [examined].
Material.-8 46,6 2.
Diagnosis. — Differs from most species in having
the single medial fascia angled under the costa.
Differs in male genitalia in having a lip-shaped bi-
lobed uncus. Differs in female genitalia in having a
broad tongue-shaped ostium in combination with a
roundish bursa copulatrix with one signum and a
broadly starting ductus seminalis which narrows
soon.
Biology. — The author collected material of this
species in Samburu National Park. The specimens
were caught at 925 m altitude, approximately 50 m
from the river, in an area with scattered Acacia spp.
trees with some shrubs and patches of grass.
Distribution. — Kenya.
44. Euchromius aris Schouten, 1988
(figs. 145, 195)
Euchromius aris Schouten, 1988: 22. Holotype: d,
‘Samburu Res. KENYA 2800’ 6 March ‘86 R. Leuchner’,
GS 420 (LACM) [examined].
Material. — 10 4,20 9.
Diagnosis. — Difficult to separate from species with
a single medial fascia and nine (or eight) black termi-
nal dots and a bluntly produced rounded frons.
Differs from these species in male genitalia in having
a gnathos without a terminal part in combination
with a slender, sharply pointed cucullus and a aedea-
gus with one group of cornuti. Differs in female gen-
italia in having the bursa copulatrix with two round-
ed signa in combination with a small dorsally bent
lip-shaped projection and the lamella postvaginalis
square.
Distribution. — Kenya.
45. Euchromius erum Schouten, 1988
(figs. 147, 196)
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“TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Euchromius erum Schouten, 1988: 21. Holotype: 9, ‘Kenya:
5 mi NE Kargi Marsabit District elev. ca 1500 feet 28
Jan.-3 Feb. 1973 Julian Donahue’, GS 365 (LACM) [ex-
amined].
Material. —3 d, 112.
Diagnosis. — Difficult to separate from species with
a single medial fascia and a bluntly projecting round-
ed frons. Differs from these species in male genitalia
in lacking the two dorsal thorns of the gnathos and
the appendix angularis in combination the presents of
heavy spines on the valvae. Differs in female genitalia
by the upper two-third of the ductus bursae being
evenly sclerotized by small spots.
Biology. — See under £. labellum.
Distribution. — Ethiopia and Kenya.
46. Euchromius locustus Schouten, 1988
(figs. 148, 197)
Euchromius locustus Schouten, 1988: 18. Holotype: 9,
“Zambia Mbala 3-8.x.1974 Locust Cont. Ctr. BM.1975-
92.’, GS 12986 (BMNH) [examined].
Material. — 2 6,4 9.
Diagnosis. — Difficult to separate from species with
a single medial fascia and a bluntly projecting round-
ed frons and nine (or eight) black terminal dots.
Differs from these species in male genitalia in having
a broad, triangular appendix angularis in combina-
tion with the absence of the two dorsal thorns of the
gnathos.
Distribution. — Tanzania, Zaire and Zambia.
47. Euchromius nigrobasalis Schouten, 1988
(figs. 149, 198)
Euchromius nigrobasalis Schouten, 1988: 20. Holotype: à,
‘Pretoria 11.x.1958 L. Vari’, GS 155 (rMsA) [examined].
Material. — 3 6,4 9.
Diagnosis. — Differs from all other species in hav-
ing a broad single medial fascia which is sprinkled
with black scales giving it a fingerprint impression.
Distribution. — Zimbabwe and South Africa.
Miyakea Marumo, 1933
Miyakea Marumo, 1933: 48. Type species: Eromene expansa
Butler, 1881: 590, by monotypy.
The genus Miyakea was described by Marumo in
1933 to accommodate Eromene expansa Butler, 1810.
Caradja (1910) described a new form of Eromene bel-
234
la (= Euchromius bellus) from Russia: Radde and
named it raddeellus. Bleszynski & Collins (1962)
made it a synonym of M. expansa (Butler, 1881). The
third species has only recently been described from
Taiwan: Euchromius lushanus Inoue, 1989. It is now
transferred to Miyakea. The female genitalia of M. ex-
pansa are depicted and described for the first time. M.
sinevi sp. n. is described from Mongolia.
Diagnosis of Miyakea
Ductus ejaculatorius enters the aedeagus terminal-
ly; dorsal thorns on the gnathos absent; tergite VIII
without sclerotized pattern; frons projecting or not;
M1 of hindwing located in the lower part of the mo-
re or less closed cell; base of M2 and M3 of hindwing
with a basal pointing vein, more or less closing the
cell (fig. 10); spinula not visible.
Morphology
Head. The frons has no diagnostic characters at
species level, it is projection or not, without point,
creamy white to brown without a ventral ridge. The
relative size of the labial palp compared to the eye dia-
meter can be used to separate the species. The anten-
na are serrate in males and setaceous in females. The
scales on the antenna are creamy white, sometimes
inconspicuous darkly ringed.
Thorax. Thorax uniformly coloured, creamy white
to brown. Patagia light brown to brown. Tegulae
creamy white with a dark patch in the middle.
Venation (fig. 10). Very similar to that of the genus
Euchromius. The differences with the latter genus are
listed under ‘Diagnosis’.
Wing pattern. The ground-plan of the pattern is
similar to that of Euchromius (fig. 7), but there are no
species with a single medial fascia. The formula of the
black terminal dots usually offers a good character to
distinguish the species. The position of the double
medial at the costa is in some cases diagnostic.
Genitalia. The male genitalia offer excellent diag-
nostic characters. The uncus is rather uniform, slen-
der and tapering to a sharply pointed tip. The gna-
thos is of equal length, the tip is minutely dentate.
The sacculus is reduced to absent. The processus bas-
alis is reduced to very prominent. The costal edge of
the valvae forming a prominent projection or not, the
cucullus is truncate. Processus inferior valvae incon-
spicuous to absent. Juxta small, rounded. Vinculum
long. Aedeagus highly diagnostic, usually with cornu-
ti. Female genitalia, papillae anales uniformly small.
Ostium armed with variously shaped projections.
Ductus bursae short, with or without a pouch-like
enlargement under the ostium. Ductus seminalis al-
ways narrow. Bursa copulatrix oblong without signa.
Intraspecific variation
The intraspecific variation of Miyakea is limited,
the male genitalia in raddeellus show some variation.
Tympanal organs
In Miyakea the tympanal organs are similar to
those of Euchromius, but the spinula on the tympa-
num is not sclerotized.
Distribution
The four species are confined to Asia, China,
Taiwan and Japan. M. expansa is endemic to Japan,
lushanus is endemic to Taiwan. M. sinevi has been
found in Mongolia. M. raddeellus has the most exten-
sive distribution, occurring in China, Russia and
Korea.
Biology
Unknown. Raddeellus has been caught at an altitu-
de of 1800 m.
Checklist of extant Miyakea Marumo
1. lushanus (Inoue, 1989)
2. raddeellus (Caradja, 1910)
3. expansa (Butler, 1881)
4. sinevi sp. n.
1. Miyakea lushanus (Inoue, 1989) comb. n.
(figs. 94, 95, 118, 169)
Euchromius lushanus Inoue, 1989: 2. Holotype: d, ‘Lushan
Nantou Formosa 29.VII.1973 Y. Shibata’, GS 13229
(inou) [examined].
Material. — 1 6, 1 ©. Taiwan: Nantou, 1 6, 1 2 (INoU).
Diagnosis. — Differs from M. sinevi by the present
of a group of three black terminal black dots. Differs
from M. expansa in not having the medial fascia con-
nected with the yellow apical marking. Differs from
M. raddeellus in male genitalia lacking the processus
basalis, in female genitalia in lacking the tongue-
shaped projection of the ostium.
External characters male, female (fig. 94). —
Wingspan 20-26 mm. Frons produced forward,
rounded without point, light brown to creamy white,
no ventral ridge; vertex creamy white, light brown in
the middle; labial palp three, creamy white at base,
very soon brown, becoming brown lead-grey, brown
from above, creamy white from below; maxillary palp
brown, terminal part creamy white; antenna creamy
white. Thorax brown; patagia light brown to brown;
tegulae creamy white with brown patch in middle.
Forewing, groundcolour creamy white densely suf-
fused with ochreous to dark brown scales, costa of an-
SCHOUTEN: Euchromius and Miyakea
terior area brown; medial fascia double slightly
arched, running to one-fourth of the dorsum; subter-
minal line brown, about midway between terminal
dots and termination of posterior area; area adjacent
to terminal dots yellow to white; seven black terminal
dots, formula 2-3-2; fringes shiny, evenly lead-grey.
Hindwing grey-brown, subterminal fascia present,
termen darkly bordered; fringes creamy white.
Male genitalia (fig. 118). — Uncus normal; gnathos
equal in length tip minutely dentate, dorsal thorns
absent; tegumen slender, appendix angularis absent;
sacculus inconspicuous, processus of sacculus absent,
processus basalis reduced to elongated rim, processus
inferior valvae absent, cucullus slightly concave,
blunt; juxta roundish; vinculum elongated; aedeagus
curved, no cornuti.
Female genitalia (fig. 169). — Papillae anales small;
membrane of tergite VIII without sclerotizations; os-
tium small with very small low projecting tooth; duc-
tus bursae short with more sclerotized lines; ductus
seminalis narrow; bursa copulatrix oblong, signum
absent.
Biology. — Unknown. Specimens were caught in
July and August at 1200 m altitude.
Distribution (fig. 95). — Taiwan.
Remarks. — M. lushanus is the sister species of rad-
deellus and/or expansa judging to the male genitalia
and falls within the genus diagnosis of the genus
Miyakea. Leaving this species in Euchromius would
cause a polyphyletic genus. Therefore /ushanus is
transferred to Miyakea Marumo, 1933.
In the original description slide number 13230 is
given as belonging to the holotype, but upon check-
ing the type material this turns out to be the female
paratype slide number. The holotype slide number is
13229.
2. Miyakea raddeellus (Caradja, 1910) sp. rev.
(hies..9,.95, 96, 1191202167)
Eromene bellus form raddeella Caradja, 1910: 115.
Lectotype: d (designated by Bleszynski (1965a: 89)),
[Russia] ‘Radde Amur 1903 Korb’. (MGAB) [not exam-
ined].
Miyakea expansa sensu Bleszynski & Collins (nec Butler,
1881) 1962: 325 [status nov. for Eromene bellus form
raddeellus Caradja, 1910 and synonymization with
Miyakea expansa (Butler, 1881)].
Material. — 10 d, 14 2. China: Changyang, 1 d
(BMNH), 1 d (rMs4); Chung Kiang [=Zhongjiang], 1 9
(BMNH); Chung King, 1 d, 5 2 (BMNH); Foochow
[=Fuzhou], 2 d, 2 2 (BMNH), 1 d (RTAS), 1 2 (TMsa);
Siang-Yang-Fu [=Xiangfan], 1 d (BMNH); Wenchow
[=Wenzhou], 1 2 (rras), 3 d, 3 2 (zrmk). Korea: Ryong
Hpieng [locality not found], 1 ® (MNHN).
235
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
@ expansa
ô W raddeellus
o A
lushanus
Figs. 93-97. Euchromius and Miyakea. — 93, E. viettei, distribution map; 94, M. lushanus, head, lateral aspect, scales removed;
95, distribution map of M. lushanus, M. raddeellus, M. expansa and M. sinevi, 96, M. raddeellus, head, lateral aspect, scales re-
moved; 97, M. expansa, head: lateral aspect, scales removed. Scale bar 0.5 mm to figs. 94, 96-97.
Diagnosis. — M. raddeellus can be separated from
M. expansa by a double medial fascia which does not
touch the yellow apical marking in combination with
seven black terminal dots. Differs from M. sinevi in
having a group of three black terminal dots. Differs
from M. lushanus in male genitalia by the presence of
a processus basalis, in female genitalia by a tongue-
shaped projection of the ostium.
External characters. male, female (figs. 9, 96). —
Wingspan 19-28 mm. Frons not produced forward,
236
rounded without point, light brown, darker in centre,
no ventral ridge; vertex light brown; labial palp two,
sides creamy white at base, becoming lead-grey,
brown from above, light brown from below; maxil-
lary palp light brown, dark brown ringed at base of
last segment, terminal part light brown; antenna
creamy white. Thorax creamy white to light brown;
patagia light brown; tegulae creamy white with dark
patch in the middle. Forewing, groundcolour creamy
white, densely suffused with ochreous to dark brown
scales, posterior area with, sometimes inconspicuous,
spot near medial fascia; medial fascia double, concave
to nearly straight, running to one-fourth to one-third
of the dorsum; subterminal line ochreous to dark
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white; seven black terminal dots, formula 2-3-2;
fringes highly shiny, evenly grey. Hindwing creamy
white to light brown, subterminal fascia usually clear-
ly present, termen darkly bordered; fringes creamy
white with faint brownish line.
Male genitalia (figs. 119, 120). — Uncus slender, ta-
pering to sharply pointed tip; gnathos equal in length,
tip minutely dentate, dorsal thorns absent; tegumen
slender, appendix angularis absent; sacculus incon-
spicuous, narrow, processus of sacculus absent, pro-
cessus basalis large, broad, various in length, tip elon-
gated, bent inward, second strongly sclerotized
projection with dentated dorsal edge formed by costal
edge of valvae, reaching beyond cucullus, processus
inferior valvae small, inconspicuous, cucullus broad,
ending truncate; juxta small, rounded, fringed with
hairy membrane; vinculum long, elongated; aedeagus
large with three to five cornuti.
Female genitalia (fig. 167). — Papillae anales small;
membrane of tergite VIII without sclerotizations; os-
tum with bifurcate projection, projection covered
with minute spines; ductus bursae short, broad,
strongly sclerotized in upper part, armed with small
spines; ductus seminalis narrow; bursa copulatrix
large, oblong, no signa.
Biology. — Unknown. This species has been caught
in April, May, June, July and September. Altitude
mentioned: 1800 m.
Distribution (fig. 95). — China, Korea and USSR.
Remarks. — All previous literature concerning ma-
terial of Miyakea expansa outside Japan refers to M.
raddeellus or M. lushanus. The type material of this
species could not be studied since the collections in
Bucarest were closed for research during my visit to
the museum.
3. Miyakea expansa (Butler, 1881)
(figs. 9, 95, 97, 121, 168)
Eromene expansa Butler, 1881: 590. Lectotype: ® (designat-
ed by Bleszynski (1965a: 89), ‘Tokei 80.97’, GS 18213
(BMNH) [examined].
Miyakea expansa (Butler, 1881) Marumo (1933: 49).
Euchromius expansa (Butler, 1881) Inoue (1982: 225).
Miyakea raddeellus sensu Bleszynski & Collins (1962: 325)
(nec Caradja, 1910).
Material. — 1 d,1 9. Japan: Otaru, 1 6 (BMNH); Tokei,
1 2 (BMNH).
SCHOUTEN: Euchromius and Miyakea
Diagnosis. — M. expansa is easily distinguished
from all other species by its six black terminal dots in
which a group of three dots is present in combination
by the outer medial fascia touching the yellow apical
marking of the forewing.
External characters male, female (figs. 9, 97). —
Wingspan 21-30 mm. Frons not produced forward,
rounded without point, light brown to creamy white,
no ventral ridge; vertex yellow-brown; labial palp
two, sides brown, brown from above, creamy white
from below; maxillary palp brown, terminal part
creamy white; antenna creamy white, inconspicuous
darkly ringed. Thorax light brown; patagia light
brown; tegulae creamy white with dark brown patch
in the middle. Forewing, groundcolour creamy white
densely suffused with ochreous to dark brown scales;
medial fascia double, slightly arched, running to one-
fourth of the dorsum; subterminal line dark brown,
about midway between terminal dots and termina-
tion of posterior area; area adjacent to terminal dots
yellow to white; six black terminal dots, formula 2-3-
1, central dots of group ‘three’ sometimes very large,
nearly double; fringes shiny, evenly lead-grey.
Hindwing grey, subterminal fascia present, termen
darkly bordered; fringes creamy white.
Male genitalia (fig. 121). — Uncus slender, tapering
to pointed tip; gnathos equal in length, tip minutely
dentate, dorsal thorns absent; tegumen slender, ap-
pendix angularis absent; sacculus narrow, processus of
sacculus absent, processus basalis large, swollen in ba-
sal half, dorsal edge very roughly dentate, processus
inferior valvae small, inconspicuous, cucullus slender
ending truncate; juxta normal sized, rounded; vincu-
lum long, elongated; aedeagus relatively large, one el-
ongated group of cornuti.
Female genitalia (fig. 168). — Papillae anales small;
membrane of tergite VIII without sclerotizations,
edges of tergite VIII free; ostium simple, with tongue-
shaped projection; ductus bursae grooved in upper
half, small pouch-like enlargement direct under os-
tium; ductus seminalis narrow; bursa copulatrix ob-
long, no signa.
Biology. — Unknown. Caught in June.
Distribution (fig. 95). — Japan.
Remarks. — M. expansa is restricted to Japan. All
references concerning material of the mainland of
Asia are most likely to belong to M. raddeellus.
Bleszynski (1965a) erroneously depicted the female
genitalia of M. raddeellus and stated it to be those of
expansa.
4. Miyakea sinevi sp. n.
(figs. 95, 97a)
Type material. — Holotype: 1 2, ‘Mongoli’, GS R.S. 767
(ZMAS).
DSTI
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Diagnosis. — M. sinevi is easily distinguished from
all other species by lacking a group of three black ter-
minal dots.
External characters female — Wingspan 20 mm.
Frons not produced forward, rounded without point,
brown with some creamy white scales, no ventral
ridge; vertex creamy white with brown scales; labial
palp three, sides brown, brown from above, brown
from below; maxillary palp brown; antenna creamy
white, inconspicuous darkly ringed. Thorax brown;
patagia brown; tegulae creamy white with a broad
dark brown patch in the middle. Forewing, ground-
colour creamy white densely suffused with ochreous
to dark brown scales; costal area of anterior part
brown; medial fascia double, slightly arched, running
to one-fourth of the dorsum; subterminal line dark
brown, about midway between terminal dots and ter-
mination of posterior area; area adjacent to terminal
dots white; eight or seven black terminal dots, formu-
la 2-2-2-2 or 2-2-2-1; fringes shiny, evenly lead-grey.
Hindwing grey-brown, subterminal fascia absent;
fringes too worn to be described.
Female genitalia (fig. 97a). — Papillae anales small;
membrane of tergite VIII without sclerotizations,
edges of tergite VIII free; ostium simple, with a broad
tongue-shaped projection, under the ostium a pouch
with a strong sclerotized part is clearly separated from
the ductus bursae; ductus bursae long; ductus semi-
nalis narrow; bursa copulatrix small, no signa.
Biology. — Unknown. Caught in June.
Distribution (fig. 95). — Mongolia.
Etymology. — This species is named in honour of
Dr. Sinev of St. Petersburg a highly esteemed special-
ist of Momphidae.
ACKNOWLEDGEMENTS
I wish to thank the following persons and institutes for
loans and gifts of material without which this study would
not have been possible.
Prof. Dr. I. Ahmad (eskk, Karachi, Pakistan), Dr. S.
Allyson (encr, Ottawa, Canada), Dr. P. H. Arnaud Jr.
(casc, San Francisco, U.S.A.), Dr. D. Azuma (AnsP,
Philadelphia, U.S.A.), Dr. M. Brancucci (NHMB, Basel,
Switzerland), Dr. D. Buckhardt (MHNG, Geneve,
Switzerland), Dr. G. W. Byers (semc, Lawrence, U.S.A.),
Dr. C. B. Cottrell (rMsA, Pretoria, South Africa), Dr. U.
Dall’Asta (mrac, Tervuren, Belgium), Dr. D. R. Davis
(UsNM, Washington, U.S.A.), Mr. G. Derra (Bamberg,
Germany), Dr. W. Dierl (zsmc, Munchen, Germany), Dr.
J. P. Donahue (Lacm, Los Angeles, U.S.A.), Dr. D. K.
Faulkner (spmc, San Diego, U.S.A.), Dr. A. Freidberg (-
TAUI, Tel Aviv, Israel), Mr. C. Gielis (Lexmond, The
Netherlands), Dr. P. Grootaert (1sNB, Brussel, Belgium),
Dr. G. F. Gross (sama, Adelaide, Australia), Dr. B.
Gustafsson (NHRS, Stockholm, Sweden), Mr. W. Hogenes
(ZMAC, Amsterdam, The Netherlands), Dr. P. Huemer
(izui, Innsbruck, Austria), Mr. K. J. Huisman (Wezep, The
Netherlands), Dr. E. Hull (Helsby, Great Britain), Dr. H.
238
Inoue (Tokyo, Japan), Dr. H. D. Jackson (NMBz, Bulawayo,
Zimbabwe), Dr. R. de Jong (RMNH, Leiden, The
Netherlands), Dr. O. Karsholt (zmuc, Copenhagen,
Denmark), Dr. R. Krause (smtp, Dresden, Germany), Mr.
B. Landry (cncı, Ottawa, Canada), Dr. J. K. Liebherr
(cuic, Ithaca, New York, U.S.A.), Dr. M. Lödl (NMHw,
Vienna, Austria), Mr. J. A. W. Lucas (Rotterdam, The
Netherlands), Dr. G. Luquet (MNHN, Paris, France), Dr. K.
Maes (Gent, Belgium), Dr. S. E. Miller (BPBM, Honolulu,
U.S.A.), Dr. E. S. Nielsen (anic, Canberra, Australia), Prof.
U. Parenti (Mızı, Torino, Italy), Dr. K. T. Park (cısk,
Chuncheon, South Korea), Dr. J. Powell (cısc, Berkeley,
U.S.A.), Mr. W. De Prins (Berchem/Antwerpen, Belgium),
Dr. J. E. Raastad (zmun, Oslo, Norway), Dr. J. E. Rawlins
(ccm, Pittsburgh, U.S.A.), Dr. F. H. Rindge (AMNH, New
York, U.S.A.), Dr. J. M. Ritchie (NMKE, Nairobi, Kenya),
Dr. R. Robineau (Tremblay-les-Gonesse, France), Prof. Dr.
R. U. Roesler (LNKD, Karlsruhe, Germany), Dr. P. H. Scott
(SBMN, Santa Barbara, U.S.A.), Dr. R. Schuster (ucpc,
Davis, U.S.A.), Dr. S. Yu. Sinev (zmas, St. Petersburg,
Russia), Dr. M. A. Solis (usNM, Washington, U.S.A.), Dr.
D. Stüning (ZFMK, Bonn, Germany), Dr. P. Y. Ustjuzhanin
(Novosibirsk, Russia), Dr. L. Vari (TMsA, Pretoria, South
Africa), Dr. A. Vives (MNMS, Madrid, Spain), Dr. C. Vogt
(mczc, Massachusetts, U.S.A.), Dr. A. Vojnits (HNHM,
Budapest, Hungary), Mr. H. W. van der Wolf (Nuenen,
The Netherlands).
I wish to thank Mr. F. Abdoel (Museon, Den Haag, The
Netherlands) for assistance with the colour plate. I am in-
depted for advise and critical remarks to Dr. J. P. Duffels
(Amsterdam), Prof. Dr. E. Gittenberger (Leiden) and Dr. R.
de Jong (Leiden).
I especially wish to thank Mr. M. Shaffer (BMNH,
London, United Kingdom) for his great assistance both dur-
ing my many visits to the BMNH and for the important pre-
liminary study he carried out on the BMNH material.
Visits to the BMNH, ZSMC and NMHW have been made
possible by grants awarded by the Uyttenboogaart-Eliasen
Stichting.
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SCHOUTEN: Euchromius and Miyakea
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Received: April 1992
Revised version accepted: September 1992
Fig. 97a. Female genitalia; ventral aspect. Miyakea sinevi. Scale bar 1 mm.
241
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
RN
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Figs. 98-100. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 98, Euchromius ornatus,
99, Euchromius cornus, 100, Euchromius californicalis. Scale bar 0.5 mm.
242
SCHOUTEN: Euchromius and Miyakea
LH ty if
ww 7
A
Figs. 101-105. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 101, Euchromius mat-
ador, 102, Euchromius gnathosellus, 103-105, Euchromius zephyrus, 103, valvae, 104, juxta separated, 105, tegumen, uncus
and gnathos separated. Scale bar 0.5 mm.
243
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
US SNN
IN
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Figs. 106-108. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 106, Euchromius tan-
alis 107, Euchromius mythus, 108, Euchromius galapagosalis. Scale bar 0.5 mm.
244
SCHOUTEN: Euchromius and Miyakea
VA | vd
TAI
4
Figs. 109-110. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 109, Euchromius li-
maellus, 110, Euchromius minutus. Scale bar 0.5 mm.
245
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
SNN
Figs. 111-113. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated. — 111, Euchromius saltalis, aedeagus
dorsal aspect; 112, Euchromius ocelleus, aedeagus lateral aspect; 113, Euchromius confusus, aedeagus lateral aspect. Scale bar 0.5
mm.
246
SCHOUTEN: Euchromius and Miyakea
Figs. 114-118. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 114, Euchromius pul-
verosus, 115-117, Euchromius vinculellus, 115, male genitalia, 116-117, processus basalis; 118, Miyakea lushanus. Scale bar 0.5
mm.
247
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 119-121. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated. — 119-120, Miyakea raddeellus, 119,
male genitalia, aedeagus lateral aspect, 120, processus basalis and valve; 121, Miyakea expansa, aedeagus dorsal aspect. Scale
bar 0.5 mm.
248
SCHOUTEN: Euchromius and Miyakea
Figs. 122-124. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 122, Euchromius ana-
piellus, 123, Euchromius bellus, 124, Euchromius bleszynskiellus. Scale bar 0.5 mm.
249
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Mag
LET
VIE
AIR
Ni
NN IR Ne
Figs. 125-127. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 125, Euchromius bles-
zynskii, tegumen, uncus and gnathos separated. Scale 0.5 mm. 126, Euchromius scobiolae. Scale 1 mm. 127, Euchromius super-
bellus. Scale bar 0.5 mm.
250
SCHOUTEN: Euchromius and Miyakea
N
a,
Figs. 128-130. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 128, Euchromius ke-
redjellus, 129, Euchromius malekalis, 130, Euchromius mouchai. Scale bar 0.5 mm.
251
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Ys BR
NIN
M
N
N NN
Figs. 131-133. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 131, Euchromius niv-
alis 132, Euchromius rayatellus 133, Euchromius gozmanyi. Scale bar 0.5 mm.
252
SCHOUTEN: Euchromius and Miyakea
Figs. 134-136. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 134, Euchromius gra-
tiosellus, 135, Euchromius jaxartellus, 136, Euchromius ramburiellus. Scale bar 0.5 mm.
255
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
À | INN »
pis:
Wh iif Si, ies EN
SME ar
Engen
Figs. 137-139. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 137, Euchromius su-
danellus, tegumen, uncus and gnathos separated; 138, Euchromius subcambridgei, tegumen, uncus and gnathos separated; 139,
Euchromius cambridgei. Scale bar 0.5 mm.
254
SCHOUTEN: Euchromius and Miyakea
Figs. 140-143. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 140-141, Euchromius
viettei, 140, male genitalia, aedeagus lateral aspect, 141, valve and lobe; 142, Euchromius klimeschi 143, Euchromius hampso-
ni, tegumen, uncus and gnathos separated. Scale bar 0.5 mm.
255
TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 144-146. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 144, Euchromius dis-
copis 145, Euchromius aris, 146, Euchromius labellum. Scale bar 0.5 mm.
256
SCHOUTEN: Euchromius and Miyakea
RE
149
Figs. 147-149. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 147, Euchromius er-
um; 148, Euchromius locustus, 149, Euchromius nigrobasalis. Scale bar 0.5 mm.
257
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 150-151. Female genitalia; ventral aspect. — 150, Euchromius circulus, 151, Euchromius cornus. Scale bar 0.5 mm.
258
SCHOUTEN: Euchromius and Miyakea
Figs. 152-153. Female genitalia; ventral aspect. — 152, Euchromius californicalis 153, Euchromius matador. Scale bar 0.5 mm.
259
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 154-156. Female genitalia; ventral aspect. — 154, Euchromius gnathosellus, 155, Euchromius zephyrus 156, Euchromius
tanalis. Scale bar 0.5 mm.
260
SCHOUTEN: Euchromius and Miyakea
Figs. 157-159. Female genitalia; ventral aspect. — 157, Euchromius mythus 158, Euchromius geminus, 159, Euchromius galap-
agosalis. Scale bar 0.5 mm.
261
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 160-162. Female genitalia; ventral aspect. — 160, Euchromius limaellus, 161, Euchromius minutus, 162, Euchromius sal-
talis. Scale bar 0.5 mm.
262
SCHOUTEN: Euchromius and Miyakea
Figs. 163-165. Female genitalia; ventral aspect. — 163, Euchromius ocelleus, 164, Euchromius confusus, 165, Euchromius pulve-
rosus. Scale bar 0.5 mm.
263
Tir SCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 166-167. Female genitalia; ventral aspect. — 166, Euchromius vinculellus, 167, Miyakea raddeellus. Scale bar 0.5 mm.
264
SCHOUTEN: Euchromius and Miyakea
|
Figs. 168-170. Female genitalia; ventral aspect. — 168, Miyakea expansa; 169, Miyakea lushanus, 170, Euchromius gratiosellus.
Scale bar 1 mm.
265
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 171-173. Female genitalia; ventral aspect. — 171, Euchromius anapiellus 172, Euchromius bellus, 173, Euchromius bles-
zynskiellus. Scale bar 0.5 mm.
266
SCHOUTEN: Euchromius and Miyakea
174
A) Q\ Z
ur WORN
ar I
CLEA INNI i
ail INN |
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NIN
NN
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Figs. 174-176. Female genitalia; ventral aspect. — 174, Euchromius bleszynskii, 175, Euchromius scobiolae, 176, Euchromius
superbellus. Scale bar 0.5 mm.
267
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
178
N JA
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a | | iN
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Figs. 177-179. Female genitalia; ventral aspect. — 177, Euchromius keredjellus, 184, Euchromius mouchai, 185, Euchromius ma-
lekalis. Scale 0.5 mm.
268
SCHOUTEN: Euchromius and Miyakea
Figs. 180-182. Female genitalia; ventral aspect. — 180, Euchromius rayatellus, 181, Euchromius nivalis, 182, Euchromius goz-
manyi. Scale bar 0.5 mm.
269
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 183-185. Female genitalia; ventral aspect. — 183, Euchromius jaxartellus, 184, Euchromius zagulajevi, 185, Euchromius
ramburiellus. Scale 0.5 mm.
270
SCHOUTEN: Euchromius and Miyakea
genitalia; ventral aspect. — 186, Euchromius donum; 187, Euchromius sudanellus, 188, Euchromius sub-
.5 mm.
Figs. 186-188. Female
cambridgei. Scale bar 0
271
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 189-191. Female genitalia; ventral aspect. — 189, Euchromius cambridgei, 190, Euchromius vietter, 191, Euchromius
hampsoni. Scale bar 0.5 mm.
272.
Figs. 192-194. Female genitalia; ventral aspect. — 192, Euchromius klimeschi, 193, Euchromius discopis 194, Euchromius label-
lum. Scale bar 0.5 mm.
Zijl)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 195-198. Female genitalia; ventral aspect. — 195, Euchromius aris, 196, Euchromius erum; 197, Euchromius locustus, 198,
Euchromius nigrobasalis. Scale bar 0.5 mm.
274
BERNHARD J. van VONDEL
Natuurmuseum Rotterdam
REVISIONIOF THE PALAFARCTIG AND ORIENTAL
SPECIES OF PELTODYTES REGIMBART
(COLEORTERASEABIRENDAE)
Vondel, B. J. van, 1992. Revision of the Palaearctic and Oriental species of Peltodytes
Régimbart (Coleoptera: Haliplidae). — Tijdschrift voor Entomologie 135: 275-297, figs 1-101.
[ISSN 0040-7496]. Published 1 December 1992.
The eight Palaearctic and Oriental species of the genus Peltodytes Régimbart are revised. P. pek-
inensis is described as new. Most primary types have been studied. P. wui Gschwendtner is con-
sidered a junior synonym of P. coomani Peschet. Lectotypes are designated for P. coomani
Peschet, P. intermedius (Sharp), P. variabilis (Clark) and P. wui Gschwendtner. A key to the
species is provided, as are distribution-maps.
B. J. van Vondel, Roestuin 78, 3343 CV Hendrik Ido Ambacht, The Netherlands.
Key words. — Peltodytes, palaearctic region, oriental region, new species.
Since Zimmermann (1924) published his key to
the Haliplidae of the world, only little attention has
been payed to the East Palaearctic and Oriental spe-
cies of Peltodytes, with only four new species described
since. Doubts about the status of these species war-
ranted a revision. The type material of most nominal
taxa was studied, but the identity of some others was
sufficiently established. Unfortunately a few types
could not be located.
Female genitalia are not described, because they
usually appear very uniform. Further research is nec-
essary to check if there are characters sufficiently reli-
able to separate the species.
MATERIAL
About 1700 specimens, including type-material
were studied from the following collections:
British Museum (Natural History), London, UK
(BMNH); B.P. Bishop Museum, Honolulu, Hawaii,
USA (BpBM); Fries Natuurhistorisch Museum,
Leeuwarden, Netherlands (FnML); IES Laboratory,
Hebrew University of Jerusalem, Israel (Hui);
Institut Royal des Sciences Naturelles, Brussels,
Belgium (tsNB); Museum National d'Histoire
Naturelle, Paris, France (MNHN); Museon, The
Hague, Netherlands (Mog); Naturhistorisches
Museum Wien, Vienna, Austria (NHMV); Natuurmu-
seum Rotterdam, Rotterdam, Netherlands (NHMz);
Oberösterreichisches Landesmuseum, Linz, Austria
(OLML); Instituut voor Bos- en Natuuronderzoek
(Rijksinstituut voor Natuurbeheer), Leersum,
Netherlands (RIN); Nationaal Natuurhistorisch
Museum (Rijks Museum van Natuurlijke Historie),
Leiden, Netherlands (RMNH); Instituut voor Taxo-
nomische Zoölogie (Zoologisch Museum), Amster-
dam, Netherlands (ZMAN); Zoologiska Museet,
Helsinki, Finland (ZMH); Museum für Naturkunde
der Humboldt-Universität, Berlin, Germany (zMHB);
Universitetets Zoologiske Museum, Copenhagen,
Denmark (zmuc); Zoologische Staatssammlung,
Munich, Germany (zsmc).
private collections:
A. L. van Berge Henegouwen (Zoetermeer), J. G.
M. Cuppen (Ede), M. B. P. Drost (Wadenoijen), Th.
G. Giessen (Gaanderen), M. Hielkema (Gouda), J.
Huijbregts (Leiden), Y. Jongema (Wageningen), P.
Kanaar (Leiderdorp), N. Nieser (Tiel), J. C. P. M.
van de Sande (Nijmegen), M. Schilthuizen (Leiden),
M. Schreijer (Alkmaar), H. J. Vallenduuk (Boxtel),
B. J. van Vondel (Hendrik Ido Ambacht), all in The
Netherlands.
METHODS
Dissecting, preparing and drawing the aedaegi fol-
lows the methods described by Vondel (1991: 76).
Morphological terms are exemplified in figs. 1 and 2.
The Palaearctic and Oriental regions are treated
here together because some species are common to
both regions. Peltodytes is not known from the
Australasian region east of Sulawesi.
Locality-names are spelled, as far as possible, ac-
cording to the Times Atlas of the World
(Comprehensive edition, 1983), but when different
from original labels, the modern name is given in pa-
rentheses. Names of countries are used with their
present-day boundaries.
25)
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
ON === DD UD D
WNT”
VE BWN:
Fig. 1. Peltodytes caesus, dorsal view. — an, antenna; as, apical spur; bp, location of basal punctures, illustrated for each species;
cl, clypeus; el, elytron; fr, frons; i.1, 1.2, 1.3, etc, first, second, third etc. interval; pr, pronotum; p.1, p.2, p.3, etc. first, sec-
ond, third, etc puncture-row; ss, sutural stria; st, setiferous striole; su, suture; ve, vertex.
Fig. 2. Peltodytes caesus, ventral view. — cp, metacoxal plate; ep, elytral epipleuron; mp, metasternal process; pe, pronotal epi-
pleuron; po, proepisternum; pp, prosternal process; ps, prosternum; s.5, fifth sternite; s.6, sixth sternite; s.7, seventh (last)
sternite.
Distribution maps are based on material examined.
Reliable literature records are included using different
symbols.
Notes on the biology are mainly based on literature
and the authors experience.
SYSTEMATIC SECTION
Genus Peltodytes Régimbart
Peltodytes Régimbart, 1878: 450. Type-species: Dytiscus cae-
sus Duftschmid, 1805, by subsequent designation
(Balfour-Browne 1936: 68).
Cnemidotus sensu Erichson, 1832: 48 (nec Cnemidotus
Illiger, 1802: 373). [Type-species not validly fixed
(Holmen, 1987: 86)].
Neopeltodytes Satô, 1963a: 21. Type-species: Peltodytes callo-
sus (Leconte), by original designation [see note below].
Cnemidotus, Aubé 1836: 36, Seidlitz 1887: 35.
Peltodytes, Zimmermann 1920: 298, Zimmermann 1924: 6,
Balfour-Browne 1936: 68, 1938: 5, 1940: 157, Guignot
276
1947: 34, Zaitsev 1953: 32, Satô 1963a: 21, 1963b: 23,
Franciscolo 1979: 67, Holmen 1987: 86, Beutel &
Ruhnau 1990: 4.
Remarks
Satö referred to the division of Peltodytes into two
groups by Zimmermann (1924) and defined the sub-
genus Neopeltodytes by the absence of the posterior
teeth on the metacoxal plates as present in the species
of Peltodytes s. str. Zimmermann used another char-
acter: an in the middle impressed metasternal process
in group I (= Peltodytes s. str. in Palaearctic, Ethiopean
and Oriental regions) or a not impressed metasternal
process in group II (=Neopeltodytes, in Nearctic and
most northern part of Neotropical regions).
Assignment of Peltodytes-specimens to one of the
subgenera meets some problems as there are Nearctic
species having a weak tooth on the metacoxal plates
or a more or less impressed metasternal process, while
some Palaearctic species sometimes show hardly de-
veloped teeth on the metacoxal plates. À more exten-
sive character-set is needed to support the subgeneric
division.
The phylogenetic position of Peltodytes is discussed
by Beutel & Ruhnau (1990).
The genus Peltodytes is represented in all faunal re-
gions, excepting the Australasian and the Neotropical
regions apart from Mexico and Cuba (Roberts 1913,
Zimmermann 1924, Guignot 1959).
Differential diagnosis
The genus Peltodytes is defined by the following
combination of characters:
Head: Labial and maxillar palpi with ultimate seg-
ment clearly longer than penultimate segment.
Behind the eyes only one genal line (fig. 51).
Elytra: along suture a stria over at least posterior
half. All puncture-rows relatively equal in strength,
no weaker secondary puncture-rows present. If inter-
mediate (secondary?) puncture-rows are present,
these are restricted to apical part and about as strong
as adjacent rows. Apical part of suture incised, expos-
ing inner margin of suture.
Underside: Epipleura reaching last abdominal seg-
ment. Metacoxal plates reaching at least sixth sternite,
laterally bordered to halfway posterior edge. Posterior
edge of each plate produced into a strong tooth, ex-
cept or hardly in nearctic species.
The wings were figured by Griffini (1897).
Sexual characters: Males have the first two tarsal
segments of fore and midlegs widened and ventrally
provided with sucker-hairs.
Within the Haliplidae Peltodytes can be recognized
by means of the following key.
7 to the genera of Haliplidae:
Last segment of palpi longer than penultimate
segment (fig. 6 and 7). Metacoxal plates at least
covering part of sixth abdominal sternite (fig. 2)
REN LE RAT Peltodytes Regimbart
— Last segment of palpi clearly shorter than penul-
timate segment. Last three abdominal sternites
freelyavisible a LR MU Lone 2
2. Head as wide as pronotum. Elytra strongly punc-
tured, not in rows. … …. Algophilus Zimmermann
— Head not as wide as pronotum. Elytra with clear
LONS CEN 3
3. Pronotum almost square, front-corners strongly
rounded. Elytra with or without longitudinal
ridges in first six intervals. ...... Brychius Thomson
— Pronotum widest at base, sides converging anteri-
orly. Elytra without longitudinal ridges in first six
IME val RR Haliplus Latreille
OL ahah ed eee te dl Apteraliplus Chandler
VAN VONDEL: Palaearctic Peltodytes
Note: Algophilus is restricted to Southern Africa.
Apteraliplus is only known from North America with
a single species: Apteraliplus parvulus (Roberts). The
separate generic status of the latter is subject to fur-
ther research.
Checklist of palaearctic and oriental Peltodytes
Peltodytes Régimbart, 1878
caesus (Duftschmid, 1805)
?curculinus (O. F. Müller, 1776)
quadrimaculatus (Drapiez, 1820)
caesus var. levantinus (Sahlberg, 1902a)
delhermi (Mazéret, 1923)
[zmpressus Fabricius sensu Panzer,
Fabricius, 1787. Misidentification]
coomani Peschet, 1923
wui Gschwendtner, 1934 syn. n.
dauricus Zimmermann, 1924
intermedius (Sharp, 1873)
rotundatus (Aubé, 1836)
conifer (Seidlitz, 1887)
pekinensis sp. n.
sinensis (Hope, 1845)
variabilis (Clark, 1863)
koreanus Takizawa, 1931
sumatrensis Régimbart, 1885
1794; nec
Key to species
1. Margins of pronotum narrow, not thicker than
antennae, hind corners hardly protruding (fig. 3)
- Margins of pronotum broad, clearly thicker than
antennae, hind corners strongly protruding (fig.
Ohne 7
2. Hind margin of metacoxal plates with a weak
bluntitooth{(fig4l0) Re 3
- Hind margin of metacoxal plates with a sharp
longatooth 325). ee ee 4
3. Fourth elytral puncture-row complete or at most
slightly interrupted in the middle. Base of prono-
tum strongly impressed, in the impression a
transverse row of strong darkened punctures. Not
with basal marks on pronotum, larger than dark-
ened punctures. Vertex at most with two vague
marks (fig. 3). Males: surface of apical parts of
Parameres(hauny (ii P. caesus
— Fourth elytral puncture-row only clearly present
in basal part, represented by about 3-5 punctures.
Base of pronotum hardly impressed, on both
sides of impression two joined strong punctures
and about three small ones, connected by a dark
mark. Vertex with two small distinct marks (fig.
74). Males: surface of apical parts of parameres
areolaten(f1 NS 4) A P. sinensis
4. Antennae slender, segments longer than wide
(ISDN EME nd 5
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8 à ;
7
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278
— Antennae short, segments 2-9 about as long as
wide (fig. 62). Males: surface of apical parts of
parameres hairy, (fig #15) P. rotundatus
5. Punctures on elytral epipleura black. Prosternal
process clearly bordered in anterior half (fig.41).
Males: surface of apical parts of parameres areo-
[ateiktie 48). er ne CAE RAR. RAR 6
— Punctures on elytral epipleura not black.
Prosternal process usually not completely and
clearly bordered in anterior half (fig. 30). Surface
of apical parts of parameres hairy (fig. 37)...........
REN NR AN eee P. dauricus
6. Pronotum without black punctures in posterior
corners between margin and widened basal punc-
tures (fig. 49). Proepisternum without strong
dark punctures, when very weak punctures
present these are not darkened. Teeth of coxal
plates usually long but with a blunt point, espe-
cially in lateral view (fig. 55 and 56). Head with a
black collar behind eyes (fig. 51)......P. pekinensis
— Pronotum with black punctures in posterior cor-
ners between margin and widened basal punc-
tures (fig. 38). Proepisternum with strong dark-
ened punctures. Teeth of coxal plates long and
sharp (fig. 43 and 44). Head without a black col-
lar directly behind eyes. … … … … … P. intermedius
7. Elytra with large dark marks and usually with
black band connecting basal punctures. Two
clear sharp dark spots on vertex (fig. 85). … … ….
EEN P. sumatrensis
— Elytra with at most a few small marks and base
usually not black. Vertex with at most two vague
SPOOLS} (ipa IG) e rer es P. coomani
DESCRIPTION OF SPECIES
Peltodytes caesus (Duftschmid)
(figs. 3-15)
? Dytiscus curculinus O. F. Müller, 1776: 73. Syntypes:
[Denmark or Norway]. [probably destroyed].
Dytiscus caesus Duftschmid, 1805: 284. Syntypes: Austria.
Typematerial probably in OLML [not examined].
Haliplus quadrimaculatus Drapiez, 1820: 349. Location of
types unknown [not examined].
Cnemidotus caesus var. levantinus Sahlberg, 1902-1903a: 17.
Syntypes: 1 ex., Israel, Jaffa, J. Sahlberg; 6 ex., Greece,
Corfu, J. Sahlberg (ZMH)(Bistrôm 1987: 33) [not exam-
ined].
Cnemidotus delhermi Mazéret, 1923: 67. Syntypes: France,
Armagnac, étangs à Espas, L. Mazéret. Location of types
unknown [not examined].
VAN VONDEL: Palaearctic Peltodytes
(Dytiscus impressus sensu Panzer 1794: t. 7. nec Fabricius,
1787. [Misidentification]. Lectotype ® of D. impressus
Fabricius, which is identical to Haliplus ruficollis (De
Geer, 1774) was designated by Holmen (1987: 133)].
Peltodytes curculinus, Zimmermann 1920: 299, 1924: 7.
Cnemidotus impressus, Seidlitz 1887: 35.
Peltodytes impressus, Zimmermann 1920: 299, 1924: 7.
Dytiscus caesus, Erichson 1832: 48.
Peltodytes quadrimaculatus, Zimmermann 1920: 299, 1924:
Us
Hatiplus caesus, Gyllenhal 1827: 234.
Cnemidotus caesus, Aubé 1836: 38, Régimbart 1877: 354,
Seidlitz 1887: 35.
Peltodytes caesus, Bedel 1881: 223, Escalera 1914: 56,
Zimmermann 1920: 299, 1924: 7, Balfour-Browne
1940: 157, Csiki 1946: 571, Guignot 1947: 36, 1959:
21, Lagar Mascaro 1951: 76, Zaitsev 1953: 34, Freude
1971: 9, Hosseinie 1974: 242, 1978: 167, Ienistea 1974:
204, Minoransky & Dzumailo 1974: 26, Ali 1976: 91,
Galewski 1976: 8, Konev 1976: 56, Mateleshko 1977:
68, Franciscolo 1979: 70, Guéorguiev 1958: 44, 1963:
215, 1981: 399, Foster 1981: 4, Angelini 1984: 48,
Holmen 1987: 88, Zakharenko & Moroz 1988: 282,
Belyashevskiy 1989: 124, Leblanc 1990: 1.C.3.
Cnemidotus caesus var. levantinus, Sahlberg 1902-1903b: 6.
Peltodytes caesus var. levantinus, Zimmermann 1920: 299,
1924:7 7; Guignot ls) 21,2 Zaitscvan 9 53:13);
Franciscolo 1979: 71.
Peltodytes caesus ssp. levantinus, Guéorguiev 1958: 44.
Remarks
P. caesus was described from Austria. Since no ot-
her species of Peltodytes are known from Austria, the
description by Duftschmid leaves hardly any doubt
about the identity of the species.
Schönherr (1808: 26) gives Dytiscus curculinus O.
F. Miiller as a synonym of D. impressus Fabricius,
which is not conspecific with P. caesus. Zimmermann
(1920) gives it again (with a questionmark) as a pos-
sible synonym of P. caesus. As Müllers’ collection does
not exist anymore it seems impossible to settle the
identity of this name.
Description
Length 3.3-3.8 mm, width 1.7-2.0 mm. Body sub-
parallel to parallel, widest in the middle (fig. 3).
Head: Yellow to brown, two large, sometimes
vague, dark blotches between eyes, sometimes conflu-
ent with slight darkening on vertex. Moderately
strongly and densely punctured, punctures on vertex
darkened. Distance between eyes 0.9-1.0X width of
one eye. Antennae yellow to yellow-red, segment 4-6
about as long as wide (fig. 4). Palpi yellow to yellow-
red.
Figs. 3-15. Peltodytes caesus, Netherlands. — 3, dorsal view; 4, antenna; 5, punctures near elytral base and suture; 6, maxillair
palpus; 7, labial palpus; 8, prosternal process; 9, lateral view of prosternal process; 10, metacoxal plate; 11, lateral view of met-
acoxal plate; 12, left paramere; 13, penis; 14, right paramere; 15, surface of apical part of parameres.
279
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N My
Figs. 16-26. Peltodytes coomani, lectotype. — 16, dorsal view; 17, antenna; 18, punctures near elytral base and suture; 19, pros-
ternal process; 20, lateral view of prosternal process; 21, metacoxal plate; 22, lateral view of metacoxal plate; 23, left paramere;
24, penis; 25, right paramere; 26, surface of apical part of parameres.
280
Pronotum: Yellow. Along base a row of widened
darkened punctures, of which strongest ones are op-
posite third elytral puncture-rows. Rest of pronotum
with sparse uncoloured punctures. Base impressed.
Lateral margin finer than antennae, not narrowed an-
teriorly, straight with convex anterior part.
Elytra: Yellow, darkened suture, discal, sometimes
vague, mark not connected to suture, vague marks on
intervals sometimes missing. Margined completely,
posteriorly not clearly concave. Posterior part not
bulbous, evenly rounded as rest of elytra. Punctures
moderately strong, about 30 punctures in first row.
Posteriorly about 6-15 punctures in second interval.
Fourth row in middle not or over a distance of at
most 5 punctures interrupted. Puncture-rows in sev-
enth and ninth interval stretched out over about pos-
terior half or even exceeding the middle. Basal punc-
tures widened( fig. 5). All punctures darkened.
Sutural striae stretched out over posterior 2/3.
Underside: Yellow to yellow-red, legs yellow to
brown, darkened towards coxae. Epipleura with un-
coloured punctures, reaching to 1/4 of last sternite.
Prosternal process posteriorly grooved in the middle
and with marginal plicae, strongly and coarsely punc-
tured, anteriorly hardly margined (fig. 8 and 9).
Metasternal process strongly grooved in the middle,
hardly punctured, plicae along coxae (fig. 8).
Metacoxal plates with a blunt tooth, just reaching last
sternite, sparsely punctured (fig. 10 and 11). Last
sternite unpunctured, strongly impressed in front
corners. Setiferous striole on dorsal face of hind tibia
about 1/3X length of tibia, longest of two tibial spurs
3/4X length of first tarsal segment.
Male: Penis and parameres as in figs. 12-14, surface
of apical part of parameres hairy (fig. 15).
Immature stages: Third stage larvae described by
Schiodte (1872: 203).
Biology
In fresh or occasionally brackish stagnant water of
ponds or ditches and slowly running streams. Among
vegetation of Ranunculus, Callitriche, Elodea, Chara
and other waterplants. Feeding on filamentous algae
and probably Characeans. The eggs are laid on the
surface of waterplants. The adults are attracted to
light (Konev 1976 and pers. obs.).
Distribution (fig. 96)
Europe northward to southern England, Denmark
and southern Sweden, eastward to Belorussia and the
Ukraine, southward to Portugal, Spain, Italy and
Greece. Asia: Turkey, Kazakhstan, southern part of
the Russian Federation, Afghanistan, Iran, Syria,
Israel. Africa: Morocco. Alı (1976) records a
Peltodytes sp. from Irak. It probably concerns P. cae-
SUS.
van VONDEL: Palaearctic Peltodytes
Material examined. — Albania: 2 ex. (NHMv) ® Austria: 76
ex. (NHMV, RMNH, ZMAN) ® Belgium: 1 ex. (Lodewijcks).-
Czechoslovakia: 3 ex. (NHMv) + Denmark: 1 ex. (ZMAN) ®
France: 33 ex. (BMNH, MOG, NHMV, RMNH, ZMAN, Jongema,
Huijbregts, Vondel) + Germany: 16 ex. (NHMv) © Great
Britain: 3 ex. (ZMAN) ® Greece: 11 ex. (NHMV, ZMAN, Berge
Henegouwen) * Hungary: 6 ex. (NHMV, RMNH, ZMAN) ®
Iran: 3 ex. (BMNH) © Italy: 4 ex. (NHMV, Jongema) ®
Jugoslavia: 19 ex. (NHMV, ZMAN) ® Kazakhstan: 1 ex. (NHMV)
e Morocco: 10 ex. (BMNH) ® Netherlands: 750 ex., all prov-
inces + Poland: 9 ex. (NHMv, RMNH) ® Portugal 11 ex.
(RMNH, ZMAN, Nieser) * Rumania: 1 ex. (NHMv) ® Russian
Federation: 1 ex. (NHMv) ® Switzerland: 1 ex. (Jongema) ®
Turkey: 31 ex. (NHMV, RMNH) ® Ukraine: 6 ex. (NHMV,
RMNH).
Peltodytes coomani Peschet
(figs. 16 -26)
Peltodytes coomani Peschet, 1923: 178. Lectotype d (here
designated), Hoa Binh, Tonkin [Vietnam], V.VI.1922,
A. de Cooman, TYPE, Museum Paris, 1945, Coll. R.
Peschet, Peltodytes coomani n. sp. R. Peschet (MNHN) [ex-
amined].
Peltodytes wui Gschwendtner, 1934: 107. Lectotype d (here
designated), Canton [China], Collector C. F. Wu, Type
Gsch., Hal. 42, Coll. Gschwendtner, Peltodytes wui
Gsch., det. Gschwendt., Type. (OLML) [examined]. syn.
n.
Peltodytes coomani, Zimmermann 1924: 208.
Description
Length 3.2-3.5 mm, width 1.9-2.1 mm. Body wi-
de, subparallel in the middle (fig. 16).
Head: Yellow-brown to yellow-red, seen from
beside brown marks behind eyes, sometimes two
small vague spots on vertex. Weakly to moderately
strongly punctured. Distance between eyes 0.7-0.8x
width of one eye. Antennae yellow to yellow-brown,
last segment usually darkened at top, segment 3-6
longer than wide, segment 7-10 more than twice as
long as wide (fig.17). Palpi yellow to yellow-brown.
Pronotum: Yellow to yellow-brown, dark mark op-
posite third elytral puncture-rows, slightly impressed
between these marks. Lateral borders slightly to clear-
ly convex, margins from very strong posteriorly to
narrow anteriorly, posterior corner clearly protrud-
ing. Sparsely, but in marginal parts sometimes coarse-
ly punctured, in basal dark marks one very large and
sometimes a few smaller punctures recognizable.
Elytra: Yellow to yellow-brown, suture completely
dark, towards base slightly narrowed, usually dark
mark between first and second and between third and
fifth puncture-row, sometimes without marks.
Posterior part bulbous, covering margins when seen
from above. Apical part of margin concave. Puncture-
rows moderately strong, basal punctures widened and
connected by a slight impression (fig. 18). Additional
puncture-row in second interval restricted to about 6
punctures in posterior part. Fourth puncture-row on-
281
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 27-37. Peltodytes dauricus, figs. 27-33, holotype, figs. 34-37, Harbin. — 27, dorsal view; 28, antenna; 29, punctures near
elytral base and suture; 30, prosternal process; 31, lateral view of prosternal process; 32, metacoxal plate; 33, lateral view of
metacoxal plate; 34, left paramere; 35, penis; 36, right paramere; 37, surface of apical part of parameres.
282
ly represented by 1 to 4 punctures in basal part. No
recognizable additional puncture-rows in seventh and
ninth interval. In impression in front of bulbous apex
a clear puncture-row from suture to lateral margin.
All punctures darkened, sometimes marginal and epi-
pleural punctures not darkened.
Underside: Yellow to yellow-brown, legs yellow-
brown, darkened near coxae. Epipleura punctured,
punctures usually brown, not as dark as elytral punc-
ture-rows. Prosternal process grooved, groove inter-
rupted in posterior part, strongly punctured (fig. 19
and 20). Prosternum with uncoloured punctures, an-
teriorly only margined close to prosternal process.
Metasternal process slightly impressed anteriorly and
strongly posteriorly, weakly punctured (fig. 19).
Metacoxal plates with a weak tooth, posterior sutural
corner weakly margined, strongly punctured except
in sutural area (fig. 21 and 22). Setiferous striole on
dorsal face of hind tibia about 1/4X length of tibia,
longest of two tibial spurs about 2/3X length of first
tarsal segment.
Male: Penis and parameres as in figs. 23-25.
Surface of apical parts of parameres areolate (fig. 26).
Immature stages: Unknown.
Distribution (fig. 101)
Vietnam, China (prov. of Guangdong)
Material examined. — Vietnam: 1 d, lectotype, 8 paralec-
totypes with same labels as lectotype (4 in MNHN and 4 in
RMNH); 29 ex, Tonkin, Hoa Binh, leg. A. de Cooman
(dates, if present, varying from 1932 to 1934) (ISNB, RMNH,
MNHN); 3 ex., Annam, Reg. de An-ninh. R. P. M. Mannier,
dr. Régimbart vidit 1903, ex. coll. Oberthür ( 1 ex. labelled:
Peltodytes variabilis Clk.) (MNHN) ® China: 1 à, lectotype of
P.wui (OLML); 2 ex., Canton [Guangzhou], Mell S. V. (1 ex.
with date x.10)(ZMHB); 2 ex., Hainan Id., S. China, SW of
Nodoa, Tan-hsien (Distr.), 17.v11.1929.
University, Fifth Hainan Expedition (BPBM).
Lingnan
Peltodytes dauricus Zimmermann
(figs. 27-37)
Peltodytes dauricus Zimmermann, 1924: 8. Holotype 2 (by
monotypy), Nikolsk, Ussurijsk, Mandl, Type, Holotypus
Staatssamml. München (zsmc) [examined].
Peltodytes dauricus Wu 1932: 341, J. Balfour-Browne 1946:
436, Zaitsev 1953: 36 (as jun. syn. of intermedius).
Description
Length 4.2 mm, width 2.4 mm. Body wide, subpa-
rallel. (fig. 27).
Head: Yellow-brown, weakly punctured. Distance
between eyes 0.9X width of one eye. Antennae (fig.
28) and palpi yellow-brown.
Pronotum: Yellow-brown, basal dark marks oppo-
site third elytral puncture-row. Impressed between
basal marks. Weakly punctured, three or four strong
dark punctures in basal marks clearly smaller than el-
van VONDEL: Palaearctic Peltodytes
ytral basal punctures, about 15 dark punctures in im-
pression between basal marks. Lateral borders convex,
margins widest in the middle, not wider than anten-
nac.
Elytra: Yellow-brown, very strong puncture-rows,
although punctures are usually separately darkened,
the strong darkening gives the elytra a striped appear-
ance. Suture darkened, small marks between first and
second, between third and fifth and between eighth
and tenth row. Fourth row missing about five punc-
tures in the middle. Additional puncture-rows
present in posterior half of seventh and ninth interval.
No or only few additional punctures in apical part of
second interval. Basal punctures in a common trans-
verse impression (fig.29). All punctures ringlike im-
pressed: centre of puncture is slightly raised. Very
small punctures along suture. Sutural stria in posteri-
or half.
Underside: Yellow-brown to brown, legs yellow-
brown to brown around coxae. Elytral epipleura yel-
low-brown, weak punctures uncoloured. Prosternal
process strongly narrowed near coxae, in anterior part
usually without a clear margin, with various impres-
sions and coarse punctures (fig. 30, 31). Prosternum
margined anteriorly, except near epipleura, at most
very weakly and sparsely punctured. Metasternal pro-
cess flat with a pit in the middle, sparsely punctured,
the part covering the mesocoxae separated by a clear
plica (fig. 30). Metacoxal plates with a strong but not
very sharp tooth (fig. 32, 33). Setiferous striole on
dorsal face of hind tibia about 1/5X length of tibia,
longest of two tibial spurs about 2/3X length of first
tarsal segment.
Male: Penis and parameres as in figs. 34-36. (illus-
trated for the first time by J. Balfour-Browne (1946).
Surface of apical parts of parameres hairy (fig. 37).
Immature stages: Unknown
Distribution (fig. 98)
Russian Federation near Wladiwostok, China
province Heilongjiang.
Material examined. — Russian Federation: ® Holotype
(zsmc) © China: 1 d, Weishaha [Weihe], Manch,
20.vii.1939, M. Weymarn; 3 ex., Hsiaoling [?], Manchuria,
vii.1939, M. Weymarn; 1 ex., Harbin, Manchuria,
22.ix.1940, M. Weymarn, Peltodytes dauricus Zimm., J.
Balfour-Browne det. 1944; 4 ex., Charbin [Harbin], 5.-
viii.1952, BM.1953-715; 3 ex., Charbin [Harbin],
DAI 52 B MAS 3715 2.) ex ME lUnekiant
[Heilongjiang], Harbin, 13.vi.1965 & 26.vi.1966, P.M.
Hammond, BM 1967-215 (BMNH).
Peltodytes intermedius (Sharp)
(figs. 38-48)
Cnemidotus intermedius Sharp, 1873: 55. Lectotype ® (here
designated), “Type [round label with red margin], Japan
283
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6
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©
©
38, dorsal view; 39, antenna; 40, punctures near elytral base and su-
Figs. 38-48. Peltodytes intermedius, Japan: Yokoshima. —
; 43, metacoxal plate; 44, lateral view of metacoxal plate; 45,
left paramere; 46, penis; 47, right paramere; 48, surface of apical part of parameres.
ture; 41, prosternal process; 42, lateral view of prosternal process
284
Lewis [label with yellow middleband], Japan [yellow oval
label], Sharp coll. 1905-313, Cnemidotus intermedius type
DS. [according to description abundant at Nagasaki]
(-BMNH) [examined].
Peltodytes intermedius, Régimbart 1899: 191, Peschet 1923:
179, Zimmermann 1919: 75, 1920: 300, 1924: 7,
Takizawa 1931: 138, Wu 1932: 341, Zaitsev 1953: 36,
Nakane 1963: 55, 1987: 27, Satô 1963b: 23, 1984: 3,
Yoon 1988: 620.
Description
Length 3.2-3.7 mm, width 1.9-2.2 mm. Body wi-
de, subparallel, widest in the middle. (fig. 38)
Head: Yellow to yellow-brown, moderately strong-
ly and densely punctured. Sometimes with a slightly
darkened collar, which does not reach the eyes.
Distance between eyes 0.8-1.0X width of one eye.
Antennae (fig. 39) and palpi yellow.
Pronotum: Yellow, basal dark marks opposite fifth
elytral puncture-row, containing 2 or 3 widened
punctures. Slightly impressed between basal marks.
Strong darkened punctures along base and in hind
corners, weak not or only sligthly darkened punctures
in anterior and discal part. Lateral borders straight to
convex, margin not wider than antennae, narrowed
anteriorly.
Elytra: Yellow, suture darkened, four larger and
some small dark marks on each elytron, of which dis-
cal one is separated from suture. Puncture-rows
strong. Fourth puncture-row widely interrupted, usu-
ally only one to three punctures remaining near base
and at most about five apically. Basal punctures of
first five rows strongly widened. Apically about 10-15
punctures in second interval. Apically remains of ad-
ditional puncture-rows in seventh and ninth interval.
All punctures darkened. Apical margin not clearly
concave. Completely margined. Striae along suture in
posterior half. Along suture weak and dense darkened
punctures. Apical part slightly bulbous.
Underside: Yellow to yellow-red, legs yellow to yel-
low-red, darkened towards coxae. Elytral epipleura
yellow, anterior part with dark punctures, reaching
middle of last sternite. Prosternal process narrowed
near coxae, moderately strongly punctured, slightly
impressed in the middle, posteriorly with a central
impression and a punctured stria on each side, anteri-
orly weakly margined (fig. 41, 42). Metasternal pro-
cess with deep pit in the middle, hardly punctured,
weak striae along coxae (fig. 41). Metacoxal plates
with small sharp tooth, not reaching last sternite, in
lateral view also sharp, posterior part between teeth
bent down to the body, densely punctured, all punc-
tures about equal in strength and uncoloured (fig. 43,
44). Setiferous striole on dorsal face of hind tibia 1/2-
1/3X length of tibia, longest of two tibial spurs about
3/4X length of first tarsal segment. Last sternite un-
punctured, impressed beside the middle.
Male: Penis and parameres as in figs. 45-47.
van VONDEL: Palaearctic Peltodytes
Surface of apical part of parameres areolate (fig. 48).
Immature stages: According to Bertrand (1972)
the larva was described by Fukuda, Kurosa and
Hayashi (publication unknown to me).
Biology
Found in ponds and in a river.
Distribution (fig.99)
Japan, China, Korea. Zimmermann (1919) gives
Taiwan, Zaitsev (1953) gives the most eastern part of
the Russian Federation.
Material examined. — Japan: Kyushu: Lectotype (BMNH);
4 ex., Nagasaki, Hilsendorf (zMHB); 10 ex., Ikenosono,
Konyama T, Kagoshima Pref., 6.x.1985, leg. E. Matsui; 10
ex., Yokoshima T, Taman-gun, Kumamoto Pref., 9.vi.1986,
leg. E. Matsui (Vondel); Tsushima: 1 ex., P. Holst; 1 ex.,
Tsu-sima, vii-x.1891 (BMNH); Honshu: 4 ex., Tokio, 25-
27.1x.1881, G. Lewis; 1 ex., Yokohama,1.i-9.ii.1881, G.
Lewis (BMNH); 1 ex., Kobe, 20.vii.1916, leg. J. E. A. Lewis
(-BPBM); 3 ex., Surr. Tokio, Bodemeyer (RMNH); 2 ex.,
Tokio; 8 ex., Yedo [river Edo], vii.1875, leg. Hilsendorf
(zMHB); Hokkaido: 1 ex, Sapporo, 5-16.viii.1880, G. Lewis
(BMNH); 5 ex., Obihiro, 24.vii.1906, H. Schoede S. G.
(zMHB); 1 ex., Hakodate, 1897, R.P. Faurie (MNHN); Japan
(Island unknown): 1 2, Japon, Det. Sharp, Lewis, inzerme-
dius, det. Sharp 1873, Peltodytes intermedius Shp, Type
(-ISNB); 2 ex., Okitsu [?], iv+v.1913, F. Muir (BPBM); 7 ex.,
Chiuzenji [?], 19-24.viii.1881, G. Lewis; 15 ex., Chiuzenji,
G. Lewis; 1 ex., Chuzai, Chiuzenji, G. Lewis; 2 ex., Sarsiew
[?], 20.ix.1909, J.E.A. Lewis; 25 ex., Lewis, 15.ix.1881 or
29.x.1881; 8 ex., G. Lewis, (some 9.11.1881); 1 ex, G.
Lewis; 1 ex., 1896, Lord Dormer (BMNH); 2 ex., Hiller S.; 2
ex, Hilsendorf (zMHB) ® China: 1 ex., 5. Shanghai (RMNH);
1 ex., Peping [Beijing], C.F. Wu (oLML) ® Korea: Kangwon-
do: 3 ex, Wonsong-kun, Kyangkyok, pond, 3.viii.1988;
Chungpuk-do: 1 ex, Chunchon-city, Chiljon-dong, 5.-
viii.1988; 3 ex., Chungwon-kun, Chisil-pond, 16.viii.1987;
4 ex., Poun-kun, Chisan-ri, 17.viii.1989; 3 ex., Sintanjin-
city, 21.vii.1987; Kyongnam-do: 2 ex., Yangson-kun,
Taesok-pond, 22.x.1988; 2 ex., Chinryang-kun, Kumsan-
pond, 11.viii.1989; Kyongki-do: Kwangju-kun, Chungang-
pond, 20.viii.1989; Chonnam-do: 2 ex., Imsil-kun,
Samgye-ri, 14.viii.1989; 5 ex., Namwon-kun, Okrim-pond,
13.viii.1989; Kyongpuk-do: 3 ex., Yongpung-kun, Hway-
on-pond, 2.viii.1988 (Vondel, ex coll. Sung Hwa Lee).
Peltodytes pekinensis sp. n.
(figs. 49-60)
Type material. — Holotype d, “Peping [Beijing],
Collector C. F. Wu, coll. Gschwendtner, intermedius Shp,
det. Gschwendtner” (OLML). — Paratypes, Id, 12: 1 9,
data as holotype (coll. Vondel); 1 d, “China, Peking
[Beijing], vii.[19]13 (ZMHB)
Description
Length 3.6-3.9 mm, width 2.1-2.2 mm. Body wi-
dest just behind shoulders, in the middle subparallel
(fig. 49).
Head: Yellow to yellow-brown, dark mark on neck
reaching downwards to halfway height of eyes (fig.
285
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
seeaeof
5 6 Su
Way
KE
N \ i \
peren,
Figs. 49-60. Peltodytes pekinensis, Holotype. — 49, dorsal view; 50, antenna; 51, lateral view of head; 52, punctures near ely-
tral base and suture; 53, prosternal process; 54, lateral view of prosternal process; 55, metacoxal plate; 56, lateral view of met-
acoxal plate; 57, left paramere; 58, penis; 59, right paramere; 60, surface of apical part of parameres.
286
51). Strongly and densely punctured, slightly im-
pressed behind front margins near antennae.
Distance between eyes 0.6-0.8X width of one eye.
Antennae yellow to yellow-brown, top of last segment
slightly darkened (fig. 50). Palpi yellow-brown to
brown.
Pronotum: Yellow. Lateral borders convex in ante-
rior third part, clear margins not wider than anten-
nae, slightly narrowed anteriorly. Sparsely punctured,
basally opposite each fifth elytral puncture-row one
strong deep puncture together with two or three
smaller punctures included in a dark mark, impres-
sion between these marks with few sometimes slight-
ly darkened punctures.
Elytra: Yellow, usually small marks in second and
fourth interval, dark suture narrowed towards base.
Strong puncture-rows, basal punctures of row 1, 2, 3
and 5 widened and connected by a weak impression
(fig. 52). Additional posterior puncture-row in sec-
ond interval reaching about as far as the middle, addi-
tional posterior puncture-rows in seventh and ninth
interval clearly present. Sutural stria in posterior half
reaching the middle, along suture small sparse punc-
tures.
Underside: Yellow, legs yellow, hind femura, coxae
and tarsi of all legs yellow-brown to brown.
Prosternal process narrowed near coxae, sharp halfcir-
cular impression at posterior margin, strongly and
coarsely punctured (fig. 53 and 54). Metasternal pro-
cess grooved and with a pit in the middle, weakly and
sparsely punctured (fig. 53). Metacoxal plates with a
strong blunt tooth easily reaching last sternite, tooth
in lateral view also strong and blunt, suture of coxal
plates posteriorly only bending weakly towards body
(fig. 55 and 56). Last sternite unpunctured. Epipleura
with clear darkened punctures. Setiferous striole on
dorsal face of hind tibia about 1/4X length of tibia,
longest of two tibial spurs about 3/4X length of first
tarsal segment. Male: Penis and parameres as in figs.
57-59. Surface of apical parts of parameres areolate
(fig. 60).
Immature stages: Unknown
Distribution (fig. 98)
Only known from Beijing, China.
Peltodytes rotundatus (Aube)
(figs. 61-73)
Cnemidotus rotundatus Aubé, 1836: 40. Syntypes: Southern
France and Italy (probably in MNHN) [not examined].
Cnemidotus conifer Seidlitz, 1887: 35. Syntypes: Sicily.
(probably in zsmc) [not examined].
Cnemidotus rotundatus, Seidlitz 1887: 35.
Peltodytes rotundatus Zimmermann 1920: 300, 1924: 8,
Falkenström 1939: 2, Guignot 1947: 37, 1959: 23, Lagar
Mascaro 1951: 76, Zaitsev 1953: 36, Freude 1971: 10,
VAN VONDEL: Palaearctic Peltodytes
Franciscolo 1972: 63, 1979: 70, Ienistea 1974: 204,
Angelini 1984: 48, Ferreras Romero 1987: 20, Leblanc
1990: 1.C.5.
Peltodytes conifer, Escalera 1914: 56, Zimmermann 1920:
299}
Peltodytes rotundatus var. conifer, Zimmermann 1924: 8,
Guignot 1947: 37, 1959: 23.
Peltodytes rotundatus ssp. conifer, Zaitsev 1953: 37,
Franciscolo 1979: 70, Guéorguiev 1981: 400, Angelini
1984: 48.
Remarks
Aube’s original description of rotundatus, in which
the specific character of the sharp tooth on the meta-
coxal plate is mentioned, leaves no doubt about the
identity of this species.
The species or subspecies or variety conifer differs
from rotundatus in having weaker elytral punctures
and a less interrupted fourth elytral puncture-row.
These characters, however, are very variable and all
kind of intermediate forms may be met with.
Description
Length 3.3-3.9 mm, width 2.0-2.2 mm. Body wi-
de, subparallel, widest in the middle (fig. 61).
Head: Yellow to yellow-brown, vertex slightly
darkened. Moderately strongly punctured, on vertex
with strong dense darkened punctures. Distance
between eyes 0.7-1.1X width of one eye. Antennae
yellow, first six segments wider than long (fig. 62).
Palpi yellow to yellow-brown (fig. 64 and 65).
Pronotum: Yellow to yellow-red. Along base a row
of darkened punctures, of which the ones opposite
fifth elytral puncture-row are widened, base between
these widened punctures strongly impressed. Lateral
and anterior part with sparse uncoloured punctures.
Lateral margins slightly concave to slightly convex,
not wider than antennae.
Elytra: Yellow with or without vague marks on
intervals, narrowly darkened suture faded towards
base, sutural striae on about posterior half.
Completely margined, margin apically abruptly bent.
Puncture-rows weak to very strong, 24-28 punctures
in first row. Posteriorly between first and second row
usually no punctures, but occasionally some punc-
tures present, remains of additional rows on posterior
part of seventh and ninth interval present. Fourth
row widely interrupted, anterior part varying from 0-
4 punctures. Widened basal punctures in a clear
transverse impression (fig. 63). All punctures dark-
ened.
Underside: Yellow to yellow-red, legs yellow to yel-
low-red, coxae brown, elytral epipleura yellow with
darkened or uncoloured punctures. Prosternal pro-
cess narrowed near coxae, posteriorly impressed, lat-
erally with plicae formed by coarse punctures, anteri-
or side clearly margined, usually strongly punctured
287
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
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EN
Way en
SN
NS
70
71
Figs. 61-73. Peltodytes rotundatus, Portugal: Algarve. — 61, dorsal view; 62, antenna; 63, punctures near elytral base and su-
ture; 64, maxillair palpus; 65, labial palpus; 66, prosternal process; 67, lateral view of prosternal process; 68, metacoxal plate;
69, lateral view of metacoxal plate; 70, left paramere; 71, penis; 72, right paramere; 73, surface of apical part of parameres.
288
(fig. 66 and 67). Metasternal process with strong im-
pression in the middle, weakly punctured, plicae
along coxae (fig. 66). Metacoxal plates with long
sharp tooth, reaching last sternite, moderately strong-
ly punctured, near suture weakly punctured (fig. 68,
69). Last sternite unpunctured, apex with micro-
punctuation. Setiferous striole on dorsal face of hind
tibia 1/4X length of first tarsal segment.
Males: Penis and parameres as in figs. 70-72.
Surface of apical part of parameres hairy (fig. 73).
Immature stages: Third stage larvae described by
Bertrand (1951: 76)
Biology
Living between filamentous algae in lakes, pools,
ponds, rivers, marshes, brooks, streamlets. Among ve-
getation of Utricularia, Nuphar and other water-
plants. Attracted to light.
Distribution (fig. 97)
Mediterranean area: Portugal, Spain including the
Baleares, France including Corsica, Italy including
Sardinia, Jugoslavia, Albania, Greece including Crete,
Morocco. Escherich (1897) gives Turkey.
Material examined. — Albania: 3 ex, Elbasan,
10+15.ix.1918, Priesner (NHMv) ® France: 1 ex., Castelnau-
le-Lez; 1 ex., Toulouse; 1 ex., Biaritz; 1 ex., Chatellerault; 4
ex.,, Carcassonne; cx, Pau; lex, Gers; lex, Ripaud,
Aude, 12.vii.1973, Davidson, at light; 2 ex., Narbonne,
Etang de Leucate, 5.viii.1956, Eckerlein; 2 ex, Bruges, Pyr.
Atl., 27.vii.1990; 11 ex., Corsica, Porto, 8.vi.1976, stream
in alder-marsh nr sea, van Nieukerken; 3 ex., Corsica,
Ghisonaccia, Fium Orbo River, 23.vi.1976, van
Nieukerken; 8 ex., Corsica, Miomo, 5km N. of Bastia,
24.vi.1976, van Nieukerken; 2 ex., Lyon; 1 ex., Lac
d’Hourtin, Medoc, 28.ix.1973, Jongema; 1 ex., Near Uzès,
Gard, 11.vii.1987, Schilthuizen; 16 ex., from unknown lo-
cality (BMNH, NHMV, RMNH, ZMAN, Jongema, Schilthuizen,
Vondel) © Greece: 5 ex., Attika; 1 ex., Kreta [Kriti], Kroeses,
14.iv.1933; 1 ex., Morea, Parnon Mt.; 1 ex., Athinai; 33 ex.,
Lakonia, 5 km SE Sparti, 27.ix.1962; 4 ex, Ilia, Olympia, 2-
3.x.1962; 18 ex., Corfu [Kérkira], Sindari, brook,
15.vii.1972, Berge Henegouwen (NHMV, RMNH, ZMAN,
Berge Henegouwen) ® Italy: 4 ex., Sardinia, Nuoro, R.
d’Oliena nr. St Giovanni, 24.viii.1987, Berge
Henegouwen; 2 ex., Sardinia, Muravera, 24.v-20.vi.1965,
Budberg; 2 ex., Pieris, Friaul, Müller; 7 ex., Riva, Tirol; 1
ex., Genova; 1 ex., Lucania face F. Basento, 22.iv.1970,
Angelini; 2 ex., from unknown locality (NHMV, RMNH,
Berge Henegouwen) © Jugoslavia: 1 ex., Hercegovina,
Dracevo, v.1911; 4 ex, Castelnuovo [Herzeg Novi],
Dalmatia (NHMv) * Morocco: 1 ex., Defilia, nr. Figuig, 5-
20.iv.1966(BMNH); 1 ex., Tiznit, 27.vii.1959, Eckerlein
(NHMV); 1 ex., Haute Atlas, Tamanar, 100 km N. of Agadir,
25.iv.1961, Dorgelo (ZMAN) * Portugal: 1 ex., betw. Setubal
and Marateca, 18.ix.1976, Berge Henegouwen;l ex., betw.
Palma and Marateca, 13.vii.1973, stagnant ditch with
Utricularia and Nuphar, muddy water, Berge Henegouwen;
1 ex., Palma, Rio San Martinho, 18.ix.1976, Berge
Henegouwen; 1 ex, Alentejo, Rio Mira, nr. S. Ana da Serva,
VAN VONDEL: Palaearctic Peltodytes
21.ix.1976, Berge Henegouwen; 5 ex, Algarve, 10 km S.
Sao Marcos da Serva, 21.ix.1976, Berge Hennegouwen; 7
ex., Alentejo, 15 km N. Santa Clara a Velho, 18.ix.1976,
Berge Henegouwen; 3 ex., Veiros, Riba d’Ana Laura,
25.1x.1976, Berge Henegouwen; 7 ex., Algarve, Guerenca,
Riba, Algibre, 16.iv.1981, Berge Henegouwen; 1 ex., Evora,
19.viii.1967; 1 ex., Evora, 4 km N. Arraiolos, 3.v.1986,
Kanaar; 6 ex., Evora, streamlet of Quintel, Pera Manca, 19.-
viii.1967, Nieser; 7 ex., Algarve, Quanteira, 24-29.xii.1967,
Nieser; 4 ex., Algarve, nr. Estof, 19.vii.1971, Nieser; 3 ex.,
Rio Antua, S. Estaroja, 22.iv.1976, Nieser (RMNH, ZMAN,
Berge Henegouwen, Kanaar, Nieser) * Spain: 4 ex., Cost
Brava, vi.1966 & vi.1967, Budberg; 1 ex., Llambillas,
Gerona, 4-5.viii.1956, Eckerlein; 7 ex., Posadas, 18-
19.iv.1925; 3 ex., Pollensa, Mallorca; 14 ex., Albufeira,
Mallorca (partly 15.vi.1913); 1 ex., Valencia; 1 ex., Palencia;
2 ex., Abadia, prov. Caceres, alt. 700 m., 5.v.1960; 1 ex., nr.
Rosal de la Frontera, Rio Chanca, stagnant pools in dry riv-
er course, vii.1973, Berge Henegouwen; 2 ex., Caceres,
Puerto Roque, Rivera David, 26.ix.1976; 1 ex., Baleares,
Pont de Giretto, S. Soller, 6.v.1983, Stock; 1 ex., Pyrenes,
San Jaume de Llierca, riv. Fluvia, 12.viii.1987, Vallenduuk
(NHMV, RMNH, ZMAN, Vallenduuk) + Unknown country: 9
ex. (NHMV, RMNH).
Peltodytes sinensis (Hope)
(figs. 74 -84)
Haliplus sinensis Hope, 1845: 15. Lectotype 9 (here desig-
nated), [China, Canton] Dr. Cantor Penang, Type,
60.15, E.I.C., Sinensis. Hope (BMNH) [examined]
Haliplus variabilis Clark, 1863: 417. Lectotype d (here des-
ignated), China, 2056, 6756, Cotype, Peltodytes variabi-
lis Clk (=sinensis Hope) (BMNH) [examined].
Peltodytes koreanus Vakizawa, 1931: 138. Syntypes: Korea:
18 ex., Suigen, 5-15.vii.1922, T. Uchida & S. Takano; 1
ex., Eitoho, 13.vi.1923, Yuuki. (not located) [not exam-
ined].
Peltodytes sinensis, Régimbart 1899: 192, Peschet 1923: 179,
Zimmermann 1920: 300, 1924: 7, Takizawa 1931: 139,
Wu 1932: 341, Gschwendtner 1934: 107, Falkenström
1936a: 2, 1936b: 79, Zaitsev 1953: 36, Satô 1960: 252,
1963b: 24, 1984: 3, Cho 1969: 165, Nakane 1987: 27,
Yoon 1988: 620.
Peltodytes variabilis Zimmermann 1920: 300, 1924: 7, Satô
1963b: 24.
Peltodytes koreanus, Satô 1963b: 24 (synonimised with P. si-
nensis), Yoon 1988: 620.
Description
Length 3.4-3.8 mm, width 1.9-2.2 mm. Body
oval, subparallel, widest in the middle (fig. 74).
Head: Yellow-brown to brown, two dark marks
between eyes, usually vaguely darkened behind eyes.
Moderately strongly punctured. Distance between
eyes 0.6-0.7X width of one eye. Antennae (fig. 75)
and palpi yellow to yellow-brown.
Pronotum: Yellow to yellow-red, dark marks oppo-
site fifth elytral puncture-rows, containing 2-4 wid-
ened strongly impressed punctures, between basal
marks impressed and with a few, sometimes darkened
punctures. In hind corners 2-5 darkened punctures,
else with uncoloured punctures, in front corners with
289
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
WU
AT
Figs. 74-84. Peltodytes sinensis, 74-80, holotype, 81-84, Korea.-74, dorsal view; 75, antenna; 76, punctures near elytral base
and suture; 77, prosternal process; 78, lateral view of prosternal process; 79, metacoxal plate; 80, lateral view of metacoxal
plate; 81, left paramere; 82, penis; 83, right paramere; 84, surface of apical part of parameres.
290
coarse punctures. Lateral margin not clearly wider
than antennae, narrowed anteriorly.
Elytra: Yellow to yellow-red, darkened suture, four
sometimes vague marks on each elytron, discal mark
separated from suture, sometimes a slight narrow
darkening along central part of base. Posterior part
slightly bulbous, apical margin at most slightly con-
cave. Margined completely. Punctures moderately
strong and all darkened. Additional posterior row on
second interval with 10-12 punctures. Fourth punc-
ture-row widely interrupted, about 3-5 punctures an-
teriorly and 1 or 2 posteriorly. Short additional poste-
rior puncture-rows in seventh and ninth intervals.
Basal punctures of especially fifth and sixth row wid-
ened, connected by a transverse impression (fig. 76).
Stria along posterior half of suture, in anterior half a
few small darkened punctures.
Underside: Yellow to yellow-red, legs yellow to yel-
low-brown, metafemur, metatarsi and all coxae dark-
ened. Elytral epipleura yellow with darkened punc-
tures, reaching to halfway last sternite. Prosternal
process narrowed near coxae, anteriorly and in poste-
rior part impressed, posteriorly with lateral striae,
strongly and sometimes coarsely punctured (fig. 77
and 78). Metasternal process with deep pit in the
middle, hardly punctured, clear striae along coxae
(fig. 77). Metacoxal plates with a dull tooth, reaching
last sternite, strongly punctured except in weakly
punctured sutural area (fig. 79 and 80). Last sternite
unpunctured, impressed near front corners.
Setiferous striole on dorsal face of hind tibia about
1/5X length of tibia, longest of two tibial spurs 3/4x
length of first tarsal segment.
Male: Penis and parameres as in figs. 81-83.
Surface of apical part of parameres areolate (fig. 84).
Immature stages: The third-stage larva is figured by
Yoon (1988).
Biology
In stagnant water, in rivers. Up to an altitude of
2100 m.
Distribution (fig. 100)
China, provinces Beijing, Fujian, Guangdong,
Henan, Hunan, Hupeh, Jiangsu, Jiangxi, Shanghai,
Sichuan, Yunnan, Zhejiang. Korea. Japan, Ryukyu
Islands. Vietnam. Taiwan. Philippines.
Material examined. — China: lectotype ® of Haliplus si-
nensis, lectotype of Haliplus variabilis, 2 paralectotypes of
Haliplus variabilis, same labels as lectotype; 1 ex.
Kwantung, Tai-ping, Lo-ting [Luoding, Guangdong],
6.vii.1932, W. E. Hoffman; 1 ex, Kwantung [Guangdong],
Yim-na, Shan [?], 1200 m., 15.vi.1936, L. Gressitt; 1 ex.,
Yunnan, Western Hills nr. Kunming, 2100 m., 7.vii.1940,
J. L. Gressitt; 1 ex., Hunan, Li, Tsinshih, T.C. Maa,
29.v.1937 (BPBM); 1 ex., Tali [Dali], Haut Yunnan; 1 ex.,
VAN VONDEL: Palaearctic Peltodytes
Nanking [Nanjing], v.1928, J.G. Needham; 3 ex., Yunnan
Fou [Kunming], H. Perrot; 5 ex., Ning-po [Ningbo,
Zhejiang], vii.1937 (isnB); 4 ex, Foochow [Fuzhou,
Fujian], vi.1935 & 1937-1938, M. S. Yang; 8 ex., Chusan
Isl. [Zhoushan], J. J. Walker; 2 ex., Honan, Loyang (Henan,
Luoyang], 6.viii.1966, P.M. Hammond (BMNH); 3 ex.,
Shanghai (zmuc); 3 ex., Yunnan-Sen, 1898, Excoffier; 1 ex.,
Kouy Tchéou [?], 1910, P.Cavaleria; 2 ex., Shanghai; 1 ex.,
Dohnu [2] (MNHN); 10 ex., Yunnan-Sen; 14 ex., Tsche-
kiang, Ning-po [Zhejiang, Ningbo]; 2 ex., Canton
(Guangzhou, Guangdong], Nordfluss, v.1909, Mell
(zMHB); 2 ex., Shanghai (RMNH) © Taiwan: 1 ex, Puli
(Mori), vi.1954, native collector; 35 ex., Taihoku,
4.iv+6.v+22.v.1928, F. C. Hadden (BrBM); 2 ex., Taipe
City, 16.viii.191, C. F. Lee (NHMV); 1 ex., Takau, Sauter
(isNB); 35 ex., Anping, iv.1910, Sauter (ZMHB) ® Korea:
7ex., Kyongki-do, Ichon-kun, Solsong-pond, 19.viii.1989;
6 ex, Chungpuk-do, Chongwon-kun, Doldari (stone-
bridge)-pond, 17.viii.1989; 1 ex., Chungpuk-do, Sintanjin-
city, 21.vii.1987; 2 ex, Chonnam-do, Yongam-kun,
Kumho-pond, 22.vii.1988; 2 ex., Kyongki-do, Kwangju-
kun, Chungang-pond, 20.viii.1989; 3 ex., Chonnam-do,
Yongam-kun, Taegan-pond, 22.vii.1988; 3 ex., Chonnam-
do, Imsil-kun, Samgye-ri, 14.viii.1989 (Vondel, ex. coll.
Sung Hwa Lee); 1 ex., Hede Quelpaert [?], 1908 (MNHN) ©
Japan (Ryukyu-Islands): 3 ex., Dana, Iheya Is., Okinawa
Pref., 20.viii.1989, leg. E. Matsui (Vondel).-Philippines: 5
ex., N. Luzon, Prov. Ifugao, Banaue, 22.vi.1988, K. Martin
(NHMv) ® Without locality: 1 ex., Boyer 800 (NHMy).
Peltodytes sumatrensis Régimbart
(figs. 85-95)
Peltodytes sumatrensis Régimbart, 1885: 55. Holotype ®:
‘Ed. Everts, (W. Dates), Medan, Deli, Sumatra;
Peltodytes sumatrensis Regb., type, Type; Museum Leiden,
Peltodytes sumatrensis Reg. (RMNH) [examined].
Peltodytes sumatrensis Peschet 1923: 179, Zimmermann
1920: 300, 1924: 9, 1927: 1.
Description
Length 3.4-3.7 mm (in original description ero-
nously given as 4.75 mm), width 1.9-2.1 mm. Body
subparallel, widest in the middle (fig. 85).
Head: Yellow to yellow-brown, behind eyes, on
posterior part of vertex, round eyes and on clypeus
darkened, between eyes two small spots. Sparsely
punctured. Distance between eyes 0.5-0.6X width of
one eye. Antennae yellow to yellow-brown (fig. 86).
Palpi yellow to brown.
Pronotum: Yellow, basal dark marks opposite fifth
elytral puncture-row. Lateral borders slightly convex,
very strong margins narrowed anteriorly. Hind cor-
ners clearly protruding. Strongly and coarsely punc-
tured along margins, opposite third to fifth elytral
puncture-row a group of two or three very large punc-
tures, between these groups some smaller punctures
in a slight impression. Except in the two basal marks
all punctures colourless. Anteriorly moderately
strongly punctured.
Elytra: Yellow to yellow-brown, darkened along su-
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TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
292
ture, along base as far as sixth punture-row (fig. 85,
87), mark on disc and three other marks on about
fifth and seventh interval. Apical part bulbous, mar-
gin near apex concave. Completely margined. Strong
puncture-rows, first three rows generally weaker than
following three rows, fourth puncture-row only rec-
ognizable by a few punctures in basal part. Between
first and second row about five punctures in apical
part. All punctures darkened. Sutural stria in posteri-
or half.
Underside: Yellow to yellow-brown, legs yellow,
hind femora, coxae, knees and tarsi yellow-brown to
brown. Prosternal process narrowed near coxae, deep
half circular impression at posterior margin, clearly
impressed in anterior 3/4, sparsely punctured (fig. 88,
89). Metasternal process strongly impressed behind
the almost flat anterior part, clear striae along meso-
coxae, weakly and sparsely punctured (fig.88).
Metacoxal plates with a weakly developed blunt tooth
reaching last sternite, in lateral view also blunt,
strongly punctured except near middle- and anterior
suture (fig. 90, 91). Last sternite impressed on both
sides, unpunctured. Epipleura reaching to halfway
last sternite, with darkened punctures in anterior half.
Setiferous striole on dorsal face of hind tibia about
1/4X length of tibia, longest of two tibial spurs about
3/4X length of first tarsal segment.
Male: Penis and parameres as in figs. 92-94.
Surface of apical part of parameres areolate (fig. 95).
Immature-stages: Unknown
Biology
In stagnant water.
Distribution (fig. 101)
Indonesia, Sumatra; Thailand. Zimmermann
(1924) gives Celebes (Indonesia, Sulawesi).
Material examined. — Indonesia: Holotype ? (RMNH); 22
ex., Sumatra, Fast coast, Medan, dated: 1904, 1906,
vii.1920, 29.ix.1921, x.1921 (RMNH, ZMAN); 2 ex., Sumatra,
Deli (ZMAN); 2 ex., Sumatra, det Regimbart 1885, Peltodytes
sumatrensis Reg. Type [does not belong to a syntype-series as
there is only the holotype] (ISNB); 1 ex., Sumatra, Samosir
Isl., 2°28’N 98°49’ E, Tuk-Tuk, L.Toba, 2800 ft. 4.11.1978,
M. Holmen, 13 (zmuc); 1 ex., East coast Sumatra,
Perbaoengan; 1 ex., Sumatra, Medan, Bindjai; 1 ex,
Bandar, Sumatra, 23.ix.1919 (RMNH); 2 ex., N. Sumatra,
Tigibinanga, West of Kabanjahe, 21.1.1990 (16), leg.
Schödl (NHMv) + Thailand: 1 ex., Bangkok, Siam,
Cnemidotus sumatrensis Rég. R. I. Sc. N. B., coll. v.
Dorsselaer, L.G.25622, Peltodytes sumatrensis var.?, det. B. J.
VAN VONDEL: Palaearctic Peltodytes
v. Vondel 1989 (isp); 1 ex., NW-Thailand, Chiangmai
Prov., Chiangdao [Ban Chiang Dao], 450 m., 5.xi.1958, in
standing stagnant water (BPBM).
ACKNOWLEDGEMENTS
I wish to express my sincere thanks to the following
persons for placing material or information at my dis-
posal: L. Baert (Brussels, snp), A. L. van Berge
Henegouwen (The Hague, Moc), M. J. D. Brendell
(London, BMNH), B. Brugge (Amsterdam, ZMAN), J.
G. M. Cuppen (Ede), K. Desender (Brussels, ISNB),
M. B. P. Drost (Wadenoijen), H. Duffels
(Amsterdam, zMAN), Th. G. Giessen (Gaanderen), F.
Gusenleitner (Linz, oLML), M. Hansen (Copen-
hagen, ZMUC), F. Hieke (Berlin, zmus), M.
Hielkema (Gouda), S. J. Hine (London, BMNH), M.
Holmen (Copenhagen, zmuc), J. Huijbregts (Leiden,
RMNH), M. Jäch (Vienna, NHMV), Y. Jongema
(Wageningen), P. Kanaar (Leiderdorp); J. Krikken
(Leiden, RMNH), P. Leblanc (Piney), Sung Hwa Lee
(Taegu), E. Matsui (Hondo City), T. Nakane
(Chiba), N. Nieser (Tiel), R. Ortal (Jerusalem, HuJr),
H. Perrin (Paris, MNHN), G. A. Samuelson
(Honolulu, BPBM), J. C. P. M. v.d. Sande
(Nijmegen), G. Scherer (Munich, zsmc), M.
Schilthuizen (Leiden); M. Schreijer (Alkmaar), H.
Vallenduuk (Boxtel). The Uyttenbogaart-Eliasen
Foundation is acknowledged for financial support.
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298
FER WILLEMSE' & KLAUS-GERHARD HELLER?
! Laurastraat 67, Eygelshoven
° Institut für Zoologie, Erlangen
NOTES ON SY SPEMATICS OF GREBIGSRECIES OF
ROZGIETMONEISGCHIER 1853) (ORDEIORIWERA:
PHANEROPTERINAE)
Willemse, F. & K.-G. Heller, 1992. Notes on systematics of Greek species of Poecilimon
Fischer, 1853 (Orthoptera: Phaneropterinae). — Tijdschrift voor Entomologie 135: 299-315,
figs. 1-43. [1ssN 0040-7496]. Published 1 December 1992.
New data on the systematics of Greek species of Poecilimon Fischer, 1853 are presented. Three
new species are described: P. gracilioides (from Ipiros), P. mariannae (from Thessalia) and P. er-
imanthos (from the Peloponnisos). P. artedentatus Heller, P. veluchianus Ramme and P. zimme-
ri Ramme are given species rank. P. mytilenensis Werner is redescribed and additional descrip-
tive notes are provided for P. soulion L. Willemse, P. aegaeus Werner and P. athos Tilmans et al.
New data on bioacoustics, ecology and faunistics for some species are recorded as well. An up-
dated check list of Greek species is included.
Correspondence. — Dr. F. Willemse, Laurastraat 67, 6471 JH Eygelshoven, The Netherlands.
Key words. — Orthoptera, Phaneropterinae, Poecilimon, taxonomy, bioacoustics, Greece.
The genus of Poecilimon Fischer, 1853 includes
over one hundred species, spread over southeast
Europe and adjacent Asia. All are short winged, usu-
ally green coloured, living in forests, shrubby vegeta-
tion or open habitats from sea-level to above the tim-
berline. Some species are widespread, but most have
restricted and some even remarkably small ranges.
Systematics of Poecilimon are difficult due to the large
number of closely similar species. Up to 1984 system-
atic studies were based entirely on morphology
(Ramme 1933, Bey-Bienko 1954, Harz 1969,
Willemse 1982, 1984).
As in most bush-crickets, males of Poecilimon pro-
duce calling songs which, in presenting an excellent
means of recognizing, provide the basis for a mate se-
lection system. Heller (1984) was the first to present
results of an elaborate study on song and morphology
of stridulatory file in Poecilimon.
Both constitute helpful distinctive characters (e. g.
file in P. ornatuslaffinis, song in chopardil veluchia-
nus). Using these two characters Heller (1984) con-
cluded that a number of previously recognized species
or groups of species are allied either markedly more or
less than formerly presumed. As a result a number of
taxonomic adjustments, mainly synonymic, were
proposed. Renewed studies (Willemse 1985a, Heller
1988, 1990) of some of the species-groups, including
morphological characters and based on more materi-
al, proved that some of these taxonomic readjust-
ments were premature. The present paper is a further
contribution towards the systematics of Poecilimon,
partly based on bioacoustics, with the description of
three new species. As far as the systematics of Greek
species of Poecilimon concern the three main prob-
lems remaining are the taxonomic ranking of the spe-
cies groups of elegans - brunneri - macedonicus, jonicus
- tessellatus - superbus, ampliatus - ebneri - klisuriensis
and the heterogeneous affinis complex. Besides new
species can still be expected especially in the compar-
atively less well explored Aegean islands.
A side effect of recent research is a considerable
amount of new faunistic data since its account in
Willemse (1984, 1985a). Some new information may
be found in Ingrisch & Pavicevic (1985) (e. g. P.
schmidti), Ponel & Hebrard (1988), Heller (1988)
and in this paper. But there is considerably more and
the senior author is preparing a communication on
important new faunistic data of Greek Orthoptera in-
cluding also Poecilimon. For convenience an updated
checklist of Greek Poecilimon taxa is included.
MATERIAL AND METHODS
Depositories and abbreviations. — British Museum
(Natural History), London (BMNH); Collection
Heller, Universität Erlangen (cH); Collection A.
Nadig, Chur (cn); Collection J. Szijj, Universität
Essen (cs); Collection J. Tilmans, Warmond (cr);
Collection Willemse, Eygelshoven (cw); Instituut
voor Taxonomische Zoölogie, Amsterdam (ITZA);
299
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 1-5. Poecilimon gracilioides. - 1, 3, dorsal view (holotype); 2, 9, lateral view (allotype); 3, stridulatory file of left elytron
d (paratype), ventral view, distal end to the left (scale 1 mm); 4, idem, detail of mid to distal third (scale 100 um); 5, base of
ovipositor, lateral view (paratype).
300
Naturhistorisches Museum, Vienna (NHMV).
Song recording and analysis. — For recordings of
the song the following equipment has been used:
Uher 4200 Report Monitor with AKG D 202 E micro-
phone with low frequencies off (after modification)
(P. gracilioides); Uher 4200 1c with Uher M 645 mi-
crophone (recordings of other species). To improve
quality recordings were made indoors. Oscillograms
were made by using a personal computer and the pro-
gram TURBOLAB (Stemmer AG). Wing movements
were registrated by an opto-electronic device (Heller
1988). Full data of the recordings can be obtained
from the authors. Stridulatory files were studied with
a light microscope and a Scanning Electron Mi-
croscope (SEM) (Institut für Zoologie I, Universität
Erlangen). A batdetector (QMC-mini) was used in the
field to locate males as indicated in the text.
Song terminology. — Calling song: the song pro-
duced by an isolated male. Impulse: the sound pro-
duced by the contact of one file tooth with the scrap-
er (= plectrum). Syllable: the sound produced by one
complete up (opening) and down (closing)stroke of
the fore wing. Hemisyllable: the sound produced by
one unidirectional movement (opening or closing) of
the fore wings. Echeme: a first-order assemblage of
syllables. Echeme-sequence: a first order assemblage
of echemes.
Presentation. — In the checklist species are present-
ed in alphabetical order but the sequence of the spe-
cies in the present study follows a preliminary ar-
rangement of relationships based on the song pattern
(Heller 1990). References listed under the species are
not always complete, full bibliography up to 1985
may be found in Willemse (1982, 1984, 1985a).
Material listed is new and does not include previous-
ly recorded specimens except for the type material of
P. aegaeus. Measurements are given in mm and con-
cern the length. Transliteration of Greek orthography
in Roman characters agrees with the system proposed
by the Permanent Committee on Geographical
Names for British Official Use, London.
TAXONOMIC PART
Updated checklist of Greek Poecilimon
Those headed in this paper are provided with an as-
terisk in the list; names between brackets indicate
places in the present paper where comparative notes
are made.
*aegaeus Werner, 1932
affinis (Frivaldsky, 1867)
anatolicus Ramme, 1933
* artedentatus Heller, 1984 stat. n.
WILLEMSE & HELLER: Systematics of Greek Poecilimon
* athos Tilmans e.a., 1989
brunneri (Frivaldsky, 1867)
chopardi Ramme, 1933 (in mariannae & veluchianus)
cretensis Werner, 1903
ebneri Ramme, 1933
*erimanthos sp. n.
deplanatus Brunner von Wattenwyl, 1891
* gracilioides sp. n. (also in veluchianus)
gracilis (Fieber, 1853) (in gracilioides)
hamatus Brunner von Wattenwyl, 1878
hoelzeli Harz, 1966 (in veluchianus)
ikariensis F. Willemse, 1982
jonicus jonicus (Fieber, 1853) (in erimanthos)
jonicus lobulatus F. Willemse, 1982 (in erimanthos, gracili-
oides, veluchianus & werneri)
jonicus tessellatus (Fischer, 1853) (in erimanthos)
klisuriensis F. Willemse, 1982
laevissimus (Fischer, 1853) (in erimanthos)
macedonicus Ramme, 1926
* mariannae sp.n. (also in propinquus & veluchianus)
miramae Ramme, 1933
*mytilenensis Werner, 1932
nobilis Brunner von Wattenwyl, 1878 (in erimanthos)
obesus Brunner von Wattenwyl, 1878 (in artedentatus)
orbelicus Pancic, 1883
ornatus (Schmidt, 1849) (in gracilioides & veluchianus)
paros Heller & Reinhold, 1992
pergamicus Brunner von Wattenwyl, 1891
pindos F. Willemse, 1982
*propinquus Brunner von Wattenwyl, 1878 (also in marian-
nae & veluchianus)
sanctipauli Brunner von Wattenwyl, 1878
schmidti (Fieber, 1853)
*soulion L. Willemse, 1987 (also in gracilioides)
syriacus Brunner von Wattenwyl, 1891
thessalicus Brunner von Wattenwyl, 1891 (in veluchianus)
thoracicus (Fieber, 1853)
*veluchianus Ramme, 1933 (also in gracilioides, mariannae
& propinquus)
* werneri Ramme, 1933 (also in erimanthos)
zimmeri Ramme, 1933 (in erimanthos, gracilioides, marian-
nae & veluchianus)
zwicki Ramme, 1939
Poecilimon artedentatus Heller stat. n.
Poecilimon obesus artedentatus Heller, 1984: 78; Willemse
1985a: 16; 1985b: 39; Heller 1988: 47; 1990: 140.
Poecilimon obesus, Werner 1929: 481; Ramme 1933: 513;
Werner 1934: 324; 1938: 167; Bei-Bienko 1954: 271;
Harz 1969: 118; Willemse 1982: 161; 1984: 34 (all par-
tim).
Material studied. — Akhaia: Boubouka-Krioneri, W of
Kalavrita; Kernitsai, N of Kalavrita 600 m (cw); Kalavrita
and surr. 700-800 m; Flamboura-Vlasia 700-800 m;
Drosato, NW of Kalavrita 900 m (cH); Arkadhia: surround.
Dhavia, E of Khrisovitsi (cH); 4 km S of Tripolis 700 m
(cw); Ilia: 4 km NW of Andritsena 650 m (cn).
Discussion
Originally artedentatus was erected at subspecific
rank for the P. obesuslike populations from the
Peloponnisos (Heller 1984; Willemse 1985a: map 3).
301
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 6-10. Poecilimon erimanthos. - 6, 3 , dorsal view (holotype); 7, 9, lateral view (allotype); 8, 5, protrusion of second ter-
gite, lateral view (paratype, Kalentzi, C. W.); 9, base of ovipositor, lateral view (paratype, A. Pandon-Alepokhori); 10, stridu-
latory file of left elytron & (paratype, Kalentzi), ventral view, distal end to the left (scale 100 um).
302
Distinction between P. obesus and artedentatus is lar-
gely based on the song and the stridulatory file.
Considering the consistency and degree of difference
and in order to balance the taxonomic interpretation
assigned to them with the taxonomic interpretation
given to similar differences observed in Poecilimon it
has been decided to raise artedentatus to species level.
It is noted here that a confusing mistake has slipped
into the key in Willemse (1985b: 38-39). The charac-
ters of the stridulatory file were exchanged erroneous-
ly and should read:
118(117)Stridulatory file with relative distance
between the teeth larger, the total number
of teeth less (some 120); etc. …
EN Poecilimon obesus Brunner 1878
Stridulatory file with relative distance
between the teeth smaller, the total number
ohteethlaren(some 280); etc...
ns Poecilimon artedentatus Heller, 1984
Poecilimon soulion L. Willemse
(fig. 11)
Poecilimon soulion L. Willemse, 1987: 173; Heller 1988: 51.
Material studied. — Type material.
Additional descriptive notes
Auditory spiracle large, aperture narrowly ellipti-
cal, dorsally reaching middle of pronotal lateral lobe.
Female stridulatory apparatus: Ventral side of left
elytron with well developed plectrum; right elytron
with stridulatory pegs on dorsal side of veinlets which
form a network bordering the widely rounded poste-
ro-apical edge of elytron.
Poecilimon gracilioides sp. n.
(figs. 1-5, 12, 24, 30)
Type material. — Holotype d: Hellas, Arta, Mt.
Tzoumerka, S of Theodhoriana, 1400-1800 m, 24. vii.
1987, Fer Willemse (cw). — Paratypes 135 49: same data (
2d cH; 1d 19 BMNH; 19 allotype, 103 29 cw).
Description
Male (fig. 1). — Medium-sized, remarkably slender,
integument moderately shiny.
Head: fastigium of vertex produced anteriorly, lat-
eral margins parallel, length and width about equal to
those of second antennal segment, longitudinally im-
pressed above.
Thorax: pronotum (fig. 24) with metazona dome-
shaped and raised above elytra, posterior margin
emarginate. Auditory spiracle large, aperture narrow
WILLEMSE & HELLER: Systematics of Greek Poecilimon
elliptical, reaching dorsally upper third of pronotal
lateral lobe. Elytra in dorsal view with basal fourth
covered by pronotum, apical margin extending be-
yond posterior margin of first tergite. Stridulatory file
of left elytron (figs. 3-4) with about 120 teeth, reach-
ing posterior margin of elytron; in ventral view arcu-
ate, in profile concave proximally and almost straight
in distal third; shortest distance between proximal
and distal most tooth 2.7-2.9 mm, greatest width in
mid part 0.19 mm; proximal two thirds with 38-45
well developed large teeth, distal third with about 80
much smaller teeth; spacing ranging from 22-24
proximally to 20-22 in mid part to about 80 teeth per
mm in distal part; change from larger, wider and less
closely set teeth to smaller, narrower and more dense-
ly arranged teeth at transition of mid to distal third of
file rather abrupt. Hind femora unarmed or with a
single ventral spine.
Abdomen: margins of tergites straight except
slightly emarginate posterior margin of first tergite.
Cercus (fig. 12) moderately long, stout, incurved and
slightly tapering in apical third, apex strongly taper-
ing into a short, strong, conical tooth near anterior
margin. Subgenital plate reaching tip of cerci, lateral
margins converging posteriorly, postero-lateral edges
slightly protruding, posterior margin straight, weak
median carina.
Coloration: yellowish-green. Vertex, pronotum,
tergites and legs finely speckled rusty brown.
Antennae yellowish brown. Fastigium of vertex rusty
red. Pronotal dorsum with a pale yellow lateral stripe,
in metazona bordered medially dark rusty brown,
sometimes a similarly coloured median streak. Elytra
yellowish with stridulatory area dark brown.
Abdomen unicolorous or with two yellowish dorsal
bands bordered laterally with black markings, some-
times a narrow black median stripe. Cercus yellowish,
tip black. Legs of general colour.
Female (fig. 2). — Pronotum almost cylindrical,
metazona very slightly inflated, scarcely raised poste-
riorly, posterior margin slightly concave. Elytra just
visible, overlapping each other medio-dorsally, apical
margin transverse, reaching anterior margin of first
tergite or shorter. Stridulatory apparatus present: ven-
tral side of left elytron with plectrum; dorsal side of
right elytron with stridulatory pegs on top of veinlets
which form a network bordering posterior margin
and postero-apical angle of elytron, strongest pegs
along posterior margin. Basal fold of dorsal margin of
lower ovipositor valve lamelliform, strongly bent ven-
trally, forming with gonangulum a rather shallow lat-
erally-facing groove (fig. 5). Coloration unicolorous
green.
Measurements. — Body d 18.0-21.0, 2 18.5-20.0;
pronotum d 4.2-4.6, © 4.5-4.8; elytron d 2.5-2.9,
2 0.1-0.3; hind femur d 14.3-15.1, 2 15.2-15.6;
303
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Figs. 11-19. Poecilimon, left cercus 3, dorsal view. - 11, P. soulion (paratype); 12, P. gracilioides (paratype); 13, P. gracilis (Mt.
Pelister, Yugoslav Makedonia); 14, P. erimanthos (paratype, Alepokhori-Greveno); 15, P. mytilenensis (Ayiassos); 16, P. mar-
tannae (paratype, Vrisia); 17, P. propinquus (Mt. Dhirfis, Evvoia); 18, P. propinquus (Mt. Othris, Magnisia); 19, P. aegaeus
(paratype, Sira). - Fig. 20. Poecilimon aegaeus, right cercus (holotype, Mikonos). - Figs. 21-22. Poecilimon, tip of & abdomen,
lateral view. - 21, 2. mytilenensis (Megalokhorion); 22, species (= ? P. mytilenensis) (Kaloni). — Figs. 23-24. Poecilimon,
Pronotum 6, lateral view. 23, P. gracilis (Florina, NW Greek Makedhonia); 24, P. gracilioides (paratype). Scales 1 mm.
304
ovipositor 8.4-8.7.
Bioacoustics. — Male calling song an isolated syl-
lable (fig. 30), produced with intervals of several sec-
onds. Two syllables produced immediately one after
another occur only sporadically. Syllable composed of
three impulse groups: a series of up to 50 faint im-
pulses, slightly increasing in amplitude, followed by a
second group of about 7 loud impulses separated
from each other by intervals of 10 to 20 ms and the fi-
nal group consisting of about 20 loud impulses with
intervals less than 3 ms (30°C). Fore wing move-
ments not yet registrated.
Distribution and ecology
Known only from Mt. Tzoumerka (1400-1800 m
altitude), NE of the town of Arta, Ipiros, western
continental Greece. Tzoumerka is the archaic name
for Mt. Athamanon or Athamanika. This large
mountain complex, highest summit 2393 m, is
zoogeographically somewhat isolated being quite nar-
rowly connected eastward with the Pindhos range
and separated westward by the Arakhthos river from
some lower mountains in western Ipiros. The new
species was found at 1400-1800 m altitude on the
northeastern slopes of one of the southern summits,
named Spilia (1932 m), S of the village of
Theodhoriana. The habitat, close to a spring, are roc-
ky slopes covered by rich vegetation of low shrubs,
herbs and grasses. The species occurs quite locally,
was not numerous and found mainly on Helleborus
leaves, together with P. ornatus, jonicus lobulatus,
zimmeri and veluchtanus.
Differential diagnosis
The species is defined by pronotum, fore wing and
cercus in the male, basal fold of the lower ovipositor
valve in the female and size, slender habitus, colora-
tion and almost unarmed hind femora in both sexes.
A key character is presented by the stridulatory file.
P. gracilioides resembles P. gracilis in size, colora-
tion and general habitus but differs from it by less
raised male pronotum (figs. 23-24), shorter male cer-
cus (figs. 12-13) and especially in the male stridulato-
ry file: in gracilis (L. Willemse 1987, Heller 1988) the
teeth are gradually narrowing and closer set near the
transition of mid to distal third of file length, in gra-
cilioides size and spacing of teeth changes abruptly at
that part of the file (figs. 3-4). The calling song resem-
bles that of P. gracilis (Heller 1984, 1988) but the dis-
tinction is not yet pointed out because variability and
influence of temperature are known insufficiently.
Readily recognizable differences with P. soulion (L.
Willemse 1987, Heller 1988) refer to a more robust
habitus, broader and less raised male pronotum, larg-
er size, more spined hind femora, slightly deeper basal
groove of the lower ovipositor valve and the colora-
WILLEMSE & HELLER: Systematics of Greek Poecilimon
tion in this species. The male cercus of both species
differs only slightly (figs. 11-12). The sudden change
in spacing of the teeth of the stridulatory file is shared
by both species. In P. soulion the file is longer (3.0-3.2
mm against 2.7-2.9 mm), the number of teeth in its
proximal two thirds slightly larger (58-62 against 38-
45) and the difference in spacing of the teeth in the
mid part of the file compared with those at its proxi-
mal (= basal) end apparently larger: about 36 and 18
per mm against about 23 and 21 teeth per mm in gra-
cilioides. Bioacoustics can not be compared as those of
P. soulion are unknown.
Etymology
Named after its resemblance to P. gracilis.
Poecilimon werneri Ramme
(figs. 31, 42)
Poecilimon werneri Ramme, 1933: 530; Harz 1969: 135;
Willemse 1985a: 17; 1985b: 41; Heller 1988: 58.
Poecilimon jonicus, Heller 1984: 95 (as synonym).
Material studied. — Aitolia-Akarnania: Menidhion 50 m;
Anoixiatikon 50 m; Loutrakion 70 m; Mitikas 10 m; 10 km
SE of Astakos 50 m (all cw); Akheloos delta, Kardhos (cs);
Akhaia/Ilia: 2 km S of Porte (N of Pinios lake) 650 m (cH).
Additional descriptive notes
Auditory spiracle large, aperture elliptical, exten-
ding dorsally to upper third of pronotal lateral lobe.
Female stridulatory apparatus: ventral side of left elyt-
ron provided with plectrum; stridulatory pegs on dor-
sal surface of small veinlets (most of them almost pa-
rallel to each other) of postero-apical part of right
elytron.
Bioacoustics. — Male calling song ( fig. 31) an iso-
lated syllable, produced with intervals of about 1 s
(ca. 29°C); syllable composed of two hemisyllables,
closing one distinctly louder than opening one; open-
ing hemisyllable (45 ms at ca. 29°C) about 25 im-
pulses; closing hemisyllable (70 ms at ca. 29°C) com-
monly two groups of impulses separated by interval of
about 25 ms (ca. 29°C), first group of 20-40 impuls-
es, second group of 1-8 or sometimes missing; maxi-
mum of frequency spectrum 29-44 kHz.
Distribution
The range, previously poorly known (Willemse
1984: map 19), is remarkably small, restricted to
coastal areas of extreme southwestern continental
Greece and opposite part of the north-western
Peloponnisos (fig. 42).
Discussion
Induced by the proposed synonymy of P. werneri
with jonicus (Heller 1984) its specific status was ar-
305
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Fig. 25-29. Poecilimon mytilenensis. — 25, 3, dorsal view (Ayiassos); 26, ©, lateral view (Playia); 27, ovipositor and its base,
lateral view (Megalokhorion); 28, stridulatory file of left elytron & (Ayiassos), ventral view, distal end to the left (scale 300
Um); 29, idem, detail of mid to distal third, distal end to the right (scale 100 um).
306
gued on morphological basis (Willemse 1985a).
Evidence for its specific status is now presented both
by their distinctive song and the syntopic occurrence
with P. jonicus lobulatus, even on a single shrub
(Cistus sp., Rubus sp.) (Menidhion and Anoixiatikon,
both Aitolia-Akarnania).
Poecilimon erimanthos sp. n.
(figs. 6-10, 14, 32-33, 42)
[P. erimanthos F. Willemse & Heller 1988]: Heller 1988:
57; 1990: 142 (bioacoustics only).
Type material. — Holotype d: Hellas, Akhaia, Mt.
Erimanthos , A. Pandhon-Alepkhori, 700 m, Fer Willemse,
17. vii. 1987 (cw). — Paratypes 76 99: same data (16 19
ITZA; 12 cn; 19 allotype, 68 72 cw); Akhaia: Mt.
Erimanthos, Alepokhori-Greveno, 800 m, 17. vii. 1987
(108 32 cw; 1d 12 BMNH); Greveno-Kalentzi, 900 m,
17. vii. 1987 (18 19 cw); Kalentzi-Drosia, 600 m, 17. vii.
1987 (28 19 cw); Kalentzi, 900 m, 17. vii. 1987 (16 19
cw), all Fer Willemse; Kalenzti, 4. vii. 1986, K.-G. Heller
(16 12 cu); Kalenzti and surr., 900-1100 m, 10. vii. 1985,
A. Nadig (16 cn); Mt. Erimanthos ab. Kalentzi, 1700-
2000 m [ but probably 1100 m near Kalentzi], 4. viii. 1970,
Willemse & Scherpbier (16 cw); Akhaia/llia: 2 km S of
Porte (N of Pinios lake), 650 m, 14. vi. 1989, K.-G. Heller
(5d cu); Ilia: 5 km E of Koumanis ( Erimanthos valley),
600 m, 16. vi. 1989, K.-G. Heller (36 12 cu); 3 km NW
of Karia, 500 m, 15. vi. 1989 , K.-G. Heller (58 112 cm);
Keramidhia, 150 m, vi. 1988, K. Reinhold (4d 19 cm);
Mt. Erimanthos, S of Kriavrisi, 800 m, 15. vii. 1987, Fer
Willemse (96 72 cw); Mt. Lambia ab. Lambia village,
1000-1150 m, 16. vii. 1991, Luc Willemse (36 22 cw).
Description
Male (fig. 6). — Small, remarkably slender, integu-
ment moderately shiny.
Head: fastigium of vertex produced anteriorly, lat-
eral margins parallel or slightly converging anteriorly,
twice as long as wide, as narrow as width of third an-
tennal segment or a little wider, longitudinally im-
pressed above.
Thorax: pronotum with metazona not widening
but slightly inflated posteriorly, posterior margin
weakly emarginate. Auditory spiracle large, aperture
widely elliptical, reaching dorsally upper fourth of
pronotal lateral lobe. Elytra with basal three quarters
covered by pronotum, apical margins reaching or just
surpassing anterior margin of first tergite.
Stridulatory file of left elytron (fig. 10) with 70-90
teeth, not quite reaching posterior margin of elytron;
in ventral view slightly fusiform and arcuate, in pro-
file distinctly bulging just beyond middle of length;
shortest distance from proximal to distal most tooth
1.0-1.2 mm, greatest width in mid third 0.05 mm; in
proximal two thirds 35-45 teeth, in distal third about
same number of distally increasingly narrowing teeth;
spacing in proximal two thirds almost regular, in dis-
WILLEMSE & HELLER: Systematics of Greek Poecilimon
tal third increasingly closer. Hind femur unarmed.
Abdomen: posterior margin of first tergite angular-
ly excised. Second tergite with a quite remarkable me-
dian protrusion (fig. 8) at least as large as antennal
scape and slightly wider than long; anterior and later-
al sides slightly flattened and arising vertically, poste-
rior side more sloping anteriorly (and thus longer
than anterior side), depressed medially and merging
with the inflated, weakly bilobate dorsal side; anterior
and dorsal surface of this protrusion bristly. Cercus
(fig. 14) slender, proximal half slightly tapering to-
wards cylindrical apical half, from base to apex in-
creasingly incurved, apex from below shortly and
from other sides gradually tapering into a sharp and
sometimes slightly irregularly shaped pointed crest.
Subgenital plate reaching mid to distal third of cer-
cus, slightly narrowing apically with obtuse median
carina ventrally; lateral margins carinate, terminating
into not or slightly protruding postero-lateral edges,
posterior margin straight to slightly emarginate.
Coloration: yellowish green. Vertex, most of dor-
sum of pronotum, tergites and legs finely speckled
rusty brown or blackish. Antennae yellowish, finely
annulated dark brown. Occiput with spots sometimes
more closely set medially, always unspotted along me-
dian line. Pronotal dorsum with yellowish median
line, prozona and posterior part of metazona reddish
brown and on either side a yellowish streak; lateral
lobes often greenish. Elytra with stridulatory area
usually brownish contrasting with yellowish to green-
ish white anterior part, apical margin sometimes dark
brown. Abdomen dorsally yellowish, laterally of sim-
ilar colour or brownish or mixed with green; tergites
with yellow median line, on either side bordered by a
black block, posterior margin yellowish; protrusion of
second tergite and its surroundings on dorsum dark
reddish brown; sternites yellowish or pale green.
Cercus yellowish, apical third dorsally blackish, tip
black. Subgenital plate pale yellowish. Femora with
black dorsal incomplete stripe, lower keels blackish,
especially distally and hind femur with black lateral
incomplete stripe as well; tibiae yellowish brown,
between spined upper edges often black.
Female (fig. 7). — Pronotum almost cylindrical,
metazona not or scarcely inflated posteriorly. Elytra
not or usually just visible, overlapping each other me-
dio-dorsally, apical margin transverse, reaching ante-
rior margin of first tergite. Stridulatory apparatus
present: ventral side of left elytron with plectrum;
dorsal side of right elytron with irregular rows of stri-
dulatory pegs on top of a number of veinlets that
reach posterior margin and postero-apical angle of
elytron, those reaching posterior margin roughly par-
allel to each other. Basal fold of dorsal margin of low-
er ovipositor valve lamelliform, moderately protrud-
ing horizontally, not or very slightly impressed
307
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
N
34
Fig. 30. Poecilimon gracilioides, oscillogram of & calling song (paratype). — Fig. 31-34. Poecilimon, d , synchronous registra-
tion of left elytron movement (above; upward is opening, downward is closing of wing) and calling song (below). - 31, P. wer-
neri (Porte); 32, P. erimanthos, isolated syllable (paratype, Kalentzi, C. H.); 33, P. erimanthos, echeme (paratype, Kalentzi, C.
H.); 34, P. mariannae (holotype). Scale 200 ms.
dorsally, forming with gonangulum a longitudinal,
deep and laterally-facing groove (fig. 9). Coloration as
male, sometimes unicolorous greenish.
Measurements. — Body d 17.0-20.0, 2 19.0-22.0;
pronotum d 4.1-4.7, ® 5.3-5.8; elytron d 0.5-1.0,
2 0.0.-0.3; hind femur d 14.0-17.1, 2 17.3-19.5;
ovipositor 9.1-9.9.
Bioacoustics. — Male calling song (figs. 32-33) an
isolated syllable (fig. 32) or a short echeme (fig. 33)
repeated at variable intervals (often about 1 s; 23-
24°C); isolated syllable consisting of very soft open-
ing hemisyllable (sometimes even undetectable) and
loud and very short (up to 5 ms; 23-24°C) closing
hemisyllable. An echeme begins with a syllable similar
to isolated one followed by a group of 10-15 impuls-
es (about 50 ms; 23-24°C), produced during second
opening of elytra while final closing movement pro-
duces no sound at all. Echemes possibly produced
mainly by males which do receive response of a fe-
308
male.
Distribution and ecology
The range of the species covers a remarkably small
area of the northwestern Peloponnisos: from the wes-
tern and southern slopes of Mt. Erimanthos (up to
1100 m altitude) down to the adjacent lowlands of
northern Ilia (fig. 42). The species was found on
thistles, blackberry bushes and Astragalus, sometimes
in a marshy spot. P. erimanthos occurred syntopically
with werneri (Akhaia/Ilia: Porte), laevissimus (Akhaia:
Kalentzi; Ilia: Koumanis), zimmeri (Ilia: Mt.Lambia)
and nobilis (Ilia: Koumanis). Attempts to find the
species together with P. jonicus tessellatus at the nor-
theastern part of its range failed up to now.
Differential diagnosis
P. erimanthos is defined by pronotum, elytron and
cercus in the male, basal fold of the lower ovipositor
valve in the female and size, slender habitus, colora-
tion and unarmed hind femora in both sexes. Most
obvious feature is the median prominence of the ma-
le second tergite by which ?. erimanthos is recogniza-
ble at a glance. So far known to us such a structure in
Poecilimon exists only in glandifer Karabag, 1950
(Karabag 1950: fig. 7) and ampliatus Brunner von
Wattenwyl, 1878 (Krauss 1878: pl. 3 fig. 3a, Harz
1969: fig. 373). The prominence in these species, ho-
wever, is located on the first tergite, partly covered by
the elytra and differently shaped. Also other features
of these species differ clearly from P. erimanthos.
Except for the prominence of the second male ter-
gite, P. erimanthos comes near the closely related spe-
cies laevissimus, werneri and jonicus (s.l.).
The male second tergite of P. laevissimus presents
no prominence but is slightly inflated, also bristly and
of similar colour medially. Other distinctive features
(Willemse 1982, Heller 1984) are summarized: male
cercus quite larger; stridulatory file less bulged; pro-
notal metazona only a little more inflated; same
coloration of body but black dots of tergites more sol-
id; antennae more widely annulated; basal fold of
lower ovipositor valve more and roundly impressed
from above, groove larger and more rounded; size of
body and legs usually larger but occasionally similar.
P. werneri differs in: slightly longer elytra, straight
profile of stridulatory file; male cercus more incurved
in apical third, tip more pointed; basal fold of lower
ovipositor valve slightly arched upward anteriorly and
groove with gonangulum more shallow and facing
more anteriorly instead laterally; tergites with similar
colour pattern but, on either side, an additional black
lateral streak (Willemse 1982, 1985a).
P. jonicus (s. 1.) can be distinguished by more inflat-
ed pronotal metazona; more visible and slightly long-
er elytra, file of similar shape but shorter, bulging
stronger and more distally, teeth less closely set; male
cercus pre-apically more angularly incurved and
slightly flattened; basal groove of lower ovipositor
valve smaller and round instead elongate and besides
in jonicus lobulatus provided with an anterior inflated
processus (Willemse 1982, 1985a).
The calling song resembles also these last three spe-
cies. In P. erimanthos the duration of the closing hem-
isyllable of the isolated syllable or of the first syllable
of an echeme respectively, is shorter than in P. jonicus
jonicus, jonicus lobulatus and werneri (fig. 31, Heller
1984, 1988). Isolated syllables of P. jonicus tessellatus,
laevissimus and erimanthos may be indistinguishable
from each other. The echeme structure in P. eriman-
thos, however, is unique. Echemes in P. laevissimus
and werneri were, so far known, not produced at all,
while in jonicus (s. |.) the second syllable of an echeme
is not restricted to an opening hemisyllable as in er7-
manthos but to a closing one or both to an opening
WILLEMSE & HELLER: Systematics of Greek Poecilimon
and closing hemisyllable (Heller 1984).
Discussion
We observed that during mating the female nibb-
led eagerly at the peculiar prominence of the second
male tergite. Further anatomy or function of the pe-
culiar prominence of the male second tergite are unk-
nown.
Etymology
A noun in apposition, named after its distribution
that centres on Mt. Erimanthos.
Poecilimon mytilenensis Werner
(figs. 15, 21, 25-29)
Poecilimon mytilenensis Werner, 1932: 3; 1933: 404;
Ramme 1933: 537; Werner 1934: 324; Harz 1969: 144;
Willemse 1982: 188; 1984: 41; 1985b: 43; Heller 1988:
69 (in Willemse 1982, 1984, 1985b & Heller 1988 as
mytilensis).
Material studied. — Lesvos: S of Ayiassos 500 m, Castanea
forest; Megalokhorion 550 m; Playia, E of Plomari; Thermi,
N of Mytilini (all rrza); Andhissa (cw).
Redescription
Male (fig. 25). — Medium-sized, integument slight-
ly glossy.
Head: fastigium of vertex with lateral margins par-
allel or slightly converging anteriorly, twice as long as
wide, as wide as second antennal segment, smooth
above.
Thorax: pronotum slightly compressed laterally
and thus a little saddle-shaped; metazona slightly
widening and raised posteriorly. Auditory spiracle
large, aperture narrow elliptical, reaching dorsally
upper third of pronotal lateral lobe. Elytra in dorsal
view with basal two thirds covered by pronotum, ap-
ical margins reaching half length of first tergite.
Stridulatory file of left elytron (figs. 28-29) with 100-
120 teeth, reaching posterior margin of elytron; in
ventral view arcuate, proximally more; in profile
slightly concave in proximal two thirds, then sharply
giving way like one step downstairs and farther distal-
ly again slightly concave; shortest distance between
proximal and distal most tooth about 2.2 mm, great-
est width near the step 0.06-0.07 mm; proximal two
thirds with 50-60 distally gradually wider and less
closely arranged teeth reaching the step and from
there, in distal third, about similar number of teeth
but now about half as wide and twice as closely set.
Hind femur unarmed.
Abdomen: margins of tergites straight; medio-an-
terior part of second tergite slightly inflated, black.
Cercus (fig. 15) slender, in proximal two thirds slight-
ly, in distal third stronger in- and a little upcurved;
309
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
Fig. 35-39. Poecilimon aegaeus. — 35, 3, dorsal view (holotype); 36, à, lateral view (holotype); 37, base of ovipositor, lateral
view (paratype, Sira); 38, stridulatory file of left elytron & (holotype), ventral view, distal end to the left; 39, idem, in profile.
- Fig. 40-41. Poecilimon mariannae. — 40, 3, dorsal view (holotype); 41, stridulatory file of left elytron d (paratype,
Metallion), ventral view, distal end to the left (scale 300 um).
310
proximal half slightly conical, distal half a little com-
pressed dorso-ventrally, apical part flattened and wid-
ened (spatulate), apical edge slightest rounded and
provided with minutely serrate and slightly down-
curved crest at medio-posterior part of edge. Male
subgenital plate (fig. 21) remarkably slender and
long; in ventral view at least twice as long as greatest
width, in distal half tapering to half greatest width or
even less; posterior margin straight, postero-lateral
edges slightly protruding; in profile distal half strong-
ly upcurved, extending far between cerci and reaching
level of last tergites, very edge of posterior margin
commonly turned down. Coloration: straw yellow,
sometimes mixed with green. Vertex, most of prono-
tum, tergites and legs finely speckled rusty brown or
reddish. Antennae yellowish, finely annulated black-
ish. Median line of vertex and pronotum and a post-
ocular stripe yellow. Pronotal dorsum usually with
black dots in prozona and along sulci, metazona often
with reddish flush. Elytra unicolorous yellowish,
sometimes a pre-apical dark brown streak. Tergites
medio-dorsally each with pair of black blocks, widen-
ing anteriorly especially in proximal tergites, not
reaching very posterior margin and bordered laterally
yellowish; lateral sides of tergites and all of sternites of
general colour. Cerci yellowish, tip dorsally black.
Subgenital plate of general colour, very hind margin
sometimes blackish. Legs with more or less distinct
black stripes.
Female (fig. 26). — Pronotal dorsum in profile
straight to commonly slightly concave. Elytra com-
pletely covered by pronotum or just visible, shortly
overlapping each other medio-dorsally. Stridulatory
apparatus present: ventral side of left elytron with
plectrum; dorsal side of posterior part of right elytron
with stridulatory pegs on top of some 5-7 almost par-
allel to each other running veinlets and on top of sev-
eral other non-directional veinlets towards postero-
apical angle. Basal fold of dorsal margin of lower
ovipositor valve lamelliform, moderately extending
laterally but strongly impressed dorsally forming with
gonangulum a round, deep and dorsolaterally-facing
pit (fig. 27). Ovipositor comparatively short.
Coloration as male but commonly unicolorous yel-
lowish.
Measurements. — Body d 18-20, 2 22-24; prono-
tum d 5.7-6.0, 2 5.4-6.0; elytron d 0.6-1.1, 2 0.1-
0.5; hind femur d 16.5-17.1, 2 16.3-17.2; oviposi-
tor 6.3-7.0.
Bioacoustics. — Unknown.
Distribution
Only known from the eastern Aegean island of
Lesvos, at least from its southern part (see remarks be-
low).
WILLEMSE & HELLER: Systematics of Greek Poecilimon
Discussion
P. mytilenensis was known only from two males (ty-
pe series) and one female (Werner 1934). The mate-
rial listed above agrees fully with the descriptions and
figures of the male holo- and paratype. Previous de-
scriptions (Werner 1932, 1933, Ramme 1933, Harz
1969) are insufficient.
Additional material before us from the island of
Lesvos: 3 km NW of Kaloni (rrza); N of Kaloni (IT-
ZA); Sikaminea (cw) is strongly reminiscent of P. my-
tilenensis, particularly the virtually identical male stri-
dulatory file. However, the male subgenital plate (fig.
22) is remarkably different: much shorter, much less
tapering posteriorly and the apical part not at all
pointing dorsally between the cerci. Some other char-
acters differ as well but not so conspicuously. It is al-
so remarkable that these specimens are from the
northern part while typical P. mytilenensis occur in
the southern part of the island. Lack of bioacoustical
particulars prevents further identification.
Poecilimon mariannae sp. n.
(figs. 16, 34, 40-41, 43)
Poecilimon cf. aegaeus Werner; Heller 1984: 104.
[Poecilimon mariannae F. Willemse & Heller, 1988 ]: Heller
1988: 68; 1990: 142 ( bioacoustics only).
Type material. — Holotype d: Hellas, Fthiotis:
Dhomokos, Metallion, 1 km South East, 450 m, K.-G.
Heller, 21. vi. 80 (cw). — Paratypes 28 1 9: same data (2d
19 allotype cH); Fthiotis: Metallion, 4 km S , 500 m, 10.
vii. 1987, F. Willemse (38 99 cw); 19. vii. 1991, F. & L.
Willemse (16 29 cw); 1 km SE of Metallion, 450 m, 29. v.
1989, K. -G. Heller (14 cy); 3 km SE of Metallion, 450 m,
29. v. 1989, K. -G. Heller (46 69 ch); Perivoli, SW of
Dhomokos, 500 m, 5. vi. 1989, K. -G. Heller (16 cn);
Larissa: 1 km NW of Nea Monastirion, 150 m, 27. v. 1990,
K. -G. Heller (45 42 cn); 2 km SW of Vrisia, 150 m, 29.
v. 1989, K. -G. Heller (56 19 cn); Vrisia near Farsala, 200
m, 10. vii. 1987, F. Willemse (24 39 cw); Kardhitsa:
Mouzaki, 15. vi. 1991, K.-G. Heller (106 59 cn).
Description
Male (fig. 40). — Small, robust, integument scarce-
ly glossy.
Head: fastigium of vertex with lateral margins.
about parallel, slightly sloping antero-ventrally, short,
as long as wide, narrower than scape, not or weakly
impressed dorsally.
Thorax: pronotum slightly saddle-shaped, a little
widening and metazona somewhat raised posteriorly;
posterior margin emarginate. Auditory spiracle tiny,
aperture split-like, greatest diameter as length of
scape. Elytra well visible, apical margin reaching or
just surpassing posterior margin of first tergite.
Stridulatory file of left elytron (fig. 41) with 62-70
teeth, almost reaching posterior margin of elytron; in
Sal
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
EP werner
%* P.erimanthos
a
16 0
@P.chopardi
BO P.marian
Fig. 42-43. Poecilimon, distribution in Greece. — 42, P. werneri and P. erimanthos; 43, P. mariannae and P. chopardi.
ventral view slightly fusiform and arcuate, in profile
evenly arcuate except very distal part; shortest dis-
tance between proximal and distal most tooth 2.0-2.3
mm, greatest width in mid part 0.08-0.09 mm; spac-
ing widest in mid third, distance between two succes-
sive teeth about 0.0416-0.0454 mm, i.e. 11 to 12
teeth per 0.5 mm, increasingly closer set both proxi-
mally and to a larger degree distally. Hind femur un-
armed.
Abdomen: tergites without particulars. Cercus (fig.
16) slender, proximal half slightly conical, distal half
almost cylindrical and gradually incurved; apex dor-
sally somewhat flattened, inner and outer margins
provided with a series of 2-6 small teeth and termi-
nating into a strong apical tooth. Subgenital plate
short, wide, lateral margins converging posteriorly,
postero-lateral edges not or barely produced and pos-
terior margin short, straight or slightly emarginate.
Coloration: straw-yellow to yellowish green.
Vertex, anterior half of pronotum, tergites and legs
finely speckled rusty-brown. Antennae finely annu-
lated. Occiput sometimes with spots closer set medi-
ally, median line always unspotted. Pronotum, on ei-
ther side, with an ill defined yellowish streak, in
metazona bordered medially reddish brown. Elytra
pale brown, stridulatory vein and pre-apical spot dark
brown. Abdomen of general colour, either provided
or not with a median blackish band and an ill defined
yellow lateral streak; median band composed of a pair
of spots along anterior margin of tergites, extending
and narrowing posteriorly without reaching posterior
margin of tergites; median line of general colour, un-
spotted. Cercus with apex black. Legs sometimes with
lower keels and a lateral incomplete stripe of hind fe-
mur blackish.
Female. — Pronotal dorsum almost cylindrical,
Sl:
metazona not or scarcely raised. Elytra completely
covered by pronotum, lateral, widely separated from
each other dorsally, no stridulatory apparatus. Basal
fold of lower ovipositor valve strong, inflated, pro-
truding ventrally and arched upward anteriorly, lon-
gitudinally impressed, forming with gonangulum an
elongate shallow anteroventrally-facing groove.
Coloration as in male, sometimes more unicolorous.
Measurements. — Body d 17.0-20.0, ® 16.0-21.0;
pronotum d 4.3-5.2, ® 5.5-6.0; elytron d 2.0- 2.4,
2 0.0-0.0; hind femur d 17.0-17.5, 2 18.8-20.0;
ovipositor 8.1-9.7.
Bioacoustics. — Male calling song consists of an
echeme (Heller 1984; 1988; fig. 34), repeated at
intervals of mostly 2-4 s (range 1-7 s; 22-24°C). Each
echeme composed of 6-8 (range 4-11) syllables pro-
duced in a frequency of about 12-18 Hz (22-26°C).
Impulse number per syllable from 6-15, first syl-
lable(s) often containing more impulses than follow-
ing ones. Impulses produced during closing move-
ment of fore wings only, opening being silent.
Maximum of frequency spectrum 20-30 kHz.
Distribution and ecology
The song of this species has been observed at the
localities of the material listed above but also at many
other localities: Dhomokos (near Metallion; A.
Stefanos; Perivoli; Makrirakhi; between Dhomokos
and Neo Monastiri); Farsala (very abundant near
Vrisia; Khalkiadhes); Larissa (Khara; Zappion; Nees
Karies; Larissa town); Kardhitsa (Sofadhes;
Kombelos; eastern side Kardhitsa town). So far the
range of the species extends from northeastern
Central Greece (Fthiotis: Dhomokos) north- and
northwestwards into Thessalia, reaching the town of
Larissa and Mouzaki, beyond the town of Kardhitsa.
Localities referring to preserved material are mapped
as solid squares, those based on acoustic observation
as open ones (fig. 43).
We found the species on thistles and other plants
mainly on the wayside and adjacent fields. It was
common in the plains S of Dhomokos, around Neo
Monastiri and N of Farsala and locally very abun-
dant.
The range of P. mariannae is overlapped by those of
veluchianus and propinquus. At most sites only one of
these species occurs, at some more than one. Near
Dhomokos, NW of Vrisia and some other localities
the new species was found syntopically with P. pro-
pinquus. Along the southern border of the plain S of
Dhomokos a narrow zone of syntopical occurrence of
P. mariannae and veluchianus was detected, the latter
inhabiting the still more southern located slopes that
border this plain. At one spot, Makrirakhi (W of
Dhomokos) we heard simultaneously the calling
songs of even all the three species. Whether P. mari-
annae occurs syntopically with chopardi and thessalicus
is still unknown. The areas where this might occur, N
and W of Larissa, have not yet been explored (fig. 43).
Differential diagnosis
The species is virtually identical with P. chopardi
except for its different song. We found only one
morphological character that might be helpful to dis-
criminate both taxa: spacing of the teeth of the stri-
dulatory file of the left male elytron is greatest in the
mid part of the file with 11-12 in the new species
against 8-10 teeth per 0.5 mm in P. chopardi. Similar
features to differentiate P. chopardi from the thessali-
cus - veluchianus - zimmeri group of species fit ma-
riannae (Willemse 1985a). Bioacoustically, however,
the new species is clearly defined by its song, differing
in several characters from P. chopardi (Heller 1984;
1988). Most reliable and recognizable feature is the
syllable number per echeme, 6-8 in P. mariannae
against 3-4 in chopardi. Echemes with a different
number of syllables are produced sometimes, viz.
down to 4 in P. mariannae and up to 5 in chopardi.
However, this happens quite exceptionally and bes-
ides irregularly by few specimens only. Also the sylla-
ble repetition rate within an echeme is different: P.
mariannae about 13 Hz and chopardi 9 Hz (both at
25°C). Different ambient temperature, however, im-
pedes the use of this character in the field. Finally, the
syllable structure is different: all intervals are equally
spaced in P. mariannae whereas each syllable consists
of two groups of impulses separated by a longer inter-
val in chopardi. Besides the number of impulses per
syllable in P. mariannae appears to be smaller than in
chopardi. Bioacoustical distinction with the P. thessa-
licus - veluchianus - zimmeri group of species is ob-
vious.
WILLEMSE & HELLER: Systematics of Greek Poecilimon
Discussion
It is noted here that most previous records of P.
chopardi (Willemse 1985a: 20) are based on morpho-
logical characters only. Confirmation by the song is
certainly needed.
Etymology
Named in honour of Marianne Volleth, who
found the first specimens.
Poecilimon veluchianus Ramme
Poecilimon veluchianus Ramme, 1933: 549; Werner 1933:
190 (as veluchiensis [sic]); Harz 1969: 150; Willemse
1985a: 20; 1985b: 42. Poecilimon thessalicus veluchianus,
Heller 1984: 103; 1988: 66.
Poecilimon chopardi, Willemse 1982: 178 (partim); 1984: 39
(partim).
Material studied. — Magnisia: Mt. Othris ab. Kokkotoi
1000-1500 m (cw); Fthiotis: 4 km E of Oiti; 1 km N of
Mesokhori (SE of Sperkhiadha) 550 m; 5 km W of
Makrakomi 350 m; N of Vitoli (Makrakomi) 330 m; 4 km
SW of Ag. Stefanos 500 m; Trilofo, NW of Lamia 800 m;
Ag. Ekaterini, 2 km N of Fourka-Pass, N of Lamia 700 m;
Brallos, S of Thermopilai pass; Fokis: 2 km E of Lilaia 350
m (all CH); Polidrosos, 3 km N of the village, wayside, near
bridge crossing Kifisos river (cw); Arta: Mt. Tzoumerka,
Athamani-Theodhoriana 1300 m; S of Theodhoriana 1400-
1800 m (both cw); Kardhitsa: Fanari (cH); Aitolia-
Akarnania: Mt. Valtou near summit Piramidha, N of
Perdhikaki 1300-1600 m (cw).
Distribution and ecology
The range is much larger than previously known
(Willemse 1985a: map 6), extending into eastern
Arta, northeastern Aitolia-Akarnania, extreme north-
ern Fthiotis as far as eastern Magnisia. The occurren-
ce in Mt. Tzoumerka (Arta) was doubtful (Willemse
1985a) but has now been established. Previous re-
cords of syntopic occurrence of P. veluchianus with
other congenerics are confused (Willemse 1982,
1985a). So far known the species may occur together
with P. ornatus, hoelzeli, gracilioides, chopardi, ma-
riannae, jonicus lobulatus, propinquus and most im-
portant zimmeri (see below). Previously we knew P.
veluchianus only from montane habitats but the spe-
cies may occur in the lowlands as well: e.g. Fthiotis:
Vitoli 330 m (Heller 1988); surroundings of Lamia
100 m and many localities in the Sperkhios valley fr-
om Sperkhiadha to Lamia.
Discussion
P. thessalicus, veluchianus and zimmeri are very sim-
ilar in their morphology and bioacoustics. While
their distribution areas do meet, no clear overlaps ex-
ist. Stressing the similarities and allopatric distribu-
tion, these taxa have been considered subspecies by
Heller (1984, 1988), while Willemse (1985a) argued
318
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
that their differences were comparatively large and
sufficiently stable to consider them species.
Recently, syntopic occurrences of P. veluchianus
and zimmeri have been established: based on the male
calling song, a 300 m wide zone was discovered, on
the transition between the plain (P. veluchianus) and
mountain slopes (P. zimmer) near Lilaia (Fthiotis), in
which both taxa occurred; likewise, in the same area
an isolated population of P. zimmeri was discovered
within the range of veluchianus, while the songs of
both taxa was heard on the boundary of both popula-
tions; syntopic occurrence of both taxa was also found
near the summit Piramidha of Mt. Valtou (N of
Perdikhaki, NE Aetolia-Akarnania) and the type lo-
cality of P. gracilioides on Mt. Tzoumerka (S of
Theodhorania, E of Arta). In all these localities no hy-
brids were found. We believe that the syntopic occur-
rence of veluchianus and zimmeri and the apparent
absence of hybrids provides a strong enough argu-
ment to consider these taxa species and not subspe-
cies.
Recent experimental studies on interbreeding
between P. thessalicus and veluchianus reveal that they
mate readily but do not have offspring (K. Reinhold,
Erlangen, pers. comm.). Though syntopic occurrence
of these taxa has as yet not been established both au-
thors now agree that both taxa are best reflected by a
species rather than subspecies level. Specimens from
Fthiotis and Magnisia are remarkably slender, small
in size and the armature of the male cercus is restrict-
ed to the inner apical margin. Their song is not dis-
tinctive from the remainder of the range. So far, these
populations are considered conspecific with P. velu-
chianus, but further studies are currently in progress.
Poecilimon propinquus Brunner von Wattenwyl
(figs. 17-18)
Poecilimon propinquus Brunner von Wattenwyl, 1878: 44
(partim); Ramme 1933: 553; Willemse 1982: 185; 1984:
40; Heller 1984: 104; Willemse 1985a: 19; 1985b: 43;
Heller 1988: 68; 1990: 144.
Material studied. — Magnisia: Mt. Othris above Kokkotoi
1100-1500 m (cw); Larissa: 2 km N of Vrisia 450 m; 4 km
N of Zappion; Fthiotis: Dhomokos 500 m (all cH); Attiki,
Skinios ; Mt. Parnis 1200 m; Avlon (all cw).
Distribution and ecology
Previously known only from the eastern
Peloponnisos, Attiki and Evvoia (Willemse 1984:
map 27), the range of this species extends remarkably
more northward reaching the districts of Larissa and
Magnisia. In Fthiotis and Larissa we found the species
on thistles and other plants on the wayside and ad-
jacent fields, on Mt. Othris mainly on ferns.
Syntopic occurrence of P. propinquus with marian-
314
nae has regularly been observed (see mariannae). P.
propinguus and veluchianus were found together in
the northern hills of the Sperkhios valley and in Mt.
Othris. In most of these places one species was ex-
tremely predominant or occurring without competi-
tors at one spot and with true syntopic occurrences at
the borderlines only.
Discussion
The apical margin of the male cercus was known to
be convex (fig. 17, Ramme 1933, Willemse 1982)
but in the northern specimens from Magnisia, Larissa
and Fthiotis it is scarcely convex or even straight (fig.
18). Bioacoustics are somewhat variable and for the
time being we consider these populations conspecific
as the song seems identical.
Poecilimon aegaeus Werner
(figs. 19-20, 35-39)
Poecilimon aegaeus Werner 1932: 2; 1933: 403; Ramme
1933: 552; Willemse 1982: 188; 1984: 41; 1985b: 42;
Heller 1988: 69.
Material studied. — Holotype d : Graecia Mykonos iv 27,
coll. Werner, Holo-Typus, Poecilimon aegaeus Werner 3
det. Werner. — Allotype 9 (juvenile): similar labels with the
date 13 iv 27 and Allo-Typus; paratypes 1d 12: Coll. Br. v.
W. Syra Erber, det. Br.v.W. Poecilimon propinquus, 5867,
Poecilimon aegaeus Werner Ramme det (all NHMW).
Additional descriptive notes
Male (figs. 35-36). — Integument glossy, general
habitus and shape of pronotum much as P. propin-
quus. Cercus incurved over all its length, apical part
about evenly tapering, inner margin with a few coar-
se teeth, outer margin with more numerous finer
teeth (figs. 19-20). Stridulatory file of left elytron of
holotype (figs. 38-39) with about 55 teeth, spacing
largest in mid part of file, 8 per mm, shortest distan-
ce between proximal and distal most tooth 3.16 mm,
greatest width 0.097 mm. Coloration much as in P.
propinquus.
Female. — Basal fold of lower ovipositor valve (Sira
©, fig. 37) robust, protruding vertically downward
and impressed in the middle forming with gonangu-
lum an elongate groove resembling P. propinquus.
Coloration also much as in that species.
Measurements (juvenile allotype omitted). — Body
8 21.1-21.2, 2 21.2; pronotum d 6.8-7.0, 2 7.0;
hind femur d 15.2-15.9, 2 17.0; ovipositor 8.7.
Bioacoustics. — Unknown.
Poecilimon athos Tilmans et al.
Poecilimon athos Tilmans, F. Willemse & L. Willemse,
1989727:
Material studied. — Type series.
Additional descriptive notes
Auditory spiracle tiny, round, diameter as that of
proximal segments of antennal flagellum. Female stri-
dulatory apparatus: elytra small, widely separated
from each other, lacking stridulatory pegs or plec-
y peg P
trum.
ACKNOWLEDGEMENTS
For loan, collecting material, discussions and further
cooperation we are much in dept to the following persons:
Dr R. Achmann, Erlangen; Dr J. P. Duffels, Amsterdam;
Professor Dr S. Drosopoulos, Athens; Dr A. Kaltenbach,
Vienna; Drs V. Kokkinos, Athens; Dr A. Nadig, Chur; Mr
K. Reinhold, Erlangen; Mr J. Smid and Mrs F. Smid-Elbers,
Arnhem; Professor Dr J. Szijj, Essen; Mr J. Tilmans and Mrs
J. Tilmans-Smid, Warmond; Dr M. Volleth, Erlangen; Drs
L. Willemse, Eygelshoven.
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Harz, K., 1969. Die Orthopteren Europas. 1: i-xx, 1-749. —
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Heller, K.-G., 1984. Zur Bioakustik und Phylogenie der
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Heller, K.-G., 1990. Evolution of song pattern in East
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C. F. Rentz (eds), The Tettigoniidae: biology, systematics
and evolution: 130-151. — Crawford House Press,
Bathurst.
Heller, K.-G. & K. Reinhold, 1992. A new bushcricket of
the genus Poecilimon from the Greek islands (Orthoptera:
Phaneropterinae). — Tijdschrift voor Entomologie 135:
163-168.
Ingrisch, S. & D. Pavicevic, 1985. Zur Faunistik,
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von Nordost-Griechenland. — Mitteilungen der
Miinchener entomologischen Gesellschaft 75: 45-77.
Karabag, T., 1950. Five new species of Poecilimon Fischer
(Orthoptera: Tettigoniidae) from Turkey. — Proceedings
of the Royal Entomological Society of London, Series B
19: 150-155.
WILLEMSE & HELLER: Systematics of Greek Poecilimon
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Sitzungsberichte der kaiserlichen Akademie der
Wissenschaften, mathematischnaturwissenschaftliche
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les Orthoptères de Grèce. — Biocosme mésogéen 5: 1-12.
Ramme, W., 1933. Revision der Phaneropterinen-Gattung
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dem zoologischen Museum in Berlin 19: 497-575.
Tilmans, J., F. Willemse & L. Willemse, 1989. The
Orthoptera of Mount athos, Greece, with description of
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Werner, F., 1929. Repulia, Amphibia, Orthoptera,
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naturwissenschaftliche Klasse [I] 147: 157-173.
Willemse, F., 1982. A survey of the Greek species of
Poecilimon Fischer, 1853 (Orthoptera, Ensifera,
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155-203.
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Received: 1 October 1991
Revised manuscript accepted: 26 August 1992
315
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
316
NEWT DESCRIBED IN) Ds CHR OOR
ENTOMOLOGIE, VOLUME 135
ACARI
Albia (Albiella) brokenensis Smit … … 105
Arrenurus (Megaluracarus) roobeeki Smit.............. 108
Arrenurus (Megaluracarus) vanderpalae Smit … … 108
Aspidiobates unidorsalis Smit … nnen 99
Australionicola Smit (new subgenus)...................... 99
IAlyarachımarpalustnsSoen ERE, Oil
Kallo bALENCOO LIN mige 99
Koenikea (Notomideopsis) purpurea Smit … 105
Recifella (Recifellida) pectinatus Smit … 103
Unionicola (Australionicola) Smit … … … 99
Unionicola (Australionicola) hammeni Smit.......... 100
Unionicola (Pentatax) davidsi Smit ...................... 100
COLEOPTERA
Cyclocephala amplitarsis Ratcliffe … …… … 179
CyclocephalataeltoidesRaidite ae en 181
@yelocephalasianı Ratclitfe nn nee 183
Cyclocephala nodanotherwon Ratcliffe................... 184
Cyclocephala pseudoconfusa Ratcliffe … … … … … … … 185
Cyclocephala rondoniana Ratcliffe … … … 185
Cyclocephala sarahae Ratcliffe … …… … … 187
Cyclocephala sarpedon Rarcliffe… … … … … … …… 188
Cyclocephala schmitzorum Ratcliffe....................... 189
Peltodytes pekinensis van Vondel. … 285
HEMIPTERA
Cylindrostethus akanthinos Chen & Nieser … … … Si
Limnometra aploa Nieser & Chen … 16
Limnometra arachnis Nieser & Chen ..................... 17
Limnometra genitalis Nieser & Chen ..................... 18
Limnometra hysterema Nieser & Chen................... 23
TimnomerralepraNiesenéciChen. "me 19
Limnometra melanochroa Nieser & Chen............... 20
Limnometra monochroma Nieser & Chen .............. 21
Limnometra pseudoinsularis Nieser & Chen ........... 21
Limnometra poliakanthina Nieser & Chen … … …. 23
Metrocoris breviculus Chen & Nieser … … … … … 156
Potamometropsis anomalis Chen & Nieser … … … … 154
Rheumatometroides drepanephoros Chen & Nieser 160
Rheumatometroides makraitos Chen & Nieser ...... 159
Rheumatometroides serena Lansbury ......... 1
Tenagogonus akanthinos Chen & Nieser .............. 150
Ventidius kurtokalami Chen & Nieser ................. 159
Ventidius xiphibion Chen & Nieser ..................... 156
VentidiuspoelAGhentaNiesers ot. seat. ee nonne 157
LEPIDOPTERA
Euchromius circulus Schouten ............................ 201
Euchromius confusus Schouten … nanne 211
Euchromius minutus Schouten … … 207
Euchromius ornatus Schouten … en 201
MiyakeassinevdSchouten em ee 238
Phyllonorycter dentifera Noreika … … … … 39
ODONATA
Oreagrion pectingi Brooks & Richards … … … … … 141
ORTHOPTERA
Poecilimon erimanthos Willemse & Heller … … … 307
Poecilimon gracilioides Willemse & Heller … … … 303
Poecilimon mariannae Willemse & Heller............ 311
Poecilimon paros Heller & Reinhold … … … … … … 163
PYCNOGONIDA
ACheACOLUMNANIISIOCk: se MOINE 116
Anoplodactylus spurius Stock … ennen 132
Buryoyderplatyspina:Stock sne Eee 113
INymphonibullatum Stock re ee 121
INymphonnnaeguipes Stock nn 124
Nymphon vulcanellum Stock … … … iii 126
Pycnogonum (Retroviger) ornans Stock … … … … …. 136
TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 135, 1992
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Tijdschrift voor Entomologie
Volume 135, no. 2
Articles
141 S.J. Brooks & S. J. Richards
A new species of Oreagrion (Odonata: Coenagrionidae)? montane damselflies from
New Guinea.
145 P. Chen & N. Nieser
Gerridae, mainly from Sulawesi and Pulau Buton (Indonesia). Notes on Malesian
aquatic and semiaquatic bugs (Heteroptera), Ill.
163 K.-G. Heller & K. Reinhold
A new bushcricket of the genus Poecilimon from the Greek Islands (Orthoptera:
Phaneropterinae).
169 P. J. van Helsdingen & A. E. Decae
Ecology, distribution and vulnerability of Macrothele calpeiana (Walckenaer)
(Araneae, Hexathelidae).
179 B. C. Ratcliffe
New species and country records of Brazilian Cyclocephala (Coleoptera:
Scarabaeidae: Dynastinae).
191 R. T. A. Schouten
Revision of the genera Euchromius Guenée and Miyakea Marumo (Lepidoptera:
Crambidae: Crambinae).
275 B.J. van Vondel
Revision of the Palaearctic and Oriental species of Peltodytes Régimbart (Coleoptera:
Haliplidae).
299 F. Willemse & K.-G. Heller
Notes on systematics of Greek species of Poecilimon Fischer, 1853 (Orthoptera:
Phaneropterinae).
© Nederlandse Entomologische Vereniging, Amsterdam
Published | December 1992 ISSN 0040-7496
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