European Journal of Taxonomy 192: 1-24
http://dx.doi.org/10.5852/ejt.2016.192
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2016 • Jasc hh of M
Research article
urn:lsid:zoobank.org:pub:5C461741-852C-4AEB-9DA3-31B92BB23777
New species of Monepidosis Mamaev, 1966 and Antipodosis gen. nov.,
a closely related genus from New Zealand (Diptera, Cecidomyiidae)
Mathias JASCHHOF
Station Linne, Olands Skogsby 161, SE-38693 Farjestaden, Sweden.
Email: [email protected]
urn:lsid:zoobank.org:author:8B4BllB4-7C33-41AC-A042-AA9903CDC4Bl
Abstract. Three new species of Monepidosis Mamaev, 1966, a Holarctic genus of Porricondylinae
(Diptera, Cecidomyiidae), are described: M. heterocera sp. nov. from Sweden and Germany,
M. scepteroides sp. nov. from Sweden, and M. shikokuensis sp. nov. from Japan. Anew porricondyline
genus, Antipodosis gen. nov., is introduced for eight new species from New Zealand, named A. australis
gen. et sp. nov., A. elongata gen. et sp. nov., A. granvillensis gen. et sp. nov., A. pure or a gen. et sp. nov.,
A. rakiura gen. et sp. nov., A. rotoiti gen. et sp. nov., A. rotoroa gen. et sp. nov., and A. waipapa
gen. et sp. nov. Male genitalic morphology indicates that Monepidosis and Antipodosis gen. nov. are
closely related, together forming the Monepidosis group of genera, which stands out from the other
Porricondylini. Monepidosis spatulata Spungis, 2006, a species originally described from Latvia and
Lithuania, is for the first time reported to occur in Sweden.
Key words. Porricondylinae, Palearctic region, Australasian region, new genus, new species.
Jaschhof M. 2016. New species of Monepidosis Mamaev, 1966 and Antipodosis gen. nov., a closely related
genus from New Zealand (Diptera, Cecidomyiidae). European Journal of Taxonomy 192: 1-24. http://dx.doi.
org/10.5852/eit.2016.192
Introduction
A recent revision of the genus Monepidosis Mamaev, 1966 recognized 14 valid species, 13 in the
Palearctic and one in the Nearctic region (Jaschhof & Jaschhof 2013; Gagne & Jaschhof 2014). Another
species of Monepidosis was described from Sweden soon after (Jaschhof & Jaschhof 2015). Previous
authors expected a considerable number of unnamed Monepidosis still to be found in the Holarctic
realm (Spungis 2006; Jaschhof & Jaschhof 2013), an expectation validated here with the description of
three new species from Sweden, Germany and Japan. Monepidosis are morphologically distinct from
the other Porricondylini (Cecidomyiidae, Porricondylinae) in that male genitalic structures, such as
gonostyli, gonocoxites, parameres, and ejaculatory apodeme, are heavily armed in a specific manner
(Jaschhof & Jaschhof 2013). Most notably, parameres are present as two pairs of large, sclerotized tusks
interlinked with each other, and the ejaculatory apodeme is developed into a long, sclerotized rod whose
apex is variously modified (Jaschhof & Jaschhof 2013: fig. 88A). Genitalic structures similar to those in
Monepidosis have now been found in eight different species of undescribed Porricondylinae from New
Zealand, which are obviously closely related to each other and thus classified together in a discrete new
genus, Antipodosis gen. nov. Monepidosis and Antipodosis gen. nov. appear to be, if not sister groups,
members of a distinct natural subset of the tribe Porricondylini, called here the Monepidosis group.
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European Journal of Taxonomy 192: 1-24 (2016)
In the present paper, the newly discovered taxa are described and named, so as to continue with the
taxonomic inventory of Palearctic and especially Swedish Porricondylinae within the framework of
‘The Gall Midge Project’ t http://www.stationlinne.se/sv/forskning/the-gall-midge-project . accessed on
25 Nov. 2015), and to take a first step into the study of porricondyline biodiversity in Australasia. This
study shows that porricondyline interrelationships are more revealing when studied globally.
Material and methods
Specimens for this study, exclusively male adults, were either picked out from general insect samples
obtained by Malaise traps or were collected by sweepnet and aspirator. Most of the specimens came
from various different projects initiated by the author, specifically ‘The Gall Midge Project’, an ongoing
venture to inventory the Cecidomyiidae of Sweden (2015-2017); the ‘Testremiinae of Japan’ project
(1998-2000; see Jaschhof 2000); and the ‘Testremiinae of New Zealand’ project (2001-2003; see
Jaschhof & Jaschhof 2003). Other specimens were provided by ‘The Swedish Malaise Trap Project’
(SMTP; see http://www.stationlinne.se/sv/forskning/the-swedish-malaise-trap-project-smtp . accessed
on 25 Nov. 2015); were received on loan from the Senckenberg Deutsches Entomologisches Institut,
Miincheberg, Germany (SDEI); or were donated by Dieter Doczkal, Zoologische Staatssammlung
Mtinchen, Germany.
Types and other voucher specimens of the species discussed here are available as microscopic
preparations (Canada balsam inclusions) and shall be finally deposited in the following collections:
Entomological Collection of Kyushu University, Fukuoka, Japan (KUEC); Naturhistoriska Rilcsmuseet
Stockholm, Sweden (NHRS); New Zealand Arthropod Collection, Tamaki, Auckland, New Zealand
(NZAC); and SDEI.
The procedures of specimen preparation and identification as well as the morphological terminology used
in adult Porricondylinae were recently described by Jaschhof & Jaschhof (2013). Several morphological
terms, some of which specific to the porricondylines treated here, are explained in Figs 2, 10 and 12 of
the present paper, with the following abbreviations used:
ctr pr
= central process
d
= dorsal
ej ap
= ejaculatory apodeme
gcx ap
= gonocoxal apodeme
gcx em
= gonocoxal emargination
gcx pr
= gonocoxal process
par
= paramere
V
= ventral
Since the male gonocoxites of almost all Porricondylinae, including Monepidosis and Antipodosis
gen. nov., are largely merged into a single structure, they are described as a unit. Body length was
measured from the head to the genitalia. Arrows (j) are used to highlight important diagnostic
characters in both the taxonomic descriptions and illustrations. The descriptive sections, titled here
‘Other characters’, mention only those characters that are not referred to in the diagnoses and differential
diagnoses, respectively. Taxa are treated in alphabetical order.
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JASCHHOF M., Taxonomy of Monepidosis and Antipodes is gen. nov.
Results
Class Insecta Linnaeus, 1758
Order Diptera Linnaeus, 1804
Suborder Bibionomorpha Hennig, 1954
Family Cecidomyiidae Newman, 1834
Subfamily Porricondylinae Kieffer, 1913
Genus Antipodosis gen. nov.
urn:lsid:zoobank.org:act:B8BF652D-AFFF-418C-829E-608741C68503
Type species
Antipodosis australis gen. et sp. nov., described below.
Diagnosis
Larvae and females of Antipodosis gen. nov. are unknown, so this genus is based solely on characters
of males. Antipodosis gen. nov. are small, inconspicuous Porricondylini, exhibiting the characters
typical of that tribe and coming under the group of genera with 14 flagellomeres and without basitarsal
spines (“group Aa” in Jaschhof & Jaschhof 2013). Antipodosis gen. nov. stands out from most other
Porricondylini in that vein CuA^ runs parallel to CuA, rather than approaching or joining it (Fig. 1A
versus IB), and the eye bridge is longer dorsally (5-10 versus 2-3 ommatidia). The structure of the
genitalia is genus-specific. Most notably, the two gonocoxal apodemes, which in other Porricondylini
are long bars separated from one another (Fig. 12), tend to merge into a single plate (Fig. 2); parameres
are typically present as two pairs (a ventral and a dorsal pair) of strongly sclerotized tusks interlinked
with each other (Fig. 2); and the ejaculatory apodeme, whose length exceeds that of the gonocoxites, is
typically a strongly sclerotized rod with apical modifications.
Fig. 1 . Male morphology of Antipodosis gen. nov. and Monepidosis Mamaev, 1966. A. Wing of
A. granvillensis gen. et sp. nov., holotype, setae omitted. B. Wing ofM shikokuensis sp. nov., holotype,
setae omitted. C. Fourth flagellomere of A. rakiura gen. et sp. nov., holotype, lateral. D. Fourth
flagellomere ofM scepteroides sp. nov., holotype, lateral. Scale lines: A-B = 0.50 mm, C-D = 0.05 mm.
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
Differential diagnosis
The genus Antipodosis gen. nov. is similar to Monepidosis (Jaschhof & Jaschhof 2013), differing from
it as follows: the eye bridge is longer; sensory hairs (= setae with hooded sockets) on the flagellomeres
are dispersed rather than aligned to form a single whorl (Fig. 1C versus ID); wings are wider (with
length/width ratios usually < 3.0); CuAj does not approach but runs parallel to CuA 0 (Fig. 1A versus
IB); the gonostylar apex bears typically a pectinate tooth (Figs 2A, 7A), but never a large plate-like
spine (Figs 10A-B, 1 IB); gonocoxites have a distinct ventral emargination (Fig. 2A) and have no central
processes (Fig. 10A); and the anterior portions of the gonocoxal apodemes are either short and separated
(Fig. 6A), or merged into a single, long plate (Fig. 2A).
Etymology
The name Antipodosis is composed of Antipod -, from the Tatin antipodes , for ‘antipodes’, and the
ending -osis, from Monepidosis , a closely related genus. Gender is feminine.
Other characters
Body length. 1.7-2.3 mm.
Head. Postfrons asetose. Antenna longer than body. Scape and pedicel usually yellowish, lighter than
flagellum. Circumfila on flagellomeres 1 to 11-14, evenly ring-shaped or slightly sinuous, in A. rotoiti
gen. et sp. nov. with short posterior extensions. Neck of fourth flagellomere longer than node; node apart
from circumfilum with microtrichia, short setae forming a basal whorl, numerous long sensory hairs
(Fig. 1C). Palpus as long as head height or longer, 4 subcylindrical segments.
Thorax. Scutum with a few lateral and dorsocentral setae. Both anepisternum and anepimeron setose.
Wing (Fig. 1A). Tonger than body. Length/width 2.5-2.9 (in A. australis gen. et sp. nov. exceptionally
3.1). Costal cell narrow, reinforced. Rs strongly oblique, in line with R_. M usually absent, in A. waipapa
gen. et sp. nov. a remnant M present at wing margin.
Legs. Densely covered with narrow scales. Claws crescent-shaped, with 1 large and 2-3 smaller teeth
basally. Empodia rudimentary, as far as known.
Abdomen. Tergites 2—5(—6) varyingly strongly desclerotized at center, setae aligned in rows along
margins; other tergites evenly sclerotized, setae aligned in 1 transverse row. Sternite 1 unsclerotized,
asetose, other sternites evenly sclerotized, setae dispersed. Pleural membrane very sparsely setose.
Genitalia (Figs 2-9). Ninth tergite either subtrapezoid or rounded posteriorly. Gonostylus either elongate,
tapered towards apex, or flattened, with broadly rounded apex; typically a pectinate tooth apically,
1-2 bristles medially; basolateral apodeme large. Gonocoxites: ventral emargination approximately
U-shaped, with glabrous, sclerotized rim; medial bridges sometimes with membranous outgrowths that
occupy parts of the ventral emargination (Figs 7A, 8A, 9A). Ventral parameres usually bent laterally
or dorsolaterally, occasionally interconnected mediobasally (Figs 3B, 9B); dorsal parameres usually
bent dorsally (thus often hardly visible in ventral view); parameral apodemes usually large. Base of
ejaculatory apodeme either widened (visible in ventral view. Fig. 5B) or flattened (visible in lateral view).
Accessory gland ducts, or their merging points with ejaculatory apodeme, distinct. Both hypoproct and
cerci present as two setose, medially merged lobes (omitted in illustrations); cerci longer than hypoproct.
Distribution and phenology
The genus Antipodosis gen. nov. is to present knowledge endemic to New Zealand. Of eight species
identified, two occur on the North Island and six on the South Island including Stewart Island. Almost
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JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
all the specimens known of Antipodosis gen. nov. were collected in native forest in austral summer
(Nov.-Jan.).
Phylogenetic remarks
The peculiarity of Antipodosis gen. nov. that gonocoxal apodemes tend to merge into a large plate
is a derived character state not known in other Porricondylini, thus an autapomorphy. From what is
currently known of world Porricondylini, it appears that Antipodosis gen. nov. is most closely related to
Monepidosis. Compelling evidence of this relationship is provided by the parameres and the ejaculatory
apodeme, whose basic structure is identical in the two genera. Both are therefore combined in what
is here called the Monepidosis group. The relationships of this group to other Porricondylini remain
obscure for the time being. With respect to male morphology, Antipodosis gen. nov. has retained more
ancestral traits than Monepidosis. For example, in Antipodosis gen. nov. the two gonocoxites are largely
separated by the ventral emargination, whereas in Monepidosis they are lengthwise connected, with
the connecting bridge even further modified to bear a pair of processes. Furthermore, in Antipodosis
gen. nov. the gonostylar apex is equipped with a pectinate tooth of fine, separate spines, which is the
structure found in many other Porricondylinae and Porricondylini, whereas in Monepidosis the spines
are merged into a large, plate-like tooth that encircles much of the gonostylus’ distal half. Concurrent
in Antipodosis gen. nov. and Monepidosis , the basic patterns of parameres and ejaculatory apodeme
undergo various modifications, which may be so pronounced that the generic affiliations of the respective
species are obscured (see Antipodosis elongata gen. et sp. nov., Fig. 3; Monepidosis shikokuensis sp.
nov., Fig. 12). One may argue that such extreme variations are likely to evolve over long periods of time
and both Antipodosis gen. nov. and Monepidosis might be phylogenetically old lineages.
Antipodosis australis gen. et sp. nov.
urn:lsid:zoobank.org:act:45076E74-E261-4A53-804F-8264851F5133
Fig. 2A-B
Diagnosis
A combination of genitalic characters is specific to A. australis sp. nov., as follows (Fig. 2A-B). Of the
gonocoxites (Fig. 2A), the processes are small but prominent (j); the ventral emargination, whose shape
is reminiscent of an inverted bell, has a broad glabrous rim that almost reaches the anterior gonocoxal
margin (j), and the apodemes are merged into a large subrectangular plate (j). The ejaculatory apodeme
has a distinct subapical extension (j, Fig. 2B).
Differential diagnosis
Antipodosis australis gen. et sp. nov. resembles A. pureora gen. et sp. nov. in having elongate, slender
gonostyli; a long, plate-like gonocoxal apodeme; two pairs of unmodified parameral tusks, and a single-
pointed, straight ejaculatory apodeme. Characters to differentiate between the two species are provided
by the gonocoxal emargination, which in A. australis gen. et sp. nov. is bell-shaped (Fig. 2A) and in
A. pureora gen. et sp. nov. U-shaped (Fig. 5A), and by the ejaculatory apodeme, whose base is flattened
in A. australis gen. et sp. nov. (not visible in the ventral view depicted in Fig. 2B) and widened in
A. pureora gen. et sp. nov. (Fig. 5B).
Etymology
The name australis is a Latin adjective meaning ‘southern’, which refers to the distribution of this
species in the two southern main islands of New Zealand.
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
Type material
Holotype
NEW ZEALAND: S, South Island, Buller, Lake Rotoroa, mixed podocarp / southern beech forest,
450 m, 15 Jan. 2001, Malaise trap “15”, Department of Conservation St. Arnaud (NZAC).
Paratype
NEW ZEALAND: 1 S, Stewart Island, Christmas Village Hut, 46.74° S, 167.97° E, 18 Jan. 2000,
Malaise trap, R.K. Didham (NZAC).
Other characters
Body length. 2.0 mm
Head. Eye bridge 5-6 ommatidia long dorsally. Circumfila on flagellomeres 1 to 13-14. Neck of fourth
flagellomere 1.8-1.9 times longer than node. Palpus as long as height of head.
Wing. Length/width 3.0-3.1.
Legs. Empodia rudimentary.
Fig. 2. Antipodosis australis gen. et sp. nov., S, holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
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JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Terminalia (Fig. 2A-B). Ninth tergite subtrapezoid, medially unsclerotized, asetose. Gonostylus
strongly tapered towards apex, pectinate tooth narrow (Fig. 2A). Gonocoxites with small protuberances
on ventroposterior margin; apodemes protrude clearly beyond ventroanterior margin (Fig. 2A). Ventral
parameral tusks bent dorsolaterally, dorsal tusks bent dorsally (Fig. 2B).
Antipodosis elongata gen. et sp. nov.
um:lsid:zoobank.org:act:CD29F5F3-E563-4898-B8ED-E3D69204939B
Fig. 3A-B
Diagnosis
Genitalic structures, such as gonocoxites, parameres and ejaculatory apodeme, are conspicuously
elongated (Fig. 3A-B), making A. elongata gen. et sp. nov. the most deviant species of Antipodosis
gen. nov. Adding to its exceptionality, ventral and dorsal parameres are largely merged with each other,
Fig. 3. Antipodosis elongata gen. et sp. nov., holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
leaving only the extreme apices separated (j, Fig. 3B). The ejaculatory apodeme is three-pointed apically
(|, Fig. 3B). The gonostylus, which is strongly flattened, has a fringe of short, thick microtrichia at the
broadly rounded apex (j, Fig. 3A).
Etymology
The name elongata , a Tatin adjective meaning ‘elongate’, refers to the genitalia of this species.
Type material
Holotype
NEW ZEATAND: <$, South Island, Buller, Lake Rotoiti, southern beech forest, 640 m, 31 Jan. 2001,
Malaise trap “2”, Department of Conservation St. Arnaud (NZAC).
Other characters
Body length. 2.2 mm .
Head. Eye bridge 7-8 ommatidia long dorsally. Circumfila on flagellomeres 1-14. Neck of fourth
flagellomere 1.5 times longer than node. Palpus E6 times longer than height of head.
Wing. Length/width 2.9.
Legs. Acropods missing, so length of empodia unknown.
Terminalia (Fig. 3A-B). Ninth tergite broadly rounded posteriorly. Gonocoxites (Fig. 3A): processes
small, subtriangular; ventral emargination large, U-shaped, with glabrous rim basally; posterior portions
extensively unsclerotized; apodemes merged anteriorly to form subrectangular plate that protrudes
clearly beyond ventroanterior gonocoxal margin (j). Apex of ejaculatory apodeme bent ventrally
(Fig. 3B). Parameres directed posteriorly, only apices bent slightly dorsally; apodemes small (Fig. 3B).
Antipodosis granvillensis gen. et sp. nov.
urn:lsid:zoobank.org:act:6CF5586C-2552-4D08-AD8F-F6F208ADE6D7
Figs 1A, 4A-C
Diagnosis
Antipodosis granvillensis gen. et sp. nov. is distinguished from the congeneric species by two unique
characters: the gonostylus bears apically a single, tooth-like spine (j. Fig. 4A), and the dorsal parameral
tusks are two-branched (j, Fig. 4B).
Etymology
The name granvillensis , an adjective, is derived from Granville State Forest in the West Coast region of
South Island, which is the type locality of this species.
Type material
Holotype
NEW ZEALAND: South Island, West Coast, Ahaura, Granville State Forest, hard beech forest, 170—
250 m, Dec. 1994, Malaise trap “2 GUL 7K”, J. Hutcheson (NZAC).
Paratype
NEW ZEALAND: 1 South Island, Buller, Lake Rotoroa, mixed podocarp/southern beech forest,
450 m, 15 Jan. 2001, Malaise trap “15”, Department of Conservation St Arnaud (NZAC).
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JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Other characters
Body length. 2.2 mm
Head. Eye bridge 6-7 ommatidia long dorsally. Circumfila on flagellomeres 1-11. Neck of fourth
flagellomere 1.4 times longer than node. Palpus 1.3 times longer than height of head.
Wing (Fig. 1A). Length/width 2.7.
Legs. Empodia rudimentary.
Terminalia (Fig. 4A-C). Ninth tergite subtrapezoid. Gonostylus elongate, slightly curved, strongly
tapered towards apex (Fig. 4A). Gonocoxites (Fig. 4A) broadly rounded ventroanteriorly; processes
inconspicuous, subrectangular (j); ventral emargination evenly U-shaped, with broad glabrous
rim basally; apodemes short, reaching just to ventral gonocoxal margin, slightly bilobed anteriorly.
Fig. 4. Antipodosis gi'anvillensis gen. et sp. nov., S. A. Genitalia, ventral, holotype. B. Apex of ejaculatory
apodeme, ventral, paratype. C. Parameres and ejaculatory apodeme, ventral, holotype. Scale lines:
0.05 mm
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
Ejaculatory apodeme with ovoid apical extension, two-pointed, the ventral point slightly bent ventrally
(|, Fig. 4B-C), basal portion poorly sclerotized. Ventral parameral tusks bent dorsolaterally, dorsal tusks
bent dorsoposteriorly; apodemes large (Fig. 4B).
Antipodosis pureora gen. et sp. nov.
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Fig. 5A-B
Diagnosis
A combination of genitalic characters is specific to A. pureora gen. et sp. nov., as follows (Fig. 5A-B). Of
the gonocoxites (Fig. 5A), the processes are small and subtriangular (j), the large ventral emargination
is perfectly U-shaped (j), and the apodemes are merged into a large plate with rounded margins (j). The
base of the ejaculatory apodeme is strongly widened (j, Fig. 5B), which is unique in Antipodosis gen. nov.
Differential diagnosis
Antipodosis pureora gen. et sp. nov. is similar to both A. australis gen. et sp. nov. and A. rotoroa gen. et
sp. nov. The three species differ from each other in the ejaculatory apodeme, which is strongly widened
basally in A. pureora gen. et sp. nov. (Fig. 5B), markedly widened subapically in A. australis gen. et
sp. nov. (Fig. 2B), and two-pointed apically in A. rotoroa gen. et sp. nov. (Fig. 8B). Other interspecific
distinctions are described under each of the species.
Fig. 5. Antipodosis pureora gen. et sp. nov., holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
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JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Etymology
The name pure ora is from the Maori language, referring to Pureora Forest, one of the largest intact tracts
of native podocarp forest in the North Island and collection site of this species.
Type material
Holotype
NEW ZEALAND: S, North Island, Taupo, Pureora Forest, Waipapa Reserve, podocarp forest, 570 m,
8 Dec. 1983, Malaise trap, J. Hutcheson (NZAC).
Other characters
Body length. About 1.7 mm.
Head. Eye bridge 4-5 ommatidia long dorsally. Apices of ante nn ae missing. Neck of fourth flagellomere
1.7 times longer than node. Palpus longer than height of head.
Wing. Length/width 2.7.
Legs. Acropods missing, so length of empodia unknown.
Terminalia (Fig. 5A-B). Ninth tergite subtrapezoid, medially unsclerotized, asetose. Gonostylus
elongate, strongly tapered towards apex, pectinate tooth narrow (Fig. 5A). Gonocoxal emargination
with glabrous rim basally; apodemes protrude beyond ventroanterior margin (Fig. 5A). Ejaculatory
apodeme pointed apically, straight, slightly widened at merging points with accessory gland ducts
(Fig. 5B). Ventral parameral tusks curved dorsolaterally, dorsal tusks directed dorsally (Fig. 5B).
Antipodosis rakiura gen. et sp. nov.
um:lsid:zoobank.org: act: 7D191604-3021-4E5E-9E8D-DDD34DC5 8310
Figs 1C, 6A-B
Diagnosis
A character readily distinguishing A. rakiura gen. et sp. nov. from all other Antipodosis gen. nov. is that
the dorsal portions of the parameres are merged into a rhomboid shaped plate (j, Fig. 6B), with the result
that only one pair of tusks remains ventrally. Other genitalic structures distinctive of this species are the
gonostylus, which is swollen on the basal two thirds and flattened on the apical third (j, Fig. 6A), and
the gonocoxites, which have prominent, rectangular processes (J,).
Etymology
The species epithet rakiura is identical with the Maori name commonly used for Stewart Island where
this species was found.
Type material
Holotype
NEW ZEALAND: S, Stewart Island, Murray Beach, 46.78° S, 168.00° E, 20 Jan. 2001, Malaise trap,
R.K. Didham (NZAC).
Other characters
Body length. 2.0 mm
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
Head. Eye bridge 4-5 ommatidia long dorsally. Circumfila present on flagellomeres 1-12. Neck of
fourth flagellomere 1.6 times longer than node. Palpus 1.2 t im es longer than height of head.
Wing. Length/width 2.8.
Legs. Empodia rudimentary.
Terminalia (Fig. 6A-B). Ninth tergite broadly rounded posteriorly, only medially sclerotized. Gonostylar
apex broadly rounded, pectinate tooth wide (Fig. 6A). Gonocoxites (Fig. 6A): ventral emargination
small, perfectly U-shaped, with glabrous rim basally; dorsal portions with subtriangular projections
on posterior margin; apodemes short, bilobed anteriorly. Ejaculatory apodeme (Fig. 6B): apex pointed,
straight; base flattened; accessory gland ducts unusually distinct. Parameral tusks curved dorsolaterally;
parameral apodemes small (Fig. 6B).
Fig. 6. Antipodosis rakiura gen. et sp. nov., <f, holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
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JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Antipodosis rotoiti gen. et sp. nov.
um:lsid:zoobank.org:act:9BA5DA6F-D8D5-4921-B55B-lCC4C739BE76
Fig. 7A-B
Diagnosis
In this unusual Antipodosis gen. nov., the flattened gonostylus has a broadly rounded apex with wide
pectinate tooth ([, Fig. 7A); the ventral parameral tusks are clearly shorter than the dorsal (j. Fig. 7B);
and the elongate gonocoxites have practically no processes (Fig. 7A).
Etymology
The species epithet rotoiti is from the Maori language, meaning ‘the little lake’, with reference to Take
Rotoiti in northern South Island where this new porricondyline was collected.
Type material
Holotype
NEW ZEALAND: c?, South Island, Buller, Lake Rotoiti, southern beech forest, 640 m, 20 Dec. 2000,
Malaise trap “2”, Department of Conservation St Arnaud (NZAC).
Fig. 1 .Antipodosis rotoiti gen. et sp. nov., c?, holotype. A. Genitalia, ventral. B. Parameres and ejaculatory
apodeme, ventral. Scale lines: 0.05 mm.
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European Journal of Taxonomy 192: 1-24 (2016)
Other characters
Body length. 2.1 mm
Head. Eye bridge 4 ommatidia long dorsally. Circumfila on flagellomeres 1-14, often with short posterior
extensions. Neck of fourth flagellomere 1.1 times longer than node. Palpus 1.5 times longer than height
of head.
Wing. Length/width 2.6.
Legs. Empodia rudimentary.
Terminalia (Fig. 7A-B). Ninth tergite subtrapezoid. Gonocoxites with asetose, subtriangular portion
ventroanteriorly; ventral emargination perfectly U-shaped, with narrow glabrous rim basally; medial
bridges with membranous outgrowths that extend into the emargination; apices of apodemes missing,
but probably merged to form a plate (Fig. 7A). Ejaculatory apodeme bent ventrally; widened subapically
at merging point with accessory gland ducts; flattened basally (Fig. 7B). Ventral parameral tusks almost
straight, directed posteriorly, dorsal tusks bent slightly dorsally (Fig. 7B).
Antipodosis rotoroa gen. et sp. nov.
um:lsid:zoobank.org:act:9C8C71A9-0BCF-4A69-BE42-8ADC45A015C7
Fig. 8A-B
Diagnosis
The gonocoxal processes are more prominent in A. rotoroa gen. et sp. nov. (j, Fig. 8A) than in any other
Antipodosis gen. nov. The gonostylus is slightly curved, tapered towards the apex, and equipped with a
long pectinate tooth (j. Fig. 8A). The apex of the ejaculatory apodeme is two-pointed (j, Fig. 8B). At
the bases of the dorsal parameral tusks are small accessory processes, which are weakly sclerotized and
blunt-ended (j. Fig. 8B).
Differential diagnosis
See A. australis gen. et sp. nov. (Fig. 2) and A. pure ora gent, et sp. nov. (Fig. 5) for two species with
genitalic structures superficially similar to that found in A. rotoroa gen. et sp. nov.
Etymology
The name rotoroa , which means The long lake’ in the Maori language, refers to Lake Rotoroa, whose
vicinity provides the only habitat yet known of this porricondyline.
Type material
Holotype
NEW ZEALAND: S, South Island, Buller, Lake Rotoroa, mixed podocarp/southern beech forest,
450 m, 15 Jan. 2001, Malaise trap “15”, Department of Conservation St Amaud (NZAC).
Paratype
NEW ZEALAND: 1 same data as the holotype (NZAC).
Other characters
Body length. 2.0 mm.
Head. Eye bridge 7-8 ommatidia long dorsally. Circumfila on flagellomeres 1-12. Neck of fourth
flagellomere 1.5 times longer than node. Palpus F3 times longer than height of head.
14
Wing. Length/width 2.6.
Legs. Empodia rudimentary.
JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Terminalia (Fig. 8A-B). Ninth tergite subtrapezoid. Gonocoxites (Fig. 8A): processes subtriangular (j);
ventral emargination large, extends beyond midlength of gonocoxites, largely occupied by membranous
outgrowths of medial bridges (j), with glabrous rim basally; apodemes merged into large, subrectangular
plate that protrudes far beyond anterior gonocoxal margin (j). Apex of ejaculatory apodeme bent ventrally
(Fig. 8B). Ventral parameral tusks only slightly bent, dorsal tusks directed posteriorly (Fig. 8B).
Fig. 8. Antipodosis rotoroa gen. et sp. nov., holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
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European Journal of Taxonomy 192 : 1-24 ( 2016 )
Antipodosis waipapa gen. et sp. nov.
um:lsid:zoobank.org:act:53252B20-F0CE-4F56-B7DF-182BBFC599FA
Fig. 9A-B
Diagnosis
This is the only species of Antipodosis gen. nov., whose gonostyli have no apical structure, such as a
pectinate or solid tooth (Fig. 9A). As another peculiarity, the apical portion of the ejaculatory apodeme
is conspicuously enlarged and spoon-shaped (j, Fig. 9B).
Etymology
The name waipapa , from the Maori language, refers to Waipapa Reserve in Pureora Forest where the
type specimens of this species were collected.
Type material
Holotype
NEW ZEALAND: 3 , North Island, Taupo, Pureora Forest, Waipapa Reserve, shrubland, 570 m, 29 Mar.
1984, Malaise trap, J. Hutcheson (NZAC).
Paratypes
NEW ZEALAND: 2 33 , same data as the holotype, but podocarp forest, 17 Nov. 1983 (NZAC).
Other characters (male)
Body length. 2.3 mm.
Fig. 9. Antipodosis waipapa gen. et sp. nov., 3, holotype. A. Genitalia, ventral. B. Parameres and
ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
16
JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Head. Eye bridge 9-10 ommatidia long dorsally. Scape and pedicel concolorous with flagellum. Neck
of fourth flagellomere 1.8 t im es longer than node. Apices of antennae missing. Palpus 1.6 times longer
than height of head.
Wing. Length/width 2.5. A short M present at wing margin.
Legs. Acropods missing, so length of empodia unknown.
Terminalia (Fig. 9A-B). Ninth tergite rounded posteriorly. Gonostylus flattened, rounded apically
(Fig. 9A). Gonocoxites (Fig. 9A): true processes missing, medial bridges with membranous outgrowths
that extend into the ventral emargination; emargination small, perfectly U-shaped, with glabrous rim
basally; apodemes short, bilobed anteriorly. Ventral parameral tusks smaller and less strongly bent than
dorsal tusks; parameral apodemes large, connected ventrally by weakly sclerotized bridge (Fig. 9B).
Base of ejaculatory apodeme flattened; apical extension covered by membranous cap (Fig. 9B).
Genus Monepidosis Mamaev, 1966
A bibliography including the synonymy of the genus Monepidosis was given in the revision of
Porricondylinae by Jaschhof & Jaschhof (2013). Species of Monepidosis can be distinguished from each
other using characters of the male gonocoxites, or more specifically, the processes at the ventroposterior
gonocoxal margin, which vary in size, shape and position. There are two kinds of processes: the
gonocoxal processes, which are outgrowths of the medial bridges, and the central processes, which are
outgrowths of the ventral bridge (Jaschhof & Jaschhof 2013; this paper, Fig. 10A). Apart from genitalic
structures, the morphology of male Monepidosis is extremely uniform. Females and larvae of most
Monepidosis species, including the species treated here, are unknown.
Monepidosis heterocera sp. nov.
urn:lsid:zoobank.org:act:A0B02B2E-6CBl-488A-8C19-DDB27BC9D43B
Fig. 10A-C
Diagnosis
A typical Monepidosis (Jaschhof & Jaschhof 2013: 185), distinguished by the following genitalic
characters in combination (Fig. 10A-B). The gonocoxal processes, which arise from a concave, asetose
area of varying size (|, Fig. 10A-B), are nearly as large and prominent (j) as the central processes.
Their apices are typically rounded (Fig. 10A), rarely more pointed (Fig. 10B), and membranous rather
than sclerotized. The central processes, separated medially by a V-shaped space, are parallel-sided,
unicolored, and end in small, strongly sclerotized hooks (j. Fig. 10A). The bases of both gonocoxal and
central processes are occasionally interconnected by a sclerotized rim, which is very pronounced in the
specimen depicted in Fig. 10B (j).
Remarks on intraspecific variability
Most of the specimens studied have rounded gonocoxal processes that arise from small depressions
(Fig. 10C), so show a combination of the character states figured in A and B. The specimens depicted
in A and B were initially suspected to belong to different species, until it became obvious that they
represent the extreme ends of the range of variation described above. I believe that most of this variation
is artificial, i.e. dependent on how a specimen is positioned in the balsam drop.
Differential diagnosis
Monepidosis heterocera sp. nov. and M. pectinatoides Jaschhof, 2013 resemble each other in having
moderately sized, prominent gonocoxal processes and wide, parallel-sided central processes. The two
17
European Journal of Taxonomy 192 : 1-24 ( 2016 )
species differ in minute details concerning those processes. As characteristics of M. pectinatoides , the
gonocoxal processes are sharply pointed and sclerotized apically, and the central processes are bicolored,
black laterally, white medially (Jaschhof & Jaschhof 2013: fig. 89B). Further, the apical portion of the
ejaculatory apodeme is more strongly bent ventrally in M. pectinatoides than in any other Monepidosis,
including M. heterocera sp. nov.
Etymology
The name heterocera is derived from the Greek words heteros, for ‘different’, and her as, for ‘horn’,
meaning ‘with different horns’, which refers to the gonocoxal processes that distinguish this species
from its closest relatives.
Fig. 10. Monepidosis heterocera sp. nov., <f. A. Genitalia, ventral, holotype. B. Genitalia, ventral
(paratype). C. Processes at ventroposterior gonocoxal margin, ventral, paratype. Scale lines: 0.05 mm.
18
JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Type material
Holotype
SWEDEN: S, Oland, Morbylanga, Skogsby, Station Linne, 56.61° N, 16.49° E, near compost pile, 1-31
Jul. 2014, Malaise trap, M. & C. Jaschhof (NHRS CEC96).
Paratypes
SWEDEN: 1 S, Oland, Gamla Skogsby (Kallcstad), 56.61° N, 16.50° E, mixed deciduous forest, 1-27
May 2014, Malaise trap, M. & C. Jaschhof (NHRS CEC97); 3 $$, same locality, 13 May-8 Jun. 2015,
Malaise trap, M. & C. Jaschhof (NHRS CEC109-110, SDEI CEC111).
Other material examined
SWEDEN: 2 $$, Oland, Gamla Skogsby (Kallcstad), 56.61° N, 16.50° E, mixed deciduous forest, 1-27
May 2014, Malaise trap, M. & C. Jaschhof (NHRS ZFMK-TIS2537355-2537356).
GERMANY: 1 S, Brandenburg, Schorfheide-Chorin Biosphere Reserve, Barnim, Klein Ziethen,
Kernberge near Serwester See, edge of pine forest, 22 Apr. 2009, Malaise trap, leg. SDEI (SDEI A7625);
1 S, Baden-Wtirttemberg, Malsch, Glaser gravel plant, edge of pine forest, 19 Mar-21 Apr. 2010,
Malaise trap, D. Doczlcal (SDEI A7624).
Other characters
Body length. 1.7-1.9 mm.
Head. Eye bridge 3-4 ommatidia long dorsally. Circumfila on flagellomeres 1-10. Neck of fourth
flagellomere 1.5 times longer than node. Palpus 1.3-1.5 times longer than height of head.
Wing. Length/width 2.8-3.0. A remnant M present at wing margin.
Legs. Empodia rudimentary.
Terminalia (Fig. 10A-C). Gonostylus flattened, with plate-lilce, irregularly serrated spine around apex
(Fig. 10A-B). Parameres present as 2 pairs of large, strongly sclerotized, dorsally curved tusks, similar
to those found in many other Monepidosis (Jaschhof & Jaschhof 2013: fig. 88A). Apex of ejaculatory
apodeme flattened, bent ventrally; base widened, sclerotized (Fig. 10A-B).
Distribution and phenology
Sweden (Oland), Germany (Brandenburg, Baden-Wtirttemberg). Adults collected in and near deciduous
and coniferous forest from March/April (southwest Germany) to July (southeast Sweden). Possibly a
thermophilic species, given that all collection sites are favoured by a warm climate.
Monepidosis scepteroides sp. nov.
urn:lsid:zoobank.org:act:60EE7BD9-7842-4026-BA78-050F926D770B
Figs ID, 11A-C
Diagnosis
Monepidosis scepteroides sp. nov. is the only Monepidosis with only one pair of parameral tusks (j,
Fig. 11C). Of the gonocoxites (Fig. 11A), the wide, shallow ventral emargination has a glabrous rim
basally (|); the central processes, which are situated posterior to that rim, form a widely open, unevenly
sclerotized V-shape (j); and gonocoxal processes are missing.
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European Journal of Taxonomy 192: 1-24 (2016)
Differential diagnosis
Monepidosis scepteroides sp. nov. is similar to M. scepteri Spungis, 2006, especially with respect to the
central processes. In M. scepteroides sp. nov., as distinct from M. scepteri , the gonocoxites are wider
than long instead of the reverse, and the plate-like gonostylar spine is a single piece rather than consisting
of two clearly separated portions (Spungis 2006: fig. 4B). Like all Monepidosis but M. scepteroides
sp. nov., M. scepteri has two pairs of parameral tusks (Spungis 2006: fig. 4A).
Etymology
The name scepteroides , an adjective, refers to the similarity of this species to M. scepteri.
Type material
Holotype
SWEDEN: Morbylanga, Gamla Skogsby (Kalkstad), 56.61° N, 16.50° E, mixed deciduous forest, 8
Aug. 2014, aspirator, M. Jaschhof (NHRS CEC94).
Fig. 11. Monepidosis scepteroides sp. nov., holotype. A. Genitalia, ventral. B. Gonostylus, ventral
(paratype). C. Parameres and ejaculatory apodeme, ventral. Scale lines: 0.05 mm.
20
JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Other characters
Body length. 1.7 mm
Head. Eye bridge 3M ommatidia long dorsally. Apices of antennae missing, circumfila on flagellomeres
1-10 at least. Neck of fourth flagellomere 1.9 times longer than node (Fig. ID). Palpus 1.3 times longer
than height of head.
Wing. Length/width 3.1.
Legs. Empodia rudimentary.
Terminalia (Fig. 11A-C). Gonostylus (Fig. 11B) strongly flattened, with long, deeply incised plate-like
spine apically and apicoventrally. Apex of ejaculatory apodeme pointed, flattened, bent ventrally; base
weakly sclerotized, widened (Fig. 11C). Parameral tusks strongly curved, directed dorsally (j, Fig. 11C).
Distribution and phenology
Sweden (Oland). Known from a single specimen collected in deciduous forest in August.
Monepidosis shikokuensis sp. nov.
um:lsid:zoobank.org:act:7F6FEC96-9091-44F0-849C-3342D230A34F
Figs IB, 12A-B
Diagnosis
Parameres and ejaculatory apodeme of this highly unusual Monepidosis are merged into a single complex
structure (Fig. 12A). The portions of parameral origin, which encircle the ejaculatory apodeme, include
vestigial tusks that are curved dorsally (j). The ejaculatory apodeme is forked into 4 long prongs (j),
of which the laterals have finely serrate apices. The structure of gonostyli and gonocoxites is also
exceptional in M. shikokuensis sp. nov. (see section ‘Other characters’, Fig. 12B).
Etymology
The name shikokuensis , an adjective, is derived from Shikoku, one of the main islands of Japan, where
this species was found.
Type material
Holotype
JAPAN: S, Shikoku, Kochi, Asakura, 33.55° N, 133.47° E, secondary growth of mixed evergreen
deciduous / coniferous forest, 4-11 Nov. 1998, Malaise trap, M. & C. Jaschhof (KUEC).
Other characters
Body length. 1.6 mm.
Head. Eye bridge 5-6 ommatidia long dorsally. Circumfila on flagellomeres 1-12. Neck of fourth
flagellomere 2.1 times longer than node. Palpus 1.3 times longer than height of head.
Wing (Fig. IB). Length/width 3.2.
Legs. Empodia rudimentary.
Terminalia (Fig. 12A-B). Gonostylus flattened, slightly curved, tapered towards apex, without apical
spine (|, Fig. 12B). Gonocoxites (Fig. 12B): processes missing; ventral emargination unusually large,
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European Journal of Taxonomy 192: 1-24 (2016)
with broad, medially raised microtrichose rim basally (|); apodemes approach each other. Base of
ejaculatory apodeme sclerotized, widened (Fig. 12A).
Distribution and phenology
Japan (Shikoku). Known from a single specimen collected in evergreen secondary growth in November.
Monepidosis spatulata Spungis, 2006
Monepidosis spatulata Spungis, 2006: 27.
Remark
Monepidosis spatulata , a species described on the basis of specimens found in Latvia and Lithuania
(Spungis 2006), is here reported for the first time from Gotland, Sweden.
ejap
gcx ap
Fig. 12. Monepidosis shikokuensis sp. nov., c?, holotype. A. Parameres and ejaculatory apodeme, ventral.
B. Genitalia, ventral. Scale lines: 0.05 mm.
22
JASCHHOF M., Taxonomy of Monepidosis and Antipodosis gen. nov.
Material examined
SWEDEN: 1 S (no. CEC98), Gotland, Eksta socken, Stora Karlso, 8-29 Aug. 2014, Malaise trap,
Hymenoptera Inventory Group 2014 (NHRS).
Discussion
The example of Monepidosis is ideally suited to demonstrate how little we know of the biodiversity
of Porricondylinae in Europe, the continent regarded quite rightly as the best-investigated for this and
the other subfamilies of fimgivorous Cecidomyiidae. Monepidosis is also exemplary for the fact that
many Porricondylinae must be regarded as rare in the sense that specimens are seldom found despite
great efforts to collect them. This scarcity, which cannot be explained due to the lack of autecological
information, makes species inventories fruitful but time-consuming ventures. For example, a nationwide
Porricondylinae inventory running in Sweden since 2009 (see Jaschhof & Jaschhof 2013) found nine
different species of Monepidosis to date, of which five were new to science (a tenth species, also
new, is known to me from two specimens that are too poorly preserved for description). Some of the
morphologically most unusual Monepidosis , such as M. shikokuensis sp. nov., came to be known not from
Europe but from Japan, where Porricondylinae have never been surveyed on a systematic basis, meaning
that random findings contribute significantly to the overall picture we have of this genus. Antipodosis
gen. nov. must be regarded as a similarly random discovery. The 13 specimens I had available to describe
Antipodosis gen. nov. and its eight species were accumulated at the time of the ‘Lestremiinae of New
Zealand’ project for curiosity’s sake rather than for taxonomy. Another 60 specimens from the same ‘by-
catch’ of New Zealand Porricondylinae belong to 41 different species of 9 genera (Jaschhof, unpublished
data), whereas only one porricondyline, Yukawaepidosis aliculata (Yukawa, 1964), was named from
the New Zealand region in the past (Gagne 1989; Gagne & Jaschhof 2014). Finally, the discovery that
Antipodosis gen. nov. and Monepidosis are closely related to each other does not necessarily indicate
straight affinities between New Zealand and Holarctic Porricondylinae. It remains unknown for the
time being whether the Monepidosis group has representatives in southeast Asia, New Guinea, New
Caledonia, or Australia, since all these regions are practically unstudied for Porricondylinae.
Acknowledgements
Research resulting in this paper was funded by Svenslca ArtDatabanken (Swedish Species Information
Centre) within the framework of Svenska Artprojektet (Swedish Taxonomy Initiative) (dha 2014-
150 4.3). My wife, Catrin Jaschhof, helped considerably with fieldwork and specimen preparation.
Many of the specimens studied here were provided by the fellow entomologists specified in the Material
examined sections. Ilea Osterblad picked out the Cecidomyiidae specimens from insect samples collected
on Stora Karlso. The manuscript was kindly read by Raymond J. Gagne before submission to the journal
and by three reviewers unknown to the author after submission; sincere thanks are given to all of them.
References
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Manuscript received: 2 December 2015
Manuscript accepted: 4 January 2016
Published on: 26 April 2016
Topic editor: Koen Martens
Desk editor: Kristiaan Hoedemakers
Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United
Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of
Denmark, Copenhagen, De nm ark
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