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HARVARD UNIVERSITY

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1976

DEEL 119 MUS. COMP, ZOOL:

i rn LIBRARY

id MAR 2/3 1977

HARVARD

SI, DSCHRIFI

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

Tijdschrift voor Entomologie, deel 119, 1976

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

BESTUUR (BOARD)

Voorzitteri(Chaisman) an se eee J.T. Wiebes

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Afleveringen 1-3 verschenen 7.VII.1976

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ERRATUM a

P. 219: 2nd line of summary, “with (hippophorbiae x euphorbiae) 9.” 9°

should read “with (hippophorbiae x hippophaes) ©.

INHOUD VAN DEEL 119

Achterberg, C. van. — A preliminary key to the subfamilies of the Braconidae (Hymenoptera)

Atyeo, W. T., & P. C. Peterson. — The feather mite genera Zumptia Gaud & Mouchet and Para-

zumptia gen. nov. (Acarina, Analgoidea) ........................

Blokhuis, B., zie Heerdt, P. F. van.

Ellis, W. N. — Autumn fauna of Collembola from Central Crete ..................

Haaften, C. van, zie Heerdt, P. F. van.

Heerdt, P. F. van, B. Blokhuis & C. van Haaften. — The reproductive cycle and age composi-

tion of a population of Pterostichus oblongopunctatus (Fabricius) in the Netherlands

(@oleopteraa@arabidae) ern. A one

Jong, R. de. — Affinities between the West Palaearctic and Ethiopian butterfly faunas .....

Kruseman, G., zie Willemse, F.

Lith, J. P. van. — New species and records of Indo-Australian Psenini (Hymenoptera, Spheci-

dacsPemphredoninde) arene nl) Tee coc) I ee a eee

Loeliger, E. A. — The enigma of Celerio hybr. Pauli Mory ......................

Meeuse, A. D. J., zie Stelleman, P.

Peterson, P. C., zie Atyeo, W.T.

Stelleman, P., & A. D. J. Meeuse. — Anthecological relations between reputedly anemophilous

flowers and syrphid flies. I. The possible role of syrphid flies as pollinators of Plan-

COLORE td SOTA AO ANTON

Willemse, F., & G. Kruseman. — Orthopteroidea of Crete ......................

DEE ooo ROIO ORE CORRI We

217

123

337

DEEL 119 AFLEVERING 1 1976

PP 7569.2,

TIJDSCHRIFT

VOOR ENTOMOLOGIE

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UITGEGEVEN DOOR LIBRA re ve

\AUG 1

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGINGY AF > È

UNIVERSIT

INHOUD

_P. F. van HEERDT, B. BLOKHUIS and CAROLINE VAN HAAFTEN. — The reproductive

cycle and age composition of a population of Prerostichus oblongopunctatus

(Fabricius) in the Netherlands (Coleoptera: Carabidae), p. 1—13, fig. 1—4,

pl. 1—2.

| Tijdschrift voor Entomologie, deel 119, afl. 1 Gepubliceerd 7-VII-1976

THE REPRODUCTIVE CYCLE AND AGE COMPOSITION OF

A POPULATION OF PTEROSTICHUS OBLONGOPUNCTATUS

(FABRICIUS) IN THE NETHERLANDS (COLEOPTERA:

CARABIDAE)

P. F. VAN HEERDT, B. BLOKHUIS and

CAROLINE VAN HAAFTEN

Zoologisch Laboratorium, Rijksuniversiteit, Utrecht

With two plates and four text-figures

ABSTRACT

The reproduction of a Dutch population of Prerostichus oblongopunctatus (F.) is analysed. Seasonal

variations in egg numbers, “corpora lutea” sizes, the relative positions of eggs in ovarioles and

oviducts, and the conditions of the eggs were studied. Three age classes (corresponding with one-, two-,

and three-year-old females) could be distinguished by measuring the sizes of the “corpora lutea”. The

duration of the reproduction and its course could be established. The survival value of a population

consisting of more than one age class is discussed.

INTRODUCTION

Previous observations by Den Boer (1968) and Van Dijk (1973) provided

information on the heterogeneity of the reproductive cycle which, in some species

of carabid beetles, may be important to survival under adverse environmental

conditions.

This heterogeneity, which presumably is partly genetically fixed, may contribute

to spreading the risk of extinction of the population. Den Boer and Van Dijk

observed that not only one-year-old females of Calathus species contributed to

reproduction, but also two- and even three-year-old ones. These authors observed

that in young females at the start of the development of the ovaries no “‘corpora

lutea” (abbreviated c. 1.) were present and that only after several weeks the first

c.l. could be found. Their results suggest that it would be possible to discriminate

between females of the old and the new generation in a field population by means

of the cl. in the ovaria.

The present work was started to investigate the validity of these facts also for

Pterostichus oblongopunctatus, a carabid beetle common in woods in the Nether-

lands. The reproductive pattern has been studied by observation of the develop-

ment of the ovaries and the maturation of the eggs. These features have been

studied during a number of successive years in order to unravel the complex

dynamics of the population concerned.

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

The area where the population of P. oblongopunctatus has been studied, occupies

2.4 hectares and was planted with oak (Quercus robur) about 90 years ago. The

oaks are sparsely intermingled with birch (Betula spec.) and have formed a dense

canopy over a well-developed shrub-stratum. Herbaceous growth is practically

non-existent, but a thick layer of partly decomposed leaf-litter (5-10 cm) makes a

good hiding place for a rich epedaphic fauna.

MATERIAL AND METHODS

Pterostichus oblongopunctatus (F.), a small bronze-black coloured beetle (9-12

mm) lives as an adult in the superficial layer of leaf-litter, preferably in broadleave«

woods. The larvae usually develop in the deeper layers. Reproduction occurs in

spring, the eggs are deposited in the litter.

Pitfall traps (Den Boer, 1968) were used for sampling: a plastic funnel (16 cm) .

fitted to a glass jar filled partially with 4% formaldehyde and dug into the soil, the

rim of the funnel being flush with the soil surface. A cover (Q 25 cm) is placed 5

cm above the funnel to prevent rain from penetrating into the trap and diluting the

formaldehyde. Three units of five traps each were dug in. Once a week the jars

were changed for fresh ones, and the specimens of P. oblongopunctatus selected

from the contents of these jars.

Fresh ovaries proved to be preferable for this study to those preserved in

formaldehyde, but the former were available only during 1972. The females were

dissected carefully and the ovaries (Fig. 1) were taken out. The development of

the ovary and the absence or presence of the corpora lutea are a criterion to the

age of the female.

terminal filament

germarium

vitellarium

clod of trophocytes

follicle epithelium plug

follicle epithelium

“corpus luteum”,

calyx

Fig. 1. Ovariole of Pterostichus oblongopunctatus, with indication of elements

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Prerostichus oblongopunctatus 3

RESULTS

In the Netherlands, this species has one generation annually. The newly hatched

adults hibernate as virgins, and copulate in spring and summer. The new

generation appears in autumn. Thus the activity pattern of the population shows

two distinct periods in spring and in autumn (Fig. 2). The spring activity starts in

100

Numbers captured

60 Pterostichus oblongopunctatus

O E

Fig. 2. The activity of Pterostichus oblongopunctatus in 1971

approximately the first week of April and is related to reproduction. The number

of beetles caught gradually diminishes towards the end of the reproduction period,

i.e. the beginning of August. The appearance in September of the new generation

probably stands for the major part of the higher catches in autumn. “Spent”

females (i.e. those that have oviposited) represent one-third of the specimens

trapped, as their activity is low at this time. Until the end of May more males than

females are caught (1971: 260 against 189). Presumably the former are more active

in this period, in search for a partner. From about the end of May until the first

weeks of August the females make up the majority (154 against 186).

The following categories of 9 in the different stages of development were

recognized (PI. 2):

(a) Without eggs and without c.l.

(b) With eggs, without c.l.

(d) With eggs under and above the c.l.

(e) With eggs above the c.l.

(f) Without eggs, with c.l. (“spent”).

In the females without c.l. in April, the eggs are still above the calyx, in the

ovarioles: a few © (PI. 2, punctated yellow) have eggs in the calyx, but have c.l. not

yet visible. Many 9 with c.l. are bearing their eggs at this time already under the

c.l. in the calyx. The females with c.l. but without eggs may have finished

oviposition at this moment (“spent”). The data suggest that at the start of the

reproduction season females with c.l. (Pl. 2, gray) may start earlier to reproduce

than females without c.l. (i.e. those belonging to the new generation).

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

According to Vlijm & Van Dijk (1967) it is possible to determine the age of the

females by the development of the ovary and the absence or presence of corpora

lutea, as follows:

First year 9:

(1) an ovary in a state of early development

(2) asmall, pointed ovary

(3) acalyx, not yet elongated

before reproduction

(4) without corpora lutea

(5) thin, yellowish-brown c.l. È

(6) thin, but distinct, dark coloured c.l. only at the end during reproduction

of the reproduction period: ““c.l. 2”.

2nd- or 3rd year 9:

(1) thin, dark coloured c.l. before the reproduction period (“c.l. 2°’)

(2) distinct, dark coloured c.l. at the beginning of the reproduction period (“*c.l.

2”)

(3) big c.l. during the reproduction period (‘‘c.l. 3”)

(4) fat, “fluffy” ovaries (the eggs and ovaria are crumbling and show a fluffy

structure).

Since the size of the c.l. has been estimated, it is hard to give a good

classification. Especially in specimens with distinct c.l. in the middle of the

reproduction period one may be uncertain of their age. In a later stage of

research, however, one of us (Blokhuis) used an ocular-micrometer to measure the

size of the c.l. Thus more accurate data could be obtained on the difference in size

of the c.l. than by mere judgement of their dimension.

The data collected of every female captured in order to judge its stage in the

reproduction (Table 1) are as follows:

(1) the number of 9 without c.l. or eggs (I)

(2) the number of 9 without c.l. with eggs (II)

(3) the number of 9 with c.l. and with eggs (III, IV, IVa)

(4) the number of © with c.l. without eggs (“‘spent’’) (V)

(5) size and colour of the c.l.

(6) the number of eggs in relation to their position in the ovary

(7) the condition of eggs and ovaries.

Table 2 shows the relation between the number of eggs per female and the size

of the c.l. If the size of the c.l. is a measure for the age of the female, it should be

possible to find a relation between age and number of eggs. The number of

ovarioles varies between 10-26. The average weekly number is about 15.4-20.0.

Probably this number is too low, as counting is made difficult by the “fluffy”

ovaries. Three females had few c.l. (resp.3,3 and 2) but many ovarioles (14, 18 and

20). In 5 females only one ovary had developed; in 2 females a double c.l. ring

could be observed.

Development of the corpora lutea.

As Miss Van Haaften (in 1970) and Blokhuis (1971) obtained almost identical

results, these can be summarized as follows:

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Pterostichus oblongopunctatus 5

Table 1. 1971: week 1-4: April; 5—8: May; 9—13: June; 14—17: July; 18—25: Aug.; 22—26: Sept.;

27—30: Oct.; 31—35: Nov.; 36—39: Dec. I, number of 9 without c.l., without eggs; II, number of 9

without c.l., with eggs; III, number of 9 with small c.l. (c.l. 1) with eggs; IV, number of 9 with distinct

c.l. (c.l. 2) with eggs; IVa, number of 9 with large c.l. (c.l. 3) with eggs; V, number of 9 with c.l. 1, c.l.

2, c.l. 3 without eggs (“spent”). Between brackets: percentages I-II (without c.l.); III (cl. 1); IV (c.l. 2);

IVa-V (c.l. 3) of the total of animals caught

Week I II III IV IVa V IVa+V

1971 nr. % nr. % nr. % boe. ele %

| 5 2 (47) = 2 (13) l — — 5 (40)

2 6 2 (67) — 2 (16.5) 2 — — — (16.5)

3 4 9 (48) — 7 (26) 5 = = 2 (26)

4 2 10 (33) 4 (11) ll (31) 6 — — 3 (25)

5 — 1 (12.5) 38725) 27:25) 2 = — — (25)

6 3 8 (27) 15 (37) 8 (22) 5 — | | (14)

7 — 5 (31) 6 (37) 3 (19) 2 = — = (13)

8 l 4 (30) 6 (35) 4 (24) 2 — — — (11)

9 — 9 (31) 11 (41) 4 (14) 4 | — — (14)

10 = 7 (20) 15 (46) 6 (17) 5 | = | (17)

11 l 3 (16) 12 (54) 2 (14) 4 | 1 — (16)

12 — — 14 (74) 2 (16) 2 — 1 | (10)

13 — 0 — 6 (55) 1 (18) 2 — | | (27)

14 3 — (11) 12 (43) 4 (21) 3 — 2 4 (25)

15 IS (65) 9 (53) 3 (18) = = — 4 (23)

16 1 — (50) — 1 (50) = — = —

17 — — 1 (11) 5 (56) — — — 3 (34)

18 1 — (14) — 3 (43) | — — 2 (43)

19 = = IS) = — — — 3 (75)

20 = = — — | — = = (50)

21 1 — (50) — — — — | = (50)

22 2 — (40) — — = = — 3 (60)

23 1 — (50) — = = = — | (50)

24 Be (75) — — — — | — (25)

25 5 — (100) — — — — — —

26 13 — (76) — — — 2 — 2 (24)

27 8 — (73) _ — — — | 2 (27)

28 7 — (70) — 2 — | 2 — (30)

29 1 — (33) — — — = = 2 (77)

30 1 — (50) = == — | = — (50)

31 1 — (50) = = = = — (50)

32 — — — = = = — —

33 1 — (100) = = = - — —

34 = = = = = = = —

35 e — = = = — —

36 => = = — = = = =

37 — = = — = = =

38 =O — = = — == ==

39 1 — (100) — — — — — —

73 60 115 70 46 9 Il 39

yvyT= sr

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

Table 2. 1971. The relation between the number of eggs per 9 and the size of the c.l.; II, no c.l.; III,

small c.l. (c.l. 1); IV, distinct cl. (cl. 2); IVa, large c.l. (c.l. 3)

Number of eggs per 9 II III IV IVa

| | 2 8 2

2 l — 17 10

3 | | 13 10

4 3 6 7 4.

5 3 2 6 3

6 6 11 5 3

7 5 10 2 |

8 8 15 | 5

9 9 16 3 2

10 3 9 2 |

Il 6 5 | —

12 6 4 | |

13 6 7 | 2

14 | 9 — |

15 — 4 | |

16 — 7 | —

17 | 2 — ==

18 — 2 | —

19 — — — =

20 — 2 — —

21 — = = —

22 — | — —

Number of 9 60 115 70 46

Number of eggs 526 1153 326 243

Mean number of eggs 8.8 10.0 4.7 973)

In the second week of April females without c.l. still constitute 44-67% of the

total catch. Their numbers decrease gradually to zero — after July 14 the results

are unreliable. Females with c.l. 1 are observed for the first time in the last week of

April. Their share begins with 6—11% and rises to 74% in the third week of June.

Later the numbers decline again, but apparently more gradually than is suggested

by Table I, for it has to be corrected by those females which have grown from c.l.

I to c.l. 2. The resulting decline is probably caused by a decrease of activity. The

number of females with c.l. 2 attain a maximum in April (30%). After April 28 their

number decreases, because either their cl. 2 may grow to c.l. 3 or by mortality of

the supposed 2nd year individuals. The number of females with cl. 3: this group is

at the onset of the activity represented to 40%, but as April wears on their

numbers decrease rapidly to 20%, in May even to 11%. After May 26 an increase

takes place again, apparently as a consequence of c.l. 2 9 developing into c.l. 3

individuals as they continue to produce eggs. The initial decrease is, obviously,

caused by the dying off of the supposed 3rd year generation (PI. 2). It must be

noted that the first females with eggs beneath the c.l. (April) are of the c.l. 3 group,

hence, the oldest females apparently take the lead in reproduction, but within a

few weeks they are already “spent” again and will probably die off soon,

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Pterostichus oblongopunctatus 7

Table 3. 1972. Size of the c.l. measured in u

From 30 March to 12 April From 31 May to 14 June

Size Number 9 Size Number 9

!) 19 — 7)

828u 2 23u |

85u 2 31u 3

108u 2 42u |

112p l 45u 2

123u l 46u 4

49u |

From 12 April to 26 April 77u 2

Size Number 9 88u I

3lu Zur

DE i From 14 June to 28 J

54u 3 rom 14 June to une

62u 2 Size Number 9

120u l — 2

29u l

From 26 April to 10 May 3lu |

Size Number 9 40n

69u 3

From 10 May to 31 May 72u l

Size Number 9 Tu 7

a Il 1081 |

31u 2 1234 2

381 1 13lu l

42 I

58, ati From 28 June to 12 July

62u 2 Size Number 9

69u l — 0

92u 3 46u |

1084 | 69u |

77u 4

1) — = without c.l. 1234 |

according to their disappearance in May (Pl. 2). A postponed reproduction of the

preceding year may be possible, so that 2nd year © may die when they are two

years old and only those 9 which failed to reproduce the previous year would

reach their 3rd year (Vlijm & Van Dijk, 1967, have supposed this to be the case in

Calathus melanocephalus).

Measurements of corpora lutea.

The estimate of the size of the c.l. being not quite reliable, those of freshly

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

1972

third

March 30 a

April 12

number of individuals

without c.l.

second year 9 9

April 13-26

first year

April 27 ot

May 10

May 11-31

June 1-14

June 15-28

number

(©)

June 29

July 12

N a O o

size in u

100 140

Fig. 3. Size of corpora lutea in 1972

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Pterostichus oblongopunctatus 9

caught 9 were measured by Blokhuis in 1972 by means of an ocular micrometer at

a magnification of 100x (Table 3). In the first weeks of April 1972 only females

with c.l. 3 (108—124u) or large c.l. 2 (77—85u) were present. One female with a c.l.

of 28u probably reproduced only a short time during the preceding year. In the

second half of April only one 9 with c.l. 3 was captured, but females with small

c.l. 2 made their first appearance (50—60u). No females with c.1. 3 could be caught

during the first part of May but the c.l. of the c.l. 2 females tend to be bigger

(SS—80u) and it is in this period that females with c.l. 1 have been collected for the

first time. During the second part of May, in June and the first two weeks of July

the c.l. 2 have reached the size which the cl. 3 showed in April, and the c.l. 1 those

of the c.l. 2. During 7 weeks (April 26-June 14) not a single female with c.l. 3 had

been caught. Hence, we may conclude that females with c.l. 3 at the onset of the

breeding season (first half of April) die after oviposition. The new 2nd year

females take their place at the end of June. These data are a confirmation of the

hypothesis that more generations are involved in reproduction (Fig. 3).

A check in 1973 on individuals of the 1971 generation which were marked as

young females in 1972, showed a size of the c.l. corresponding with those of

second year Q (cf. Fig. 4). Consequently the age of a female can be estimated

approximately by these figures though a certain overlap exists. It is evident that at

a certain moment of development, Ist year c.l. 2 and 2nd year c.l. 2 do not differ in

1973

third year 99

March

April

second year 99

April 16-19

first year 9 9

April 26

May 4

N a HD oo

May 14

size in u

number

0 50 100 140

Fig. 4. Size of the corpora lutea of individuals marked in 1972, recaptured in 1973

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

size. A more accurate way to determine the ages of the © taking part in

reproduction is the marking-recapture method which we intend to use in the .

future.

A serious obstacle in the interpretation of “corpora lutea” (sizes as well as

numbers) as indicators of age is the phenomenon of egg resorption (Joly, 1949, and

others). The precise origin of the cl. seems to be unknown.

Egg production.

When dissected, only 6 out of the 291 egg-bearing females appeared to have

white, glossy, elastic eggs, one female had brown, hard, malformed eggs. The eggs

of all the other females proved to be “fluffy” and soft and were coloured light

yellow to orange. It is possible that this coloration is due to the preservation agent,

i.c. formaldehyde. i

In the first week of May all females are bearing eggs. This situation is reached

within one month. Decrease to zero takes more time: it takes more than 3 months

before the last egg has been deposited (Table 4).

Table 4. 1971. The number of egg bearing 9. Week nr. 1: March 31-April 7. Week nr. 20: August 11-

August 18

Week Total 9 Q + eggs %

| 31 March — — —

2 7 April 15 5 33

3 14 April 12 6 50

4 21 April 27 21 78

5 28 April 36 31 86

6 5 May 8 8 100

7 12 May 43 36 84

8 19 May 16 16 100

9 26 May 17 16 94

10 2 June 29 28 97

Il 9 June 35 33 94

12 16 June 24 21 88

13 23 June 19 18 95

14 30 June 11 9 82

15 7 July 28 19 68

16 14 July 17 12 fil

17 21 July 2 | 50

18 28 July 9 6 67

19 4 Aug. 7 4 OL

20 11 Aug. 4 l 25

In 1971, the average number of eggs was 7.7. Computing the number for each

group of females shows that females without c.l. and those with small and thin c.l.

have an average of 9.6 eggs; c.l. 2 females have 4.6 and c.l. 3 females 5.8 (Table 5).

It can thus be concluded that the females without c.l. and with cl. I produce the

majority of the eggs. The groups with c.l. 2 and c.l. 3 contribute only one third of

the total.

By means of Wilcoxon’s Two Sample Test the zero-hypothesis of a similar egg-

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Pterostichus oblongopunctatus Il

Table 5. The average number of eggs in 1970 and 1971

1970 1971 Corrected average numbers (1971)

Generation 1967 |

Generation 1968

Generation 1969 4.3 4.6 (4.7 for c.l. 2-group © and c.l. 3-group 9

without in April)

I) 5.8 (6.3 for c.l. 3-group © in April)

Generation 1970 — 9.6

N.B. Generation 1968 corresponds with the c.l. 3 group in April 1971. Generation 1969 corresponds

with the c.l. 2 group and the c.l. 3 group together, without the individuals with c.l. 3 caught in

April. Between brackets the corrected averages of the number of eggs. After April the c.l. 3

group should be added to the c.l. 2 group because of the development of c.l. 2 to c.l. 3 in the

course of the summer. The average of 5.5 has been computed by adding the number of eggs of

the generations 1967 and 1968 which until that time could not yet be discerned. Only the c.l. 3 9

caught in April probably have an age of three years (or more).

production in two groups is tested against the alternative hypothesis of a different

egg-production (a= 0.05; u= 1.96). It can be concluded that group II and III do

not differ significantly. This could be expected as these groups belong to the same

generation: 1970-groups II and IV differ significantly (u= 6.4; p<0.0001) as do

groups II and IVa (u= 4.76), but groups IV and IVa do not differ significantly (u=

1.22).

Miss van Haaften, in 1970, arrived at the same results with the Median Test.

CONCLUSIONS AND DISCUSSION

Composition of the population.

The results show that the population consists of 3 generations which are all

nearly simultaneously active (Table 6, which has been derived from Tables | and

5). In June and July only 2 generations are present, as the 3rd generation dies off

after oviposition in April-May and the new generation hatches only in September.

Survival value.

As the larval stage is supposed to be rather vulnerable, it is evident that, if one

generation in its larval stage is severely damaged by adverse environmental

(humidity or temperature: too high or too low) and/or biotic (disease, parasites,

predators) influences, the next year still two additional generations survive to

assure the persistence of the population: the last 4 years about 40% of the females

taking part in reproduction consisted of 2 or 3 year old (Table 7).

At the beginning of 1971 (March 31-May 5), the rate of 2nd and 3rd year egg-

bearing females stood at 44%, the total number captured, i.e. 2nd and 3rd year 9

without eggs included, even at 51%. As reproduction goes on these numbers

decline respectively to 25% 9 with eggs and to 24% of all females captured in the

period May 6-June 2 and to 30% © with eggs and 36% © of all © captured in the

period June 3-July 7.

The ratio of this last period is too high owing to a failure in the correct

estimation of the size of the c.l. which in the future can be eliminated by the

micrometer method (cf. p. 9).

12 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

Table 6. Reproductive activity in 1971

Date Age Reproductive activity Generation

April Ist year without c.l., without eggs 1970

Ist year without c.l., with eggs 1970

Ist year with c.l. 1, with eggs 1970

2nd year with c.l. 2, with eggs 1969

3rd year with c.l. 3, with eggs 1968

3rd year with c.l. 3, “spent” 1968

May as April but 3rd year 9 disappear

June Ist year with c.l. 1, with eggs 1970

Ist year spent with c.l. l and c.l. 2 1970

2nd year with c.l. 3 and eggs 1969

2nd year spent with c.l. 3 1969

July Ist year with c.l. 1, c.l. 2 and eggs 1970

Ist year spent with c.l. 2 1970

2nd year spent with c.l. 3 1969

August Ist year with c.l. 2 and eggs 1970

l st year spent with c.l. 2 1970

2nd year spent with c.l. 3 1969

September Hatching of the new generation 1971

Ist year spent with c.l. land cl. 2 1970

2nd year spent with cl. 3 1969

October New generation 1971

Ist year spent with c.l. I 1970

2nd year spent with c.l. 3 1969

The first year generation (1970) produced a large number of eggs (average 9.6).

The second year generation (1969) had an average production of 4.7, the third year

generation (1968) one of 6.3. According to Miss Van Haaften, the generations of

1969 and 1968 produced about the same average (4.3 and 5.5, respectively). Data

of 1972 have already confirmed that the population increased with about 10%: the

generation of 1970 may have had a big part in this increase.

Survival, however, is not dependent of meteorological factors only: predation,

disease and the available amount of food will be important too, but reliable data

are, thus far, lacking.

Comparison with other carabid beetles.

It is a special advantage that Van Dijk (l.c.) studied the age composition of

Calathus melanocephalus (L.) which is, contrary to P. oblongopunctatus, a ‘‘winter

Table 7. Composition of the population in 1969, 1970 and 1971

Year Locality % more than Method

one year old

1969 Baarn 49% by estimating the size of the c.l. (by De Bruyn)

1970 Baarn 32% by estimating the size of the c.l. (by Van Haaften)

1971 Baarn 39% by estimating the size of the c.l..(by Blokhuis)

1969 Wijster 37% capture-recapture

(Van Dijk, personal communication)

VAN HEERDT, BLOKHUIS & VAN HAAFTEN: Pterostichus oblongopunctatus 13

breeder” (i.e. the larvae develop during winter time). In the former species young

adults hatch in spring and only mature in the course of July, while in P.

oblongopunctatus the adults hatch in September, mature in the course of the

autumn and start oviposition towards the end of April. In P. oblongopunctatus, egg-

production by Ist year females as well as by 2nd and 3rd year females starts earlier

in the season but finishes about July 21, whereas in C. melanocephalus it continues

well into September. As a matter of fact, P. oblongopunctatus has a lead over C.

melanocephalus as at a certain moment (end of April, beginning of May) three

generations may reproduce simultaneously, contrary to C. melanocephalus where at

best two generations do so.

ACKNOWLEDGEMENTS

We are grateful to Dr. P. J. den Boer and Dr. T. S. van Dijk (Dr. W. Beyerinck

Biologisch Station, Wijster) for their stimulating interest and useful advice in our

investigations. We thank Dr. W. J. Sluiter for reading the manuscript and Miss M.

A. de Bruyn for putting some preliminary results at our disposal.

REFERENCES

Boer, P. J. den, 1967. Zodlogisch onderzoek op het Biologisch Station te Wijster. — Med. Bot. Tuin

Belmonte Arb. 11.

———, 1968. Spreading of risk and stabilization of animal numbers. — Acta Biotheoretica 18:

165—194.

Dijk, T. S. van, 1973. The age-composition of populations of Calathus melanocephalus L. — Oecologia

12: 213—240.

Joly, P. 1949. La fonction ovarienne et son controle humoral chez les Dytiscides. — Arch. Zoöl. exp.

84: 49— 164.

Vlijm, L. et al., 1961. Ecological studies on Carabid beetles I: Calathus melanocephalus. — Arch. Neerl.

Zool. 14: 410— 422.

Vlijm, L., & T. S. van Dijk, 1967. Ecological studies on Carabid beetles II: General pattern of popula-

tion structure in Calathus melanocephalus (L.) at Schiermonnikoog. — Z. Morph. Oekol. Tiere

58: 396— 404.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 1, 1976

Pil

a b

Ovaria and oviducts of Pterostichus oblongopunctatus. a, corpora lutea; b, egg in pedicel

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TIJDSCHRIFT

VOOR ENTOMOLOGIE

“LIBRARY

(AUG 4 7 *

DE NEDERLANDSE ENTOMOLOGISCHE VER NIGINGS Y

UITGEGEVEN DOOR

INHOUD

TELLEMAN and A. D. J. MEEUSE. — Anthecological relations between repu-

__ tedly anemophilous flowers and syrphid flies. I. The possible role of

| syrphid flies as pollinators of Plantago, p. 15—31, fig. 1—2, pl. 1—3.

chrift voor Entomologie, deel 119, afl. 2 Gepubliceerd 7-VII-1976

ANTHECOLOGICAL RELATIONS BETWEEN REPUTEDLY

ANEMOPHILOUS FLOWERS AND SYRPHID FLIES

I. THE POSSIBLE ROLE OF SYRPHID FLIES AS

POLLINATORS OF PLANTAGO

P. STELLEMAN and A. D. J. MEEUSE

Hugo de Vries Laboratorium, University of Amsterdam

With two figures and three plates

ABSTRACT

Regular visits by syrphid flies of the genera Melanostoma and Platycheirus to the inflorescences of

the reputedly anemophilous Plantago lanceolata L. for the purpose of pollen consumption have often

been reported. An effective pollen transfer could be established in the present study. Observations

made in the field indicated that the flies feed on Plantago laceolata pollen in the early morning. Several

aspects of their behaviour already suggest them to be effective pollen vectors. By means of Scanning

Electron Microscope observations the presence of pollen grains of P. lanceolata on the body of the flies

could be demonstrated. Experiments in which artificially dyed pollen of P. lanceolata was deposited on

inflorescences in anthesis, while the stained pollen was subsequently recovered from the stigmas of

other spikes, demonstrate that they are efficient pollinators of this plant. The amount of pollen

transferred was estimated and, at least in certain habitats, is thought to be responsible for a

considerable part of the geitonogamous and heterogamous pollinations. The technique employed may

prove to be useful for the study of comparable cases in plants and animals.

I. OBSERVATIONS IN THE FIELD AND PRELIMINARY STUDIES

The taxon Syrphidae (Diptera) almost exclusively consists of species which in

the adult stage are typically anthophilous and feed chiefly on pollen or nectar.

Differences in body size, and in the length of the proboscis, are considerable, so

that a rather wide range of flowers can be visited: Among the Diptera, the

Syrphidae are in general the most important pollinators, although their signifi-

cance is not as great as that of the social hymenopterous group of the Apidae

(Kugler, 1970; Faegri & Van der Pijl, 1971).

As may be expected, syrphid flies normally visit flowers or inflorescences of

typically entomophilous plants, but observations in the field have shown that some

reputedly anemophilous plants are visited regularly by various species of

Syrphidae for the purpose of pollen consumption. However, the published records

provide relatively few reliable data. The most important ones, to be mentioned

presently, refer mostly to visits by representatives of the closely related genera

Melanostoma and Platycheirus of the subfamily Syrphinae.

One of the earliest reports is by Miller (1873), who noted that in the

surroundings of Lippstadt (W. Germany) insects often gather or consume pollen

on the spikes of Plantago lanceolata. Beside Hymenoptera, he reported several

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

Syrphidae among which Melanostoma mellinum (Linnaeus) is specially mentioned

as a frequent pollen-eating visitor. Muller pointed out that this fly even shows a

preference for wind-pollinated flowers: he found this insects as a regular pollen

consumer on Plantago media L., Artemisia dracunculus L., Scirpus lacustris L., and

several grasses such as Anthoxanthum odoratum L., Poa annua L., and Festuca

pratensis Huds. It was also found on a whole range of entomophilous plants where,

apart from pollen, it sometimes also ingested nectar. Müller believed that visits to

anemophilous plants by this type of insects might result in cross-pollination.

Representatives of the closely related Platycheirus were not reported as visitors of

wind-pollinated taxa.

Ludwig (1881, 1884) noticed a very large number (several thousands) of syrphids

of the genera Melanostoma, Platycheirus and Melithreptus (Sphaerophoria) on the

panicles of Molinia coerulea (L.) Moench in Saxony (E. Germany). An appreciable

number were caught by the proboscis between the paleae or the anthers and in

that case they usually appeared to be infected by a parasitic fungus (Entomophtora

spec.), while living flies were often found to be present also. Ludwig suggested that

the purplish-mauve colour of the anthers acts as a strong attractant for the visitors.

He observed the same flies on Phleum pratense L., Helictotrichon pubescens (Huds.)

Pilger, Dactylis glomerata L., and Plantago lanceolata.

The extensive monograph of Knuth (1898—1905) records a number of cases of

Melanostoma mellinum visiting various anemophiles. Apart from citing earlier

authors he also reported personal observations made near Kiel and in the North

Frisian islands (W. Germany), where he frequently noted several specimens of this

fly on inflorescences of Alopecurus pratensis L., Phleum pratense, and Anthoxanthum

odoratum. He also mentioned visits of this species to Plantago arenaria W. & K. (in

a botanical garden) and to Sanguisorba minor Scop. Knuth did not attribute any

significant role to these visits in effective pollen transfer. In addition he mentioned

an appreciable number of visits of Melanostoma mellinum (and other species of the

genus) and of Platycheirus species to entomophilous flowers.

Drabble & Drabble (1927) reported that at one time (in Great Britain) they saw

inflorescences of Phleum pratense literally covered with Melanostoma mellinum and

Hylemia variata (Fallén) (Diptera, Anthomyidae), which were busy gathering

pollen. They believed that these insects are undoubtedly effective pollinators.

Dactylis glomerata was also visited for foraging purposes but its inflorescences do

not seem to be as attractive as those of other grasses.

Porsch (1956) gave an extensive survey of Coleoptera and Diptera recorded as

pollen consumers on anemophilous plant species, but his lists are chiefly compiled

from earlier publications and do hardly contribute any new data on syrphids.

For our present study the publication by Van der Goot & Grabandt (1970)

proved to be important. They studied the pollen content of the digestive tract of

several anthophilous species of Syrphidae belonging to the genera Melanostoma,

Platycheirus, and Pyrophaena. The senior author had repeatedly (in the Nether-

lands) observed these flies visiting the inflorescences of Gramineae and of

Plantago lanceolata. In order to assess their possible preference for anemophilous

plants the ingested pollen was studied. It appeared that the diet of several species,

viz., Platycheirus clypeatus (Meigen), P. fulviventris Macquart, P. scambus (Staeger)

STELLEMAN AND MEEUSE: Anthecological relations 17

and P. angustatus Zetterstedt, consisted almost exclusively, and that of others, such

as Melanostoma mellinum (L.) and M. scalare (Fabricius), partly, of pollen of the

above-mentioned anemophiles. (It may be mentioned in passing that Grabandt has

found pollen of Plantago and of grasses in related syrphids collected in Canada;

pers. comm.). In all other species of syrphid flies examined the diet normally

consists of pollen of entomophilous plants, anemophilous pollen apparently only

being taken occasionally.

All observations cited and pollen analyses clearly indicate a certain degree of

consistency in the visits of the Melanostoma-Platycheirus group to several

anemophiles, more particularly to Plantago lanceolata and a number of grasses.

The question arises whether these flies can indeed act as effective pollinators for

these reputedly anemophilous plants and if so, what the effectiveness of pollen

transfer by insects is in comparison to the anemophilous pollination. As mentioned

above, some authors suggest that pollen transfer by insects may be of some

importance, but this conclusion is solely based on recorded observations of visits.

In our opinion such listings of visitors, although indispensable, are only indicative

of a possible pollen transfer, and only an experimental analysis can lead to more

definite conclusions. Such an approach has hitherto never been attempted, as far

as can be ascertained.

The studies reported in the present paper were primarily intended to

demonstrate the possible incidence of an effective pollen transfer by syrphid flies

from one inflorescence to another, both through field studies and experimentally.

Three methods were employed:

(1) Field studies, mainly centred around the patterns of behaviour of the

syrphids, as far as important in connection with a possible pollen transfer;

(2) the use of the SEM technique to study the precise localisation of pollen

grains attached to the flies after a visit;

(3) experiments with stained (marked) pollen to demonstrate actual pollen

transfer in the field (chapter II).

For practical reasons it was deemed necessary, for the time being, to limit our

investigations mainly to P/antago lanceolata. The results obtained by Van der Goot

& Grabandt (1970) rendered the choice of the potential pollen vectors an easy

one; the following and (locally) relatively common species seemed to be the most

likely candidates: Melanostoma mellinum, Platycheirus clypeatus, P. fulviventris, and

P. scambus.

Since it is often impossible to identify each species with certainty in the field,

especially flying specimens and those paying a fleeting visit to an inflorescence

being hard to recognize, the flies were, generally speaking, treated as a group. This

admittedly is a somewhat weak element in our deductions, but we had no option.

Exact identification is always possible when the specimen is caught and can be

examined, but we refrained from this practice as much as possible. Not only is it a

time-consuming procedure, but we also did not want to extract so many

individuals from the ecosystem within the scope of our investigations. Identifica-

tions of killed specimens were reduced to an unavoidable minimum. Fortunately

we have obtained the strong impression from our observations in the field that the

behaviour of all species included in our study is very similar, and for the purpose

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

of the present investigation identical. Whenever the terms “syrphids” or “flies”

are used in the following text, unless stated otherwise, always representatives of

the Melanostoma-Platycheirus group are meant.

The report covers investigations carried out in 1973 and 1974; the field work was

carried out from about the beginning of May till the end of September. Low-lying

sites near water offered good opportunities. The main areas for study selected

were the Naardermeer and (less intensively) the Ankeveen broads. Incidental

observations were also made elsewhere in the province of Noord-Holland. The

SEM observations were made at the Laboratorium voor Electronenmicroscopie of

the University of Amsterdam.

Observations in the field

Observations in the field indicate that syrphids habitually visit inflorescences of

Plantago lanceolata. Spikes in the male stage of anthesis (P. lanceolata is

protogynous) are predominantly visited in the early hours of the morning after

sunrise. There is a marked correlation between the beginning of the (male)

anthesis and the activity of the flies. Both processes seem to be temperature-

dependant and commence in the temperature range between 10°—13°C, but may

to some extent also be incited by other factors. As soon as in a population of P.

lanceolata the male anthesis has begun, i.e., when the first stamens have emerged,

a few syrphids begin to arrive and descend on the inflorescences to start feeding

on pollen. Normally these events take place between 5.00 and 7.00 hrs, but on

relatively cool mornings, or when bad weather conditions prevail, the period may

shift to a later time in the morning.

In the beginning the male phase of anthesis develops slowly and the number of

visitors remains limited. When the temperature rises and the weather is not too

unfavourable, the anthesis rapidly becomes more general and the frequency of the

visits also increases appreciably in a short time span. Every day there is a relatively

short period of an optimum of visits which seldom exceeds a couple of hours and

falls between about 6.00 and 11.00 hrs, but the frequency usually diminishes

already before 10.00 hrs and only rarely stays high after 11.00. After the decrease

the visits diminish in number, and soon become more or less incidental to cease

completely at about 11.00 hrs; the flies become less active after feeding and

descend to various parts of plants to rest, but especially in the beginning they often

perform cleansing and preening movements with their legs.

The sequential phenomena just described are probably connected with the

following circumstances:

(i) More or less constant and at least fairly favourable weather conditions during

the early morning induce a certain regularity in both the course of anthesis and the

frequency of the visits until the optimum is attained;

(ii) Dissipation of the pollen from the anthers principally by air currents (wind

power!) results in the exhaustion of the pollen in the anthers which reduces their

importance as a source of food;

(iii) The flies become satiated after having consumed a large quantity of pollen

and start their resting phase.

STELLEMAN AND MEEUSE: Anthecological relations 19

Unfavourable weather in the morning, but sometimes during the previous

evening or night, may bring about appreciable deviations from the regular pattern

of anthesis and the visiting rhythm. Our observations concerning the male phase of

anthesis agree almost completely with the results of Hyde & Williams (1946)

obtained at Cardiff (Wales).

Continuous observation of a group of flowering plantains during which all visits

were recorded revealed that some of the spikes in anthesis are markedly more

frequently visited than most other ones and that a sometimes appreciable number

of spikes is not visited at all. The cause of this selective preference is obscure.

Quite regularly two or three flies are present on the same spike, but larger

numbers are usually not encountered: eight specimens were seen on a single spike

on only one occasion.

The pattern of behaviour of the flies suggests that some activities may be of

importance for a possible transfer of pollen. This requires some presuppositions,

VIZ.,

(a) that pollen grains indeed become attached to the body of the fly, and

(b) that the pollen can subsequently be transferred to the stigmatic area of a

different flower.

Some characteristics of behaviour established on the basis of numerous

observations are the following.

It appears that as a rule the syrphids fly in the direction opposite to the

prevailing movement of the air immediately above the stand of vegetation. This

mode of flight was used to advantage in experiments to be described in the second

chapter.

The approach flight towards a spike in anthesis is not necessarily followed by a

landing on the same spike; quite frequently the insect hovers in the air in front of it

for a short while to proceed to a different inflorescence where the movement may

be repeated before the fly decides to descend on a third spike, etc. The final

landing is preceded by a series of brief thrusting movements aimed at the

inflorescence. The fly usually alights directly upon the anther-bearing zones, but

not rarely on the portion of the spike above the stamens in which only the female

phase of anthesis has proceeded (i.e., on the stigma-bearing zone); in the latter

case the fly walks downwards towards the stamen-bearing part of the inflo-

rescence.

_ When manipulating the anthers to feed on the pollen, the fly is usually attached

to some place on the spike by means of its second and third pairs of legs and

clutches an anther with its front legs. It subsequently places its proboscis in the

dehiscence fissure of a theca and works it over along its full length. As soon as the

theca seems to become empty the anther is given a half turn by the front legs so

that the opposite slit comes within reach of the tongue. After having thus

systematically emptied an anther the fly grabs another one but may also move

owards the stigma-bearing zone of the spike or take off. The syrphids seem to

nrefer freshly dehisced anthers, but somewhat older ones are also used for

foraging, especially later in the morning. Pollen consumption is repeatedly

interrupted by preening (preening and cleaning of the body also take place when

the insects are resting after feeding as we have seen).

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

The duration of a single visit to a flowering spike varies considerably. From over

900 observations it could be deduced that visits lasting from less than 30 seconds to

2 minutes constitute about 50% of all vistis, and that 87% of the visits do not last

longer than 15 minutes. Visits of longer duration were recorded now and then; two

exceptional cases of visits, lasting 56 and 58 minutes, respectively, were noted.

During the working over of the anthers the flies may take up different positions.

This may be of importance in connection with the possible contamination of the

insect body with pollen and in some instances also with the possible deposition of

pollen grains adhering to the body on pollen-receptive stigmas. The most

characteristic postures are the following:

(i) the insect sits inside the stamen zone; contamination of different portions of the

body with pollen is possible (PI. 1 Fig. 1);

(ii) the fly takes up a practically vertical position distally of the stamen zone with

the head facing the anthers downwards; pollen may adhere to the frontal part of:

the body, and the second and third pairs of legs may come into contact with

receptive stigmas and thus transfer attached pollen grains (PI. 1 Fig. 2);

(iii) the insect is seated on the already wilted portion of the inflorescence below

the stamen-bearing zone in a vertical position with the head in the upward

direction; pollen may adhere to the frontal parts of the body but also to its under

surface (PI. I Fig. 3);

(iv) the fly clings in a nearly horizontal position to one or more stamens in the

lower part of the whorl of stamens; contamination of particularly the lower surface

of the body with pollen is possible (PI. 1 Fig. 4);

(v) in the case of spikes in which the zone of male anthesis has almost shifted to

the top: the insect sits on the tip of the inflorescence in a more or less horizontal

position; pollen grains will become attached to mainly the frontal parts of the

body, and deposition can take place as long as receptive stigmas are present (PI. |

Fig)!

More or less intermediate positions occur frequently, and during feeding the fly

may shift its position.

The syrphids are repeatedly found on the stigma-bearing zone of the inflo-

rescence, more particularly in the following situations:

(a) the fly alights on this part and after a shorter or longer sojourn proceeds to the

stamen-bearing zone; sometimes it stays for a while, usually cleaning and preening

its body; (b) after feeding on the pollen the insect walks from the stamen-bearing

zone towards the stigma-bearing part and usually stays there for some time

preening its body; it sometimes returns to the male zone to resume pollen

consumption, but may also fly away (PI. 1 Fig. 6).

If the female phase of anthesis has not terminated, the legs or the ventral side of

the body may get in touch with receptive stigmatic surfaces, which may result in

pollen transfer. In the first situation described above (a) the chance of cross-

pollination is greater than in the second (because in the latter case the pollen is

likely to originate from the same spike).

SEM studies of the captured flies

After the presence of pollen grains on the bodies of flies captured immediately

STELLEMAN AND MEEUSE: Anthecological relations 21

after a visit to a Plantago inflorescence had been established by examination under

a binocular dissecting microscope, a more detailed study was made by means of

SEM microphotographs. For practical reasons only attention was paid to the

ventral surface of the body (including the extremities). The study served a dual

purpose: the pollen grains present could be identified more accurately, and the

distribution of the pollen across the fly could be studied.

Methods

The flies were captured when they were resting on vegetative parts of plants and

never when they were sitting on inflorescences so as to avoid possible contamina-

tion with the large quantities of pollen shaken out of the anthers when one is

handling the killing jar. For each fly a separate, clean jar was used to avoid

indirect contaminations as much as possible. After a captured fly had become

sufficiently desiccated it was mounted with its dorsal side on an object support by

O7 À

Fig. I. “Pollen map” of a syrphid fly (see text). The black dots represent pollen grains of Plantago lan-

ceolata (dots situated outside the outline of the body represent pollen grains attached to setae protru-

ding from the surface of the fly body)

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

means of silver cement and coated with gold in an Edwards (model E 12 E)

vacuum-coating apparatus. The coated specimens were systematically scanned for

adhering pollen grains at suitable magnifications in a Cambridge Mark II

Stereoscan (acceleration potential 10 KV). A schematic outline drawing of a fly

was used to map the localisation and the species of the identified pollen grains; for

each specimen a so-called pollen map (an example of which is shown in Fig. 1) was

made in this way. Special and characteristic situations were photographically

recorded.

Results

In all 32 syrphids were studied which belonged to all four species mentioned in

the introduction. On all but one of these specimens the presence of pollen grains

could be demonstrated. This pollen was almost exclusively of Plantago lanceolata; :

only in two cases a few gramineous grains were noted. The number of grains per

fly varied a great deal: from a single one to a recorded maximum of 179 with a

mean of 44. The localisation of the pollen on the ventral body surface was as

follows: on the head only a few grains were observed, if any, and even around the

mouth there were very few or none. The thorax carried the greatest number as a :

rule, especially in the pleural parts and on the hairs found in this part of the body.

The abdomen bore fewer grains, mainly on the belly. Of the leg parts the femur

carried the largest number of grains; between the three pairs of legs no important

differences were found.

Discussion

The scanning technique demonstrated conclusively that pollen grains of

Plantago lanceolata adhere to the bodies of the syrphid flies. The relatively few

number of specimens studied do not permit an accurate assessment of the

distribution of the pollen on the bodies of the flies, but it seems as if the head does

not carry many grains. The grains are sometimes found on the glabrous cuticle,

but more often adhere to the dermal setae and do not always touch the cuticle.

The attachment is almost certainly attributable to a cementing substance, whose

presence (in small amounts) in Plantago was shown by Pohl (1930) and Knoll (1930)

almost simultaneously. On SEM microphotographs (compare Plate 2) the pollen

grains, when lying close together, are frequently connected by a mass forming a

conspicuous meniscus. Also between a grain sticking to a hair and the support

such a substance is present. Pollen grains pretreated with ethanol before SEM

examination do not show this phenomenon, so that it appears to be highly

probable that the connecting matter is hardened pollen ‘Kitt’? covered by a gold

film. The almost complete absence of grass pollen on the flies is of special interest.

Conceivably, its relatively great rarity has something to do with the incidental

nature of visits to species of Gramineae for feeding purposes. It is also known that

the surface of gramineous pollen grains is only very thinly covered with Pollenkitt,

although the absolute quantity may vary appreciably from species to species (Pohl,

1930); this small amount of cementing substance may account for the poor sticking

capacity of the grains (or for a short duration of their attachment during the flight

STELLEMAN AND MEEUSE: Anthecological relations 23

of the insect). One must also bear in mind that the handling of the flies during the

capture and their subsequent preparation may rub off a number of grains from the

fly body, so that the SEM images do not necessarily present the situation

immediately before the insects were caught: grass pollen may be rubbed off more

easily than the plantain pollen.

Concluding remarks

Field observations of syrphid flies regularly feeding on P/antago lanceolata

indicate that these insects may easily become contaminated with pollen. More

casual visits to the inflorescences of grasses with easily accessible anthers may also

result in the adherence of pollen to the body.

The efficiency of an insect as a pollinator is largely dependent on the constancy

of its habits and its patterns of behaviour. If it frequently moves from one plant in

anthesis to the inflorescence of a different individual of the same plant species the

chance of transfer of specific pollen is appreciable. However, the efficacy of

zoophilous pollination is also determined by the amount of pollen carried by the

animal vector on its body, which amount differs from plant species to plant species

owing to the varying quantities of pollen and to the presence or absence of other

aids for the adherence of the grains to the insect body (such as a rough or

spinulose ectexine). As far as can be deduced from the available data, grass pollen

is less effectively transferred than plantain pollen, which may be interpreted as a

greater reliance upon pollination by syrphid flies of Plantago lanceolata as

compared to the grasses, at least in the area under investigation. This needs

verification, and this relation should also be studied in other habitats.

The different amounts of plantain and grass pollen recorded on captured flies

support the idea of a greater dependence of Plantago lanceolata on the flies in its

pollination strategy. The repeated visits of syrphids to different individuals in

stands of plantain render an efficacious pollen transfer highly probable. It also

enabled the demonstration of the transfer of pollen from one plant to another to

be dealt with in the next chapter.

II. EXPERIMENTS WITH STAINED POLLEN OF Pantago lanceolata L.

After the transfer of pollen of Plantago lanceolata by certain syrphid flies had

been shown to be highly probable (see the first chapter), experiments were carried

out to demonstrate the actual translocation of pollen grains from one plant to

another by means of pollen marked by artificial staining. It has already

conclusively been shown by means of SEM microphotographs that pollen grains

adhere to various parts of the fly body, but although there were cogent indications

of the role of the insects as effective pollen vectors, there remained some

uncertainty whether a deposition by the flies of pollen grains on the receptive

stigmatic surfaces of a flower in the female stage of anthesis actually takes place.

By using artificially coloured grains partly replacing the normal pollen the

identification of pollen grains deposited on other inflorescences was made

possible, so that the displacement of this pollen could be traced. In addition, it was

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

hoped that from such experiments certain ideas concerning the efficacy of a biotic

pollen transfer under field conditions could be gleaned.

The marking of pollen by staining

Some preliminary tests had shown that pollen of P/antago lanceolata can readily

be dyed by means of aqueous solutions of biological stains such as 0.5 %

Methylene Blue and 0.5 % Neutral Red. Attempts to dye pollen grains still

enclosed in the ripe anthers by spraying the inflorescences with such staining

solutions were not very successful. Only a relatively small amount of pollen

present near the slit of dehiscence appeared to have absorbed the dye, but the

solution did not penetrate far enough. The anther wall may be more or less water-

repellent. It was decided to use a technique in which pollen collected in the field

was dyed in the laboratory as follows.

Pollen from spikes bearing numerous stamens in anthesis was harvested by

beating the spikes over a metal tin. To obtain a quantity sufficient for subsequent

processing at least 20-30 inflorescences must be shaken out. A small quantity of

the collected pollen is poured on a disc of filter paper covering the bottom of a

petri dish and spread out evenly by means of a fine paint brush. The dish is sprayed

with the dye solution from a flower sprayer until the filter paper has assumed the

colour of the solution. The (open) dish is left to dry in a suitable place at room

temperature, which takes 2-3 hrs. To assess the condition of the grains the dish is

placed under a dissecting microscope and the grains are touched with a fine

preparation needle: if they are loose and can be easily shifted (or even jump away)

the pollen is sufficiently desiccated. It is subsequently transferred to a glass tube

_ with a brush and can be kept for some time until needed.

Pollen treated in this way was examined for some properties required for the

purpose of the present investigation:

(i) Stained and normal (= untreated) pollen grains observed under a micro-

scope at magnifications of 100—200 diam. appear to be always clearly distinguish-

able from one another;

(ii) When marked pollen was poured on to receptive stigmatic surfaces and the

stigmas were subsequently examined under a dissecting microscope, such treated

pollen grains appeared to adhere firmly to the stigmatic papillae and were clearly

distinguishable in situ from untreated grains;

(iii) When living syrphid flies of the taxa studied (Melanostoma, etc., see the first

chapter of this paper) were placed in a tube with dyed pollen, left in the tube for

several minutes, and narcotised for examination under the microscope at a

magnification of about 40 diam., they proved to carry a substantial amount of

pollen on their bodies; the localisation of the pollen grains was very much the

same as in the case of contamination with untreated pollen.

The treated grains apparently do not differ appreciably from undyed ones in

some properties essential for their transfer by flies. To remove possible remaining

doubts, the power of adherence of untreated and of dyed pollen was tested by

locking up syrphids in a petri dish containing a small amount of a mixture of

untreated and dyed grains in a known quantitative relation. After a brief sojourn

STELLEMAN AND MEEUSE: Anthecological relations 25

the flies were narcotised and their bodies were examined for adhering pollen.

From counts of the two kinds of grains for each fly (42 specimens) the relation

between untreated/dyed pollen was estimated.

The result was as follows:

mean ratio untreated/dyed pollen in the mixture: 4.41 + 0.12 [mean ratio

untreated/dyed pollen on the fly bodies: 5.18 + 0.54].

Experiments in the field: methods

For the experimental set-up the following reasoning was followed (see also the

situation sketch, Fig. 2): When of two groups of flowering plants in a sufficiently

direction

Saw BE Da =

wind

a e A

tential testslid. d

P “receptor spikes er spikes

min. 40 cm

Fig. 2. Situation sketch of a field experiment (explanation in text). For the sake of clarity the rosette of

leaves is drawn as if they are adpressed to the ground (as in Plantago major and P. media)

dense stand of Plantago lanceolata one (A) is used as the donor group by applying

dyed pollen to the spikes in the manner to be described below, and the other

group (B) is left undisturbed, a syrphid fly visiting one or several donor spikes is

likely to become covered with pollen including a number of treated grains. If it

subsequently alights on untreated spikes there is a distinct possibility of dyed

pollen becoming deposited on the stigmas of these spikes, which can be

established by examining such spikes under a dissecting microscope (PI. 3 Fig. B).

During the practical execution of the experiments the possible transfer of wind-

borne pollen had to be taken into account. The relative position of the group of

donor spikes was chosen in such a way that it always remained below the wind in

respect of the untreated spikes present in the neighbourhood. A very favourable

circumstance is that the syrphids tend to fly against the direction of the air current

over the site, in the figure from group A towards group B. It goes without saying

that this increases the efficacy and the reliability of the experiment.

In order to establish if nevertheless an undesired pollen transport of stained

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

pollen took place, test slides were placed between the two groups which were

coated with egg-white glycerol on the one side to catch air-borne pollen grains;

these slides were placed at a height a little below the average height of the donor

spikes, with the smeared side facing the latter. The distance between the groups of

donor plants and receptor plants must not be too short (at least 40 cm) so as to

avoid the risk of an undesired pollen transfer by direct contact between plants or

by air turbulence. The area chosen for such experiments was always limited

enough to be easily surveyable from one observation post and thus to permit the

reliable recording of visits to receptor spikes by flies coming from donor spikes by

following the flight of the latter.

After some trials a method was worked out to bring about a contamination of

flies with dyed pollen which agrees to all intents and purposes with the mode of

contamination with untreated pollen during a normal visit to an inflorescence.

Dyed pollen is placed on a flowering spike as follows: A sufficient quantity of dyed :

pollen is put in a glass tube of about 10 cm long and 1.5—2 cm in diam. By shaking

and tapping the tube whilst turning it about its longitudinal axis in a nearly

horizontal position the pollen is evenly distributed over the inner wall. Such a tube

is placed over a spike in male anthesis with a well-developed zone of stamens, the

inflorescence during this and the subsequent operations being held in a horizontal :

position by bending its stalk carefully. By pushing the spike to and fro and, if

necessary, by turning the tube a bit, some of the dyed pollen is rubbed off and

sticks mainly to the anthers (but also to other parts of the inflorescence such as the

stigmatic zone). The spikes must be dry; damp ones moisten the inside of the tube

too much so that the pollen sticks too firmly to the tube and becomes useless.

Such a treatment of the inflorescences does not deter flies from visiting them

normally, even if the whorl of stamens has assumed a somewhat unnatural colour.

The dyed pollen grains are as readily consumed as untreated ones; in a number of

cases a marked bluish-green discolouration of the abdomen of a fly was observed

after the animal had been feeding copiously on spikes treated with stained pollen.

It may, therefore be taken for granted that the deposition of marked pollen grains

on a spike does not affect the normal behavioural pattern of the flies in the least.

The added pollen is also dispersed in a normal way by air currents as could be

deduced from the gradual disappearance of the greyish discolouration, after some

time, of a spike treated with pollen dyed blue, even if no insect visits took place.

Examination of flies captured after they had visited a donor spike revealed the

presence of coloured pollen grains on the fly body (PI. 3 Fig. A).

Experiments in the field: results

Altogether 22 experiments were carried out, the duration of each single

experiment being 60 to 150 minutes. The number of (treated) donor spikes was 3-6

and the receptor spikes usually numbered 3—4. Of about 90 spikes which were

gathered in the field and might carry stained pollen grains ultimately only 67 could

be used for the records, because some mishaps occurred which rendered them

unusable (some became soaked with water, etc.).

The following qualifications need some explanation.

STELLEMAN AND MEEUSE: Anthecological relations 27

Fresh stigmas: stigmas with a shiny, somewhat vitreously white appearance; the

distal parts may be suffused with brown.

Old stigmas: stigmas which have mostly or completely turned brown and have

often become desiccated.

The duration of the receptive phase of the stigmas is not exactly known, but

nevertheless the distinction made above, based on a different appearance,

corresponds most probably with the receptive phase and the phase of post-

anthesis of the stigmas.

After the experiments were concluded, a publication by Zeisler (1938) came to

our notice in which a simple method is described to assess the degree of

‘maturation’ (receptiveness) of the stigmas by a chemical reaction (with H,O,).

If this test is applicable to Plantago stigmas, some relevant additional

information may thus be acquired.

Whole spike: includes bracts, perianth members, stigmas, and a zone of

stamens.

Stigma ratio (s.r.): number of stigmas with stained pollen grains

total number of stigmas

Control with test slides: The pollen counts were made of a rectangular area of 10

mm by 20 mm in the middle of the slide at a magnification of 100—200 diam. Not

in a Single instance stained pollen was recorded, but a few undyed grains were

usually present; the latter may have been deposited by air turbulence or

Table 1. Presence or absence of dyed pollen on spikes of Plantago lanceolata L.

Marked pollen present in Marked pollen absent in

On fresh stigmas 43 spikes 24 spikes

On fresh and/or old stigmas 48 spikes 19 spikes

On whole spikes 50 spikes 17 spikes

convection currents, or as fall out of the local pollen rain from somewhat higher

levels. In a number of cases test slides were also placed in a downwind position

and these almost invariably had caught large quantities of both stained and

unstained pollen.

The (67) receptor spikes were examined for the presence of marked pollen

grains and the results are as follows (Table 1).

Table 2. Distribution of dyed grains over spikes of Plantago lanceolata L.

Number of dyed grains Number present

On fresh stigmas On old stigmas On whole spike

(0) (24) (19) (17)

1-10 25 15 11

11-20 8 14 11

21-50 7 14 18

51-100 2 2 5

more than 100 1 3 5

43 48 50

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

The following survey shows the number of individual grains in the cases in

which dyed pollen had been transferred (Table 2). In the accompanying tabel

(Tabel 3) the stigma ratio of 41 spikes with transferred dyed pollen grains, and the

Table 3. Stigma ratio (s.r.) and number of grains on 41 spikes with transferred dyed pollen grains

ST. number of SIT. number of Sir number of

grains grains grains

3/9 - 8 10/10 - 136 Vile = 3

1/10 - 4 2/2 - 19 6/10 - 36

6/1 A= 63 7/9 - 47 ar "=" 20

MOSE 8 3/6 - 19 18/8 682

Al anr 4/8. - 26 2/6 - 2

8/10 - 47 SIATE 12 6/9 - 26

1/29 - 2 VA - 2 4/11 - 4

2/4 - 9 W237 4 2/26 - 9

4/8 - 31 1/21 - 12 1/8 - 1

2/11 - 7 2/4 - 3 W/22 RE 1

5/17 - 10 4/14 - 14 3/3) 2219

2/3 - i vl - 2 3/40 6

Diss 5 2/14 - 5 VO er IO

Aje RO 3/21 - 13

number of grains on the stigmas is shown. Of two of the 43 spikes with stained

pollen grains the number of fresh stigmas was inadvertently not counted.

Discussion

The examination of the test slides indicates that an abiotic transportation of

pollen against the direction of the wind is negligible. The conclusion that can be

drawn from the results of the first survey (Table 1) is that syrphid flies are

instrumental in the transfer of pollen from one spike to another and can do this

repeatedly. The absence of marked pollen grains on potential receptors can have

several reasons, the most important being:

(a) that the spikes were not visited by flies, or

(b) that the spikes were visited, but by flies that did not carry stained pollen

grains.

One can only speak of an effective pollination if pollen is deposited on fresh

(and presumably receptive) stigmas. This was the case in 20 experiments and in

64% of the total number of spikes examined, so that the general conclusion

remains valid. One of the surveys (Table 2) gives a good idea of the quantities of

marked pollen grains transferred by the animal pollen vector, but the results do

not require a special discussion.

From the viewpoint of efficacy of pollen transfer the number of stigmas on

which dyed pollen had been deposited is important. Table 3 shows that there are

appreciable differences in this respect, both in stigma ratio and in the number of

stained grains. This is to be expected if one considers that a number of mutually

independent factors is involved in the pollen transfer to the stigmas, viz.,

(a) the total number of visits to a spike,

STELLEMAN AND MEEUSE: Anthecological relations 29

(b) the duration of these visits,

(d) the amount and the location of the pollen on the body of the visitor, and

(e) the number and the length of the stigmas on the particular spike.

The first two factors could be quantified by the registration of the visits, but

there was no correlation with the quantity of deposited marked pollen. Matters

were complicated by the uncertainty in many cases whether a visitor was indeed a

carrier of dyed pollen. The dyed grains were especially deposited on the tips of the

stigmas as may be expected because these protruding parts are the most likely to

come in touch with the insect body. It is also noteworthy that stained grains were

localised on the stigmas in groups.

Another survey (Table 4) demonstrated that of all pollen-bearing stigmas only

Table 4. Distribution of stained pollen grains over the stigmas

36 stigmas (24.3%) with 1 grain

24 stigmas (16.2%) with 2 grains

24 stigmas (16.2%) with 3 grains

64 stigmas (43.3%) with 4 or more grains

24% had caught a single dyed grain, whereas in 76% of the cases two or more

grains were present. Since the gynoecium contains 2(-3) ovules the theoretical

chance of efficacious fertilization is rather great.

The experiments described in the previous chapter render it highly probable

that pollen treated with stains has practically the same properties as untreated

pollen at least as far as its power of adhesion to the insect body and of its

successful translocation is concerned. It is, in our opinion, not necessary to put its

equivalence with regard to adhesion to the stigmatic surface to the test: the sticky

stigmas readily retain all sorts of particles (some of which are larger than the

pollen grains) and their capacity to capture pollen grains by their adhesive

properties certainly exceeds the affinity of the pollen grains to the surface of the

insect body. Various considerations render the same chances of being transferred

by biotic agencies. Considering that untreated pollen seems to adhere slightly

better to the insect body, the transfer of untreated pollen by flies can be accepted

beyond reasonable doubt, even if the transfer of untreated grains from a donor

spike to a receptor spike could not be directly demonstrated: untreated pollen

found on receptor spikes may have been deposited by a fly coming from a donor

spike but just as well from various other sources outside the experimental area.

One must bear in mind that the duration of the experiments was usually shorter

than the time-span of the daily visiting period (although it always included the

phase of optimum activity of the flies), so that an — albeit relatively small—part of

the potential pollen transfer was not recorded. This only means that the rate of

biotic pollen transfer is undoubtedly slightly higher than it appeared to be under

our experimental conditions. The fact that the flies often move about on the same

spike (and preen themselves) favours the incidence of geitonogamy, and so does

their habit of frequently moving towards inflorescences in the immediate

neighbourhood (which may well belong to the same individual of Plantago

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976

lanceolata), but our experiments have shown that effective cross-pollinations are

by no means rare.

As far as the relative importance of efficiency of the biotic and the

anemophilous form of pollen transfer is concerned, the present experiments do

not permit a clear verdict. A specially adapted method (by means of fertilisation

experiments?) would have to be worked out for the purpose.

An ecologically interesting and relevant circumstance is that the syrphid flies of

the Platycheirus-Melanostoma group as a rule visit the Plantago inflorescences

during the first hours of the morning (see the first chapter). At that time of the day

the wind is often hardly noticeable or at least very weak, so that presumably the

relative share of the biotic pollen transfer in respect of that by air currents will be

greater than later during the day. Local differences in the relative rates of the two

types of pollination associated with different environmental conditions and habitat

variation (such as landscape and vegetation type and microclimatological

differences) cannot be precluded. The question in how far these variations may be

correlated with, or reflected in, modifications of the inflorescence is of interest.

Delpino (1870, cited and discussed in Muller, 1873) thought he could recognise

forms with differences in structural adaptation, which was accepted with some

diffidence by the latter author.

One should never loose sight of the fact that a biotic transfer of pollen of

Plantago lanceolata as described in the present paper can only take place where

this plantain and the potential pollinators occur sympatrically, which need not

always be the case. Field observations and experiments in other areas are

indicated, and it will be necessary to extend the investigation by comparing the

anthecological syndromes of other species of Plantago with the situation in P.

lanceolata.

The technique for the dying of pollen for purposes of specific identification after

a possible transfer can probably be improved and extended to be used effectively

for the study of other cases in which different taxa and plants and other (potential)

pollen vectors are involved. Coldwell (1951) has developed a useful method to

study the dispersal pattern of air-borne pollen (of different species of conifers) by

using pollen labelled with radioactive substances (isotopes). This technique is

rather laborious, however, and other workers who have attempted to apply this

method have not been successful (B. J. D. Meeuse, priv. comm.). The use of

artificially stained pollen permits a more or less small-scale approach to various

anthecological problems, more particularly in border-line cases between en-

tomophily and anemophily.

ACKNOWLEDGEMENTS

The authors are highly appreciative of the active assistance of Mr. V. S. van der

Goot (Amsterdam), especially with the identification of the syrphid flies. The

technical assistance of Mr. C. Bakker (Werkgroep Scanning-Electronenmicro-

scopie, Amsterdam) and of Messrs. F. D. Boesewinkel, L, Dijkhuizen, C. L.

Hansson, H. J. Koerts Meyer, A. Smit and J. Vuijk (Hugo de Vries Laboratorium)

is thankfully acknowledged.

STELLEMAN AND MEEUSE: Anthecological relations 31

The permission granted by the „Vereniging tot Behoud van Natuurmonumenten

in Nederland” to do field work in the Naardermeer nature sanctuary deserves

grateful memorisation.

REFERENCES

Coldwell, R. N., 1951. The use of radioactive isotopes in determining spore distribution patterns. —

Amer. J. Bot. 38: 511—523.

Delpino, F., 1870. Applicazione della teoria Darwiniana ai fiori ed angli insetti visitatori dei fiori. Dis-

corso pronunziato del Dr. Erm. Mueller di Lippstadt. Versione dal tedesco, e annotazioni. —

Bullet. della Soc. Entom. Ital. 2: 140—159.

Drabble, E., & H. Drabble, 1927. Some flowers and their Dipteran visitors. — New Phyt. 26: 115—123.

Faegri, K., & L. van der Pijl, 1971. The principles of pollination ecology (2nd. ed.). — Pergamon Press,

Oxford.

Goot, V.S. van der, & R. A. J. Grabandt, 1970. Some species of the genera Melanostoma, Platycheirus

and Pyrophaena (Diptera, Syrphidae) and their relation to flowers. — Ent. Ber. 30: 135—143.

Hyde, H. A., & D. A. Williams, 1946. Studies in atmospheric pollen III. Pollen production and pollen

incidence in ribwort plantain (Plantago lanceolata L.). — New Phyt. 45: 271—277.

Knoll, F., 1930. Über Pollenkitt und Bestäubungsart. — Z. Bot. 23: 609—675.

Knuth, P., 1898—1905. Handbuch der Blitenbiologie I—III. — Engelmann, Leipzig.

Kugler, H., 1970. Blütenökologie (2nd. ed.). — Fischer, Stuttgart.

Ludwig, F., 1881. Weitere biologische Mitteilungen. I. Molinia coerulea als Fliegenfangerin. — Bot.

Centralbl. 8: 87.

———, 1884. Uber den Fliegenbesuch von Molinia coerulea. — Bot. Centralbl. 18: 123.

Müller, H., 1873. Die Befrüchtung der Blumen durch Insekten. — Engelmann, Leipzig.

Pohl, F., 1930. Kittstoffreste auf der Pollenoberflache windblitiger Pflanzen. — Beih. bot. Centrbl. 46,

- (I): 286—305.

Porsch, O., 1956. Windpollen und Blumeninsekt. — Osterr. Bot. Z. 103: 1—18.

Zeisler, M., 1938. Uber die Abgrenzung der eigentlichen Narbenflache mit Hilfe von Reaktionen. —

Beih. bot. Centrbl., sect. A, 58: 308—318.

Tui '

‘ ‘ t

dur.

bal

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976 Pl. 1

Different positions of syrphid flies on inflorescences of Plantago lanceolata (explanation in text)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976 PINS

3 le

SEM microphotographs of pollen grains of Plantago lanceolata on the bodies of syrphid flies.

1, X 600; 2, X 720; 3, X 900; 4, X 900

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 2, 1976 Pl. 3

A. Part of the head of a syrphid fly to which normal (light coloured) and stained (dark coloured) pollen

grains are attached, X 50. B. Stigma of an inflorescence of Plantago lanceolata collected after a visit by

a fly, showing the presence of untreated (light coloured) and stained (dark coloured) pollen

grains, X 50

‘

Na Seiad

a > È n

è x

DEEL 119 AFLEVERING 3 1976

TIJDSCHRIFT

VOOR ENTOMOLOGIE

MUS. COMP, Zels:

a LIBRANY

4 UITGEGEVEN DOOR

7 (AUG 1 7 15,5

i HARVARD

3 DE NEDERLANDSE ENTOMOLOGISCHE VERENIGINERSITY.

bee INHOUD

c. VAN ACHTERBERG. — A preliminary key to the subfamilies of the Braconidae

_ (Hymenoptera), p. 33—78, fig. 1—123.

Gepubliceerd 7-VII-1976

A PRELIMINARY KEY TO THE SUBFAMILIES OF THE

BRACONIDAE (HYMENOPTERA)

C. VAN ACHTERBERG

Rijksmuseum van Natuurlijke Historie, Leiden

With 123 figures

Abstract

A preliminary key to the subfamilies of the Braconidae is given. The subfamilies are redefined in

the key and in a short description. Especially the apomorphous characters, which may be used to

define the subfamilies, are discussed.

INTRODUCTION

The family Braconidae can be separated from its sister-group, the Ichneu-

monidae, by at least two synapomorphous characters. The most important is the

rigidly joined 2nd and 3rd tergites of the metasoma and, secondly, the absence

of the second recurrent vein. The rigid plate formed by the 2nd and 3rd tergites

may be weakly sclerotized (as in the Aphidiinae) or may be completely

fused, e.g., in the Hybrizontinae and Acaeliinae.

The first worker, who tried to give a classification of higher groups was

Wesmael (1835), whose classification concerned only the Palaearctic region. The

Braconidae were divided into two divisions by Wesmael (p. 11): the “braconides

endodontes” and the “braconides exodontes’’. The latter group is what is now

called the Alysiinae. The “‘endodont’’-Braconidae were divided into four sub-

divisions (p. 14): (1) the “polymorphes”, (2) the “‘cryptogastres’’, (3) the “areo-

laires”, and (4) the “cyclostomes”. The “polymorphes” contain the Aphidiinae,

Euphorinae, Helconinae, Ichneutinae and Opiinae. The “cryptogastres” are an

aggregate of Cheloninae and convergent groups, belonging to the Helconinae and

Meteorideinae. The ‘‘areolaires”’ embrace the convergent groups Microgaster-

inae, Acaeliinae, Orgilinae and Agathidinae. Finally the ‘“‘cyclostomes”’ contains

the Doryctinae, Rogadinae and Braconinae, as treated in the present paper. The

first reviser of this system was Foerster (1862), who gave a very elaborate

sub-division and defined 26 ‘‘subfamilies’’, to which he added the suffix ‘‘-oidae’’.

The following systems were based more or less on the Foerster-system, but

several “subfamilies” were given tribal rank in one large subfamily. E.g., the Bra-

conoidae, Euspathioidae, Hecaboloidae, Doryctoidae, Hormioidae, Rogadoidae

and Rhyssaloidae became in Fahringer (1925) the tribes Braconini, Spathiini,

Hecabolini, Doryctini, Hormiini, Rhogadini, Exothecini and Pambolini in one

subfamily Braconinae, in effect the old “cyclostomes” of Wesmael. The “‘Sigal-

phoidae” became a part of the Helconinae, while the “Chelonoidae” are the

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

present Cheloninae and Meteorideinae. The ““Microgasteroidae” contain the

Microgasterinae and Acaeliinae, and the Agathidinae were divided by Foerster in

the “Agathidoidae” and the “Eumicrodoidae”, while the last mentioned also

contains the Orgilinae p.p. The “Pachylommatoidae” and “Aphidioidae” are

called Hybrizontinae and Aphidiinae, respectively, in the present paper. The

“Euphoroidae” and “Perilitoidae” form the Euphorinae, from which quite

recently Zemiotes Foerster was excluded and added to the Zelinae (Mason,

1973). The “Brachistoidae” of Foerster are of composite nature, they belong

partly to the Helconinae and partly to the Zelinae (as defined in the present

paper). The “Livphronoidae”, together with Pygostolus Haliday from the “Bla-

coidae”, form the tribe Centistini in the Euphorinae. The remainder of the “Bla-

coidae” belongs to the Helconinae. The “Ichneutoidae” and “Helconoidae”

became the Ichneutinae and Helconinae p.p., respectively. Foerster’s “Macro-

centroidae” contain the Macrocentrinae and Zele Curtis of the Zelinae; this:

grouping is still widely used, e.g., by Eady & Clark (1964) and Capek (1970). The

‘“Diospiloidae” belong to the Helconinae and Orgilinae. Except for Gnaptodon

Haliday (which is included in the Rogadinae), his “Opioidae’” are fully com-

parable with the Opiinae. Ultimately his ““Alysioidae” and “Dacnusoidae” are

united by Griffiths (1964: 831) in the subfamily Alysiinae; this was confirmed.

through the study of the larvae by Capek (1970: 861).

The system used by Marshall (1885: 9) is essentially that of Wesmael, but he

added as a sixth division the “Flexiliventres” for the Aphidiinae, because of their

flexible and weakly sclerotized metasoma. In 1891 he added as a seventh division

the “Pachylommatidae”, now called Hybrizontinae. These mainly large groups

were subdivided according to the system of Foerster with some modifications,

but also resulting in 26 “subfamilies” (for which Marshall used the suffix ‘‘-ides’’)

for the Palaearctic region.

The key by Ashmead (1900: 111) to the subfamilies of the Braconidae is the first

general key, intended to be used in more than one region. Ashmead separated the

Alysiinae as a family Alysiidae, while the remaining genera were divided among 17

subfamilies. His division and interpretation of the genera was often incorrect,

because he used a modification of the Foerster system without examining the

types of many genera.

The second and latest general key to the subfamilies, which was used more

extensively, is the key published by Szépligeti in 1904. It divides the Braconidae

into 31 subfamilies, one of which (the Lysiognathinae) belongs in the Ichneumo-

nidae. From the remaining 30 subfamilies the following are removed and reduced

to lower rank in the subfamilies mentioned in parentheses: Cenocoelininae

(Helconinae), Gnathobraconinae (Rogadinae), Aphrastobraconinae (Braconinae);

Exothecinae (Rogadinae), Spathiinae (Doryctinae), Hecabolinae (Doryctinae),

Pambolinae (Rogadinae), Hormiinae (Rogadinae), Sigalphinae (Helconinae),

Calyptinae (Helconinae), Liophroninae (Euphorinae), Blacinae (Helconinae),

Cardiochilinae (Microgasterinae), Diospilinae (Helconinae), Meteorinae (Eupho-

rinae) and Dacnusinae (Alysiinae). The remainder, 14 subfamilies, are augmented

to 22 in the present paper, because of three newly described subfamilies (Yp-

sistocerinae Cushman, 1923; Telengainae Tobias, 1962; Mesostoinae Van Achter-

C. VAN ACHTERBERG: The subfamilies of the Braconidae 35

berg, 1975) and a rearrangement of five groups according to Capek (1970 and

1973) and Tobias (1967). These five subfamilies are the Acaeliinae and Zelinae

(according to Tobias), the Orgilinae, Meteorideinae and Neoneurinae (modified

after Capek).

The two subfamilies erected by Fahringer (1936: 586) in his fairly general (but

rather confusing) key are not accepted in the present paper. His Aneurobraco-

ninae (containing the genus Aneurobracon Brues) is provisionally treated in this

paper as a tribe of the Agathidinae, while the Pseudodicrogeniinae (containing

only the genus Pseudodicrogenium Fahringer) is included as a tribe in the Braconi-

nae. The morphology of both types examined clearly justify this transfer (Fig.

120—122).

After Szépligeti, several others have given subfamily-keys for the Palaearctic

region (e.g., Fahringer, 1925; Tobias, 1971), while Marsh (1963: 522) gave a key

for the Nearctic region. This key was disregarded in his later key to the Nearctic

genera (1971: 841), because “there is some disagreement as to the limits of the

various subfamilies in the Braconidae,....’’.

For the terminology used, see Van Achterberg, 1976.

DISCUSSION

The main reason for the confusion in the systematics of the Braconidae is

formed by the many convergent evolutionary trends, occurring in this group.

Some of the most important trends are the following:

|. The reduction of the veins of the wing, e.g., the Blacini in the Helconinae

versus the Centistini in the Euphorinae.

2. The forming of rows of setae, usually one row per tergite. In this respect I

disagree with Griffiths (1964: 842), who stated that “the arrangement of hairs in

single rows on the gastral segments, being found in almost all other Alysiinae and

many other Braconidae is beyond doubt plesiomorph’’. I am convinced that the

opposite is true: many species with many plesiomorphous character-states have

the tergites evenly setose. Still more convincing is the fact that most Ichneu-

monidae (the sister-group, in most aspects clearly less evolved) have the meta-

somal tergites evenly setose.

3. The formation of a carapace, formed by the three basal tergites of the

metasoma, e.g., in the Helconinae (Brachistini), Cheloninae, Microgasterinae

(Fornicia Brullé), and Rogadinae (Tobias & Dudarenko, 1974).

4. The ovipositor becomes shorter, sometimes more or less curved, and its

sheaths become often wider.

5. The reduction of the segments of the palpi; the plesiomorphous condition

of the maxillary and labial palp is 6 and 4 segments, respectively. Lower numbers

occur in e.g., the Braconinae, Alysiinae, Helconinae, Neoneurinae and Hybri-

zontinae.

6. The selection of hosts other than Coleoptera larvae, especially Lepidoptera

and Diptera.

7. The development of endoparasitism.

8. The parasitism of adult insects by the Neoneurinae, Aphidiinae and Eupho-

rinae.

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

9. The reduction of the number of antennal segments occurs in almost all

groups.

10. The development of a petiolus at the basis of the first metasomal tergite:

e.g., Doryctinae (Spathiini), Euphorinae (Meteorini) and Zelinae (Zemiotini).

11. The forming of a hypoclypeal depression and a more or less concave

clypeus: e.g., Braconinae, Opiinae, and Euphorinae (Cosmophorini).

12. The twisting of the mandible: e.g., Macrocentrinae, Braconinae and, very

extremely, in the Alysiinae.

13. The formation of a dorsope and/or a laterope, while often also the TA

carinae become more developed, until beyond the middle of the first tergite: e.g.,

Helconinae, Rogadinae, Alysiinae and Euphorinae.

14. The spiracle of the first metasomal tergite becomes situated near the middle

of the tergite: e.g., Euphorinae and Zelinae (Zemiotini).

15. The radial cell becomes short and/or narrow, a tendency occurring in

almost all subfamilies.

16. The first metasomal tergite becomes depressed, especially laterally: e.g.,

Mesostoinae, Braconinae, Microgasterinae, Acaeliinae and the tribe Proteropini

of the Ichneutinae.

17. The convergent development of an “ophionoid facies” (Gauld & Huddles-:

ton, 1976) an adaption for nocturnal activity, occurring in the Zelinae, Euphori-

nae, Macrocentrinae and Rogadinae.

18. The mandibles of the larvae become slender and toothless: e.g., Euphorinae,

Neoneurinae, Aphidiinae, Alysiinae, Opiinae, Microgasterinae p.p., and Rogadi-

nae p.p.

19. The reduction of the cephalic structures of the larvae, especially of the

hypostoma: e.g., Euphorinae, Orgilinae, Helconinae (Blacini & Brachistini),

Neoneurinae, Ichneutinae (Muesebeckiini), Opiinae, and Doryctinae.

20. The labial and stipital sclerites of the larvae become long and slender: eg,

Euphorinae, Helconinae, Zelinae, and Opiinae.

Mainly owing to the research by Capek upon the cephalic structures of the

final instar larvae, I have tried to construct a key to the subfamilies proposed

by Capek and Tobias. Some changes seem inevitable to me, e.g., the fusion of the

subfamilies Rogadinae and Exothecinae. There are too many genera in these

groups, that connect the groups in more than one character. The mummification

of caterpillars by Rogas and closely allied genera is not sufficient (however

peculiar it may be) to separate them as a subfamily. The Centistinae of Capek

(1970) are included in the Euphorinae, according to Capek (1973). The Adeliini

s.l., a tribe of the Microgasterinae in Capek (1970), are redefined and partly

maintained as the tribus Miracini in the Microgasterinae; the other part is

treated as a separate subfamily, the Acaeliinae, according to Tobias (1967 and

1971, who used the name Adeliinae). The Orgilinae (the tribe Orgilini of the

Agathidinae in Capek, 1970) and the Braconinae (the tribe Braconini of the

Braconinae in Capek, l.c.) are treated as separate subfamilies. The three other

tribes (Exothecini, Hormiini and Pambolini) of the Braconinae in Capek are

included in the Rogadinae. The Spathiinae of Capek (1970) is at most a tribe

of the Doryctinae, as pointed out by Capek (1973: 267).

C. VAN ACHTERBERG: The subfamilies of the Braconidae 37

More problematic is the position of the genera Zele (and Zemiotes), Charmon

(= Eubadizon of Capek), Acampsis and Sigalphus. Capek (1970) included Zele

and Macrocentrus in his Macrocentrinae following, for example, Nixon (1938) and

Eady & Clark (1964). But Watanabe (1969: 319) considered the most adequate

location to be in a tribe Zelini of the subfamily Helconinae; also Capek (1973)

gave them a tribal rank. The larvae are distinct from the larvae of the Helconi-

nae: the labial sclerite is pentagonal, while it is transverse (Cenocoeliini) or

longitudinal (height larger than width in other tribes) in the Helconinae; they

are endoparasites of Lepidoptera, while Helconinae are almost exclusively

endoparasites of Coleoptera; and the imagines of Zele and its relatives have

several synapomorphous characters in respect to the Helconinae as treated in the

present paper. It is therefore that I follow Tobias (1967, 1971), who gave this

group subfamily rank, and I also include Charmon and Zemiotes.

The adult morphology of Charmon (e.g. the genitalia of the male, cf. Fig. 36, 40

in Tobias, 1967), its biology (also endoparasites of Lepidoptera) and the

regular shape of the emergence opening from the cocoon (irregular in the Orgili-

nae, its near relatives) indicate the relationship with Zele. The cephalic structures

of the final instar larvae of Charmon (Fig. 15 in Capek, 1970) are different from

Zele, because of the absence of the hypostomal spur, but the slender shape of

the hypostomal parts and of the stipital sclerite make a relationship with the

Zelinae more likely than with the Orgilinae (cf. Fig. 16, 17 in Capek, 1970) as

treated in this paper.

Still more complicated is the placement of Acampsis, Sigalphus and Meteoridea.

The larvae of Acampsis and Sigalphus have slender mandibles with a very wide

base (Fig. 34, 35 in Capek, 1970), unlike the basally slender mandibles of the

Cheloninae (Fig. 36, l.c.). Capek overlooked this difference, because he included

them in the Cheloninae (1970: 871), even in his key to the larvae (1973: 261)

where he mentioned as the key factor for the Cheloninae “. . . (mandibles) without

or only with a small base”. Thus owing to the shape of some cephalic structures

of the larvae and because of many differences in the adult morphology (e.g.,

nervellus broken, postpectal carina absent), I agree with Tobias (1967: 659),

who gave this group subfamily rank. Unfortunately Tobias (l.c.) used the name

“Sigalphinae”, a name for a long time (incorrectly) used for a group now included

in the tribus Brachistini of the Helconinae. Capek (1970: 871) erected the sub-

family Meteorideinae for Meteoridea, because the biology of this genus is rather

peculiar. It is (at least partly) comprised of gregarious endoparasites of Lepi-

doptera-larvae, which let pupate the host larva and thereafter the parasites spin

their cocoons inside the host cocoon. The stout stipital sclerite, the very wide base

of the mandible of the larvae with its apical half slender and toothed (cf. Fig. 12,

34, 35 in Capek, l.c.), combined with similarities in the morphology of the adults

(e.g., nervellus broken, rather short radial cell, first discoidal cell petiolate and

Ovipositor sheath wide) give some indications about its relationship with the

“Sigalphinae” of Tobias. Of the characters mentioned at least the slender apical

half of the mandibles of the larvae, the stout ovipositor sheath of the adults and

the rather short radial cell are synapomorphous. Therefore I propose to unite the

Meteorideinae of Capek and the Sigalphinae of Tobias into the Meteorideinae s.l.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

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40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

The division of the Braconidae given by Wesmael (1835), Foerster (1862),

Ashmead (1900), Szépligeti (1904), and Muesebeck & Walkley (1951) are

compared with the division given in the present paper in Table 1.

KEY TO THE SUBFAMILIES OF THE BRACONIDAE S.L.

1. Mandible unidentate (Fig. 10); antenna inserted on the top of the head

(Fig. 8, 9); ovipositor issuing near middle of metasoma (Fig. 8); whole body

densely setose (Fig. 8); associated with termites ...... Ypsistocerinae (p. 44)

— Mandible with 2—7 teeth; antenna inserted below top of head, at height of the

eyes (Fig. 5, 11, 25); ovipositor issuing near apex of metasoma (Fig. 11, 68, 72,

89, 95, 108, 109); body less setose; associated with other groups ........... 2

2. Mesoscutum protruding above pronotum (Fig. 11); epistomal (medially) and

scutellar sutures absent (Fig. 12, 15); ovipositor bent upwards (Fig. 11); clypeus.

straight medially, above an indistinct hypoclypeal depression (Fig. 12) ......

sy MAA I SRO ee a I RR aN PAR: Mesostoinae (p. 44)

— Mesoscutum not strongly protruding above pronotum, if exceptionally slightly

protruding, then ovipositor straight or bent downwards; epistomal and

clypeal sutures almost always complete; hypoclypeal depression absent, if

present, then clypeus more or less concave and differently shaped (Fig. 27,

IMI AEREE La NR O Ce OES Go 06 3

3. Hypoclypeal depression comparatively wide, deep and more or less round

dorsally (Fig. 27, 28, 35, 37, 41), exceptionally the face takes part in the

hypoclypeal depression (tribe Pseudodicrogeniini), resulting in a concave face

and a flat clypeus (Fig. 121, 122); metasoma rather often sculptured, some-

times its second tergite with a semi-circular suture (Fig. 38) .............. 4

— Hypoclypeal depression absent, if present (Fig. 5, 52), then more or less

shallow, narrower and straight dorsally or nearly so; face and clypeus more

or less convex, not concave (Fig. 85, 86); metasoma (if intermediate) often

smooth behind the first tergite (Fig. 63), its second tergite without semi-

circular suture, at most with a more or less chevron-shaped impression (Fig.

63) PORRO jones OI NI IO VIII no oc 8

4. Antenna situated at a protuberance, in front of the eyes (Fig. 97—99);

venation of wings reduced (Fig. 105); maxillary palp with 4 segments; para-

sites of adult Coleoptera ............ tribe Cosmophorini of the Euphorinae

— Antenna more or less situated between the eyes (Fig. 25, 27, 34, 42), without

distinctly developed protuberance; venation usually more complete (Fig. 19,

23, 31, 36, 43, 44); maxillary palp with 5 or 6 segments; parasites of larvae of

various groups Of INSECKS” Aln LE 5

5. First metasomal tergite strongly flattened basally and laterally (Fig. 22, 26);

dorsope absent or nearly so, if present, then dorsal carinae are situated later-

ally, above the spiracles; prepectal carina absent; occipital carina absent

dorsally; maxillary palpus with 5 segments, if with 6 segments (tribe Histero-

merini, Fig. 25) then hind femur strongly compressed, disk-shaped (cf. Fig. 20)

and fore tibia with a cluster of spines on two-third of its outer surface (Fig. 21);

10.

C. VAN ACHTERBERG: The subfamilies of the Braconidae 41

nervulus interstitial or nearly so (Fig. 23) or antefurcal (Fig. 19), very excep-

tienallypostfurcal(Fig--120) … Lu wrs corset 2m Lia Braconinae (p. 45)

First tergite not or slightly flattened, usually with dorsope and/or dorsal

carinae (Fig. 33, 38, 40, 46); dorsal carinae are removed from the spiracles;

occipital carina usually (partly) present dorsally; prepectal carina variable;

maxillary palp with 6 segments; hind femur not disk-shaped, at most weakly

compressed; fore tibia with a row of spines (Fig. 32) or without spines (Fig.

39); nervulus usually postfurcal (Fig. 31, 36, 43, 44) or absent, exceptionally

ater AS BEE EE 6

. First metasomal tergite immovably joined to the second tergite, without dorsal

carinae (Fig. 29, 30); three basal tergites about as long as half of metasoma ..

RT RS ER NE bla à Telengainae (p. 45)

First tergite flexibly joined to the second tergite, almost always with dorsal

carinae, at least basally (Fig. 40); if exceptionally the first tergite is immovably

joined to the second tergite, then the formed carapace is about as long as the

FIESCHI SIRO MORE E ROLO OO eased la RC spule eye ms OT fl

. Fore tibia with short, often rather thick spines (Fig. 32), if intermediate, then

occipital and prepectal carinae complete; hind coxa usually with an anterio-

ventral tubercle; if nervulus is present, then sides of first brachial cell parallel

(IE LG, a N RP EP er Doryctinae (p. 45)

Fore tibia without spines (Fig. 39); occipital carina often partly, or com-

pletely absent; prepectal carina variable; hind coxa without tubercle; if

intermediate then first brachial cell widened apicad (Fig. 43) and nervulus

E SER e ata man. Lera con ori à kin Rogadinae (p. 45)

. Mandibles with inner side out, their tips not touching when closed (Fig. 48,

50); mandibles usually with 3—7 teeth or lobes (Fig. 49, 51), seldom with

medial tooth large and both lateral teeth small ........... Alysiinae (p. 46)

Mandibles normal, their tips touching when closed (Fig. 5, 52, 85, 86, 90);

mandibles with two teeth, exceptionally with three teeth ................. 9

. Spiracles of first metasomal segment on its weakly sclerotized pleuron

(Fig. 56, 59); prepectal and occipital carinae completely absent ............

EREN O N E Rl Dae Microgasterinae (p. 46)

Spiracle of the first segment on its strongly sclerotized tergite (Fig. 61, 63,

77, 80, 82, 83); prepectal and occipital carinae variable ................. 10

Subdiscoidella present, often consisting of a more or less disconnected and

yellowish or brownish stripe; nervellus often weakly pigmented posteriorly

in respect to its anterior half (Fig. 65, 67, 73); if intermediate, then nervellus

Biken EME MOA) ip Aden. ts Adha ET libere hei lana BS 11

Subdiscoidella absent (Fig. 74, 75, 79, 81, 84, 92); nervellus not broken,

seldom its posterior half weakly pigmented in respect to the anterior half

IPAM ODOM) oer TE ete PT are. er 12

. First discoidal cell sessile or subsessile (Fig. 65); cu 1 often absent; occipital

carina completely absent; second tergite often with a chevron-shaped impres-

sion (Fig. 63); fore side of radial cel shorter than the pterostigma (Fig. 65) ...

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fe ait ale hI NEN SCORES OEY LR SO eee et E Agathidinae (p. 47)

First discoidal cell distinctly petiolate and cu 1 always present (Fig. 64, 67,

73); second tergite without chevron-shaped impression; fore side of radial

cell equal in length to the length of pterostigma or somewhat longer (Fig. 64,

67, 73); occipital carina present, at least laterally ..... Meteorideinae (p. 47)

. Postpectal and prepectal carinae completely present (Fig. 71); first metasomal

tergite immovably joined to the second tergite, forming with the third

tergite arigid'carapace (Fig. 72)) RR RR EIRE Cheloninae (p. 47)

Postpectal carina absent (Fig. 87, 91), exceptionally a ventral remnant present;

prepectal carina variable; first tergite usually movably joined to the second

tergite (Fig Wo) oO AR OS PE Re De, PO 13

. First metasomal tergite flattened laterally and concave medially, almost

invisible and immovably joined to the second tergite, forming with the third.

tergite a shield that covers slightly less than the basal half of the metasoma

(Fig. 61); cuqu | from the pterostigma, usually far from r 1 (Fig. 62); transverse

carina of propodeum strongly developed; occipital carina completely present

N eae wearer pny es a ARE PRE SEON Acaeliinae (p. 47)

First tergite distinctly convex (Fig. 53, 80, 83), if seldom somewhat flattened, :

then occipital carina absent; first tergite usually flexibly joined to the second

tergite; if the first tergite is immovably connected, then the three basal tergites

cover almost the complete length of the metasoma; cuqu | from the radius

(Fig. 75, 79, 118); transverse carina of propodeum less developed ........ 14

. Metasoma inserted above the hind coxae (Fig. 77, 78) .................. 15

Metasoma inserted between the hind coxae, at least partly (Fig. 87, 91,

103,106, 109) U ER ees EN VERE RA Fee Ea IAA 18

. Metasoma inserted far above hind coxae (Fig. 78); occipital carina present

dorsally: 42%: #3 PI ae eee tribe Cenocoeliini of the Helconinae

Metasoma inserted near hind coxae (Fig. 77); occipital carina absent dorsally

E ERI We Pr tah AIA CR EREN 16

. Trochantellus with apical teeth externally (Fig. 77); submediellan cell large

(Fig. 75, 81); middle lobe of mesoscutum more or less protruding (Fig. 76)

ha PO ia OI Ra I LIE Macrocentrinae (p. 48)

Trochantellus without apical teeth; submediellan cell small (cf. Fig. 23);

middle lobe of mesoscutum not protruding "MERE 17

Metapleural flange absent but with a rather wide and thin carina (cf. Fig.

91); radial cell comparatively wide (cf. Fig. 101); occipital carina present

laterally’. „2405 AEEA et Rs UN ET ARRE Orgilinae (p. 50)

Metapleural flange present (cf. Fig. 78, 87); radial cell very narrow (cf. Fig.

65); occipitalicarimajabsent) at Re RR re Agathidinae (p. 47)

. Maxillary palp short, with 2 or 3 segments (Fig. 3, 5); first brachial cell almost

square (Fig. 4) or rqu present (Fig. 7); parasites of ants ................. 19

Maxillary palp usually longer, with 4—6 segments; first brachial cell elongate

and rqu absent;’parasites of other groups ee 20

. First brachial cell almost square (Fig. 4); fore basitarsus very slender (Fig. 2);

20.

21.

DO.

23,

24.

25:

C. VAN ACHTERBERG: The subfamilies of the Braconidae 43

clypeus very narrow, longer than wide (Fig. 3); ovipositor straight; malar

Spacsiconeave posterioriy til Ai. aren MG oe Hybrizontinae (p. 48)

First brachial cell elongate (Fig. 7); fore basitarsus stout (Fig. 6); clypeus

wide, wider than long (Fig. 5); ovipositor strongly curved ventrad; malar

space without concavity posteriorly ................. Neoneurinae (p. 49)

Nervellus absent and radial cell (so far present) moderately long (Fig. 93,

96); first brachial cell comparatively narrow (Fig. 93, 96); metasomal tergites

weakly sclerotized (Fig. 95); first metasomal tergite usually slightly or not

widened apicad; clypeus comparatively small (Fig. 94); pleural suture weakly

developed; occipital carina present, at least laterally; parasites of aphids

en TUTI ORA, STO AS Aphidiinae (p. 49)

Nervellus present, if indistinctly developed or absent, then radial cell very

short (Fig. 118), first brachial cell and clypeus wider; tergites distinctly

sclerotized; first tergite and occipital carina variable; pleural suture usually

widgland'crenulate; parasites of other groups !.....2..... en. 21

Basal vein strongly curved at its anterior end (Fig. 88); occipital carina com-

LEE AREA oi che ii N ee à Ichneutinae (p. 49)

Basal vein scarcely or not curved anteriorly (Fig. 92); occipital carina usually

PES EI SRI ee TRIES PR NEE IE 22

First tergite of metasoma flattened basally and laterally, without dorsal

carinae (Fig. 82); occipital carina absent; anterior tentorial pits very large

(Fig. 90); radial cell short and comparatively high (Fig. 92); basal vein more

HS Een bent: parasifes'ofsaw-flles n. she dateren aten mon an sn

en ern hr ne EN tribe Proteropini of the Ichneutinae (p. 49)

First tergite scarcely or not flattened (Fig. 53, 83); occipital carina and/or

dorsal carinae present; anterior tentorial pits much smaller (Fig. 52, 115);

radial cell longer, if short, then narrow (Fig. 118); parasites of other groups

nt ARE RI RA SIR ARTE, E I RN RESINE E ERA ue is 23

Prepectal carina absent and metasoma short, oval (Fig. 53), its first tergite

flexibly joined to the second tergite; hypoclypeal depression present or absent;

notauli and precoxal suture more or less reduced; parasites of Diptera .....

rade leo venin Siani ze Le Opiinae (p. 46)

Prepectal carina almost always present; metasoma usually more slender

(Fig. 111—113), sometimes tergite immovably joined to the second tergite;

notauli and precoxal suture usually less reduced; almost always parasites of

BEBEREEBUNS(O. citi hanstes beneeglial id adik ask) aioe seo;

First metasomal tergite petiolate (Fig. 111, 113), sometimes petiolus rather

short (Fig. 112) or length more than 3.2 times its apical width (Fig. 114, 116);

spiracle of first tergite usually situated medially or behind the middle of the

E a la dii ici ae 25

First tergite sessile (Fig. 80, 83), shorter than 3.2 times its apical width;

spiracle usually situated in front of the middle of first tergite (Fig. 80, 83)

io net ie outa ror ammel Lis ten til LIL) ell 26

Radiellan cell widened distad (Fig. 107), sometimes with an interradiella;

whole surface of 4th and Sth tergites of metasoma densely setose (Fig. 111);

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

GUqui2ipresenti HN Ble NCIS ENNE tribe Zemiotini of the Zelinae (p. 50)

— Radiellan cell narrowed or scarcely widened distad, without interradiella (Fig.

117—119); if cuqu 2 present, then 4th and Sth tergites with one row of setae

pertergite (Fig. 112,113). rn en peewee tld: "Euphorinae (p. 50)

26. Laterope deep (Fig. 109) and cuqu 2 absent, if intermediate, then ovipositor

sheats wide, shorter than three times its maximum width (Fig. 109) .......

EN EE A Er tribe Centistini of the Euphorinae (p. 50)

— Laterope shallow (Fig. 87) or absent, if present then cuqu 2 present and/or

ovipositor sheaths very slender, much longer than three times its maximal

Widths siii ee en RENEE en CRC 27

27. Dorsal carinae of first tergite present, at least basally (Fig. 80, 83); if cuqu

2 absent and dorsal carinae of first tergite weakly developed, then dorsal

surface of propodeum distinctly shorter than its posterior surface (Fig. 87) and

radial cell comparatively wide (Fig. 84); mainly parasites of larvae of

Coleoptera’: cna ae ae ee een Helconinae (p. 48)

— Dorsal carinae absent (Fig. 91, 103, 106), if present then cuqu 2 absent, dorsal

surface of propodeum scarcely separated from its posterior surface and of

more or less equal length (Fig. 91, 103, 106) and radial cell narrow (Fig.

[01):;parasites of Lepidoptera-larvaen RO eae 28:

28. Anal lobe of hind wing distinctly developed and usually with a more or less

developed aqu’(Fig. 100, 104), if aqu’ absent, then metapleuron with a thin,

more or less protruding carina (Fig. 103); distal abscissa of radius more or

lessicurved (Fig: 1004104) ER Re ren Zelinae (p. 50)

— Anal lobe of hind wing comparatively narrow, without aqu’ (Fig. 101, 102);

if anal lobe is intermediate, then metapleuron without a thin carina, at

most with a small flange (Fig. 106); distal abscissa of radius straight (Fig. 101,

ID e ee Orgilinae (p. 50)

SHORT DESCRIPTIONS OF THE SUBFAMILIES

Ypsistocerinae (Fig. 8—10)

Small subfamily, which contains the two genera Ypsistocerus Cushman and

Termitobracon Brues. They live probably as parasites in the nests of termites in

the Neotropical region. This group possesses many apomorphous characters, e.g.,

the unidentate mandibles (Fig. 10); the far retracted hypopygium (Fig. 8); the

densely setose body (Fig. 8); the highly inserted antenna (Fig. 9) and the strongly

reduced palpi and eyes (Fig. 9, 10).

Mesostoinae (Fig. 11—18)

Small subfamily, containing only the genus Mesostoa Van Achterberg from

the Australian region, of which the biology is unknown. As pointed out by Van

Achterberg (1975: 158) almost all characters are apomorphous, e.g., the absence of

the occipital and prepectal carinae (Fig. 11); the absence of the precoxal and

scutellar sutures (Fig. 11, 15); the flattened first metasomal tergite without carinae

C. VAN ACHTERBERG: The subfamilies of the Braconidae 45

(Fig. 14); the smooth propodeum; the concave frons (Fig. 18); the upcurved

ovipositor and the compressed legs (Fig. 16).

Braconinae (Fig. 19—27; 120—122)

Large subfamily, consisting of solitary or gregarious ectoparasites of larvae of

holometabolous insects. The less evolved species often parasitize Coleoptera, the

more evolved species also Lepidoptera, Hymenoptera-Symphyta and Diptera.

According to Capek (1970: 862) the host larva is paralysed at egg-deposition,

the parasite-larva feeds on the paralysed host and forms its delicate cocoon at a

sheltered place. The cephalic structures of the larvae are remarkably homo-

geneous. For the most parts none is reduced except for the mandibles, as in the

main part of the Rogadinae as defined in the present paper. Therefore Capek

enlarged the concept of the Braconinae, in my opinion incorrectly because it is

based on an aggregate of plesiomorphous characters. It only indicates that this

group is less evolved than could be expected from the morphology of the adults.

The adults have several apomorphous characters, e.g., the flattened first tergite

(Fig. 22, 26); the reduction of the occipital and prepectal carinae and the maxillary

palp consisting of 5 segments (except for the Histeromerini).

Telengainae (Fig. 28— 30)

Small subfamily containing only the Palaearctic genus Telengaia Tobias. The

biology is unknown. The shape of the metasoma is peculiar (Fig. 29, 30), but I

am not sure about its position; it may be only a member of the Rogadinae.

Doryctinae (Fig. 31—36)

Rather large subfamily, consisting of ectoparasites of larvae of (wood-boring

and bark-mining) Coleoptera. Less common also other hosts in plant tissues are

attacked. The host-larva is paralysed before egg-deposition. The cephalic

structures of the larvae are similar to those of the Braconinae. The morphology

of the adults and their biology indicate their comparatively close relationship

to the ancestral stem of the Braconidae. Some of the few apomorphous characters

are the spines of the fore tibia (Fig. 32); the often large hypoclypeal depression

(Fig. 35) and the usually distinctly developed dorsope (Fig. 33).

Rogadinae (Fig. 37—44, 46)

As treated in the present paper a large group and rather heterogeneous

because in this group the transition to endoparasitism has taken place. This is

accompanied with some pecularities as the tooth-less mandibles of the larvae (but

already in the Braconinae occur larvae with only a few teeth on their mandibles)

and the mummification of the host-caterpillar. In at least one genus closely related

to Rogas Nees (viz., Bucculatriplex Viereck) the pupation takes place in the host-

pupa in stead of in the mummified host-larva. Some species are solitary or

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

gregarious ectoparasites of larvae of Lepidoptera (Hormius Nees and its relatives;

also Oncophanes Foerster, but O. campsolechiae Watanabe (probably belonging to

Clinocentrus Haliday) is reported to be a gregarious endoparasite of larvae of Lepi-

dopterous leaf-rollers; their woolly cocoon is formed outside the host). Others are

solitary ectoparasites of Hymenoptera-Symphyta (Phanomeris Foerster s.l.) or

solitary endoparasites of mining larvae of holometabolous insects (Colastes

Haliday), while Rogas Nees and its allied genera are mainly solitary or gregarious

endoparasites of larvae of Lepidoptera, which pupate within the mummified larva

or in the pupa. Some apomorphous characters are the often deep hypoclypeal

depression; the reduction of the occipital carina in several genera; the endopara-

sitism of the main part of the subfamily and the often distinctly developed dorsope

of the first metasomal tergite.

Alysiinae (Fig. 45, 47—51)

One of the largest subfamilies of the Braconidae of which the larval cephalic

structures, e.g., the simple and smooth mandibles, resemble those of the larvae of

the Opiinae. Almost all species are solitary endoparasites of larvae of Diptera, but

some are gregarious (Aphaereta Foerster). The egg is usually deposited in the larva,

seldom in the egg (Polemochartus Schulz) or probably sometimes in the (pre-)pupa

(Aphaereta Foerster). The delicate cocoon is made in the puparium of the host.

The most striking apomorphous character of this group is the rotation of the

mandibles of the adults in a way that the outer side becomes the inner side.

Other apomorphous characters are the absence of the prepectal and the occipital

carinae, together with the development of additional teeth on the mandible and of

dorsope.

Opiinae (Fig. 52, 53)

Rather large subfamily, consisting of endoparasites of larvae of Diptera;

pupation in the puparium of the host. Ciosely related to the Alysiinae but the

mandibles are normally attached (but more or less twisted) and the occipital

carina is almost always present laterally. Some of its apomorphous characters

are the smooth mandibles of the final instar larvae, the absence of the prepectal

carina, the comparatively short and stout metasoma, the more or less reduced

notauli, precoxal and pleural suturae and the tendency to develop a hypoclypeal

depression.

Microgasterinae (Fig. 54—60)

Large subfamily, consisting of endoparasites of larvae of Lepidoptera. Often

gregarious; the eggs are deposited in the egg of the host or in the early instar

larvae. In the Microgasterini and Cardiochilini the larvae pupate outside the host;

the gregarious species often spin together in a common web. In the Miracini

(which are endoparasites of leaf-mining Lepidoptera) the larvae pupate in the

host-cocoon. The wing venation is often strongly reduced (Microgasterini, Mira-

C. VAN ACHTERBERG: The subfamilies of the Braconidae 47

cini), but the most important apomorphous characters are the position of the

spiracle of the first metasomal segment and the absence of the prepectal and of

the occipital carinae.

Agathidinae (Fig. 64—66)

Rather large subfamily containing solitary or gregarious endoparasites of larvae

of Lepidoptera. The egg is deposited in the early instar larva, the parasites pupate

outside the host larva. The presence of a more or less distinct subdiscoidella

(Fig. 65) and the sometimes comparatively high insertion of the metasoma are

remarkable, but are probably old plesiomorphous characters; also the larvae are in

several aspects (e.g., the shape of the mandibles) rather slightly derived. Some

apomorphous characters are the small second cubital cell (or cuqu 2 absent),

posterior half of nervellus more or less weakly pigmented; the absence of the

occipital carina; the short radial cell (Fig. 65); and the second metasomal tergite

often has a more or less chevron-shaped impression (Fig. 63).

Meteorideinae (Fig. 63, 67, 68, 70, 73)

Small subfamily, solitary or gregarious endoparasites of larvae of Lepidoptera.

According to Capek (1970: 859) Acampsis Wesmael and Sigalphus Latreille may

deposite their eggs in the egg of the host and pupate outside the host larva,

while Meteoridea Ashmead pupates inside their host pupa. This group shows many

plesiomorphous characters in both the larval and the adult morphology. Some of

the few apomorphous characters are its endoparasitism, the comparatively wide

ovipositor sheaths, the more or less developed dorsal carinae and deep laterope of

the first metasomal tergite.

Cheloninae (Fig. 69, 71, 72, 74)

Rather large subfamily, which consists of solitary endoparasites of larvae of

Lepidoptera. The egg is deposited in the egg of the host. The delicate cocoon is

attached to the remains of the host larva. This group possesses many apomorphous

characters, e.g., the metasomal carapace, the short and high radial cell, the

presence of the postpectal carina, the often densely setose eyes, the basally slender

mandibles of the final instar larvae and the often indistinct hypostomal suture.

The deposition of the egg in the host egg is clearly an adaptation to the habits of

its hosts, which live at more or less hidden places.

Acaeliinae (Fig. 61, 62)

Small subfamily, its members closely resemble the Miracini of the Microgasteri-

nae (cf. Fig. 62 with Fig. 55). They share also their hosts, viz., both are endo-

parasites of larvae of leaf-mining Lepidoptera, also the pupation is in the cocoon

of the host; this may account for their resemblance. The larvae are distinguishable

from the larvae of the Microgasterinae by the presence of a mandibular base (cf.

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 27, 28 in Capek, 1970). The flattened shield formed by the three basal tergites

of the metasoma is peculiar, they are fused almost invisibly (Fig. 61). Some

other apomorphous characters are the strongly developed transverse carina of the

propodeum, the position of the cuqu 1 and r 1 and the reduction of several other

veins, e.g. the nervellus (Fig. 62).

Macrocentrinae (Fig. 75—77, 81, 86)

Subfamily of moderate size, containing solitary or gregarious endoparasites of

larvae of Lepidoptera. The parasite pupates outside the host. Polyembryony is

known of the gregarious species, of which the specimens spin together in a

common web. The most strikingly apomorphous character of this group is the

toothed trochantellus (Fig. 77), others are the specialized depressed area at the

second tergite laterally; the high attachment of the metasoma, the reduction of the’

occipital carina, the more or less protruding middle lobe of mesoscutum (Fig. 76)

and the usually absent dorsal carinae of the first metasomal tergite (Fig. 77).

Hybrizontinae (Fig. 1—4)

Small subfamily with one genus, Hybrizon Fallen, which contains endoparasites

of larvae of several ant-genera. The naked pupae remain in the host nest. Pos-

sesses many apomorphous characters, e.g., the short palpi (Fig. 3), the concavity

behind the malar space, the slender legs, the peculiar venation of the wings

(Fig. 4), the slender clypeus and probably completely grown together 2nd and

3rd tergites (Fig. 1).

Helconinae (Fig. 78—80, 83—85, 87)

Large subfamily of which almost all species are endoparasites of larvae of

Coleoptera. The few exceptions may be Blacus Nees (some derived species have .

been bred from larvae of Diptera) and Dyscoletes Haliday (bred from larvae of

Mecoptera). This subfamily is difficult to characterize by apomorphous charac-

ters, except for the endoparasitism. Formerly this name often served for a hetero-

geneous group, clearly used as a rest group of remaining genera. Actually the.

group of genera included in this paper can be characterized by several more or less

“reticulate”’ occurring apomorphous characters. The most important apomor-

phous characters in the Cenocoeliini are the highly inserted metasoma (Fig. 78),

the concave frons, as occurs also in the Helconini and less distinctly in the

Brachistini. The Helconini (Fig. 83) have usually the dorsal carinae distinctly

developed, often reaching behind the middle of the first tergite as in the Blacini

(Fig. 80) and Brachistini. A dorsope is present in the Blacini (Fig. 80) and Ceno-

coeliini (Fig. 78), and less commonly in the Helconini (Fig. 83). The first discoidal

cell is (sub-)sessile in the Helconini (p.p.), Brachistini (Fig. 84) and Blacini (but

seldom shortly petiolate). The reduction of the veins aqu 1 + 2 and aqu’ occurs

in the Helconini, Brachistini and Blacini, as also the reduction of the sculpture on

the metasoma and of the precoxal suture. The reduction of veins leads to the loss

C. van ACHTERBERG: The subfamilies of the Braconidae 49

of the cuqu 2 in the Brachistini (Fig. 84) and Blacini. In this subfamily the tribe

Helconini is clearly less derived than other tribes; it possesses many plesio-

morphous character-states. The final instar larvae have also many plesiomorphous

characters (Fig. 9 in Capek, 1970), which supports the hypothesis about the

evolution of the Braconidae from exoparasites of sheltered living larvae of

Coleoptera.

Ichneutinae (Fig. 82, 85, 88—90, 92)

Rather small subfamily, consisting of endoparasites of larvae of sawflies

(Ichneutini, Proteropini) and of lepidopterous leafminers (Muesebeckiini). The

egg is deposited in the egg of the host but the larva developes after pupation of

the host. The cocoon is formed inside the cocoon of the host. Rather hetero-

geneous group, e.g., the final instar larvae of the Muesebeckiini have (at least in

one genus) slender and smooth mandibles, stipital sclerite robust and hypostoma

absent, while the Ichneutini have robust and toothed mandibles (as the Pro-

teropini) and the cephalic sclerites present and robust (Fig. 33 in Capek, 1970).

The cephalic sclerites are also present in the Proteropini but are very slender

(Fig. 32, l.c.). Also the imagines differ considerably, but with the information

available it seems better to unite the three tribes in one subfamily. The most

important apomorphous characters are the absence of the occipital carina, and

also more or less of the prepectal carina in the Muesebeckiini and Proteropini;

the short and comparatively high radial cell in the Ichneutini and Proteropini

(Fig. 88, 92) (which resembles the Cheloninae), the short ovipositor with its more

or less widened sheaths (Fig. 89), the more or less strongly curved basal vein

and the absence of aqu’ (Fig. 88, 92).

Neoneurinae (Fig. 5— 7)

Small subfamily, consisting of endoparasites of adults of worker ants; the

development is in the gaster of the host. The cocoon is attached to the remains

of the host. The most important apomorphous characters are the short palpi

(Fig. 5), the peculiar venation of the wings (Fig. 7), the concave coxae dorso-

apically and the strongly bent ovipositor.

Aphidiinae (Fig. 93—96)

Rather large subfamily, containing solitary endoparasites of aphid nymphs and

adults. Praon Haliday and Dyscritulus Hincks emerge from their host and spin

their cocoon directly beneath the aphid, whereas all other genera pupate within

the mummified host skin, mounted on the surface of the plant. The cephalic

structures of the larvae are like those of other Braconidae, according to Capek

(1970, p. 848). The apomorphous characters of this subfamily are especially

distinct in the larvae, because of the long and funnel-shaped spiracles without a

division into atrium and closing apparatus, of the simple and smooth mandibles

50 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

and of the disk-shaped antennae. The apomorphous characters of the adults are

the reduction of the wing venation (Fig. 93, 96), the specialized female genitalia

and the weakly sclerotized metasoma (Fig. 95).

Zelinae (Fig. 91, 100, 103, 104, 107, 111)

Small subfamily, which contains solitary endoparasites of larvae of Lepidoptera.

The pupation is outside the host larva. The apomorphous character of the larvae

in respect to the Orgilinae is the slender stipital sclerite; Zemiotes Foerster has also

smooth mandibles and Zele Curtis has the hypostomal spur absent. The adults

have few apomorphous characters, e.g., the first metasomal tergite is slender and

without dorsal carinae (Fig. 103), even petiolate in Zemiotes Foerster (Fig. 111).

Orgilinae (Fig. 101, 102, 106)

Small subfamily containing species with often many apomorphous characters.

They are endoparasites of larvae of Lepidoptera, which pupate outside the host

larva; the emergence opening of the cocoon is irregular. The larvae are character-

ized by the absence of the hypostoma (Fig. 16, 17 in Capek, 1970). Apomorphous

characters of the adult parasites are the absence of the aqu’ (Fig. 10}, 102); the

more or less reduced metapleural flange (Fig. 106) and occipital carina dorsally;

the more or less small radial cell of fore wing and anal lobe of the hind wing (Fig.

101, 102); cuqu 2 absent (Fig. 101) or if present, forming a small second cubital cell

(Fig. 102).

Euphorinae (Fig. 97, 99, 105, 108—110, 112—119)

Large and diverse subfamily, consisting of solitary or gregarious endoparasites

of larvae of Lepidoptera and Coleoptera (Meteorus Haliday), of solitary or

gregarious endoparasites of adult Coleoptera (and more seldom of their larvae),

(e.g., Perilitus Nees s.l., Ropalophorus Haliday in Curtis, Cryptoxilos Viereck, Stre-

blocera Westwood), of adult parastic and aculeate Hymenoptera (Syntretus Foers-

ter, the only exception to the rule that Braconidae are primary parasites), of adult

Neuroptera (Chrysopophthorus Goidanich), of nymphal and adult Heteroptera

(Wesmaelia Foerster, Aridelus Marshall, Leiophron Nees, Holdawayella Loan) and

of nymphal and adult Psocoptera (Leiophron Nees). In Meteorus Haliday the

cocoon of some species hangs from a long thread. The most important

apomorphous character of the larvae are the smooth and short mandibles. The

apomorphous characters of the adults are rather “‘reticulate”’ as in the Helconinae,

e.g., the wing venation is reduced in the Cosmophorini (Fig. 105), the Euphorini

(Fig. 118, 117, 119) and to a lesser degree in the Centistini, the parasitism of adult

insects also occurs in these tribes; a dorsope occurs in the Centistini (p.p.),

Euphorini (p.p.) (Fig. 110) and in the Meteorini (p.p.) (Fig. 112, 113); the notauli

and the precoxal suture are reduced in the Centistini and in the Euphorini; the

comparatively wide ovipositor sheaths, together with a more or less robust and

bent ovipositor in the Centistini (Fig. 109) and Euphorini (p.p.; Fig. 108), and the

C. VAN ACHTERBERG: The subfamilies of the Braconidae SI

spiracles of the first metasomal tergite are situated in the middle of the tergite or

behind the middle in the Euphorini (Fig. 110, 114, 116) and in the Meteorini (p.p.;

Fig. 112, 113).

The possible relations between the subfamilies are depicted in Fig. 123. The

following groups may be recognized:

Group A: The old “‘cyclostomes” of Wesmael, together with the later formed

subfamilies Telengainae and Mesostoinae. They share the hypoclypeal depression,

the apically more or less concave clypeus (Fig. 27, 28, 35, 41, 12, 37, 121) and the

more or less flattened first metasomal tergite (Fig. 14, 22, 26, 30, 33, 38, 40).

Group B: Specialized endoparasites of larvae of Diptera with the pupation in

the host-puparium. The larvae have smooth, sickle-shaped mandibles and the

labial sclerite absent or at least broadly interrupted ventrally. The adults have the

prepectal carina absent and a more or less oval metasoma (Fig. 53).

Group C: Specialized endoparasites of larvae of Lepidoptera; the egg is deposit-

ed in the eggs of the host or in the early instar larvae. The larvae of the Acaeliinae

and of the Microgasterinae-Cardiochilini have the tips of the mandibles bifid; the

larvae of the Microgasterinae and of the Cheloninae have mandibles without or

only with a small, scarcely differentiated base. The adults often have the eyes

densely setose, the radial vein is often shortened (Fig. 54, 55, 57, 58, 62, 74) and

the first metasomal tergite is more or less flattened in the Microgasterinae and

Acaeliinae (Fig. 56—61).

Group D: Specialized endoparasites of larvae of Lepidoptera; the larvae share

the robust mandibles with a long, toothed blade (but the blade is intermediate

in Meteoridea Ashmead). The imagines have the radial cell rather small (Fig. 65,

67, 73) and the laterope deep and usually large (Fig. 68, 70).

Group E: Specialized endoparasites of larvae of Lepidoptera; the larvae have

the labial sclerite transverse and the epistomal arch and hypostoma are absent.

The adults share the tendency to loose the dorsal carinae of the first metasomal

tergite.

Group F: Generally very specialized endoparsites, especially of adult insects.

The larvae have toothless, more or less wedge-shaped mandibles. The adults share

the tendency to have the spiracle of the first metasomal tergite situated near the

middle (Fig. 95, 109, 110, 112—114, 116); the wing venation is often very

specialized (Fig. 7, 93, 96, 105, 117—119).

The position of the Hybrizontinae within the Braconidae is rather uncertain, the

situation of the spiracle near the middle of the tergite (Fig. 1) it shares with group

F, but this may be a convergential development.

The Ichneutinae, Macrocentrinae, and Helconinae form separate groups on

their own. The larvae of the Ichneutinae show some similarity with the larvae in

Group D, but the mandibles have a more or less developed triangular base. The

adults also show some similarity, e.g., the reduction of the wing venation and the

flattened first metasomal tergite in several groups.

The larvae of the Macrocentrinae have a transverse labial sclerite with two

processes ventrally, and polyembryony occurs in the genus Macrocentrus Curtis.

The adults differ from the Helconinae by the shape of the trochantellus (Fig. 77),

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

of the first and second metasomal tergites of the mesosoma (Fig. 76), and of the

head (Fig. 86).

Finally the Helconinae; both in biology and morphology a slightly derived

group. The larvae have wedge-shaped and toothed mandibles with a more or less

distinctly differentiated base and the adults show the tendency to develop long

dorsal carinae and distinct dorsope (Fig. 78, 80, 83).

ACKNOWLEDGEMENTS

I wish to express my sincere thanks to Dr. J. Decelle (Tervuren) and Dr. J. Papp

(Budapest) for the loan of types and to Mr. T. Huddleston (London) for the

correction of the English text.

LITERATURE

Achterberg, C. van, 1975. A new genus, Mesostoa gen. nov., from W. Australia, belonging to a new sub-

family (Hym., Braconidae). — Ent. Ber., Amst. 35: 158-160, Fig. 1-8.

———, 1976. A revision of the tribus Blacini (Hym., Braconidae, Helconinae). — Tijdschr. Ent. 118

(7): 159-322, Fig. 1-476.

Ashmead, W. H., 1900. Classification of the Ichneumon flies of the superfamily Ichneumonoidea. —

Proc. U.S. nat. Mus. 23: 1-220.

Capek, M., 1970. A new classification of the Braconidae (Hym.) based on the cephalic structures of the

final instar larvae and biological evidence. — Can. Ent. 102 (7): 846-875, Fig. 1-58.

———, 1973. Key to the final instar larvae of the Braconidae (Hym.). — Acta Inst. forest. zvol.: 259-

268, | Fig.

Cushman, R. A., 1923. A new subfamily of Braconidae (Hym.) from termite nests. — Proc. ent. Soc.

Wash. 25 (2): 54-56, Fig. 1-3.

Eady, R. D., & J. A. J. Clark, 1964. A revision of the genus Macrocentrus Curtis (Hym., Braconidae) in

Europa, with description of four new species. — Entomologist’s Gaz. 15: 97-127, Fig. 1-92.

Fahringer, J., 1925-28. Opuscula braconologica, Palaearktischen Region, 1: 1-606, Fig. 1-12, | table.

———, 1936. Uber einige merkwirdige und seltene Hymenopteren-Gattungen aus Afrika. — Fest-

schrift 60. Geburtstage Prof. Dr. Embrik Strand 1: 568-590 + Fig. 1-27.

Foerster, A., 1862. Synopsis der Familien und Gattungen der Braconen. — Verh. naturh. Ver. preuss.

Rheinl. 19: 224-288, 1 Fig.

Gauld, I. D., & T. Huddleston, 1976. The nocturnal Ichneumonoidea of the British Isles, including a

key to genera. — Entomologist’s Gaz. 27: 35-49, Fig. 1-20.

Griffiths, G. C. D., 1964. The Alysiinae (Hym., Braconidae) parasites of the Agromyzidae (Dipt.). |.

General questions of taxonomy, biology, and evolution. — Beitr. Ent. 14: 823-914, Fig. 1-38, 2

graphs, 2 tables.

Hedgvist, K.-J., 1955. Studien über Braconiden. I. — Ent. Tidskr. 76 (2-4): 92-98, Fig. 1-2.

Marsh, P. M., 1963. A key to the Nearctic subfamilies of the family Braconidae (Hym.). — Ann. ent.

Soc. Am. 56: 522-527, Fig. 1-29.

———, 1971. Keys to the Nearctic genera of the families Braconidae, Aphidiidae and Hybrizontidae

(Hym.). — Ann. ent. Soc. Am. 64: 841-850, Fig. 1-11.

Marshall, T. A., 1885. Monograph of British Braconidae, 1. — Trans. ent. Soc. London (1): 1-280 + 6

plates.

———, 1891. Les Braconides. In: André, E. (ed.), 1891-96. Species des Hyménoptères d'Europe et

d’Algérie, 5: 1-628 + 19 plates.

Mason, W. R. M., 1973. Recognition of Zemiotes (Hym., Braconidae). — Proc. ent. Soc. Wash. 75 (2):

213-215.

Muesebeck, C. F. W., & L. M. Walkley, 1951. Family Braconidae. In: C. F. W. Muesebeck et al. (ed.),

Hymenoptera of America north of Mexico, Synoptic catalogue. — Agriculture Monogr. 2: 90-

184.

C. VAN ACHTERBERG: The subfamilies of the Braconidae 53

Nixon, G. E. J., 1938. Notes on the taxonomy and synonymy of Zele Curtis and Macrocentrus Curtis

(Hymenoptera, Braconidae). — Bull. ent. Res. 29: 415-424, Fig. 1-2.

Shenefelt, R. D., & C. F. W. Muesebeck, 1957. Ashmead’s Meteoridea (Hym., Braconidae). — Proc.

ent. Soc. Wash. 59 (3): 129-134, Fig. 1-4.

Szépligeti, G. V., 1904. Hymenoptera, Fam. Braconidae. In: Wytsman, P. (ed.), 1902-32. Genera Insec-

torum, 22: 1-253 + 32 Fig.

Tobias, V. I., 1962. A new subfamily of braconids (Hym., Braconidae) from Middle Asia. — Trudy zool.

Inst. Leningr. 30: 268-270, Fig. 1-4. (Russian)

———, 1967. A review of the classification, phylogeny and evolution of the family Braconidae (Hym.).

— Ent. Obozr. 46 (3): 645-669, Fig. 1-43. (Russian)

———, 1971. Review of the Braconidae (Hym.) of the USSR. — Trudy zool. Inst. Leningr. 54: 156-

268, Fig. 1-112. (Russian)

Tobias, V. I., & G. P. Dudarenko, 1974. General trends in evolution of Braconidae (Hym.) abdomen.

— Vest. Zool. 1974 (3): 65-72, Fig. 1-2. (Russian)

Watanabe, C., 1937. A contribution to the knowledge of the braconid fauna of the Empire of Japan. —

J. Fac. Agric. Hokkaido Univ. 42 (1): 1-188 + i-iv + Fig. 1-15.

———, 1968. Notes on the genus Cosmophorus and Orgilus in Japan with description of a new species

(Hym., Braconidae). — Insecta matsum. 31: 1-6 + Fig. 1-11.

———, 1969. Notes on the genera Zele Curtis and Xiphozele Cameron with special reference to the

species in Japan (Hym., Braconidae). — Proc. ent. Soc. Wash. 71 (3): 318-328, Fig. 1-9.

Wesmael, C., 1835. Monographie des Braconides de Belgique. — Nouv. Mém. Acad. sci. R. Bruxelles

9: 1-252 + 1 plate + 2 tables.

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 1—4, Hybrizon buccata (de Brébisson), 9, Netherlands, Meijendel. 1, basal half of metasoma,

lateral aspect; 2, fore leg, lateral aspect; 3, head, frontal aspect; 4, wings. Fig. 1, 3: 2.1 times scaleline;

Fig. 2, 4: scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 55

1.0 mm

Fig. 5—7, Neoneurus auctus (Thomson), 9, Lappland, Enontekiò. 5, head, frontal aspect; 6, fore leg,

lateral aspect; 7, wings. Fig. 8, Ypsistocerus manni Cushman, 9; habitus, lateral aspect. Fig. 9, 10,

Ypsistocerus vestigialis Cushman. 9, head, frontal aspect; 10, mouth parts, ventral aspect. Fig. 5: scale-

line; Fig. 6, 7: 1.2 times scale-line; Fig. 8: 0,5 times scale-line. Fig. 8-10 after Cushman, 1923

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

FRONTAL ASPECT

LATERAL ASPECT

Fig. 11—18, Mesostoa compressa Van Achterberg, 9, holotype. 11, habitus, lateral aspect; 12, head,

frontal aspect; 13, wings; 14, abdomen, dorsal aspect; 15, mesonotum, dorsal aspect; 16, hind leg,

lateral aspect; 17, antenna, frontal aspect; 18, head, dorsal aspect. Fig. 11, 13, 16, 17: scale-line; Fig.

12, 14, 15, 18: 1.5 times scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae SY

Fig. 19-21, 24, Histeromerus mystacinus Wesmael, 9, Netherlands, Asperen. 19, wings; 20, fore leg,

anterio-lateral aspect; 21, detail of fore tibia, posterio-lateral aspect; 24, hind tarsus, lateral aspect.

Fig. 22, 23, Bracon urinator Fabricius, 9, Switzerland, Saas-Fee. 22, first metasomal tergite, dorsal

aspect; 23, wings. Fig. 19: scale-line; Fig. 20, 24: 1.2 times scale-line; Fig. 21, 22: 2.5 times scale-line;

Fig. 23: 0.6 times scale-line

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

1.0 mm

Fig. 25, 26, Histeromerus mystacinus Wesmael, 9, Netherlands, Asperen. 25, head, lateral aspect; 26,

propodeum, first and second metasomal tergites, dorsal aspect. Fig. 27, Bracon urinator Fabricius, 9,

Switzerland, Saas-Fee; head, frontal aspect. Fig. 28-30, Telengaia ventralis Tobias. 28, head, frontal

aspect; 29, metasoma, lateral aspect; 30, metasoma, dorsal aspect. Fig. 25-27: scale-line; Fig. 28, 30:

after Tobias, 1962; Fig. 29: after Tobias & Dudarenko, 1974

C. VAN ACHTERBERG: The subfamilies of the Braconidae 59

Fig. 31-33, Doryctodes imperator (Haliday), © , Netherlands, Cadier. 31, wings; 32, fore tibia, anterio-

lateral aspect; 33, first and second metasomal tergites, dorsal aspect. Fig. 34-36, Dendrosoter

protuberans (Nees), 9, Netherlands, Best. 34, head, dorsal aspect; 35, head, frontal aspect; 36, wings.

Fig. 31, 33: 0.6 times scale-line; Fig. 32: 2.5 times scale-line; Fig. 34-36: scale-line

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 37, 38, Gnaptodon pumilio (Nees), 9, Netherlands, Waarder. 37, head, frontal aspect; 38, meta-

soma, dorsal aspect. Fig. 39, 40, 43, Rhyssalus clavator Haliday, 9, Netherlands, Wijster. 39, fore

tibia, posterio-lateral aspect; 40, basal half of abdomen, dorsal aspect; 43, wings. Fig. 41, 42, Rogas

excavatus (Telenga), 4, Italy, Riva s. Garda. 41, head, frontal aspect; 42, head, dorsal aspect. Fig.

37-39: 2.5 times scale-line; Fig. 40-42: 1.2 times scale-line; Fig. 43: scale-line

C. vAN ACHTERBERG: The subfamilies of the Braconidae 61

Fig. 44, Rogas unipunctator (Thunberg), 9, Netherlands, Oostkapelle; wings. Fig. 45, 51, Polemochartus

liparae (Giraud). 45, 3, Netherlands, Haaren, three basal segments of metasoma, dorsal aspect; 51,

©, Netherlands, Eindhoven, mandible, ventro-lateral aspect. Fig. 46, Rogas drymoniae Watanabe, 1937;

habitus, dorsal aspect. Fig. 47-49, Tanycarpa punctata Van Achterberg, 9, holotype. 47, propodeum,

first and second metasomal tergites, dorsal aspect; 48, head, frontal aspect; 49, detail of mandible,

lateral aspect. Fig. 50, Chaenusa bergi (Riegel), 9, paratype; head, frontal aspect. Fig. 44, 45: 0.5

times scale-line; Fig. 46: 0.1 times scale-line; Fig. 47, 51: 2.5 times scale-line; Fig. 48: 1.2 times

scale-line; Fig. 50: 1.8 times scale-line; Fig. 49: 3.2 times scale-line

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 52, 53, Opius (Apodesmia) polyzonius Wesmael, 9, Netherlands, Asperen. 53, head, frontal aspect;

54, metasoma, dorsal aspect. Fig. 54, Apanteles falcator (Ratzeburg), 9, Netherlands, Waarder; wings.

Fig. 55, Mirax cremastobombyciae (Fullaway), 9, holotype; wings. Fig. 52: 2.5 times scale-line; Fig. 53,

54: scale-line; Fig. 55: 1.7 times scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 63

Fig. 56, Mirax cremastobombyciae (Fullaway), 9, holotype; basal half of abdomen, dorsal aspect.

Fig. 57, Microplitis theretrae Watanabe, after Watanabe, 1937; habitus, dorsal aspect. Fig. 58, Micro-

gaster takeuchii Watanabe, after Watanabe, 1937; habitus, dorsal aspect. Fig. 59, Apanteles falcator

(Ratzeburg), 9, Netherlands, Waarder; metasoma, dorsal aspect. Fig. 60, Cardiochiles japonicus Wata-

nabe, after Watanabe, 1937; habitus, dorsal aspect. Fig. 61, Acaelius spec., 9, Netherlands, Oost-

voorne; metasoma, dorsal aspect. Fig. 56, 61: 2.5 times scale-line; Fig. 59: scale-line; Fig. 57,

58: 0.2 times scale-line; Fig. 60: 0.1 times scale-line

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 62, Acaelius spec., 9, Netherlands, Oostvoorne; wings. Fig. 63, 65, Earinus nitidulus (Nees), 9,

Netherlands, Schaarsbergen. 63, first and second metasomal tergites, dorsal aspect; 65, wings. Fig. 64,

Meteoridea japonensis Shenefelt & Muesebeck, after Shenefelt & Muesebeck, 1957; wings. Fig. 62: |

4.0 times scale-line; Fig. 63, 65: scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 65

Fig. 66, Braunsia matsumurai Watanabe, 9, after Watanabe, 1937; habitus, dorsal aspect. Fig. 67, 68,

Acampsis alternipes (Nees), 9, Netherlands, Den Haag. 67, wings; 68, habitus, lateral aspect. Fig. 69,

Chelonus (Microchelonus) tosensis Watanabe, after Watanabe, 1937; 9, habitus dorsal aspect; à, apex

of metasoma, apical aspect. Fig. 70, Siga/phus irrorator (Fabricius), 9, Netherlands, Naaldwijk; meta-

soma, lateral aspect. Fig. 66, 0.1 times scale-line. Fig 67, 68, scale-line. Fig. 69, 0.4 times scale-line. Fig.

70, 0.6 times scale-line; t = tooth

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 71, 72, 74, Ascogaster instabilis Wesmael, 9, Netherlands, Asperen. 71, mesosoma, ventro-lateral

aspect; 72, metasoma, lateral aspect; 74, wings. Fig. 73, Sigalphus irrorator (Fabricius), 9, Netherlands,

Naaldwijk; wings. Fig. 71, 72, 74: scale-line; Fig. 73: 0.5 times scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 67

Fig. 75—77, Macrocentrus thoracicus (Nees), 9, Netherlands, Meijendel. 75, wings; 76, mesoscutum,

lateral aspect; 77, propodeum and first metasomal segment, lateral aspect. Fig. 78, 79, Cenocoelius

analis (Nees), 9, Netherlands, Wijster. 78, propodeum and first metasomal segment, lateral aspect;

79, wings. Fig. 80, Blacus (Ganychorus) striatus Van Achterberg, 9, holotype; propodeum and first

metasomal tergite, dorsal aspect. Fig. 75: 0.5 times scale-line; Fig. 76-79: scale-line; Fig. 80: 2.5 times

scale-line

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 81, 86, Austrozele brevicaudis (Szépligeti), 2, lectotype. 81, wings; 86, head, frontal aspect. Fig. 82,

Proterops nigripennis Wesmael, 9, Netherlands, Kralo; basal half of metasoma, dorsal aspect. Fig. 83,

Elachistocentrum similis (Szépligeti), 9, lectotype; propodeum and first metasomal tergite, dorsal

aspect. Fig. 84, Eubazus (Brachistes) lapponicus (Thomson), 9, lectotype: wings. Fig. 85, Ichneutes

spec., 9, Netherlands, Waarder; head, frontal aspect. Fig. 81: 0.4 times scale-line; Fig. 82: scale-line;

Fig. 83: 1.8 times scale-line; Fig. 84: 0.7 times scale-line; Fig. 85: 1.2 times scale-line; Fig. 86: 0.8

times scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 69

5 D TT 2a,

È S

Fig. 87, Eubazus (Foersteria) tibialis (Haliday), 3, Netherlands, Wijster; propodeum and first meta-

somal segment, lateral aspect. Fig. 88, 89, /chneutes spec., 9, Netherlands, Waarder. 88, wings; 89,

habitus, lateral aspect. Fig. 87-89: scale-line

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 90, 92, Proterops nigripennis Wesmael, 3, Netherlands, Kralo. 90, head, frontal aspect; 92,

wings. Fig. 91, Charmon extensor (Linnaeus), 9, Netherlands, Naardermeer; propodeum and first

metasomal segment, lateral aspect. Fig. 90: 2.5 times scale-line; Fig. 91: scale-line; Fig. 92: 0.6 times

scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae ZU

Fig. 93—95, Ephedrus plagiator (Nees), 9, Netherlands, Maastricht. 93, wings; 94, head, frontal aspect;

95, metasoma, lateral aspect. Fig. 96, Aphidius ervi Haliday, 9, Netherlands, Asperen; wings. Fig. 97,

Cosmophorus cembrae Ruschka, after Hedqvist, 1955; anterior part of head, lateral aspect. Fig. 98,

Cosmophorus klugi Ratzeburg, after Watanabe, 1968; head, dorsal aspect. Fig. 99, Cosmophorus regius

Niezabitowski, id. Fig. 93, 95, 96: scale-line; Fig. 94: 2.5 times scale-line

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 100, Charmon extensor (Linnaeus), 9, Netherlands, Naardermeer; wings. Fig. 101, Orgilus

laevigator (Nees), 9, Netherlands, Waarder; wings. Fig. 102, Microtypus wesmaeli Ratzeburg, 9,

Netherlands, Crailo: wings. Fig. 100, 102: 0.6 times scale-line; Fig. 101: scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 73

10 mm

Fig. 103, 104, Zele infumator Lyle, 9, Netherlands, Crailo. 103, Propodeum, lateral aspect; 104, wings.

Fig. 105, Cosmophorus regius Niezabitowski, after Watanabe, 1968; wings. Fig. 106, Microtypus wesmaeli

Ratzeburg, 9, Netherlands, Crailo; propodeum, lateral aspect. Fig. 103, 104: 0.6 times scale-line;

Fig. 106: scale-line

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig. 107, Zemiotes deceptor (Wesmael), 9, Netherlands, Wageningen; wings. Fig. 108, Leiophron (Leio-

phron) apicalis Haliday, 9, Netherlands, Wijster; apex of metasoma, lateral aspect. Fig. 109, Ancylo-

centrus ater (Nees), 9, Netherlands, Waarder; metasoma, lateral aspect. Fig. 110, Streblocera macro-

scapa (Ruthe), 9, Netherlands, Waarder; first metasomal tergite, dorsal aspect. Fig. 107: 0.6 times

scale-line; Fig. 108-110: 2.5 times scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae 75

ee seal oo tne 7

1.0mm

111

Fig. 111, Zemiotes deceptor (Wesmael), ©, Netherlands, Wageningen; metasoma, dorsal aspect. Fig.

112, Meteorus cf. sulcatus Szépligeti, 9, Netherlands, Wijster; metasoma, dorsal aspect. Fig. 113,

Meteorus ictericus (Nees), 9, Netherlands, Waarder; metasoma, dorsal aspect. Fig. 114, Syntretus

cf. conterminus (Nees), 9, Netherlands, Waarder; first metasomal tergite, dorsal aspect. Fig. 115,

Perilitus (Microctonus) cf. deceptor Wesmael, 9, Netherlands, Putten (G.); head, frontal aspect. Fig.

116 Chrysopophthorus spec., 9, Costa Rica, Turrialba; first metasomal tergite, dorsal aspect. Fig. 111:

0.6 times scale-line; Fig. 112, 113, 116: scale-line; Fig. 114, 115: 2.5 times scale-line

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

Fig.117, Perilitus (Microctonus) cf. deceptor Wesmael, 9, Netherlands, Putten (G.); wings. Fig. 118,

Leiophron (Leiophron) apicalis Haliday, 9, Netherlands, Oostvoorne; wings. Fig. 119, Chrysopophthorus

spec., 9, Costa Rica, Turrialba; wings. Fig. 117, 118: 1.2 times scale-line; Fig. 119: scale-line

C. VAN ACHTERBERG: The subfamilies of the Braconidae i

Fig. 120—122, Pseudodicrogenium monstrosum Fahringer. 120, fore wing, paralectotype; 121, head,

frontal aspect, lectotype; 122, head, lateral aspect, lectotype. Fig. 120, 122: scale-line; Fig. 121: 2.0

times scale-line

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 3, 1976

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\ e es = =

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TIE SY M = 5

ASS È, BER \ HH \

>= > \ bl x

! H DENE (dp) in

' (©) x x >

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rare) AN DS ON eee ees

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Sa 35

i 3 a = — o a ©

2, OE N o O df

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È ae ©) 6 Hd O

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Fig. 123. Dendrogram indicating the possible phylogenetic relationship and the hosts of the subfamilies

belonging to the Braconidae. A(!) = parasites of nymphs and adults of aphids; C = parasites of

larvae of Coleoptera; C! = parasites of adult Coleoptera; D = parasites of larvae of Diptera; F =

parasites of ant-larvae; F! = parasites of adult ants; H = parasites of larvae of Hymenoptera; H! =

parasites of adult Hymenoptera; HE(!) = parasites of nymphs and adults of Heteroptera; I =

associated with termites; L = parasites of larvae of Lepidoptera; M = parasites of larvae of

Mecoptera; N! = parasites of adult Neuroptera; P(!) = parasites of nymphs and adults of Psocoptera;

S = parasites of larvae of Symphyta; ? = host unknown

ÉD AT 568.2

DEEL 119 AFLEVERING 4 1976

TIJDSCHRIFT

VOOR ENTOMOLOGIE

US. AR ARY

DEC 8199

ARR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGINGS! 2

UITGEGEVEN DOOR

INHOUD

_ J. P. van LITH. — New species and records of Indo-Australian Psenini (Hymen-

optera, Sphecidae, Pemphredoninae), p. 79—122, fig. 1—53.

| Tijdschrift voor Entomologie, deel 119, afl. 4

Gepubliceerd 25-X-1976

NEW SPECIES AND RECORDS OF INDO-AUSTRALIAN

PSENINI (HYMENOPTERA, SPHECIDAE,

PEMPHREDONINAE)

J.P. VAN LITH

Allard Piersonstraat 28c, Rotterdam

With 53 text-figures

ABSTRACT

The following new species and subspecies are described and illustrated: Psen (Psen) nitidus

sabahensis, Sabah, nitidus binghami, Sikkim, elisabethae umboiensis, New Guinea, elisabethae bougain-

villensis, Solomon Is., Psen (subgenus?) anodontotus, New Guinea, Psen (Mimumesa) oresterus, West

Pakistan, Psenulus major, Laos, decipiens, Laos, laosensis, Laos, armipes, Philippine Is., tectus, Laos,

anomalus, Vietnam, vaneuensis, Laos, puncticeps rufipes, Sumba, exiguus, Laos, Malaya, suturalis, Laos,

crabroniformis wapiensis, Laos, crabroniformis nathani, South India, /imbatus, Laos, impressus, Philippine

Is., pictus, New Guinea, ornatus sumbaensis, Sumba, fyanensis, Vietnam, Laos, nigrolateralis, Laos,

vientianensis, Laos, leucognathus, South India. Many new records of known species are given, in some

cases also a description of the hitherto unknown opposite sex. Psen (Mimesa?) inflatus is trans-

ferred to the subgenus Psen.

During the past decade, many Psenini have been collected in the Indo-

Australian area. Special mention should be made here of the rich material

gathered in Laos by native collectors for the Bishop Museum, Honolulu. Thus far

no Psenini were known from that country. Now 17 Laotian species and subspecies

can be listed, of which 12 are described here as new. Ten of these forms are

recorded from Laos only. Much new information has also been obtained with

regard to the distribution of earlier described species.

The male genitalia have been figured as much as possible. It is obvious that

comparison of the parameres is of good assistance in the classification in species

groups or subgenera. Since my last key to the Indo-Australian species of Psenulus

(1962) the number of forms has more than doubled and now amounts to 134. A

new key for Psenulus is now in preparation. Notwithstanding the progress made

our knowledge of the Indo-Australian Psenini is still very incomplete; much more

collecting in this region is highly desirable. Of only 36 forms both sexes are known.

For 62 females and for 36 males the opposite sex has not yet been found or

recognized. For the genus Psen the figures are somewhat better (40, 31 and 18,

respectively). We know very little about their bionomics. Best known in this

respect is Psenulus sogatophagus Pagden, 1933. Dr. D. H. Murphy, Singapore,

kindly provided some information regarding nesting and prey of Psenulus

trimaculatus.

I am much obliged to the Bishop Museum and all other friends and institutions

who sent me their material for study. Their names are given below together with

the symbols used in the text. Special thanks are due to Prof. J. van der Vecht, who

stimulated me long ago to study the Psenini and continues to show interest, and to

80 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Mrs. E. van der Vecht-Bourguignon who collected two new species during their

visit to New Guinea in 1972.

AMNH — The American Museum of Natural History, New York, N.Y., U.S.A.:

Mrs. M. Favreau, J. G. Rozen Jr.

BISH — Bernice P. Bishop Museum, Honolulu, Hawaii, U.S.A.; Miss A.

Manning, F. J. Radovsky

BM — British Museum (Natural History), Department of Entomology,

London, U.K.; L.A. Mound, C. R. Vardy

CGR — C.G. Roche, Kuala Lumpur, Malaysia

CNC — Entomology Research Institute, Canada Department of Agriculture,

Ottawa, Canada; J. Barron, L. Masner, C. M. Yoshimoto

ELW — Laboratorium voor Entomologie, Wageningen, The Netherlands; via -

J. van der Vecht

HT — H. and M. Townes, American Entomological Institute, Ann Arbor,

Michigan, U.S.A.

MA — Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Af-

deling Entomologie, Amsterdam, The Netherlands; J. P. Duffels, W. .

N. Ellis

ML — Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands; J.

van der Vecht

NMB — Naturhistorisches Museum, Basle, Switzerland; via J. van der Vecht

UI — University of Idaho, Department of Entomology, Moscow, Idaho,

U.S.A.; A. R. Gittins

US — University of Singapore, Singapore; D. H. Murphy

USNM — National Museum of Natural History, Smithsonian Institution,

Washington, D.C., U.S.A.; P. D. Hurd, K. V. Krombein, A. S. Menke

Psen Latreille

Subgenus Psen

Psen (Psen) emarginatus Van Lith

Van Lith, 1959: 43—44 (Java); 1968: 101—102 (N. Borneo).

New record from Borneo: 19 and 1g, Sabah, Mt. Kinabalu, 5000 ft, 1—5 May

1973, K. M. Guichard (CGR).

Psen (Psen) rufoannulatus Cameron

Cameron, 1907: 90 (India: Simla).

Van Lith, 1965: 58—60; 1973: 125—126 (Nepal).

New record from Nepal: 18, Kathmandu, 1300—1400 m, 7—12 May 1960, J.

and M. Sedlacek (BISH).

Psen (Psen) nitidus nitidus Van Lith

Van Lith, 1959: 28— 30 (Java, Bangka, Krakatau, Sri Lanka); 1968: 105—106 (S. India, NE. Sumatra).

First record from Malaya: 1 ©, Johore, Kr. Pulai, 21 Dec. 1967 (CGR).

J. P. van LITH: Indo-Australian Psenini 81

Fig. 1—2. Psen (Psen) nitidus binghami subsp. nov., g, holotype, outer parts of genitalia, dorsal aspect,

paramere, ventral aspect. Fig. 3—4. Psen (Psen) elisabethae umboiensis subsp. nov., 3, holotype,

genitalia, ventral and dorsal aspect. Fig. 5. Psen (Psen) elisabethae bougainvillensis subsp. nov, 3,

holotype, 8th sternite, lateral aspect. Fig. 6—8. Psen (Psen) inflatus Van Lith, 3, genitalia in dorsal,

ventral and lateral aspect. Fig. 9. Psen anodontotus sp. nov., pygidial area of 9, holotype. Fig. 10—12.

Psen (Mimumesa) oresterus sp. nov., 10, 9, pygidial area; 11—12, 3, paramere, outer and inner aspect

82 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Psen (Psen) nitidus takasago Tsuneki

Tsuneki, 1967: 8—9 (Taiwan).

Van Lith, 1968: 106.

New record from Taiwan: 29, Sauter coll. (AMNH).

Psen (Psen) nitidus sabahensis subsp. nov.

Two females from North Borneo differ from the female of the nominate form

and from the female of the subspecies takasago in having dark brown instead of

yellowish-brown, hind tarsi. The apex of the antennae below is blackish-brown.

Fore tibiae and fore and mid tarsi are reddish-brown, slightly darker than in other .

forms of P. nitidus.

Scutum more strongly punctate than in nominate form. Pygidial area broader,

its base more shining and punctures along lateral margins larger. Pubescence of

scutum somewhat brownish, greyish-white in the nominate form.

Male unknown.

Borneo: 29, holotype (Roche No. 15222) and paratype (Roche No. 15221),

Sabah, Mt. Kinabalu, 5000 ft, 1—5 May 1973, K. M. Guichard (CGR). The

holotype has been deposited by Mr. Roche in the British Museum (Natural

History), London.

Psen (Psen) nitidus binghami subsp. nov.

Male. — Resembling nominate form but differing in having dark brown hind

tarsi, brown mid tarsi, strongly punctate scutum and brownish pubescence on

thorax and gaster.

Genitalia (Fig. 1, 2) as in nominate subspecies.

Sikkim: 14 , holotype, coll. Bingham 1902—120 (BM).

Psen (Psen) elisabethae elisabethae Van Lith

Van Lith, 1959: 31—34 (Java, Sumatra); 1968: 106 (Vietnam).

New record from North Sumatra: 19, Sibolangit, Lortzing (ELW ex coll.

Roepke).

New record from West Java: 1g, Mt. Gedeh, 14 Aug. 1965, J. E. Lukavsky

(CNC).

Psen (Psen) elisabethae madrasiensis Van Lith

Van Lith, 1968: 107 (S. India).

First records from Laos: Vientiane Prov., Ban Van Eue, 19, 15 Febr. 1966,

Malaise trap, | 3, 29 March 1966, native collector (BISH).

The face of this female is distinctly silvery pubescent. Pygidial area narrow, as in

nominate subspecies.

J. P. van LITH: Indo-Australian Psenini 83

Psen (Psen) elisabethae umboiensis subsp. nov.

Male. — Much resembling nominate form. Length about 10 mm. Femora and

tibiae black, tarsi yellowish-brown, antennae and tegulae black. Petiole about

twice as long as first tergite. Last antennal segment over twice as long as broad at

base. Pubescence of face and tempora silvery-white, mostly appressed.

Pubescence of rest of body whitish, thorax dorsally more greyish. Genitalia (Fig. 3,

4) large, brown.

NE. New Guinea: | 3, holotype, Umboi I., about 8 km WNW. of Lab Lab, 300

m, 8—19 Febr. 1967, G. A. Samuelson, Malaise trap; 18, paratype, Umboi I., |

km N. Awelkom, 600 m, 21—28 Febr. 1967, G. A. Samuelson, Malaise trap

(BISH).

This subspecies is distinguished from the nominate form by the larger size,

longer antennae and dark legs and tegulae.

Psen (Psen) elisabethae bougainvillensis subsp. nov.

Male (holotype). — Resembling preceding form. Length over 11 mm. Legs

and tegulae black, tarsi yellowish-brown. Petiole about 1% times as long as first

tergite. Last antennal segment over 2% times as long as broad at base. Punctation

fine. Pubescence as in preceding form. Eighth sternite: Fig. 5.

Solomon Is.: 1g, holotype, Bougainville, Kukugai Vill., 150 m, Nov. 1960, W.

W. Brandt (BISH).

The genitalia of this specimen are like those of the subspecies umboiensis. It is

distinguished from the nominate subspecies by the dark legs, shorter petiole,

longer antennae and large size. It differs from umboiensis by the shorter petiole

and longer antennae.

Psen (Psen) coriaceus Van Lith

Van Lith, 1959: 34—36 (Mindoro, Luzon, Negros, Mindanao).

New record from Luzon: 1 9, Baguio, June 1917, F. X. Williams (BISH).

Psen (Psen) paulus paulus Van Lith

Van Lith, 1968: 113—114 (New Guinea).

New records from NE. New Guinea: 19, Mt. Missim, 980 m, Malaise trap, 20

July 1969, J. L. Gressitt and Y. Hirashima, 13 , Mt. Missim, 1300 m, Malaise trap,

7—21 Dec. 1966, G. A. Samuelson, 24, Wau, 1000 m, 23 Dec. 1969, M. Sedlacek

(BISH). |

SE. New Guinea: 13, Mamai Pltn, E. of Port Glasgow, 150 m, light trap, 17

Febr. 1965, R. Straatman (BISH).

Psen (Psen) bryani Perkins & Cheesman

Perkins & Cheesman, 1928: 28—29 (Samoa).

Van Lith, 1968: 116.

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

New records from Samoa: 19, Tutuila I., 9 Aug. 1957, W. R. Kellen, 24,

Tutuila, Vatia, O—200 m, 9 March 1971, N. L. H. Krauss (BISH).

Psen (Psen) hakusanus seminitidus Van Lith

Gussakovskij, 1934: 7 (Mimesa kohli, nec Psen kohli Fox, 1898; China); 1937: 653 (Psen kohli; China,

Tibet).

Van Lith, 1965: 40 (Psen (Psen) seminitidus new name); 1968: 119.

Tsuneki, 1966: 9—10 (Psen (Psen) hakusanus seminitidus new status; Taiwan); 1967: 2—3; 1971: 17.

New record from Taiwan: 19, Taiwan C., Alisan, 2400 m, 3—9 July 1972, T. C.

Maa (BISH).

Psen (Psen) orientalis Cameron

Cameron, 1890: 269 (Madras, India); 1902: 289 (Psen reticulatus; India).

Van Lith, 1965: 42—44; 1968: 120—121.

New record from Kashmir: 19, Gulmarg, summer 1913, F. W. Thomson (BM).

Psen (Psen) politiventris politiventris Rohwer

Rohwer, 1921: 321 (Psen (Mimesa) politiventris; Luzon).

Van Lith, 1959: 36—37 (Psen (Psen) politiventris); 1965:48.

New records from Luzon: Mountain Prov., Abatan, Buguias, 60 km S. of

Bontoc, 1800—2000 m, 13, 27 May 1964, 19, 15 June 1964, H. M. Torrevillas

(BISH).

Psen (Psen) alishanus Tsuneki

Tsuneki, 1967: 3—5 (Taiwan); 1971: 1, 17.

New records from Taiwan: 29, Taiwan C., Alisan, 2400 m, 3—9 July 1972, T. C.

Maa (BISH).

Psen (Psen) refractus meridianus Van Lith

Van Lith, 1965: 51—52 (S. India).

New record from South India: 19, Anamalai Hills, Cinchona, 3500 ft, May

1969, P. S. Nathan (MA).

Psen (Psen) aureohirtus aureohirtus Rohwer

Rohwer, 1921: 322—323 (Psen (Mimesa) aureohirta; Philippine Is.).

Van Lith, 1959: 49—50 (Psen (Psen) aureohirtus ); 1965: 56; 1968: 121.

New record from Luzon: 19, Mountain Prov., Abatan, Buguias, 60 km S. of

Bontoc, 1800—2000 m, 9 May 1964, H. M. Torrevillas (BISH).

J. P. van LITH: Indo-Australian Psenini 85

Psen (Psen) ruficrus Van Lith

Van Lith, 1965: 62—63 (New Guinea); 1968: 122— 124.

New records from NE. New Guinea: 30 km E. of Goroka, 2000—2100 m, 19,

15—21 April 1968, 13, 22—31 May 1968, J. and M. Sedlacek (BISH).

Psen (Psen) sedlaceki Van Lith

Van Lith, 1968: 124—125 (New Guinea).

New record from NE. New Guinea: 1g, Morobe District, Ulap, 800— 1100 m,

Sept. 1968, N. L. H. Krauss (BISH). This is the second male recorded, the female

is still unknown.

Psen (Psen) inflatus Van Lith

Van Lith, 1968: 129—130, 9 (Psen (Mimesa?) inflatus; Central New Guinea).

New records: NE. New Guinea, Wau, 1g, Jan. 1970, 19, 2100-2300 m, 3 Aug.

1971, Sedlacek (BISH); 24, Wau, 2100—2360 m, Mt. Kaindi, 14 Sept. 1972, J. van

der Vecht (ML).

Female. — Somewhat darker than the form from Central New Guinea. Head,

scutum and mesopleura including hypo-epimeral area black, with indistinct bluish-

black shine, propodeum with steel-blue or violet-blue reflections. Dorsum and

ventral plate of petiole black, sides and underside of petiole dark reddish-brown.

Gaster black with metallic-blue shine, apex reddish-brown. Femora with light

metallic shine, hind tibiae dorsally dark brown, reddish-brown below. Apical third

of fore wings infuscate. Interantennal tooth blunt in dorsal aspect, as in the

females from Central New Guinea. Petiole about ten times as long as wide in the

middle, in dorsal aspect.

First description of male. — Resembling female. Scape and following segment

of antennae and basal half of underside of flagellum reddish. In two of the males

the epicnemium has a violet-blue metallic shine, in one of these two males the

mesopleura including hypo-epimeral area with the same shine. The male caught in

January has a completely brownish-black petiole, in the two other males the

petiole is reddish-brown with dark dorsal line.

Antennae long, third segment about four times, segments 4—6 over 2% times,

segments 7—10 about 2% times, segments 11-12 over twice as long as broad at apex,

segment 13 over 2% times as long as broad at base. No tyloidea. Gastral sternites

3—4 with long, dark brown fasciculate hairs in middle of hind margin. Genitalia

(Fig. 6—8) long, brownish-yellow, inner apex of stipes with long and broad

membranous extension or flag.

Because of the characteristic fasciculate hairs on sternites 3—4 of the male and

the structure of the genitalia this species has now been placed in the subgenus

Psen, although the epicnemial region in some respects more resembles that of

Mimumesa. The cone-shaped, laterally extending hypo-epimeral areas and the high

transverse carinae on the ventral side of the thorax distinguish this form from any

other species of Psen sl.

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

The specimens from NE. New Guinea differ from the two females from Central

New Guinea mainly in the metallic shine of the propodeum not being bronze or

brassy, but steel-blue or violet-blue. There are no distinct structural differences.

The north-eastern form may represent a different subspecies. The metallic

reflection varying in intensity and also the colour of the petiole being variable in

the males, a decision must be postponed until more material, especially from

Central New Guinea, is available.

Psen (?Psen) anodontotus sp. nov.

Female (holotype). — Head and thorax shining black, sides and back of thorax

with some steel-blue shine. Labrum and median part of mandibles dark reddish, .

palpi pale brown, pronotal tubercles dark brown, tegulae reddish-brown. Apex of

scape of antennae below and flagellum below brownish. Femora dark brown,

reddish on foreside, also fore trochanters reddish. All tibiae and tarsi reddish,

except fore and mid basitarsi which are pale yellowish. Petiole black, apex dorsally

and ventral plate and rest of gaster reddish. Veins of wings black. i

Frons impunctate and shining, on both sides of median carina somewhat

convex; vertex shining with a few punctures. Back of head more densely punctate.

Frontal carina very slightly broadening downwards, not ending in a tooth between

antennae. Clypeus densely finely punctate, apical margin shining, very slightly

emarginate, almost blunt, basal half of disk convex. Occipital carina below rather

high, ending normally in hypostomal carina. Scape long and slender, about five

times as long as broad, flagellum clavate, third segment about 3% times, fourth

segment about twice as long as broad at apex, following segments gradually

thickening, segments 8—11 as long as broad or slightly less, segment 12 about 1%

times as long as broad at base. Mandibles bidentate at apex.

Dorsum of pronotum narrow, lateral corners sharp. Scutum and scutellum

shining, sparsely but distinctly punctate, metanotum finely punctate. Dorsal and

hind part of propodeum forming a distinct angle. Enclosed area of propodeum

triangular, median two longitudinal carinae much diverging, lateral parts with only

a few oblique carinae. Behind propodeal enclosure a broad smooth and shining

area, back of propodeum with coarse reticulate carination, sides almost

smooth. Metapleura shining. Mesopleura and mesosternum shining with a few fine

punctures. Hypo-epimeral area strongly convex, smooth and shining. Anterior

oblique suture foveolate, surface of anterior plate dull. Interepicnemial area

roundly depressed, depression margined laterally by a distinct carina, which

continues into the inner carina of the epicnemial areas; at the junction of these

carinae a short backwards bent carina defining inner third of lower part of

epicnemial area. Outer epicnemial carina slightly bent backwards, epicnemial

areas not closed below. Acetabular carina a little behind the carina margining the

interepicnemial area, slightly longer than half the distance between outer

epicnemial carinae. Behind acetabular carina a few weak transverse carinae,

median mesosternal carina distinct, ending before apex. Metasternum emarginate

(angle about 135 degrees), with transparent margin. Legs rather thick. Upper two-

thirds of back of hind femora smooth and shining, separated from lower part,

J. P. VAN LITH: Indo-Australian Psenini 87

which bears a few hairs, by a narrow dull stripe consisting of very fine punctures,

each bearing a short fine hair. Second recurrent vein of fore wings ending just in

third submarginal cell.

Petiole cylindrical, slender, about 1% times as long as first tergite, dorsally

smooth. Gaster shining, hardly punctate, broad hind margins of tergites smooth,

impunctate. Pygidial area (Fig. 9) shining, elongate-triangular, apex rounded,

lateral carinae distinct but short, along margin a few punctures.

Head and thorax whitish pubescent, pubescence on clypeus silvery, mostly short

and appressed, near apex of clypeus a few long stiff yellowish hairs. Tempora

somewhat appressed whitish pubescent. Petiole ventro-laterally with fine and long,

obliquely downward directed hairs. Pubescence of legs yellowish-golden, on gaster

sparse, denser on sixth sternite.

Length about 7.5 mm.

Male unknown.

East New Guinea: 19, holotype, Wau, 1500 m, McAdam Park, on Hibiscus

flowers, 17 Sept. 1972, Mrs. E. van der Vecht (ML).

P. anodontotus differs from the other species of the subgenus Psen, where it is

provisionally placed, in having short, clavate antennae, in the interepicnemial area

which is depressed and margined below by a fine rounded carina in front of the

acetabular carina, and by the inner carinae of the epicnemial areas. The inter-

antennal carina does not end in a sharp tooth or elevated area, as is usual in the

subgenus Psen. The structure of the epicnemial region is similar to that of the

palaearctic species of the subgenus Mimumesa Malloch. The petiole differs,

however, in being cylindrical instead of carinate.

Subgenus Mimumesa Malloch

Psen (Mimumesa) oresterus sp. nov.

Female. — Length about 6.5 mm. Rather resembling the palaearctic P. littoralis

(Bondroit). Apical three antennal segments pale brown, rest of underside of

antennae dark brown. Tarsi brown, tarsal segments 2—S of fore legs pale brown.

Punctation of head between ocelli and oculi as in P. littoralis, vertex with some

fine transverse striation. Scutum shining, distinctly finely punctate, hind margin

with short longitudinal striae; scutellum punctate, posteriorly hardly striate.

Metanotum dull. Central area of propodeal enclosure without median longitudinal

carina (holotype) or with irregular median longitudinal carina (paratype). Pygidial

area (Fig. 10) as in P. littoralis.

Male. — Same size. Rather resembling P. unicolor Van der Linden, but vertex

not densely punctate. Antennae brown below, apical three segments paler. All

tarsi and apex of fore tibiae pale brown.

Antennal segments 4—12 with linear tyloidea, tyloides on segment 12 about ?/,

of total length of segment and situated on basal half. Scutellum posteriorly slightly

striato-punctate. Central area of propodeal enclosure with median longitudinal

carina. Genitalia: Fig. 11, 12.

West Pakistan: Quetta, Baluchistan, 29, holotype and paratype, | 3, allotype,

June 1902, 1 3, paratype, Aug. 1903, C. G. Nurse (BM).

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

The last mentioned male has a deformed petiole, with apex about half as broad

as propodeum and about twice as long as it is wide at apex.

Subgenus Mimumesa?

Psen (Mimumesa?) tridentatus tridentatus Van Lith

Van Lith, 1959: 55—56 (Sumatra); 1965: 67 (Malaya).

First record from Vietnam: 19, 17 km S. of Dilinh, 1300 m, 6—13 Oct. 1960,

C.M. Yoshimoto (BISH).

First record from Laos: 13, Vientiane Prov., Ban Van Eue, 29 March 1966,

native collector (BISH). .

The genitalia of this male agree with those of a male of P. tridentatus

chrysomallus from Burma. Basiparameres and parameres chestnut brown, valves of

aedeagus (Fig. 15) yellowish-brown, volsellae dark brown. Basiparameres and

parameres (Fig. 13, 14) heavily built, parameres thick, inner margin with fringe of

stiff hairs.

The subgeneric position of P. tridentatus, P. auratus and P. multipunctatus is still

uncertain. Their genitalia are very similar to those of Mimumesa, in which

subgenus they were originally placed, but they are much heavier. They are much

different from the genitalia of the subgenera Psen, Punctipsen and Pseneo. The

petiole is remarkable because of the structure of the dorsal side. It has an irregular

apical groove and large lateral punctures prolonged into short grooves. The

propodeum is conspicuously longitudinally striate. The acetabular carina is

complete, as in Mimumesa and Pseneo. In P. tridentatus the clypeal margin is more

or less tridentate. Although in Pseneo the acetabular carina, clypeal margin and

pygidial area are very similar and the sternal apical tufts are equally lacking in the

Fig. 13—15. Psen tridentatus tridentatus Van Lith, 3, 13—14, genitalia in dorsal and lateral aspect; 15,

penis valve. Fig. 16. Psen tridentatus chrysomallus Van Lith, ©, pygidial area

J. P. van LITH: Indo-Australian Psenini 89

males, I do not believe that P. tridentatus should be considered to belong to this

genus. Probably a new subgenus should be erected for the three species in

question. This subgenus would mainly be characterized by the structure of

propodeum and petiole, the complete acetabular carina and the lack of sternal

tufts. I hope to return to this problem later.

Psen (Mimumesa?) tridentatus chrysomallus Van Lith

Van Lith, 1965: 68, 3 (Burma).

Fortunately also the female of this subspecies has turned up. It was collected a

few days earlier than the two males described in 1965, at the same locality.

It resembles the female of the nominate form, but is distinguished by the larger,

size, the paler base of the antennae and the stronger sculpture of scutum and

propodeum.

Anterior margin of clypeus as in male (Van Lith, 1965: Fig. 77). Frons below

median ocellus with two small low tubercles, each at a distance from median

ocellus and from median frontal carina about as large as diameter of ocellus. A

slightly raised elongate smooth area along oculi. Vertex not densely, partly very

finely, punctate. Third antennal segment in frontal view about four times as long

as broad at apex, following two segments about twice, sixth segment about 1%

times as long as broad, segments gradually diminishing in length, segment 11 about

as long as broad at apex and last segment about 1% times as long as broad at base.

Scutum more closely punctate than in nominate form, especially in the middle

of the scutum, with only a few small interstices between the punctures. In the

nominate form the smooth spaces between the punctures are much larger in the

middle of the scutum, about four or five times as large as diameter of punctures.

Anterior margin of scutum much finer punctate in both forms, but in the female

from Burma distinctly finer than in the nominate form. Enclosed area of

propodeum not depressed, behind this area the number of longitudinal carinae is

greater than in the nominate form, as a consequence the smooth areas in the

antero-lateral corners of the back of the propodeum are smaller. Punctation of

tergites fine and rather dense, interstices on apical half about three or four times

as large as punctures, interstices on margin of tergites 4—5 larger. The fine

punctures on the fifth tergite are intermixed with a few larger punctures, it has a

broad smooth margin. Petiole as in nominate form. Pygidial area also similar,

lateral keels high, apex with small emargination, apically an indistinct median

longitudinal keel (Fig. 16).

Face with dense, appressed silvery pubescence and also with long silvery hairs.

Pubescence of gaster paler than in male. Apical margin of first two tergites with a

row of long, sidewards directed, pale golden hairs. The hairs should be examined

from above, because they may appear to be much darker when seen from aside.

Apical margin of first two tergites laterally and margins of tergites 3—4 with long

whitish hairs, also pubescence on disk of all tergites very pale.

Length about 13 mm.

Upper Burma: 19, Nam Tamai Valley, 3000 ft, lat: N 27°42’, long. E 97°54’, 12

Aug. 1938, R. Kaulback (BM).

90 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Psen (Mimumesa?) multipunctatus Van Lith

Van Lith, 1959: 55 (Negros); 1965: 68 (Psen (Mimumesa) auratus multipunctatus; Luzon).

New record from Negros: 1 9, Negros Or., L. Balisasayao, 1—7 Oct. 1959, L. W.

Quate (BISH).

New record from Luzon: 19, Mount Prov., Ifugao, Mayoyao, 1000—1500 m, 11

July 1966, H. M. Torrevillas (BISH).

Because of the strong punctation of the gaster this form is now recognized as a

distinct species. The male is still unknown.

Psenulus Kohl

Psenulus bicinctus Turner

Turner, 1912b: 363—364 (Assam).

Van Lith, 1972: 159—160 (Assam, Sikkim, Nepal).

First record from Burma: 4g, NE. Burma, Kambaiti, 7000 ft, 1 May 1934, R.

Malaise (BM).

Apparently the colour of the gaster is somewhat variable. In two of the males

the first tergite is reddisch, the two other males have this tergite largely brownish or

blackish. Third tergite with dark brown median area only, fourth tergite dark

brown with reddish apical margin.

Petiole nearly twice as long as first tergite, longer than in the female, where it is

at most 1% times as long as first tergite.

Genitalia (Fig. 17) pale yellowish-brown, parameres deeply emarginate on inner

side, a triangular tooth on apex of inner side of basiparameres.

Psenulus orinus Van Lith

Van Lith, 1973: 131—133 (Nepal).

New record from Nepal: 19, Bokaihunde, 20 km N. of Trisuli (Nawakot), 2100

m, 13—17 Nov. 1965, L. W. Quate (BISH).

The genitalia closely resemble those of the preceding species. Both species are

placed in the group of Psenulus rufobalteatus Cameron. The genitalia of rufobal-

teatus have not yet been examined.

Psenulus major sp. nov.

Male. — Length about 10—10.75 mm. Black, with bluish shine. Base of

flagellum below, base of mandibles, labrum, palpi and fore tibiae and tarsi reddish-

brown; mid tarsi, base of hind tibiae, tegulae, apex of petiole and sides of first

tergite brown. Wings distinctly fuscate, veins brown.

Interantennal carina sharp, ending below antennae in a low transverse carina.

Clypeus shining, densely superficially punctate, anterior margin with deep, almost

triangular emargination and short lateral teeth. Frons densely minutely punctate,

vertex shining, sparsely finely punctate, tempora densely very finely punctate.

J. P. van LITH: Indo-Australian Psenini 91

Antennae long, segments rounded below, about twice as long as broad at apex or

1% times as long as broad in the middle. Segments 4—12 with narrow tyloidea, on

most segments almost as long as segment, indistinct carina on segment 3. Occipital

carina ending in hypostomal carina.

Lateral corners of pronotum obtuse. Scutum shining, distinctly punctate,

interstices up to a few times size of punctures, sometimes in rows, punctures much

finer and sparser on outer side of subpleural sutures. Prescutal sutures distinct,

reaching hind margin. Scutellum shining, distinctly punctate, much sparser on

anterior half. Enclosed area of propodeum triangular, shining; central area wide,

with indistinct median carina, in lower part with a few transverse arched carinae.

Back of propodeum coarsely reticulate, carinae almost parallel on upper part,

horizontal part behind enclosed area smooth and shining. Mesopleura, hypo-

epimeral area, mesosternum and anterior plate of mesepisternum shining, sparsely

finely punctate. Anterior oblique suture foveolate, widened upper part with

longitudinal carinae. Legs slender. Petiole almost cylindrical, about twice as long

as first tergite, apex above with short, elongate-triangular groove. Gastral

segments shining, densely finely punctate. Eighth gastral sternite with long spine.

Genitalia (Fig. 18) dark brown, parameres broad with rounded apex, apex of

basiparameres on inner side with large triangular tooth.

Pubescence of face and frons greyish-brown, dense and short, with some longer

hairs, laterally somewhat depressed but not on clypeal disk so that the sculpture is

easily seen. Thorax and gaster brownish-grey pubescent, mesosternum whitish.

Petiole latero-dorsally with a row of mostly short hairs, latero-ventrally with long

erect hairs, ventral plate with dense short pubescence.

Laos: 1g, holotype, Vientiane Prov., Ban Van Eue, 29 March 1966; 64, 15

March 1966, 33, 29 March 1966, 1¢, 30 March 1966, 1¢, 30 March 1967, all

paratypes, same locality, some specimens collected in Malaise trap, native

collectors (BISH).

The punctation of the scutum varies somewhat and is in some paratypes finer

and sparser than in the holotype.

P. major is easily distinguished from other black Psenulus by the narrow

interantennal carina, the long and narrow tyloidea, large size and long petiole.

Psenulus decipiens sp. nov.

Male (holotype). — Length about 8 mm. Head and thorax black. Mandibles

except tips yellowish-red, palpi yellowish-brown. Scape of antennae and underside

of flagellum yellowish-brown, flagellum dorsally brown. Pronotal tubercles brown.

Legs including trochanters yellowish-red, fore and mid femora below somewhat

brown. Gaster including petiole yellowish-red. Veins of wings dark brown.

Anterior margin of clypeus bidentate, distinctly emarginate. Interantennal

carina narrow, ending below antennae in transverse carina. Frons and vertex

sparsely finely punctate. Postocellar area not raised. Occipital carina ending

before hypostomal carina. Antennae about as long as head and thorax together,

segments rounded below, segment 3 about 2% times, segments 4—12 about twice

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Fig. 17—20. Male basiparameres and parameres, dorsal aspect, of various species of Psenulus. 17, P.

bicinctus Turner, Burma; 18, P. major sp. nov., paratype; 19, P. decipiens sp. nov., holotype; 20. P.

laosensis sp. nov., paratype. Fig. 21—22. Psenulus armipes sp. nov., 9, holotype, 21, face, lower part; 22,

thorax, dorsal aspect

J. P. van LITH: Indo-Australian Psenini 93

as long as broad at apex, last segment over 2% times as long as broad at base. No

distinct tyloidea.

Pronotal corners rounded. Scutum shining, distinctly irregularly punctate, on

anterior half weaker and more sparsely. Prescutal sutures as long as scutum, less

distinct on posterior half. Scutellum and metanotum shining, sparsely punctate.

Enclosed area of propodeum narrow, longitudinal groove narrow, back of

propodeum shining, below and laterally irregularly carinate, also posterior half of

sides. Mesopleura, hypo-epimeral area, anterior plate of mesepisternum and

mesosternum shining, minutely punctate. Anterior oblique suture foveolate,

widened upper part with a few oblique carinae. Legs slender. First recurrent vein

of fore wings ending well before end of first submarginal cell. Petiole cylindrical,

little longer than first tergite, apex with small triangular groove. Gastral tergites

minutely sparsely punctate.

Pubescence of face and dorsal side of pronotum silvery, mostly appressed.

Pubescence of rest of body yellowish-grey.

Genitalia (Fig. 19) resembling those of P. laosensis, basal part yellowish-brown,

greater part of parameres dark brown, apex rounded, inner corner of basi-

parameres with low tooth.

Laos: 18, holotype, Vientiane Prov., Ban Van Eue, 15—31 May 1965, native

collector (BISH).

P. decipiens seems to be closely related to P. major and P. laosensis but differs,

besides in other characteristics, in the absence of distinct tyloidea.

Psenulus laosensis sp. nov.

Male. — Length about 8 mm. Head and thorax black; mandibles except apices

yellowish, palpi brownish-yellow, antennae dark brown, scape and basal half of

underside of flagellum reddish-brown, pronotal tubercles yellowish-brown. Legs

including trochanters and gaster including petiole yellowish-red. Veins of wings

brown.

Anterior margin of clypeus bidentate, distinctly emarginate. Interantennal

carina very slightly broadened dorsally, with fine groove, carina ending below

antennae in transverse carina. Occipital carina ending before hypostomal carina.

Frons and vertex minutely sparsely punctate. Postocellar area not raised.

Antennae longer than head and thorax together, segments laterally somewhat

flattened, underside rounded. Segment 3 in lateral view over twice, segments

4—10 about twice, segments 11—12 over twice as long as broad at apex, segment

13 nearly three times as long as broad at base. Segments 4—12 with oblique long

narrow tyloidea, longest on median segments but not reaching base or apex.

Pronotal corners rounded. Scutum shining, minutely punctate, prescutal sutures

distinct on anterior half, less distinct on posterior half. Scutellum and metanotum

shining. Enclosed area of propodeum short triangular; upper part of back of

propodeum shining, lower part and posterior part of sides reticulate, median

longitudinal groove narrow. Mesopleura, hypo-epimeral area, anterior plate of

mesepisternum and mesosternum shining, indistinctly punctate. Anterior oblique

suture foveolate, widened upper part with a few oblique carinae. Legs slender.

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

First recurrent vein of fore wings ending in second submarginal cell. Petiole

almost cylindrical, nearly 1% times as long as first tergite, a small triangular

depression at apex. Gastral tergites minutely punctate.

Basal part of genitalia (Fig. 20) reddish-brown, parameres dark brown, broad

with rounded apex, inner shoulder of basi-parameres with low tooth.

Pubescence of face and pronotum pale golden, mostly appressed, of rest of body

yellowish-brown.

Laos: 18, holotype, Vientiane Prov., Ban Van Eue, 29 March 1966, native

collector; 43, paratypes, same locality, 15—30 March 1966, one in Malaise trap,

native collector (BISH).

P. laosensis is probably closely related to the entirely black P. major, perhaps

also to P. decipiens.

Psenulus armipes sp. nov.

Female (holotype). — Length about 8 mm. Head black; basal half of mandibles

yellow, palpi straw-yellow. Scape of antennae yellowish, following segments.

dorsally brown, ventrally yellowish-brown. Thorax yellow, lower part of pronotum

black, propleura dark brown. Posterior ‘/, of area between prescutal sutures black,

posterior corners of scutum black (Fig. 22). Metapleura and an adjacent mark on

mesopleura brown, a brown streak on mesopleura below. Interepicnemial area

and median longitudinal band on mesosternum brown. Enclosed area of propo-

deum, median longitudinal groove and narrow apical margin of propodeum black.

Fore and mid legs including coxae yellow, hind legs yellowish-red, hind coxae

yellowish. Petiole dark brown, ventral plate of petiole and gaster reddish.

Head thick. Frontal carina raised and widened between antennae, broadened

part about twice as long as broad, ending below antennae in a short, arched,

transverse carina. Clypeus flat, superficially finely punctate, median part of

anterior margin raised, widely emarginate, lateral teeth large and protruding in

lateral view. In frontal view a median tooth on a lower level is visible (Fig. 21).

Frons shining, depressed medially. Vertex shining, finely punctate, a deep oblique

groove along lateral ocelli. Occipital carina ending in hypostomal carina.

Mandibles with tooth near upper base, apex bidentate, lower teeth blunt (worn

off?). Fore margin of labium blunt, laterally with tooth. Antennae short, clavate,

third segment about three times as long as broad at apex, following segments

gradually decreasing in length, segments 9—11 about as long as broad at apex, last

segment about 1% times as long as broad at base.

Pronotal angles with small tooth. Scutum shining, prescutal sutures sharp and

narrow, reaching hind margin of scutum, apically slightly widened and deeper.

Enclosed area of propodeum triangular with distinct median carina, lateral carinae

oblique. Back of propodeum smooth, longitudinal groove narrow. Anterior

oblique suture narrow, widened upper part smooth. Legs rather thick, base of hind

tibiae flattened and with thin spine about twice as long as width of base of tibia.

On outer side of hind tibiae a few stout reddish thorns. Petiole about 1% times as

long as first tergite, cylindrical, flattened dorsally, base with lateral ridge. Pygidial

area elongate-triangular, apex rounded.

J. P. van LITH: Indo-Australian Psenini 95

Face with silvery, mostly appressed, pubescence, rest of body yellowish-grey

pubescent, pubescence on mesosternum and sixth sternite more dense and golden.

Petiole laterally with long erect hairs.

Male unknown.

Philippine Is.: 19, holotype, Misamis Or., Dinawihan Gingoog, 26 km E. of

Gingoog City, 100—300 m, 15 August 1965, H.M. Torrevillas, Malaise trap

(BISH).

P. armipes belongs to the group of P. scutatus. It differs from P. scutatus in colour

as well as in the shape of the clypeal margin.

Psenulus tectus sp. nov.

Female (holotype). — Length about 8.5 mm. Head black, margin of clypeus

somewhat reddish transparant. Basal #, of mandibles yellow, apex dark reddish.

Labrum reddish-brown. Palpi yellowish-brown. Scape of antennae yellow, apex

brown on inner side; flagellum (missing after third antennal segment) brown

above, yellowish-brown below. Thorax black, following parts yellow: dorsal side of

pronotum, pronotal tubercles, broad line along lateral sides, posterior ?/, of inter-

prescutal area, axillae, scutellum except a narrow brown mark along posterior

margin, metanotum, propodeum except a broad black band above (with two

lateral downward projections) and a narrow band at apex, anterior plate of

mesepisternum, a vague large spot on mesopleura below, behind anterior oblique

suture, a small spot below fore wings and a larger one below hind wings. Petiole

yellowish-red, gaster pale red. Fore and mid legs yellow, back of femora more or

less brown, hind legs yellowish-red with brown tarsi. Tegulae pale yellow, veins of

wings brown.

Head large. Frontal carina raised, broadened and excavate between antennae,

ending below antennae in a transverse carina (Fig. 23). This transverse carina is

situated on a raised area which in dorsal view is protruding like a sloping roof (Fig.

25); in lateral view this area extends far beyond the surface of the clypeus (Fig.

24). Protruding median part of anterior margin of clypeus broad, indistinctly

tridentate, disk of clypeus superficially punctate. Frons and vertex shining, almost

impunctate, tempora slightly dull, finely punctate. Occipital carina ending in

hypostomal carina. Apex of mandibles bidentate. Third antennal segment over

three times as long as broad at apex.

Pronotal corners obtuse. Scutum finely aciculate, very finely and sparsely

punctate. Prescutal sutures narrow, indistinctly foveolate, reaching hind margin.

Scutellum and metanotum shining, almost impunctate. Enclosed area of pro-

podeum triangular, median longitudinal groove on back of propodeum narrow,

back laterally with shallow, finely transversely striate groove. Sides of propodeum,

mesopleura and mesosternum shining, almost impunctate. Anterior oblique suture

narrow, foveolate, upper part striate. Petiole cylindrical. nearly twice as long as

first tergite, a short groove at apex. Pygidial area elongate-triangular, apex blunt

(Fig. 26). Femora rather thick, base of hind tibiae with smooth narrow groove,

margined on inner side by a narrow high carina, base also with long thin spine

which is twice as long as width of base. First recurrent vein ending in first

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

submarginal cell, second recurrent vein in third submarginal cell.

Face with silvery, mostly appressed pubescence. Pubescence of rest of body

yellowish-grey. Petiole dorso-laterally, laterally and ventrally with a row of long

erect hairs.

Male unknown.

Fig. 23-26. Psenulus tectus sp. nov., 9, holotype, 23—25, head in frontal, lateral and dorsal aspect; 26,

pygidial area. Fig. 27—29. Psenulus anomalus sp. nov., Q, holotype, 27, thorax; 28, head in dorsal

aspect; 29, pygidial area

J. P. vAN LITH: Indo-Australian Psenini 97

Laos: 19, holotype, Savannakhet Prov., Savannakhet, 15 April 1967, native

collector (BISH).

P. tectus is one of the first two representatives on the Asiatic continent of the

group of Psenulus scutatus, thus far known from the Philippine Islands, Borneo

and Java. It is easily distinguished from the other species of this group by the roof-

like protuberance below the antennae.

Psenulus anomalus sp. nov.

Female (holotype). — Length about 6.5 mm. Head black; mandibles yellow with

dark tips, palpi yellowish-brown. Dorsal half of scape brown on inner side,

flagellum dorsally brown, yellowish below. Thorax (Fig. 27) black with following

parts yellow: dorsal side of pronotum, pronotal tubercles, lateral margins of

scutum, a large mark between prescutal sutures near hind margin, scutellum

except a narrow brown mark along posterior margin, a very small spot on axillae,

metanotum except narrow brown hind margin, back of propodeum except broad

basal band and narrower apical margin (sides of propodeum black), upper third of

anterior plate of mesepisternum. Fore and mid legs yellow, underside of fore

femora brown, hind legs reddish. Petiole black, rest of gaster including ventral

plate reddish. Veins of wings brown.

Head large. Frontal carina raised and excavate between antennae, ending below

antennae in a projecting short transverse carina (Fig. 28). Clypeus shining,

indistinctly sculptured on surface. Anterior margin of clypeus somewhat emar-

ginate, left part deformed in the type, probably with large, blunt median tooth.

Frons and vertex smooth and shining. Tempora dull. Occipital carina ending in

hypostomal carina. Antennae short, clavate, segments 11-12 nearly twice as broad

as long. Mandibles with blunt apex.

Pronotal corners obtuse. Scutum very finely aciculate with some fine punctures.

Prescutal sutures narrow, finely foveolate, reaching hind margin. Between

prescutal sutures before hind margin a number of short rugae. Parapsidal sutures

long. Scutellum and metanotum smooth and shining. Enclosed area of propodeum

rather narrow, the small central area connected with fore margin by a short carina

(petiolate). Back and sides of propodeum smooth, median sulcus narrow, between

back surface and sides a narrow area of fine irregular carination. Mesopleura

smooth and shining, mesosternum more distinctly punctate. Anterior oblique

suture foveolate, also widened upper part. Legs rather thick, basal ?/; of hind tibiae

with smooth groove, margined on inner side by a sharp carina, base of this groove

with a narrow spine about twice as long as width of base. First recurrent vein of

fore wings interstitial, second recurrent vein ending well in third submarginal cell.

Petiole almost cylindrical, nearly 1% times as long as first tergite, a small

triangular depression at apex. First tergite longer than broad at apex. Gaster very

finely aciculate, indistinctly punctate. Pygidial area (Fig. 29) triangular.

Pubescence of face and tempora silvery, mostly appressed. Pubescence of rest

of body whitish, dense on mesosternum; epicnemial areas below with a small,

densely pubescent spot. Petiole with long lateral and ventral erect hairs, dorso-

laterally a row of fine short hairs, intermixed with a few long hairs.

Male unknown.

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Vietnam: 19, holotype, 15—35 km NW. of Phan Rang, 8—16 Nov. 1960, C.M.

Yoshimoto (BISH).

Although the clypeal margin is deformed, the other characteristics, especially

the shape of the pygidial area justify the naming of this form as a distinct species.

P. anomalus is the first representative of the group of P. scutatus in Vietnam and

the second known species of this group on the Asiatic continent.

Psenulus quadridentatus Van Lith

Van Lith, 1962: 37—38 (Malaya); 1972: 162 (Vietnam); 1973: 133 (Nepal).

New record from Malaya: 19, Pahang, Cameron Highlands, 5000 ft, 29 April

1973, C.G. Roche (CGR). |

Psenulus chillcotti Van Lith

Van Lith, 1973: 133—136 (Nepal).

First record from Burma: 19, NE. Burma, Kambaiti, 7000 ft, 1 May 1934, R.

Malaise (BM).

Psenulus vaneuensis sp. nov.

Male (holotype). — Length 5—6 mm. Head black; labrum reddish, palpi

yellowish-brown, flagellum of antennae dark brown, underside and scape reddish-

brown. Thorax black, pronotal tubercles yellowish-red. Legs including trochanters

and apex of coxae reddish, underside of fore and mid femora brown. Gaster

including petiole red.

Clypeus convex, surface densely superficially punctate, anterior margin biden-

tate. Interantennal carina broadened, this part more than twice as long as wide,

carina ending in a transverse carina below antennae. Frons and vertex shining,

superficially finely punctate, tempora more densely punctate. Occipital carina

ending in hypostomal carina. Antennae moniliform, segment 3 nearly twice as long

as broad at apex, following segments gradually decreasing in length, segments

8—11 about as long as greatest width, segment 12 slightly longer than broad,

segment 13 nearly twice as long as broad at base. Segments 3—12 with distinct,

dark, oblong tyloidea, about twice as long as broad, small on segment 3.

Pronotal corners obtuse. Scutum shining, densely, mostly finely, punctate.

Prescutal sutures foveolate, about as long as anterior % of scutum. Scutellum

shining, finely punctate. Metanotum dull, densely minutely punctate. Enclosed

area of propodeum rather narrow, lateral parts with oblique carinae. Back of

propodeum dorsally smooth and shining, declivous part finely punctate, apex with a

few rugae, dorso-laterally a broad, coarsely foveolate groove. Posterior part of

sides of propodeum irregularly carinate. Mesopleura shining, finely punctate,

interstices a few times size of punctures. Hypo-epimeral area minutely punctate.

Mesosternum densely finely punctate. Anterior oblique suture broad, foveolate,

also widened upper part. Petiole thick, about as long as first tergite, dorsally

J. P. vAN LITH: Indo-Australian Psenini 99

convex, ventrally keeled, sides somewhat depressed. Gaster shining, minutely

punctate.

Genitalia (Fig. 30) yellowish-brown, apex of basiparameres on inner side

projecting as a low tooth, apex of parameres rounded.

Pubescence greyish-white.

Female unknown.

Laos: 14, holotype, Vientiane Prov., Ban Van Eue, 15 May 1966, native

collector (BISH).

P. vaneuensis seems to be closely related to P. chillcotti, which also belongs to

the group of P. quadridentatus. The genitalia, however, are different (cf. Van

Lith, 1973).

Psenulus puncticeps (Cameron)

Cameron, 1907: 91 (Psen puncticeps; India).

Rohwer, 1923: 595—596 (Diodontus antennatus; Singapore).

Van Lith, 1962: 44—46 (Psenulus antennatus; Malaya, Java, Bali); 1973: 136—137 (Psenulus

puncticeps; Nepal).

First records from Laos: 1 9, Sayaboury Prov., Sayaboury, 17 Febr. 1965, native

collector, 19, Vientiane Prov., Ban Van Eue, 750 m, forest streambed, 10-11

April 1965, J.L. Gressitt, Malaise trap (BISH).

In these females the pronotal tubercles are yellowish, the petiole is reddish-

brown or dark brown, with reddish apex and ventral plate.

First record from Flores: 19, C. Flores, Moni, Wolowaru, 11 Nov. 1949, Bühler

and Sutter (NMB).

This female is rather dark. Pronotal tubercles blackish-brown on fore part,

yellowish-brown behind. First tergite almost entirely black, also petiole with

ventral plate. Apical % of hind tibiae brownish-black, mid tibiae slightly brownish.

Genitalia of a male from Malang, Java: Fig. 31, 32.

Psenulus puncticeps rufipes subsp. nov.

A male from West Sumba much resembles the males of the nominate form from

Java and Bali. It differs in having the hind legs including trochanters completely

reddish and there is no yellowish ring on the base of the hind tibiae. Fore and mid

legs yellowish-red, femora brownish below. Ventral plate of petiole and first tergite

reddish. Posterior half of pronotal tubercles brownish.

Antennal segments less rounded than in the nominate form, segments 3—10

with small but distinct tyloidea. The males from Java and Bali have indistinct

tyloidea on segment 4—9, which have not been mentioned earlier.

W. Sumba: | 3, holotype, Pogobina, 16 Sept. 1949, Bühler and Sutter (NMB).

Psenulus exiguus sp. nov.

Female (holotype). — Length about 6 mm. Head and thorax black, mandibles

dark brown, palpi brown. Antennae dorsally dark brown, ventrally yellowish-

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Fig. 30—36. Male genitalia of various species of Psenulus. 30, P. vaneuensis sp. nov., holotype, dorsal

aspect; 31 —-32, P. puncticeps (Cameron), Java, dorsal and ventral aspect; 33, P. crabroniformis wapiensis

subsp. nov., paratype; 34, P. exiguus sp. nov., paratype, Malaya; 35—36, P. suturalis sp. nov., holotype,

lateral and dorsal aspect

J. P. van LITH: Indo-Australian Psenini 101

brown. Pronotal tubercles brownish-yellow. Femora brown, tibiae and tarsi

yellowish-brown. Petiole dark brown. Gaster dark reddish, first tergite and apical

half of tergites 2—3 and base of fourth tergite blackish (irregular, discoloured?).

Raised part of interantennal carina broadened and excavate. No distinct

transverse carina. Median part of clypeal margin with two small teeth. Frons and

vertex shining, finely but distinctly, rather densely, punctate. Tempora finely

striate. Occipital carina ending in hypostomal carina. Mandibles tridentate at

apex. Antennae short, clavate, segments 6—10 shorter than broad at apex, last

segment about 1% times as long as broad at base.

Pronotal corners obtuse. Scutum shining with some irregular, mostly fine,

punctation. Prescutal sutures indicated on posterior part of scutum by a few

irregular rows of punctures. Hind margin of scutum with short rugae. Enclosed

area of propodeum narrow, central area and sulcus on back of propodeum rather

wide, sulcus with some transverse carinae. Back of propodeum shining, laterally

an irregularly carinate edge. Mesopleura shining, almost impunctate. Anterior

oblique suture narrow, foveolate, widened upper part with a few transverse

carinae. Legs normal. First recurrent vein of fore wings interstitial, second

recurrent vein ending in third submarginal cell. Petiole subquadrate in cross-

section, about 1% times as long as first tergite. First tergite about as long as broad

at apex, tergites densely finely punctate. Pygidial area indistinct.

Pubescence of face, tempora and pronotum silvery, mostly appressed. Thorax

whitish pubescent, mesosternum very densely so, also a small round patch on

epicnemial areas below. Gaster yellowish pubescent.

Male. — Resembling female. Length 6—6.5 mm. Tergites 1, 3 and following

brown. Apical outer half of hind tibiae brown. Normal transverse carina below

antennae. No distinct tyloidea. Back and dorsal part of propodeum with fine

oblique striation.

Genitalia (Fig. 34) yellowish-brown, parameres largely transparent. Apex of

basiparameres with inconspicuous small tooth, apex of parameres rounded.

Laos: 19, holotype and 13, allotype, Sayaboury Prov., Sayaboury, 13 April

1966, native collector (BISH).

Malaya: 1g, paratype, Selangor, Ulu Langat, 300—390 m, 13 June 1958, T.C.

Maa (BISH).

The male from Malaya is slightly different in colour. Petiole and first tergite,

apical % of tergite 2, apical half of tergite 3 and tergites 4—7 dark brown. Only

back of propodeum with oblique striae. |

P. exiguus belongs to the group of P. puncticeps. It differs from P. puncticeps

which has been taken at the same locality in Laos, by the small size and the

extension of the dark brown colour.

Psenulus suturalis sp. nov.

Male (holotype). — Length about 8.5 mm. Head black; mandibles dark reddish,

palpi brown. Antennae dark brown, reddish-brown below. Thorax black, hind

margin of pronotal tubercles and tegulae reddish-brown. Foreside of fore and mid

femora, whole fore and mid tibiae and tarsi and greater part of hind tibiae reddish-

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

brown, rest of legs brown. Veins of wings dark brown. Petiole black, gaster

including ventral plate of petiole reddish.

Raised part of interantennal carina narrow, ending below antennae in a long

transverse carina. Clypeus densely superficially punctate, anterior margin with two

small teeth. Frons, vertex and tempora smooth and shining. Occipital carina

ending in hypostomal carina. Antennae long, third segment more than twice as

long as broad at apex, segments 4—12 about twice as long, segment 13 more than

twice as long as broad at base. No tyloidea, segments 4—10 with indistinct

tubercles.

Pronotal corners about rectangular. Scutum, scutellum and metanotum shining,

sparsely finely punctate. Prescutal sutures almost reaching hind margin, broad,

foveolate. Enclosed area of propodeum and sulcus on back of propodeum deep.

Behind enclosed area of propodeum some indistinct longitudinal striation, back

and sides of propodeum very coarsely reticulato-carinate. Mesopleura and

mesosternum smooth and shining, sparsely minutely punctate. Anterior oblique

suture broad and coarsely foveolate, also widened upper part. Legs normal,

slender. First recurrent vein of fore wings ending in first submarginal cell, distance

from second submarginal cell about % of abscissa of radius in the second cell.

Second recurrent vein ending in third submarginal cell. Petiole broad, widening

towards apex, slightly longer than first tergite. Petiole dorsally with two parallel,

broad, rounded carinae, laterally with deep groove, ventral side flattened. First

tergite longer than broad at apex. Tergites shining, sparsely finely punctate.

Genitalia (Fig. 35, 36) long and narrow, reddish-brown, broadened part of

parameres almost black. Parameres on inner side basally broadened into a large

triangular tooth, parameres towards apex gradually broadening, apex with small

tooth on inner side. Between basiparameres and parameres a high, rounded ridge

on dorsal side.

Pubescence of face silvery-white, mostly appressed. Tempora with long whitish

pubescence. Vertex and thorax with greyish-brown pubescence, dense on

mesosternum. Gaster with yellowish-brown pubescence. Petiole dorso-laterally,

laterally and ventrally with long erect hairs.

Female unknown.

Laos: 1 à, holotype, Vientiane Prov., Ban Van Eue, 15—31 May 1965, native

collector (BISH).

P. suturalis is distinguished by the coarse and long prescutal sutures, the

structure of the petiole and the genitalia. It does not fit well into any of the Indo-

Australian species-groups described thus far.

Psenulus erraticus basilanensis (Rohwer)

Rohwer, 1921: 318—319; 1923: 595 (Diodontus basilanensis; Basilan, Singapore).

Van Lith, 1962: 49— 50 (Psenulus erraticus basilanensis); 1970: 95—96 (Tawi Tawi).

New record from Singapore: | &, University of Singapore, Campus, 8 May 1972

(US).

Pubescence of face silvery, of frons, thorax and gaster yellowish-golden (fresh

J. P. van LITH: Indo-Australian Psenini 103

specimen), hairs long on metanotum. Petiole with lateral row of short hairs, sides

with long hairs directed obliquely downwards.

This is the second known male of P. basilanensis. Both have been collected in

Singapore.

Psenulus crabroniformis crabroniformis (Smith)

Smith, 1858: 107 (Mellinus crabroniformis; Borneo).

Van Lith, 1962: 51—53 (Psenulus crabroniformis); 1972: 168—170 (Thailand).

Tsuneki, 1974: 637—638 (Malaya).

New record from Malaya: 1 9, Pahang, Genting Serpah, 2100 ft, 2 Dec. 1973, C.

G. Roche (CGR).

New record from Thailand: | 9, 50 km W. of Tak, 900 m, 7—8 April 1966, J.

and M. Sedlacek (BISH).

First records from Laos: 6 9, Sayaboury Prov., Sayaboury, 12 Febr., 2 March

and 13 April 1966, two in Malaise trap; 2 9, Vientiane Prov., Ban Van Eue, 29

March and 31 May, 1966, native collector (BISH).

The petiole is reddish, base somewhat paler. In the female from Pahang the

sides of the petiole are slightly darkened apically. OOD: POD = 9:7 (OOD is

distance between oculi and ocelli, POD is distance between posterior ocelli).

Psenulus crabroniformis wapiensis subsp. nov.

Four males from Laos differ from the nominate form (cf. Van Lith, 1962) in the

sculpture of the thorax. The punctures are strong, the interstices often smaller

than the punctures; in two specimens there are short transverse rugae across the

prescutal sutures. The mesopleura are distinctly punctate, interstices mostly larger

than punctures. In one specimen (15 March 1967) the apical half of the petiole is

dark brown.

Genitalia (Fig. 33) with slender parameres, deeply emarginate at apex and with

some long hairs at apex. |

Female unknown.

Laos: 2 3, holotype and paratype, Wapikhamthong Prov., Wapi, 30 March

1967, 2 6, paratypes, 15 March and 15 April 1967, same locality, native collector,

light trap (BISH).

Psenulus crabroniformis nathani subsp. nov.

Female (holotype). — Differs from the nominate form from Thailand and

Malaya by the darker petiole (more than apical half black, base yellowish), the

almost entirely black instead of reddish hind femora and the dorsally dark brown

apical half of the hind tibiae.

Pubescence of face and tempora silvery-white, of vertex greyish-brown, of

thorax greyish or whitish. OOD : POD = 9:7.

Male unknown.

South India: | 9, holotype, Anamalai Hills, Cinchona, 3500 ft, May 1967, P.S.

Nathan (MA).

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Psenulus crabroniformis sumatranus (Ritsema)

Ritsema, 1880: 225—226 (Psen sumatranus; Sumatra).

Van Lith, 1962: 53—54 (Psenulus crabroniformis sumatranus; Sumatra, Java).

New records from West Java: | 9, Mt. Gedogan, Djampang Tengah, Oct. 1937,

1 9, Mt. Melang, Djampang Wetan, 3—7000 ft, Dec. 1937, 1 9, Mt. Malang,

3—4000 ft, K. M. Walsh (BM).

In the Javanese form OOD: POD appears to be 10.5: 7.

Psenulus philippinensis philippinensis (Rohwer)

Rohwer, 1921: 317—318 (Diodontus philippinensis, Luzon).

Van Lith, 1962: 54—56 (Psenulus philippinensis philippinensis).

New record: | 9, Philippine Is., Luzon, Mt. Limay, Baker (USNM).

Thus far the nominate form is restricted to Luzon, where in total nine females

have been collected.

Psenulus philippinensis dapitanensis (Rohwer)

Rohwer, 1923: 594—595 (Diodontus dapitanensis; Mindanao).

Van Lith, 1962: 56 (Psenulus philippinensis dapitanensis; Mindanao, Bohol).

New records from Philippine Is.: 1 9, Biliran, Baker (USNM); 1 3, Negros,

Cuernos Mts, Baker (USNM).

First description of male. — Length about 8.5 mm. Resembling female. Face

with golden pubescence. Pubescence of thorax golden-brown. Hind femora

dorsally not brownish, as in the female from Biliran, but all legs entirely yellowish-

red, also antennae except scape which is more yellowish, flagellum dorsally not

brownish, as in female.

Antennal segments 3—10 about 1% times, segments 11—12 about 1% times as

long as broad at apex, last segment twice as long as broad at base. No tyloidea.

Scutum resembling that of nominate subspecies, but with tendency to transverse

rugosity.

Genitalia (Fig. 37—38) slender, yellowish-brown. Parameres with small trian-

gular flag about halfway inner side, apex with large tooth on inner corner and a

few long hairs. A few bristles on underside of parameres near apex. Apex of

seventh sternite with small triangular emargination. Apical margin of sixth sternite

(Fig. 40) very finely serrate. Eighth sternite Fig. 39.

Notwithstanding the sculpture of the scutum I consider this male to belong to

dapitanensis, mainly because of the blackish apical % of the petiole.

Psenulus maculatus maculatus Van Lith

Van Lith, 1962: 61—62 (Malaya).

New record: | ©, Singapore, Changi Beach, 7 July 1972 (US).

J. P. van LITH: Indo-Australian Psenini 105

—

SA TIZI

pesi

& fi

Fig. 37—40. Psenulus philippinensis dapitanensis (Rohwer), 3, Negros, 37—38, genitalia, dorsal and

lateral aspect; 39, 8th sternite, lateral aspect; 40, 6th sternite, lateral aspect. Fig. 41—42. Psenulus

trimaculatus Van Lith, 3, paratype, 41, basiparameres and parameres, dorsal aspect; 42, pronotum and

scutum. Fig. 43. Psenulus varius Van Lith, 9, thorax, dorsal aspect. Fig. 44. Psenulus limbatus sp. nov.,

©, holotype, thorax, dorsal aspect

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976.

Psenulus trimaculatus Van Lith

Van Lith, 1962: 80—81 (Malaya).

New records: 3 3, Singapore, Bukit Timah Forest, 26 April 1973, D. H. Murphy

(US); | &, Malaya, Kepong, 130—300 m, 13—21 March 1966, light trap, J.

Sedlacek (BISH). i

These males are darker than the holotype from Penang. The lateral marks on

the scutum reach the black hind margin (Fig. 42), the median mark is connected

with the hind margin by a narrow or indistinct line. Axillae black, with smaller or

larger yellow spot. Third antennal segment over twice as long as broad in frontal

view, segments 4—6 about twice, segments 7—10 about 1% times, segments

11-12 about twice as long as broad at apex, last segment about 2% times as long

as broad at base. No tyloidea.

Genitalia (Fig. 41) straw-yellow, apex straight.

The marking of the scutum much resembles that of P. peterseni Van Lith, from

the Philippine island Tawi-Tawi, of which only females are known. However,

peterseni has the anterior plate of the mesepisternum largely yellow instead of

entirely black, whilst the hypo-epimeral area is partly black, instead of entirely

yellow as in the male of P. trimaculatus. Also the petiole of P. peterseni is different.

Both species belong to the group of P. sandakanensis.

With regard to the three males from Singapore I received the following note

from Dr. D. H. Murphy, Singapore: “Specimens of Psenulus reared from nests in

vertical dead sapling in heavily shaded forest valley near small stream. Nests were

in holes apparently made by larvae of Therates dimidiatus Dei. (Coleoptera:

Cicindelidae) and several similar holes were still occupied by Therates larvae. Prey

residues from one chamber were examined and contained recognisable fragments

of Aphididae only, including siphons probably belonging to Greenidoidea sp. The

forest belongs to “Tropical Moist Forest” (Holdridge classification). Nests were

between 3 and 4 feet above ground level.”

Dr. Murphy kindly also sent me one of the nests. It contains three empty cells,

each about 8 mm long, diameter over 3 mm. The head end of the cells is closed by

a brown lid, on the outer side with some fine white silk threads; the inner side is

lined with a white layer which continues for about | cm on the sides. Between the

first and second cell and between the second and third cell there is also a thin

partition of macerated pith.

Psenulus varius Van Lith

Van Lith, 1962: 69—70 (Malaya); 1972: 176—177.

New records: 5 9, Malaya, Penang, Tanjong Bungah, Sungei Kechil, 21—24

Jan. 1964, | 9, Malaya, Penang, Penang Hills, Sungei Pinang, 16 May 1964, H. T.

Pagden (BM).

There is some variation in the marking of the scutum (Fig. 43). In five of the six

females the narrow line which connects the median mark with the hind margin is

more or less indistinct, interrupted or even absent. The black mark of the

J. P. van LITH: Indo-Australian Psenini 107

scutellum is either triangular or reduced to a median longitudinal mark.

The first recurrent vein of the fore wings is sometimes almost interstitial.

Psenulus limbatus sp. nov.

Female (holotype). — Length about 8 mm. Head black; apical margin of

clypeus and labrum reddish, mandibles except apices yellow, palpi yellowish.

Scape of antennae and underside of segments 2 and 3 yellow, rest of underside of

flagellum yellowish-red. Thorax black except the following parts. Dorsal side of

pronotum and pronotal tubercles yellow. Scutum yellow with three broad black

bands reaching fore and hind margin (Fig. 44). Axillae yellow, scutellum with

yellow hind margin, interrupted in the middle. Metanotum, back of propodeum,

upper % of anterior plate of mesepisternum and a vague spot on mesopleura near

foreside yellow. Fore and mid legs and hind coxae yellow, hind femora and tibiae

yellowish-red with brownish outer side, hind tarsi brown. Gaster including petiole

yellowish-red. Veins of wings brown.

Anterior margin of clypeus hardly emarginate, with two small teeth; distance

between these teeth about ¥, of total distance there between the eyes. Interanten-

nal carina dorsally sharp, ending below antennae in a transverse carina. Frons,

vertex and tempora shining, almost impunctate. Postocellar area not raised.

Occipital carina ending in hypostomal carina. Third antennal segment about 2%

times, segments 4—6 about twice, segments 7—11 about 1% times as long as broad

at apex, segment 12 about 2% times as long as broad at base.

Scutum, scutellum and metanotum shining, distinctly finely punctate. Enclosed

area of propodeum triangular, dorsal part of pronotum shining, back and posterior

half of sides of propodeum densely finely reticulate, median groove narrow.

Mesopleura, hypo-epimeral area, anterior plate of mesepisternum and mesoster-

num shining, indistinctly punctate. Legs slender. First recurrent vein of fore wings

interstitial. Petiole cylindrical, little longer than first tergite. Pygidial area long and

narrow, apically with almost parallel carinae.

Pubescence of face silvery, mostly appressed, of rest of body whitish.

Laos: | 9, holotype, Vientiane Prov., Gi Sion Vill., de Tha Ngone, 21—28 Febr.

1966, native collector (BISH).

Because of the transverse carina below the antennae P. limbatus has been placed

in the group of P. esuchus. It resembles somewhat the male of P. fyanensis from

Vietnam and Laos. However, fore and mid femora and tibiae are entirely yellow,

the anterior plate of the mesepisternum is more distinctly yellow, and the

propodeum is less dark. Certainty regarding the status of these two forms can be

obtained only when the opposite sexes are found.

Psenulus impressus sp. nov.

Male. — Length about 6 mm. Head black; mandibles yellow except dark tips,

palpi pale yellow. Scape of antennae yellow, flagellum yellowish-brown below,

dark brown above. Thorax yellow with narrow black mark along hind margin of

scutum, this mark anteriorly roundly extended. Black parts: posterior corners of

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

scutum, anterior and lateral depressed parts of scutellum, enclosed area of

propodeum and median longitudinal sulcus on back of propodeum, a small mark

below fore wings, interepicnemial area and a narrow median line on mesosternum.

Fore and mid legs including coxae and hind coxae yellow, rest of hind legs

reddish-yellow, gaster including petiole reddish, apical spine brown. Veins of wings

dark brown.

Clypeus superficially punctate, margin depressed, slightly triangularly emar-

ginate, two short triangular teeth. Raised part of interantennal carina narrow,

ending below antennae in a transverse carina. Frons and vertex shining, with very

fine and dense, hairbearing punctation. Interocellar area much raised, bare and

shining. Upper part of back of head sloping obliquely downward from hind margin

of posterior ocelli to occipital carina. Occipital carina ending in hypostomal

carina. Antennae long and slender, segments 3—12 about twice as long as broad at

apex, last segment about 2% times as long as broad at base. No tyloidea.

Pronotal corners obtuse. Scutum with large superficial punctures, interstices

mostly smaller than punctures. Prescutal sutures distinct on anterior half of

scutum. Area around median scutal lines and parapsidal sutures impressed, which

is best seen in anterior aspect. Scutellum and metanotum sparsely superficially

punctate. Propodeal enclosed area with large central hectagonal area, lateral parts

with few oblique carinae. Back of propodeum coarsely reticulato-carinate, upper

part finely obliquely striate. Anterior oblique suture narrow, foveolate, widened

upper part smooth. Legs slender. First recurrent vein of fore wings ending in

second submarginal cell, nearly interstitial, second recurrent vein ending in third

submarginal cell. Petiole cylindrical, longer than first tergite, gaster slender.

Pubescence on face, tempora and frons silvery, mostly appressed on clypeus, on

ventral side of thorax dense, whitish, on rest of head and body yellowish-grey or

yellowish-golden.

Genitalia (Fig. 46) small, pale brownish-yellow. Parameres dark brown, largely

transparent, apex blunt.

Female unknown.

Philippine Is.: 1 &, holotype, Mt. Montalban, Rizal, Wa-wa Dam, 150—200 m, 2

March 1965, L. M. Torrevillas, 2 &, paratypes, same locality, 3 and 14 March

1965, H. M. and L. M. Torrevillas (BISH).

Psenulus interstitialis interstitialis Cameron

Cameron, 1906: 222—223 (Psenulus? interstitialis; New Guinea).

Van Lith, 1962: 84—86 (Psenulus interstitialis interstitialis); 1972: 182—184 (New Guinea, Papua).

As stated earlier (Van Lith, 1972), this is a variable form which is also shown by

the undermentioned specimens.

Lower half of anterior part of pronotum usually black, in some specimens

entirely yellow. Scutum entirely yellow or with central square black mark.

Antennal flagellum sometimes completely yellowish-red, usually dorsally more or

less darkened. There does not seem to be any correlation between the varieties

and their collecting localities; males from Mamai Plantation have either an

J. P. van LITH: Indo-Australian Psenini

109

Fig. 45—47. Male genitalia of various new species of Psenulus. 45, P. vientianensis, holotype; 46, P.

impressus, paratype; 47, P. fyanensis, holotype. Fig. 48—50. Thorax in dorsal aspect of various new

species of Psenulus. 48, P. fyanensis, 3, holotype; 49, P. nigrolateralis, 9, holotype; 50, P. pictus, 3,

paratype

entirely yellow scutum and pronotum or scutum with black mark and lower part of

pronotum black.

New records from NE. New Guinea: | 9, May River, 6 June 1963, R.

Straatman, light trap; 1 9, Wau, Bulolo River, 850—900 m, 24 Aug. 1965, J.

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Sedlacek, Malaise trap; | 3, Bulolo, 750 m, 5 Oct. 1965, J. Sedlacek, Malaise trap;

I 8, Wau, 1400 m, 27 Sept. 1969, A. B. Mirza; | 3, Sepik, Angoram, 20—30 m,

14—16 Aug. 1969, J. L. Gressitt; 1 9, Baiyer River, 1000 m, 1—4 Sept. 1969, Y.

Hirashima (BISH).

SE. New Guinea: | 9 and 3 3, Mamai Pltn, E. of Port Glasgow, 150 m, 16 Febr.

1965; 2 3, same locality, 27 Febr. 1965, 1 &, Milne Bay, abt. 10 m, March 1965, all

R. Straatman, light trap; 1 3, Cape Killerton, 0—5 m, 6—13 May 1965, W. A.

Steffan, Malaise trap; 1 9, Central Dist., 3.2 km S. Vanapa River, Brown Road,

22—24 May 1965, W. A. Steffan and Y. M. Huang (BISH).

Papua: | 9, Laloki Exp. Station, 10—17 June 1971, J. Stibick; 1 &, N. Dist.,

Onombususu, 26—28 Aug. 1971, J. Stibick (HT). Both specimens from Papua have

an entirely yellow scutum.

Two females from NE. New Guinea, Green River - Sepik River junction, 200 m,

22 June and 27—28 June 1965, collected by R. Straatman with light trap (BISH)

have yellowish-red thorax, antennae and fore and mid legs and resemble the

subspecies salomonensis (Van Lith, 1972). As there are a few minor differences

their identification is still uncertain.

Psenulus pictus sp. nov.

Five males, closely related to P. interstitialis, are much more richly marked with

black, especially on scutum and mesosternum.

Black parts: a large longitudinal mark on scutum (Fig. 50) connected by a

narrow line with triangle against posterior margin and two narrow lateral marks,

covering the parapsidal sutures and connected with hind margin of scutum,

enclosed area of propodeum with adjacent lateral spots on declivous part, median

longitudinal groove and apical margin of declivous part, depressed lateral parts of

scutellum and metanotum, epicnemium except lateral margin, mesosternum,

marking roundly dilated on to mesopleura and connected with broad hind margin

of mesopleura, which also covers hind margin of hypo-epimeral area.

Fore and mid legs including coxae and trochanters and hind coxae yellow, tarsi

of fore and mid legs and hind legs except coxae orange-red. Gaster and petiole

orange-red. Antennal segments 3—13 dorsally black, underside orange-red, often

brownish on apical half. Veins of wings yellowish-brown.

Black marks on mesopleura in some specimens somewhat reduced. Lateral

marks on scutum, in one case also median mark, not connected with hind margin.

Length about 8.5 mm.

Various species belonging to the group of P. interstitialis are subject to much

variation, especially P. interstitialis, but no transitional forms have been found at

Wau, where also P. interstitialis occurs. After some hesitation I decided to

consider the described males as a distinct species. The study of more material,

especially of the female, should confirm this opinion.

NE. New Guinea: 4 3, holotype and paratypes, Wau, McAdam Park, 1500 m,

17 Sept. 1972, flying at Hibiscus, Mrs. E. van der Vecht (ML); 1 3, Wau, 1250 m, 4

May 1965, J. and M. Sedlacek (BISH).

J. P. VAN LITH: /ndo-Australian Psenini 111

Psenulus araucarius Van Lith

Van Lith, 1972: 187 (Central New Guinea).

This close relative of Psenulus interstitialis has now also been collected in NE.

New Guinea: | 9, Mt. Missim, 980 m, 20 July 1969, J. L. Gressitt and Y.

Hirashima (BISH).

The third antennal segment is over three times as long as broad at apex, the

pygidial area is broader than in P. interstitialis and the base of the hind tibiae is

smooth and flattened. This is the second known female, the male has not yet been

found.

Psenulus ornatus sumbaensis subsp. nov.

Female (holotype). — Length about 9 mm. Head black, labium reddish,

mandibles yellow with dark tips, palpi yellowish. Scape of antennae yellow,

flagellum reddish below, dark brown above. Thorax black with following yellow

parts: dorsum of pronotum, pronotal tubercles, two partly indistinct lines on

scutum, slightly broadened posteriorly and not reaching hind margin, (an indistinct

brownish mark along lateral margins), two elongate marks along hind margin

of scutellum, metanotum except small triangular mark on hind margin, two large

marks on back of propodeum, each consisting of a pair of marks broadly

connected below. Tegulae reddish. Fore and mid legs including coxae and

trochanters, and hind coxae, yellow; fore and mid tarsi and entire hind legs

reddish. Veins of wings brown. Gaster including petiole orange-reddish.

Raised part of frontal carina between antennae not broadened, no transverse

carina below antennae. Clypeus very finely punctate, lower part more shining,

margin depressed, protruding median part with two small teeth, distance between

these teeth about Y, of total distance there between the eyes. Frons and vertex

shining. Apex of mandibles bidentate, a distinct reddish-brown inner tooth.

Labium slightly emarginate. Antennae slightly clavate, third segment in frontal

view about three times, segments 4—6 over 1% times, segments 7—8 about 1%

times as long as broad at apex, segments 9—11 about as long as broad, last

segment about 1% times as long as broad at base. Occipital carina ends normally in

hypostomal carina.

Pronotal corners obtuse. Scutum, scutellum and metanotum shining, almost

impunctate. Prescutal sutures short and fine. Enclosed area of propodeum

triangular, ill-defined posteriorly, lateral parts with oblique striae. Propodeal

longitudinal groove narrow. Upper half of declivous part of propodeum smooth

and shining, lower half finely, mostly transversely, reticulato-carinate, sides more

coarsely so. Metapleura impunctate, mesopleura including hypo-epimeral area

and mesosternum shining, very finely sparsely punctate. Anterior oblique suture

foveolate, widened upper part smooth. Apex of mid tibiae anteriorly with

longitudinal ridge, posteriorly with three or four short reddish thorns. Base of hind

tibiae with short longitudinal area consisting of fine red thorns. Apex laterally

with three short red thorns. Inner tibial spur of hind legs bent downwards, about

30 degrees. First recurrent vein of fore wings interstitial, second recurrent vein

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

ending in third submarginal cell. Petiole about seven times as long as wide in the

middle, in dorsal aspect, one and a half times as long as first tergite, cylindrical,

dorsally somewhat flattened, a triangular pit at apex, sides with a narrow strip of

fine punctures. Gaster shining with fine hair-bearing punctures. Pygidial area

narrow, shining, with distinct lateral carinae.

Pubescence of head silvery, on clypeus somewhat appressed but leaving

sculpture visible. Pubescence of thorax greyish, on mesosternum whitish and

denser, lower part of epicnemial areas with a large round patch of appressed

whitish pubescence. Pubescence of gaster yellowish-grey. Petiole with long, erect,

yellowish-golden lateral hairs. Apical margins of gastral sternites 4—5 with dense

short pubescence.

Male unknown.

West Sumba: | 9, holotype, Rua, 1 Sept. 1949, Buhler and Sutter (NMB).

This female is undoubtedly closely related to P. ornatus Ritsema from East Java

of which only a single male is known. However, the petiole of sumbaensis has no

darkened apex and the legs are more reddish. P. ornatus baliensis (Van Lith, 1962)

is very similar and has also an entirely reddish petiole but the black marking on

dorsal and ventral side of its thorax is much less extensive.

Psenulus ornatus baliensis Van Lith

Van Lith, 1962: 88—89 (Psenulus interstitialis baliensis; Bali).

Tsuneki (1971: 3—5) places P. kankauensis Strand, 1915, P. tristis Van Lith, 1962,

P. elegans Van Lith, 1962 and P. pempuchiensis Tsuneki, 1971 as subspecies of P.

ornatus Van Lith, 1962.

He was certainly right, the more so as there is also a difference in the colour of

the veins of the fore wings. In P. interstitialis and its subspecies and closely related

species, all restricted to the Papuan area, the veins are yellowish or pale yellowish-

brown. In the other species of the group of P. interstitialis, including P. luzonensis

and P. ornatus, the wing veins are brown.

Also P. baliensis has dark wing-veins and is now reclassified as a subspecies of P.

ornatus.

Psenulus fyanensis sp. nov.

Male (holotype). — Length about 7.5 mm. Head black; mandibles except dark

tips and palpi yellow. Scape of antennae yellow, basal half of underside of

flagellum yellowish, rest of antennae brown. Thorax black with following parts

yellow: dorsal side of pronotum, pronotal tubercles, long narrow mark along

tegulae and two median lines which are broadened near hind margin (Fig. 48),

axillae, scutellum except median longitudinal brown mark, metanotum, four large

marks on back of propodeum. Fore and mid legs yellow, trochanters and femora

somewhat brown below, hind legs reddish-brown. Gaster including petiole

reddish. Veins of wings dark brown.

Median part of anterior margin of clypeus straight, lateral teeth indistinct,

separated by less than % of total distance between the eyes. Raised part of

interantennal carina narrow, ending below antennae in a transverse carina. Frons

J. P. van LITH: Indo-Australian Psenini 113

and vertex indistinctly punctate, postocellar area not raised. Occipitalcarinaending

in hypostomal carina. Antennae long, segment 3 about 2% times, segments 4—12

about twice as long as broad at apex, segment 13 about 2% times as long as broad

at base. No tyloidea.

Pronotal corners rounded. Scutum and scutellum shining, sparsely finely

punctate. Metanotum densely superficially punctate. Behind enclosed area of

propodeum a narrow shining area, back and posterior half of sides of propodeum

coarsely reticulate. Hypo-epimeral area, anterior plate of mesepisternum and

metapleura smooth and shining, mesopleura with distinct but superficial and

rather widespread punctures, punctation on mesosternum denser. Anterior

oblique suture foveolate, widened upper part with transverse carinae. Legs

slender. First recurrent vein of fore wings ending in second submarginal cell, near

radial sector. Petiole cylindrical, about 1% times as long as first tergite which is

about 1% times as long as broad at apex. Gaster indistinctly punctate.

Genitalia (Fig. 47) yellowish-red, slender, inner part of parameres somewhat

transparent, apex straight with slightly projecting outer tip.

Pubescence of face silvery, mostly appressed, of rest of body whitish.

Female unknown.

Vietnam: | g, holotype, Fyan, 900—1000 m, 11 July—9 Aug. 1961, N.R.

Spencer (BISH).

Laos: | &, paratype, Vientiane Prov., Ban Van Eue, 31 May 1966, native

. collector, Malaise trap (BISH).

The male from Laos has a much darker scutellum and the first recurrent vein of

the fore wings is interstitial.

The shape of the parameres points to a close relationship with P. interstitialis

and P. luzonensis.

Psenulus nigrolateralis sp. nov.

Female. — Length about 8 mm. Head black; anterior margin of clypeus reddish,

mandibles except dark tips yellow, palpi yellowish-brown, labrum reddish. Scape

of antennae yellow, underside of flagellum yellowish-brown, upper side dark

brown. Pronotum dorsally yellow. Scutum (Fig. 49) yellow with three black lines,

median line anteriorly narrowed, posteriorly not distinctly connected with hind

margin. Axillae black. Scutellum and metanotum yellow. Enclosed area of

propodeum black, also adjacent black spot laterally. Back of propodeum yellow,

longitudinal groove and two spots at apex of propodeum black. Sides of thorax

including anterior margin of sides of propodeum, and underside of thorax black.

Fore and mid legs including coxae and hind coxae yellow; hind legs including

trochanters yellowish-red. Gaster including petiole reddish. Veins of wings dark

brown.

Median part of clypeal margin protruding, indistinctly bidentate, teeth sepa-

rated by about Y, of total distance between the eyes. Interantennal carina narrow,

no transverse carina below antennae. Occipital carina ending in hypostomal

carina. Frons and vertex shining, minutely punctate, postocellar area not raised.

Third antennal segment about 2% times, segment 4 about twice as long as broad at

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

apex, following segments gradually decreasing in length, segment 11 about 1%

times as long as broad at apex, last segment about twice as long as broad at base.

Pronotal corners rounded. Scutum rather densely finely punctate, intermixed

with larger punctures. Scutellum and metanotum sparsely finely punctate. Back of

propodeum almost smooth, dorso-laterally some fine reticulation. Mesopleura and

mesosternum shining, sparsely finely punctate. Anterior oblique suture foveolate,

widened upper part smooth. Petiole cylindrical, about 1% times as long as first

tergite. First tergite little longer than broad at apex. Gaster minutely punctate,

pygidial area narrow, carinae on apical half almost parallel. Legs slender. First

recurrent vein of fore wings interstitial.

Pubescence of face silvery, mostly appressed. Pubescence of rest of body

whitish. Epicnemial areas below with round, densely, yellowish-white pubescent .

patch. Sternites 4—5 apically with dense short whitish pubescence. Petiole dorso-

laterally and ventro-laterally with long erect whitish hairs.

Male unknown.

Laos: | 9, holotype, Vientiane Prov., Ban Van Eue, 15 Jan. 1966; 1 9, paratype,

Sayaboury Prov., Sayaboury, 15 Jan. 1966, native collectors (BISH).

This form certainly belongs to the group of Psenulus interstitialis and may be a

dark subspecies of P. luzonensis (Rohwer).

Psenulus vientianensis sp. nov.

Male. — Length about 8 mm. Head black; mandibles except dark tips yellow,

palpi pale yellowish-brown. Labrum reddish-yellow. Scape of antennae yellow,

underside of antennae yellowish-brown, upper side dark brown. Thorax black with

following parts yellow: dorsal side of pronotum, pronotal tubercles, axillae,

metanotum and four elongate marks on propodeum. Scutum near pronotal

tubercles with small yellowish-red mark and near hind margin with two very small

yellow marks. Scutellum brown and black, outer lateral margins yellow. In upper

corner of anterior plate of mesepisternum a vague yellowish spot. Fore and mid

legs yellow, underside of femora largely brown. Hind trochanters, femora and tarsi

brown, hind tibiae more reddish. Gaster including petiole reddish. Veins of wings

dark brown.

Median part of anterior margin of clypeus indistinctly bidentate, teeth separated

by about % of total distance between the eyes. Raised part of interantennal carina

narrow, ending below antennae in a transverse carina. Frons and vertex hardly

punctate, postocellar area not raised. Occipital carina normally ending in

hypostomal carina. Antennae slender, segment 3 over twice, segments 4— 12 about

twice as long as broad at apex, last segment over 2% times as long as broad at base.

No tyloidea.

Pronotal corners rounded. Scutum and scutellum shining, finely sparsely

punctate. Metanotum densely, superficially, punctate. Behind enclosed area of

propodeum a smooth margin which is broader laterally, this smooth area defined

postero-laterally by foveolae and a high carina. Back of propodeum coarsely

reticulate, also greater part of sides of propodeum. Anterior plate of mesepister-

num, mesopleura, hypo-epimeral area, metapleura and mesosternum smooth and

J. P. van LITH: Indo-Australian Psenini 115

shining, hardly punctate. Anterior oblique suture foveolate, widened upper part

almost smooth. Petiole cylindrical, about 1% times as long as first tergite, apex

dorsally with small triangular depression. First gastral tergite about 1% times as

long as broad at apex. Gaster minutely punctate. Legs slender. First recurrent vein

of fore wings ending in first submarginal cell or interstitial.

Genitalia (Fig. 45) long and slender, yellowish-brown, inner part of parameres

somewhat transparent, apex straight with outer corner distinctly projecting

backward.

Pubescence of face silvery, mostly appressed, of rest of body whitish.

Female unknown.

Laos: I 3, holotype, Vientiane Prov., Ban Van Eue, 15—31 May 1965; 1 &,

paratype, same locality, 30 Nov. 1965, native collectors (BISH).

This species undoubtedly belongs to the group of Psenulus interstitialis. The

genitalia differ little from those of P. fyanensis, but the colour is darker. Like some

of the related forms described here as species, it may be a subspecies, perhaps of

P. ornatus. The material is too scanty and the study of the genitalia still too

insufficient to allow more definite conclusions.

Psenulus continentis Van Lith

Van Lith, 1962: 97—98 (Malaya).

New record: | 9, Singapore, Campus of University, 5 May 1972 (US).

This female agrees with the description of the holotype which was captured in

Penang, in a car. The axillae are completely yellow. Anterior oblique suture black,

foveolate, widened upper part almost without sculpture.

Psenulus pulcherrimus pulcherrimus (Bingham)

Bingham, 1896: 443 (Psen pulcherrimus; Tenasserim).

Van Lith, 1962: 101 (Psenulus pulcherrimus); 1969: 200 (Vietnam); 1973: 140—141 (subspecies?;

Nepal).

As in the case of a series of specimens from Nepal the variability of P.

pulcherrimus is again shown by 26 specimens from Laos and one male from

Thailand. Most often the scutum has a large postero-median yellow mark and two

narrow marks along the sides. In one third part of the specimens from Laos the

lateral marks are reduced or absent and in two of the females the scutum is

entirely black. On the contrary in three males from Laos and in the male from

Thailand the scutum is not only yellow marked postero-medially and laterally, but

also anteriorly. The anterior marks range from two small separate triangular spots

(Fig. 51) to a more or less broad band (Fig. 52), in one case distinctly connected

with the lateral marks (Fig. 53). Extensive yellow marking of the scutum does not

run parallel with the colour of the scutellum as the two females with entirely black

scutum have the scutellum completely or nearly completely yellow. On the

contrary, in four males with much yellow marking of the scutum the anterior half

or fourth of the scutellum is black. Hind femora, first tergite and apex of petiole

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Fig. 51—53. Dorsal and posterior side of thorax of Psenulus pulcherrimus pulcherrimus (Bingham), 3,

51—52, Laos; 53, Thailand

are not darkened. It seems that scutum and propodeum are richer coloured in the

males, in the females the scutellum is more yellow.

Darkening of fore and mid femora occurs in the Laos material only in a few

females, that of the first tergite in one female and six males. In half of the females

and males the apex of the petiole and the hind femora are darkened. Both colour

forms occur in the same province, even a female and a male taken on the same

date in the same locality of the province of Wapikhamthong are different. Six of

the seven specimens from Sayaboury have a darkened apex of petiole and hind

femora whilst all of the ten specimens from Ban Van Eue in Vientiane Province

have entirely reddish petiole and hind femora. In the material from Nepal the hind

femora are always much darker brown, almost black and the first tergite is

darkened.

In the male from Singapore recorded below the petiole is dark brown at the

apex and the ventral plate is dark brown except a reddish mark near the posterior

margin. The base of the first tergite is brownish-black, the hind femora except

J. P. van LITH: Indo-Australian Psenini 117

apex are dark brown, also the apical half of the hind tibiae. Yellow marks on

propodeum large. Scutellum with narrow black anterior margin.

In the female from Malaya the apex of the petiole, the base of tergite I and % of

the hind tibiae are brown.

First records from Laos: Vientiane Prov., Gi Sion Vill., de Tha Ngone, | 9,

9— 16 Jan. 1966, 1 &, 7—21 Febr. 1965; Ban Van Eue, 4 ©, 15—31 May 1965, 15

Febr. 1966, Malaise trap, 29 March and 30 June 1966, 6 &, 29 April 1966, light

trap, 15 May, 15—31 May and 31 May 1966, Malaise trap, 15 Dec. 1966;

Sayaboury Prov., Sayaboury, 3 ©, 12 Febr. and 13 April 1966, 4 &, 15 Jan. and 2

March 1966, 2 3, 30 Sept. 1966, light trap; Wapikhamthong Prov., Wapi, 3 9, 15

March, 30 March and 31 May 1967, 3 g, 30 March, 15 April and 31 May 1967, .

light trap; Sedone Prov., Pakse, 1 9, 31 May 1967; Savannakhet Prov.,

Savannakhet, 1 3, 15 April 1967, all by native collector (BISH).

First record from Thailand: | 3, Trang Prov., Khaophappha Khaochang,

200—400 m, 3 Jan. 1964, G. A. Samuelson (BISH).

First record from Malaya: 1 9, Kedah, Sungei Toh Pawang Estate, 10 Aug.

1966, H. T. Pagden (BM).

First record from Singapore: | 3, Postal District 22, 26 Sept. 1970, C.G. Roche

(CGR).

Psenulus melanonotus Van Lith

Van Lith, 1969: 203 (West Sumbawa).

First record from Flores: 1 3, C. Flores, Moni, Wolowaru, 11 Nov. 1949, Bühler

and Sutter (NMB).

This male is darker than the two males from Sumbawa. Scape of antennae pale

brown instead of yellow, pronotum dorsally brown, no large yellowish-white spots,

hind margin of scutellum brownish with small yellow spot, metanotum pale yellow

with brown fore margin and vague median brown spot instead of being completely

ivory-coloured, propodeum with two vaguely defined circular yellow spots. Only

hind margin of pronotal tubercles yellowish-brown. Fore legs yellow, femora

brown below; mid legs yellow, trochanters brown below, femora except apical

foreside brown; hind legs brown, only dorso-basal % instead of % of hind tibiae

pale yellow.

Propodeum behind enclosed area very finely striate instead of smooth.

The differences between this male and the two from Sumbawa do not seem to

be of great importance.

Psenulus leucognathus sp. nov.

Male. — Length about 6 mm. Head black; scape and mandibles ivory-white,

apex of mandibles, also labrum, reddish, palpi and underside of flagellum pale

yellowish. Thorax black, with following ivory-white parts: pronotum except

central and lateral parts, pronotal tubercles, two lateral marks on metanotum and

two small marks on back of propodeum. Tegulae reddish transparent. Trochan-

ters, dorsal side of fore and mid femora, fore and mid tibiae except a brown mark

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

on back, tarsi of fore and mid legs and basal % of hind tibiae ivory-white, rest of

hind legs dark brown. Petiole dark brown, hind margin of ventral plate and gaster

red. Veins of wings black.

Raised part of interantennal carina narrow, ending below antennae in a

transverse carina. Anterior margin of clypeus bidentate, distance between apices

of teeth about % of total width of margin. Frons and vertex shining. Postocellar

area narrow. Occipital carina ending in hypostomal carina. Base of antennae

somewhat flattened, segments rounded below, segment 3 about twice, segments

4—9 over 1% times and segments 10—12 about 1% times as long as broad at apex,

last segment about twice as long as broad at base. No tyloidea.

Pronotal angles obtuse. Punctation of scutum distinct, irregular, interstices

centrally a few times as large as punctures, laterally punctures finer. Scutellum and.

metanotum sparsely punctate. Enclosed area of propodeum triangular, with

oblique carinae, back and sides of propodeum coarsely reticulate, behind enclosed

area a narrow smooth area. Mesopleura and mesosternum finely sparsely

punctate, hypo-epimeral area smooth. Anterior oblique suture foveolate, widened

upper part with a few transverse carinae. Legs normal. Petiole about as long as

first tergite, cylindrical, dorsally somewhat flattened. Gaster slender, first tergite

longer than broad at apex.

Face and tempora densely, mostly appressed, silvery pubescent, rest of body

whitish pubescent.

Female unknown.

South India: 1 g, holotype, Kerala State, Walayar Forests, 200 ft, Sept. 1959,

P.S. Nathan (UI, deposited on permanent loan in USNM).

P. leucognathus belongs to the Psenulus pulcherrimus-group. It is very close to P.

melanonotus from Sumbawa, from which it differs in the red gaster and the lesser

extent of the whitish marking. Both species are easily distinguished from other

species of the group by the ivory-white mandibles and whitish marks on pronotum,

metanotum and propodeum.

Psenulus extremus Van Lith

Van Lith, 1966: 45—46 (Psenulus carinifrons extremus; New Guinea); 1969: 206 (Psenulus extremus).

New records from NE. New Guinea: 1 9, East Highlands Aiyura, 1800— 1900 m,

6 Jan. 1965, J. Sedlacek (BISH); 1 3, Huon Gulf, Morobe District, 22 May—19

June 1937, J. L. Froggatt (BM).

The male is slightly paler coloured than the allotype and paratype recorded in

1966. Pronotum, posterior half of scutellum and metanotum brown. Pronotal

tubercles yellowish-brown. Fore and mid tibiae entirely yellow. Basal % of hind

tibiae yellow in dorsal aspect, also hind basitarsi yellow.

Antennal segments 7—12 with low, yellowish-brown, longitudinal ridge (ty-

loidea).

Psenulus carinifrons carinifrons (Cameron)

Cameron, 1902: 288—289 (Psen carinifrons; India).

Van Lith, 1962: 103—104 (Psenulus carinifrons); 1966: 43 (Psenulus carinifrons carinifrons).

J. P. van LITH: Indo-Australian Psenini 119

Female (first description). — Length 7 mm. Head and thorax black; scape of

antennae yellow, underside of flagellum yellowish-brown. Mandibles yellow with

dark red tips. Palpi straw-yellow. Pronotum, pronotal tubercles, scutellum and

metanotum yellow. Fore and mid legs including trochanters yellow, tarsi

somewhat brownish. Hind trochanters brownish-yellow, femora and tarsi dark

brown, tibiae including apical spurs yellow, in lateral view apical Y, of hind tibiae

dark brown. Petiole dark brown, ventral plate, first tergite and second segment

except narrow apical margin reddish, rest of gaster blackish-brown. Veins of wings

dark brown.

Frons slightly convex, distinctly finely punctate, interstices mostly as large as

punctures, vertex still more finely punctate, interstices larger.

Scutum finely punctate, interstices larger than punctures. Scutellum finely

sparsely punctate. Back of propodeum smooth behind enclosed area, finely

obliquely striate on declivous lateral parts, upper margin of sides with coarse

reticulate carination. Mesopleura shining, sparsely punctate.

Pakistan: 1 ©, Karachi, date in leadpencil indistinct (2.9.03?), T. R. Bell (BM).

With note ‘out of reedgrass stems; larva feeds on minute froghoppers’.

Two females from Laos are very similar but their gaster is slightly darker.

Petiole black, narrow apical margin of ventral plate of petiole reddish. In the

female from Tha Ngone the first tergite is dark reddish-brown, second sternite

brown, second tergite basally with lateral, round, red mark, hind margin on both

sides with narrow reddish mark. Hind trochanters and femora black, hind tibiae

yellow, slightly more than apical % black. Length 6.5 mm. In the female from

Sayaboury the first tergite is black, base of second tergite laterally with round red

mark, slightly more than apical % of hind tibiae black.

Laos: 1 9, Vientiane Prov., Tha Ngone, 28 Dec. 1965; 1 9, Sayaboury Prov.,

Sayaboury, 12 Febr. 1966, native collector (BISH).

Psenulus carinifrons rohweri Van Lith

Van Lith, 1962: 108 (pro parte; Java, Kangean Is.); 1966: 45; 1969: 205—206 (Luzon, Formosa).

First record from Sumba: 1 9, W. Sumba, Kodi, 3 Aug. 1949, Buhler and Sutter

(NMB).

The apical third of the hind tibiae is black in dorsal aspect. Hind tarsi including

basitarsi dark brown.

Psenulus carinifrons malayanus Van Lith

Van Lith, 1969: 206 (Malaya, Singapore, Sumatra, Borneo).

New records from Borneo: 2 9, 9 g, Sarawak, Kuching, 14 Jan. 1968, C. G.

Roche (CGR).

Singapore: | g, Postal District 17, 17 Jan. 1971, C. G. Roche (CGR).

Malaya: | 9, Penang I., Baker (USNM); 1 9, 2 g, Perak, Tapah Hill Forest

Reserve, 2700 ft, 21 Oct. 1973, C. G. Roche (CGR).

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

Psenulus carinifrons scutellatus Turner

Turner, 1912a: 54 (Psenulus? scutellatus; NE. Australia); 1916: 128 (Neofoxia scutellatus).

Van Lith, 1962: 108 (Psenulus scutellatus); 1966: 43—44 (Psenulus carinifrons scutellatus); 1969:

203—205 (New Guinea, Ambon, Buru, Mindanao); 1970: 103.

New record from Buru: | 9, Station 9, 18 May 1921, L. J. Toxopeus (MA).

Psenulus sogatophagus Pagden

Pagden, 1933: 97—101 (Malaya).

Van Lith, 1966: 42 (Thailand, Assam?)

First records from Laos: | 9, Vientiane Prov., Gi Sion Vill., de Tha Ngone,

24—31 Oct. 1966, native collector; 1 3, Sedone Prov., Paksong, 17 May 1965, P.

D. Ashlock (BISH).

Psenulus xanthognathus xanthognathus Rohwer

Rohwer, 1910: 660 (Psenulus (Neofoxia) xanthognathus; Luzon).

Van Lith, 1962: 104—107 (Psenulus carinifrons xanthognathus; Luzon); 1966: 44—45; 1969: 206

(Psenulus xanthognathus xanthognathus); 1972: 196.

New records from Luzon: Laguna, 1 9, Los Baños, Mt. Makiling, Baker, 1 5,

Los Bafios, Baker (USNM), Laguna, Agricultural College, 2 9, 2 4,6 Sept. 1931,

F. C. Hadden (BM); I 3, Albay Prov., Libon, Caguscos, 200 m, 12 May 1965, H.

M. Torrevillas (BISH); 2 3, Mount Prov., Ifugao, Mayoyao, 1000—1500 m and

1200— 1500 m, 9 July and 3 Sept. 1966; 3 9 and 8 6, Rizal, Mt. Montalban, Wa-

wa Dam, 150—200 m, 25 Febr.—19 March 1965, H. M. Torrevillas (BISH).

Antennal segments 5—12 or 6—11 of male with indistinct, low, somewhat

shining tyloidea.

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—_— —, 1902. Descriptions of new genera and species of Hymenoptera collected by Major

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—— —, 1906. Hymenoptera of the Dutch Expedition to New Guinea in 1904 and 1905, Part

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———, 1972. Contribution to the knowledge of Oriental Psenulus (Hymenoptera, Spheci-

dae, Psenini). — Tijdschr. Ent. 115: 153—203.

———, 1973. Psenini from Nepal (Hymenoptera, Sphecidae). — Tijdschr. Ent. 116:

123—143.

Pagden, H. T., 1933. Two new Malayan Sphecoids. — Trans. R. ent. Soc. Lond. 81:

93—101.

Perkins, R. C. L. & L. Evelyn Cheesman, 1928. Insects of Samoa, part V, fasc. 1, Hymenop-

tera 1—58.

Ritsema, Cz., C., 1880. On two new exotic species of fossorial Hymenoptera. — Notes Ley-

den Mus. 2: 225—226.

Rohwer, S. A., 1910. Some new Hymenopterous Insects from the Philippine Islands. —

Proc. U.S. natn. Mus. 37: 657—660.

———, 1921. Descriptions of new Philippine wasps of the subfamily Pseninae. — Philipp.

J. Sci. 18: 309—323.

———, 1923. New Malayan wasps of the subfamily Pseninae. — Philipp. J. Sci. 22:

593—601.

Smith, F., 1858. Catalogue of the Hymenopterous Insects collected at Sarawak, Borneo,

Mount Ophir, Malacca and at Singapore, by Mr. A. R. Wallace. — J. Proc. Linn.

Soc. 2: 43—130.

Tsuneki, K., 1966. Contribution to the knowledge of the Pemphredoninae fauna of Formosa

and the Ryukyus (Hymenoptera, Sphecidae). — Etizenia 14: 1—21.

———, 1967. Studies on the Formosan Sphecidae (III). The subfamily Pemphredoninae

(Hymenoptera). — Etizenia 24: 1—11.

———, 1971. Studies on the Formosan Sphecidae (XIII). A supplement to the subfamily

Pemphredoninae (Hym.) with a key to the Formosan species. — Etizenia 57: 1—21.

———, 1974. A contribution to the knowledge of Sphecidae occurring in Southeast Asia

(Hym.). — Polskie Pismo ent. 44: 585—660.

Turner, R. E., 1912a. Notes on Fossorial Hymenoptera. — IX. On some new Species from

the Australian and Austro-Malayan Regions. — Ann. Mag. nat. Hist. (8) 10: 48—56.

———, 1912b. Notes on Fossorial Hymenoptera. — X. On new Species from the Oriental

and Ethiopian Regions. — Ann. Mag. nat. Hist. (8) 10: 361—377.

———, 1916. Notes on Fossorial Hymenoptera. — XIX. On new Species from Australia.

— Ann. Mag. nat. Hist. (8) 17: 116—136.

122

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 4, 1976

INDEX

The names of new species and subspecies are marked with an asterisk

alishanus

anodontotus

anomalus

antennatus

araucarius

armipes

auratus

aureohirtus

baliensis

basilanensis

bicinctus

binghami

bougainvillensis

bryani

carinifrons

chillcotti

chrysomallus

continentis

coriaceus

crabroniformis

dapitanensis

decipiens

Diodontus

elegans

elisabethae

emarginatus

erraticus

esuchus

exiguus

extremus

fyanensis

hakusanus

impressus

inflatus

111

112

102

118

115

103

104

112

102

107

118

112

107

interstitialis

kankauensis

kohli

laosensis

leucognathus

limbatus

littoralis

luzonensis

maculatus

madrasiensis

major

malayanus

melanonotus

Mellinus

meridianus

Mimesa

Mimumesa

multipunctatus

nathani

Neofoxia

nigrolateralis

nitidus

oresterus

orientalis

orinus

ornatus

paulus

pempuchiensis

peterseni

philippinensis

pictus

politiventris

Psen

Pseneo

Psenulus,

108

112

117

107

112

104

119

117

103

120

113

111

112

106

104

110

pulcherrimus

puncticeps

Punctipsen

quadridentatus

refractus

reticulatus

rohweri

ruficrus

rufipes

rufoannulatus

rufobalteatus

sabahensis

salomonensis

sandakanensis

scutatus

scutellatus

sedlaceki

seminitidus

sogatophagus

sumatranus

sumbaensis

suturalis

takasago

tectus

tridentatus

trimaculatus

tristis

umboiensis

unicolor

vaneuensis

varius

vientianensis

wapiensis

xanthognathus

115

119

995

110

106

120

104

111

101

106

112

106

114

103

120

| De T568 oy

DEEL 119 AFLEVERING 5 1976

4 UITGEGEVEN DOOR MUS. COMP. ZOOL. .

È LIBRARY

U 91975

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING |

> HARVARD

UNIVERSITY

INHOUD

F. WILLEMSE and G. KRUSEMAN. — Orthopteroidea of Crete, p. 123—164, fig.

1—13, pl. 1—10.

Tijdschrift voor Entomologie, deel 119, afl.5 _ Gepubliceerd 25-X-1976

ORTHOPTEROIDEA OF CRETE

F. WILLEMSE and G. KRUSEMAN!)

With 13 text-figures and ten plates

ABSTRACT

Previous records and 3300 newly collected specimens formed the basis of this survey of the

Orthopteroidea (= Orthoptera-Saltatoria) of Crete. The following new taxa, all from Crete, are

described: Platycleis (P.) grisea cretica, Eupholidoptera forcipata, E. latens, E. pallipes and E. gemellata.

The following species are new to the fauna of Crete: Phaneroptera n. nana Fieber, Homorocoryphus

n. nitidulus (Scopoli), Sepiana sepium (Yersin), Heteracris I. littoralis (Rambur) and Tropidopola

longicornis (Fieber) (ssp.n.?). The occurrence of Platycleis (P.) escalerai I. Bolivar could be confirmed.

However, previous records of Acrometopa servillei (Brullé), Eupholidoptera chabrieri (Charpentier),

Troglophilus cavicola (Kollar), Omocestus petraeus (Brisout), and Oedipoda miniata (Pallas) are

considered unreliable. In all, at least 63 species are listed. The typically insular fauna of Crete is closely

related to the fauna of the Cyclades and the southern Sporades, while the relationship between Crete

and Anatolia appears to be closer than that between Crete and the Greek mainland. Besides, there

exists an affinity to the Sicilian fauna, although apparently less close than to the fauna of the eastern

part of the Mediterranean Region.

CONTENTS

AECOCU CC TEAM A ai A et A Eee. ES, es Sh. ae 123

MISHOHSPEGIESTOUNAOMOEREtE MEA RR ee cor 124

SYSEC MACHO AGE eat it een EN RE sent: 125

Teiigonione se sa 6 FO Se pon EE cae Olona illa 125

GRO Re ee Se) she 2 lue cles mY Spade et 138

Trilkialboaten ini TN EEE EER 141

WE LRIP OI CA ao obtain Gel ue lcm IAT kek ic 141

INGIICOMOPIOLA CA MO ERE nese ML. en zoe Gee gapen en RA MENT RTE 142

Zones eres Le e i i] aa iena 153

Lit oft locales. "me EA GT RER fins ee al Gwe EEND ces te 155

FEIERTE TE nn Re inn Lollo a dors SSR 163

INTRODUCTION

The latest survey of the Orthopteroidea of Crete was given by Ramme (1927).

The present authors, together with their families, collected new material (3300

specimens), from 44 localities, which served as a basis for the present critical

study. About half of this material has been deposited in the Instituut voor Taxo-.

nomische Zoölogie, University of Amsterdam, while the other half, including the

!) Addresses: of the senior author, Laurastraat 67, Eygelshoven, The Netherlands; of the junior author,

Instituut voor Taxonomische Zoölogie, University of Amsterdam, The Netherlands.

123

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

types of the new taxa, is preserved in the collection of the senior author.

References given under the species names usually concern records of original

material. As to the localities of the species, the reader is referred to the map

(Fig. 13). Each locality or group of adjacent localities has been given a number,

which refers to the “List of Localities’, p. 155. The list contains the data

on the localities visited recently as well as those of previous records, .as far

as traceable.

The classification of the higher taxa of the Acridomorphoidea in the present

paper is that of Dirsh (1975).

The collecting of additional material from another 13 localities by the following

persons is gratefully acknowledged here: A. C. & W.N. Ellis, Amsterdam; W. H.

Gravestein, Amsterdam; J. A. W. Lucas, Rotterdam; S. J. van Ooststroom,

Oegstgeest; J. H. Woudstra, Zaandam; valuable information was provided by A.

Kaltenbach, Vienna and D. K. McE. Kevan, Quebec.

LIST OF SPECIES FOUND ON CRETE

Phaneroptera n. nana Fieber

Tylopsis lilifolia (Fabricius)

Acrometopa cretensis Ramme

Poecilimon cretensis Werner

Conocephalus (Xiphidion) discolor

Thunberg

Homorocoryphus n. nitidulus (Scopoli)

Tettigonia viridissima (Linné)

Decticus albifrons (Fabricius)

Platycleis (P.) grisea cretica subspec.

nov.

P. (P.) intermedia (Serville)

P. (P.) affinis Fieber

P. (P) escalerai I. Bolivar

P. (Incertana) incerta Brunner v.W.

Sepiana sepium (Yersin)

Eupholidoptera astyla (Ramme)

E. cretica Ramme

E. forcipata spec. nov.

E. latens spec. nov.

E. pallipes spec. nov.

E. gemellata spec. nov.

Rhacocleis germanica (Herrich Schaef-

fer)

Uromenus (Bolivarius) elegans

(Fischer)

Dolichopoda paraskevi Boudou-Saltet

D. spec. Boudou-Saltet

Troglophilus spinulosus Chopard

T. roeweri Werner

Gryllus bimaculatus De Geer

Acheta domesticus (Linné)

Tartarogryllus bordigalensis (Latreille)

Modicogryllus geticus Vasiliu (?)

Gryllomorpha dalmatina (Ocskay)

G. cretensis Ramme

Discoptila lindbergi Chopard

Arachnocephalus vestitus Costa

Myrmecophilus (Myrmophilina) ochra-

ceus Fischer

Trigonidium cicindeloides Rambur

Oecanthus pellucens (Scopoli)

Gryllotalpa gryllotalpa (Linné)

Tetrix depressa (Brisout)

Paratettix meridionalis (Rambur)

Paranocarodes fieberi (Brunner v.W.)(?)

Orchamus raulinii (Lucas)

O. y. yersini (Brunner v.W.)(?)

Pyrgomorpha c. conica (Olivier)

Tropidopola longicornis (Fieber) (sub-

spec. nov.?)

Calliptamus italicus (Linné)

C. b. barbarus (Costa)

Heteracris |. littoralis (Rambur)

Pezotettix giornae (Rossi)

Anacridium aegyptium (Linné)

Ochrilidia pruinosa Brunner v.W.

O. tibialis (Fieber)(?)

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 125

Dociostaurus maroccanus (Thunberg) A. t. thalassinus (Fabricius)

Chorthippus (Glyptobothrus) b. brunneus Acrotylus I. longipes (Charpentier)

(Thunberg) A. patruelis (Herrich Schaeffer)

C. (G.) biroi (Kuthy) A. insubricus inficitus (Walker)

Truxalis nasuta (Linné) Oedipoda caerulescens (Linné)

Locusta migratoria Linné O. venusta Fieber

Oedaleus decorus (Germar) Sphingonotus spec.

Aiolopus strepens (Latreille) Acrida turrita (Linné) (?)

SYSTEMATIC PART

Tettigonioidea

Tettigoniidae

Phaneropterinae

Phaneroptera nana nana Fieber, 1853

Localities. N. Rethimnis: 49 (1 ©).

Distribution. P. nana is distributed throughout the Mediterranean Region,

western Arabia, Africa and Madagascar. The range of the nominate subspecies

covers the northern part of this area, coinciding almost exactly with the Mediter-

ranean type of vegetation and Mediterranean crops.

Discussion. Recently P. nana sparsa Stal has been found in Spain (Ragge,

1965). It should be noted here that the comparative length of the fore wing and

hind femur of our specimen agrees with the nominate subspecies and not with

sparsa.

Up to now not recorded from Crete.

Tylopsis lilifolia (Fabricius, 1793)

Tylopsis liliifolia: Werner, 1903: 68; Kuthy, 1907: 553; Werner, 1927: 428; Ramme, 1927: 188.

Localities. N. Chaniou: 9 (1 9); 15e (1 & ); 16 (1 & 1 9); 17b (Werner, 1927),

SO): 20 (1 3); 22b (Kuthy, 1907); 31 (23. 2 9); N. Rethimnis: 40

ee 19), 46" (Kuthy, 1907), 49 (2.9); 50b (1 9); 56 39);

me itakhou: 61 (2 6 1 9); 63 (2 5 1 9); 65 (2 G); 69 (18); 7la (2 9), d Gea

3 2); 74 (1 3); 83a (Werner, 1903); 97a (1 9); N. Lassithiou: 107b (1 6); 116

(2 3); 118 (Ramme, 1927); 122e (1 ©).

Distribution. The range of this species covers most of the Mediterranean

Region.

Acrometopa cretensis Ramme, 1927

Acrometopa macropoda: Kuthy, 1907: 553 (misidentification).

Acrometopa servillea: Werner, 1927: 428 (misidentification ?).

Acrometopa cretensis Ramme, 1927: 122, Fig. Se-8e, Pl. f. Sb-c.

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Localities. N. Chaniou: 6a (Ramme, 1927); 13 (1 9); 19 (Werner, 1927)?;

N. Réthimnis: 46 (Kuthy, 1907; Ramme, 1927); N. Iräkliou: 71a (1 9): 97a (1 3);

N. Lassithiou: 118 (Ramme, 1927); 120(Ramme, 1927).

Distribution. This species is known from Crete and some islands of the

Cyclades: Kimolis, Polivos, fos, Antiparos, Kythnos and Kéa (Werner, 1934, 1937).

Discussion. A. servillea has been recorded only once from Crete (Werner, 1927,

referring to a juvenile male). Most probably the specimen belongs to cretensis,

which was described shortly after Werner’s paper had been published. For the

time being, we remove servillea from the faunistic list of Crete.

Poecilimon cretensis Werner, 1903

Poecilimon jonicus cretensis Werner, 1993: 67, Fig.

Poecilimon distinguendus Kuthy,, 1907: 554.

Poecilimon cretensis: Ramme, 1927: 186; Harz, 1969: 137, Fig. 292—293, 323, 418—419.

Localities. N. Chaniou: I (1 4); 13(2 3); 22d (1 3); N. Rethimnis: 38a (Werner,

1903); 52b (Kuthy, 1907); 53 (93 3 43 9); 54a (Kuthy, 1907), b (Ramme, 1927);

N. Irakliou: 7la (31 & 24 9); 78 (Ramme, 1927); 83b (3 & 49); 86c (Kuthy,

1907); 91 (Kuthy, 1907); 93 (Ramme, 1927); N. Lassithiou: 107a (Ramme, 1927);

118 (Ramme, 1927); 127 (Harz, 1969); 128 (Ramme, 1927).

Distribution. Known only from Crete and one island of the Cyclades: Naxos

(Ramme, 1927).

Discussion. This species occurs from the lowlands up to above the timberline

(2200 m, 52b). Specimens from above the present-day timberline show extensive

black pigmentation.

Conocephalinae

Conocephalus (Xiphidion) discolor Thunberg, 1815

Niphidium [sic] fuscum: Kuthy, 1907: 553.

Localities. N. Chaniou: 22b (Kuthy, 1907); N. Irakliou: 71b (1 ©), d (10 & 13 9

2 juv.).

Distribution. This species is found in the British Isles and throughout Europe,

extending into Palaearctic Asia and N. Africa.

Homorocoryphus nitidulus nitidulus (Scopoli, 1786)

Localities. N. Réthimnis: 41 (1 juv.).

Distribution. This species is widely spread throughout the southern part of

Europe, in subtropic and tropical Africa, and in Palaearctic Asia.

Discussion. Up to now not recorded from Crete.

WiLLEMSE & KRUSEMAN: Orthopteroidea of Crete 127

Tettigoniinae

Tettigonia viridissima (Linné, 1758)

Tettigonia viridissima: Werner, 1927: 428; Ramme, 1927: 188.

Localities. N. Chaniou: 1 (1 juv.); 15e (4g 29); 26b (Werner, 1927);

N. Irákliou: 71a (1 g observed); 78 (Ramme, 1927); N. Lassithiou: 121 (Ramme,

1927).

Distribution. This species occurs in most of the Palaearctic Region.

Decticinae

Decticus albifrons (Fabricius, 1775)

Decticus (Locusta) albifrons: Lucas, 1854: 167.

Decticus albifrons: Werner, 1903: 68; Ramme, 1927: 188.

Localities. N. Chaniou: 10 (Lucas, 1854); N. Réthimnis: 38a (Werner, 1903);

40 (19); 49 (4 3 Œ 9); 50b (1 g); N. Irákliou: 71d (1 3 1 9); 86a (Lucas, 1854);

N. Lassithiou: 111b (Ramme, 1927); 112 (Ramme, 1927).

Distribution. The range of this species covers the Mediterranean Region and

extends into southwestern Asia.

Platycleis (Platycleis) grisea cretica subsp. nov.

(PI. 1 Fig. 2, Pl. 2 Fig. 5)

The material from Crete was compared with over 300 specimens of the grisea-

complex, originating from more than 50 localities (lowland up to 2200 m) and

representing nominate grisea (Fabricius), grisea transiens Zeuner, including

topotypes of the latter, and grisea monticola Chopard. Special attention was given

to specimens from the high mountains of Greece.

The grisea material, except for that from Crete, is characterized as follows:

elytron more than three times as long as pronotum (from 3.2 to 5.1 times), even if

elytron does not reach hind knee completely; Rs vein (nomenclature as proposed

by Ragge, 1955) distinctly separated from MA and with some accessory posterior

branches; exceptionally, Rs fused with MA for a short distance, but then these

veins distinctly diverging again apically; elytron strongly elongate, anterior and

posterior margins roughly parallel, apex widely rounded; hind wing hardly shorter

than elytron, MA with a number of accessory posterior branches; hind femur

invariably slender, distinctly attenuate apically.

As to the characters of the tegmina and the hind femur, it is apparent that the

material from Crete (PI. 1 Fig. 2, PI. 2 Fig. 5) and that from Greece (PI. 1 Fig. 1, PI.

2 Fig. 4) is different (compare a study on Platycleis (Tessellana) by Kaltenbach,

1964). However, the abdominal terminalia and the superficially studied stridula-

tory apparatus appear to be similar. We consider the population of Crete to belong

to the grisea-complex but to represent a distinct subspecies.

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Material studied: 3 holo-, 9 allo-, 5 3 9 9 paratypes, labelled: Hellas, Kriti,

Idi Oros, Kolita-Psiloritis 1700-2100 m, 28-29. vii.1973; additional paratypes: Lefka

Ori, Linoseli above Xiloskalo 1800-1900 m, 5.viii. (1 ©) & Omalos 1000 m, 4.viii.

(14 2 9) 1973, all F. Willemse c.s.

Description.

3-9 (Pl. 1 Fig. 2, PI. 2 Fig. 5). Within the grisea-complex distinct by shorter

tegmina and hind femur. Elytron not or slightly extending beyond tip of abdomen,

by far not reaching hind knee, not more than 2-3 times as long as pronotum;

basally, including the stridulatory apparatus in the male, as wide as usual; apically

strongly narrowing towards narrowly rounded or subacute apex; R vein divided

into RI and Rs, bifurcation located as usual; Rs after a short distance completely,

fused with MA (in 18 specimens) or nearly touching MA (in 2 specimens); elimina-

tion of the area between Rs and MA, together with shortening of the longitudinal

veins and reduction of the areas posteriorly of R, results in brachyptery. Hind

wing short, only slightly longer than half the elytron; MA without accessory

branches, area between MA and MP strongly reduced. Hind femur comparatively

short, slightly attenuate apically. Male cercus of usual shape, slightly shorter than

in nominate grisea. Epiphallus as in grisea-complex. Female abdominal terminalia

about as in grisea transiens, hind margin of last abdominal sternite more or less

elevated in the middle. Coloration as usual.

Measurements (length in mm). [di Oros (6 & 10 9): body 3 17.0-18.1, 9

17.3-19.0; pronotum ¢ 5.1-5.9, © 5.2-5.6; elytron 3 10.6-12.7, 9 10.3-12.8; hind

femur 3 14.8-15.1, © 13.9-15.4; ovipositor 9.9-10.8; Linoseli (1 9): body 17.1;

pronotum 5.6; elytron 13.1; hind femur 15.8; ovipositor 10.3; Omalös (1 3 2 9):

body & 17.2, 9 17.6-17.9; pronotum ¢ 5.0, © 5.2-5.5; elytron & 14.2, 9 14.8-15.0;

hind femur & 15.1, © 16.1-16.9; ovipositor 10.1-10.2.

Localities. N. Chaniou: 13; 15e; N. Rethimnis: 53.

Distribution. Known only from the western and central mountains of Crete.

Discussion. The available material is uniform as to its general appearance. The

tegmina and the hind femora of the few specimens from Omalós are slightly

longer (1-3 mm) than those of the specimens from above the timberline, which

differences may be connected with differences in altitude. The venation and the

shape of the tegmina are uniform, except for the course of Rs in both elytra of the

male from Omalés and in the right elytron of the female from Linoséli. In these

elytra Rs and MA are separate, altough very close, almost touching each other.

Galvagni (1959) described an other short-winged form in Platycleis (Platycleis),

viz., P. (P.) concii. This species occurs in the Madonie mountains of Sicily, from

1000 up to 1800 m altitude. Comparison of concii (Pl. 1 Fig. 3, Pl. 2 Fig. 6) with

grisea cretica reveals the shape and venation of the tegmina to be nearly the same.

However, the long and attenuate hind femur, the more robust general appearance

with wider head and thorax and the larger measurements in the former are quite

distinct.

Previous records of grisea from Crete are doubtful: Griffini (1894: 92) records a

single male from Laki (17a) and Ramme (1927: 188) a single male from Ierapetra

(111b). Both records probably refer to intermedia. Distinction between the males

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 129

of grisea and intermedia was hardly possible in the time of Griffini’s paper, nor in

that of Ramme’s. Reliable records of intermedia (based on the female) from the

two localities are now available: from Ierápetra by Ramme himself (1951), and

from Laki in the present material. We assume that the only representative of the

grisea-complex in Crete is grisea cretica.

Details of the localities and the habitat of grisea cretica are discussed below

under Eupholidoptera forcipata and pallipes. Flight was not observed.

Platycleis (Platycleis) intermedia (Serville, 1839).

Platycleis grisea: Griffini, 1894: 92 (misidentification ?); Ramme, 1927: 188 (idem).

Platycleis intermedia: Kuthy, 1907: 553; Ramme, 1951: 245, 247.

Localities. N. Chaniou: 15b (Kuthy, 1907); 16 (2 3); 17a (Griffini, 1894) ?, c

(2 2); 31 (6 5 3 2); N. Réthimnis: 40 (2 9); 46 (Kuthy, 1907); 48 (44 3 9); 49

(13 49); 50b (lg 59); 55 (1 g); N. Iräkliou: 61 (1 9); 71a (2 3); 90c (1 8

1 92); 91 (Kuthy, 1907); 97a-b (11 g 1 2); N. Lassithiou: 108 (18); 109 (1 9); 111a

(Ramme, 1951), b (Ramme, 1927) ?; 119 (2 9); 122d (1 9).

Distribution. The range of this species covers S. Europe and N. Africa, and

extends far into Palaearctic Asia.

Discussion. Previous records of grisea by Griffini and Ramme which are here

referred to intermedia, are discussed under grisea cretica. It should be noted here

that the specimens of intermedia from Crete usually are smaller than those from

the mainland of Europe, and thus resemble grisea (especially the male).

Platycleis (Platycleis) affinis Fieber, 1853

Platycleis affinis: Werner, 1903: 68.

Localities. N. Réthimnis: 40 (1 9); 50b (24 5 9); N. Irákliou: 86b (Werner,

1903).

Distribution. This species occurs in central, but mainly in southern Europe,

extending into northern Africa and far into western Asia.

Platycleis (Platycleis) escalerai I. Bolivar, 1899

Platycleis escalerai (?): Ramme. 1927: 143, 188.

Localities. N. Irákliou: 61 (2 ¢ 2 9); 71d (1 ©); N. Lassithiou: 111b (Ramme,

1927).

Distribution. This species is distributed throughout southeastern Europe and

the adjacent part of western Asia.

Discussion. While Ramme was not certain of his identification of a single male

from Ierápetra, the present material is proof of the occurrence of this species in

Crete.

Platycleis (Incertana) incerta Brunner von Wattenwyl, 1882

Incertana incerta: Zeuner, 1941: 37, Fig. 31.

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Localities. Crete (Zeuner, 1941); N. Chaniou: 15e (1g 3 9); 16 (1 &); 31

(2 3); N. Réthimnis: 40 (1 & 1 9); 41 (98 22); 48 (1 6 19); 49 (2 g); 50b

(3 62 9); 56 (2 9); N. Irakliou: 57 (4 &); 61(1 9); 63 (1 3); 71d (18 g 24 ©).

Distribution. Known from the Balkan peninsula, Crete, some Aegean islands

and Turkey.

Sepiana sepium (Yersin, 1854)

Localities. N. Réthimnis: 40 (1 9); 50b (1 & 1 ©); N. Irakliou: 71d (2 & 2 9).

Distribution. The range of this species covers southern Europe, including some

Mediterranean islands, and extends into Turkey.

Discussion. Up to now not recorded from Crete.

Eupholidoptera Ramme, 1951

Up to now, two species of Eupholidoptera were known from Crete. In the

present material another four, apparently new, species are recognized. The.

following descriptions of the new species were made after comparison with the

type-species, E. chabrieri (Charpentier).

Key to the males of the Eupholidoptera species from Crete

I. Cercus and'subgemitalfplatewathitee the RED e e e 2

— Cercus and subgenital plate without teeth ................ 3

2. Tip of epiphallus with a lateral spine on each side (PI. 7 Fig.35—36) .....

Peete, Uae SE Gag I MI EIA CIONI nae Sure, eee ee oe VANONI pallipes spec. nov.

— Tip of epiphallus laterally rounded and without spines (PI. 7 Fig. 37—38)

SOL TOD TONE, Ms Che? A ONY Oe ete TE AIR ICE gemellata spec. nov.

3. Stylus of subgenital plate long, about as long as cercus (Fig. 7); epiphallus

symmetrical, apical parts close together, partly fused (Fig.9) cretica Ramme

— Stylus of subgenital plate short, shorter than half length of cercus; epiphallus

asymmetrical, or if symmetrical, then with apical parts divergent nd

4. Median excision of hind margin of subgenital plate very wide and deep and

more than half the length of the plate (PI. 5 Fig. 21); epiphallus symmetrical,

large, robust, apical parts divergent (PI. 6 Fig. 29—31) . . forcipata spec. nov.

—,, Medianiexcision!less wide and deep? Teo een 5

5. Hind margin of last abdominal tergite with a narrow median excision (Fig. 1);

epiphallus asymmetrical, apical parts partly fused (Fig. 4) . . astyla (Ramme)

— Hind margin with a wider median excision (Pl. 5 Fig. 19); epiphallus sym-

metrical, apical parts divergent (PI. 7 Fig. 32—34) ...... latens spec. nov.

A key to the females of Eupholidoptera from Crete is not given. Distinctive

characters are apparent only in forcipata, while the females of cretica and gemellata

are unknown.

Kuthy (1907: 553) recorded Thamnotrizon chabrieri from Amari (46). Ramme

(1927: 194) briefly discussed this record, but Kuthy’s material could not be traced.

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 131

As to the range of this species, its occurrence in Crete is not probable and con-

firmation is needed. For the time being, the species is omitted from the faunal list

of Crete.

Eupholidoptera astyla (Ramme, 1927)

(Fig. 1—5)

Pholidoptera astyla Ramme, 1927: 133, 196, 198, Fig. 11d, 13—14; Ramme, 1930: 799, 821, Fig. 5—7, 9;

Ramme, 1939: 94—96, 100, Fig. 27.

Eupholidoptera astyla: Ramme, 1951: 198, 203, 206, 209, 211, Fig. 51; Harz, 1969: 362, 377, Fig. 1109,

1130, 1159—1161.

Localities. N. Lassithiou: 197a; 111b; 112 (all Ramme, 1927).

Distribution. This species is known only from the original material: a male

from Naxos (the Cyclades), and three females and the tip of a male abdomen

from eastern Crete. The male from Naxos has been selected as lectotype.

Therefore Naxos is the type-locality and not Crete, as indicated by Harz (1969).

Eupholidoptera cretica Ramme, 1951

(Fig. 6—9)

Eupholidoptera cretica Ramme, 1951: 198, 202, 203, 211, Fig. 47, 51; Harz, 1969: 362, 377, Fig.

1111-1112.

Localities. N. Chaniou (12b) (Ramme, 1951).

Distribution. The species is known only from the male holotype. As far as could

be traced, Sanmaria, in the original description, was a misspelling of Samaria.

Eupholidoptera forcipata spec. nov.

(PI. 2 Fig. 7—8, PI. 4 Fig. 14, PI. 5 Fig. 18, 21, PI. 6 Fig. 25, 29—31, PI. 8 Fig. 39)

Material studied: & holotype, 9 allotype, 45 & and 59 © paratypes, labelled:

Hellas, Kriti, Idi Oros, Kolita-Psiloritis 1700-2100 m, 28-29.vii.1973, F. Willemse

C.S.

Description.

& (PI. 2 Fig. 7). Robust. Pronotum wide, scarcely or not widening posteriorly,

metazona comparatively short, hind margin slightly convex. Legs short and thick.

Last abdominal tergite (Pl. 5 Fig. 18) curved strongly downwards; hind margin

with a deep and wide median excision, which is transversely concave in the

middle and straight at sides; from this excision, at either side, projects a large,

triangular lobe with slightly wrinkled surface and strongly toothed apex, which

points ventro-laterally.

Cercus (PI. 6 Fig. 25) without tooth, short, about as long as last abdominal

tergite measured in the middle; basal half roughly cylindrical and straight, apical

half narrowing at inner side and slightly upcurved, apex obtusely pointed.

Subgenital plate (PI. 5 Fig. 21) without spines, very large, slightly wider than

long and slightly tapering apically, distal half strongly slanting upwards and, in

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

9 10 11

Fig. |—5. Eupholidoptera astyla (Ramme). 1—4, male; 1, last abdominal tergite: 2, subgenital plate;

3, cercus, dorsal view; 4, epiphallus; 5, female, subgenital plate. Fig. 6—9, E. cretica Ramme, male.

6, last abdominal tergite; 7, subgenital plate; 8, cercus, dorsal view; 9, epiphallus. (All after Ramme).

Fig. 10—11. Tropidopola longicornis (Fieber) subspec. nov.?, apex of phallus of two males from

Amnissös

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 133

closed position, covering distal part of last abdominal tergite; anterior margin

with an obtuse-angled excision; posterior margin very wide, with a deep and

rectangular excision; lateral parts of plate angularly bent dorsad along a low and

smooth lateral ridge from which the stylus arises distally. Stylus short, not

exceeding one fourth of cercal length, about two or three times as long as wide.

Epiphallus (Pl. 6 Fig. 29—31) strongly sclerotized, large; apical parts fused and

narrow in proximal half, strongly divergent, hook-like and recurved in distal half,

extending far over last abdominal tergite.

General colour yellowish brown. Clypeus, frons and genae with several sym-

metrically arranged black points. Vertex and occiput with a pair of black, medial

fasciae, composed of transverse stripes and usually separated from each other by a

yellow median line. Above the eye a longitudinal black stripe, behind the eye a

similar but wider one. Pronotal dorsum, in middle of prozona, with black markings

in varying degree, metazona sometimes with green flush. Pronotal lateral lobe with

wide, black, dorsal fascia, in prozona not sharply delimited ventrally, in metazona

sharper and strongly narrowing posteriorly. Elytron and last abdominal tergite

completely black. First abdominal tergite partly black, other abdominal tergites

faintly dotted with dark brown. Abdominal sternites orange brown, subgenital

plate orange yellow, the latter with lateral parts black. Cercus dark brown. Fore

and middle legs with numerous black points and stripes. Hind femur with a short,

longitudinal, black stripe dorso-basally, usually composed of a series of transverse

stripes; outer side with a large, roughly triangular or V-shaped black streak near

the middle and with more or less numerous blackish or dark brownish transverse

stripes. Apical half of hind femur with a short, dorsal, black stripe. Hind knee and

postgenicular part of hind tibia black, except dorsally.

Q (PI. 2 Fig. 8). Slightly larger than male. Elytron almost completely covered

by pronotum. Cercus very slightly curved upwards, short, conical, apex pointed.

Abdominal sternites simple. Subgenital plate (Pl. 8 Fig. 39) much wider than long,

distal half strongly slanting upward; posterior margin slightly converging towards

a deep but narrow median excision, which reaches at least middle of plate and

has the angles widely rounded. Ovipositor comparatively short, straight or slightly

curved upwards apically. Coloration as in male.

Measurements (length in mm): body & 21.0—24.0, © 18.5—24.0; pronotum ¢

9.2—10.1, 9 9.0—9.2; elytron & 5.4—6.5, 9 3.0—4.0; hind femur & 15.4—16.9,

Q 17.0—17.6; ovipositor 16.0—17.2.

Localities. N. Réthimnis: 53.

Distribution. Known only from Mt. Idi, central Crete.

Discussion. E. forcipata is well characterized by its abdominal terminalia and

close to anatolica (Ramme). In the male of the latter species the posterior margins

of the last abdominal tergite and that of the subgenital plate are excised in a much

different way, while the stylus is more elongate and the proximal half of the apical

parts of the epiphallus is wider.

The type-locality, Kolita-Psiloritis, can be reached by foot from Kamäres

(520 m), a village on the southeastern slopes of Mt. Idi. From there a track follows

the main water-pipe of the village and leads to several springs. The last and

highest of these springs is named Skaronerò (1650 m) and is the type-locality

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

of E. gemellata (see below). At a short distance and after passing the present-day

timberline, a small plateau with some shepherd’s huts is reached, named Kolita

(1750 m). During the night spent up there we experienced considerable night-

frost. E. forcipata was mainly collected in the surroundings of the huts. It lives

rather hidden in low, dense, prickly shrubs, e.g. Astragalus spec., and is found

together with Poecilimon cretensis and Platycleis grisea cretica. The collecting was

considerably facilitated by the use of firm gloves. At times the sound of E. forci-

pata was heard, during the warmest hours of the day. As in most of Eupholi-

doptera species, the sound much resembles that of Ephippiger species.

Eupholidoptera latens spec. nov.

(PI. 3 Fig. 9—10, PI. 4 Fig. 15, PI. 5 Fig. 19, 22, PI. 6 Fig. 26, Pl. 7 Fig. 32—34, PI.

8 Fig. 40)

Material studied: 3 holotype, 9 allotype, 2 3 3 9 paratypes, labelled: Hellas,

Kriti, Lefka Ori, refuge near Koukoule 1600-1800 m, 6.viii.1973; additional para-

types: 1 3, Lefka Ori, Linoseli above Xiloskalo, 1800-1900 m, 5.viii.1973, and 1 g,

Lefka Ori, Omalos, 1000 m, 4.viii.1973, all F. Willemse c.s.

Description.

3 (PI. 3 Fig. 9). Pronotum with metazona comparatively short, posterior margin

slightly convex.

Last abdominal tergite (PI. 5 Fig. 19) strongly curved downwards; posterior mar-

gin with a wide, concave, moderately deep, median excision, which is sharply

toothed laterally, the teeth pointing ventrally.

Cercus (PI. 6 Fig. 26) without tooth, long, about twice as long as last abdominal

tergite (measured in the middle), slender; basally almost three times as wide as

apically, slightly curved inward; apex obtusely pointed.

Subgenital plate (Pl. 5 Fig. 22) without spines, much longer than wide, strongly

tapering distally; distal part slanting upwards, in closed position scarcely covering

last abdominal tergite; ventral surface with median keel; lateral margin bent

upwards proximally, strongly inflated distally and there forming a smooth, wide

range, ridges on either side converging towards apex of plate; apex with narrow,

triangular, median excision, which does not reach farther than one-fourth of

length of plate. Stylus (PI. 4 Fig. 15) short and thick, length about one-fifth of

cercus, straight, cylindrical, about two or three times as long as wide, inserted

pre-apically at ventral surface of subgenital plate and pointing ventrally (i.e.

perpendicularly to the subgenital plate).

Epiphallus (PI. 7 Fig. 32—34) strongly sclerotized, of moderate length, apical

parts fused and wide in proximal half, moderately divergent, hook-like and weakly

recurved in distal part; the latter extends over last abdominal tergite.

Coloration as in forcipata, but black markings, especially those of hind femur,

less conspicuous.

pronotum or almost so. Cercus straight, short, conical, apex pointed. Abdominal

sternites simple. Subgenital plate (PI. 8 Fig. 40) about as long as wide; posterior

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 135

margin obliquely convergent towards a triangular, median excision, which reaches

about one-third of length of plate, its posterior angles narrowly rounded. Ovi-

positor slightly curved apically. Coloration as in male.

Measurements (length in mm): body & 18.0-23.0, 9 17.8-18.2; pronotum &

8.7-9.4, © 8.2-8.5; elytron & 4.5-6.0, @ 1.0-2.0; hind femur 3 16.0-17.5, 9 16.1-

16.9; ovipositor 13.5-14.4.

Localities. N. Chanfou: 13; 14; 15e.

Distribution. Known only from the Lefka Range, western Crete.

Discussion. This species is well characterized by the abdominal terminalia of the

male, and is easily distinguished from the other species of the genus, with the

exception of astyla. The peculiar position of the male styli resembles that of

astyla. The latter species differs from /atens, in the male sex, in the smaller

excision of the posterior margin of the last abdominal tergite (Fig. 1), the more

inwards curved cercus (Fig. 3), the less tapering subgenital plate (Fig. 2), and the

quite distinct epiphallus (Fig. 4). Distinction of /atens and astyla in the female sex

is not obvious. The slight differences in shape of the subgenital plate (Fig. 5, PI. 8

Fig. 40) cannot be considered reliable, as the variation in both species is insuffi-

ciently known.

The type-locality can be reached, by car, from the village and high plateau,

both named Omalös (1000 m). Via a rather bad track of about 5 km, a refuge

»Kalérji” belonging to the Hellenic Alpine and Ski Federation is reached. It lies

close to one of the peaks of the Lefka Range, named Koukoulé. The specimens

were sparsely found in the surroundings of the refuge, where they live remarkably

hidden in dense shrubs, e.g. Juniperus. Catching them appeared to be a rather

exhaustive task. The adjacent locality Linoséli is discussed below under E. pallipes.

E. latens also occurs on the slopes bordering the Omalòs plateau, where it lives in

shrubs of Quercus coccifera, and again is very difficult to find. Using its song, which

much resembles that of Ephippiger, as a guide, only one single male could be

caught during twilight.

A single female from Mt. Idi, found above Kamares (1000 m) on our way to

Kolita (see under E. forcipata), much resembles /atens. However, without the male,

we are not certain as to its true identity.

Eupholidoptera pallipes spec. nov.

(Pl. 3 Fig. 11—12, PI. 4 Fig. 16, PI. 5 Fig. 20, 23, Pl. 6 Fig. 27, Pl. 7 Fig. 35—36, PI.

8 Fig. 41)

Material studied: 3 holotype, 9 allotype, 5 & paratypes, labelled: Hellas, Kriti,

Lefka Ori, Linoséli above Xiloskalo, 1600—1800 m, 5.viii.1973, F. Willemse c.s.

Description.

3 (PI. 3 Fig. 11). Eyes prominent. Pronotum not widening posteriorly; lateral

edges of dorsum widely rounded; metazona very short, with slightly flattened

dorsum and without median keel or almost so; posterior margin slightly convex.

Last abdominal tergite (Pl. 5 Fig. 20) moderately curved downwards; posterior

strongly produced apically with a narrow, roughly rectangular or heart-shaped

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

median excision, which is strongly toothed laterally, teeth pointing towards each

other or more to the ventral side.

Cercus (PI. 6 Fig. 27) of moderate length, about as long as last abdominal

tergite; relatively thick, basally'twice as wide as apically, slightly curved inwards,

with a narrow, inwards curved, inner tooth at short distance from basis; apex of

cercus obtusely pointed, apex of tooth sharply spined.

Subgenital plate (PI. 5 Fig. 23) about as in the type-species; longer than wide,

moderately tapering distally, in closed position not covering last abdominal

tergite; posterior margin with a moderately deep and narrow, concave, median

excision and, at either side, a single spine situated between median excision

and insertion of stylus. Stylus as in the type-species, at least half as long as cercus.

Epiphallus (PI. 7 Fig. 35—36) weakly sclerotized, narrow and long; apical parts

completely fused along whole length, slightly recurved, slightly tapering distally

but tip widening again and, at either side, with a fine lateral spine; apical parts

considerably projecting, reaching level of apex of cerci.

General colour pale yellow brown. Frons above clypeal margin with a pair of

large black spots, which may be fused into a single transverse fascia. Vertex of

general colour, occiput and area along dorsal margin of eye completely black.

Pronotum of general colour, or, usually, with small black spot in central part of

metazona of lateral lobe. Elytron and abdomen as in the type-species. Legs uni-

colorous, hind femur without black markings, hind knee sometimes blackish.

Q (PI. 3 Fig. 12). Elytron completely covered by pronotum. Cercus curved

slightly inwards, short, conical, apex pointed. Subgenital plate (PI. 8 Fig. 41) as

long as wide, hind margin slightly convergent towards a wide, concave, median

excision, which reaches about one-fourth of length of plate, its apical angles

widely rounded. Ovipositor slightly curved apically. Coloration as in male.

Measurements (length in mm): body 4 18.0-21.0, 9 16.0; pronotum 4 7.4-7.9,

Q 7.5; elytron 4 3.8-5.0, 9 1.5; hind femur g 14.0-15.0, © 16.5; ovipositor 15.0.

Localities. N. Chaniou: 13.

Distribution. Known only from the Lefka Range, western Crete.

Discussion. The species is well characterized by the external abdominal termi-

nalia, which resemble those of the type-species, together with the unique epi-

phallus of the male and the rounded pronotum and unicolorous legs in both sexes.

The type-locality is accessible by foot from a refuge named Xilöskalo (1200 m),

situated at the southernmost point of the Omalós plateau. From here a mountain

track in the direction of one of the peaks of the Lefka Range, named Gigilös

(2080 m), leads to a spring and saddle both named Linoséli. The latter is the

precise locality of the type-series. The habitat is similar to that of forcipata. The

song has not been heard.

Eupholidoptera gemellata spec. nov.

(PI. 4 Fig. 13, 17, PI. 5 Fig. 24, Pl. 6 Fig. 28, Pl. 7 Fig. 37—38)

Material studied: 4 holotype, labelled: Hellas, Kriti, Idi Oros, Kamares-Kolita,

Skaronero 1650 m, 28.vii.1973, F. Willemse c.s.

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 137

Description.

& (PI. 4 Fig. 13). Resembling pallipes, but epiphallus distinct. Apical parts of

epiphallus (Pl. 7 Fig. 37—38) not fused along their whole length; distal thirds

separated, although close together and wider; tip simply rounded without lateral

spines.

Coloration differing from pallipes in black spots on pronotum, which are larger

and fused over the pronotal dorsum into a single, wide, transverse black band,

and also in the absence of black spots on frons.

Q. Unknown.

Measurements (length in mm): body 18.0; pronotum 7.5; elytron 3.2; hind femur

16.0.

Locality. N. Irakliou: 67.

Distribution. Known only from Mt. Idi, central Crete.

Discussion. The single available specimen lacks the left hind leg and the left

fore leg is slightly deformed. Although much resembling pallipes, the different

shape of the epiphallus justifies its specific distinction. The differences in the male

cercus (Pl. 6 Fig. 27—28) in pallipes and gemellata should not be considered

reliable, as variation in both species is insufficiently known.

The type-locality is described under E. forcipata. The specimen was found on

Quercus coccifera, about 15 m above the spring.

Rhacocleis germanica (Herrich-Schaeffer, 1840)

Rhacocleis germanica: Ramme, 1927: 188; Uvarov, 1942: 312.

Localities. N. Chaniou: 12a (Uvarov, 1942); 17c (2 3); 20 (1 9); 31 (1 9); 35d

(23 2 9); N. Réthimnis: 40 (1 ©); 44a (Ramme, 1927); 49 (2 3 7 © 2 juv.); 50b

(1 Q);N. Irakliou: 71d (1 ©); 74(1 3 1 9); 86e (Uvarov, 1942).

Distribution. The range of this species extends from southern France to

southern Slovakia, Moldavia and western Turkey. It is known from Corsica,

Sicily, Kerkyra (= Corfu) and Crete, but not from the other Aegean islands.

Ephippigerinae

Uromenus (Bolivarius) elegans (Fischer, 1853)

Ephippigera idomenaei Lucas, 1854: 165, PI. 2 Fig. 1; Werner, 1903: 68, Fig.; Kuthy, 1907: 553.

Steropleurus siculus: Ramme, 1927: 188.

Steropleurus idomenaei: Uvarov, 1942: 320, PI. 26 Fig. 31.

Localities. N. Chaniou: 10 (Lucas, 1854); 12a (Uvarov, 1942); 31 (2 ¢ 2 9);

N. Rethimnis: 38a (Werner, 1903); N. Irakliou: 69 (1 g); 7la (18 39); 72

(Ramme, 1927); 86a-b-c (Lucas, 1854; Werner, 1903; Kuthy, 1907); 83a

(Werner, 1903); 90b (1 3); 97b (2 3); N. Lassithiou: 106 (Ramme, 1927); 107a

(Ramme, 1927); 109 (1 ©); 111b (Ramme, 1927); 113 (Ramme, 1927); 119

(1 3); 120a (Ramme, 1927); 126 (Ramme, 1927).

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Distribution. The range of this species covers the mainland of Italy (from

Toscana to Calabria), while it also occurs on the islands Corsica, Elba, Giglio,

Sardegna, Sicily and Crete.

Grylloidea

Rhaphidophoridae

Dolichopodinae

Dolichopoda paraskevi Boudou-Saltet, 1973

Dolichopoda spec. Chopard, 1957: 26. (? partim)

Dolichopoda paraskevi Boudou-Saltet, 1973: 58, Fig. A-K.

Distribution. Recorded only from the type-locality: N. Irákliou: 92a-b (Chopard,

1957 ?; Boudou-Saltet, 1973).

Dolichopoda spec. Boudou-Saltet, 1973

Dolichopoda spec. Boudou-Saltet, 1973: 59, Fig. L-N.

Distribution. Known only from the type-locality: N. Lassithiou: 102b. This

species was described but not yet named by lack of a male.

Dolichopoda spec. Chopard, 1957

Dolichopoda spec. Chopard, 1957: 26; Boudou-Saltet, 1973: 57.

Localities. N. Lassithiou: 98a-b & 99a-b (Chopard, 1957; Boudou-Saltet, 1973).

Specimens from caves, which have not been identified due to their juvenile stage.

Troglophilinae

Troglophilus spinulosus Chopard, 1921

Troglophilus spinulosus Chopard, 1921: 147, Fig.; Chopard, 1957: 26.

Localities. N. Chaniou: Sa (Chopard, 1957); 23a (Chopard, 1957), c (Chopard,

1957); N. Réthimnis: 37b (Chopard, 1921); N. Lassithiou: 102a (Chopard, 1957).

Distribution. Known only from Crete.

Discussion. The species was described after a juvenile male. The precise type-

locality is unknown, and placed tentatively under 37b.

Troglophilus roeweri Werner, 1927

Troglophilus roeweri Werner, 1927: 429, Fig. 1—2, 6; Chopard, 1957: 26.

Distribution. Known only from the type-locality: N. Chaniou: 23d.

Discussion. The record of Troglophilus cavicola from Chania (22a) in Werner

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 139

(1903: 69) refers, according to that author (1927: 429), to spinulosus or roeweri.

According to Chopard (1957: 26) both names could be synonymous.

Troglophilus spec.

Troglophilus: Boudou-Saltet, 1973: 57.

Localities. N. Chaniou: 4; 5b; 15f; 23a-b-c; N. Lassithiou: 102b (all Boudou-

Saltet, 1973).

Discussion. This record refers to a list of caves from which Troglophilus is

known without further specification.

Gryllidae

Gryllinae

Gryllus bimaculatus De Geer, 1773

Liogryllus bimaculatus: Ramme, 1927: 189.

Localities. N. Chaniou: 24 (Ramme, 1927); 28c (Ramme, 1927); N. Réthimnis:

50b (2 9); N. Irakliou: 68 (1 9); 97b (5 juv.).

Distribution. The range of this species covers southern Europe and extends

into Africa and Asia.

Acheta domesticus (Linné, 1758)

Gryllus domesticus: Ramme, 1927, 189.

Localities. N. Chaniou: 33 (Ramme, 1927); N. Réthimnis: 54b (Ramme, 1927).

Distribution. Occurring throughout most of the world.

Tartarogryllus bordigalensis (Latreille, 1804)

Gryllus bordigalensis: Kuthy, 1907: 553; Ramme, 1927: 189.

Localities. N. Chaniou: 22b-c (Kuthy, 1907; Ramme, 1927).

Distribution. This species is distributed throughout the Mediterranean Region,

its range extending into western Asia.

Modicogryllus geticus Vasiliu, 1970

Gryllus algericus: Kuthy, 1907: 553. (?)

Locality. N. Chaniou: 22b (Kuthy, 1907).

Distribution. Known from Roumania, Yugoslavia and European Turkey.

Discussion. Kuthy’s record probably refers to a/gericus Brunner von Wattenwyl,

1882 (nec Saussure), which was described by Kis (1967) under the name chopardi.

The latter name being preoccupied, it has been changed into geticus. Confirmation

is needed, especially because Kuthy’s material might belong to another species of

this genus, for instance algirius (Saussure).

140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Gryllomorpha dalmatina (Ocskay, 1832)

Gryllomorpha dalmatina: Werner, 1927: 431; Chopard, 1957: 28.

Localities. N. Chaniou: 18 (Werner, 1927); N. Lassithiou: 102a (Chopard, 1957).

Distribution. The range of this species covers the Mediterranean Region.

Gryllomorpha cretensis Ramme, 1927

Gryllomorpha cretensis Ramme, 1927: 189.

Localities. N. Réthimnis: 44 (Ramme, 1927); N. Irákliou: 57 (1 ©).

Distribution. Known only from Crete.

Discussion. This species was known only from its type-specimen (9). Our adult

specimen agrees with Ramme’s description, except for the coloration of the head,

which is not unicolorous. The vertex between the eyes and the fastigium are dark

brown, while the occiput, genae and face are much paler. The measurements

(length in mm) are as follows: body 9.6; pronotum 1.6; hind femur 6.0; hind tibia

4.5; ovipositor 6.4.

Discoptila lindbergi Chopard, 1957

Discoptila lindbergi Chopard, 1957: 26, Fig. 1, 3.

Localities. N. Irakliou: 58a-b; 60; 88; N. Lassithiou: 98a; 99a, c; 102a; 115 (all

Chopard, 1957).

Distribution. Known only from Crete.

Mogoplistinae

Arachnocephalus vestitus Costa 1855

Arachnocephalus vestitus: Kuthy, 1907: 554.

Localities. N. Chaniou: 22b (Kuthy, 1907); N. Irakliou: 57 (1 9);65(1 ©).

Distribution. This species occurs in the Mediterranean Region.

Myrmecophilinae

Myrmecophilus (Myrmophilina) ochraceus Fischer, 1853

Myrmecophila ochracea: Kuthy, 1907: 554; Ramme, 1927: 190.

Localities. N. Chaniou: 3 (Ramme, 1927); N. Iräkliou: 86c (Kuthy, 1907);

N. Lassithiou: 122f (2 9).

Distribution. The range of the species covers the Mediterranean Region.

Trigonidiinae

Trigonidium cicindeloides Rambur, 1839

Trigonidium cicindeloides: Kuthy, 1907: 553; Ramme, 1927: 189.

Localities. N. Chaniou: 8 (1 9); 22b (Kuthy, 1907); N. Iräkliou: 71c (1 & 7 juv.);

N. Lassithiou: 120a (Ramme, 1927).

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 141

Distribution. Widely distributed in the Mediterranean Region, Asia, Africa and

Madagascar.

Oecanthinae

Oecanthus pellucens (Scopoli, 1763)

Oecanthus pellucens: Kuthy, 1907: 553; Ramme, 1927: 189.

Localities. N. Chaniou: 15b (Kuthy, 1907); 16 (1 9); 22b (Kuthy, 1907);

N. Réthimnis: 56 (1 9); N. Irákliou: 57 (1 9); 84 (3 9); 85b (1 9); 90b (1 5);

97b (1 9); N. Lassithiou: 121 (Ramme, 1927); 122e (1 ©).

Distribution. The range of this species covers central Europe, the Mediter-

ranean Region and N. Africa.

Gryllotalpoidea

Gryllotalpidae

Gryllotalpa gryllotalpa (Linné, 1758)

Gryllotalpa vulgaris cophtha: Werner, 1903: 69.

Gryllotalpa vulgaris: Kuthy, 1907: 554; Ramme, 1927: 190.

Gryllotalpa gryllotalpa: Werner, 1927: 431.

Localities. N. Chaniou: 19 (Werner, 1927); 22b (Kuthy, 1907); N. Réthimnis:

47b (Werner, 1903).

Distribution. Widely spread throughout Europe, northern Africa and Asia.

Discussion. The locality Kolomodis, recorded by Ramme (1927), could not be

traced and is omitted from our list of localities.

Tetrigoidea

Tetrigidae

Tetriginae

Tetrix depressa (Brisout, 1848)

Tettix depressus: Kuthy, 1907: 552.

Locality. N. Chaniou: 22b (Kuthy, 1907)

Distribution. The range covers the Mediterranean Region and extends far into

Palaearctic Asia.

Paratettix meridionalis (Rambur, 1838)

Paratettix meridionalis: Kuthy, 1907: 552; Ramme, 1927: 190.

Localities. Nordkiste (Ramme, 1927); N. Chaniou: 22b (Kuthy, 1907); N.

Réthimnis: 50b (4 g 5 9); N. Irákliou; 69 (4 & 2 juv.); 83c (9 g 15 © 1 juv.); 86c

(Kuthy, 1907); N. Lassithiou: 117 (1 & ).

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Distribution. This species is distributed throughout the Mediterranean Region,

extending into western Asia.

Acridomorphoidea

Pamphagoidea

Pamphagidae

Pamphaginae

Paranocarodes fieberi (Brunner von Wattenwyl, 1882)

Paranocarodes fieberi: Ramme, 1951: 283.

Discussion. This species has only been recorded once (Ramme, 1951), referring

to a female labelled: Kreta, Frivaldsky. Further details on the specimen are not

given. Demirsoy (1973) distinguishes several subspecies in fieberi, from Turkey.

His study deals only with Anatolian material and that from the Aegean islands is

not discussed. The occurrence in Crete needs confirmation.

Orchamus raulinii (Lucas, 1854)

Acinipe raulinii Lucas, 1854: 167, Pl. 2 Fig. 2.

Pamphagus raulinii: Brunner von Wattenwyl, 1882: 201; Kuthy, 1907: 553.

Pamphagus yersinii: Werner, 1903: 67 (misidentification?).

Orchamus raulinii: Ramme, 1927: 192, Uvarov, 1942: 347; Descamps & Mounassif, 1972: 254; Harz,

1975: 109, Fig. 256, 275, 363—364, 367, 371, 334— 391.

Localities. N. Chaniou: 15a (Werner, 1903) ?; 24 (Ramme, 1927); N. Rethim-

nis: 54a (Kuthy, 1907; Ramme, 1927); N. Iräkliou: 86a (Lucas, 1854); N.

Lassithiou: 122g (Harz, 1975).

Distribution. So far known only from Crete.

Discussion. The types being lost, Harz selected neotypes from Sitia (122g).

However, his neotypes should be disregarded because their designation does not

agree with Article 75 of the International Code of Zoological Nomenclature.

According to Ramme (1927), Werner’s record of yersini refers to raulinii.

Orchamus yersini yersini (Brunner von Wattenwyl, 1882)

Porthetis raulinii (nec Lucas, 1854): Yersin, 1860: 529, PI. 10 Fig. 26—28.

Pamphagus versini Brunner von Wattenwyl, 1882: 200.

Orchamus versini: Ramme, 1927: 192; Uvarov, 1942: 347; Ramme, 1951: 411; Descamps & Mounassif,

1972: 252, Fig. 10—11; Harz, 1975: 108, Fig. 267, 276, 365—366, 369—370, 372—375.

Locality. “Candia” (Brunner von Wattenwyl, 1882; Ramme, 1951; Descamps &

Mounassif, 1972; Harz, 1975).

Distribution. The nominate subspecies is recorded from Crete, some Aegean

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 143

islands (Kos; Marathokampos, Samos; Karpathos), but mainly from Syria and the

Lebanon. The subspecies hebraeus Uvarov, 1942, is known from Israel.

Discussion. The record from Crete is based on the locality labels of the ¢ holo-

type and © allotype: Candia, Prof. Zeller, don. Dohrn 1855. However, it is

astonishing that since Brunner’s record of 1882 or the year of the locality label,

1855, no further material from Crete has become available. An explanation was

given by Uvarov (1942) who assumed that the types bear not the correct locality

labels and thus yersini does not occur in Crete. It is not within the scope of the

present paper, to unravel this problem. In any case, confirmation of the occur-

rence of yersini in Crete is needed.

Pyrgomorphidae

Pyrgomorphinae

Pyrgomorpha conica conica (Olivier, 1791)

Pyrgomorpha grylloides: Griffini, 1894: 92; Werner, 1903: 67; Kuthy, 1907: 553; Ramme, 1927: 192.

Localities. N. Chaniou: 20 (27 & 16 9); 22b (Kuthy, 1907); 25a (Griffini, 1894);

N. Réthimnis: 38a (Werner, 1903); 54b (Ramme, 1927); N. Irákliou: 7la-b (8 &

13 2); 73 (Ramme, 1927); 74 (1 9); 76 (Ramme, 1927); 78 (Ramme, 1927); 81

(1 2); 83a (Werner, 1903); 85a-b-c (3 3 7 2); 86c (Kuthy, 1907), d (Ramme,

1927); 90a-b (2 2); 91 (Kuthy, 1907); 93 (Ramme, 1927); N. Lassithiou: 104 (1 9);

105b (2 3 2 2); 108 (1 3); 109 (2 3); 111b (Ramme, 1927); 119 (19); 122b-c (6 3

119).

Discussion. A revision of Pyrgomorpha is currently in progress (Kevan, 1971 and

1974). Mr. Kevan studied a sample of our material and kindly informed us that

we are dealing with the nominate subspecies.

Acridoidea

Catantopidae

Tropidopolinae

Tropidopola longicornis (Fieber, 1853) subspec. nov. ?

(Fig. 10,11, PI. 8 Fig. 42—43)

Localities. N. Irákliou: 71b (24 4 9),c(1&),d(1g 3 9, reared to adult stage

in October); 84 (3 3); 86f (1 3); 90b (12 g 109); N. Lassithiou: 122e (11 3

10 2).

Discussion. Tropidopola has not yet been recorded from Crete. The latest

revision of the Mediterranean species was given by La Greca (1964). He men-

tioned the following taxa: cylindrica cylindrica (Marshall, 1836) from the western

Mediterranean area; graeca graeca Uvarov, 1926, from mainland Greece, southern

and western Anatolia, and Cyprus; graeca transjonica La Greca, 1964, from

Taranto, Apulia, S. Italy; longicornis longicornis Fieber, 1853, from Egypt (type-

locality proposed by Uvarov, 1926: 173. which is not in accordance with Fieber’s

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

original data); and /ongicornis syrica (Walker, 1871) from Syria and Palestine. Our

material does not fit any of these taxa. The shape of the head (PI. 8 Fig. 42—43)

and of the male cercus resemble /ongicornis rather than graeca. The antennae are

shorter than in nominate /ongicornis and resemble those of /ongicornis syrica. The

longest middle segments are between 1.2 to 1.6 times as long as wide. However,

the phallic complex (studied in nine males) differs from that of /ongicornis in the

apex which is comparatively more elongate, slender, while the outline of the apical

penis valves (in lateral aspect) is from hardly (Fig. 10) to moderately (Fig. 11)

sigmoid and not angulate or incised as figured by La Greca (1964: Fig. 37—38).

After a study of material from other parts of the Mediterranean Region, it became

apparent that, due to insufficient knowledge of the individual variation, some

characters are not reliable. We consider the Tropidopola population of Crete to.

constitute a distinct geographical race, allied to nominate /ongicornis.

Calliptaminae

Calliptamus italicus (Linné, 1758)

Caloptenus italicus: Griffini, 1894: 92 (?); Werner, 1903: 67 (2); Kuthy, 553 (2).

Calliptamus italicus: Werner, 1927: 431; Jago, 1963: 320.

Calliptamus italicus grandis: Ramme, 1927: 193.

Localities. Kriti (Jago, 1963); N. Chaniou: 26b (Werner, 1927); 31 (39);

N. Réthimnis: 48 (3 & 1 9); 49 (18 5 2); 50b (1 6 1 2); N. Irakliou: 61 (1 5);

Tla (35 29), d (3 & 39); N. Lassithiou: 106 (Ramme, 1927); 107a (Ramme,

1927); 109 (2 g); 111b (Ramme, 1927); 114 (Ramme, 1927); 119(1 3).

Distribution. The range of this species covers southern Europe and Turkey,

from where it extends into central Asia.

Discussion. Griffini (1894), Werner (1903) and Kuthy (1907) mentioned only

italicus and not barbarus. However, the latter species is by far more common in

Crete as well as throughout southern Europe, where italicus ranks second (cf.

Jago, 1963: 320). Therefore their records are not included in the locality list above.

Calliptamus barbarus barbarus (Costa, 1836)

Calliptamus siculus: Ramme, 1927: 193.

Calliptamus barbarus barbarus; Ramme, 1951: 311; Jago, 1963: 329, 334.

Localities (summary). N. Chaniou: 6; 9; 14—17; 20; 31; N. Réthimnis: 37—40;

44; 48—50; 56; N. Irakliou: 57; 61; 63—65; 67; 69; 71; 74; 82; 84; 87; 89—90;

96—97; N. Lassithiou: 105—107; 116; 120—122; 124—125.

Distribution. Widely distributed throughout the Mediterranean Region, its

range extending far into Palaearctic Asia.

Discussion. The data of our material of barbarus (154 & 205 9) agree with

Jago’s remark that barbarus in Crete occurs up to 1700 m. The same can be said of

italicus, which lives up to 1100 m. In the same paper, Jago mentioned the

population of barbarus in Crete to have bright orange legs, and “many specimens

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 145

show separate inner femoral spots”. Our specimens have indeed bright orange

inner sides of the hind femora and pale orange hind tibiae, but the inner femoral

spots are usually fused.

Eyprepocneminae

Heteracris littoralis littoralis (Rambur, 1838)

Localities. N. Chaniou: 20 (1 3 4 juv.); N. Irakliou: 71a (24 1 9), d (3 juv.);

74 (48 5 32 2).

Distribution. The range of the nominate subspecies covers southern Spain (type-

locality), northern, western and eastern Africa, and extends into S.W. Asia.

Discussion. Our material was compared with that from southern Spain. The

colour of the hind tibiae and the shape of the male subgenital plate agree with

the nominate subspecies rather than with littoralis similis (Brunner von Wattenwyl,

1861) or other subspecies. At Pitsidia (74) the species is abundant and lives in the

dunes under extremely dry and hot conditions.

Till now not recorded from Crete.

Catantopinae

Pezotettix giornae (Rossi, 1794)

Platyphyma giornae: Griffini, 1894: 92.

Pezotettix giornae: Kuthy, 1907: 553; Ramme, 1927: 193.

Localities. N. Chaniou: 9 (1 ©); 15b (Kuthy, 1907), e (18 2 9); 16(2 3 39);

17a (Griffini, 1894); 20 (1 9); 31 (1 9); 36 (Ramme, 1927); N. Rethimnis: 37c

(Ramme, 1927); 39 (Ramme, 1927); 40 (3 5 2 9); 4103 a 3 2); 49 (1a 39); 56

(43 2 9); N. Irdkliou: 57 (5 g 2 9); 61 (1 9); 63 (23); 65 (lg 79); 82464

89).

Distribution. Widely distributed in the southern part of central Europe and

the Mediterranean Region.

Cyrtacanthacridinae

Anacridium aegyptium (Linné, 1764)

Acridium (Gryllus) lineola: Lucas, 1854: 169.

Acridium aegyptium: Griffini, 1894: 92; Werner, 1903: 67; Kuthy, 1907: 553.

Anacridium aegyptium: Werner, 1927: 431; Ramme, 1927: 193.

Localities. N. Chaniou: 2 (Lucas, 1854); 6b (1 3 2 9); 21 (Werner, 1903); 22c

(Ramme, 1927); 26b (Werner, 1927); 28a-b (Lucas, 1854; Griffini, 1894);

30 (Werner, 1903); 35d (2g 1 juv.); 36 (Ramme, 1927); N. Réthimnis: 41

(1 juv.); 50b (2 g); N. Irakliou: 57 (1g); 63 (1 juv.); 65 (1 9); 7la (1 9), d

(23 2 juv.); 74 (1 juv.); 78 (Ramme, 1927); 83c (1 4 1 juv.); 86a (Lucas, 1854),

c.d (Kuthy, 1907; Ramme, 1927), f (14 2 9); 90b (2 5); 97a-b (2 9); N. Lassi-

thiou: 104 (1 9); 122a (Lucas, 1854); 123 (1 ©).

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Distribution. The range of this species covers southern Europe, large parts

of Africa, and S.W. Asia.

Acrididae

Gomphocerinae

Ochrilidia pruinosa Brunner von Wattenwyl, 1882

Ochrilidia pruinosa Brunner von Wattenwyl, 1882: 92.

Platypterna pruinosa: Kuthy, 1907: 552; Ramme, 1927: 191, 194; Salfi, 1931: 284, Fig. 48—57.

Localities. N. Chaniou: 20 (1 juv.); 22b-c (Kuthy, 1907; Ramme, 1927); 35b

(63 3 2), d (1 juv. 9, reared to adult stage in November); N. Iräkliou: 84 .

(2 3); 85c (46 19 1 juv.); 89 (10 3 99 4 juv.); 90b (13 1 9); N. Lassi-

thiou); 123(2 3); 125a (63 & 12 9).

Distribution. This species, described from Rhödos, has been recorded from

Crete, several islands of the Cyclades and Sporades, Cyprus and Mediter-

ranean E. Turkey.

Discussion. The present material has been compared with a large topotypic

series from Rhödos, which agrees fairly well with the descriptions and figures

of Brunner von Wattenwyl and Salfi. The black spot on the lower inner lobe

of the hind knee is uniformly present in the topotypic material and in material

from Crete. However, in the specimens from Crete, the typical pruinose

coloration is lacking in both sexes, except for the males from Frango Kastello

(35b), in which it is present although less conspicuous than in the topotypic

specimens. Also the proportions of the body, tegmina and legs in the speci-

mens from Crete are slightly more robust than in the topotypes. We are not

convinced that the Ochrilidia population of Crete is identical with pruinosa.

From Crete another species of Ochrilidia has been recorded, viz., tibialis

(Fieber). At present its identity is not traceable, as will be pointed out below.

Some references are given:

Platypterna tibialis Fieber, 1853: 98; Ramme, 1927: 194; Salfi, 1931: 255, 322, Fig.

179—181.

Ochrilidia tibialis: Brunner von Wattenwyl, 1882: 91, Fig. 22; Werner, 1901:

272; Johnston, 1956: 716; Harz, 1975: 603, Fig. 2169—2178.

Fieber’s description is unsatisfactory. The locality reads: ,,Griechenland.

Straube. Fieb.”. Brunner von Wattenwyl’s description gives some useful details.

As to the distribution he recorded: ,,Vorkommen: Griechenland (Fieb.), Candia

(c.m.), Spanien (Mus. Genf.). — Ausserdem in Aegypten und Syrien (c.m.).”.

Salfi gave a full description and figures of the specimen on which Brunner von

Wattenwyl’s record “Candia” was based. Salfi presumed this specimen to

represent Fieber’s type. However, Mr. Kaltenbach (Vienna Museum) kindly

informed us (in litt. 17.X.1973): „Unser stark beschadigtes und vielfach zusammen-

geleimtes Exemplar von Platypterna tibialis Fieb. angeblich aus der Coll. Fieber

tragt an der Nadel 5 Etiketten mit folgender Beschriftung: 1. 9481. — 2. Coll. Br.

v. W. ex Coll. Fieber Kreta. — 3. det. Br. v. W. Ochrilidia tibialis. — 4. M. Vienna.

— 5. Platypterna tibialis Fieb. 9 — M. Salfi det. Die handgeschriebenen Original-

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 147

Fig. 12. Greek area in 1850: Greece within the black lines: G. Br., Ionian islands belonging to the

United Kingdom; remaining part, Turkey (after A. Stieler, Hand Atlas, Gotha, 1864)

etiketten Brunners (von Fieber ist anscheinend kein Etiket erhalten !) sind in

Brunner’s Inventar eingeklebt und zeigen die Beschriftung: 1. 9481 Platypterna. —

2. tibialis Fryvw. 9481 Kreta. ,,Fryvw.” steht wahrscheinlich p. err. für ,,Fieb.”!.

Nach dem von Ihnen angegebenen Zitat Brunner’s ist es durchaus möglich, dass

das Tier Fieber’s gar nicht erhalten ist und die gedruckten Etiketten an der Nadel

(erst nach dem Tod Brunner’s gedruckt) auf Grund eines Missverstandnisses den

Vermerk ‚ex Coll. Fieber” enthalten. In Wirklichkeit handelt es vielleicht um das

Tier aus Brunner’s eigener Sammlung. Unser 9 hat helle Kniee wie Salfi angibt.

Sonst ist die Art nur durch ein Expl. aus Syrien bei uns vertreten.”

Thus, the validity of the “Kreta” labelled specimen as holotype is at least

doubtful. Furthermore, as far as we could trace, additional material which both

agrees with Brunner’s specimen and originates from Crete or Greece has never

more been found since Brunner. Therefore, the occurrence within the region of

Greece of an Ochrilidia species which lacks the black spots of the lower inner lobe

of the hind knee is doubtful. Unless such material will be found and a neotype can

be selected, Fieber’s tibialis is not identifiable. It is pointed out here, that in

Fieber’s time ,,Griechenland” did not cover the actual area of Greece (Fig. 12).

Attempts to find out the origin of Fieber’s specimen, which he obtained from

Straube, a dealer, were not successful.

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Dociostaurus maroccanus (Thunberg, 1815)

Stauronotus maroccanus: Kuthy, 1907: 552.

Dociostaurus maroccanus: Ramme, 1927: 191.

Localities. N. Rethimnis: 54b (Ramme, 1927); 55 (1g); N. Iräkliou: 7la

(21 4 28 9); 75 (Ramme, 1927); 80 (Ramme, 1927); 86c-d (Kuthy, 1907;

Ramme, 1927); 90a (13 19); N. Lassithiou: 105b (34 3 9); 109 (6g 19);

110(Ramme, 1927); 111b (Ramme, 1927).

Distribution. This species occurs in southern Europe and northern Africa,

its range extending far into Palaearctic Asia.

Chorthippus (Glyptobothrus) brunneus brunneus (Thunberg, 1815)

Stenobothrus bicolor: Werner, 1903: 67; Kuthy, 1907: 552.

Stauroderus bicolor: Werner, 1927: 431; Ramme, 1927: 191.

Localities. N. Chaniou: 11 (Werner, 1903); 15c (Werner, 1927), e (1 g); 17b

(Werner, 1927); 22b (Kuthy, 1907); N. Réthimnis: 46 (Kuthy, 1907); 54c (1 3);

55 (4 9); 56 (3 3 10 9); N. Irakliou: 57 (3 3 2 9); 64 (1 3); 65 (48 5 9); 83b

(1 & 1 9); 86c (Kuthy, 1907); 91 (Kuthy, 1907); 97a-b (6 3); N. Lassithiou: 105b

(1 3); 120b (2 9).

Distribution. Widely distributed throughout Europe and western Asia.

Discussion. The specimens from Crete are slightly smaller than those from the

mainland of Europe and Turkey, and the elytra are slightly less attenuate.

Werner (1903: 66) recorded a male from Chania and a male from Réthimnon

under Stenobothrus petraeus Brisout. His material could not be traced in the

Vienna Museum (Kaltenbach, in litt. 8.vii.1975). According to Werner’s paper, he

had no other material of petraeus available. He mentioned that the hind tibiae

of these males were yellowish red, which disagrees with his identification. As far

as could be traced, there are no further records of petraeus from Crete, the

Cyclades and the Sporades, nor from the Peloponnese. For the time being, we

therefore omit Omocestus petraeus (Brisout) from our list.

From Omalds (15), Griffini (1894: 92) recorded a juvenile specimen under

Stenobothrus spec. ?. From this locality both brunneus and biroi are known.

Chorthippus (Glyptobothrus) biroi (Kuthy, 1907)

(Pl. 10 Fig. 47—48)

Stenobothrus biroi Kuthy, 1907: 552, 554.

Stauroderus biroi: Ramme, 1927: 194.

Chorthippus (Glyptobothrus) biroi: Harz, 1975: 876, Fig. 3157—3158, 3285—3291.

Localities. N. Chaniou: 9 (2 9); 13 (18 3 26 2); 14 (5 g 2 9); 15b (Kuthy, 1907),

ei (lo 10°9); 16 (143509); Mic (9) (Harz, 1975, MINCE MAO) Nt

Rethimnis: 40 (1 4);48 (2 g 3 9); 49 (16 g 8 9);50b (18); 53(3g 1 9); 5602

3 9); N. Irakliou: 57 (1 9); 65 (33 8 9); 67 (3g 19); 82 (1 9); 87 (2 3); 90c

(33 4 2); 97b (1g 3 2), c(2 3); N. Lassithiou: 101c (Harz, 1975); 116(4 3 8 9);

NEON

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 149

Distribution. So far known only from Crete.

Discussion. This species occurs from lowland up to 2000 m. The shape of the

lateral pronotal keels is rather variable. The colour of the hind tibia ranges from

pale yellowish to blackish brown. Opportunity is taken here to give some figures

of the general appearance of this little known species (Pl. 10 Fig. 47—48).

Truxalinae

Truxalis nasuta (Linné, 1758)

Tryxalis procera: Lucas, 1854: 167 (misidentification ?).

Tryxalis variabilis; Lucas, 1854: 167.

Tryxalis unguiculata: Brunner von Wattenwyl, 1882: 90; Griffini, 1894: 91; Werner, 1903: 66; Kuthy,

1907: 552.

Acridella variabilis: Werner, 1927: 431.

Acridella nasuta. Ramme, 1927: 190.

Truxalis nasuta: Dirsh, 1951: 206, Fig. 139—149, 217, map 9.

Localities. Crete (Dirsh, 1951); Candia (Brunner v.W., 1882); N. Chaniou: 22a-b

(Werner, 1903; Kuthy, 1907); 24 (Ramme, 1927); 25a-b (Griffini, 1894; Werner,

1927); 26a-b (Werner, 1927; Ramme, 1927); 28b (Griffini, 1894); 35a (Dirsh, 1951);

N. Réthimnis: 37c (Ramme, 1927); 38a-b (Werner, 1903; Ramme, 1927); 46

(Kuthy, 1907); 47a (Griffini, 1894); 54b (Ramme, 1927); N. Irákliou: 7la-b (1 3

3 0); 77 (Ramme, 1927); 83a (Werner, 1903); 85a (1 3); 86a-b-c-d (Lucas, 1954;

Werner, 1903; Kuthy, 1907; Ramme, 1927); 90a (1 3); N. Lassithiou: 104 (1 3);

105a (Dirsh, 1951), b (1 9); 111b (Ramme, 1927); 112 (Ramme, 1927); 114

(Ramme, 1927); 122b-c (2 3).

Distribution. The range of this species covers the southern parts of the main-

land of Europe, the Mediterranean islands, and northern Africa, and extends into

Palestina and Syria.

Discussion. The record of procera Klug, 1830, by Lucas (1854) is most probably

erroneous, as Crete is widely separated from the area of distribution of that

species (Dirsh, 1951: 183, map 8).

Oedipodinae

Locusta migratoria Linné, 1758

Oedipoda Gryllus migratoria: Lucas, 1854: 170.

Pachytilus danicus: Kuthy, 1907: 553.

Locusta migratoria ph. danica: Ramme, 1927: 191.

Localities. N. Irákliou: 71d (2 3); 86a (Lucas, 1854), c (Kuthy, 1907); N. Lassi-

thiou: 120a (Ramme, 1927).

Distribution. According to Harz (1975: 466) the population of Crete belongs to

subspec. cinerascens (Fabricius, 1781), which is distributed throughout southern

Europe.

150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Oedaleus decorus (Germar, 1826)

Oedaleus nigrofasciatus: Werner, 1903: 67.

Oedaleus decorus: Ramme, 1927: 191.

Localities. N. Réthimnis: 38a (Werner, 1903); N. Irakliou: 86b (Werner, 1903),

d(Ramme, 1927); 83a (Werner, 1903); N. Lassithiou: 106 (Ramme, 1927).

Distribution. Widely distributed throughout S. Europe, N. Africa and Asia.

Aiolopus strepens (Latreille, 1804)

Epacromia strepens: Kuthy, 1907: 552.

Aeolopus strepens; Ramme, 1927: 191.

Aiolopus strepens: Hollis, 1968: Fig. 51.

Localities (summary). N. Chaniou: 15; 17; 22; 34; 36; N. Rethimnis: 37—43;

45; 48; 50; 56; N. Irakliou: 57; 63; 65; 67; 69; 71; 74; 83—87; 89—90; 97;

N. Lassithiou: 105; 107; 117; 122; 125.

Distribution. The range of this common species covers most of the Mediter-

ranean Region.

Discussion. Our material (109 3 96 9) was found from the lowland up to

1700 tn.

Aiolopus thalassinus thalassinus (Fabricius, 1781)

Oedipoda Acridium laeta: Lucas, 1854: 170.

Epacromia thalassinia: Kuthy, 1907: 552.

Aeolopus thalassinus: Ramme, 1927: 191.

Aiolopus thalassinus thalassinus: Hollis, 1968: Fig. 84.

Localities. N. Chaniou: 20 (2 ¢); 22b (Kuthy, 1907); 32 (Lucas, 1854); N. Réthim-

nis: 38b (Ramme, 1927); 50a (2 9); N. Iräkliou: 71a-b-c-d (13 & 16 9); 84 (1 3);

85a (5 5 1 9); 86a (Lucas, 1854), d (Ramme, 1927); 90b (2 9); 95 (Ramme, 1927);

97b (13 1 9), d(1 9); N. Lassithiou: 108 (1 3); 109 (3 3); 120b (1 3); 122b(3 3

49),e(23 6 9).

Distribution. The nominate subspecies occurs in southern Europe, the whole of

Africa, its range moreover extending far into southwestern Asia.

Discussion. A record from Mt. Idi (52a) (Lucas, 1854) appears doubtful because

the preferred habitat of this species differs very much from that offered by Mt. Idi.

Acrotylus longipes longipes (Charpentier, 1843)

Acrotylus longipes: Kuthy, 1907: 553.

Localities. N. Chaniou: 20 (4 5 6 ©); 22b (Kuthy, 1907); N. Irákliou: 71d (1 9);

74 (22 5 19 2); 85a (8 3), c (13 3 13 2); 89 (14 3 14 2); 90b (20 3 14 9); 97d

(21 3 16 9).

Distribution. Widely distributed in southeastern Europe, southwestern Asia, and

Africa.

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 151

Discussion. The species lives predominantly along the seashore. The hind wing

in our material is yellow or colourless, but not orange.

Acrotylus patruelis (Herrich-Schaeffer, 1838)

Acrotylus patruelis: Griffini, 1894: 92; Ramme, 1927: 192.

Localities. N. Chaniou: 7 (Ramme, 1927); 13 (1 9); 28b (Griffini, 1894); N.

Réthimnis: 37c (Ramme, 1927); 42a-b (Ramme, 1927); 43 (Ramme, 1927); 50b

(5 3 1 2); 56 (24 1 2); N. Irakliou: 65 (1 3); 7la (2 3), d (1 8); 87 (1 3); 97d

(2 3 2 2); N. Lassithiou: 104 (1 3); 105b (1 3); 109 (16 19); Ille (16 19);

177(1 9); 119(1 9); 122d(1 g); 125b (1 g).

Distribution. The range of this species covers the Mediterranean Region, south-

western Asia and most of Africa, including Madagascar. In Crete, the species

occurs up to 1800 m (13).

Acrotylus insubricus inficitus (Walker, 1870)

Acrotylus insubricus: Kuthy, 1907: 552; Ramme, 1927: 192.

Acrotylus insubricus inficitus: Maran, 1958: 177.

Localities. Kreta (Maran, 1958); N. Chaniou: 3 (Ramme, 1927); 20 (1 g); 22b

(Kuthy, 1907); 24 (Ramme, 1927); 36 (Ramme, 1927); N. Réthimnis: 38b (Ramme,

1927); 39 (Ramme, 1927); 44b (1 9); 50b (3 3); 51 (Ramme, 1927); N. Irakliou: 64

(1 g); 7ld (1 92); 75 (Ramme, 1927); 82 (1 9); 83c (1 & 1 9); 84 (5 & 2 2); 85c

(1 5); 86d (Ramme, 1927), f (1 9); 87 (63 2 2); 89 (2 2); 90b (2 3), c (Sg 19);

97c (2 9); N. Lassithiou: 116 (3 4 2 9); 120a (Ramme, 1927); 124 (1 ©).

Distribution. According to Mafan, this subspecies occurs in central and south-

western Asia, the southern part of European USSR, northeastern Africa, and

Crete. Our material agrees with his description of the subspecies.

Oedipida caerulescens (Linné, 1758)

(Pl. 9 Fig. 44)

Oedipoda Gryllus coerulescens: Lucas, 1854: 170.

Oedipoda caerulea [sic] : Griffini, 1894: 92.

Oedipoda coerulescens: Kuthy, 1907: 553; Werner, 1927: 431; Ramme, 1927: 191.

Localities (summary): N. Chaniou: 2; 7; 9; 13—17; 22; 25—28; 31; N. Réthimnis:

37; 39—41; 44; 48—50; 56; N. Irákliou: 57; 61; 63—65; 67; 69; 71; 74; 82; 86—87;

90; 94; 97; N. Lassithiou: 105; 107; 109—111; 119; 122.

Distribution. This species is widely distributed in Europe, northern Africa and

western Asia.

Discussion. This common species (studied: 148 3 136 9) occurs in Crete up to

1800 m. The black fascia of the hind wing, although with a wide overlap, is less

strongly developed in the specimens from Crete than in those from the mainland

of Greece (PI. 9 Fig. 44).

152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Oedipoda venusta Fieber, 1853

(Pl. 10 Fig. 45—46)

Oedipoda venusta Fieber, 1853: 23; Brunner von Wattenwyl, 1882: 161; Werner, 1903: 67 (?); Kuthy,

1907: 553; Ramme, 1927: 192; Harz, 1975: 489, Fig. 1615 (Mioscirtus v.), 1747.

Oedipoda gratiosa: Werner, 1903: 67 (misidentification ?).

Localities. Creta (Brunner v. W., 1882); N. Chaniou: 13 (7 g 6 9); 14(3 9); 15b

(Kuthy, 1907), e (83 5 9), g (Harz, 1975); 16 (33 4 9); N. Rethimnis: 38a

(Werner, 1903) ?; 52b (Kuthy, 1907); 53 (16 GS 12 9); 54a (Kuthy, 1907); 55

(6 3 6 9); N. Irákliou: 67 (11 g 8 9); Timbáki, see discussion (Harz, 1975); 83a

(Werner, 1903) ?; 94 (Ramme, 1927); N. Lassithiou: 100 (Ramme, 1927); 101b

(2 9); 103 (Ramme, 1927); 107b (2 9); 122c (13 g 13 ©).

Distribution. Despite Fieber’s original record “Griechenland”, this species is,

as far as known, confined to Crete.

Discussion. The present material agrees with Ramme’s comment that the tip of

the hind wing in many specimens is but scarcely infuscate (PI. 10 Fig. 45—46). The

upper keel of the hind femur is lowered in its apical part, although not as con-

spicuous as in some other species of Oedipoda. Attention is drawn to this character

because venusta has been placed in some other genera in the older literature (Scin-

tharista, Mioscirtus, Microscirtus, Morphacris). This was apparently due to errone-

ous descriptions, reading in Fieber “Hinterschenkel oben mit ganzem Kiel” and in

Brunner von Wattenwyl “femora postica carina superiore haud interrupta”.

Among the localities listed above, the name Timbaki refers to: Tympaki, 29.-

31.vii.1958, H. Eckerlein (Harz, 1975: Fig. 1747). The locality could no more be

included and numbered in the list of localities and on the map, the manuscript

being finished.

The male type is damaged and preserved in the Vienna Museum. Harz designa-

ted neotypes, which should be disregarded being not in agreement with Article 75

of the International Code of Zoological Nomenclature.

The records of venusta and gratiosa by Werner (1903) refer to juvenile specimens

from the same localities. His material could not be traced in the Vienna Museum

(Kaltenbach, in litt. 8.vii.1975). Identification of juvenile specimens in Oedipoda

species is not reliable. As gratiosa (= miniata Pallas) has been only recorded once

and the record is doubtful, we omit this species from the faunal list of Crete.

Sphingonotus spec.

Oedipoda Gryllus coerulans: Lucas, 1854: 170.

Sphingonotus caerulans: Griffini, 1894: 92.

Sphingonotus coerulans: Kuthy, 1907: 552; Ramme, 1927: 192.

Sphingonotus rubescens rubescens: Mistshenko, 1936: 170.

Localities. Crete (Mistshenko, 1936; 3 9); N. Chaniou: 9 (1 9); 14(7 & 10 9);

15b (Kuthy, 1907), e (16 & 12 9); 17c (1 3); 20 (2 3); 28b (Griffini, 1894); N.

Rethimnis: 37a (Lukas, 1854), c (Ramme, 1927); 38b (Ramme, 1927); 42a (Ramme,

1927); 45 (Ramme, 1927); 50b (20 g 14 9); 53 (10 3 5 Q); N. Iräkliou: 67 (6 3

19); 7la (1 3), c (1 9); 74 (1 9); 79 (Lukas, 1854); 86c-d (Kuthy, 1907; Ramme,

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 153

1927); 90a-b (2 & 2 9); 93 (Ramme, 1927); 97d (20 3 17 9); N. Lassithiou: 101a

(Ramme, 1927); 105b (4 3 8 2); 108 2 3); 109 (2 3 49); Ille (1 g 2 9); 113

(Ramme, 1927); 120b (2 3 1 9); 122b-c-d-e (9 g 11 ©).

Discussion. Sphingonotus is common throughout Crete, from the seashore up to

1800 m. The identification of our material offered considerable difficulties. It

agrees partly with caerulans (L.) (especially with caerulans exornatus Nedelk.),

partly with nominate rubescens (Walker). However, the greater part of the material

is intermediate between these taxa. Especially in longer series from one locality

all transitional forms are present. We agree with the critical remarks by Ramme

(1951: 406) and abstain from identification of the present material from Crete.

We assume that the previous records of Sphingonotus from Crete refer to material

similar to ours.

Acridinae

Acrida turrita (Linné, 1758) (?)

Tryxalis nasuta (nec Linné); Kuthy, 1907: 552.

Localities. N. Chaniou: 20 (1 3 1 juv.); 22b (Kuthy, 1907); 35b (1 juv.); N.

Rethimnis: 40 (2 juv.); 41 (1 juv.); 50b (1g 2 juv.); N. Irákliou: 64 (1 9); 71d

(2 juv.); 74 (2 juv.); 84 (12 3 7 9); 85c (6 3 1 2); 86c (Kuthy, 1907); 87 (43 1 9);

89(11 g 10 9); 90b (9 3 3 2); 97c (2 juv.), d (4 g).

Distribution. Widely distributed in Africa and recorded from Sicily and Vul-

cano I.

Discussion. Our material has been identified with the help of the latest

revision of the genus (Dirsch, 1954). Judging from the position of the transverse

sulcus of the pronotum, which is clearly behind the middle of the pronotum, the

population of Crete should belong to turrita. However, we are not certain because

it is difficult to distinguish it from bicolor (Thunberg) (recently synonymized with

ungarica Herbst), which is the common species of this genus in the Mediterranean

Region. i

In the older literature there is considerable confusion about the nomenclature

in Acrida and Truxalis species. If the original material cannot be re-examined,

old records should be disregarded. However, in this particular case Kuthy’s

records are interpretable: he recorded both Tryxalis nasuta L. and T. unguiculata

Rambur, which represent Acrida turrita (L.) and Truxalis nasuta (L.), respectively

(compare Brunner von Wattenwyl, 1882: 87—90).

CONCLUSIONS

Among the present material from Crete, the following new taxa are described:

Platycleis (Platycleis) grisea cretica subspec. nov., Eupholidoptera forcipata, E. latens,

E. pallipes and E. gemellata. As far as we could trace, the following species are new

to the fauna of Crete: Phaneroptera n. nana Fieber, Conocephalus (Xiphidion)

discolor Thunberg, Homorocoryphus n. nitidulus (Scopoli), Sepiana sepium (Yersin),

Heteracris |. littoralis (Rambur) and Tropidopola longicornis (Fieber) (subspec.

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

nov.?). The occurrence of Platycleis (Platycleis) escalerai Bolivar could be confirmed.

The identity of the Acrida and Sphingonotus material from Crete could not be

established.

For various reasons, previous records of Modicogryllus geticus Vasiliu, Parano-

carodes fieberi (Brunner v. W.) and Orchamus y. yersini (Brunner v. W.) are doubt-

ful and need confirmation. According to Chopard (1957) Troglophilus roeweri

Werner could be synonymous with Chopard’s 7. spinulosus. It is pointed out that,

for the time being, the identity of Ochrilidia tibialis (Fieber) is not clear. Previous

records of Acrometopa servillea (Brullé), Eupholidoptera chabrieri (Charpentier),

Troglophilus cavicola (Kollar), Omocestus petraeus (Brisout) and Oedipoda miniata

(Pallas) are considered unreliable.

In all, we listed at least 63 species of Orthopteroidea (= Orthoptera-Saltatoria)

occurring in Crete. The fauna of Crete is typically insular, i.e. comparatively

poor in species. The range of about one third of the species is, as far as known,

confined to the island, or covers Crete and some neighbouring islands of the

Cyclades. A list of these species is given here:

Acrometopa cretensis Ramme

Poecilimon cretensis Werner

Platycleis (Platycleis) grisea cretica subspec. nov.

Eupholidoptera astyla (Ramme)

E. cretica Ramme

E. forcipata spec. nov.

E. latens spec. nov.

E. pallipes spec. nov.

E. gemellata spec. nov.

Dolichopoda paraskevi Boudou-Saltet

D. spec. Boudou-Saltet

Troglophilus spinulosus Chopard

T. roeweri Werner (synon.?)

Gryllomorpha cretensis Ramme

Discoptila lindbergi Chopard

Orchamus raulinii (Lucas)

Tropidopola longicornis (Fieber) subspec. nov. (?)

Oedipoda venusta Fieber

Chorthippus (Glyptobothrus) biroi (Kuthy)

Little can be said about the relationship of these species and subspecies with

other members of the genera concerned. The position of Acrometopa cretensis is

somewhat isolated in the genus, as is discussed by Ramme (1927). The nearest

relative of Poecilimon cretensis is P. inflatus Brunner von Wattenwyl, which occurs

in southwestern Anatolia, opposite the southern Sporades. Platycleis (Platycleis)

grisea cretica comes close to the nominate form and can be considered to be

related to the Sicilian species P. concii Galvagni. The affinity between the

numerous Eupholidoptera species is, with few exceptions, far from clear. Orchamus

raulinii presumably is more closely allied to those species of the genus, which

occur in some Aegean islands, Anatolia, Cyprus, Syria, the Lebanon and Israel.

Our Tropidopola material from Crete comes close to the Egyptian species. T. longi-

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 155

cornis (Fieber), but as pointed out in this paper, a revision of the Mediterranean

species is needed and the suggested resemblance with /ongicornis might be in-

correct. The relationship with congeneric species in cases like Oedipoda venusta,

Chorthippus (Glyptobothrus) biroi and the different diverse cave-Orthoptera is

doubtful, except for Dolichopoda paraskevi. This species comes near D. naxia

Boudou-Saltet, which occurs in Naxos, one of the Cyclades. This agrees with the

close resemblance between the fauna of the Cyclades and that of Crete, which is

also proved by the distributional ranges of Acrometopa cretensis, Poecilimon creten-

sis and Eupholidoptera astyla.

The number of Eupholidoptera species confined to Crete is remarkably large,

which is very interesting from a zoogeographical point of view. There is a con-

spicuous difference between the rich fauna of Anatolia and the neighbouring

Aegean islands, where the largest number of Eupholidoptera species occur, and the

poor fauna of the Peloponnese and Kithira more to the south, which have one

species each. This could be an argument that the faunas of Crete and Anatolia

would be more closely related than those of Crete and the Peloponnese. However,

as to the number of congeneric species, the opposite holds true for Poecilimon.

While in Crete this genus is represented by one species only (although very close

to the Anatolian P. inflatus), in Anatolia and the Balkan Peninsula, including the

Peloponnese, an overwhelming number of Poecilimon species occur.

The non-endemic species of Crete are, with few exceptions, widely distributed

species: they are found all over the Mediterranean Region, or in its western or

eastern parts, or throughout southeastern Europe, including the Peloponnese, the

Aegean islands and northwestern Anatolia. A particular case is formed by the only

member of the Ephippigerinae in Crete, Uromenus (Bolivarius) elegans (Fischer),

which occurs in Corsica, Sardegna, Italy, Sicily and Crete. Actually, Crete forms

the southeasternmost part of the range of this subfamily, which has its centre of

distribution in southwestern Europe and northwestern Africa. Neither in the

Peloponnese, nor in the Aegean islands or Anatolia a member of this subfamily is

known to occur. Unfortunately, the distribution in the Greek and Anatolian

regions and often the taxonomy of the faunistically more important species are far

from sufficiently known.

With so many gaps in our knowledge, it may be clear that only general conclu-

sions can be drawn as to the relationship of the orthopterous fauna of Crete. The

closest relationships are found with the fauna of the Cyclades and the southern

Sporades. The affinity to the fauna of Anatolia appears to be greater than to that

of the Greek mainland, especially of the Peloponnese. Besides there is a

resemblance to the fauna of Sicily, although less clear than to that of the eastern

part of the Mediterranean Region.

LIST OF LOCALITIES

(numbers refer to the map, Fig. 13)

The localities and data of the material studied in the present paper and that

recorded previously, are listed below under regional headings: Nomös Chaniou

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

156

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WILLEMSE & KRUSEMAN: Orthopteroidea of Crete

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158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

nr. 1-36; Nomös Réthimnis nr. 37-56; Nomös Iräkliou nr. 57-97; Nomos Lassi-

thiou nr. 98-128.

The Greek orthography may be transcribed in Roman characters in diverse

ways. In our list we use the spelling as printed in the Tourist Map of Crete, copy-

right Chr. Z. Mathioulakis, Athens. This map is included in H. Guanella, Kreta,

Ein Reiseführer, 3. Auflage 1972, Flamberg Verlag, Zürich. Other transcriptions

used previously follow the ones used here.

The names of most of the collectors are abbreviated as follows:

(A) K. Attems (Li) K. Lindberg

(B) L. Birò (Lu) J.A.W. Lucas

(B-S) P. Boudou-Saltet (S) A. Schultz

(E) A.C. & W. N. Ellis (O) S.J. van Ooststroom

(Ga) W.H. Gravestein (We) erner

F.W

(Ge) O. Grebenchikoff (Wi) F. Willemse c.s.

(K) M.C. &G. Kruseman (Wo) J.H. Woudstra

Nomós Chaniou:

1. Falásarna, 7.v.1973 (O & Ga).

. Kisamos: plateau de Kissamos (Luc).

. Meráda: Dorf Merades sw. von Kastelli Kisamu, 7.iii.1925 (S).

. Spilia Aghia Sofia: cavité Aghia Sophia, Topolia (B-S).

. Ellinöspilio,

a. Hellinospilo, 25.iv.1955 (Li);

b. cavité Hellinospilo (B-S).

6. Rodopou: |

a. Rhodopu, 8.x:1926 (S);

b. 1.v.1973 (O).

7. Moni Odigitrias: Kloster Gonia, Halbinsel Spatha, 8.x.1925 (S).

8. Gerani, 6.v.1973 (Ga).

9. Vasiliana, 700 m, 7.viii.1973 (Wi).

10. Paleochora: environs de Sélino (Luc).

11. Aghia Roumeli: Hagia Rumeli, 10.v.1900 (A).

12. Samaria:

a. (Ge);

b. Sanmaria, 13.vi.1942, Kl. Zimmermann.

13. Linoséli, along mountain track between Xilöskalo and summit Gigilós (Léfka Ori), 1600-1800 m,

5.viii. 1973 (Wi).

14. Koukoulé, surrounding of mountain hut ““Kalérji” below summit Koukoule (Léfka Ori), 1600-1800

m, 6.viii.1973 (Wi).

15. Omalós:

a. Homalos, 1050 m, 8.v.1900 (A);

b. 1050 m, 18.viii.1906 (B);

c. Homalos-Ebene, 1000 m, vi.1926 (We);

d. 4.v.1973 (Ga);

e. Omalós plateau, 1000 m, 4.viii.1973 (Wi);

N B © D

grotte Omalos Katavothron (B-S);

1000 m, 31.vii.1938 (Ge).

áki-Omalós, between the villages, 750 m, 4.viii.1973 (Wi).

23.

24.

25.

26.

36.

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 159

. Laki:

a. Lacus(C);

b. Lakkos, in Schluchten, 520 m, vi.1926 (We);

c. 450 m, 3.viii.1973 (Wi);

d. Lakki, 500-1000 m, 30.vii.1939 (Ge).

. Mesklä, Schlucht nach Theriso, 300-540 m, vi.1926 (We).

. Fournés: Phurnes, Tal des trockenliegenden Platanos, 100 m, vi.1926 (We).

. Makris Tichos, 5 km West of Chania, dunes, 0-5 m, 7.viii.1973 (Wi).

. Perivolia, sidwestlich von Kanea, v.1900 (A).

. Chanià:

a. Kanea, v.1900 (A);

b. Canea, 1906 (B);

c. Canea, 31.iii.1925 (S);

d. v.1968 (Wo).

Katholiko:

a. grotte de Catholivo ou d’Aguiou, 21.iv.1955 (Li) & cavité Catholivo ou Aghiou (B-S);

b. cavité Panagia (B-S);

c. Achyrospilo, 21.iv.1955 (Li) & cavité Achyrospilo (B-S);

d. Arkalo Spileo, Halbinsel Akrotiri, 200-300 m, v (? vi) 1926 (We); (precise location unknown).

Moni Tzagaroliou: Kloster Aja Trias, Halbinsel Akrotiri, 26-28 .i1.1925 (S).

Akrotiri:

a. (C);

b. Akrotiri-Ebene, 30-70 m, v.1926 (We).

Aptera:

a. Abdera, Eparchie Apokoronas, 9.x.1925 (S);

b. Ruinen und Gewölbe des Metellus Creticus, 200 m, v.1926 (We).

. Nisos Agion Theodöron: ile D. Teodore (C).

. Eparchia Apokorönou (approximately):

a. plateau d’Apokorona (Luc);

b. Apocorona (C);

c. Apokorona, 9.x.1925 (S).

. Georgioupolis: cavité (B-S).

. Alikampos: Ali Kampos, v.1900 (A).

. Askyfou, 750 m, 2.viii.1973 (Wi).

. Chöra Sfakion: pentes de Sphakia (Luc).

. Komitädes, 18.iii.1925 (S).

. Kapsodäsos, 19.iii.1925 (S).

. Frango Kastello:

a, Frankokastelli, 25.v.1938, R. E. Gathorne-Hardy;

b. 5.v.1973 (Ga);

c. idem (QO);

d. beach and dunes, 0-4 m, 3.viii.1973 (Wi).

Skaloti, 19.iii.1925 (S).

Nomös Réthminis:

37.

38.

39.

40.

41.

Goniä:

a. environs de Gonia (Luc);

b. 23.iii.1904, D. M. A. Bate (location correct ?);

c. Megali Episkopi bis Gonia, 12.x.1925 (S).

Réthimnon:

a. Rethymno, 21.v.1900 (A);

b. 13.x.1925 (S).

Prasiés: Prasses, 23.x.1925 (S).

Sélia, 450 m, 2.viii.1973 (Wi).

Spili, 600 m, 2.viii.1973 (Wi).

160

33.

54.

55:

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Patsós:

a. Patssos, 22.x.1925 (S);

b. Potami, nw. von Patssos, 22.x.1925 (S).

. Louträ: Lutra, 14.x.1925 (S).

. Moni Arkadiou:

a. Kloster Arkadi, 16.x.1925 (S);

b. 8.v.1973 (O).

. Moni Asomaton: Kloster Assomatos, 17.x.1925 (S).

. Amari, 1906 (B).

. Vizari:

a. Visari (C);

b. Visari, 24.v.1900 (A).

. Ano Méros, 500-700 m, 1.viii.1973 (Wi).

. Chordakion, 450 m, 1.viii.1973 (Wi).

. Aghia Galini:

a. 0-40 m, x.1972 (E);

b. 0-40 m, 31.vii.1973 (Wi).

. Kouroutes: Kurutes, am Fuss des Psiloriti, 18.x.1925 (S).

. fdi Oros (approximately):

a. pentes d’Ida (Luc);

b. Mons Ida, 2200 m, 1906 (B).

Kolita-Psiloritis, between hamlet Kolita and summit Psiloritis (fdi Oros), 1700-2100 m, 28-29.vii.

1973 (Wi).

Nidha plateau:

a. Antrum Jovis, 1200-1500 m, 1906 (B);

b. Nidka-Hochebene und Andiskari, 11.v.1925 (S);

c. near the Idéon Antron, 1370 m, 11.vi.1972 (K).

Anogia, 800 m, 10-15.vi.1972 (K).

Drosia, 250 m, 23.x.1972 (E).

Nomös Irakliou:

ST,

59%

60.

61.

62.

63.

64.

65.

66.

67.

68.

69.

70.

12:

13.

Marathos, 450 m, 26.x.1972 (E).

Spilios Camilari:

a. 5.iv.1955 (Li);

b. grotte annexe de Camilari, 2.iv.1955 (Li).

Krousönas: Krussona, 9.v.1925 (S).

Sarchos: grotte de Sarkhos, 10.iv.1955 (Li).

Siva, 300 m, 27.iii.1973 (Wi).

Asites: Assitaes, M. Holtz.

Stavrakia, 300 m, 27.vii.1973 (Wi).

Tsagaraki, 15.x.1972 (E).

Aghia Varvara, 600-750 m, 21.x.1972 (E).

Megali Vrisi: Megali Wryssi, 23.v.1925 (S).

Kamäres-Kolita, along mountain track between village Kamäres and hamlet Kolita ({di Oros), 520-

1650 m, 28-29.vii.1973 (Wi).

Kamares, 520 m, 28-29.vii.1973 (Wi).

Vorizia, 450 m, 27.vii.1973 (Wi).

Zarós: Saro, 15.v.1925 (S).

Phaistos:

a. near the excavations, 100 m, 23-26.v & 18.vi.1972 (K);

b. 1-2 km South of the ruines, wet meadows in the plain, 40 m, 24.v.1972 (K);

c. idem, 16 & 18.x.1972 (E);

d. idem, 30.vii.1973 (Wi).

Aghios Ioanis: Ajos Joannis, 16.v.1925 (S).

Mires: Myràs, 16.v.1925 (S).

74.

79°

76.

17.

78.

TR)

80.

81.

82.

83.

84.

85.

86.

87.

88.

89.

90.

Ole

92.

93.

94.

95.

96.

97.

98.

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 161

Pitsidia, dunes, 0-60 m, 30.vii.1973 (Wi).

Siva: Siwa, Ep. Pyrgiotissa, 7.v.1925 (S).

Moni Odigitrias: Kloster Hodigitria, Ep. Kanurion, 18.v.1925 (S).

Lasea, 18.v.1925 (S).

Moni Apezanon: Kloster Apeganas, Ep. Kanurion, 20.v.1925 (S) (correct ?).

Mesaras: plaint de Messara (Luc) (approximately).

Vourvoulitis: Wurwulitis, 22.v.1925 (S).

Loukia, 29.iv.1973 (Ga).

Archanes, 600-750 m, 19.x.1972 (E).

Knossos:

a. 29.v.1900 (A);

b. wayside, 17 & 21.v.1972 (K);

c. along brooklet, South of the ruines, 100 m, 10.viii.1973 (Wi).

Gazion, 7 km West of Iraklion, x.1972 (E).

Therisos, also Lindo Beach or Lido, 3 km West of Iräklion, 0.3 m:

a. 19.v.1972 (K);

b. 14.x.1972 (E);

c. 25.1x.1973 (K).

Iraklion:

. environs de Candie (Luc);

. Kandia, 1900 (A);

. Herakleion, 1906 (B);

. 1-24.vi.1925 (S);

(Ge);

. 5-12.iv.1975 (Lu).

Kornarou, 6 km East of Iraklion, 10-15 m, 27.ix.1973 (K).

Neraidispilo ou grotte Ilithias, 4.iv.1955 (Li).

Florida Beach, 6 km East of Iraklion, 0-3 m, 26.ix.1973 (K).

A mnissòs:

a. beach, 0-5 m, 20.v. & 16.vi.1972 (K);

b. 27.x.1972 (E);

c. 5-50 m, 29.ix.1973 (K).

Nisos Dia: Insula Dhia, 1906 (B).

Skotinon:

a. Hagia Paraskevi a Scotino, 5.v.1955 (Li);

b. grotte Aghia Paraskevi, prés du village de Skotino, ix.1971 (B-S).

Moni Agaräthou: Kloster.Anagarathos, 27-28.v.1925 (S).

Xidas: Xyda, 27.vi.1925 (S).

Kastamonitsa, 27.vi.1925 (S).

Chersonisos: Chersonissos, 24.vii.1925 (S).

Malia:

maonoc»e

a. inland ofthe ruines, 10-15 m, 18.v. & 9.vi.1972 (K);

b. 22-23. & 29.x.1972 (E);

c. along highway, 10 m, 9.viii.1973 (Wi);

d. beach, 0-5 m, 28.ix.1973 (K).

Nomòs Lassithiou:

Vrachasion:

a. grotte Peristera, 11.v.1955 (Li);

b. idem (B-S).

Milatos:

99.

a. grotte de Milatos, 12.v.1955 (Li);

b. idem (B-S);

c. grotte de Saint-Constantin, 12.v.1955 (Li).

100. Kato Metochi, 28.vi.1925 (S).

162 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

101. Psichron:

a. Psychro, 29.vi.1925 (S);

b. 850 m, 19.vi.1972 (K);

c. Psychro, 900 m, 7.viii.1939 (Ge).

102. Diktéon Antron:

a. Dictéon antron, 7.v.1955 (Li);

b. grotte Diktaion Andron, ix.1971 (B-S).

103. Katharò plateau: Hochtal Catharo, 30.vi.1925 (S).

104. Kritsá, 330 m, 29.v.1972 (K).

105. Aghios Nikélaos:

a. St. Nikolo, v.1904 W.M.;

b. 0-5 m, 28-31.v.1972 (K);

c. 3 km North and South of the town, 0-50 m, 9.viii.1973 (Wi);

d. 31.iii.1975 (Lu).

106. Males: Malas, 1.vii.1925 (S).

107. Anatoli:

a. Anadoli, 1.vii.1925 (S);

b. 400 m, 9.viii.1973 (Wi).

108. Graligia, 0-5 m, 3.vi.1972 (K).

109. Xerökampos, 0-5 m, 2.vi.1972 (K).

110. Kalögeri: Kalogeros, 2.vii.1925 (S).

111. Ierapetra:

a. Jerapetra;

b. idem, 30.vi.1925 (S);

c. 0-5 m, 1.vi.1972 (K).

112. Kato Choriò: Kato Chorion, 3.vii.1925 (S).

113. Pachiamos, 4.vii.1925 (S).

114. Sfaka: Sphaka, 5.vii.1925 (S).

115. Tourtouli: Megali Katofyngui, 9.v.1955 (Li).

116. Sikia, 400 m, 5.x.1973 (K).

117. Aghios Geörgios — Epäno Episkopi, 300 m, 5.x.1973 (K).

118. Chamézion: Chamesi, 6.vii.1925 (S).

119. Skopi, 100 m, 7.vi.1972 (K).

120. Piskokéfalon: i

a. Piskokephalon, 7.vii.1925 (S);

b. 5 m, 7.x.1973 (K).

121. Poúsa Eklisiá: Russaklisia, 15.vii.1925 (S).

122. Sitia:

. plateau de Sitia (Luc);

. beach near the town, 0-3 m, 5.vi.1972 (K);

. above the cemetery, 30 m, 6.vi.1972 (K);

. hills West of the town, 50 m, 2.x.1973 (K);

. wet meadows East of the Town, 2-3 m, 4.x.1973 (K);

v. Oertzen;

g. 2-16.v.1942, Wettstein.

123. Sitia, 5 km East of the town, beach, 0-5 m, 3.x.1973 (K).

124. Palékastron, 100 m, 3.x.1973 (K).

125. Vai:

a. mouth of the brook with Juncus maritimus, below zone of Phoenix theophrasti, 0 m, 3.x.1973

(K);

b. above the zone of Phoenix, 5 m, 3.x.1973 (K).

126. Acra Sideros: Kap Sidero, 14.vii.1925 (S) (approximately).

127. Nisos Gianisäda: Insel Janisada.

128. Nisos Eläsa: Insel Elasa, 1887, v. Oertzen.

0 EO: 0

WILLEMSE & KRUSEMAN: Orthopteroidea of Crete 163

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Chopard, L., 1921. Description d’une espèce nouvelle du genre Troglophilus (Orth. Phasgonuridae). —

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Demirsoy, A., 1973. Revision der anatolischen Pamphaginae (Saltatoria, Caelifera, Pamphagidae). —

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Descamps, M., & Mounassif, M., 1972. La complexe Orchamus, Paracinipe, Acinipe et Pamphagus (Acri-

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Dirsh, V., 1951. Revision of the group Truxales (Orthoptera, Acrididae). — Eos. Madr., Tomo extraor-

dinario 1950: 119—247, Fig.

—_ _, 1954, Revision of species of the genus Acrida Linné (Orthoptera, Acrididae). — Bull. Soc.

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Fieber, F., 1853. Synopsis der europäischen Orthopteren. — Lotos 3: 90—104, 115—129, 138—154,

168—176, 184—188, 201—207, 232—238, 252—258.

Galvagni, A., 1959. Descrizione e corologia della Platycleis concii n.sp., elemento brachittero raccolto

sulle alte Madonie (Sicilia) (Orthoptera, Tettigoniidae, Decticinae). — Memorie Mus. civ.

Stor. nat. Verona 7: 259—268, Fig.

Griffini, A., 1894. Catalogue des Orthoptéres recueillis dans l’île de Candie. — Miscnea ent. 2: 91—92.

Guanella, H., 1972. Kreta, ein Reiseführer, 3. Auflage: 1—416, Fig., map. — Zurich (Flamberg).

Harz, K., 1969. Die Orthopteren Europas. Vol. 1. — Series entomologica, 5: i—xx, 1— 749, Fig. — The

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—— —, 1975. Die Orthopteren Europas. Vol. 2. — Series entomologica, 11: 1—939, Fig. — The Hague

(Junk).

Hollis, D., 1968. A revision of the genus Aiolopus Fieber (Orthoptera: Acridoidea). — Bull. Br. Mus.

nat. Hist. (Ent.) 22: 309—355, Fig.

Jago, N., 1963. A revision of the genus Calliptamus Serville (Orthoptera: Acrididae). — Bull. Br. Mus.

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Johnston, H., 1956. Annotated catalogue of african grasshoppers: i—xxii, 1—833. — Cambridge (Uni-

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Kaltenbach, A., 1964. Orthogenese und geographische Verbreitung bei westpaläarktischen Gottesan-

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Kevan, D. K. McE., 1971. The type-species of the genus Pyrgomorpha Audinet-Serville and proposed

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———, 1974. The identity of Truxalis fuscus Palisot de Beauvois (Orthoptera: Acridoidea) and the ty-

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153—161, Fig.

Kis, B., 1967. Gryllus (Modicogryllus) chopardi — eine neue Orthopteren Art aus Rumänien. — Rei-

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La Greca, M., 1964. Le Tropidopola (Orthoptera, Catantopidae) italiane con osservazioni sulle species

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164 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

Mistshenko, L., 1936. Orthoptera palaearctica critica XII. Revision of palaearctic species of the genus

Sphingonotus Fieber (Orth. Acrid.). — Eos. Madr. 12: 65—282, Fig.

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Fig.

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Mitt. zool. Mus. Berl. 24: 41—50, 150, Fig.

———, 1951. Zur Systematik, Faunistik und Biologie der Orthopteren von Südost-Europa und Vor-

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Salfi, M., 1931. Orthoptera palaearctica critica X. Revision du genre Platypterna Fieber (Acrid.). —

Eos. Madr. 7: 255—347, Fig.

Uvarov, B., 1926. Orthoptera palaearctica critica II. Genus Tropidopola St. (Acrid.). — Eos. Madr. 2:

149— 177, Fig.

———, 1942. New and less known southern palaearctic Orthoptera. — Trans. amer. ent. Soc. 67:

303—361, Fig.

Werner, F., 1901. Die Dermapteren- und Orthopterenfauna Kleinasiens. — Sber. Akad. Wiss. Wien

110: 259—306, Fig.

———, 1903. Ueber die von Herrn Dr. Karl Grafen Attems aus Kreta mitgebrachten Orthopteren. —

Verh. zool.-bot. Ges. Wien 1903: 65—69, Fig.

———, 1927. Zoologische Streifzüge in Attika, Morea und besonders auf der Insel Kreta. I. Orthop-

teren. — Abh. naturw. Ver. Bremen 26: 426—431.

———, 1934. Dritter Beitrag zur Kenntnis der Tierwelt der Ägäischen Inseln. — Sber. Akad. Wiss.

Wien 143: 313—337, Fig.

———, 1937. Ergebnisse der vierten zoologischen Forschungsreise in die Ägäis (1936). — Sber. Akad.

Wiss. Wien 146: 89—118, Fig.

Yersin, A., 1860. Note sur quelques Orthopteres nouveaux ou peu connus d’Europe. — Ann. Soc. ent.

Fr. (3) 8: 509—537, Fig.

Zeuner, F., 1941. The classification of the Decticinae hitherto included in Platycleis Fieb. or Metriop-

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976

=: “iii

PIÙ

Fig. 1—3. Platycleis. 1, P. grisea transiens Zeuner, topotype (Mt. Chelmos, 1700—2000 m, 1.viii.1970,

F. Willemse c.s.). 2, P. grisea cretica subsp. nov., holotype. 3, P. concii Galvagni, topotype (Contrada

Quacella, Madonie, 20.vii.1963, La Greca)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 Pl. 2

rires PTT I trans

Fig. 4—6. Platycleis species. 4, P. grisea transiens Zeuner, topotype (cf. PI. 1 Fig. 1). 5, P. grisea

cretica subsp. nov., allotype. 6, P. concii Galvagni, topotype (cf. PI. 1 Fig. 3). Fig. 7—8, Eupholidoptera

forcipata spec. nov., 7, holotype, 8, allotype

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 Pl. 3

Fig. 9—12. Eupholidoptera species. 9—10, E. latens spec. nov., 9, holotype, 10, allotype. 11—12, E. palli-

pes spec. nov., 11, holotype, 12, allotype

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 PI. 4

Fig. 13—17. Eupholidoptera species. 13, E. gemellata spec. nov., holotype. 14—17, tips of male

abdomen, lateral view. 14, E. forcipata spec. nov., paratype; 15, E. latens spec. nov., paratype; 16,

E. pallipes spec. nov., paratype; 17, E. gemellata spec. nov., holotype, epiphallus removed

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 Pl. 5

Fig. 18—24. Eupholidoptera species. 18—20, last abdominal tergite of males. 18, E. forcipata spec. nov,

paratype; 19, E. latens spec. nov., paratype. 20, E. pallipes spec. nov. paratype. 21—24, male

subgenital plate, ventral view. 21, E. forcipata spec. nov., paratype; 22, E. latens spec. nov, paratype;

23, E. rallipes spec. nov., paratype; 24, E. gemellata spec. nov., holotype

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 BR 6

RR i

Fig. 25—31. Eupholidoptera species, males 25—28, cercus, dorsal view. 25, E. forcipata spec. nov.,

paratype; 26, E. latens spec. nov., paratype; 27, E. pallipes spec. nov., paratype; 28, E. gemellata spec.

nov., holotype. 29—31, E. forcipata spec. nov., paratype, epiphallus

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 PI. 7

Fig. 32—38. Eupholidoptera species, males, epiphallus. 32—34, E. latens spec. nov. paratype; 35—36,

E. pallipes spec. nov, paratype; 37—38, E. gemellata spec. nov, paratype

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 P1.8

as a a =

Fig. 39—41. Eupholidoptera species, females, subgenital plate. 39, E. forcipata spec. nov., paratype; 40,

E. latens spec. nov., paratype; 41, E. pallipes spec. nov., paratype. Fig. 42—43, Tropidopola longicornis

(Fieber) subsp. nov.?, fastigium verticis. 42, 3, Amnissós; 43, 9, Phaistös

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 PI. 9

Fig. 44. Oedipoda caerulescens (Linné), 9: upper series, mainland of Greece (left: Peloponnese, Mikra

Mantinea, S. of Kalamata, 25 m, 20.vii.1971; middle: Peloponnese, Marmara, Erymanthos River,

600 m, 30.vii.1970; right; Sterea, Mt. Iti, near refuge above Ypati, 1800-2000 m, 11.13.viii.1973); lower

series, Crete (left: 48; middle: 17c; right; 50b)

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 5, 1976 PI. 10

Fig. 45—46. Oedipoda venusta Fieber. 45, 9, Anatoli (107b); 46, g, Linoséli (13). Fig. 47—48.

Chorthippus (Glyptobothrus) biroi (Kuthy). 47, 9, Linoséli (13); 48, 3, Omalós (15e)

VER

ven

Fly deat

| ED-T5682

DEEL 119 AFLEVERING 6 1976

TIJDSCHRIFT

VOOR ENTOMOLOGIE

MUS. COMP, ZOO

UITGEGEVEN DOOR LIBRARY >

DEC 91970

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGINGO

i UNIVERSITÀ

INHOUD

R. DE JONG. — Affinities between the West Palaearctic and Ethiopian butterfly

_ faunas, p. 165—215, fig. 110.

- Tijdschrift voor Entomologie, deel 119, afl. 6 Gepubliceerd 25-X-1976

AFFINITIES BETWEEN THE WEST PALAEARCTIC AND

ETHIOPIAN BUTTERFLY FAUNAS

R. DE JONG

Rijksmuseum van Natuurlijke Historie, Leiden

With ten figures

ABSTRACT

A survey is given of the species and genera common to the West Palaearctic and Ethiopian Regions.

The possibilities of a faunal exchange in the past, especially during the Pleistocene, are analysed and

related to the present distribution of West Palaearctic species in the Ethiopian Region and vice versa.

It is demonstrated that faunal exchange across the Saharo-Arabian desert zone was not infrequently

possible, but most invading species died out subsequently. Palaearctic species in the Ethiopian Region

had a better chance to survive than Ethiopian species in the Palaearctic Region. Although the bare

condition of the desert zone at present keeps the Palaearctic and Ethiopian Regions apart, it is

concluded that the main factor inhibiting large-scale faunal exchange during the Pleistocene has been

the repeated change of climate.

CONTENTS

1. Introduction

General ear: earner. OEM ME ING ri. RIESE MAS 166

iitreyPalacarctic-Ethiopianiboundary "er ion i ee dert ed eek 167

2. Range, ecology and relationship of genera and species of butterflies common to the

WestRalaearcticandEthiopian Regions e eenen ee ep ee ee 168

3. The affinities between the West Palaearctic and Ethiopian butterfly faunas ......... 181

Direct affinities

Palaearctic influence in the Ethiopian Region ...................... 182

‘ Ethiopian influence in the Palaearctic Region . ..................... 183

Indirect affinities

Indo-Etmopian'andieremicispecies annen enen sl ee 184

Remainingigeneranges sn 2 e eas oc wee as a cosh, we SEE dad 185

Summaryzandkeonelusionster © Sey cee fee o 185

AMM EiStoricalaspectsmen ks het AR coe seep TERN EI AIRIS I AER es 186

Ecological history of the northern half of Africa and the Arabian peninsula. + + + - ..... 186

ICI VOEDEREN RS SPERO RE RT CTS Te BENE WI AD Ue Debt: 186

Bleistocene un vn a ee Sete) «eset sy ey aie coined: 188

The direct West Palaearctic-Ethiopian affinities

HRESOUTR WA GRIGRS” a ob ele Slee eeeee SSNS ee. 191

Mhemnortnwarditraversene nore eee ee eek yh eran ce el ee cn ay eR) eet eee et 197

The indirect West Palaearctic-Ethiopian affinities

iitetindosEthiopianspeciess is Heyes. EU EME eat Ss SEEN TE 203

Bietesemichsneciëshejdied. nto dotate et babies: (EER oe pepe dats 204

STRING? CCL Vel er aa nr Aen oe einen MERCURE Ra nA) mes inl un a fai 205

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

5. Barriers outsidethe desert zone. e 0 e E 205

The West Mediterranean: 0 + 2 ar sf Sb cu he sl tee A O 205

ThesouthermiRed Sea: 2180 Pa Dina ESE VR ee Ne: 208

6. ‘Summary and'conclusions (i piu = ee use ee RICE EEE 209

References oh e late SIRIO re 212

1. INTRODUCTION

General

This study is a zoogeographic one. It tries to describe the affinities between the

butterfly faunas of the West Palaearctic and Ethiopian Regions and to explain

these affinities with the aid of possible past distributions. As it deals with :

distributions, the ranges of the species have to be known and in this respect the

butterflies form an attractive group, because possibly no insect group is better

known.

This study also deals with history and, therefore, is concerned with evolution.

In this respect, butterflies are probably scarcely better known than any other

group of insects, as by far the greater part of the extensive literature on butterflies

is faunistic (i.e., a description of the composition of species of a certain area) or a

detailed analysis of the wealth of forms. At least the lower classification of the

butterflies is still largely typological and evolutionary studies are almost absent.

This is felt as a hindrance for the present paper, as an evolutionary study may

reveal much of the probable geographic history (cf. De Jong, 1974). Nevertheless,

there is no reason to suppose that an evolutionary study of the butterflies will

necessitate many alterations in the classification and for the present paper I

assume that the generally accepted genera are monophyletic. Further, if a genus

has many representatives in region A, one of which occurs (possibly in another

form) in region B, where it is the sole representative of the genus, it is concluded

that this particular species invaded B from A. In some cases, viz. migrants, there is

direct evidence of such invasions. In other cases, evidence or at least support

should be looked for in an evolutionary study and in this way the present paper

may serve to indicate some of the many gaps in our knowledge.

I have attempted to give the total number of species known for each genus, but

have often failed to find agreement in literature. This is due partly to (sometimes

excessive) splitting of genera and species, partly to the fact that, usually, regional

faunas instead of whole genera have been studied and I cannot help thinking that

various degrees of “intuition” have played a part. It is, for instance, difficult to

estimate the number of species in the genus Lycaena (the Coppers). Kostrowicki

(1969) lists 30 species in the Palaearctic Region, but generally this genus is divided

into four genera (Heodes, Lycaena, Thersamonia and Palaeochrysophanus) in the

West Palaearctic. For Central Europe, Forster & Wohlfahrt (1955) place the

species helle and phlaeas in Lycaena s.s. and dispar in Thersamonia, while Higgins &

Riley (1970) consider dispar congeneric with phlaeas. If there is so much

disagreement in the best known part of the world, how about the Asiatic species,

for which Verity (1943) introduced two more genera. In such cases it seems most

practical to record the number of species in the collective genus, in the above case

DE JONG: West Palaearctic and Ethiopian butterfly faunas 167

Lycaena s.l. In the text it is specified how the total number of species in a genus has

been arrived at. It is not of crucial importance for the present study. So far as the

genera can be considered monophyletic, large genera may be preferable for the

present kind of study, as they may give a good impression of the relative

development of a group of species in various regions, e.g. the very strong

development of the Coppers (Lycaena s.l.) in the Holarctic Region as opposed to

the very poor representation outside this area. On the other side, however, it is

also indicative to observe that the Coppers in the Holarctic Region can be divided

into six genera, while only one of these genera, with a single group of species,

occurs in the Ethiopian Region. An advantage of the use of many small genera is

that it is easier for a non-specialist to trace closely related species (if the small

genera are considered monophyletic, what can only be assumed as they are usually

based typologically).

The affinities between the West Palaearctic and Ethiopian butterfly faunas have

been studied up to the generic level, i.e. only common genera have been

considered. The affinities based on related instead of common genera have been

regarded too remote to be studied successfully on the basis of the present-day

knowledge of both the phylogeny of the butterflies and the geographic and

climatic changes in the more remote past. Another limitation to the present paper

is the area studied. As is known many Oriental genera penetrate the Palaearctic

Region in East Asia. Some of these genera (e.g. Graphium, Appias, Mycalesis,

Deudorix) also occur in the Ethiopian Region and in such cases it is probable that

the affinities between the Palaearctic and Ethiopian Regions are indirect, via the

Oriental Region. To exclude such cases, only the butterfly genera occurring in the

West Palaearctic Region, west of about 50° E Long. (thus including Europe and

the Middle East), have been considered.

The Palaearctic-Ethiopian boundary

There is some disagreement about the boundary between the Palaearctic and

Ethiopian Regions. Darlington (1957), for instance, delimited the Ethiopian

Region as the whole of Africa except the parts of Morocco and Algeria north of

the Atlas Mountains, while De Lattin (1967) sets the boundary along the southern

edge of the Sahara. There is no reason to enter here into a discussion on the use of

distinguishing zoogeographic regions, but as descriptive, geographic terms they

have at least practical value, provided they are clearly defined. The obstacle in

defining the boundary between the Palaearctic and Ethiopian Regions is the

position of the Sahara. Although often the peculiar character of its fauna is

recognized, the Sahara is still usually considered a transition zone. Geographically

the Sahara is, indeed, the transition between the tropical parts of Africa and the

warm-temperate parts of the northern part of the Old World. Ecologically,

however, speaking in terms of major habitats, the Sahara is a life zone, a biome,

comparable in this respect to, e.g., the taiga or the sea. I never saw the taiga

considered the transition between the deciduous forests to the south and the

subarctic birch forest to the north. And there is apparently little meaning in calling

the sea the transition zone between two land masses cr in regarding Africa the

168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

transition zone between the Atlantic and Indian Oceans, just because it lies in

between.

The zoogeographic distinction of the Palaearctic and Ethiopian Regions is based

on a different faunal composition. This difference can, at least partly, be explained

as the result of a separated faunal development, i.e. there has been a barrier

preventing free faunal exchange. Evidently the main barrier has been and is the

Sahara.

These considerations lead to the assumption that the boundary between the

Palaearctic and Ethiopian Regions does not run north or south of or straight

across the Sahara, but is the Sahara itself. Studying the affinities between the

butterfly faunas of the Palaearctic and Ethiopian Regions is studying the function

of the Sahara as a barrier, at present and in the past. As a consequence, only those .

genera and species are considered common to both regions, which occur north as

well as south of the Sahara.

The area studied has been extended to the east to cover the Arabian peninsula,

extending the barrier to the Saharo-Arabian desert zone and regarding Yemen the

north-eastern outpost of the Ethiopian Region.

2. RANGE, ECOLOGY AND RELATIONSHIP OF GENERA AND SPECIES OF BUTTERFLIES

COMMON TO THE WEST PALAEARCTIC AND ETHIOPIAN REGIONS

Papilio. A cosmopolitan genus with a large development in the tropics, but with

only three species in the West Palaearctic, viz. machaon L., hospiton Géné, and

alexanor Esper. The Ethiopian fauna comprises about 50 species. Over the whole

more than 200 species are known. The West Palaearctic species are placed by

Munroe (1960) in the machaon group, which ranges throughout the Holarctic

Region and into Central America. Together with the East Palaearctic xuthus L.,

they are thought to be closely related to the demoleus group, which occurs in the

Ethiopian and Oriental Regions and extends eastward to Papua.

Pieris. Much disagreement exists in the delimitation of the genera Pieris and

Pontia. Part of the disagreement is shown in Table 1. The number of species

allocated to Pieris varies from about 18 to about 26, see e.g. Talbot (1932a) and

Bernardi (1947). Of these six to nine are confined to the New World. In the Old

World the genus is almost strictly Palaearctic, but some species extend far into the

Oriental Region (canidia Sparrman, melete Ménétriés) or occur near the edge of

the Palaearctic (naganum Moore). One species, rapae L., has been imported as far

as New Zealand (Laidlaw, 1970). Apparently, an Oriental development of the genus

is absent. In the Ethiopian Region there is a single, endemic species, viz.

brassicoides Guérin, which is restricted to the montane grasslands of Ethiopia and

N. Tanzania (Carcasson, 1964). The latter population was distinguished as ssp.

meridionalis Joicey et Talbot. According to Lagnel (1966) brassicoides can be

considered to form a superspecies together with brassicae L. (Canary Islands, N.

Africa and W. Europe to Yunnan) and deota De Niceville (Issyk-Kul to Ladak).

The common European brassicae (Large White) can be found everywhere in

flowery places from lowlands up to 2000 m (Fig. 3).

Pontia. Part of the disagreement on the delimitation of this genus is shown in

DE Jong: West Palaearctic and Ethiopian butterfly faunas 169

Table 1. We are here mainly interested in the first four species on which there is

no disagreement with regard to the genus they belong to and which as a group

occur in the Ethiopian and Palaearctic Regions. But to place their evolution in a

wider perspective, also the other, probably closely related species listed in Table |

Table 1. Allocation of species to Pontia and related genera by various authors

Talbot Verity Bernardi Dos Passos Higgins & Riley

(1932a) (1947) (1947) (1964) (1970)

daplidice Pontia Pontia Pontia = Pontia

glauconome Pontia Pontia Pontia — —

distorta Pontia Pontia — — —

helice Pontia Pontia — = =

chloridice Pontieuchloia Pontieuchloia Pieris — Pontia

beckeri Pontieuchloia Pontieuchloia Pieris Pieris Pontia

protodice Pontieuchloia Synchloe Pieris Pieris —

sisymbrii Pontieuchloia —Synchloe Pieris Pieris —

callidice Synchloe Synchloe Pieris — Pontia

dubernardi Synchloe Synchloe Pieris — —

davidis Synchloe Synchloe Pieris — =

nigricans Synchloe Synchloe Pieris — —

have to be considered and without choosing for one of the classifications

mentioned the following observations may be important for the present study:

a. All species of Table 1 are confined to some part of the Holarctic Region,

except the first four species, two of which are Ethiopian, while the other two occur

in both the Ethiopian and Palaearctic Regions.

b. Pontieuchloia is considered intermediate between Pontia and the almost

exclusively Holarctic Euchloe (Verity, 1947).

c. According to Verity (1947) the last three species of Table 1 are intermediate

between Synchloe and the Palaearctic genus Aporia.

All these observations point to an almost entirely Holarctic group of species

with a slight development in the Ethiopian Region. As related genera occur in the

Palaearctic and not in the Ethiopian Region, the Palaearctic origin of the

Ethiopian members seems obvious.

The ranges, etc., of the first four species of Table 1 are as follows:

daplidice L. — From France to Japan (Higgins & Riley, 1970); Canary Islands

(Van Regteren Altena, 1949); N. Africa south to Tibesti (Bernardi, 1962). In all

these regions there is little geographic variation (though various subspecies have

been described), possibly due to the fact that the species, at least in Europe, is

strongly migratory. A separate subspecies (aethiops Joannis et Verity) is recorded

from Ethiopia, Somalia and Yemen (Gabriel, 1954). P. daplidice prefers rough

grounds and flowery meadows.

glauconome Klug. — An eremic species occurring from Mauretania to Ethiopia,

Kenya, Somalia and Sinai and through Arabia and Mesopotamia to NW. India.

The African and Arabian populations are thought to be subspecifically distinct

(ssp. glauconome Klug) from the more eastern populations (ssp. iranica Bien.).

distorta Butler. — Restricted to Somalia and Kenya (Aurivillius, 1910—25).

Probably also a more or less eremian species.

helice L. — A common species throughout S. Africa (Swanepoel, 1953), north to

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

S. Botswana and S. Rhodesia (Van Son, 1949). Further, in the montane grasslands

of E. Africa (Kivu, Tanzania, Uganda, Kenya) as a separate subspecies (johnstoni

Crowley) (Fig. 5).

Euchloe. This genus is divided by Bernardi (1945) and Verity (1947) into two

subgenera, viz. Elphinstonia and Euchloe. Higgins & Riley (1970) consider

Elphinstonia a separate genus, but their allocation of species to Euchloe and

Elphinstonia is different from Bernardi’s and Verity’s. This difference of opinion is

unimportant for the present study and we will distinguish a single genus, like

Bernardi did, but without bothering about subgenera.

The genus Euchloe comprises ten species. Three species are confined to N.

America; the remainder are Palaearctic, but two species also occur in the .

Ethiopian Region, viz.:

belemia Esper. — SW. Europe, Canary Islands, N. Africa (also Tibesti) to Iran

and Baluchistan (Higgins & Riley, 1970). In the Ethiopian Region it is known from

Ethiopia and Somalia (Talbot, 1932b; Carpenter, 1935). The Ethiopian populations

are distinguished as ssp. abyssinica Riley. This species prefers rough, flowery

places, usually at low levels. It is a strong flyer; in S. Spain I had no difficulty in

distinguishing this species among other spring species like Pontia daplidice, Euchloe

ausonia and Pieris rapae, by its flight only.

falloui Allard. — More eremic than belemia, ranging far into desert oases and

occurring throughout N. Africa, including Tibesti, Fezzan, Tripoli, and Egypt, but

in the north-west mainly south of the Atlas Mountains (Higgins & Riley, 1970). In

the Ethiopian Region it has been found in Somalia. According to Talbot (1932a)

the Somalia population represents the same subspecies as which flies in S. Algeria

(obsolescens Rothschild). Gabriel (1954) recorded this species from SW. Arabia,

with the comment that this population may represent a separate subspecies.

Colotis. An Indo-African genus, with 40 Ethiopian species (Carcasson, 1964) and

seven species in India and Ceylon (Wynter-Blyth, 1957). Almost all species occur

in open and usually dry country and many species live partly or exclusively in

semidesert environments. There are no endemic Palaearctic species, but some

species penetrate into the southern parts of the Palaearctic. They are:

evagore Klug. — Throughout the Ethiopian Region (Williams, 1969) and in

Yemen (Gabriel, 1954); as a separate subspecies, nouna Lucas, in Morocco,

Algeria, Tunis and S. Spain (Higgins & Riley, 1970) (Fig. 7).

fausta Oliv. — India to Egypt and SW. Arabia; north to Israel and Jordania

(Hemming, 1932) and, as a migrant, in Lebanon and sometimes Turkey (Larsen,

1974). Although it is a common species in Lebanon, fausta appears unable to

hibernate there.

calais Cramer. — Through most of the Ethiopian Region outside forest areas. In

the Oriental Region in India and Ceylon. It occurs in Yemen (Gabriel, 1954) and

seems to be resident as far north as Ennedi (Bernardi, 1964) and the Air

Mountains (Rothschild, 1921) in the Sahara, and the Jordan Valley (Larsen, 1974).

Talbot (1934) considered the E. Mediterranean form indistinct from the Indian

one, but according to Gabriel (1954) and Larsen (1974) the African form flies

there.

De JONG: West Palaearctic and Ethiopian butterfly faunas 171

phisadia Godart. — NW. India, Arabia to Nubia, Ethiopia, Somalia, Kenya

Senegal (Talbot, 1934) and Rio de Oro (Bernardi, 1966). Also known from Ennedi,

Tibesti, Ahaggar and Air (cf. Table 5) in the Sahara. Talbot mentioned

Madagascar but this seems an improbable locality. In Israel and Jordania it flies as

ssp. palaestinensis Staudinger (Hemming, 1932; Talbot, 1934).

chrysonome Klug. — According to Talbot (1934) from S. Israel and Jordania

through Arabia, Nubia and Somalia to Tanzania and N. Rhodesia; further in N.

Nigeria and Rio de Oro (Bernardi, 1966) and mentioned from Ennedi, Tibesti,

Ahaggar and Aîr in the Sahara (cf. Table 5).

Anaphaeis. A palaeotropical genus with two species in the Indo-Australian

Region and seven in Africa and Madagascar. There are no endemic Palaearctic

species. The species aurota Fabr. occurs throughout Africa south of the Sahara, in

a variety of habitats, from forest edges to semidesert bush (Williams, 1969).

Further north it occurs in Sudan, Egypt and Arabia (Gabriel, 1954) and, as a

migrant, in Lebanon (Larsen, 1974) and Jordania (Hemming, 1932). It extends

further east through India to Ceylon (Wynter-Blyth, 1957).

Catopsilia. This genus has a peculiar distribution. According to Talbot (1932a)

there are three species in Madagascar (and Mauritius) and five in the Indo-

Australian Region, while one species, viz. florella Fabr., flies throughout the

Ethiopian Region and extends via Egypt and Arabia to India, China and Hainan.

Only since 1966 it is known from the Canary Islands where it has become

established probably as a result of migration (Manley & Allcard, 1970). It has been

found as a migrant in Lebanon (Larsen, 1974) and Iraq (Wiltshire, 1957).

Colias. This genus reaches its greatest development in the Holarctic Region

where about 50 species occur (36 in the Palaearctic). In the Neotropical Region 12

species are known. Although 14 species have been found in India, they cannot be

regarded Oriental residents, as they are confined to the higher altitudes of the

Himalayas, with the exception of erate Esper (Wynter-Blyth, 1957). The literature

is somewhat confused about the identity of the Indian erate, but that is not

important here. The Indian erate occurs in NW. India (Chitral to Kumaon) and in

the S. Indian mountains.

In the Ethiopian Region two species of Colias are known, viz.:

erate Esper. — Palaearctic, from E. Europe to Japan and Formosa (Higgins &

Riley, 1970) and in India (see above). Further in Ethiopia, Somalia, Sudan, Egypt

and SW. Arabia (Gabriel, 1954) (ssp. marnoana Rogenhofer).

electo L. — According to Williams (1969) distributed over the greater part of

Africa south of the Sahara, in grasslands, along forest edges, etc., but I have not

found data on its occurrence in W. Africa. Outside Africa, it hi been found in

Yemen en Saudi-Arabia (Gabriel, 1954). Aurivillius (1910—25) and Van Son (1949)

regarded electo conspecific with crocea Geoffroy, but various authors (e. g.

Lempke, 1944 (1946); Jarvis, 1953) concluded on typological grounds that crocea

and electo are specifically distinct. C. crocea is a well-known migrant in Europe

which occurs from the Azores, Madeira and Canary Islands through N. Africa and

S. and C. Europe to Iran (Higgins & Riley, 1970). In N. Africa crocea goes as far

south as Tibesti (Bernardi, 1962).

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Jarvis (1953) suggested that the E. African form of electo, pseudohecate Berger,

could be a species separate of electo, but we will not busy ourselves with such

speculations.

Note. I have no special knowledge of the genus Colias, but it appears to me that

the Ethiopian species can be considered to belong to two species groups, viz.:

a. the hyale group, comprising hyale L. (C. Europe to Altai; strongly migratory),

australis Verity (S. and C. Europe to S. Russia and Turkey; possibly migratory) and

erate Esper (E. Europe to Japan and Formosa; south to Ethiopia, Somalia, S.

India).

b. the crocea group, comprising crocea Geoffroy (W. Palaearctic to Iran;

strongly migratory), fieldii Ménétriés (N. India to N. Burma, at higher altitudes)

and electo (Ethiopian Region to Arabia) (cf. Lempke, 1944 (1946); Jarvis, 1953).

Classified in this way it is striking that these groups are the only groups of Colias

species in the Old World represented outside the Palaearctic Region, and that

both possess at least one species that is strongly migratory. It seems inevitable to

consider the Ethiopian species Palaearctic invaders.

Danaus. A pantropical genus with about 60 species. A few species occur in the

Nearctic Region, a few others penetrate the Palaearctic Region in the east and

two have been found in the W. Palaearctic, mainly as migrants. They are:

plexippus L. — An American species that since 1880 is resident in the Canary

Islands. It also occurs in the Azores. Rare vagrants reach Portugal, Spain, France,

Ireland and England (Higgins & Riley, 1970). This species has extended its range

from America particularly in western direction, largely aided by man, and it now

occurs in Hawaii, New Zealand, Australia, Papua, Indonesia, Formosa and the

Andaman Islands (Klots, 1951; Higgins & Riley, 1970). It is absent from Africa and

continental Asia.

chrysippus L. — A very common species throughout the Ethiopian Region, in

open and bush country, gardens, woodlands, margins of forests (Williams, 1969).

Resident in the Canary Islands. North to Lebanon and thence eastward

throughout tropical Asia to Australia; vagrants occasionally found in Morocco, S.

Italy and Greece (Larsen, 1974). In Africa it occurs in two subspecies:

a. ssp. alcippus Cramer. — W. Africa south of the Sahara, north to S. Morocco

and also found in Aîr, Tibesti and Ennedi in the Sahara.

b. ssp. chrysippus L. — Whole of Africa, except the west; eastern Mediterranean

and thence eastward. According to Bernardi (1962) this form also occurs in the

Canary Islands, but Manley & Allcard (1970) list the form of that area as a distinct,

endemic subspecies (kanariensis Fruhst.).

Libythea. A peculiar, cosmopolitan genus with about ten species (the two New

World species are classified by Dos Passos, 1964, in a separate genus).

Pagenstecher (1901) distinguished two subgenera in the Old World, viz. Libythea

with six species, in Europe, Asia and Australia; and Dichora with three species, in

the Ethiopian Region (one confined to Madagascar and Mauritius). Williams

(1969) considered one of the Ethiopian species of Pagenstecher (/aius Trimen) a

subspecies of the Ethiopian /abdaca Westwood, and according to Corbet &

Pendlebury (1956) labdaca is the Ethiopian representative of the Oriental narina

De Jong: West Palaearctic and Ethiopian butterfly faunas 173

Godart, so that the distinction of two subgenera is insignificant.

In the West Palaearctic only celtis Laich. occurs. Its range extends from S.

Europe and N. Africa to Japan and Formosa. From the above it seems improbable

that there is a direct relationship between the West Palaearctic and Ethiopian

species.

Charaxes. A palaeotropical genus with 86 species in Africa south of the Sahara

(Carcasson, 1964) and more than 20 in the Indo-Australian Region. There is only a

single species occurring in the Palaearctic Region and this species also flies

throughout the Ethiopian Region, viz. jasius L. (Fig. 9). It was apparently by

mistake that Carcasson called Charaxes a Palaearctic genus.

The distribution of jasius in the Palaearctic is confined to the coastal areas

around the Mediterranean. South of the Sahara, jasius occurs in two distinct

geographic forms all over the continent, in savanna country, woodlands and bush.

Neptis. Apart from a large development in the Ethiopian Region (47 species;

Carcasson, 1964), there are many species confined to the Indo-Australian area (31)

and to the Palaearctic (18), while 20 species occur in both the Palaearctic and

Oriental Regions (Eliot, 1969). Most Palaearctic species are confined to East

Asia, often to the contact zone with the Oriental Region; only two species occur as

far west as Europe:

sappho Pallas. — A woodland species, distributed from C. Europe (Salzburg) to

the Amur region, Japan, Formosa and S. Vietnam, in a number of subspecies.

rivularis Scopoli. — Also a woodland species, from the Southern Alps

(Piedmont) to Kamchatka, Kurile Is., Japan, Formosa and Szechwan, in a number

of subspecies.

There is no apparent reason to suppose a close relationship between the two

mentioned species and the Ethiopian members of the genus. On the contrary, the

European species appear Asiatic invaders in Europe, judged from their present

distribution.

Hypolimnas. A palaeotropical genus with a twenty odd species. Two species

occur in Africa as well as in the Indo-Australian area and one of these, misippus L.,

is rarely found as a migrant in Lebanon, probably coming from Egypt, where it is

common (Larsen, 1974). This species has a very extensive range. It occurs from

Madagascar and Africa (where it is common everywhere in open country, bush,

etc.; Williams, 1969) to India, SE. Asia, New Guinea, Solomons Islands, and

Australia. It was supposedly introduced into the Antilles from Africa in slave ships

some centuries ago and has been found not only in the West Indies, but also north

into the USA and in northern S. America (Common & Waterhouse, 1972).

Vanessa. This genus was recently dealt with by Field (1971), who thought it

necessary to split Vanessa in three genera, viz. Vanessa, Bassaris and Cynthia. In

my opinion this splitting is somewhat superfluous, as for the reflection of the

affinities within the genus the distinction of species groups or subgenera would

have done and nomenclatorial troubles would have been avoided. I deal with the

genera of Field here as subdivisions of the genus Vanessa s.l. This genus is, with 16

species, distributed all over the world. Four species have been found in the West

174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Palaearctic and one of these is the sole representative of the genus in the

Ethiopian Region. They are:

atalanta L. — According to Field (1971), a Holarctic species that is introduced

in Hawaii, but Higgins & Riley (1970) thought its Palaearctic range not to extend

eastward beyond Iran. No subspeciation within the Palaearctic. It is a well-known

migrant in Europe and in the northern part of its range it reintroduces itself each

year by migrations from the south.

Together with the following and three other species, atalanta belongs to Vanessa

s.s., which is distributed in the Holarctic and Oriental Regions.

indica Herbst. — This species shows a peculiar distribution: it is widely

distributed in India, China, Korea and Japan, and quite separated from this range

it occurs in Madeira and the Canary Islands (distinguished as ssp. vulcanica .

Godart).

cardui L. — The most cosmopolitan butterfly, found everywhere, except in

Australia and New Zealand (where it is replaced by kershawi M’Coy) and in arctic

and antarctic regions. It is rare in S. America south of Venezuela. This species is

strongly migratory and this may be the reason why there is no geographic

variation, though subspecies have been described (see Field, 1971). In much of

Europe, cardui cannot hibernate, but can produce a summer generation. It is the

only Vanessa species in Africa. With the Australian kershawi and seven New World

species cardui forms what Field called the genus Cynthia.

virginiensis Drury. — One of the New World species of Field’s Cynthia, occurring

from Canada to Colombia and Venezuela and possibly much further south (Field,

1971). Field recorded this species to have become established by migration in

Hawaii, the Azores, Madeira and the Canary Islands, but Higgins & Riley (1970)

and Manley & Allcard (1970) only mentioned the Canary Islands, as far as the

Palaearctic is concerned. Since 1948 many specimens have been found in Portugal

and some also in Spain, as far as the Pyrenees (Gomez Bustillo & Rubio, 1974). No

subspecies are known.

Precis. In the Old World this genus is almost entirely confined to the tropics, but

in the New World it extends over North and South America. From the Ethiopian

Region 29 species are known, from the Indo-Australian Region 11, some of which

have a palaeotropical distribution. Only two species have been found, at least one

as a migrant, in the Palaearctic region, viz.:

hierta Fabr. — An abundant species throughout the Ethiopian Region in open

country (Williams, 1969); Arabia (Gabriel, 1954); eastward through India (where it

is very common; Wynter-Blyth, 1957) to Ceylon and Hongkong (Fruhstorfer,

1912). In the Palaearctic it has only been found in the Lebanon, some 60 years ago.

Larsen (1974) thinks that it may have been imported, rather than having made its

own way to the Lebanon.

orithya L. — Another palaeotropical species, abundant in open, dry country

throughout the Ethiopian Region (Williams, 1969) and extending eastward through

Arabia and India to Australia (Gabriel, 1954). It is an immigrant in Iraq, where it

prefers oases and river banks (Wiltshire, 1957), but I have not found indications

about the origin of the Iraqi immigrants.

De JONG: West Palaearctic and Ethiopian butterfly faunas 175

Issoria. As delimited by Warren (1956) this genus comprises three species

groups, a Palaearctic, an Ethiopian, and a S. American one. Only one of the three

Palaearctic species occurs in the W. Palaearctic, viz. lathonia L. (Canary Islands,

N. Africa and W. Europe to Szechwan and through the Himalayas to Bhutan;

Higgins & Riley, 1970; Fruhstorfer, 1912); the other two Palaearctic species are

confined to Asiatic mountains.

The Ethiopian species group is composed of three species, viz. hanningtoni

Elwes, smaragdifera Butler, and excelsior Butler, distributed over the mountains of

E. Africa, south to Malawi, and the Cameroon highlands.

The S. American group numbers five species and is mainly confined to Chili

(southward to Tierra del Fuego) and higher parts of the Bolivian and Peruvian

Andes (Lehmann, 1913).

The affinities between the species groups are rather obscure. Warren (1956: 390)

writes: “It has been shown that the subdivision of Issoria is unjustified, the various

types of the tenth abdominal segment testify to a close affinity which the world-

wide distribution and complete segregation of many species has failed to

obliterate”. However, such a remark suggests that the groups are conjoined on

account of the common possession of plesiomorphous character conditions, a

poor basis for monophyly.

Although the phylogenetical affinities are obscure, the geographical affinities

seem somewhat clearer. As related genera are absent in Africa but numerous in

the Palaearctic Region, the African species group appears to be an Ethiopian

offshoot of Palaearctic origin. Moreover, the most usual foodplants of the

Palaearctic Issoria and related genera, Viola species, are also the foodplants of

the Ethiopian species. The genus Viola is with 400 species mainly distributed in

the temperate regions.

It seems possible that the Ethiopian and S. American groups are relics of an

once larger, Holarctic group, driven away to outlying districts by a strong

development of other (? new) genera (Argynnis, Brenthis, Boloria, etc.) in the

Holarctic Region, or they could maintain themselves in Africa and S. America,

because they had already colonized the mountains of these continents before more

successful genera in the Holarctic almost completely replaced their parental stock.

Melitaea. One species, abyssinica Oberthür, belonging to the otherwise comple-

tely Palaearctic didyma group (27 species; Higgins, 1941) is the sole representative

of this genus in the Ethiopian Region. It occurs in Ethiopia (Carpenter, 1935) and

as a separate subspecies (scotti Higgins) in SW. Arabia (Gabriel, 1954).

Ypthima. This genus numbers more than 80 species in the Ethiopian, S. and E. °

Palaearctic and Oriental Regions, and a single species further east, in New Guinea

and Australia (Common & Waterhouse, 1972). The only representative in the W.

Palaearctic is asterope Klug. It is distributed throughout the Ethiopian Region, in

wooded areas, bush, savanna, grasslands (Williams, 1969), and through Arabia and

Syria to India, W. and C. China (Gabriel, 1954). As contrasted with what was

claimed by Larsen (1974), asterope does not reach Australia (cf. Common &

Waterhouse, 1972). In the W. Palaearctic, asterope occurs as far north as the

Adana district in Turkey (Higgins, 1966; Larsen, 1974).

176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Hipparchia. There is some disagreement on the delimitation of this genus. It has

many Palaearctic species. A single species, tewfiki Wiltshire, occurs in SW. Arabia

(Gabriel, 1954), where it is endemic. Its closest relative is uncertain (possibly

statilinus Hufn.?), but in view of the distribution of the genus, tewfiki undoubtedly

has a Palaearctic ancestor.

Lasiommata. Like the related genera, which formerly were united with this

genus into the collective genus Pararge, Lasiommata is purely Palaearctic, except

for two species, felix Warnecke, which is confined to SW. Arabia, and maderakal

Guérin, an endemic species from Ethiopia (Fig. 2). Both species look very much

like maera L., which extends from N. Africa and W. Europe to the Himalayas and

De Lattin (1967) stated that these species are closely related. Indeed, I suppose the .

specificity of felix and maderakal is typologically, not biologically based.

Virachola. This genus, which could be considered a subgenus of Deudorix, is

Indo-African and has 27 species in the Ethiopian Region (Stempffer, 1967). One of

these species extends northwards into the Palaearctic:

livia Klug. — An eremic species, known from Tanzania, Somalia, Nubia, Arabia,

Israel (Gabriel, 1954), and attaining its northern distribution limit in Lebanon

(Larsen, 1974). It is migratory.

Apharitis. This genus was erected by Riley (1925) to receive nine eremic species,

distributed from Ghana through the southern Sahara and Arabia to N. India. Their

ranges are as follows (data from Riley, 1925, unless stated otherwise):

gilletti Riley. — Somalia.

nilus Hewitson. — S. Sudan to N. Nigeria and N. Ghana.

buchanani Rothschild. — N. Nigeria, S. Sahara.

acamas Klug. — C. Sahara (Ahaggar Mountains), Sudan, Egypt, SW. Arabia,

north through Israel, Jordania and Lebanon to Cyprus and Taurus (Hemming,

1932; Larsen, 1974), east to Chitral.

myrmecophila Dumont. — Tunisia; Jordania and SE. Arabia (Hemming, 1932).

epargyros Eversmann. — Kurdistan to Baluchistan and northward to the Kirghis

Steppes and Kuldja.

cilissa Lederer. — N. Syria, Kurdistan; Israel (Larsen, 1974).

maxima Staudinger. — N. Syria, Kurdistan.

lilacinus Moore. — Punjab to Assam.

From these data it appears that Apharitis is neither an Ethiopian nor a

Palaearctic genus, but belongs to the eremic zone between the Ethiopian and

Palaearctic Regions and penetrates into those regions.

Tarucus. This Indo-African genus was revised by Bethune-Baker (1918), who

distinguished 20 species. Afterwards three species have been added. Stempffer

(1967) lists 12 species from the Ethiopian Region. According to Clench (1965)

“these small blues are essentially desert or subdesert species”. Nevertheless, some

species can live in the Palaearctic Region:

theophrastus Fabr. — From S. Spain through N. Africa, Turkey, Arabia and Iraq

to India (Higgins & Riley, 1970), Burma and Ceylon (Swinhoe, 1905—10). In

DE JONG: West Palaearctic and Ethiopian butterfly faunas 177

Africa distributed from the Mediterranean to Senegal, N. Nigeria and Ethiopia

(Clench, 1965).

rosaceus Austaut. — Algeria, Tunis, and desert oases in N. Africa and Arabia,

widely distributed in W. Asia to Iraq and Iran (Higgins & Riley, 1970). In Africa

rosaceus extends south to Senegal, N. Nigeria and Kenya (Clench, 1965).

balkanicus Freyer. — Coastal regions of Algeria and Tunis, and from the

Adriatic coast through the Balkans and Turkey to Lebanon and Iran (Higgins &

Riley, 1970). It also occurs in SW. Arabia (Gabriel, 1954). According to Larsen

(1974) balkanicus occurs through “most of eremic Africa”, but I have not found

any confirmation of this statement.

The genus extends as far south as S. Africa, where three species occur

(Swanepoel, 1953); in the Oriental Region it reaches Borneo.

Lampides. A monotypic genus. Its single species, boeticus L., is strongly

migratory and occurs with little or no geographic variation throughout the hotter

parts of the Old World. In the east it extends to Australia, where it is common

(Common & Waterhouse, 1972) and since 1965 it has been found in New Zealand

(Laidlaw, 1970). It is common throughout Africa and is found as a resident in S.

Europe, but migrants have been observed as far north as N. Germany.

Cyclyrius. Only two species are placed in this genus, viz. mandersi Druce, which

is confined to Mauritius, and webbianus Brullé, which flies in the Canary Islands.

This peculiar and apparently relic distribution becomes somewhat more under-

standable if we know that Cyclyrius is thought to be closely related to the next

genus, Syntarucus (see Stempffer, 1967; Eliot, 1973), which may have replaced

Cyclyrius in continental Africa.

Syntarucus. Stempffer (1967) lists 11 species from the Ethiopian Region, five of

which are restricted to islands. In the Oriental Region a single species, plinius

Fabr., occurs, from India to Java (Swinhoe, 1910—11; Wynter-Blyth, 1957). One

of the Ethiopian species, pirithous L., is also found (subspecifically indistinct)

throughout S. Europe and much of Turkey. Larsen (1974) considered it an “Afro-

eremial species”, but it is certainly not strictly “eremial”’. According to Boorman

(1970), it is a common species of forest and savanna areas in W. Africa, and

Swanepoel (1953) states that pirithous (he used the junior synonym telicanus)

occurs literally everywhere in S. Africa.

It is interesting to note that no Syntarucus species are known from the Canary

Islands and Mauritius, where the Cyclyrius species are found, although many

islands around Africa have been colonized by Syntarucus species.

Chilades. There appears to be much disagreement on the delimitation of this

genus, also in recent times. Clench (1965) stated: “This small genus of about eight

species, most of which are Australian, has two species occurring in Africa. ..”.

Stempffer (1967) placed seven Ethiopian species in this genus, although he, in the

description of the genus Freyeria, stated that Chilades occurs from Egypt to the

New Hebrides. However, neither D’Abrera (1971) nor Common & Waterhouse

(1972) mentioned this genus from the Australian Region. There is only a single

Oriental species of which I am sure, viz. laius Cramer, as it is the type species of

178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

the genus; it occurs in India, Ceylon, Burma and S. China (Swinhoe, 1905—10).

A single species, galba Lederer, is found in the Palaearctic Region. It is confined

to the Middle East and has been found in S. Turkey, Lebanon, Jordan, Israel,

Arabia, Iraq (Larsen, 1974). According to Wiltshire (1957), it is an oasis and desert

insect. Due to the chaotic classification of Chilades, it is impossible to state

whether galba is more closely related to Ethiopian or to Oriental species.

Freyeria. As with the foregoing genus it is obscure which species can be assigned

to this genus. In Africa there are two species (Stempffer, 1967). One of them is

confined to Madagascar (minuscula Aurivillius), the other (trochilus Freyer) is

widespread in the Ethiopian Region, occurring in the savannas from W. to E.

Africa and southwards to the Cape; it is also known from S. Cameroon and Gabon

(Clench, 1965). Outside the Ethiopian Region trochilus flies from Egypt and Arabia

to Ferghana and through the Oriental Region to the Philippines and Australia

(Common & Waterhouse, 1972). In the W. Palaearctic it occurs through Lebanon

and Iraq north to Turkey and the Balkans. In Europe it is found on barren stony

ground (Higgins & Riley, 1970), and Larsen (1974) and Wiltshire (1957) mention

comparable habitats in Lebanon and Iraq.

Azanus. Another Lycaenid genus common to the Ethiopian and Oriental

Regions. Stempffer (1967) records seven Ethiopian species, while four species are

mentioned by Swinhoe (1910—11) from India to Sumatra. One of these species,

jesous Guérin, occurs in the Palaearctic, in addition to an Afro-Oriental

distribution. Its distribution is as follows: Ethiopia to the Cape, Madagascar; in N.

Africa in Morocco and Egypt; Arabia to Lebanon; further east in India, Ceylon

and Burma (Higgins & Riley, 1970; Larsen, 1974; Wynter-Blyth, 1957). Larsen

called it an Afro-eremic species, but he was probably unaware of its occurrence in

the Oriental Region.

Zizeeria. A small genus of four or five species (Chapman, 1910; Common &

Waterhouse, 1972); two species are confined to the Oriental Region, one to the

Australian Region, and the fourth, knysna Trimen, is palaeotropical, occurring

from S. Africa to Oceania. This widespread species is also found in the W.

Palaearctic (Fig. 8). It can be devided into two forms, which only differ in the male

genitalia and are, by some authors, considered separate species. These forms are:

knysna Trimen. — Throughout the Ethiopian Region, including Madagascar,

Mauritius, Seychelles, north to SW. Arabia in the east (Gabriel, 1954), to Canary

Islands and through Algeria and Morocco to northern Spain (Gomez Bustillo &

Rubio, 1974) in the west.

karsandra Moore. — E. Algeria, Tunisia, Sicily, Crete (Higgins & Riley, 1970)

and from Lebanon to Oceania (Larsen, 1974) and Australia (Common &

Waterhouse, 1972).

The habitat of this species is described by Higgins & Riley (1970) as “moist

places beside streams”, and Wiltshire (1957) observes that “it is not a desert insect,

and favours roadsides, paths, lawns and gardens, though not exclusive to such”.

Lycaena. There is some disagreement on the subdivision of this genus. In its

broadest sense it comprises about 30 Palaearctic (Kostrowicki, 1969) and 16

DE JONG: West Palaearctic and Ethiopian butterfly faunas 179

Nearctic species (Dos Passos, 1964). It is not exclusively Holarctic: there is one

species in the mountains of Guatemala, three species occur completely inexplica-

bly in New Zealand (Laidlaw, 1970), and there is a slight Ethiopian representation.

Verity (1943) needed six genera to class the Palaearctic species and his

classification was largely, but not entirely, adopted by Higgins & Riley (1970) so far

as the European species are concerned. The Ethiopian representatives belong to

Lycaena s.s., which has about ten Palaearctic species and a number of Nearctic

ones. The Ethiopian species are (Fig. 4):

phlaeas L. — Its range comprises almost all of the Holarctic Region. It is a very

adaptable species; in the Palaearctic phlaeas is found from the oases in the desert-

like plains of Iraq (Wiltshire, 1957) to the shore of the Arctic Sea in Norway

(Nordstròm, 1955). In the Ethiopian Region it occurs above 2000 m in SW. Arabia

(ssp. shima Gabriel) and in the mountains of Ethiopia (ssp. pseudophlaeas Lucas).

abottii Holland. — Confined to the highlands of Kenya and N. Tanzania,-where

it flies in grassy places along streams, etc. The larvae feed on Rumex (Williams,

1969), one of the foodplants of phlaeas. According to Stempffer (1967), abottii

could be a subspecies of phlaeas.

orus Cramer. — Restricted to S. Africa, where it inhabits the south-western and

southern coastal regions as far as Port Elizabeth; it occurs from sea level up to

moderate altitude; the larvae feed on Polygonum (Clark & Dickson, 1971), one of

the foodplants of phlaeas.

clarki Dickson. — Also restricted to S. Africa, where it is mainly an inland insect

(Clark & Dickson, 1971). Up to 1971 it was considered conspecific with orus and

the remarks by Swanepoel (1953) about the habitat of orus (“a highland butterfly

occurring mostly in places where winter is most severe”) may refer to clarki. The

larvae of clarki feed on Rumex (cf. abottii and orus).

With regard to the southern extensions of the otherwise Holarctic genus

Lycaena s.l., Stempffer (1967) remarked: “One can understand how, during the

colder geographical [I suppose he meant geological] periods, the genus Lycaena

managed to reach South Africa by way of the mountains of Abyssinia and East

Africa. It is much more difficult to explain how it reached New Zealand, since in

Asia it extends no further than the Himalayas and Sze Chuan being effectively

replaced by Heliophorus in further Asia”.

Carcharodus. In its broadest sense this genus comprises about six Palaearctic

species; in its most restricted sense it is monotypic, its single species, alceae Esper,

occurring through most of the western part of the Palaearctic Region, east to C.

Asia and N. India. In SW. Arabia it is found as a separate subspecies, wissmanni

Warnecke.

Spialia. The 27 species of this genus can be classified into seven species groups,

which all occur in the Ethiopian Region, while three groups are also represented

outside this area (De Jong, 1974b, and in press). Two groups have members

in the West Palaearctic Region:

sertorius group (Fig. 10). — Comprised of the species mafa Trimen (from S.

Africa through E. Africa north to Ethiopia and SW. Arabia), sertorius Ochs. (W.

Europe and NW. Africa), orbifer H.-S. (E. Europe to E. Asia) and galba Fabr.

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

(India to Ceylon and Burma; Hainan). The Palaearctic members, sertorius and

orbifer, are supposed to be the northern offshoot of an originally Ethiopian stock;

both can be divided into a number of geographic forms which are believed to

reflect the isolation during glacial periods (De Jong, 1974b). The species of this

group can usually be found in hot localities, in open and flowery places; I found

mafa in Kenya up to an altitude of 2000 m. i

phlomidis group. — Mainly distributed in the dry regions bordering the desert

belt that separates the Palaearctic and Ethiopian Regions. Three species are

confined to the region north of the desert belt, viz. phlomidis H.-S., osthelderi

Pfeiffer, and geron Pungeler, while the fourth species of this group, doris Walker, is

found to the north and south of this zone and is, virtually, capable of living in the

desert, as far as its foodplant, Convolvulus lanatus, grows. In the Ethiopian Region :

doris is known from Sudan, Ethiopia and Somalia. It also occurs in SW. Arabia. In

the Palaearctic Region doris is found in Morocco and from Egypt to Pakistan.

Mainly on the basis of its genitalia doris is supposed to be the most recent member

of the group (De Jong, 1974b). Although there appears to be a close relationship

with the Ethiopian spio group, it cannot be stated that the ph/omidis group has an

Ethiopian origin (see also Chapter 3, Summary and conclusions, p. 185). As doris

seems to be the most recent member of the group and its relatives are Palaearctic,

it is concluded that doris invaded SW. Arabia and E. Africa from the north.

Gegenes. A small genus of four species. Two species, niso L. and hottentota Latr.,

are restricted to the Ethiopian Region, where they are widely distributed; one,

nostrodamus Fabr., is distributed disjunctly around the Mediterranean, eastward to

NW. India and southward to Aden, Omdurman and Kassala, and the fourth,

pumilio Hoffm., is found over the combined areas of its congeners, but is lacking

from Tunisia to the Sinai. All species are usually met with in hot and dry localities,

and nostrodamus and pumilio appear to be the most eremic species, which may

count for the fact that they occur in the Palaearctic as well as the Ethiopian

Region and have a patchy distribution around the Mediterranean. Although the

Ethiopian species (hottentota and niso) can live in dry conditions, they are usually

found in more or less grassy places. Indeed, in E. Africa niso is perhaps the

commonest species of Hesperiidae and I found it in Kenya most numerous along

paths in more or less wooded areas. Possibly their preference for grassy places has

forced niso and hottentota to remain restricted to Africa south of the Sahara.

Recently, Larsen (in litt.) found differences between the East and West

Mediterranean pumilio populations and he thinks these populations to be

specifically distinct. This interesting observation has also some bearing on the

reconstruction of the geographic history of the genus. On the (unfirm) basis of the

distribution of related genera (cf. Evans, 1937, 1949) I have concluded that the

genus originally formed part of the old, Tertiary steppe fauna that invaded Africa

from the east. As the extant species seem to fall apart into two groups, viz.

pumilio/nostrodamus and niso/hottentota, there appears to have been an Ethiopian

and a South Palaearctic development of the genus, which may have been initiated

by the desiccation of the area that is now covered by the Saharo-Arabian desert.

In this reconstruction the differentiation of pumilio in the Mediterranean area may

DE JONG: West Palaearctic and Ethiopian butterfly faunas 181

be the result of glacial isolations, and both pumilio and nostrodamus are northern

invaders in Africa.

An alternative hypothesis, adhered to by Larsen, assumes that pumilio is an

Ethiopian species that invaded the Mediterranean along two routes, giving rise to

an East and a West Mediterranean form. If the ancestor of the pumilio/nostrodamus

stock was Ethiopian, also nostrodamus is an Ethiopian element in the Palaearctic

and one could imagine that Gegenes expanded twice to the Mediterranean, the

first time bringing the ancestor of nostrodamus to the north, the second time giving

rise to the two different pumilio forms. This reconstruction sounds very plausible,

but does not take into account that the pumilio/nostrodamus ancestor must have

originated somehow and somewhere. As the Gegenes species can live in steppe-like

habitats, the genus has undoubtedly formed part of the steppe fauna that in

Tertiary times was distributed from Africa to India, whatever the geographic

origin of the Gegenes ancestor. In that case, however, it is much more easily

imaginable that pumilio/nostrodamus originated north than south of the Sahara (see

above). Consequently, I consider provisionally both pumilio and nostrodamus

northern invaders in Africa. I cannot consider the wide distribution of pumilio

south of the Sahara (ssp. gambica Mab.) an objection to this hypothesis, for why

should not a steppe species expand itself so widely over Africa?

Borbo. A palaeotropical genus; 17 species are restricted to the Ethiopian

Region, three are only found in the Indo-Australian area, and one species,

borbonica Boisd., is distributed throughout the Ethiopian Region, in Madagascar

and various islands in the Indian Ocean (Bourbon is type locality), in N. Africa

(Morocco, Algeria, Egypt), the Middle East (Syria, Lebanon, Israel), and is also

found in Spain, in Gibraltar and Tarragona (Gomez Bustillo & Rubio, 1974). The

Mediterranean populations have been distinguished as ssp. zelleri Lederer. B.

borbonica is a rather eclectic species which can live in eremic as well as in more

humid and wooded areas. I found it in W. Kenya along paths in the Kakamega

Forest.

Pelopidas. This genus is mainly Indo-Australian, but among its nine species two

(mathias F. and thrax Hbn.) are found in the Ethiopian as well as in the Oriental

Region and these species are the sole representatives of the genus in Africa. Only

thrax reaches the W. Palaearctic: Turkey (Higgins, 1966), Syria, Lebanon (Larsen,

1974), Jordan (Hemming, 1932), Cyprus, Israel (Evans, 1949); further from Egypt

through Arabia and Iraq to NW. India, Burma and Malaya (Evans, 1949), and

through most of the Ethiopian Region (Evans, 1937). In the W. Palaearctic the

same subspecies (ssp. thrax F.) occurs as in NW. India, the Ethiopian subspecies is

different (ssp. inconspicua Bertolini). It is a known migrant (Larsen, 1974), but that

does not mean that it is not resident in the W. Palaearctic. As B. borbonica, thrax

seems to be a species that has not a very restricted habitat preference.

3. THE AFFINITIES BETWEEN THE WEST PALAEARCTIC AND ETHIOPIAN BUTTERFLY

FAUNAS

From the data given in Chapter 2 it is obvious that the Palaearctic-Ethiopian

affinity cannot be simply described as a north-south connection, but that it is of a

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

rather complex nature. In this chapter we will try to unravel the various types of

affinities. The first division to be made is into direct and indirect affinities.

The term ‘direct affinity” is applied to those cases which concern a penetration

from region A into region B, disregarding a possible secondary radiation in region

B. In the present instance two such direct affinities are possible, viz. a penetration

from the Palaearctic into the Ethiopian Region and vice versa.

By the term “‘indirect affinity” is meant that region A or B, or both, received the

taxon which it has in common with the other region, from a source outside the

other region. In some cases this source can be indicated (allowing a subdivision of

this category), in other cases taxa are placed under this heading by lack of

knowledge. F

Direct affinities

Palaearctic influence in the Ethiopian Region

The species of this group are listed in Table 2. Some features of this group are:

a. all species belong to genera which are not represented in the Oriental Region,

or at most penetrate slightly into that region (Pieris, Colias, Gegenes);

Table 2. Species of (ultimately) Palaearctic origin in the Ethiopian Region. An asterisk indicates that

the species is not specifically distinct from Palaearctic species

SW. Arabia E. Africa S. Africa

Pieris brassicoides x

*Pontia daplidice x x

Fe glauconome x x

distorta x

helice x x

*Euchloe belemia x

* falloui x x

*Colias erate x x

electo x x

Issoria hanningtoni x

smaragdifera x

excelsior x

Melitaea abyssinica x x

Hipparchia tewfiki x

Lasiommata maderakel x

felix x

*Tarucus balkanicus x

*Lycaena phlaeas x x

abottii x

orus x

clarki x

*Carcharodus alceae x

* Spialia dorus x x

*Gegenes ncstrodamus x 1)

È pumilio x x

1) occurs in Sudan

DE JONG: West Palaearctic and Ethiopian butterfly faunas 183

b. most Ethiopian representatives are conspecific with Palaearctic species or are

so Closely related that they can be considered to form a superspecies together with

Palaearctic species;

c. in the Ethiopian Region most species are confined to the north-eastern,

eastern and/or southern part, and no species are confined to W. Africa;

d. none of the species are forest species, though they may fly in close proximity

to the forest; for the rest, their habitat preferences are divergent, ranging from

eremic (e.g., Euchloe falloui) to montane grasslands (e.g., Pieris brassicoides).

Ethiopian influence in the Palaearctic Region

There is only a small number of originally Ethiopian species occurring in the

Palaearctic Region, but as will be shown it is impossible to delimit this group

precisely. The species are listed in Table 3. Their common features are:

a. all species belong to genera which have at least one endemic species in the

Oriental Region;

b. all species fly north as well as south of the Sahara and the northern

populations are usually subspecifically not differentiated from the southern ones

(Spialia sertorius and orbifer are exceptions; they are considered to form a

superspecies with the Ethiopian mafa and the Oriental galba);

c. in the Palaearctic the species are confined to the Mediterranean area (again,

Spialia sertorius and orbifer are exceptions);

d. they are no forest species, but for the rest their habitat preference is

divergent. Generally speaking the species live in Africa in various types of savanna

country. The pierid and lycaenid species are the most eremic ones.

Table 3. Species in the Palaearctic Region originating from the Ethiopian Region. An asterisk indica-

tes that the species is not specifically distinct from its Ethiopian relative, two asterisks indicate that the

species occurs with different subspecies in the Palaearctic and Ethiopian Regions

West Mediterranean East Mediterranean

**Colotis evagore : x

* calais x

* chrysonome x

*Catopsilia florella p.p. !) >)

*Danaus chrysippus alcippus x 2)

**Charaxes jasius x x

*Virachola livia x

* Syntarucus pirithous x x

*Zizeera knysna knysna x 2)

Spialia sertorius x

orbifer x

**Borbo borbonica x x

1) only in Canary Islands

2) see Table 4

Of the listed species we can be reasonably sure that they are Ethiopian intruders

in the Palaearctic Region, as they otherwise occur only in Africa (except Spialia).

Among the species listed in Table 3, four have an Indo-African distribution. Their

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Ethiopian origin has been concluded from their geographic variation, except

Catopsilia florella, which has no subspecies and of which the Canary Islands

population can only have originated from W. Africa. There are, however, more

species with an Indo-African distribution which also occur in the Palaearctic

Region, but without a subspecific indication as to their geographic origin. For that

reason they will be dealt with in the next paragraph.

Indirect affinities

Indo-Ethiopian and eremic species

As it appears impossible to divide sharply between species with an Indo-

Ethiopian and an eremic distribution, these species (so far they occur in the

Palaearctic Region) have been listed together in Table 4. Their common feature is

Table 4. Species in the Palaearctic Region with an Indo-Ethiopian or eremic distribution

W. Medi- E. Medi- Africa S. of migratory

terranean terranean equator

Colotis phisadia

fausta

Anaphaeis aurota

Catopsilia florella i)

Danaus chrysippus chrysippus i)

Hypolimnas misippus

Precis orithya

Ypthima asterope

Apharitis acamas

myrmecophila

Tarucus theophrastus

rosaceus

Lampides boeticus

Chilades galba

Freyeria trochilus

Azanus jesous

Zizeeria knysna karsandra x

Pelopidas thrax thrax

TAUX EX NC ORO

DCE IEC IST X

PLS RS

X MX MX MX MX MX MX XX MX MX MX MX MX MX MX MX MX

1) see also Table 3

2) in the Ethiopian Region only Z. knysna knysna

that it is improbable that they have an Ethiopian or Palaearctic origin. For some

species (Colotis fausta, Pelopidas thrax) an Oriental origin appears obvious. For

other species the region of origin is quite uncertain and it is possible that all

species belong to the fauna that has always lived in the contact area between the

Palaearctic, Oriental and Ethiopian Regions, not a mixed fauna, but a more or less

independent, intermediate fauna (without sharp boundaries). This is most obvious

in the eremic species, as they are confined to the said region. The less confined to

an eremic habitat, the larger is the distribution, and some species can live in such a

variety of habitats, including the eremic, that they could extend their range

throughout the warmer parts of the Old World (e.g., Danaus chrysippus,

De Jong: West Palaearctic and Ethiopian butterfly faunas 185

Hypolimnas misippus, Lampides boeticus). Whatever their place of origin, none of

these species can now be considered a Palaearctic intruder in Africa or an

Ethiopian intruder in Eurasia.

Two other features call for attention, viz. the small number of species of this

category in the W. Mediterranean and the large number of migratory species. The

latter feature may be partly responsible for the slight subspecific differentiation of

the species. The occurrence of two of these species in the W. Mediterranean, viz.

Catopsilia florella and Danaus chrysippus chrysippus, appears to be a direct result of

their migratory habit. Apart from these two species only five species of Table 4

occur in the W. Mediterranean, while all species are known from the E.

Mediterranean (none of the species occurs outside the Mediterranean except the

migratory Lampides boeticus). A possible historical explanation for this difference

is given in Chapter 4.

Remaining genera

Five genera remain to be dealt with. Two of them, viz. Papilio and Neptis, are

very rich in species in tropical environments, while few species occur in

temperate regions. Whatever their geographic history may have been, there are

no indications that they crossed the eremic zone between the Palaearctic and

Ethiopian Regions in more or less recent times as no species occur north as well as

south of the Saharo-Arabian desert zone. It is possible, if not probable, that the W.

Palaearctic members of these genera are originally invaders from the E.

Palaearctic (and neighbouring Oriental) Region.

The distributions of the genera Libythea and Cyclyrius show a relict character and

little or nothing can be said on their history.

The genus Vanessa is the only one in this group that has the same species

(cardui) flying in both the Palaearctic and Ethiopian Regions. Although this very

strongly migratory species seems to cross the Sahara rather regularly to the north,

it is very unlikely that it should be considered an Ethiopian intruder in Europe.

Although its geographic origin is uncertain, it may be suggested that cardui is a

New World species that by its strong migratory behaviour conquered the world.

Summary and conclusions

Summarizing, four kinds of relationship between the West Palaearctic and

Ethiopian Regions can be distinguished:

. by southward traverse across the Saharo-Arabian desert zone,

. by northward traverse across the Saharo-Arabian desert zone,

. through the Oriental Region,

. through the eremic fauna.

Apparently the Sahara has been successfully crossed several times by various

species, from north to south and vice versa. Taking the view that generally

speaking the existence of species common to the regions north and south of the

Sahara indicates a more recent contact than that of common genera without

common species, the traverses seem to cover a long period. The more remarkable

2 © œ 8

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

is the very small number of successful traverses. Successful north-south traverses

number about 17 (cf. Table 2) and most traverses are of such a recent age that the

species concerned scarcely had time for differentiation (although only ten

originally Palaearctic species in the Ethiopian Region are considered conspecific

with Palaearctic species). If we bear in mind that the West Palaearctic comprises

more than 400 butterfly species, it is apparent that the Sahara is an efficient barrier

for the West Palaearctic butterfly fauna in general.

For the Ethiopian butterflies the Sahara appears to act as a still stronger barrier.

Successful northward traverses number only twelve (cf. Table 3). As the Ethiopian

butterfly fauna comprises almost 2700 species, it is evident that a successful

crossing is a very rare event (but in Chapter 4 it will be shown that unsuccessful

crossings may have been rather frequent). In all cases the same species occurs

north as well as south of the desert zone and the species are confined in the

Palaearctic to the Mediterranean (with the exception of Spialia sertorius/orbifer).

This suggests that the average age of the northward traverses is still younger than

that of the southward ones.

Thus there appear to exist two remarkable differences between the southward

and northward traverses, viz.:

a. in relation to the total number of butterfly species in the West Palaearctic and

Ethiopian Regions the southward traverses are much more frequent than the

northward ones;

b. judged from the differentiation of the species after the traverse the southward

traverses cover a much longer period than the northward ones.

A possible explanation for these differences is given in the next chapter.

The Indo-Ethiopian genera pose a separate problem. Not all of these genera

have representatives in the W. Palaearctic. The complex nature of the Indo-

Ethiopian relationship falls outside the scope of this paper, but in the next chapter

a few words have to be said on this subject in order to get a better insight in the

Euro-Ethiopian affinities.

4. HISTORICAL ASPECTS

In this chapter we will try to relate the data amassed in the foregoing chapters to

ecological changes in the northern part of Africa in the past. As the Euro-

Ethiopian affinities in the butterflies are mainly at the species level, we are

primarily interested in the ecological changes that took place during the

Pleistocene. Affinities at the genus level may point to an older, Tertiary

connection. Very little is known of this period and various authors do not always

agree. Nevertheless, the little that is known may help to explain the distribution

patterns found.

Ecological history of the northern half of Africa and the Arabian peninsula

Tertiary

For our purpose it does not seem relevant to consider the period prior to the

Miocene as it is hardly believable that a contact broken since then is at present

still discernible at the genus level.

DE JONG: West Palaearctic and Ethiopian butterfly faunas 187

From Middle Miocene to Late Pliocene there was a connection with the

Oriental Region via Arabia. The Red Sea in its present form came into existence

only at the end of the Pliocene or the beginning of the Pleistocene; prior to that

time it was a large inland sea (Ekman, 1953). As the Persian Gulf also appears to

be rather recent and the climate in this area in Miocene and Pliocene was often

more humid than at present (e.g., at the end of the Miocene and various times in

the Pliocene Egypt has experienced periods of considerable rainfall, see Moreau,

1952), a close contact between and intermingling of the Oriental and Ethiopian

faunas was possible. This intermingling was largely unidirectional and consisted

mainly in a large invasion of Oriental faunas (De Lattin, 1967). In the Pliocene the

correspondence has been still larger than at present, as many groups became

extinct in the Oriental Region (shown by the famous fossil Siwalik fauna of N.

India).

I wonder if it is correct to speak of Oriental species invading Africa, as is usually

done. It concerns mainly steppe species which lived in a region that is now partly

Oriental, partly Palaearctic. The uplift of the Himalayas (of which the Siwalik

Range forms the southern border), the main boundary between the Palaearctic

and Oriental Regions, “‘is believed (...) to have taken place in four major impulses,

respectively during the Eocene, the Middle Miocene, the Upper Pliocene and in

the Late Pleistocene” (Mani, 1968). This means that it is not relevant to speak of a

Palaearctic and an Oriental Region as biogeographical regions in the Tertiary.

Thus, it is misleading to speak of an Oriental invasion into Africa, and it seems

more suitable to term it an Asiatic invasion.

More temperate Asiatic species, especially those which could not live in a steppe

or savanna environment, had at this period no chance of reaching Africa as the

mountains by way of which they could cross the steppe region were still in the

process of uplifting and there were not yet mountains in East Africa high enough

to support temperate species.

The progress of the Alpine orogeny in Late Tertiary and especially the gradual

rise of the Taurus-Armenia-Zagros mountain system, the falling temperature and

the opening of the southern end of the Red Sea (not before the middle of the

Pliocene; Ekman, 1953), must have hampered the free exchange of steppe species

between Asia and Africa. Nevertheless, during humid periods the passage, though

difficult, must have been possible.

There is no full agreement on the Tertiary connections between Africa and

Europe in the W. Mediterranean. According to Moreau (1966) there was a

landbridge in the Tunisia-Sicily area in the Pliocene. Besides such a landbridge

Verity (1940) mentioned a direct connection between Sardegna and Africa in the

early Pliocene, but it is not clear how far such a connection is supported by

geological evidence.

Although Moreau (1966) stated that there is no evidence of a connection across

the Straits of Gibraltar, Ekman (1953) postulated a Middle Tertiary landbridge

between Cape Spartel and Cape Trafalgar, although in the Pliocene the

Mediterranean became connected to the Atlantic, subsequently through the valley

of the Guadalquivir, the valley of Fez and at last across the Straits of Gibraltar. A

similar sequence of events was given by Verity (1940) and Kostrowicki (1969). It is

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

clear that such a connection in this area cannot have been of much importance as

a direct passage, as only those species occurring in the northernmost tip of NW.

Africa when it was conjoined to Spain, could freely move into Africa when this

part became connected to the rest of NW. Africa. But apart from such a

connection, the Straits of Gibraltar are only a narrow gap (at present 13 km), and

anyone who from Tarifa, the southernmost point of Spain, has looked at the

mountains of Tanger will be convinced that in due course many species must have

been able to cross the gap.

In the meantime the Atlas Mountains were uplifted, rendering temperate

species a possible habitat far to the south. However, during the Tertiary the

lowland was probably still too warm to give temperate species the opportunity to

expand to the south. Moreover, the Sahara south of the Atlas Mountains would

have been a strong barrier for temperate species, although it may have been

covered over large areas by mediterranean shrub and savanna-like vegetation,

possibly continuously so along the Atlantic coast to the W. African savannas. But

warmth-loving species of open formations may have been able to cross the Sahara

by this way (to the north as well as to the south), thus forming a connection

between the European and Ethiopian faunas.

In sum, interchange of species between the W. Palaearctic and Ethiopian

Regions during the Miocene and Pliocene was possible through Arabia and the

Middle East and, probably to a lesser extent, across the W. Mediterranean and

through NW. Africa. Only species which could live in a steppe or savanna-like

environment could make use of these connections.

The above description suggests that we know a lot of the Late Tertiary ecology

in the area concerned, but considering that this period covered about 25 millions of

years during which large mountain massifs were uplifted and ecological conditions

must have changed at a large scale, it is clear that we know extremely little.

Pleistocene

About the relatively short Pleistocene period (about 2 millions of years;

considered here to extend up to the present day, after Moreau, 1966, and

Hamilton, 1974) we are rather better informed, though most data refer to the last

30,000 years.

The glaciations of the Palaearctic brought many temperate species far to the

south where they could survive warmer interglacial periods in the S. Palaearctic

mountains. Thus, an ecologically wider array of species stood, as it were, waiting to

cross the Sahara as soon as conditions became favourable.

At present the Sahara is too dry to be crossed by Palaearctic as well as

Ethiopian species. During a more humid period an interchange of species living in

steppe and savanna and perhaps mediterranean shrub would be possible, but for

species not capable of living in such an environment a lower temperature is

required. Recent investigations, particularly pollen analysis, have revealed that

there have, indeed, been colder and more humid periods. The history of the

vegetation of East Africa was summarized by Van Zinderen Bakker (1971) and

Hamilton (1974). It appears that from 30,000 to 25,000 BP (Kalambo Interstadial,

DE JONG: West Palaearctic and Ethiopian butterfly faunas 189

corresponding with the European Paudorf Interstadial) the temperature was about

2° C lower than at present. The temperature decreased between 25,000 and

12,500 BP to 6° C lower than now. This period is known as the Kenya Glacial and

coincides with the last maximum of the Wurm Glaciation in the Palaearctic. After

a rapid increase the temperature c. 10,500 BP was similar to today. A further

synchronization in the temperature fluctuation between Europe and East Africa is

the occurrence of a short cold phase just before the Postglacial Climatic Optimum,

about 4500—7500 BP (Neolithic), when temperatures were about 2° C higher than

today.

This synchronization suggests-that similar fluctuations took place in the

intervening area. This is very important. The lower limit of the montane zone in

East Africa, at present at 1500 m, is supposed to have been depressed at least by

1000 m during the maximum of the Kenya Glacial and Moreau (1966) calculated

that even only 12,000 BP, with a temperature 2.5° C lower than today, the lower

montane limit would have been at 1000 m. If we suppose that a similar descent of

the montane zone took place in the area between East Africa and the Palaearctic,

the enormous impact on the distribution of montane habitats is easily seen from

the surface relief shown in Fig. 1. In fact an almost continuous montane block was

le] LI d DI y è ng ER Di = Tr

TTI] 500 -1000 m o F TH ; =

BEER 1000-2000 m N b È “ella si

EEN >2000m

Fig. 1. Surface relief in northern Africa

formed from Asia Minor to East Africa, and from NW. Africa diagonally across

the Sahara to Central Africa. Indeed, pollen analysis has shown that the Ahaggar

Massif during much of the later part of the last glaciation was clothed with a rich

mediterranean vegetation and even the European lime (Tilia), now absent in N.

Africa, occurred there (Moreau, 1966).

There is, however, one factor which must have greatly influenced the

availability of montane habitats for Palaearctic species trying to extend their range

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

southward, viz. humidity. Hamilton (1974) observed, that at least in the East

African mountains the last glacial was dry, contrary to the common opinion that it

was a pluvial period. Moreover, according to Moreau (1966) Arabia has probably

never been better than semiarid at any time during the Pleistocene, except the

mountain areas of Yemen and Gebel Akhdar and the somewhat elevated southern

rim of Arabia, where wooded steppe and savanna may have existed, possibly with

patches of dry evergreen forest, during glaciations and the humid Neolithic. _

Although the duration, extension and chronology of dry and humid periods are

still largely unknown, it is clear that drought must have hampered the southward

extension of Palaearctic species, if it occurred during the only periods the

temperature was low enough for such an extension, i.e. during glacial periods.

For Ethiopian species trying to extend their range northward, drought as well as

low temperatures would have been unfavourable. How far the warm interglacial

periods have been favourable for northward extensions is unknown at present

(apart from a humid period in the latter Middle Pleistocene; Moreau, 1963), but at

least postglacially there has been a favourable period, viz. the humid period during

the Neolithic, in Europe known as the Postglacial Climatic Optimum or the

Atlantic (cf. Moreau, 1955). This humid period, with a temperature about 2° C

higher than now, attained its maximum about 5,000 years BP. At that time the

western Sahara (and also the mountains of Air and Tibesti) was covered by

mediterranean vegetation (scrub and dry woodland) up to the southern limit of the

present Sahara, where it was in contact with the Ethiopian savanna vegetation

(Moreau, 1966). Braestrup (1947) pictured the rivers at that time running across

the western Sahara. It is clear that especially the coastal belt along the Atlantic

requires only a small degree of climatic improvement in order to become viable as

a passage for northern and southern species. The area north-east of the line

Tripoli-Khartoum, however, appears to have always been drier and during the

Neolithic only the Nile Valley could act as a corridor in this area (Moreau, 1963).

However, the Neolithic period is believed to have been humid in the Sudan and

Ethiopia and Moreau (1966) mentioned for Arabia, at least for its southern part,

a somewhat richer vegetation, especially in mountainous areas, during this period.

For temperate Palaearctic species trying to go southward the Postglacial

Climatic Optimum must have been a bad period. As a consequence of the higher

temperature (about 2° C) the lower limit of the montane zone was about 400 m

higher. Mountains below 1900 m lost their montane habitat and their function as

stepping stones, montane habitats became more isolated and the land in between

too hot.

By a sudden deterioration late in the Neolithic, caused by unknown factors, the

entire mediterranean vegetation in the western part of the Sahara became extinct

and from that time on the present bare condition of the western Sahara was

developed. A similar condition has existed earlier in the Pleistocene, as at some

stage in the late Pleistocene, prior to 22,000 BP the dunes of the Sahara extended

even 300 miles south of the present limit of moving sands in West Africa. It is

obvious that the present and similar conditions do not facilitate the passage in

both directions.

To summarize the Pleistocene vicissitudes, it can be said that during glaciations,

De Jong: West Palaearctic and Ethiopian butterfly faunas 191

conditions were favourable for a southward traverse, as far as drought did not

prevent it. For the last time such conditions existed from about 25,000 to 18,000

BP. Northward traverses were mostly feasible during humid periods, the last of

which occurred only about 5000 BP.

The distribution of mountain ranges suggests that the route from the Middle

East to the Ethiopian highlands and further south was much easier for temperate

species than from the NW. African mountains southward. It is not clear whether a

western or an eastern traverse was easier for Mediterranean and Ethiopian

species; possibly both traverses were equally easy (or difficult), though not

necessarily at the same time.

The direct West Palaearctic - Ethiopian affinities

The southward traverse

The Palaearctic species in the Ethiopian Region are largely absent from West

Africa, suggesting that these species used mainly the eastern traverse. It should,

however, be borne in mind, that East Africa offers the best opportunities for more

or less temperate species to survive in its large mountain areas, and the possibility

that also the western traverse was used but the species became extinct there,

cannot be ruled out in advance. Indeed, there are indications that some species

crossed the Sahara in its western part, but if they ever reached the Ethiopian

Region, they died out entirely there. The southward movement in the western

traverse is shown by the species at present occurring in the mountain areas in the

Central Sahara, viz. Ahaggar (3000 m), Air (2000 m), and Tibesti (almost 3500 m).

Table 5 gives an impression of the butterfly composition of these mountains.

Although the data of these areas are incomplete, it is remarkable that in Tibesti, at

the greatest distance from the Atlas Mountains, still six species of the families

Pieridae and Nymphalidae occur that can be considered northern immigrants.

Bernardi (1962) also listed Vanessa cardui as a Palaearctic species in Tibesti, but to

me the geographic history of cardui seems obscure; moreover, the lack of

subspecific variation makes it impossible to decide on the origin of the Tibesti

population.

Four out of the six Palaearctic species in Tibesti have also been found in the

Sahara between the Atlas Mountains and In-Salah (= Ain-Salah), more than

halfway Ahaggar, mainly in river beds and oases (Rothschild, 1913). These species

have also been found in the northern part of Rio de Oro (Bernardi, 1966). One of

them, Pontia glauconome, is actually an eremic species. It is obvious that a more

humid period can easily bring these species across the Sahara and their isolation in

the mountains in the Central Sahara (if they are isolated there at all) is not older

than about 5000 years (Postglacial Climatic Optimum). This may also count for the

fact that they are subspecifically indistinct from specimens from Algeria.

Of the two remaining Palaearctic species in Tibesti, Vanessa atalanta is a well-

known migrant, as is the cosmopolitan Vanessa cardui. It is not impossible that at

least the latter species crosses the Sahara regularly in present times.

Melitaea didyma appears to be the only species which requires a colder period for

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Table 5. Composition of butterfly species in various regions of the Sahara. Data extracted from Roth-

schild (1913, 1915, 1921), Riley (1934), Bernardi (1962) and Bernardi & Stempffer (1951)

NW. Sahara Ahaggar Air Tibesti Palaearctic

Papilio machaon x

Pieris rapae

Pontia daplidice

glauconome

Euchloe ausonia

falloui

belemia

Elphinstonia charlonia x x x

Colotis calais

phisadia

chrysonome

halimede

eupompe

liagore

evagore x

aurora

evippe

eris

Pinacopteryx eriphia

Calopieris eulimene

Anaphaeis aurota

Nepheronia buqueti x

Colias croceus x

Catopsilia florella

Eurema brigitta

Vanessa atalanta

cardui x x x

Precis oenone

Byblia ilithyia x

Melitaea didyma x x

deserticola x x

Danaus chrysippus

Ypthima asterope

Virachola livia x

Tolaus nursei

Apharitis acamas x

Anthea crashayi x

Cacyreus lingeus x

Castalius cretosus x

Tarucus theoprastus x x

rosaceus

Lampides boeticus

Chilades eleusis

Azanus ubaldus

Zizeeria knysna

Aricia cramera

Lycaena phlaeas

Gegenes nostrodamus

x X MX X

x xX X MX

NN KOK OK OG a

EX eX DI DER DC AXE GK MEN x

EX EX OS

PDC KE EX

x x MX X

DE JONG: West Palaearctic and Ethiopian butterfly faunas 193

extending its range so far to the south (the mention of this species from the Sahara

between the Atlas and Ahaggar by Rothschild, 1913, concerns the related species

M. deserticola, see Higgins, 1941). It is however surprising that this geographically

confusingly variable species in Tibesti occurs in a form which is subspecifically

indistinct from the Algerian form, thus suggesting 4 mucn more recent contact

than the seemingly required Last Glacial.

From Ahaggar three Palaearctic species are known that have not been found in

Tibesti, viz. Papilio machaon, Euchloe ausonia and Euchloe falloui. Subspecifically

they are scarcely or not at all distinct from the Algerian representatives of the

respective species, suggesting that they reached the Ahaggar in relatively recent

times. As in the case of Melitaea didyma in Tibesti we are forced to suppose a more

recent range extension than during the Last Glacial. As P. machaon and E. falloui

have actually been found in the NW. Sahara (Rothschild, 1915), they may even

now reach the Ahaggar at times. The Postglacial Climatic Optimum must have

been favourable for their range extension, though the temperature was higher than

at present, as they could follow a route mainly over higher land (500—1000 m)

(Fig. 1). In the same way Euchloe ausonia and Melitaea didyma may have reached

the Central Saharan mountains, though for them the passage was more difficult

and their occurrence so far to the south appears the result of a chance

colonization.

There are no traces of a penetration from NW. Africa still further across the

Sahara. About 400 km south-east of Tibesti extends the Ennedi mountain range. No

Palaearctic species are known from this area. except the eremic Pontia glauconome

(Bernardi, 1964).

Although the Air mountains are much closer to Ahaggar than Tibesti, they have

only a single Palaearctic species, viz. Pontia daplidice. This paucity may be

attributed to their much smaller extent and less elevation. It is noticeable that

there are no traces left of butterflies colonizing the Central Saharan mountains

during the Last Glacial. If they ever did, they died out entirely.

While in the Saharan mountains the affinities with the Palaearctic Region are at

the species and even subspecies level, the picture in East Africa is quite different.

Of the 25 species in the Ethiopian Region thought to be ultimately of Palaearctic

origin (Table 2), less than half (11) are considered specifically indistinct from

Palaearctic species and only four or five of them are regarded subspecifically

identical. This suggests that the contact with East Africa is at least partly of a

much older date, and the various grades of relationship between the Ethiopian and

Palaearctic members (subspecific, specific, superspecific, generic) indicate a

repeatedly broken contact.

The generally older age of the Palaearctic-E. African contact is due to the fact

that only few of the species concerned could use this eastern traverse across the

eremic regions during the humid Neolithic, viz. Pontia daplidice and glauconome,

Euchloe belemia and falloui, Colias erate and Tarucus balkanicus. Except the last

one, these species have also been supposed to have extended their range

southward during the same period through the western traverse. Their slight

differentiation in the Ethiopian Region is apparently due to the rather recent date

of their isolation.

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

Fig.2

Lasiommata maera g

L. maera

@ Lu felix

A L. maderakal

Fig. 3

Superspecies Pieris brassicae

Z 77 P. brassicae

@ P.brassicoides

Fig.4

Lycaena phlaeas groyp

WL L. phlaeas

HL. abottii

L. clarki

| Fig.5 6

MI L.orus

|cFoota helice

Fig. 2—5. Butterfly species with Palaearctic ancestry in the Ethiopian Region

De JonG: West Palaearctic and Ethiopian butterfly faunas 195

As the other species need a lower temperature, their isolation must date back at

least as far as the last phase of the Last Glacial, i.e. 18,000—25,000 years ago. The

differentiation of the Lycaena phlaeas group can only be attributed to at least two

subsequent southward movements, the first of which (giving rise to L. orus/clarki)

may also have brought Pontia helice to S. Africa. It is impossible to date these

movements more or less exactly, but there is no reason to consider them otherwise

than coinciding with various phases of the Last Glacial or with an earlier Glacial.

The systematic isolation of the /ssoria hanningtoni group may also be due to a long

geographic isolation, from before the Last Glacial.

The history of Pontia distorta is still obscure. It is confined to the dry savanna of

Somalia and Kenya and may have come there during a dry period across the

Ethiopian mountains, but before further suggestions are made, it is advisable to

clear up the relationship of distorta to other Pontia species.

A glance at Fig. 1 shows that in the eastern traverse two routes were available

for the temperate species (i.e. the species which needed a cold period for their

southward movement), viz. east and west of the Red Sea. The eastern route with

its large mountainous areas seems to offer the best opportunities, though using this

route the species had to jump the southern end of the Red Sea and a lowland area.

From the species known to occur in the SW. Arabian mountains it is clear that

many species used this route indeed and it seems possible that most Palaearctic

species which reached Ethiopia, did so by way of this route. For species that

needed a humid rather than a cold period for range extension the jump over the

southern end of the Red Sea must have been still easier.

In table 2 are indicated the species of Palaearctic origin occurring in SW.

Arabia. The data are extracted from Gabriel (1954), who mentioned 18 Palaearctic

species in SW. Arabia, but he did not mean to state a geographic origin, only to

indicate that the species concerned also occurred in the Palaearctic Region. Two

of the species listed in table 2 belong to genera which are unrepresented in Africa

south of the Sahara (Eumenis, Carcharodus). Apparently, it was too difficult for

these species to make the jump to the Ethiopian highlands (or they became extinct

there). The other species are represented by closely related species (Pararge felix),

or subspecies (Melitaea abyssinica, Lycaena phlaeas) or by identical subspecies

(remaining species) in Ethiopia. These data strongly suggest, that the gap between

the SW. Arabian and Ethiopian mountains was not insurmountable for many

Palaearctic species (see also the next paragraph and Chapter 5, the paragraph on

the Red Sea). Of the Palaearctic species in East Africa only Pieris brassicoides and

the Issoria hanningtoni group have no relatives in SW. Arabia, so that it is quite

impossible to say which route they used to reach Ethiopia.

Summary and conclusions. Two traverses have been available for a southward

movement across the Saharo-Arabian desert zone, viz. a western traverse, from

the Atlas Mountains across the W. Sahara and the mountains of the C. Sahara, and

an eastern traverse, at either side of the Red Sea. During the cold Glacial Periods

and phases, the last of which reached its maximum only 18,000—25,000 years ago,

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

temperate species could extend their ranges southward through the mountains of

the western and eastern traverses, but only traces of a repeated use of the eastern

traverse have been left. The western traverse was a dead-end road as there was no

link with an Ethiopian mountain range that could support temperate species

during warmer times. Even in the C. Saharan mountains no traces are left of

glacial range extensions by butterflies from the north.

The very low number of glacial invaders in East Africa and SW. Arabia (about

eight) can have various causes:

a. there has been no continuous passage for temperate species, but the passage

served as a filter or even the SW. Arabian and E. African mountains were reached

by chance colonization;

b. many species which actually reached SW. Arabia/E. Africa, died out during _

hotter and possibly drier times;

c. many species were in principle well able to make the traverse, but their

foodplants failed to do so or at least to establish themselves permanently in SW.

Arabia/E. Africa;

d. many species could not compete with Ethiopian species already living in the

more temperate habitats.

It seems to me that the main cause for the low number of glacial invaders is a

combination of the first two (maybe three) mentioned. This combination was

apparently still more effective in the C. Saharan mountains, where glacial invaders

appear to be absent at present.

It would be interesting to know the foodplants of all Palaearctic species in the

Ethiopian Region. For the Lycaena phlaeas group Rumex species have been

recorded as foodplants and the /ssoria hanningtoni group is said to live on Viola

species. As both plant genera are presumably Palaearctic intruders it is unlikely

that the butterflies living on it experienced much competition on the part of the

Ethiopian species. This may explain their relative success in the Ethiopian Region.

Butterflies capable of living under higher temperatures could move southward

during humid periods, the last of which occurred only about 5000 years ago. This

relatively recent contact is reflected by the fact that they are slightly or not at all

differentiated in their present C. Saharan and E. African habitats. They were not

especially dependent on the mountainous traverses of the more temperate species,

but as according to Moreau (1966) the eastern Sahara has always been drier than

the western part, we may also speak here of a western and an eastern traverse,

keeping in mind that the western traverse did not only comprise the mountainous

regions, while the eastern traverse may also have included the Nile Valley.

Also for these species the western traverse was a dead-end road. Although the

southward penetration has left various traces in the C. Saharan mountains, all

species which ever reached the Ethiopian Region by this way have become extinct

there. It is even questionable whether these species were ever able to establish

themselves well in the probably rather saturated W. African savanna fauna. This

can, however, not be the only reason for their absence in W. Africa as some of

these species could establish themselves firmly in E. Africa. Apparently the

mountains of E. Africa offer better opportunities for survival of these species too,

than the possibly too hot, low-lying steppe and savanna along the southern edge of

DE JONG: West Palaearctic and Ethiopian butterfly faunas 197

the Sahara in W. Africa. Also the sudden disappearance of the Mediterranean

vegetation from W. Africa shortly after the Postglacial Climatic Optimum (see

Moreau, 1966) may have played a part in the absence of Mediterranean butterflies

in W. Africa or at least there may be a common cause.

Also the Palaearctic species capable of living in steppe and savanna environ-

ments number very low in E. Africa and SW. Arabia (about six) and in this case

again, the cause for the low number may have to be looked for in a combination of

desiccation and disappearance of foodplants, assuming that there have ever been

more species penetrating so far to the south.

In sum, a southward movement of Palaearctic butterflies across the Saharo-

Arabian desert zone has occurred various times during the Pleistocene, but

penetration into the Ethiopian Region has only been successful (i.e. with still living

progeny) through the eastern traverse, along either side of the Red Sea. The

effectiveness of the Red Sea itself, especially at its southern end, as a barrier will

be dealt with in Chapter 5.

The northward traverse

For Ethiopian species the Glacial Periods have probably been too cold for a

successful northward expansion and a movement in that direction must generally

have been confined to periods which were more humid and possibly also warmer

than the present one. The last of such periods, the Postglacial Climatic Optimum,

occurred only about 5000 years ago and it is to be expected that species, taking

this opportunity, have differentiated in the Palaearctic Region very little or not at

all. Indeed, eight of the thirteen species of table 3 are not subspecifically distinct in

the Palaearctic Region from their Ethiopian representatives and thus, for these

species a northward expansion during the Neolithic comes into consideration.

Migratory species (Catopsilia florella, Danaus chrysippus) may have reached the

Palaearctic still more recently; C. florella is known from the Canary Islands only

since about ten years.

If the Palaearctic representative is subspecifically distinct from the Ethiopian

stock, this does not mean a priori that the isolation is of an older age. It is quite

well possible that Colotis evagore developed its Palaearctic race, nouna, during the

last 5000 years. However, for the two other species concerned, viz. Charaxes jasius

and Borbo borbonica, the situation is somewhat different as they occur dis-

continuously in the same subspecific form in the West as well as in the East

Mediterranean. The absence of differentiation in this area suggests that the

distribution has been continuous up to a rather recent time or that there has been

a rather recent range expansion within the Mediterranean area. If an Ethiopian

species succeeded in expanding its range northward throughout most of the

Mediterranean, it is improbable that during a subsequent isolation the Mediter-

ranean range would remain continuous, and thus the development of a single

Mediterranean subspecies is improbable. As a consequence, Charaxes jasius and

Borbo borbonica probably became isolated (and differentiated) in the Mediter-

ranean before the Neolithic and extended their range in this area during this

humid period.

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 6, 1976

The two Palaearctic Spialia species, sertorius and orbifer, are specifically distinct

from their Ethiopian relatives. They are believed to have originated from a

common Ethiopian stock, of which the Ethiopian mafa and the Oriental galba

constitute the remainder of the progeny. As the geographic variation of sertorius

and orbifer reflects the influence of at least the Last Glacial and the origin of

sertorius and orbifer themselves is the result of glacial isolations, the northward

expansion of the Ethiopian stock must have occurred earlier. S. sertorius and

orbifer are not restricted to the Mediterranean Region, but can also live under

much cooler conditions, while mafa ascends at least to 2000 m in the East African

mountains. Therefore, it is possible that the northward expansion occurred during

one of the earlier Glacial Periods, as a result of the lowering of the montane zone.

As for the southward movement, two routes were available for the northward

movement, a western and an eastern one, but the Ethiopian species did not need

mountains on their way, only hot and not too dry country. Thus one would expect

that during the Postglacial Climatic Optimum many Ethiopian species expanded

their ranges northward over a broad front, with the exception of the eastern

Sahara, where conditions seem to have always been unfavourable. Nevertheless, at

present very few Ethiopian species live in the Palaearctic Region. There is another

remarkable feature: almost all Ethiopian penetrations into the Palaearctic Region

appear recent, mostly not older than about 5000 years. Only the Spialia species

indicate a penetration in the early Pleistocene.

The mainly recent arrival of the present Ethiopian species in the Palaearctic

Region may be due to the fact that former penetrations (before the Last Glacial)

were largely obliterated by the dramatic climatic change during the Last Glacial.

The relatively low number of Ethiopian species in the Palaearctic Region at

present indicates that postglacially the passage to the north was difficult or that

only few species managed to survive till the present day.

An indication of a northward movement over a broad front is found in the

present butterfly fauna of the western Sahara and the Saharan mountains. The

butterflies must have reached the mountains during a more humid period and they

now live there largely isolated, being surrounded by large stretches of uninhabi-

table land. As they are scarcely, if at all, differentiated from tropical species, the

isolation must be recent and as for the Ethiopian species in the Palaearctic

Region, the Postglacial Climatic Optimum comes into the picture as a favourable

period for northward range extension.

In view of the isolation and rather severe conditions the number of tropical

butterflies in the Saharan mountains is not unimportant. Table 6 and Fig. 6 give

the numbers and percentages of tropical (i.e. originating from the area south of the

Sahara) and non-tropical species in various regions in the Sahara and NW. Africa.

As is to be expected, the mountains closest to the southern edge of the Sahara

(Air, Ennedi) contain the largest percentages of tropical species. Remarkable,

however, is the steep decline in the percentage of tropical species in Ahaggar and

further to the north, the more so as in northern Rio de Oro, further north than

Ahaggar and close to the southern fringe of the Palaearctic Region, the situation is

quite different and the ratio tropical-nontropical species is about the same as in

Tibesti. As there were ample opportunities for northward range extension only

DE JONG: West Palaearctic and Ethiopian butterfly faunas 199

Table 6. Number of tropical and non-tropical species in various regions in the Sahara and NW. Africa.

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4 3 E. A. LOELIGER. — The enigma of Celerio hybr. Pauli Mory, p. 217-219, pl. 1.

«| Tijdschrift voor Entomologie, deel 119, afl. 7 Gepubliceerd 25-X-1976

THE ENIGMA OF CELERIO HYBR. PAULI MORY

E.A. LOELIGER

Hofdyck 48, Oegstgeest, Netherlands

With one colour-plate

INTRODUCTION

In the summer of 1897, the well-known Swiss entomologist M. Paul observed on

Hippophae rhamnoides in the vicinity of Sion, Wallis, Switzerland, a sphinx

caterpillar resembling Celerio euphorbiae. Unfortunately, the detailed description

of the caterpillar has been lost. Breeding of the caterpillar was successful,

however, and on September Ist of the same year the moth emerged. It was put at

the disposal of Prof. Standfuss for his collection (Fig. 1; at present kept in the

Entomology Institute of the Eidgenòssische Technische Hochschule at Zurich,

Switzerland). Shortly thereafter, the medical student Mory published a photo-

graph together with a detailed description of the moth. The author, on the basis of

a comparative study, thought that it could be the product of a natural cross-

breeding of Celerio euphorbiae 3 with Celerio hippophaes 9, since called Celerio

hybr. Pauli Mory (1901). Several years later, Denso (1911) denied the probability

suggested by Mory for two reasons. First, caterpillars from experimentally

obtained crossbreeding of Celerio euphorbiae 4 with Celerio hippophaes 9,

although clearly resembling those of Celerio euphorbiae could only be bred ab ovo

on Euphorbia cyparissias or other Euphorbia species. Second, the resulting moths

differed in various ways from Paul’s specimen. Since it remained possible that the

latter was a bastard resulting from a cross between Celerio livornica and Celerio

hippophaes, Denso gave the experimentally obtained bastard of Celerio euphorbiae

& with Celerio hippophaes 9 the name Celerio hybr. euphaes (see Denso, 1911).

Paul’s finding remained enigmatic. With the present report we offer a possible

solution of the problem of a Celerio euphorbiae-like caterpillar to be found on

Hippophae rhamnoides, a solution which might serve as a model holding equally

well for bastards of Celerio livornica crosses with Celerio hippophaes.

EXPERIMENT

The materials used for the experiment which gave the desired result were the

genuine species Celerio euphorbiae and Celerio hippophaes, found in the summer of

1973 in the Wallis region of Switzerland. Brought to Holland, the pupae hatched

as early as the latter part of 1973, and cross-breeding of Celerio hippophaes 3 with

Celerio euphorbiae Q (Celerio hybr. hippophorbiae) under strictly aseptic conditions

217

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 7, 1976

was successful on Euphorbia cyparissias and/or polychroma. In 1974, the following

cross-breeding with this hybrid succeeded:

1. Celerio hybr. hippophorbiae 3 x Celerio hippophaes 9, and

2. Celerio hybr. hippophorbiae 3 x Celerio euphorbiae 9.

Caterpillars of variety 1 were highly variable and accepted only Hippophae

rhamnoides; some of them strongly resembled the caterpillars of Celerio hippophaes,

but none those of Celerio euphorbiae. Variety 2, on the other hand, was very similar

to Celerio euphorbiae, and accepted only leaves of Euphorbia cyparissias and/or

polychroma. Most of the pupae of both varieties hatched in September of 1974.

Probably supported by excellent climatological conditions, cross-breeding of

Celerio hybr. (hippophorbiae x euphorbiae) 3 with Celerio hybr. (hippophor-

biae x hippophaes) 9 was again successful three times. The three female moths’

together yielded about 150 fertile eggs. Approximately 25 per cent of the

caterpillars accepted Hippophae rhamnoides leaves and developed extremely well;

the rest did equally well on Euphorbia polychroma. Most of these caterpillars

resembled another successfully obtained bastard, the result of cross-breeding of

Celerio hybr. (hippophorbiae x hippophaes) 3 with (hippophorbiae x hippophaes) 9.

However, a far from neglegible number of not only the caterpillars raised on

Euphorbia polychroma but also of those kept on Hippophae rhamnoides, were almost

indistinguishable from Celerio hybr. hippophorbiae, i.e. resembled Celerio euphor-

biae rather closely so that observers less familiar with the detailed pattern of this

Celerio species would probably have concluded that these were Celerio euphorbiae

caterpillars of the red (Fig. 3), or the yellow, or even of the black variety (Fig. 4).

DISCUSSION

Cross-breeding is rather common in nature (Mory, 1897). Moreover, the two

species in question, Celerio euphorbiae and Celerio hippophaes, live near each other

in the foreland of the Rhone River in the Wallis region. In captivity, Celerio hybr.

euphaes Denso is one of the most easily obtained bastards. Ab ovo caterpillars of

this crossing, however, do not accept Hippophae rhamnoides. Hence, in nature this

bastard is doomed to death. However, the reciprocal bastard, Celerio hippophorbiae

which is obtained just as easily in captivity, grows very well on the fodder plant of

the maternal family. Caterpillars of both of these primary bastards produce highly

fertile moths. Secondary bastards are therefore easily obtained: whereas the

variety Celerio hybr. hippophorbiae 3 x Celerio hippophaes 9 lays the eggs on and

accepts Hippophae rhamnoides, the reciprocal bastard, Celerio hybr. hippophorbiae

3 x Celerio euphorbiae 9, does the same with the various Euphorbia species

growing in the Wallis region. Of these two, the rather sophisticated recombination

of Celerio hybr. (hippophorbiae x euphorbiae) & with (hippophorbiae x hippophaes)

Q in captivity has been shown to lay eggs on Hippophae rhamnoides, which is

accepted by approximately one-fourth of the emerging caterpillars, even those

strongly resembling Celerio euphorbiae. This tertiary bastard now might have been

responsible for Paul’s finding of an euphorbiae-like caterpillar on Hippophae

rhamnoides. The fact that ab ovo caterpillars of the two primary bastards Celerio

hybr. euphaes and Celerio hybr. hippophorbiae do not accept Hippophae rhamnoides,

E. A. LOELIGER: Celerio hybr. Pauli 219

whereas the cross Celerio hybr. (hippophorbiae x euphorbiae) 3 with (hippophor-

biae x hippophaes) © does so, including euphorbiae-like specimens, must be

explained by chromosomal crossing over.

Unfortunately, the adults of our tertiary bastards (Fig. 2) are all distinctly

different from Celerio hybr. Pauli (Fig. 1). We must consequently agree with

Denso’s criticism and reconsider the possibility of a bastard with another Celerio

species, most likely with Celerio livornica, particularly because of the white flames

on the thorax. Such flames were not present on any of the specimens bred by us. If

ab ovo caterpillars of the primary bastard of Celerio livornica and Celerio hippophaes

too should prove unable to grow on Hippophae rhamnoides (which is certainly also

the case for Celerio hybr. vespertilioides and Celerio hybr. irene), our observation

would offer a model for further experiments with these two species of the genus

Celerio, to elucidate the enigma of Celerio hybr. Pauli Mory. Celerio livornica is a

frequent and well-known guest of the Wallis region, coming from the Mediter-

ranean area, and its fodder plants are even more ubiquitous than those of Celerio

euphorbiae.

ACKNOWLEDGEMENT

The author wishes to thank Dr. A. Diakonoff, Leiden, The Netherlands, and Dr.

F. Karrer, Zofingen, Switzerland, for their interest and help in finding the relevant

literature.

SUMMARY

A tertiary bastard was obtained by crossing Celerio euphorbiae with Celerio hippophaes, i.e. Celerio

hybr. (hippophorbiae x euphorbiae) 3 with (hippophorbiae x euphorbiae) 9. An appreciable proportion

of the ab ovo caterpillars of this bastard, many of which clearly resemble those of Celerio hybr.

hippophorbiae (and hence of Celerio euphorbiae), accept Hippophae rhamnoides, which for ab ovo

caterpillars of the primary bastard Celerio hybr. euphaes and Celerio hybr. hippophorbiae is never the

case. Although the emerging moths are distinctly different from Celerio hybr. Pauli, our experiments

might serve as a model for cross-breeding experiments between Celerio hippophaes and other Celerio

species such as Celerio livornica, in order to explain the enigmatic observation made by Paul in 1897.

LITERATURE

Denso, P., 1911. Celerio hybr. hippophaés 4 x euphorbiae 9 und hybr. euphorbiae 4 x hippophaés 9. —

Entomol. Zeitschr. 25: 151—153.

Mory, E. von, 1901. Ueber einige neue schweizerische Bastarde des Sphingiden-Genus Deilephila und

die Entdeckung abgeleiteter Hybriden in der Natur, sowie Beschreibung einer neuen Varietat

von Deilephilia vespertilio Esp. — Mitt. Schweiz. entomol. Ges. 10 (1): 333—360.

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Fig. 1. Celerio hybr. Pauli Mory; original specimen in the Standfuss collection at Zurich (courtesy of

Prof. Dr. W. Sauter, director). Fig. 2. Cross-breeding of C. hybr. (hippophorbiae x euphorbiae)

& x (hippophorbiae x hippophaes) 9, supposed to be C. hybr. Pauli. Fig. 3 and 4. Caterpillars of C. hybr.

(hippophorbiae x euphorbiae) 3 x (hippophorbiae x hippophaes) 9, supposed to be C. hybr. Pauli, red

variety on Euphorbia polychroma (3) and black variety on Hippophae rhamnoides (4), respectively

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HARVARD

UNIVERSITY

INHOUD

i W. N. ELLIS. — Autumn fauna of Collembola from Central Crete, p.221—326, fig.

| 1—46

} .

| | Tijdschrift voor Entomologie, deel 119, afl. 8 Gepubliceerd 29-XII-1976

AUTUMN FAUNA OF COLLEMBOLA FROM CENTRAL

CRETE!)

WILLEM N. ELLIS

Instituut voor Taxonomische Zoölogie (Zoölogisch Museum), Amsterdam

With 46 text-figures

ABSTRACT

A systematic and faunistic account of the Collembola occurring in central Crete is presented.

All material was collected in the second half of October, 1972, by various techniques, including

portable Berlese funnels. In all, 93 species are recognized. Of these, 15 represent new taxa: the new

species Mesaphorura critica, Onychiurus xenonis, Acheroxenylla cretensis, Hypogastrura tethyca, Xenyllodes

minitaurus, Pseudachorutes (Pratanurida) mucronata, Neanura cretensis, Lathriopyga anthrenoidea, Crypto-

pygus triglenus, Clavisotoma albertinae, Dimorphotoma porcellus, Troglopedetes cretensis, and Pseudosi-

nella paprivata, and two new subspecies Odontella nana orientalis and Sminthurinus alpinus bisetosus.

The new genus Acheroxenylla (monotypic for A. cretensis n. sp.) closely resembles Xenylla but has only

2+2 ocelli. Heteromurus sexoculatus Brown, 1926, and Seira graeca Ellis, 1966, are revalidated.

Onychiurus sublatus Gisin, 1957, O. gisini Haybach, 1960, are synonymized with O. prolatus Gisin, 1956,

as well as the two subspecies O. p. conlatus Gisin, 1962, and O. p. trilatus Gisin, 1963. A key is given

to recently described species of Mesaphorura, and the better-known European Seira. Critical discus-

sions of many species are included, and mass occurrences of Clavisotoma albertinae and Dimorpho-

toma porcellus are reported.

INTRODUCTION

Although Crete has received much attention from collectors, its collembolan

fauna has been completely neglected. To make a start on the faunistic and system-

atic exploration of the springtail fauna of this splendid island, the author and his

wife made a collecting trip between Oct. 14th and Oct. 30th in 1972. Since we had

our base in Iraklion, mainly this region was sampled. This is the lowest, and

possibly not the most interesting part of the island, and extensive collecting in

the mountains is still urgent. Nevertheless, an interesting fauna was found.

The time of collecting coincided with the transition from the summer drought to

the autumn rains; this is the short season when geophytes produce their leaves,

and annuals germinate, twining the landscape from brownish-yellow to green

within a couple of weeks. All of the collected material is treated here, and all

specimens are kept (mostly on slides, mounted separately in Marc André II unless

otherwise mentioned) in the collections of the Zodlogisch Museum, Amsterdam.

The material was collected manually, with the sweeping net, or more usually

with a set of portable Berlese funnels based on the model advocated by J. T.

!) Manuscript received: 15 August 1975

222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Salmon. Unless specified otherwise, all collecting sites were at altitudes of less

than 400 m above sea-level. To avoid undue length, the list of references is in

general restricted to descriptions published since the appearance of Salmon’s

Index to the Collembola (1964).

This is my third contribution to the study of the Greek fauna (the first, on two

species from the mainland, was published as early as 1966, the second, concerning

the fauna of Rhodes, appeared in 1974). Since more are planned, zoogeographical

and ecological discussions are kept to a minimum in the present paper. These

subjects will receive the attention they deserve after the fauna of more localities

has been treated.

Again, help without which this paper could not have been completed was

generously given by colleagues, among whom special mention should be made .

of J.-M. Betsch (Brunoy), R. Dallai (Siena), P. N. Lawrence (London), P. F.

Bellinger (Northridge, California), Z. Massoud (Brunoy), A. Szeptycki (Krakow),

and R. Yosii (Kyoto). The assistance of our technician, L. B. Panday, and our

secretaries T. Dovale and E. C. Gräper is gratefully acknowledged. Prof. J. H.

Stock and Prof. J. T. Wiebes are sincerely thanked for their critical reading of the

manuscript.

List of samples

1. Kriti, Thérisos, 14.x.1972, (ca. 5 km W. of Iraklion), upper 5 cm of rather dry,

comparatively loose loam covered by Capparis spec., on slope along street;

author’s collection number 972.215.

2. Tsagaräk (a village 17 km S of Iraklion, along the Giöfiros River), 15.x.1972,

crumbly, dry loam, sampled at a depth of 10 cm in an olive grove without

undergrowth; 972.217.

3. Tsagarak, litter and topsoil (small hard lumps of dry loam) under an isolated

Quercus coccifera L. in phrygana (i.e., a very common low vegetation type,

comparable to the French garigue, consisting mainly of small shrubs and

geophytes); 972.234.

4. Festos, 16.x.1972, rather damp, crumbly loam in trodden meadow on small

island in the Geropotamos River; 972.197.

5. Festös, rather dry crumbly loam with many roots and many stones in small

bush along Geropótamos, consisting of Vitis, Punica, and Ficus with little

undergrowth of geophytes; 972.227.

6. Festös, along Geropotamos: loose soil and root-mat of an opulent vegetation

comprising various herbs and tall grasses (ca. 60 cm) with a somewhat ruderal

appearence; 972.239.

7. Festös, along Geropötamos; trodden, compact, rather moist loam with many

pebbles, under a low vegetation, mainly clover; 972.236.

8. Festós, along Geropótamos: swept from low grass of meadow; 972.213.

9. Fortétsa (about 5 km SE of Iraklion), 18.x.1972, after two days of heavy

rain, wet swollen loam of vineyard; 972.224.

10. Fortétsa, wet sandy loam, trodden and ruderal, with some perennial grasses;

972230!

31.

32.

W. N. ELLIS: Collembola from Central Crete 223

. Fortétsa, moist compact loam under large almond tree in fallow field; many

goat droppings; 972.208.

. Ioùchtas, a hill near Archanes, 10 km SSW of Iraklion, 19.x.1972, 620 m, loose

warm-brown terra rossa under low bushes in phrygana, west exposition;

972.240.

. Toúchtas, 700 m, loose terra rossa half under Quercus coccifera on yoke of the

hill; 972.221.

. Ioùchtas, 760 m, loose terra rossa in phrygana on west exposition; 972.229.

. Iouchtas, 600-700 m, swept from bushes (mainly Pistacia lentiscus L.); 972.210.

. Amnisòs, 5 km E of Iraklion, 20.x.1972, stony yellowish compact loam under

Sarcopoterium bush in degraded phrygana on weak east slope; 972.238.

. Amnisòs, rather dry, loose, humus-rich sand on 3 m high cliff, along the sea,

under shabby herbaceous vegetation; 972.241.

. Amnisòs, loose loam at north foot of a hedge of Arundo donax L. (a tall grass,

2-4 m high) along field, 972.198.

. Agia Varvara, 21.x.1972, 700 m, pleurocarpous mosses and Selaginella denticu-

lata (L.) Link, on short, almost vertical slope, northwest exposition, in phry-

gana; 972.199.

. Agia Varvara, 750 m, humus-rich very stony loam with a scanty grass and

Juncus vegetation; the spot seems to be a spring in the spring season; 972.228.

. Agia Varvara, 750 m, opulent cushion of pleurocarpous moss on north bank

of dry rivulet in degraded phrygana; 972.223.

. Agia Varvara, 600 m, swept from low bushes (Sarcopoterium) in phrygana;

972.203.

. Rethimnon, 22.x.1972, verge of a road near the city, comparatively damp,

heightened rather recently with coarse (beach-?) sand, overgrown with scanty

grass; 972.225.

. Rethimnon, comparatively damp plastic yellow loam at the foot of an earthen

wall about 2 m high across a field, north exposition; 972.216.

. Drosia, between Iräklion and Perama, about 15 km E of the latter, 23.x.1972,

crumbly loam, abundant mosses and litter under large Quercus coccifera, at

base of north side of an east-west running ravine; 972.212.

. Drosia, upper 2 cm of bare compact loam, with a thin carpet of ephemerous

liverworts (Riccia, Fossombronia, Targionia) in strongly degraded phrygana on

south wall of the same ravine as 25; 972.243.

. Drosia, litter under large Quercus coccifera on south slope; 972.226.

. Drosia, large isotomid, walking in large numbers freely on stones, soil, etc.,

on south slope, collected manually; 972.214.

. Knossos, vicinity of the excavations, 24.x.1972, crumbly yellow loam, rarely

trodden, under large peach tree on north slope of brook near the ‘‘guest-

house” (Xénon), undergrowth mainly Oxalis pes-caprae L.; 972.242.

. Knossos, loose loam, sparsely grown with grass and Oxalis pes-caprae at foot of

a 4 m-high cliff along road; 972.219.

Knossos, collected manually under stones, pieces of dead wood, etc.; 972.195.

Iraklion, 25.x.1972, after heavy rains, loam, litter and rotting leaf bases under

vigorous ruderal vegetation (Ecballium elaterium (L.) Rich.); 972.233.

224

33%

34.

35:

36.

38.

39:

40.

41.

42.

43.

44.

45.

46.

47.

48.

49.

50.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Iraklion, loam of ruderal field, overgrown mainly with grass and Carpobothrus

along road; 972.232.

Iraklion, bank of loam overgrown with strongly ruderal vegetation; 972.244.

Marathos, 15 km W of Iraklion, 26.x.1972, litter under Pistacia lentiscus in

phrygana on weak north slope; 972.209.

Marathos, upper 2 cm compact bare loam, with ephemerous vegetation of

liverworts (Fossombronia) and seedlings, along roadside; 972.231.

Marathos, litter under large Ceratonia siliqua L. in field; 972.237.

Marathos, collected manually from stones and pieces of wood, and swept

from Pistacia lentiscus; a Seira on an olive trunk; 972.196.

Marathos, large extremely numerous isotomids, walking on stones and on

bare soil; 972.204.

A mnisòs, 27.x.1972, upper cm loam along beach, with very spare vegetation of

Salicornia fruticosa (L.) L., some annual mosses (Bryum spec.), near Tamarix

bush; 972.200.

A mnisòs, loamy sand of fallow field with a mat of Malva spec.; 972.235.

Gazi, 7 km W of Iraklion, 28.x.1972, loamy clay under Ecballium in fallow field

500 m from the shore; 972.206.

Malia, 29.x.1972, brown, crumby, stony loam grown with a poor grass vegeta-

tion under large Ceratonia tree; 972.205.

Malia, litter of Quercus coccifera in well-developed phrygana; 972.211.

Malia, terra rossa, acrocarpous mosses and Selaginella deep in a rock-fissure

at foot of north slope of ravine; 972.222.

Malia, collected manually under stones in phrygana; 972.201.

Agia Galini, 14 km NW of Festòs, 30.x.1972, beach sand and Posidonia (a kind

of sea grass) debris in a layer of about 2 cm; 972.218.

Agia Galini, Tamarix litter and fine sand, overgrown with Malva and Oxalis

pes-caprae just above the beach; 972.207.

Agia Galini, yellowish-brown loam of lucerne field; 972.220.

Agia Galini, collected manually, under large pebbles on the beach; 972.202.

Table 1. Greek orthography and transcription of the locality names used

Konm Kriti ’Auviodg Amnisös

Oéproos Thérisos ‘Ayla Bopßäga Agia Varvära

Toayapäx Tsagaräk Péôvuvov Réthimnon

‘Hoáxherov Iráklion Aooovd Drosia

Fiépvoos Giófiros Tléoapa Pérama

Deorôc Festös (in Times Atlas Phaistos) Kvmoég Knossos

Teow-Ilötauog Geropótamos Méeatos Marathos

®ootétoa Fortétsa Tatuov Gazi

Ioúyras Touchtas Madara Malia

"Aoxaves Archanes ‘Ayla TaAfvn Agia Galini

W. N. ELLIS: Collembola from Central Crete 225

ACCOUNT OF THE SPECIES

ONYCHIURIDAE

Metaphorura affinis (Borner, 1902) (Fig. 1)

Material: see table 2.

Discussion. The rather extensive material of this common European species that

I could study is strongly heterogenous in some characters of the apex of the

abdomen. The material can be separated into two forms, A and C, connected

by a few intermediate specimens I shall refer to as B. For the number and distri-

bution of these forms, see Table 2.

The forms can be segregated as follows. Specimens of form A have the dorsal

granulation on abd, so strongly developed that the secondary granulations touch

each other, giving a roughly hexagonal outline (Fig. 1d). Between the macro-

chaetae p, on abd,, ten such coarse granules are present. The granules also have a

distinctly flattened apex, as can be seen in profile. The same type of integumentary

ornamentation is visible on the dorsal surface of head, thorax, and abd,_,. On

abd, the granules are not flattened or hexagonal, but more or less globular.

Laterodorsally, abd, shows an area where these granules are grossly enlarged

(Fig. lc). Furthermore, specimens of this type invariably have the sensilla s on abd,

very short — in fact, reaching only to about the middle of the pseudocellus of abd,

(Fig. 1b, c). Finally, the anal spines are slightly heavier and more strongly yellow-

tinged than those in the types B and C.

Specimens of type C have the integumentary granulation rather fine and

regular, the granulations not touching each other and having a globular profile.

Only the granulation of abd, is much coarser, but the two areas with strongly

swollen granula are absent. Moreover, seta s on abd, is distinctly longer, reaching

well beyond the posterior margin of the corresponding pseudocellus (Fig. 1a).

As mentioned above, the anal spines seem to be more slender and paler. About

17-18 granulations are present between the p, on abd,.

The intermediate type B has the granulations enlarged, though not so strongly

Table 2. Distribution of types A, B, and C within the samples of Metaphorura affinis

A B C sample nr. locality

DIO gl uv. SÌ Festos

O2 guy: 6 Festos

1595 1 juv. l juv. 24 Rethimnon

19, I juv. 48 Agia Galini

39,13 49 Agia Galini

l ©, 2 juv. 30 Knossos

39 29 Knossos

19 16 A mnisòs

1 juv. 9 Fortétsa

19 14 loúchtas

DOF 2 juv. 20 Agia Varvara

SHOR i uve 45 Malia

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 1. Metaphorura affinis (Börner). a, abd, of type C 9; b, abd, , of type A 9; c, abd, ‚of type À ©

(on abd, setae a, omitted); d, hexagonal arrangement of secondary granulations between p,-p, on abd,

of type A 9 (very strong magnification)

W. N. ELLIS: Collembola from Central Crete 227

as in type A (between the p, on abd, there are about 12 granulations), and they are

only slightly hexagonal in outline and flattened at the top. On abd, the strongly

enlarged lateral granulations are completely or almost absent. Seta s on abd, is just

as short as in type A. Evidently, type B is much closer to type A than to type C,

and it could be argued that A and B should be united.

Borner (1902) described his Tullbergia affinis from many specimens from Sicily

and Calabria, and called the integumentary reticulation ,,fein und gleichmaszig,

nur auf dem Kopfe ein klein wenig kraftiger”. His material thus seems to agree

with my form C.

I have checked the other material of this species in our collection. Material

from Iceland, Sweden, Austria, and the CSSR proved to be referable to form B;

material from Yugoslavia, as well as the specimens recorded by me from Rhodes,

are identifiable as belonging to C.

As far as I know, form A, which seems to deviate the most from typical affinis,

cannot be correlated with any species description. The drawing made by Hand-

schin (1929) of abd, of his Tullbergia bipartita shows a perfectly simple granulation,

only compatible with type C.

Cassagnau (1963) described a comparable diversity in granulation in material of

M. bipartita (Handschin, 1920) from North Africa: some specimens, considered

to be aberrants, showed a remarkably coarse granulation (consisting of rounded

papillae) all over the body (also the ventral face?), whereas abd, bore the two types

of enlarged granulations described above. Moreover, the unpaired, conical organ

on abd, was reduced to at most a vague boss. The last observation differs widely

from what I found in Cretan material, where the unpaired cone is well-developed

in all forms.

I admit that the complex situation described above could be explained by

assuming the presence of two species, affinis and bipartita, in the present material.

But I did not succeed in discerning the particular bilobation in the elements of the

postantennal organ said to be characteristic for bipartita; I even wonder whether

these structures are visible at all with light microscopy, and I am inclined to share

Gisin’s (1944) opinion that the two taxa are synonymous.

As is evident from Table 2, the three types cannot be simply related to sex or

maturity. However, since some very young specimens have seta s strongly de-

veloped on abd, and a reticulation of type C, it is possible that type A concerns

specimens that are not only sexually mature but also completely full-grown.

Cassagnau (l.c.) brought the described phenomena into tentative corres-

pondence with ecomorphosis. This possibility merits closer consideration,

although the seemingly normal fat body, the full intestine, and the apparent

sexual maturity of many specimens of form A all argue the contrary. However,

this suspicion and the fact that intermediate forms between the extremes occur in

Crete and elsewhere, make it impossible, at present, to attribute specific status

to types A and C. Their virtually sympatric occurrence within Crete is sufficient

reason not to create subspecies.

One female from sample 29 is aberrant in that it completely lacks the anal

spines. The median process is present however.

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Metaphorura spec. (Fig. 2)

Material: one immature specimen from sample 30.

Discussion. The single specimen, measuring 0.6 mm, is so immature that no

trace of a genital orifice is visible. It is very unfortunate that more and better

material is not at hand, since the specimen shows some interesting features.

The 6th abdominal segment has 2+2 spines, and therefore identification as a

Stenaphorura seems appropriate (Fig. 2a). The relatively small postantennal organ,

with a small number of elements (Fig. 2b), the structure of ant,, and the pseudo-

cellar formula 11/122/22221 point toward identity with Tullbergia (Stenaphorura)

gisini Selga, 1963, described from the Sierra de Guadarrama. Seemingly, the

chaetotaxy of the Greek specimen is in good agreement with the Spanish material.

However. there are the following important differences:

(i) the Greek specimen has an unguiculus (of about 1/3 unguis);

(ii) the accessory sensilla in antennal organ III is lacking;

(iii) the ant. org. III is covered by three high, thin skin flaps (Fig. 2 c);

(iv) the postantennal organ is simple in outline;

(v) the apex of abd, bears a tubercle which is completely comparable to that of

Metaphorura (though slightly lower than is usual in M. affinis which in this respect

is notoriously variable).

A median tubercle as in Metaphorura is described by Gisin (1963a) in his Tull-

bergia novemspina from Yugoslavia; judging from the figures given by Gisin,

however, this species differs not only by having 4+4 lateral spines on abd,, but

also by a very different chaetotaxy of abd,

Neotullbergia tricuspis (Borner, 1902)

Tullbergia ramicuspis Gisin, 1953, 1960, Dallai, 1973 (nec T. tricuspis: Gisin, 1944, 1960).

Material: 1 © from sample 24.

Discussion. The species has been recorded from central and southern Europe

(Spain, Italy, France, Yugoslavia). The single specimen from Crete agrees well

with the description of ramicuspis by Gisin (1953), but the anal spines may be a bit

more slender.

The complicated synonymy of this species has been clarified by Huther (1961).

However, another problem of synonymy still has to be settled. Bonet (1944)

recognized in Lipura pusilla Giard, 1895 (description corrected in Giard, 1896;

note also that Bonet mentions wrong publication dates) a Neotullbergia. He was

not able to locate this species, but among material from Chili Rapoport & Rubio

(1963) found a species that fitted Giard’s description. These authors overlooked

Giard’s remark that he knew L. pusilla not only from material from Chili but also

from Cape Gris Nez in NW France. This fact, the scanty description, and the small

size (0.6 mm) of pusilla, all favour the supposition that pusilla might be, at least

partially, immature tricuspis. To preclude the annoying complications that would

result, I herewith restrict the type locality of Giard’s species to Santa Rita, Chill.

W. N. ELLIS: Collembola from Central Crete 229

Fig. 2. Metaphorura spec. a, abd,_,; b, postantennal organ and neighbouring pseudocellus; c, antenna,_,

230 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Mesaphorura critica n. sp. (Fig. 3)

Material: sample 6: 1 9; 20: 6 9; 23: 1 9; 24: 5 9 and 2 juv.; 25: 3 Q and 2 juv.;

27:1 9; 33:1 9; 49:2 9. The © from sample 23 is selected as holotype.

Description. Habitus as usual in the genus; total length about 0.4 mm. Granula-

tion fine and regular, a bit coarser on abd,.

Fig. 3. Mesaphorura critica n. sp. a, ant,; b, abd,_;; c, ventral tube; d, th,; e, chaetotaxy of anal region

W. N. ELLIS: Collembola from Central Crete 231

Ghactotaxwth/amis;:th;:as, 1m; 03,184 S Pio Py24; thas tha(Fig. 3d); abd,:

np; abd, idem; abd, idem; abd,: a,_,, m,, p,.: abd,: a;,-24 Py 5 Pacs

Distinct macrochaetae on abd, are a, and p,, on abd, a, and p,. Anal flaps without

setae |’, (Fig. 3 e).

Rosette-shaped pseudocelli distributed as 11/011/10011; those on th, and th, are

situated between setae m, and p..

Antennae with the normal sensory complement. Sensilla b thick (Fig. 3a). Ant:

head Q = ca. 0.65. Postantennal organ about 1.5-2 times pseudocellus. Labium

with 4+4 setae. On th, , the “Lateralsensille’” s, distinguished by Rusek (1971b)

not differentiated from a normal hair; sensillae s’ present on these segments.

Sensilla s on abd, swollen, 9.8 » (for comparison: the lenght of seta p, on this

segment is 16 u).

Claw toothless, 8.5 u, unguiculus needle-like, 3 u.

Semicircular ridges on abd, evident, anal spines 6 u, curved (Fig. 3b).

Ventral tube with 4 +4 apical setae, 1 laterobasal, and 1 lateral seta (Fig. 3c).

Discussion. The material under consideration is exactly intermediate between

M. italica and M. sylvatica, both described by Rusek (1971) from Italy, the CSSR,

and Bulgaria. The material is identical to M. italica, except for the lack of seta

a, on abd,; otherwise it is identical to M. sylvatica, but the pseudocellus on th, is

situated between setae m, and p,.

The above situation might either be interpreted as an indication of synonymy

of italica and sylvatica or necessitate the description of a new species. The perfect

uniformity of all specimens studied from Crete finally persuaded me to choose the

second alternative.

The specific name refers to the ambiguous state of the new species, alluding at

the same time to the Greek name Kriti.

Mesaphorura italica (Rusek, 1971)

Material sample 11:,lijuv.;17:4 9; 23: 1 9; 24:1 9525:3 9; 30:1 9; 3521 9%

The material is in full agreement with Rusek’s detailed description. The species

was subsequently recorded by Rusek (1973b: Italy) and by Ellis (1974: Rhodes).

Mesaphorura krausbaueri Borner, 1901

Material: sample 40: 11 9 and 2g ; 43:1 9.

Description. The material differs consistently from the description given by

Rusek (1971) in one respect: in all specimens studied seta I’, is present on the anal

flaps. However, this is not only the case in Greek material: a casual study of some

Dutch specimens showed that this seta is also present there.

Since the delimitation of this species has been drastically narrowed by Rusek’s

work, it is impossible to say much about its geographical distribution. As con-

ceived of at present, it is known from the CSSR, Italy, the Netherlands, and Crete.

The occurrence of males in the Cretan material is noteworthy.

To synthetize the knowledge of the krausbaueri group, I constructed the

following, simple key to the species.

232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Key to the species of the Mesaphorura krausbaueri group

1. abd, in frontal row with 1+1 microchaetae between bordering macrochaeta

(a) inithis case): 5 se LIRICA ee ee ee hygrophila (Rusek, 1971)

— 2+2 microchaetae between bordering macrochaetaa, ............ 2

— 3+3 microchaetae between bordering macrochaetaa, ............ 3

2. pseudocellus on th, behind or between p; and p, ... sylvatica (Rusek, 1971)

— this pseudocellus,betweenm, and ps „en Lern oOo: critica n. sp.

3, /thywith.2=# 2,pseudocelli 5. «clio PRO SSR 4

—, th, with. L#.I,pseudocelli, … up er un oge HAT an eke D

all sensillae strongly developed; in particular sensilla b on ant, inflated

ke morator en sensibilis Rusek, 1973

— most sensillae slender; sensilla bon ant, thin .... tenuisensillata Rusek, 1974 i

5. pseudocelli on th, ‚situated between m, and pjy ...... italica (Rusek, 1971)

— these pseudocelli between or behind p, and p, 2 tot Beyer 6

6. on abd, p, is a macro-, p,a microchaeta ........ krausbaueri Börner, 1901

— on this segment p, is a micro-, p,a macrochaeta ....... yosiii (Rusek, 1967)

Protaphorura prolata (Gisin, 1956)

Onychiurus prolatus Gisin, 1956.

O. sublatus Gisin, 1957. — nov. syn.

O. gisini Haybach, 1960. — nov. syn.

O. prolatus conlatus Gisin, 1962. — nov. syn.

O. prolatus trilatus Gisin, 1963b. — nov. syn.

Material: sample 9: 4 9 and 1 5; 18: 8 9 and 10 3 ; 20: 1 juv.; 21:1g and

1 juv.; 24:9 9, 11 & and 12 juv.; 30: 8 juv.; 31: 1 9; 32:6 9 + 36 ex. in alcohol;

34:5 9,34 and juv.; 36: 1 3 and 1 juv.; 37: 12 9 and3g ; 38:6 9,73 and 12

juv.; 44: 1 juv.

Discussion. The extensive material (in all 125 specimens) is very homogenous,

and variation is not much greater between samples than within samples. The

largest specimens measure 1.8 mm, but adult specimens are normally 1.3 mm long.

The dorsal pseudocellar arrangement was found to be very constant, viz.,

33/022/33343. In fact, not a single deviation from this formula was noted. The

lack of a tooth on the unguis is also constant. Absence of seta m on th, was

doubtful only once (it did occur in one adult specimen, and only asymmetrically).

Seta i is sometimes missing, especially in immature material. Much more variable

is the number of microchaetae in hind row of th,. Rather often, only two were

found (although almost always asymmetrically, the higher number being present

on the other half of the segment). At the base of the ventral tube, 2+2 setae are

usually present, but an arrangement 1+1 occurs relatively frequently. Abd, inva-

riably lacks seta s’.

The shape of the anal spines, in itself difficult to assess exactly, is rather

variable, ranging from rather stout to rather slender. The arrangement of the

W. N. ELLIS: Collembola from Central Crete 233

four prespinal setae on abd, cannot be regarded as a useful character, gradual

transitions being demonstrable between almost parallel and forming a blunt angle.

The M/s ratio on abd, (following Gisin’s convention, in which the length of the

anal spines is set at 10) is variable. Some random measurements, all made in

adult specimens, are: sample 9: 12/7; sample 18: 11/6, 11/6, 12/6, 12/6, 13/6; sample

32: 24/11; sample 34: 18/10; sample 37: 18/11; sample 44: 18/12, 20/12, 20/10.

Formidable as this may seem, it only means a shift in the length ratio of M and

s from 1.5 to 2.2; often it is also difficult to measure the exact length of hairs very

accurately, because they do not lie parallel to the optical plane. The same holds

even more strongly for the measurement of the anal spines.

The circumstances compelled me to recognize a rather iconoclastic set of new

synonymies.

Onychiurus sublatus Gisin, 1957, was described after an unstated number of

specimens from Austria and England. The species was considered to differ from

Onychiurus prolatus (1) because of its smaller size, viz., 1.4-1.85 mm as against

2.3-2.6 mm in prolatus, (2) because the anal spines and claw are plumper in subla-

tus, and (3) because M/s = 24/7-8 (in prolatus originally given as 21/9, but in

Gisin (1960) “as in latus”, which would mean 29/10).

Onychiurus gisini Haybach, 1960, described from Austria, was contrasted with

O. cancellatus Gisin, 1956, because the prespinal setae of abd, were said to

delineate two strongly convergent lines. Apart from this character, the doubtful

value of which is discussed above, there is no essential difference from O. prolatus.

Onychiurus prolatus conlatus was described by Gisin (1962) after material from

caves in eastern Switzerland (prolatus prolatus was described originally from caves

in the Savoy Alps and the Swiss Jura). Differentiating characters are (1) M/s ratio

19/14 (instead of 21/9), (2) a difference in shape of abd, (as seen in profile) and

(3) the four prespinal microchaetae of abd, forming two slightly converging lines.

(Gisin adds here that he had also observed this in a typical population of

prolatus!)

Onychiurus prolatus trilatus was described by Gisin (1963b) from caves in France

(Dröme). The subspecies was characterized by having two instead of one median

hair on the tergum of abd,; however, at least three adult specimens from Crete

show this same character.

Onychiurus pseudoghidinii Dallai, 1969 (Fig. 5a)

Material: sample 4: 1 adult and 2 subadult 5 .

Discussion. The adult specimen clearly demonstrates the (only) character that

distinguishes pseudoghidinii from ghidinii Denis, 1938, viz., the third pair of pseudo-

celli at the ventral face of the head (Fig. 5a). However, in one of the subadult

specimens these pseudocelli are greatly reduced, and in the other they are absent.

In my opinion, this casts some doubt on the validity of Dallai’s species, but in

view of the paucity of my material I must limit myself to drawing attention to the

imminent synonymy.

O. pseudoghidinii is described from the Isle of Montecristo (about 50 km S of

Elba); ghidinii, described from the north of Italy, is also known from France,

234 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 4. Onychiurus xenonis n. sp. a, th,; b, abd, ,; c, ventral aspect of head; d, posterior margin of head

and th,

W. N. ELLIS: Collembola from Central Crete 235

Switzerland, Bulgaria, Morocco, Madeira, and the Azores. Material from the last

locality has been described in detail by Hüther (1970). The material now under

consideration coincides completely with Huther’s description, except that the

granulation of the Greek material seems to be somewhat less developed.

In one specimen antennal organ III was guarded in both antennae by only four

papillae. The other two specimens had the normal number (five).

Onychiurus cf. stachianus Bagnall, 1939

Material: sample 6: 1 © ; 24: 1 juv.

Description. The single adult specimen measures 1.2 mm. It agrees with the

description of stachi Denis, 1938 (nec Bagnall, 1935, rebaptized stachianus by

Bagnall (1939)), except that the ungues have no lateral teeth. In the absence of

adult males the identification can, of course, only be tentative.

The species was described from the Postumia cave in Yugoslavia; it has been

recorded from the CSSR by Rusek (1959), from England by Goto (1953), and from

Spain by Selga (1962a). Confusion with O. pseudostachianus Gisin, 1956 (= stachia-

nus sensu Gisin, 1952) in these records cannot be ruled out, however.

Onychiurus xenonis n. sp. (Fig. 4)

Material: sample 29: holotype 9 and 6 9, 3 4, paratypes.

Description. Length only 0.6 mm. White, integument finely and regularly

granulated. Antennal bases not differentiated. Differentiation into micro- and

macrochaetae weakly developed; some setae swollen to the shape of sensillae.

No anal spines.

Antenna, with small retractile papilla and sensilla in a groove above ant. org. III.

The latter consists of 5 setae, 5 slender papillae, 2 straight rods, and 2 curved

smooth sense clubs. Postantennal organ compound, with about 12 primary tuber-

cles. Claw without inner or lateral teeth, unguiculus gradually tapering to a slender

apex, reaching tip of unguis. No trace of furca.

On head and on all body segments except th, and abd,, one or two pairs of

hairs are swollen and have acquired the shape of sensillae or small spinules.

On the head two such pairs are present, not far from the hind margin (Fig. 4d).

Also th,_, each have a pair of such setae, which are situated laterally, just dorso-

caudad to the lateral sensilla, which is apparently characteristic for all Onychiuri-

dae and Hypogastruridae (Fig. 4a). Abd,_, each have two pairs, one of which is

situated in about the middle of the width of the segment anterior to the second

pseudocellus from the median, whereas the second pair of sensillae lies antero-

laterally from the third pseudocellus (Fig. 4b). Abd, has only one pair, situated

in front of the third pair of pseudocelli, and abd, again has two pairs, one

near the anteromedian (first) pair of pseudocelli, the second anterior to the fourth

pair of pseudocelli. On the sternite of abd,, two indistinct pairs of these sensillae

are present in front of the pseudocelli. Ventral tube with 6 + 6 setae.

Pseudocelli: Dorsal arrangement of pseudocelli 32/133/33354, ventral arrange-

ment 4/000/2222. The 4th pseudocellus under the head is situated near the lateral

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 5. Onychiurus pseudoghidinii Dallai, a, pseudocelli on ventral face of head. Acheroxenylla cretensis n.

sp. b, ventral chaetotaxy of abd,_y; c, antennal organ III; d, abd, dorsally; e, ant,_,; f, claw of P,

W. N. ELLIS: Collembola from Central Crete 237

border of the labium. The pseudocellus is slightly oval and easily overlooked

(Fig. 4c).

In the male specimens no male organ is discernible.

Discussion. The new species belongs to the Onychiurus (s. str.) complex of

species, having normal unguiculi, smooth sense clubs in the antennal organ, and

at the same time lacking anal spines. The species is conspicuous in having no less

than 4 pseudocelli on the ventral face of the head, and also in possessing a

number of spine-like ‘“‘sensillae’”. This last character suggests a relationship to

O. edinensis Bagnall, 1935, according to Murphy (1960) the valid name for O.

spinularius Gisin, 1952. This species, however, possesses strong anal spines.

The fourth ocellus at the ventral face of the head is situated close to the lateral

edge of the labium, and is easily overlooked. Nevertheless, xenonis is not easily

confused with the species with which it seems otherwise to be most related with

respect to the arrangement of the pseudocelli, e.g. O. dunarius Gisin, 1956.

HYPOGASTRURIDAE

Acheroxenylla n. gen.

Diagnosis. Homochaetotic Hypogastruridae. Antennae with retractile papilla,

3 outer and | inner cylindrical sense hairs. Postantennal organ, unguiculus, furca,

and retinaculum absent. Ocelli 2+2. Tibiotarsi with 2 tenent hairs. Anal spines

present.

Type species: Acheroxenylla cretensis n. sp.

This is another small genus of xenylline stock. Clearly, it is a close relative of

Xenylla, as is evident from the absence of a postantennal organ, number and

location of sense hairs on ant, and of tenent hairs on tibiotarsus, absence of the

unguiculus, and the general appearance of the chaetotaxy.

However, in Xenylla total absence of a furca is rare; Salmon (1944) created the

genus Propexenylla for one species having this character. But there is a complete

gradation between fully developed furca (for Xenylla, anyway), through X. boerneri

Axelson, to X. acauda Gisin and Propexenylla atrata Salmon. Consequently,

I agree with Da Gama (1969) and place atrata in Xenylla.

The reason why I nevertheless separate A. cretensis from Xenylla lies in the

number of ocelli. In almost all species of Xenylla (roughly 60) the number of ocelli

is 5+5; in a few instances it is 4+4. I feel that inclusion of cretensis would unduly

enlarge the scope of Xenylla in this respect.

Within the genera derived from Xenylla having fewer than 5+5 eyes, Ache-

roxenylla comes closest to Acherontiellina Salmon, 1964. This genus was recently

redefined by Djanaschvili (1971a, b) as having cylindrical sensillae on ant,,

possessing anal spines, and completely lacking the furca. Yet cretensis is placed in

a genus of its own, to avoid disturbing the homogeneity of Acherontiellina and its

near relative Acherontiella Absolon, 1913, both of which comprise only eyeless

forms.

238 TIJDSCHRIFT VOOR ENTOMOLOGiE, DEEL 119, AFL. 8, 1976

Key to the genera of the Xenylla group:

Po ocell SAE Led RO EL Biscoia Salmon, 1962

SOMME ee eo omen ae ee et ur Xenylla Tullberg, 1869

OE IT NI Acheroxenylla n. gen.

OCTO OC LTT SOIN CSO

furca completely absent asc e REA AE PSR REN

furca more or less'reducedbut present’ O arme TT

anus nearlyventrals sn tse. Xenyllina Delamare Deboutteville, 194

anusterminal An eten N I N

sensillae on ant, about spherical, anal spines absent .................

Re ee TA DIES, N ONU SE ed Acherontiella Absolon, 1913 :

— sensillae on ant, about cylindrical, anal spines present ................

ER MALDI E ONERI, (ROS AIA RE RR I kene RE Acherontiellina Salmon, 1964

5. mucro not separated from dens, or absent; sensillae on ant, cylindrical .....

LR IVI NS Acherontides Bonet, 1945

— mucro separated from dens; sensillae on ant, conical, sometimes oval .....

SIE Pseudacherontides Djanaschvili, 1971

I Se

B OO Un © D

This key is essentially based on the work of Djanaschvili (1971a, b). I must add

that I am not certain as to the inclusion of Biscoia in this group. The high eye-

number, the presence of an unguiculus, the well-developed dens, and the peculiar

antennal organ III, argue against its inclusion. I have nevertheless included the

genus in this key, because it is reported to lack the postantennal organ.

Acheroxenylla cretensis n. sp. (Fig. 5 b-f, 6-8)

Material: sample 1: 21 9, 32 3 and 2 juv.; 11: 18 9, 13 g and 5 juv.; 17:7 9;

26:6 9, 1g and 1 juv.; 27: 1 g and 3 juv. Moreover preserved in alcohol, not

studied in closer detail, from sample 1: 196 specimens, 11: 210 specimens (mixed

with some Acherontiellina that cannot be distinguished habitually), and from

sample 17: 27 specimens. The holotype is a g from sample 1.

Description. Length 0.7 mm. White; pigment absent, except for a few pigment

granules clustered around the separate ocelli and occurring only in a few speci-

mens. Integument coarsely granulate, especially on dorsal parts of thorax and

abdomen. Hair cover not differentiated in micro- and macrochaetae, only setae s

distinctly longer than the others. Anal spines short and curved, on hardly differ-

entiated papillae. Hairs smooth and simple, except the row of 3+3 setae on abd,,

which are conspicuously serrate on their anterior surface (Fig. 5d).

Antennae about 5/6 times head diagonal. Ant, with large retractile papilla, and

3 outer as well as | inner nearly cylindrical, swollen sensory hairs. Between the

group of 3 there is at least one other, very small, sensilla, difficult to observe:

Of the cover of apparently normal setae on ant,, one subapical seta is inserted on

some sort of wart flanking the pit of the retractile papilla (Fig. Se). Antennal organ

ITI consists of two long and slender sensillae, each guarded by a seta. Between the

W. N. ELLIS: Collembola from Central Crete 239

long sensillae there is an integumentary flap that protects two small sensillae (Fig.

SC):

Fig. 6. Acheroxenylla cretensis n. sp. dorsal chaetotaxy of head and thorax

240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

ER A N

— A \

SS | [ N Sr zl wa Pe: bi

= Î ER

Fig. 7. Acheroxenylla cretensis n. sp. dorsal chaetotaxy of abdomen

W.N. ELLIS: Collembola from Central Crete 241

Lo. REC

Fig. 8. Acheroxenylla cretensis n. sp. a, ventral chaetotaxy of head; b, ventral chaetotaxy of abd,_,; c,

ventral chaetotaxy of abd,_,

242 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Postantennal organ lacking. Only 2 + 2 small, widely separated ocelli.

Feet normal, short, unguiculus absent, unguis without any tooth, two dorsal

tenent hairs (with one normal, short hair between them) very faintly clavate

(Fig. 5f). Ventral tube with 4+4 setae (Fig. 8b).

Chaetotaxy. Cephalic chaetotaxy is interpreted along the lines indicated by

Cassagnau (1974), who in essence restored the system of Yosii (1960). Rows p and

c are complete, there is but one seta v, setae d,_, and a, are present, the sd row

is very short, consisting of but two setae, allegedly sd, and sd,. The affinity with

Xenylla is evident, only the sd row is still more reduced. Th, with 3+3 setae.

Th,_, with a,, a, a,_,, m,, m,, m,, and m,, and the p row complete: p, _,. Seta p, is

ss. On th,, moreover, a pair of setae m, is present (Fig. 6).

Abd,_,: seta p, is ss. Abd,: in frontal row only a, and a,, in median row only m,, .

posterior row complete p,_,. Abd,: only a,, p,_, present; p, = ss. Abd, with 343,

anteriorly serrate setae (Fig. 7).

Ventral chaetotaxy: Head with a,, m,, and p, present, m,_, lacking (Fig. 8a).

All thoracal sternites and abd, without setae. Abd,: a,, p,, p3_, (Fig. 8b). Abd:

As 6, My, Py, Pz, Ps_g- It is not possible to analyse the setae on abd, (Fig. 8c).

Subcoxae 1-3 with 1, 2, 3 setae, respectively.

Acherontiellina bougisi rhodia (Ellis, 1974) n. comb. (Fig. 9 a, b)

Acherontiella bougisi rhodia Ellis, 1974.

Material: sample 4: 8 9,94, and | juv.; 6:8 9 andlg;7:5 9 andlg ;9:24&

and 1, juv.; 10:2 95 11 40,17, andıl juv.s 16.292294 9and 1 30229

and 64 532: 2 9,13, and 1 juv.; 33.14 and juv.. 34.37 9 ande

Discussion. The present material is in good agreement with the two specimens

from Rhodes on which I based the subspecies. The larger material now available

permits me to add some supplementary details and to analyse some aspects of the

variability.

The internal tooth at the unguis is usually clearly visible. Generally, though

perhaps not in all specimens, the two tenent hairs are feebly clavate.

As I have already indicated, the dorsal chaetotaxy is identical to that of bougisi

bougisi, discussed and illustrated by Thibaud (1967). However, some variability

interferes with this agreement. In particular, seta a, on the head is rather often

lacking (in 27% of 60 observations). The row of setae sd, basically consisting

of three setae sd, , is reduced in some cases to two hairs sd,_,. In 3% of the

observations this reduction of sd occurred on only one side, in another 4% it

occurred symmetrically (n = 61). In one specimen seta v was lacking on both

sides, and in 5% (3 cases) only unilaterally (n = 61).

The ventral chaetotaxy can be summarized as follows. The head possesses a,,

m,, and m, as well as generally a pair of setae p,. In 16% seta p, is lacking on one

side, in 5% on both (n = 71) (Fig. 9b). The thoracic sternites are without setae.

The ventral tube has 4+4 setae as in Xenylla. Ventral chaetotaxy of abd,_, is very

irregular, in abd, a, is distinct, and there is a very unstable row p, consisting of

5+5 setae; either p, or p, is particularly strong and seems to have a sensory

W. N. ELLIS: Collembola from Central Crete 243

Fig. 9. Acherontiellina bougisi rhodia (Ellis). a, ventral chaetotaxy of abd,_,; b, ventral chaetotaxy of

head. Hypogastrura cf. gisini Strenzke. c, unguis P,; d, mucro exterior; e, dorsal cephalic chaetotaxy; f,

lateral aspect of mucrodens; g, posterior face of mucro; h, i, j, k, postantennal organs

244 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

function (Fig. 9a). In abd, setae a, ; are evident, but it was not possible to discern

a row m or p, nor was the position of a pair of sensory hairs sufficiently stable

to establish a fixed scheme.

By transferring bougisi to the genus Acherontiellina, I adopt the classification

proposed by Djanaschvili (1971a, b), which is assimilated into the key to the

genera of the Xenylla group (p. 238).

Xenylla maritima Tullberg, 1869

Material: sample 3:3 9,14, and 4 juv.; 50: 17 9 and4 + 8 ex. in alcohol.

Xenylla maritima is a well-known species, reported from all European countries

and, with varying certainty, elsewhere. The material agrees completely with the

detailed chaetotactical description by Da Gama (1969).

Ceratophysella succinea (Gisin, 1949) (bona species?)

Material: sample 23: 2 juv.; 24: 14 ; 26: 3 Q and 9 juv.; 32: 1 9 and | juv.;

37:1 9;41:2g and 7 juv.; 44: 2 juv.; 48:2 9,44, and 8 juv. + 15 ex. in alcohol;

49:49,1%,and 1 juv.

Discussion. The material is easily placed in species group A,, the denticulata

group, of Bourgeois & Cassagnau (1972): on abd, are present: a, and a,, m, and m,,

and p,_,, where p, and p, are macrochaetae and p, is ss. Within this group, the

material could be attributed to C. engadinensis (Gisin, 1949), characterized by

having 7+7 setae on the dentes and normal anal spines; or alternatively to C.

succinea, which was differentiated on the basis of its having only 6+6 setae on the

dentes and strong, yellow anal spines.

Since Cassagnau and his collaborators’ work on ecomorphosis and epitocal

processes in Hypogastruridae has shown that characters derived from integu-

mentary details other than chaetotaxy are unstable and of limited use, the last-

mentioned characters must be approached with caution. However, the number of

dental setae was not more informative in the present material. In fact, all grada-

tions between 5 +5 and 7+7 setae where encountered: 5+5:1x;5+?1x;5+6:2

x;54+7:1x;6+6: 12x ;64+2?:2x;7+6: 10x ;7+7: 12x;7+?: 4x. It was not pos-

sible to correlate the number of dental setae with the size of the specimens or their

sexual maturity. For 41 observations, I obtained a correlation coefficient of r =

0.255; for a 5% significance, a value of 0.321 would be required. In sample 26, no

specimen with more than 6 setae per dens were found, but in the other, larger

samples all values occurred.

Evidently, some doubt concerning the separation of succinea and engadinensis

is warranted. However, in this genus, perhaps more than in any other collembolan

group, the results of rearing experiments must be decisive and, basing myself on

material from a different faunal region, I do not dare to synonymize two species

described after Swiss material. I prefer to identify my Greek material with suc-

cinea, because the specimens generally have honey-coloured anal spines, which

are indeed rather long, and moreover because succinea has already been mention-

ed from Greece (Evvia) by Bourgeois & Cassagnau (1972).

W. N. ELLIS: Collembola from Central Crete 245

Fig. 10. Hypogastrura cf. gisini Strenzke. dorsal thoracic chaetotaxy

Seta a’, on abd, (i.e., the seta differentiating C. denticulata from engadinensis),

which Bourgeois & Cassagnau found in 33% of their adult spécimens of succinea

from Evvia, only occurred sporadically in my material; it was found in two im-

matures from sample 26, and in a female from 49, each time on one side only.

C. succinea has been recorded from Spitsbergen, Jan Mayen, Germany, Swit-

zerland, Austria, the USSR, Bulgaria, and Yugoslavia. There are records of

engadinensis from Switzerland, Austria, Poland, France, and Spain.

Ceratophysella gibbosa (Bagnall, 1940)

Hypogastrura occidentalis Gisin, 1958.

Material: sample 32: 1 © ; 34: 1 juv.; 41: 8 juv.; 42: 1 juv.

This species, described from the British Isles, is, according to Yosii (19662)

“almost cosmopolitan”. H. occidentalis was described from Madeira.

Ceratophysella armata (Nicolet, 1841)

Material: sample 35: 4 juv.; 36: 1 juv.; 43: 5 juv.

The species has been recorded from most European countries. It is difficult

to assess the distribution of the species, because the classical species armata

has been subdivided during recent decades.

246

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

II VN per

e sc 1 \ È SIR FE

6 ia % ER iD

A ial N alia ap

hat | iin tye N

4 V A LE. | ihe

2 \ | | ee

eee arsenals medie et À

/ 4 È |, | 5 \f

4 4 [APT ra]

Fig. 11. Hypogastrura tethyca n. sp. dorsal chaetotaxy of thorax and abd, ,

W. N. ELLIS: Collembola from Central Crete 247

Hypogastrura cf. gisini Strenzke, 1955 (Fig. 9 c-k, 10, 12 a-e)

Material: sample 3: 1 9; 10: 10 9, 2g, and 1 juv.; 11:1g;13:1g and | juv.;

Se audea 3232.12.95; 43 and 3 juv.; 26: 2 9; 27: 1g ; 32:49 and lg:

333819 „Sc, andıl juv.; 36:5 ©.

Description. Total length 0.9-1.1 mm. Sparse greyish-blue pigment, scattered

lightly, even eye patches not completely obscured. Integument moderately and

rather evenly granulate. Posterior face of dens not much more strongly granulated.

Anal spines short (about 6-7 u, or %-% unguis P,), curved, on papillae which are

as high and only free at the base (Fig. 12a). Eyes 8+8.

Antennae about as long as diameter of head, with retractile apical papilla, 1

inner and 3 outer slightly swollen and curved sense hairs. Antennal organ III

normal. Postantennal organ small, about two times as large as next ocellus, with

4 (occasionally 3) short and wide lobes, about as large as central part, broadly

attached. Accessory tubercle often invisible (Fig. 9h-k).

Tibiotarsi with I, 1, 1, weakly clavate tenent hairs. Unguis with small inner

tooth; no lateral teeth visible; unguiculus about half as long as unguis, with a very

weak basal lamella, tapering apically (Fig. 9c). Ventral tube with 4+4 setae;

retinaculum with 3 +3 teeth.

Dens with 7+7 setae, posterior face not strongly granulated, about 2.5-3 times

mucro (Fig. 9f). Mucro of assimilis-type, with globular apical tooth and a rather

high outer lamella (Fig. 9d, g).

Chaetotaxy. All hairs comparatively long, smooth. No differentiation into

micro- and macrochaetae. Chaetotaxy of head complete (row p, c, d,_,, sd ,_«, ap,

oc Va). \horacic chaetotaxy: th,: 3+3 setae; th,: a,_,, m,_,, P,_;. p,—ss. Th, is

as th,, except that m, and m, are missing (Fig. 10). Subcoxae 1, 2, 3.

Abdomen,_, with two rows. Abd, with complete a and p rows, and ss in p,. The

m row is represented by m,, m,, and m,. However, irregularities occur rather

frequently, especially in m,, but even the p row may be affected, Abd, with a,_,

and p,_, (p,;=ss), and also an m row normally composed of 3+3 setae. Here as

well, irregularities occur rather frequently, and affect especially the presence of m,

(Fig. 12b-d).

Ventral chaetotaxy of head complete (a,, m,, m,, p,). Th,_, without setae.

Discussion. The Greek material comes reasonably close to gisini Strenzke,

which has been described from sandy beaches along the German North Sea and

Baltic coasts. There is only one striking difference: gisini has an unguiculus

¥%-Y% the length of the unguis, whereas in the Cretan material it is much longer.

It is unfortunate that the chaetotaxy of gisini is not known; I have not been able

to examine type material.

The material also shows a striking resemblance to H. capitata Cassagnau &

Delamare Deboutteville, 1955, from the Lebanon. This species differs in pos-

sessing a strong tooth at the unguis, an unguiculus with a distinct lamella, a dens

with 6 setae and a proportionally shorter mucro, and feebly clavate body setae.

Since not all of these characters are very decisive, a redescription of capitata is

necessary.

Another species, H. christianseni Yosii, 1960, from the United States, could

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 12. Hypogastrura cf. gisini Strenzke. a, anal spine in lateral view; b, dorsal chaetotaxy of abd,_;; c,

d, dorsal chaetotaxy of abd, of two different specimens. Hypogastrura tethyca n. sp. e, ventral tube in

lateral view; f, tibiotarsus and claw of P,; g, posterior face of dens; h, oblique view of mucrodentes of

another specimen

W. N. ELLIS: Collembola from Central Crete 249

according to its description be conspecific with the present material. Upon my

request, Professor Yosii kindly gave me the type-series of this species. A study of

this material revealed seta m, to be present on th,, contrary to the original descrip-

tion. At the time of that publication, the improbability of the absence of this hair

was of course not yet known. This finding seemed to remove the last point of

difference, but on the same occasion the number of retinacular teeth, stated

incorrectly to be 3+3, proved in reality to be 4+4. This, as Yosii remarks, brings

christianseni into close proximity with assimilis Krausbauer, 1898; it remains

distinct because it bears an m row on abd,.

Hypogastrura manubrialis (Tullberg, 1869)

Material: sample 23: 2 9, 3g, and 3 juv.; 31: 1g; 32: 6 2, 4g, and 1 juv.;

41:5 9,23 ‚and 1 juv.; 42:33 (47:19.

Discussion. H. manubrialis is easily distinguished from the foregoing species by

the larger number of antennal sense hairs, the different structure of the PAO

(the lateral tubercles are located above their insertion on the central tubercle, and

thus seem mutually unconnected), the retinaculum with 4+4 (in a few instances

3 +4) teeth, the slender acute mucro, the coarsely papillate posterior face of dens,

and the small almost straight anal spines. Bourgeois & Cassagnau’s (1972)

observation that m, is missing in th, is confirmed.

The species has been recorded from most European countries, and also from

most continents.

Hypogastrura tethyca n. sp. (Fig. 11, 12 e-h, 13)

Material: sample 10: 1 9; 30: 3 9, 4g, and 1 juv.; 31: 10 9 and 84 ; 32: 2 9;

37: 14 and 1 juv.; 43: 1 9 andlg;44:5 9, 126, and 6 juv.; 45: 1g and I juv.

Holotype is a 4 from sample 44.

Description. Length of largest specimens 1.9-2.0 mm. Bluish-grey pigment

forms irregular mottling, and an irregular panther-pattern all over the body, but

less pronounced on ventral parts and extremities. Dens in particular almost

without pigment.

Integument finely and regularly granulate.

Antennae about 0.9 times head diagonal. Ant, with retractile, entire papilla,

3 outer and | inner slender, strongly curved sense hairs, ventrally about 20 short

stiff apically truncate hairs form a kind of sensory rasp. Ant, with 2 short curved

sensillae and 2 long and rather slender sense hairs (Fig. 13 e).

Eyes 8 +8 on intensely pigmented eye patch. Postantennal organ distinctly wider

than nearest ocellus, consisting of 4 bladders, the anterior two the longest.

Accessory boss present (Fig. 13f, g). Rim of labrum with 4 flat protuberances

(Fig. 13 d); labral setae 4/5, 5, 4.

Tibiotarsi with 1, 1, 1 long, feebly but distinctly clavate tenent hairs. Unguis

with distinct internal tooth, lateral teeth absent or extremely small. Unguiculus

with large basal lamella and a filament about 2/3 internal length of unguis

(Fig. 12 f).

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 13. Hypogastrura tethyca n. sp. a, b, c, dorsal chaetotaxy of abd, of three specimens; d, labral

margin seen from ventral; e, antennal organ III; f, g, postantennal organs of two specimens; h, anal

spines in oblique view

W. N. ELLIS: Collembola from Central Crete 251

Ventral tube with 4+4 setae (Fig. 12 e). Retinaculum with 4+4 teeth, rarely

3+4.

Dens 100 u, mucro about 36 u. Dens with 7 setae. Posterior face of dens finely

granulated. Mucro elongate, with only very weak inner and outer lamellae

(Fig. 12 g, h).

Anal spines well developed, slightly curved, ca. 19 u, on (almost touching) anal

papillae of 22 u (Fig. 13 h).

Chaetotaxy composed of strong, comparatively stiff, at their anterior face

rugose to slightly serrate, setae. The setae are all alike, except that in posterior

direction the dorsal setae become stronger. Setae sensuales easily recognized by

their length and structure.

Cephalic chaetotaxy with complete rows p and c, v=2, d and sd = 5, oc = 3,

a, present. Th, 3+3 setae. Th, complete, p, = ss; th, as th, but m, and m, missing

(Fig. 11). Abd,_, with two rows of setae. Abd, difficult to interpret because of

many irregularities (Fig. 13 a-c). The p row is always complete. Between setae p,

(ss) the a row normally shows only 3+3 setae, but not infrequently 3+4 or 4+4;

the m row consists of 3+3 or 2+2 setae. Abd, with 2+2 setae in the a row be-

tween p,-p,, and never with an m row (Fig. 11).

Ventral chaetotaxy of head with a,, m,_,, p,. Thorax ventrally without setae.

Diagnosis. I find it very difficult to give this species a place in the genus. Using

the key given by Gisin (1960) one arrives blindly at Hypogastrura sahlbergi (Reuter,

1895) or H. capitata Cassagnau & Delamare, 1955. The new species differs from

the former by the different shape of the anal spines and their papillae, by the

structure of the setae — short, slender, and pointed in sahlbergi, long stiff bristles

in tethyca — and above all by having not 7-8 setae at each side of the ventral tube,

but the normal number 4+4. H. capitata, seemingly more closely related to

H. gisini, has 3+3 teeth on the retinaculum, clavate body setae, and a different

mucro.

The strong basal lamella of the unguiculus is reminiscent of such species as

tullbergi, viatica, etc., but these have more than 1 clavate tenent hair on the

tibiotarsi. H. monticola Stach, 1946, has the single tenent hair not clavate, very

short anal spines, and a “‘spoon-like’’ mucro. H. packardi (Folsom, 1902) has

capitate body setae, an extremely heavy clavate tenent hair, and, even in the

forma dentata, too short an unguiculus; according to Bourgeois & Cassagnau,

the chaetotaxy is differentiated into micro- and macrochaetae. H. aterrima Yosii,

1972, from Japan, has 3+3 teeth on the retinaculum and a strong differentiation

into micro- and macrochaetae. H. paradoxa Yosii, 1965, also from Japan, seems in

some respects close to tethyca: 1 clavate tenent hair on tibiotarsus, same shape

of anal spines and claw, etc.; however, that species forms a link with Ceratophysella

in having a small exsertile sac between ant, and ant, as well as a peculiar chae-

totaxy, an m row being fully present on abd, and abd,. In H. theeli (Tullberg,

1876) s. Yosii, 1972 = H. trybomi (Schott, 1893) s. Hammer, 1953, the tenent hair

is not clavate. H. macrotuberculata Hammer, 1953, has a highly divergent mucro

with a high lamella.

The species is dedicated to Tethys, goddess of the sea, who embodies at the

252) TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 14. Odontella nana orientalis n. ssp. a, eye patch and postantennal organ; b, ventral chaetotaxy of

abd,_,; c, ant,_,; d, atypical dens; e, typical mucrodentes; f, ventral chaetotaxy of head; g, mouth cone

W. N. ELLIS: Collembola from Central Crete 253

same time the very sea that shaped the present zoogeographical pattern of the

Greek Collembola.

Hypogastrura vernalis (Carl, 1901)

Material: sample 10:49 and 1g ; 17:29 and lg.

Recorded from Scandinavia to the Mediterranean and from the British Isles to

Poland. Along the Mediterranean it has been found from the Iberian peninsula to

Yugoslavia.

NEANURIDAE

Odontella nana orientalis n. ssp. (Fig. 14, 15, 16 a-b)

Material: sample 29: 19 and 7g ; holotype isa.

Description. Mean length of the males is 0.60 mm (s = 0.016, s; = 0.006 mm),

the single female measures 0.89 mm; statistically, this difference is strongly

significant (t=16.353, df=6, P<0.001).

Greyish-blue pigment sparse on dorsal parts of body and head and, even more

so, on feet and furca. Eye patch strongly pigmented. Integument with primary

granules arranged in a tetragonal lattice. The skin is strongly folded in an irregular

way. Habitus normal for the genus, with very short conical antennae.

Mouth-parts normal for the genus (Fig. 14 g), labium with some short and 1 +1

long hairs. Ocelli 5+5, c somewhat smaller than the others. Postantennal organ a

pretty four-pointed star, perfectly symmetrical (Fig. 14 a). Ant, with a sensilla at

outer face. Ant. organ III two short cones, and laterally from them two bent,

swollen claviform clubs. Ant, with 8 sense hairs and an apical cupola (Fig. 14 c).

Claw slender, with strong basal tooth on inner lamella, and a pair of teeth

basally on lateral lamellae. No unguiculus. No clavate tenent hair (Fig. 16 b).

Ventral tube with 3+3 setae. Retinaculum tridentate. Dentes with 5+5 setae

(Fig. 14 e) but one specimen had 4+5 (Fig. 14 d). Mucro typical. Male genital

tubercle small, with 10 setae. Female genital cleft guarded by no more than 3

setae. Ventral flaps of anal segment with some remarkably strong setae. Anal

spines very short and conical, on very low papillae. The anal spines are often

present on one side only, and in one instance one was cleft. The presence of the

spines is evidently an unstable character, although their shape is very constant.

Chaetotaxy (Fig. 15), composed of fine, rather short, smooth setae which are

never clubbed. Cephalic chaetotaxy rather reduced: present are p,_,, C,, (p, may

sometimes seem to be c, as well), d,, d,_,, sd,_,, OC,_;; a, is absent. Th, with the

peculiar number of 4+4 setae. Th,_, identical, with a,, a,_, (other setae of a row

difficult to interpret, see Fig. 15), m,, m,, p,_,, p,=ss, p.. Cassagnau (1974) gives a

somewhat different interpretation of the thorax of O. nana Cassagnau, 1954.

According to this author, a, is present in th,, but absent in th,. Some individuals on

a slide with specimens of nana, preserved in the collection of the Laboratoire

d’Ecologie générale, Brunoy, unfortunately without indication of locality, showed

a much more median position of the hair on th, labelled by me as a,, i.e., almost

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

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x 4 RER TOR aE ON

Seend bb bar ss | it

Lg er

A ess

x 6 TS

” Pr ERS DES EN

SA ; fee

Fig. 15. Odontella nana orientalis n. ssp. dorsal chaetotaxy

W. N. ELLIS: Collembola from Central Crete 259

anterior to p,. Otherwise, they seemed identical to the Greek specimens in

chaetotaxy.

Abd ,_, alike, with two rows, consisting of a,, a, ,, Pı_» Pa_s; Ps = SS. Abd, as

abd,_, but with the insertion of an m row with m, and m,. Abd, with a,, a,, p, ;;

Pass.

Ventral chaetotaxy of head with a, ,, m,_,, no posterior setae (Fig. 14 f).

Thoracic sternites without setae. Abd, with 1 +1 setae near the ventral tube. Abd,

with 2+2 setae in the posterior, and 1+1 in the anterior row. Abd, and the

manubrium with a complicated array of setae (Fig. 16 b).

Discussion. Judging by the shape of the postantennal organ, the comparatively

large number of sense hairs on ant,, and the presence of anal spines, the Cretan

material belongs in the group of O. stachi Denis, 1947, described after one

specimen from Burgundy, O. nana Cassagnau, 1954, described after a single

specimen from the Pyrenees, and O. vallvidrerensis Selga, 1966, proposed for a

richer material from NE Spain. It differs from vallvidrerensis in having 5 + 5 instead

of 4+4 setae on the dentes, and from stachi in having a distinct lateral tooth on the

unguis. It differs from all three by the very low, blunt anal spines, and above all the

absence of the unguiculus. The latter character relates the present material to O.

sublamellifera Denis, 1948, from Vietnam, but that species lacks the lateral teeth

on the unguis and has a somewhat different, smaller, postantennal organ. In all

other body characters it agrees closely with O. nana, and I prefer not to exaggerate

the importance of the differences by making another new species.

If I may take the slide from Brunoy as a reference for nana s. str., another

difference would be the absence of seta m, on abd, in nana nana.

Xenyllodes minitaurus n. sp. (Fig. 16 c-e, 17, 18, 19 a-d)

Material: sample 30: 1g ; 36:19 and 13 (holotype).

Description. Total length of the only female 0.7 mm; the males are 0.5 and 0.6

mm. Habitus slender, strongly resembling that of a Mesaphorura. Basal skin

reticulation in a normal, hexagonal lattice (very different from the quadrangular

lattice in Odontella); skin folded in high conical papillae, which are absent only on

the feet and furca. No pigment, but below each eye there is a small aggregation of

colourless granules.

Ant, with small sensilla in external face. Ant. org. III with two small curved

sensillae, and between them two almost imperceptible sense rods (Fig. 18 d). Ant,

with retractile end bulb, and 6 swollen sense hairs. Hairs at ventral face mainly

with a blunt apex (Fig. 16 d).

Eyes 2+2, separated by roughly their own diameter. Postantennal organ broadly

and a bit irregularly triangular, sunken below the integument in a cavity

communicating with the surface via a triangular fissure (Fig. 19 a, b). Mouth parts

not studied. Mouth cone typical (Fig. 18 c). Tibiotarsi with some long hairs, which

are possibly finely clavate (Fig. 19 d). Unguis strongly curved, without lateral or

internal teeth. Unguiculus present, needle-shaped (Fig. 19 c).

Ventral tube with 3 + 3 setae.

Retinaculum bidentate. Manubrium with 2+2 setae; dentes almost quadrangu-

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

256

al areas; b, claw of P..

e, anal spines

& ©

SUE

X 5

S &

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nee

a 8

= 8

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= &

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W. N. ELLIS: Collembola from Central Crete 257

lar, smooth, with 2+2 setae. Mucro exceedingly long and crook-shaped (Fig. 16 c).

Male genital papilla with about 11 setae; female genital orifice with 3+3 setae

(Fig. 18 a). Two anal spines present, strong and curved, roughly 1.5 times unguis

(Fig. 16 e).

Chaetotaxy composed of fine, smooth setae, nowhere differentiated (Fig. 17).

Head lacking seta a,, with setae p,_,, C,, C3_4 d,, d,_,, Sd,_,, oc,_,. As in Odontella,

th, has 4 +4 setae. Chaetotaxy of th, and th, similar: a ,_,, m,, m,, p,_,, P3_4- It is not

possible to recognize a seta sensualis in p,. Abd,_,with a,; a,_, p,_,. Abd, lacks a

da,

5 n

ta =

di a Sl ros 74 Azar EK

ari IN SNA

din lives ssi £ Ts =

Se 8 A \ =) 4 =, ;

2 En ASI rp

A Ve

Ri Shp Westen oF poe pus ' ee

ve BN ee Sane See Jl | | Li

A: A \ \ À NR N LE he

a cr ae RED cai = ZE N a ne!

Leg }

= TA A Va e ri

S 5

i SE la elas x

ort a = le / —

A Be e UE

é VA NEA Di ha ES

peri er m È pee o fo

Bo > |

ern E ne

pi; £3 DAT ee:

) ris ee a

dl eee

MZ row lk: REL:

ein ZN À Mie. =

be LENS SA Re

Ri 3 X en

Fig. 17. Xenyllodes minitaurus n. sp, dorsal chaetotaxy

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

258

illa nn

MN 10)

he” ama na nee O CAPA ala [

lé

Ji e SC

= >; 7 Cai

| N —

nn AGODA

a | da

Poca

v > 2)

UO, got Sr,

“poe a VUL uvY UV ES, x N FAN \ aby,

SABOT |

4} a,

COS /

(0)

Fig. 18. Xenyllodes minitaurus n. sp. a, female genital orifice; b, antennal organ III; c, mouth cone; d,

ventral chaetotaxy of abd,_,; e, ventral chaetotaxy of head

W. N. ELLIS: Collembola from Central Crete 259

row m, and only has a,, a,_,, P,_,, p, . Abd, with a,_,, p,_,. Sete sensualis is, as

usual, p, in abd,_, and p; in abd,. Abd, with one unpaired seta.

Ventral chaetotaxy of head with a,, m,, and p,, but apparently lacking m, (Fig.

18 e). Abd, with 1 +1 seta beside the ventral tube. Abd, with 1 anterior seta, and

2+3 (2 specimens) or 3+3 (1 spec.) setae in posterior row. For the chaetotaxy of

abd,_,, I refer to Fig. 18 d.

Discussion. Presently, only two species of Xenyllodes having 2+2 eyes are

known, viz., bayeri Kseneman, 1935, from Central Europe, and X. ghilarovi

Martynova, 1964, very briefly described from the USSR (Kursk). The new species

differs from bayeri by being much more slender, the large, triangular, sunken

postantennal organ, the dentes not being tuberculate, and the subempodial setae

being much longer. X. ghilarovi is differentiated from bayeri rather concisely:

“Postantennal organ large, triangular. Body without pigment’. The only accom-

panying drawing shows two eyes set fairly close to each other (separated by less

than half eye diameter) and the postantennal organ superficial.

The name was chosen not only because part of the material was collected in

Knossos, the palace of the Minotauros, but also because I was reminded of this

legendary being by the shape of the mucro, which has some resemblance to the

emblem of the holy bull.

Brachystomella parvula (Schaffer, 1896)

Material: sample 16: 29 and 1g ; 17: 1 juv.; 27:79, 4g, and 2 juv.; 36: 49; 41:

40 and I juv.; 42: 19; 43: I juv.; 45: 3 juv.

This material will be dealt with by Dr. Z. Massoud and Dr. J. Najt, who are pre-

paring a revision of this species.

Friesea afurcata Denis, 1926

Introductory remarks. The Cretan material contained a fine series of this spe-

cies. It was, however, by no means homogeneous material and at a first glance

seemed to comprise three species, but since the differences were not considered

sufficiently important, I preferred to treat the variants as three types of the same

species. They are possibly to be explained by differences in post-adult develop-

ment; I found no indications for an explanation in terms of ecomorphosis.

It is interesting to compare this variability with the results of Grow & Christian-

sen (1974), which unfortunately did not become available to me until after the

completion of the present study.

Friesea afurcata Denis, 1926, type I (Fig. 19 e-g, 20)

Material: sample 41: 119 and 35 ; 49: 19; 19 from sample 41 was partially, and

29 and 23 were wholly depigmented by treatment with HCIKCIO,. One of the

depigmented 9 was squashed for a study of the mouth parts.

Description. Mean total length of 109 from sample 41 is 1.63 mm (s=0.14 mm,

s; = 0.04 mm); mean length of 3g is 1.31 mm (s=0.28 mm, s; = 0.16 mm). Dif-

ference in size between the sexes significant (t=2.853, df=11, 0.02>P>0.01). The

single Q from sample 39 measures 1.2 mm. Irregular bluish-grey pigmentation dis-

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 19. Xenyllodes minitaurus n. sp. a, eyes and postantennal organ in perpendicular view; b, the same,

oblique view; c, claw of P,; d, tibiotarsus of P,. Friesea afurcata Denis type I. e, tibiotarsus of P,; f,

dorsal chaetotaxy of abd,_,; g, dorsal chaetotaxy of head and thorax. (Encircled hair bases refer to

setae missing in the specimen from which the drawing was made.)

W. N. ELLIS: Collembola from Central Crete 261

persed all over the body, less densely in ventral region. Eye patches dark. Integu-

mentary granulation rather fine and regular, in abd, a bit coarser.

Ant, with retractile apical papilla, distinct strongly curved swollen sense hairs,

and a number of setiform sense hairs. Ventro-distally there is a group of about

eight short blunt sense hairs. Suture between ant, and ant, only visible ventrally

(Fig. 20 a). Ant, with antennal organ normal; two small, parallel, rectangularly cur-

ved sense clubs in a shallow groove, guarded by two thickened sense hairs ca. 7—8

u long (Fig. 20 c). Eyes 8+8. Maxilla typical (Fig. 20 f); mandible with 3 strong ba-

sal teeth, 3 smaller distal teeth with a small fourth one, and a tooth associated with

the corpus of the mandible (Fig. 20 g).

Unguis with a tooth in the middle of the internal lamella, which is, however, so-

metimes indistinct or missing; lateral carinae without teeth. Unguiculus absent.

Two inner and two outer tenent hairs with a barely perceptible distal dilatation

(Fig. 19 e).

Ventral tube with 4+4 setae; retinaculum and furca completely absent. Male

genital papilla with about 25 setae, female genital orifice with about 15 associated

hairs. Anal spines 4, strong, almost straight, arranged in a square, totally

differentiated from normal hairs (Fig. 20 b).

Dorsal chaetotaxy (Fig. 19 f, g) composed of coarse, long, distinctly rugose setae

(ventral chaetotaxy composed of smooth smaller hairs). Cephalic chaetotaxy with

setae p,_4, C;_4 29 d,_, (d, is unpaired), sd,_,, oc,_,. No setae v. Th, with 4+4 setae.

iether esa, m, m, Pp, >, Pae) Pa = SS. Th, differs from th, only but

constantly by lacking a,. Subcoxae with 1, 2, 2 setae. Abd,_, with a,, a,_5, P1_s;

p, = ss. Chaetotaxy of abd, is rather irregular, especially in the a row. It seems as

though normally a,_, are present. In the posterior row the normal situation is: p,_,,

P4_s; Ps = ss. In abd, a,_, are present, as well as p,_,. Very often not p, (the normal

case) but p, = ss; in the present material this only happens asymmetrically, but

this might explain the situation illustrated by Da Gama (1964) for F. afurcata,

where p, is ss at both sides of abd,.

Ventral chaetotaxy of head with only a, and m, (Fig. 20 e); thoracic sternites and

abd, not chaetose. Abdominal sternites 2-4 with a large number of setae which are

difficult to homologize (Fig. 20 d).

Diagnosis. On the basis of the keys of Gisin (1960) and Massoud (1967) one

would identify the present species as F. afurcata Denis, 1926. This species was

described from Italy, and was later reported from Germany and Austria. The

species was recorded from the Lebanon by Cassagnau & Delamare Deboutteville

(1955). The types have been studied in as much detail as was still possible by Da

Gama (1964) in connection with a redescription of F. ladeiroi Da Gama, 1959.

Quite recently, Dallai (1973), in connection with the description of the related F.

lagrecai, provided additional details after topotypical material of afurcata. From all

this it can be inferred that the Cretan material is, to say the least, very close to

afurcata; possibly important differences are in the tenent hairs — in the Greek

specimens much less distinctly clavate than as drawn by Dallai — and in the

chaetotaxy of abd,. Although some ambiguity exists, partly due to different

interpretations of the chaetotaxy, Cretan afurcata seem to have one more seta in

the prow.

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 20. Friesea afurcata Denis, type I. a, ant,_,; b, lateral aspect of abd,; c, antennal organ III; d,

ventral chaetotaxy of abd,_,; e, ventral chaetotaxy of head; f, mandible; g, maxilla

W. N. ELLIS: Collembola from Central Crete 263

Another species which seems very close to the present material is Colonavis

grandis Salmon, 1949 (= Friesea salmoni Massoud, 1967, nom. nov. pro F. grandis

(Salmon, 1949) nec Mills, 1934) from Campbell Island.

Friesea afurcata Denis, 1926, type II (Fig. 21,22 a)

Material: sample 16: 19; 26: 19 and 23 ; 36: 159, 103, and 1 specimen of

unknown sex - squashed; 43: 19.

Discussion. This material differs from what I have provisionally called „type I”

in the following characters:

(a) in the chaetotaxy of th, and th, the m row has no setae — normally! In at

least one specimen th, has a distinct m,, in a symmetrical position (Fig. 21 d);

(b) in abd,, p, = ss (Fig. 21 a);

from normal setae;

(d) the abdominal setae are more slender, shorter, and much less rugose; often it

is hard to find any rugosity;

(e) the integument of abd, is distinctly more coarsely granulated than that of the

rest of the body; this is very striking, and visible at rather low magnifications;

(f) the ventral chaetotaxy of abd, and abd, is much less dense (Fig. 22 a);

(g) lateral sensillae of ant. organ III longer compared to body of ant, and to

length of guarding setae (Fig. 21 b);

(h) the specimens are much smaller; the mean total length of the females from

sample 36 is 0.71 mm (s = 0.05 mm, s; = 0.02 mm), of the males 0.62 mm (s = 0.06

mm, s;=0.02 mm); the difference in size between the sexes is significant

(t = 3.481, df = 23, 0.01<P<0.001); the specimens from the smaller samples lie in

the same range.

The last character could be interpreted as an indication of immaturity of the

material, but most of the males show active internal genitalia and testicular tissue.

However, the otherwise close similarity to afurcata (number of anal spines, total

reduction of retinaculum and furca, number of eyes, etc.) is such that I prefer

provisionally to put more emphasis on the relationship between the forms around

afurcata rather than to separate them by creating different species. Moreover,

some of the characters may be related to allometric growth (structure of setae,

anal spines, chaetotaxy of abdominal sternites 1 and 2), and I cannot exclude the

possibility that material of type I is somewhat older than that of type II. It may be

significant that Gisin (1960) mentioned for afurcata the unusual wide length

interval of 0.8-1.6 mm.

Friesea afurcata Denis, 1926, type III (Fig. 22 b, c)

Material: sample 2: 13 ; 40:49 and 4g .

Discussion. This material comes closest to “type II”. The mean size for the 9 is

0.7 mm, for the g 0.6 mm. This sex difference is not significant. The sensillae

lateral to antennal organ III are long, even somewhat undulate. The tibiotarsal

tenent hairs are even more weakly differentiated. The integument of abd, is

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 21. Friesea afurcata Denis, type II. a, dorsal chaetotaxy of abd,_,; b, antennal organ III; c,

mandible; d, dorsal chaetotaxy of head, thorax, and abd,: an x indicates a hair missing in the actual

specimen, an encircled seta is lacking in the remainder of the specimens

W. N. ELLIS: Collembola from Central Crete 265

differentiated in the same way, i.e., coarsely granulated. However, I am forced to

place it in a type of its own — if not a species — since the setae a, on abd, are

spiniform, and just as strongly as a, or p,. This brings the present material close to

F. ladeiroi Da Gama, 1959, described from Madeira, and differentiated from F.

afurcata by having six anal spines (shaped just like those of my afurcata III), the

setae almost completely smooth, and only one tenent hair which is not clavate. In

“he LORS

Fig. 22. Friesea afurcata Denis, type II. a, ventral chaetotaxy of abdomen. Friesea afurcata Denis, type

III. b, dorsal chaetotaxy of abd,_,; c, dorsal chaetotaxy of thorax

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

1964, Da Gama added to these characters: absence of setae a, in abd, and perhaps

the absence of a, in th,_,. F. afurcata III remains distinct from ladeiroi in possessing

a, on abd,, and moreover in having a,, but missing m,, in th,_, (I admit that if

inserted a bit more caudad, the seta which is present might just as well be called

m,; but a, and m, are not present at the same time anyway).

Friesea decipiens Steiner, 1958 (Fig. 23 a, b)

Material: sample 25:59 and 32.

Discussion. This species, described from Spain has recently been found by

Dallai (1969 b, 1973) and Rusek (1973 b) in Italy. The species seems to have a

Mediterranean distribution. Supplementary to the description by Steiner and the

drawings by Rusek, I give some descriptive notes.

Mean length of the females 0.7 mm (s = 0.07 mm, s; = 0.03 mm); mean length

of the males 0.6 mm (s = 0.02 mm, s; = 0.01 mm). Cephalic chaetotaxy as

illustrated for F. afurcata. Thoracic chaetotaxy differs by the absence on th,_, of

m,; at the same time, a, is displaced somewhat caudad, being inserted almost at

the place of m,. Abd,_, with a,, a,_,, Pı_» P4_s; Pa = SS. Abd, with 3 +3 setae in the

a row before the 4+4 in the p row. Abd, lacks a, and p, is ss. Seta p, rather long

and stiff; seta a, and a, distinctly longer than drawn by Rusek. The median anal

spine is usually smaller than the lateral ones, and mostly but not always, almost

straight (Fig. 23 b). Ventral chaetotaxy of head with a, and m,; thorax and abd,

ventrally achaetose. Ventral chaetotaxy of abd, , rather imperfectly symmetrical

(Fig. 23 a). All body setae very fine and smooth, except p, on abd, which is a bit

coarser though still smooth. Subcoxae 1, 2, 2.

The material is in good agreement with the description by Steiner except that

the unguis is provided with a distinct inner tooth.

Tremoisea cf. ossica Cassagnau, 1973 (Fig. 23 c-e)

Material: sample 45: 13 .

Discussion. It was a pleasant surprise to find a representative of the genus

Tremoisea, described in 1973 by Cassagnau for a species from Kerkira and another

from the Greek mainland, not far from Larissa. The single specimen, measuring

1.5 mm, had its intestine filled with some dark-blue (clay?) material, showing up an

unexpected array ‘of what seem to me to be intestinal diverticula (Fig. 23 e).

However interesting these may be, they impeded closer study of the specimen. But

all essential traits necessary for the generic allocation are distinct: the strange

mouth cone (Fig. 23 d), the multidentate mandible and the elongate Frieseinae-

type maxilla, the complete eye patch, albeit lacking the postantennal organ, the

abundant chaetotaxy, and so on. However, it seems to me that the chaetotaxy of

my single specimen, as far as this could be studied, was not perfectly symmetrical.

From the structure of the unguis (Fig. 23 c) it is evident that in any case the

specimen comes close to 7. ossica, described from Thessalia.

W. N. ELLIS: Collembola from Central Crete 267

Fig. 23. Friesea decipiens Steiner. a, ventral chaetotaxy of abd,_,; b, dorsal chaetotaxy of abd, «.

Tremoisea cf. ossica Cassagnau. c, claw of P,; d, mouth cone; e, habitus of cleared specimen (to show

intestinal diverticula)

Pseudachorutella cf. asigillata (Börner, 1901) (Fig. 24)

Material: sample 21: 1 juv., depigmented by HCI-KCIO, treatment; 35: uv

squashed for observation of mouth parts.

Description. Length 0.6 mm; irregular not very strong greyish-blue pigment on

dorsum and extremities. Habitus normal. Skin granulation moderate. Antenna,

268 TIJDSCHRIFT VOOR ENT JMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 24. Pseudachorutella cf. asigillata (Borner). a, claw of P,; b, posterior face of mucrodens; c, mucro;

d, dorsal chaetotaxy of head and thorax; e, ant,_,; f, dorsal chaetotaxy of abd, ,

W. N. ELLIS: Collembola from Central Crete 269

with apical retractile papilla superficially trilobed, and with 8 rather slender,

strongly curved sense hairs; no sensory rasp. Ant, with 2 small sensillae (I could

not make out whether they are straight or curved) guarded by two comparatively

slender sense hairs; a small sensilla in a groove ventrally in ant, (Fig. 24 e). Eyes

8 + 8 in a well pigmented eye spot. Postantennal organ absent. Mouth cone long;

mandible in one specimen three-toothed; maxilla with two lamellae, one with a

small distal hook, at least the last third free. No differentiated tenent hairs.

Unguiculus absent, unguis rather slender, with a small inner tooth, but without

lateral teeth (Fig. 24 a).

Ventral tube with 4+4 setae. Retinaculum tridentate. Dens with 7+7 setae,

mucro elongate, straight, with a narrow inner lamella gradually narrowing towards

apex (Fig. 24 b, c). No anal spines.

Chaetotaxy composed of short, smooth, nowhere clavate setae; setae sensuales

well differentiated. Cephalic chaetotaxy with p and c row difficult to separate, d,_,

(d, unpaired), sd,_,; a, absent; oc,_, (Fig. 24d). Th, with 3 + 3 setae. Th, with a, „

ESD Pas; Py = ss. Th, similar but lacking a, Abd,_, with a, a,_,, Pisi

Be — 58,

Abd, with a,, a,, m,, Pı_» Pa_s; Ps = SS. Abd, with a,_,; p,_,; pz = ss (Fig. 24 f).

Discussion. This material, which is very similar to that which I recorded earlier

(Ellis, 1974) from Rhodes, and also shares the inconvenience of being very young,

is a bit difficult to interpret. It differs in one chaetotactic detail from asigillata as

drawn by Da Gama (1964) viz., in the presence of a seta p, in abd,. Moreover, the

shape of the straight mucro, with its narrow, apically unnotched lamella, has little

resemblance to that of asigillata. Finally, the observation of a three-toothed

mandible is suggestive — but nothing is known about possible variability in this

structure. I am awaiting adult material before making a decision.

Pseudachorutes dubius Krausbauer, 1898

Material: sample 27: 1 juv.; 35: 1g .

The species has already been recorded from Greece (Evvia, Cassagnau, 1971;

Rhodes, Ellis, 1974).

Pseudachorutes libanensis (Cassagnau & Delamare Deboutteville, 1955)

n. comb. (Fig. 25, 26 a-c)

Aethiopella libanensis Cassagnau & Delamare Deboutteville, 1955.

Material: sample 17: 1 juv.; 32:49 and 34 ; 48:29.

Description. Mean length of the six females 0.8 mm (s = 0.16 mm, s; = 0.07

mm), that of the 3 males 0.7 mm (s = 0.10 mm, s; = 0.06 mm).

Skin granulation normal for the genus, greyish-blue pigment, not particularly

dark, scattered over dorsal parts of body.

Ant, with apical bulb “apple-like”: almost globular, but with a slight apical

invagination (Fig. 25 h). Outer side of this antennomere with 2, inner side with 6

moderately thick curved sense hairs. Antennal organ III consists of two small,

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

270

W. N. ELLIS: Collembola from Central Crete 271

Fig. 26. Pseudachorutes libanensis (Cassagnau & Delamare). a, antennal organ III; b, dorsal chaetotaxy

of abd, _,; c, ventral chaetotaxy of abd,_,. Pseudachorutes (Pratanurida) mucronata n. sp. d, ant, 4

Fig. 25. Pseudachorutes libanensis (Cassagnau & Delamare). a, maxilla; b, mandible; c, mouth cone; d,

postantennal organ; e, dorsal chaetotaxy of head and thorax; f, furca; g, anterior face of mucrodens; h,

la apical papilla of antenna

272 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

straight (or almost so) sensillae, guarded by two long, slightly wavy sense hairs

(Fig. 26 a). Ocelli 8 + 8. Postantennal organ with about 8 elements (Fig. 25 d).

Mouth cone as long as typical for the genus (Fig. 25 c). Mouth parts studied only

in transparency, normal for the genus with a needle-shaped maxilla (Fig. 25 a) and

a two-toothed mandible (Fig. 25 b).

Tibiotarsus without differentiated tenent hairs. Unguis without teeth, or (seen

once) with a very feeble tooth on inner lamella. Unguiculus absent.

Ventral tube with 4+4 setae; retinaculum tridentate. Dens with 6+6 setae (Fig.

25 f, g). Mucro crooked, and with inner lamella more developed than outer one.

Dens with characteristic elongate area on anterior surface where the skin

reticulation is missing.

Chaetotaxy composed of very short, smooth, never capitate setae. On the head.

are present p,_, (p, is sometimes missing), c,_,, d,_y, Sd,_,; a, and v are missing. Ind

row no unpaired setae. Th, with 3+3 setae. Th,_, identical, with a,, a, 5, P,_>, P4_s;

p, = ss (Fig. 25 e). In abd,_, are present a,, a,, P,_,, P3, ps. In abd, we have a,, a,, as,

and p,_,, P4_5- Abd, with a,_, and p,, p,; p, = ss (Fig. 26 b). No anal spines.

Discussion. The present material agrees closely with the description of.

Aethiopella libanensis, based on a single specimen from Beirut, except of course in

the structure of the postantennal organ, which is said to be moruliform in the type

specimen. However, even the description leaves some uncertainty as to this point,

and in one of the two aspects of the postantennal organ drawn by the authors, it is

almost completely pseudachorutiform. I am therefore convinced that libanensis

was incorrectly allocated, presumably because it was described after a defective

specimen.

The species comes close to Pseudachorutes parvulus Borner, 1901, P. subcrassus

Tullberg, 1871, P. crassus Da Gama, 1964, P. geronensis (Massoud, 1963), and P.

pratensis Rusek, 1973. P. libanensis differs from the first three species in the

absence of seta a, in th,. and also in the combination of characters: no unpaired

seta in d row on head, and abd, carrying setae a, ,, p,, P;. The main differences

with respect to geronensis are the normal mouth cone (that of geronensis is very

short for the genus), the straight sensillae in the ant. org. III, and the structure of

the mucro. Possibly, the chaetotaxy of abd, is also different in geronensis. P.

libanensis is most closely related to P. pratensis Rusek, 1973. The chaetotaxy of

these two species seems to be identical. Differences concern the larger number of

sense hairs in ant, (4 in pratensis), the shape of the sensillae in the antennal organ

III (short, distally swollen clubs in pratensis, straight rodlets in libanensis), and

especially the form of the mucro, which is normal for the parvulus group in

pratensis but hooked in libanensis. Useful distinctions are perhaps offered by the

slightly larger number of elements in the PAO and the distinct tooth on the unguis

in pratensis.

Pseudachorutes (Pratanurida) mucronata n. sp. (Fig. 26 d, 27, 28 a-c)

Material: sample 17: 1 9 (holotype); 36: 19.

Description. The holotype female measures only 0.4 mm; the other specimen is

0.6 mm. Both specimens have the genital orifice open, but provided with only a

few setae, and it is possible that they are not completely mature. Pigmentation

W. N. ELLIS: Collembola from Central Crete

273

\ = —

x | \

SÒ fe i At

VES

= 4 NON

Fig. 27. Pseudachorutes (Pratanurida) mucronata n. sp. a, dorsal chaetotaxy of abd, ,; b, dorsal

chaetotaxy of head and thorax; c, mandible; d, maxilla; e, mucrodens in lateral aspect; f, ventral

chaetotaxy of mouth region; g, furca and retinaculum

274 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

greyish-blue, but particularly strong. Eye patch darker. Integument finely and

regularly reticulate, as in Pseudachorutes s.s.; no Neanura-type reticulation present.

Habitus as in a normal Pseudachorutes.

Antenna, with retractile apical papilla undivided; interior face with 4, outer face

with 2 curved and thickened sense hairs; no sensory rasp (Fig. 26 d). Antennal

organ III two minute, erect clubs, almost without integumental plica, guarded by

two strong sense hairs; antenna, moreover with a sensilla in a groove (Fig. 28 c).

Eyes 8+8. Postantennal organ with 7-8 elements arranged in a circular rosette

(Fig. 28 a, b). Mouth cone short (Fig. 27 f). Mouth parts (seen in transparency):

mandible with two teeth only (Fig. 27 c), maxilla needle-shaped with two lamellae

(Fig. 27 d).

Unguis without inner or lateral teeth; no unguiculus, no differentiated tenent

hairs.

Ventral tube with 4+4 setae; retinaculum bidentate. Furca present but reduced.

Dentes rather short, almost triangular, with 4+4 setae; mucro present, reduced to

no more than a hook-like continuation of the dens, not visibly separated from the

latter (Fig. 27 e, g). Female genital orifice small, with 5 setae. No anal spines.

Chaetotaxy composed of short, smooth setae. It is almost impossible to

homologize the setae, partly because only two specimens are available, but

especially because the sensory setae are not visibly differentiated. Cephalic

chaetotaxy with rows p and c incomplete; setae v and a, are lacking; d,_, (d, is

unpaired, called d, in the papers of Da Gama et al.), sd,_,. Th, with 3+3 setae. Th,

with 4 setae in the a row and 3 setae in p row in discal area; th, identical except

one seta missing in a row. In th,_,, neither shows a distinct seta p, (Fig. 27 b).

Subcoxae 1, 2, 3.

Discussion. The new species fits easily into the genus Pratanurida, created in

1973 by Rusek for his new P. cassagnaui from Central Europe. That species was

redescribed shortly afterward by Dunger, 1974. Briefly, Pratanurida is Pseuda-

chorutes with the furca reduced. Pseudachorutes mucronatus is easily distinguished

from cassagnaui by its mucronate dens having 4+4 setae, the less coarse

reticulation, the absence of seta a, on the head, the apparently real absence of

setae p, in th,_,, and perhaps also in the higher number of postantennal elements:

7-8 as against 4 (Rusek) or (4) 5-6 (Dunger) in cassagnaui.

For a discussion concerning the position of Pratanurida and Stachorutes Dallai,

1973, intermediate between Pseudachorutes and Micranurida, I refer to Rusek

(1973a) and Dunger (1974). I might only add that a tendency toward reduction of

the dens is not altogether absent in Pseudachorutes, as indicated by P. boerneri

Schott, 1902; this is why I consider Stachorutes a good genus, but Pratanurida a

subgenus.

Neanura granulata Cassagnau & Delamare Deboutteville, 1955 (Fig. 28 d)

Material: sample 6: 19 and lg ; 35:19.

Description. Since the present material differs somewhat from the original

description, which was based on 10 specimens from the Lebanon, I shall describe

the Cretan specimens in some detail.

. ELLIS: Collembola from Central Crete

Fig. 28. Pseudachorutes (Pratanurida) mucronata n. sp. a, b, postantennal organs; c, antennal organ III.

Neanura granulata Cassagnau & Delamare. d, chaetotaxy of tubercles di-di on abd,. Lathriopyga

anthrenoidea n. sp. e, f, apex and base of a long macrochaeta; g, small macrochaeta; h, maxilla; i, left

antenna,_,; k, unguis P,; 1, mandible

275

276 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Length of male 1.1 mm; females 0.9 (sample 6) and 1.4 mm (sample 35). (This is

at least 1 mm smaller than the original material, and some difference must possibly

be explained by allometric growth). Colour in alcohol totally white, except the

2+2 lively blue eyes. Skin moderately granulate for the genus, more so on the

tubercles, which are well differentiated by their coarser granulation and by their

being elevated and provided with re-inforcement ridges. Antennae typical for the

genus, apical bulb only indistinctly trilobate. Mouth parts also normal with needle-

shaped maxilla and simple tridentate mandible. Claw toothless, no tenent hairs.

Ventral tube with 4+4 setae, rudiment of furca barely distinguishable, only

three setae present at this location. Sixth abdominal segment completely visible

from above.

Hairs of varying length, distinctly rugose with a pronounced double contour.

In the following Table 3 distinction is made between long (L) and short (S)

macrochaetae, microchaetae (m), and setae sensuales (s). A macrochaeta is

considered short if it is at most half as long as a seta in the same or a bordering

tubercle. This denotation is only useful for the dorsal chaetotaxy; it loses its clarity

for e.g. the lateral cephalic chaetotaxy. Fusion of tubercles is indicated by

parentheses around the setal code. The anteromedian cephalic tubercle is not

attached to the central one (LSS+LSS); central tubercle (LSS+S+LSS); ocular

tubercle LLs; lateral tubercles of head LLLSmmm.

Note. In their drawing of abd, the authors of the species indicate that the fusion

of the dorso-internal tubercles on that segment is rather incomplete, although they

state in the text that the tubercles are fused along the median line. However,

Massoud (1967), who studied the types, keys granulata with the species having

these tubercles free, and this may raise some doubt about my identification.

The microchaetae on the dorso-internal tubercles of abd, are extremely small,

sometimes even shorter than the diameter of a skin granule, and difficult to detect

(Fig. 28 d). This not only differs from what the authors depicted, but may also

confuse identifications when only the gross seta number is indicated.

Other points of divergence from original description are the full separation

between central and anterior cephalic tubercle (perhaps not a very important

Table 3. Number of setae on dorso-internal (di), dorso-external (de), dorso-lateral (dl), and lateral (1) tu-

bercles in Neanura granulata

di de dl |

head (posterior row) ES LS LLSS

thorax 1 N SS L

2 LSS LSs LLSs

3 LSS LSsm | LSSs

abdomen | LS LSsm LS LSs

2 LS LSsm LS LSs

3 LS LSsm LS LSs

4 LS LSs LSS LSs

5 (LSm + LSm) (LLSSss)

6 (LLLLSSS)

——66&6&6F6Ftty=ttt BS mm mnt-rr——————————————__—__

W. N. ELLIS: Collembola from Central Crete Die

Fig. 29. Neanura cretensis n. sp., dorsal chaetotaxy

278 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

point, since the authors report having encountered difficulties in delimiting the

tubercles) and the presence of an unpaired seta d, in the central tubercle.

If this description is compared with that of N. tetrophthalma (Stach, 1929) given

by Dunger (1966), it is evident that the differences between the two species are

feeble indeed. The most useful criterion seems to be the relative lengths of hairs,

as Dunger has already remarked, especially with respect to the central cephalic

tubercle (tetrophthalma has seta sd, long and stout in the central tubercle, the

others are very thin and short; d, is unpaired — as in the present material!).

Neanura cretensis n. sp. (Fig. 29)

Material: sample 27: 19, holotype.

Description. Length only 0.7 mm; nevertheless, the specimen seems fully adult, .

the genital orifice having about 16 setae on anterior lip. Blue pigment dispersed in

irregular dots all over the body; eye patches intensely black. Integument between

the tubercles moderately granulated, in the tubercles rather strongly granulated,

posteriorly even more so. Especially the posterior tubercles are very strong. Ocelli

2 +2. Sixth abdominal segment completely visible from above.

Antenna normal, apical papilla distinctly trilobed, sense hairs distinct. Mouth

cone long, mouth parts not dissected, apparently normal for the genus. Unguis

toothless, unguiculus absent. Ventral tube with 4+4 setae. Furca rudiment not

recognizable.

Tubercles. Central and anterior cephalic tubercles free. Dorso-internal tubercle

on head hind margin without rosette, as is the dorso-internal one in th,. Dorso-

internal tubercles on abd, fused, dorso-external ones fused with the lateral

tubercles.

Chaetotaxy (Fig. 29). Macrochaetae without double contour, almost without

serrations or granulations, not constricted at base. True microchaetae, which are

distinct in granulata, are almost lacking. Again with the denotation: L: long

macrochaeta, S: short macrochaeta (relative to macrochaetae in the same or

bordering tubercles), m: microchaeta, s: sensory seta, the chaetotaxy is as

indicated in Table 4. Head. central tubercles (LSS + S + LSS), anterior tubercle:

(LSS + LSS), ocular tubercle: LLS, laterali LLLmmm. Between central tubercle

and antennal base a free seta S.

Discussion. This is again a species closely resembling N. tetrophthalma (Stach,

1929). This and some related species have been treated recently by Dunger (1966).

N. cretensis differs from N. tetrophthalma tatricola (Stach, 1951) by having well-

pigmented body and eyes; it differs from N. t. lusatica Dunger, 1966, by having the

macrochaetae neither granulate nor basally constricted, and also in the different

length proportions of the macrochaetae (most distinct in central cephalic tubercle

and abd,). N. cretensis differs from typical tetrophthalma by having the unguis

toothless, and from all mentioned taxa by having the tubercles much more

developed.

Lathriopyga anthrenoidea n. sp. (Fig. 28 e-l and 31)

Material: sample 25: 1 juvenile, holotype, and 3 more juveniles; 43: 1 juv.

Description. Length of the largest specimen, the holotype, is 1.4 mm; others

W. N. ELLIS: Collembola from Central Crete 279

Table 4. Number of setae on dorso-internal (di), dorso-external (de), dorso-lateral (dl), and lateral (1) tu-

bercles in Neanura cretensis n. sp.

di de

head (posterior row) LS LS

thorax | N LSs

2 LSS LSs

3 LSS LSs

abdomen | LS LSs

2 LS LSs

3 LS LSs

4 LS LSs

5 (L+L) LLSS

LLSSss

6 LLLSS

dl |

LS LSs

LS LLs

LS SSss

LLS SSSSSss

range from 1.1.-1.3 mm (a mean is of course not meaningful in immature material).

Habitus normal for the genus, due to the very long macrochaetae not unlike that

of a carpet beetle larva, Anthrenus. Abd, completely hidden. Pigmentation wholly

absent except for the 2 + 2 strongly dark-blue pigmented eyes. Skin granulation

basally not very strong, well spaced. Tuberculation in the rosettes not very strong

though distinct.

Fig. 30. Lathriopyga longiseta (Caroli), dorsal chaetotaxy of head and th,

280 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Ant, with trilobed, not very distinct apical papilla and 8 sense hairs. Antennal

organ III two sensillae, strongly bent in the same direction, guarded by a strong

ventral and a weaker dorsal sense hair (Fig. 28 i). (Skin granulation of antenna

rather uneven. Dorsal part of ant,, and whole basal part of ant, rather coarsely

granulate, other parts, especially dorsal part of ant, weakly granulate).

Eyes 2 + 2. Mouth cone long; mouth parts not dissected; in transparency a 6-

toothed mandible is visible, with basal tooth strongly developed (Fig. 28 1), and a

needle-shaped maxilla with two lamellae which are closely appressed (fused or

not) and provided with some apical denticulation (Fig. 28 h).

Feet plump, with a distinct tooth on inner lamella; lateral lamellae toothless

(Fig. 28 k). Subcoxae with 1, 3, 3 setae. Ventral tube with 4 + 4 setae. Furca

rudiment a median boss with some 8 setae.

Vestiture consisting of very heavy, blunt, distinctly rugose, apically winged

macrochaetae of variable length, often conspicuously long, a few minute

microchaetae, some sensory setae, and ventrally shorter, undifferentiated setae.

The dorsal chaetotaxy is summarized in Table 5 (cf. also Fig. 31).

Table 5. Number of setae on dorso-internal (di), dorso-external (de), dorso-lateral (dl), and lateral (1) tu-

bercles in Lathriopyga anthrenoidea n. sp.

di de dl |

head (posterior row) LS L LLSSS

thorax | L LS L

2 LSS LSs LSSs

3 LSS LSSs LSSs

abdomen | LS LSSs LS LSs

2 LS LSSs LS LSs

3 LS LSSs LS LSs

4 LS LSs LSSs SSSss

LS LLSSSS

On the head all tubercles are free and well developed. The apical tubercle has

setae (LS+SL), the central tubercle has (LSS+SSL) and lacks an unpaired seta,

ocular tubercle has LLm, lateral tubercle LLLSSmm.

Discussion. The complete absence of fusion of tubercles makes this species very

distinctive. This is why I venture to describe the new taxon on the basis of the

present meagre material. Also, the distinct tooth on the unguis (which is, however,

much less strongly developed than in /ongiseta (Caroli, 1912) and hellenica Ellis,

1974) is a rather divergent character. The thoracic chaetotaxy points to an affinity

with the L. phlegraea group, but there the dorso-internal tubercles are fused in

abd,.

Lathriopyga longiseta (Caroli, 1912) (Fig. 30)

Material: sample 35: 13 .

The single specimen, measuring 1.1 mm, agrees well with the original

description and the more detailed description given by Da Gama (1964). The

281

W. N. ELLIS: Collembola from Central Crete

Fig. 31. Lathriopyga anthrenoidea n. sp., dorsal chaetotaxy

282 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

species has been recorded from Italy, Corsica, Madeira, Yugoslavia, and — since

the synonymy with L. hystrix (Bagnall, 1940) was established by Lawrence in 1970

— England. The near relative L. hellenica Ellis, 1974, was described from Rhodes.

ISOTOMIDAE

Coloburella linnaniemii (Denis, 1937)

Material: sample 21: 2 juv.

Their immaturity taken into consideration, the two specimens are in good

agreement with the redescription of C. linnaniemii given by Rusek (1972). The only

notable deviation is the somewhat longer furca — the tip of the mucro reaches —

onto abd, over about % segment length. The species is now known from Italy and

Crete; the near relative C. cassagnaui Rusek, 1972, occurs in the Pyrenees.

Tetracanthella cf. hygropetrica Cassagnau, 1954 (Fig. 32a)

Material: sample 19: 9 immature specimens, only some of recognizable sex (29,

200):

Discussion. A gradual intergradation seems to exist from 7. tuberculata

Cassagnau, 1954 (described from the Sierra de Guadarrama, since then recorded

from Portugal, the Pyrenees, Corsica, Italy, Yugoslavia, and the USSR), through

T. hygropetrica luxemburgensis Stomp, 1968 (Luxemburg) and 7. h. matthesii Da

Gama, 1959 (Madeira), to 7. h. hygropetrica (described from the Pyrenees,

recorded from Portugal and Spain as well as from Italy). In fact, the best argument

in favour of the validity of luxemburgensis and matthesii is their isolated

occurrence.

The present material is, due to its immaturity, difficult to identify. It has a

manubrium/dens ratio of about 6/5 (Fig. 32a), which comes closest to the

Portuguese hygropetrica hygropetrica described by Da Gama (1964). In the present

material reticulation is mixed and fine, mostly as narrow as the diameter of the

hair rings, locally narrower. A smooth area occurs only in postero-median part of

abd,.

Isotomodes trisetosus Denis, 1923

Material: sample 13:19.

In her. monograph of the genus, Da Gama (1963) gives records from South-

Central Europe, Madeira, the Azores, and Peru.

Folsomides parvulus Stach, 1922 (Fig. 32b, c)

Material: sample 16: 39 and 2 juv.; 29: 4 juv.; 30: 59 and 14 juv.; 35: IQ and 5

juv.; 36: 29; 37: 19; 38: 19; 43: 139 and 13 juv.; 44:69 and 6 juv.; 45:19.

Discussion. My description and notes on variability in material from Rhodes

(Ellis, 1974) are also applicable to the present material. The eye number is

invariably 2 +2; the eyes are sometimes devoid of pigment, especially in juveniles

W. N. ELLIS: Collembola from Central Crete

Fig. 32. Tetracanthella cf. hygropetrica Cassagnau, 1954. a, posterior face of furca and retinaculum.

Folsomides parvulus Stach. b, ventral face of ant; c, manubrium of holotype. Folsomides nanus Ellis. d,

ventral face of ant,; e, ventral aspect of abd,_,. Folsomia ksenemani Stach. f, ventral face of ant, (sensilla

indicated by x is absent in smaller immatures); g, chaetotaxy of anterior face of manubrium, with

indication of the order of appearance of the setae in the ontogeny

283

284 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

and small adults. Posterior face of dens with 3+3 setae, except in three specimens

(two juveniles from sample 16, and a 9 from sample 30) in which I found only

2+3.

Dr. A. Szeptycki kindly loaned me the holotype of the species under

consideration. It is mounted on a slide bearing two labels, both with a blue margin

and handwriting: a small one stating “Unicum”, and a larger one “Folsomides

parvulus n.g.-n.sp. Berekalja 1915”. The specimen is an adult 9 and measures

590 u. The condition is rather poor, but I could study the furca in some detail.

The dens has at one side distinctly 3 setae; in the other dens the two distal setae

are visible, but due to a dirt particle observation of the basal seta is impossible.

The posterior face ofthe manubrium bears 6 + 6 setae (Fig. 32c).

Again, the species seems to be parthenogenetic.

Folsomides americanus Denis, 1931

Material: sample 13: 19 and 1 juv.; 26:49 and 1 juv.; 27:39 and 6 juv.; 43: 4

juv.

Discussion. The difference between americanus and parvulus is based on one

character only. The posterior pair of ocelli is lost in americanus. Not only pigment

granules but also a lens are missing, and the primary granulation on the location of

the second pseudocellus in parvulus is completely regular.

After some hesitation (due in part to the apparent coexistence of americanus and

parvulus in sample 43), I prefer to separate americanus as a distinct species. One of

the arguments for doing so is that parvulus shows no trend indicating obliteration

of the posterior ocellus: posterior and anterior ocellus always show an almost

equally dense pigmentation and are equally large; moreover, there are no

asymmetries.

The ocelli of many immatures of americanus are not or only weakly pigmented,

as in parvulus and ‘‘anophthalamis’’ Hepburn & Woodring, 1964.

Although the limited number of specimens does not permit of definite

statements, here too there is an indication of parthenogenesis.

The species seems to occur in all subtropical regions in the Americas and

Europe. It has already been recorded from the Lebanon (Cassagnau & Delamare

Deboutteville, 1955), Spain (Selga, 1973), and France (Cassagnau & Rouquet,

1962).

Folsomides navacerradensis Selga, 1962

Material: sample 36:49, Sg , and 11 juv.

Discussion. The specimens agreed well with the description by Selga, apart from

the variability in dental chaetotaxy, as described by Petersen (1965). The number

of setae on the posterior face of the manubrium fluctuated (in adult specimens)

between 7+7 and 12+12; on dentes both 3 and 4 setae occurred. The Cretan

material thus cannot be related to F. navacerradensis pratensis Palissa & Zivadino-

vic, 1974, from Bosnia. Here, only 3 setae are constantly present on the dens, and

W. N. ELLIS: Collembola from Central Crete 285

the manubrium has at most 8 pairs of setae. In two specimens (a juvenile and a g )

one of the dentes lacked the mucro. This is the distinctive feature of F. meridio-

nalis Dallai, 1973, described from the Eolic Islands.

Folsomides marchicus (Frenzel, 1941)

Material: sample 24: 29.

The specimens did not deviate from the original description. Pigment was fairly

well developed; posterior face of dens with 3 +3 setae in both specimens.

The species has been recorded before from Spain and Ibiza, France, Italy,

Germany, Switzerland, the CSSR, and Yugoslavia.

Folsomides nanus Ellis, 1974 (Fig. 32 d, e)

Material: sample 12: 2 9 and 1 juv.; 25: 19; 30: 29; 36: 189 and 23 ; 43: 29,

3g „and 3 juv.; 45: 169,65 ‚and 4 juv.

Discussion. Mean length of the females 0.70 mm (s = 0.08, s; = 0.01, n= 41),

that of the males 0.55 mm (s = 0.04, s; = 0.01, n = 11). The specimens are in good

agreement with the material of nanus from Rhodes, with one exception. The dens

shows no trace of a mucro and is very blunt in the ventral view. In the material

from Rhodes, the dens is always more or less contracted apically to a vestige of a

mucro in the ventral view. Moreover, in the present material the manubrium has

at its posterior face a slightly higher number of setae, the modal value being 6+6,

often asymmetrically 5+6 or 6+7 (Fig. 32 e).

Since the Cretan material was collected in the autumn, seasonal variation may

partially explain the difference, and I therefore refrain provisionally from giving

the Cretan material subspecific status.

Folsomia ksenemani Stach, 1947 (Fig. 32 f, g)

Material: sample 12: 2 juv.; 13: 33 juv. (2 immature 9); 14: 3 juv.; 19: 10 juv. (1

immature 9); 21: 1 juv.; 25: 39 juv.

Discussion. The number of ocelli is constantly 1+1, and I have no reason to

agree with Christiansen (1959) and other authors that ksenemani represents

aberrant one-eyed specimens of a two-eyed species such as penicula Bagnall, 1939.

In the antennal organ I, two small sensillae (but in the smaller immatures

constituting the majority of the present material only one) are guarded by a sense

hair which is barely differentiated from a normal hair (Fig. 32 f). Moreover, a

small basal spine is present here; the dorsal homologues being two comparable

spiniform setae. Thoracal sternites without setae. Ventral tube anteriorly bare,

posteriorly usually with 4 setae, in lateral flaps usually with 3+3, sometimes with

3+4 and in one example even with 4+4 setae. All this applies to immature

material. |

The number of setae on anterior face of manubrium varied strongly. Table 6

gives the frequencies I could observe. Although the present material includes no

mature specimens, the higher setal numbers already outvalue the limit given by

286 TIJDSCHRIFT VOOR ENTOMOLOGir, DEEL 119, AFL. 8, 1976

Table 6. Number of anterior setae of the manubrium of Folsomia ksenemani

——————___=—+— ——————m m-mmmr____—_—_——rrrc"- td

Sample nr. Number of setae

4 5 6 7 8 9 10 11 12 13 14 15

12 - I | -

13 3 2 3 5 7 4 fl I |

14 - - l | | -

19 1 2 l l l 2 2

21 - - - l - - - - - - -

DS - - - - 2 4 3 4 21 2 3 -

Total l - 5 4 7 10 14 10 30 2 4 |

Stach (1947) for ksenemani, and consequently the material should be identified as

F. pseudodiplophthalma Stach, 1947. These two species have, however, been

synonymized on rather convincing grounds by Poinsot (1972). Acting furthermore

as “first revisor”, she has invalidated pseudodiplophthalma, and I follow her in the

synonymy. The low values are, however, no less instructive, because they suggest

an impending synonymy of ksenemani with (the senior name!) monophthalma

Bagnall, 1939, described from Romania, and, according to Lawrence (1973) an

immature! Topotypical material might show that this suspicion is justified.

The gradual rise in setal number on the manubrium follows a rather regular

pattern. In Fig. 32 g I have numbered the setae according to the order in which

they appear. Only level 4 and 5 are interchangeable, and I often found a specimen

with both arrangements asymmetrically.

Cryptopygus thermophilus (Axelson, 1900)

Isotomina salaymehi Christiansen, 1959. — nov. syn.

Material: sample 4: 39, 33, and 6 juv. + 10 ex. in alcohol; 7: 29, 63 , and 3

juv.; 18: 19 and 5 juv.; 20: 19, 1g, and 10 juv.; 32: I juv.; 41:29 and 2 juv. + ca.

90 ex. in alcohol; 48: 39, 1g, and 2 juv. + 100 ex. in alcohol.

Discussion. The unguis is toothed in almost all specimens. Tenent hairs not

clavate. Proportional width of postantennal organ rather variable. Ventral tube

with 4+4 setae on lateral flaps, and 1+1 posterior setae. Thoracic sternites

without setae.

The species described by Christiansen, recorded from several localities in the

Lebanon, would differ from thermophilus by a pair of blunt sensory setae on abd,, a

faint subdivision in the distal part of the tibiotarsus, a small ocellus D, and by

having the longest setae on the abdomen three times longer than shortest ones. All

these characters can be found in material referable to thermophilus, not only in the

present material, but also in specimens from western Europe. The sensory hair of

abd, is typical for the whole species group.

The species seems to be cosmopolitan.

Cryptopygus ponticus (Stach, 1947)

Material: sample 9: 1 juv.; 10: 19 and 6 juv. + 68 ex. in alcohol; 11: 19, 55 ,

and 2 juv. + 23 ex. in alcohol; 16: 1g ; 17: 3 juv.; 23: 4 juv.; 29:19, 18, and 2 juv.;

W. N. ELLIS: Collembola from Central Crete 287

BD ovand 2 juv.; 34:19 and | juv.; 36:19 and 23 ; 37:29; 38:19; 41:19, 1g,

and | juv.; 42: 1g and 5 juv.; 43:23 and 8 juv.

Discussion. Tooth on internal ungual lamella distinct. Lateral flaps of ventral

tube constantly with 4+4 setae. Posterior face of ventral tube in adult specimens

always with more than 1 +1 setae: modal value 2 +2, less frequent values 1 +2 or

2+3. (This is a useful additional difference from C. thermophilus, but may not be

valid everywhere, since Yosii (1966b) described ponticus from Afghanistan with

1+1 posterior setae). Juveniles often have only 1 +1 posterior setae; in adults the

longest, distal setae seem to be the homologues of the pair present in C.

thermophilus.

Cryptopygus triglenus n. sp. (Fig. 33)

Material: sample 23: 2 juv.; 26:59, 52 , and 3 juv.; 27:59 and 5 juv.; Holotype

isag from sample 26.

Description. Mean length of the 5 males 0.76 mm (s = 0.03, s; 0.01), that of the

10 adult females 0.92 mm (s = 0.08, s; 0.03).

Pigment sparsely distributed in grey clouds over head and body; eye patches are

well pigmented. Integument finely and regularly granulate. Hair cover consists of

moderately fine smooth setae.

Ant: Head = 1.08. Ant. 1:2:3:4 = 36:54:57:100. Ant, without apical retractile

bulb, only a semiglobular protuberance; subapically, a small sensilla in a deep

groove, guarded by a short strongly bent seta; the segment is covered by normal

hairs and by many sense hairs. Ant, with ant. org. III composed of two flexed

sensillae guarded by two sense hairs; external sensilla in the middle of this segment

spine-like (Fig. 33 d). Antennal organ I with two long sense hairs, the external one

the shortest, and a small spine-like setula near base of segment (Fig. 33 e).

Eyes 3+3. Two ocelli set close together in one intense pigment spot, a third one

lies about 1.5 eye-diameters more caudad, on its owh pigment bed. Postantennal

organ oval, with a thickened anterior “rim” and a weak transverse subdivision

(Fig. 33 b). Labral chaetotaxy 4/5, 5, 4. Mandible normal, maxillar head with

rather narrow lamellae which do not reach beyond the claw.

Thoracal sternites without setae. Unguis with a distinct inner tooth, without

lateral or dorsal teeth; unguiculus with weak inner lamella, without apical filament

(Fig. 33 f). No clavate tenent hairs. Apical part of tibiotarsus weakly separated

from main part by indistinct furrow.

Abdominal sternites in mid-line bare. Ventral tube without anterior setae,

posteriorly with about 2 +2 setae (the distal pair the strongest), in lateral flaps with

4+4 setae. Retinaculum quadridentate with 1 seta on corpus. Male and female

genital orifices normal. Abd, and abd, completely fused. On abd, ventrolaterally,

an indistinct sense hair.

Manubrium anteriorly with 1 +1 strong subapical setae, posteriorly with about

20+20 setae in a very symmetrical arrangement (Fig. 33a). Two pairs of

manubrial spines. Dentes finely corrugated, anteriorly with 15 strong setae,

posteriorly with 1, 1, 1, 2, 2 fine setae (Fig. 33 c). Mucro bidentate, not lamellate.

Discussion. As far as I know, the genus includes only two species with 3 + 3 eyes:

TISDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

288

DELDEN

AAL

> nm

Fig. 33. Cryptopygus triglenus n. sp. a, posterior chaetotaxy of manubrium; b, eye-patch and

postantennal organ; c, posterior and anterior aspects of mucrodens; d, antennal organ III; e, ventral

aspect of ant,; f, claw of P,

W. N. ELLIS: Collembola from Central Crete 289

Fig. 34. Cryptopygus debilis (Cassagnau). a, posterior aspect of manubrium; b, mucro; c, location of

sensillae and some setae on ant,; d, anterior aspect of manubrium, posterior (left) and anterior face

(right) of mucrodens; e, claw of P,; f, antennal organ I; g, receptaculum seminis; h, dorsal aspect of

postantennal organ and ant,

290 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

C. trioculatus Izarra, 1972, from Argentina, and insignis Massoud & Rapoport,

1968, from Patagonia. The former belongs to the series with the dens much

reduced (anteriorly only one, posteriorly two setae) and is undisputedly very

different from triglenus. C. insignis, however, comes close to triglenus, but differs in

the tridentate mucro and the somewhat shorter dens with a different chaetotaxy,

especially posteriorly, and, finally, in the enormous postantennal organ.

C. triglenus obviously is a near relative of C. ponticus, and differs from that

species only in its eye number. It is not without some hesitation that I propose this

new species on the basis of a single difference. That for the time being I

nevertheless consider triglenus a good species is explained by the fact that no

intermediate eye numbers (3+4, 4+4 or 4+5) have been detected in the rich

material available.

The specific name means “with three jewels”.

Cryptopygus debilis (Cassagnau, 1959) (Fig. 34, 35 a, b)

Material: sample 13: 19 and 12 juv.; 14: 2 juv.; 25: 6 juv.; 35: 69 ; 36: 1 juv.; 44:

16 juv.; 45:19 and I juv.

Description. Isotomina debilis was described from the Central Pyrenees, from

two alpine localities. This makes my identification less likely. It is unfortunate that

the original description is very short, and I add, therefore, a complete description

of my material here.

Mean length of the 7 adult females 0.53 mm (s = 0.02, s; = 0.01). No trace of

pigment. Integument basically normal, with a fine primary granulation. However,

abd „and abd , , show numerous, irregularly distributed circular structures which

I cannot interpret (Fig. 35a). They are about as large as hair bases, sometimes

smaller, and consist of a raised ring with a depressed centre — resembling an

erythrocyte. They occur in juveniles and adults, and are not correlated with

moulting. I would consider this phenomenon to be an artefact if it was not always

limited to the same segments and did not occur in material from all localities. I

have never seen a comparable structure, save perhaps in entomobryoid pseudo-

pores. Hairs normal, smooth.

Antennae: head = 1.2; ant. 1:2:3:4 = 30:50:55:100. Ant, without exsertile

papilla, subapical sensilla remarkably large, guarded by an equally proportionally

strong, bent hair. Some sense hairs, difficult to identify, occur on this anten-

nomere, as well as 3 outer and 1 inner swollen, curved sense hairs (Fig. 34c). Ant,

with antennal organ III composed of two short sensillae, guarded by two

apparently normal hairs. External sensilla of this antennomere in the shape of a

curved spine. Antennal organ I consists of a thick outer and a thin (and a bit

shorter) inner sense hair (Fig. 34f).

Eyes absent. Postantennal organ usually very large, almost without a rim (only

some thickening in ventral corner). Some specimens (juveniles) have, however, a

somewhat less disproportionate PAO. Mouth parts normal. Labral chaetotaxy is

313,904.

W. N. ELLIS: Collembola from Central Crete 291

9) 7 5 \\ Bates CARA NA \

A; NS ALY, JAMMIN)

Nin dai

end

O 9 Yj N fated Yuan IN UN Mia a

) // / ory 7 4 \ \\

arts () VV Lo i LIDI US

Fig. 35. Cryptopygus debilis (Cassagnau). a, sensilla, some setae, and cuticul amentation of abd,; b,

chaetotaxy of abd, ,. Clavisotoma albertina ateral aspect of furca; d, anterior (left) and

ens

e n. sp. c, l

posterior (right) aspect of mucro

292 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Thoracal sternites bare except 1+1 seta on th,. Claws simple, unguis without

teeth, unguiculus with proportionally strongly developed outer lamella, tenent

hairs not clavate; subsegmentation of tibiotarsi not seen (Fig. 34e).

Ventral tube anteriorly without setae, 4+4 setae on lateral flaps, posteriorly

usually with 1+2 setae (if more adult material had been available this might have

been a higher figure). Retinaculum quadridentate, with one seta on corpus.

Abdominal sternites bare in median line. |

Manubrium posteriorly with about 16 +16 or 17 +17 setae (Fig. 34a), anteriorly

in all except two adult specimens with 2+2 setae, arranged in a distal pair and

another pair in about the middle of the manubrium (Fig. 34d). The same

arrangement was found in only one juvenile. The remaining juveniles and one

adult had only the distal pair of setae. The last adult specimen, which was at the

point of ecdysis, had 1 + 1 setae on the old skin, and 2 +2 on the new one.

Manubrium distally with a pair of weak though distinct manubrial spines

(Fig. 34a). Dentes posteriorly with about 25 corrugations, basal and apical part not

corrugated (Fig. 34d). Anteriorly there are about 30 setae, posteriorly 1, 1, 2, 2 fine

setae. Mucro bidentate (Fig. 34b).

Abdomen 5 and 6 completely fused. Abd, with 1 +1 short clavate sensillae. They

are usually relatively very thick, but in some cases are slightly more slender (Fig.

35a, b).

Since part of the juvenile material consisted of immature males, there is no

question of regarding the species as parthenogenetic.

Since Mr. P. N. Lawrence is preparing a revision of the genus Cryptopygus, I

leave it to him to compare this species with the other members of the genus.

Proisotoma minuta (Tullberg, 1871)

Material: sample 4: 19; 6:19; 23:19, 1g, and 2 juv.

The species seems to have an almost cosmopolitan distribution.

Ballistura schoetti (Von Dalla Torre, 1895) (Fig. 37f)

Material: sample 40: 19 and 16 juv.; and also some 120 specimens, apparently

all immatures, in alcohol.

The only adult specimen measured 1.0 mm, and was still very pale. This

halophilous species is distributed along the coasts of the Atlantic and the

Mediterranean.

Clavisotoma albertinae n. sp. (Fig. 35c, d, 36, 37a-e)

Material: sample 26: 4 juv.; 32: 159, 133, 3 juv., and | specimen of unknown

sex; 36: ca. 550 ex. in alcohol; 37: 1 juv.; 38: 109, 11g, 11 juv., and ca. 50 ex. in

alcohol; 39: ca. 300 ex. in alcohol, not sorted according to sex; from 43: 19; 50:

19. All material (except the specimens in alcohol) mounted individually on slides

in Marc Andre II; some specimens depigmentated. Holotype is a 4 from sample

32, all other specimens being paratypes except a very immature specimen from 32,

which cannot be attributed to the species with sufficient certainty.

W. N. ELLIS: Collembola from Central Crete 293

Fig. 36. Clavisotoma albertinae n. sp. a, maxillar head; b, articulation of mandible against the clypeus; c,

mandible; d, antennal organ III; e, antennal organ I; f, claw of P,

294 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Description. Total length 1.1 mm, the sexes not differing markedly in size.

Habitus stout, characteristic for the genus. Pigmentation moderate, bluish-grey.

The whole body and all the extremities are pigmented; head and dorsum distinctly

darker than other parts. Integument smooth, hairs short and stout, curved.

Antenna: head = 0.85. Ant. 1:2:3:4 = 45:48:60:100. Ant, without apical papilla, but

with a small subapical globule in a deep pit. Antennal organ III consists of two

short blunt and strongly curved rods, guarded by two also proportionally rather

short and blunt sense hairs. Another short and curved sensilla at outer surface of

same segment (Fig. 36d). Antennal organ I (ventrally on ant,) composed of two

comparatively long blunt sense hairs and a short but otherwise seemingly normal

hair (Fig. 36e). Eyes 8 +8, all well developed though not all equally large (G and H

are smaller), in a black eye patch (Fig. 37b). Postantennal organ shortly oval, with ,

an irregular, sometimes rather thick rim; its longest axis is about as long as the

diameter of ocellus B (Fig. 37e).

Labral chaetotaxy 2/5, 5, 4. Mandible stout, but essentially normal for the

family, with a strongly chitinized outer condyle (Fig. 36 b, c) which is apposed to

the interior surface of the clypeus. Maxilla essentially typical with large, three-

toothed claw, globular body, and two dorsal and two ventral fringed lamellae,

which seem to have a tendency to split (Fig. 36a).

Tibiotarsi of P, with one, those of P,_, with two clavate tenent hairs. Unguis with

distinct inner tooth, without lateral or dorsal teeth. Unguiculus with strongly

curved inner lamella, tapering to an apical filament (Fig. 36f). Thoracic sternites

bare.

Ventral tube with 4+4 setae on lateral flaps, and 1 + 1 posterior setae (Fig. 37d).

Retinaculum tridentate, with 1 seta on corpus (Fig. 37c). Abdominal sternites

bare.

Manubrium anteriorly bare, posteriorly with many (40-60) setae. Antero-distally

a pair of strongly sclerotized acute manubrial hooks (Fig. 37a). Anterior face of

dens with a large, somewhat variable number (8-15) of setae in distal half;

posterior face with about 13 setae and the bosses characteristic for the genus.

Mucro typical for the genus, large, two-toothed, strongly lamellate (Fig. 35c, d).

Male genital orifice on a tubercle surrounded by a dense row of about 30-40 fine

hairs.

Discussion. An inspection of the synoptic key I prepared for Clavisotoma (Ellis,

1970), shows that a combination of characters: presence of tooth on the unguis,

8+8 eyes, tenent hairs 1, 2, 2, retinaculum tridentate, terminal filament present,

has not previously been found in this genus. C. albertinae is furthermore highly

characteristic by virtue of the large number of setae occurring antero-distally on

the dens. C. albertinae seems most closely related to hankoi (Stach, 1930),

described from Hungary and recorded from Switzerland (Gisin, 1960).

In Marathos (samples 37, 38, 39) we witnessed a true mass occurrence of this

species, albeit limited to only 20-30 square metres (as far as could be explored),

where countless numbers were walking about on the almost bare, moist clay of a

piece of waste land and on stones lying at its edge. We presume that the animals

had been (dormant?) living deep in the soil during the dry season, and came to the

W. N. ELLIS: Collembola from Central Crete 295

Fig. 37. Clavisotoma albertinae n. sp. a, manubrium-dens junction; b, eye patch; c, retinaculum; d,

ventral tube; e, postantennal organ and anterior ocelli. Ballistura schoetti (Von Dalla Torre). f, ventral

aspect of ant,. Dimorphotoma porcellus n. sp. g, maxillar head; h, lateral aspect of mucrodens; i, ventral

aspect of mucrodens; j, claw of P,

296 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

surface — or were actually even driven to the surface — by the soaking and

swelling of the clay of the subsoil after the torrential rains of the last few days.

It is a pleasure for me to dedicate this species to my wife Albertine, who not

only participated actively in the collecting but also has the gifted collectors knack

of finding unusual things in unexpected places. It was she who discovered the mass

occurrence of the species under consideration.

Dimorphotoma porcellus n. sp. (Fig. 37 g-j, 38, 39, 40 a-d)

Material: sample 25: 79 and lg ; 26: 939 and 693 ; 27: 19; 28:99 and 195.

Moreover as alcohol material, not sorted according to sex, about 700 specimens

from sample 26. Holotype isa 3 from sample 26. i

Description of female. Total length 1.4 mm (sd = 0.11 mm, n = 12). Habitus

stout, about as in Clavisotoma. Pigmentation sparse: greyish-blue pigment tinges

(without strong concentrations or mottlings) the whole dorsal part of the body,

especially the thoracic tergites, and the extremities. Only the enveloping cells

around the internal part of the eyes are strongly pigmented. Integument smooth,

primary granulation very fine, except in the ‘‘crown” on abd, (see below). Hair

cover dense, consisting of fine, smooth acute setae, only on abd, ‚a bit longer, and

sometimes with a few serrations at anterior face; some special setae around the

“crown”.

Abd, well separated from abd,. Seen from above, abd, seems fairly small due to

pronounced cryptopygy.

Ant/head ratio is 1.05. Ant 1:2:3:4 = 42:65:66:100. Ant, without apical exsertile

papilla, but with a slender spine-like sensilla subapically, a small curved sensilla in

a shallow groove, many seemingly normal hairs, and dorso-apically (ca. 20) slender

curved hair-like but blunt sensillae. Ant, with ant. org. III consisting of two small

bent sensillae flanked at both sides by a somewhat larger almost straight hair,

furthermore a sensilla in the middle of outer face (Fig. 40d); ant, with normal

setae, one very slender curved blunt sense hair apically in latero-ventral position,

and proximo-ventrally as well as proximo-dorsally, a very fine and short hairlet;

ant, somewhat swollen, proximo-dorsally with a fine setula, ventrally with antennal

organ I consisting of a fine sense hair, a shorter curved sense hair, and a short

sensilla between them (see Fig. 40b).

Antennal bases swollen, bare; the frontal region seems somewhat jammed

between the antennal bases, the hairs pointing in all directions (Fig. 39a). PAO

broad-oval, twice as long as diameter ocelli A or B, mostly simple but sometimes

with a constriction in the middle of its long axis. Ocelli 8+8, all except G and H

equally large (Fig. 39a). Labral chaetotaxy 2/5, 5, 4, distal two rows on low papillae

(Fig. 40c). Labium normal; labial palp, see Fig. 39b. Mandible normal for the

family (see drawing of Clavisotoma albertinae, Fig. 36c). Maxilla also normal, with

tridentate claw, globular body, and 2+2 lamellae which do not reach beyond the

apex of the claw (Fig. 37g).

Feet 1, 2, 3 with 1, 2, 2, weakly clavate tenent hairs. Unguis without inner of

lateral teeth. Unguiculus about 0.5 times unguis, blunt, without apical filament

(Fig. 37j). Thoracic sternites | and 3 bare, that of th, with 1+ 1 seta.

W. N. ELLIS: Collembola from Central Crete

Fig. 38. Dimorphotoma porcellus n. sp., habitus of a male

297

298 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Ventral tube anteriorly without seta, laterally with 4 +4, posteriorly with 5+ 5 or

6+6. Abdomen ventro-medially without setae. Retinaculum tridentate, with one

seta on corpus.

Manubrium: dens: mucro = 10: 6: 1. Furca moderately long, reaching to the

distal border of abd,. Manubrium anteriorly with 1+1 (seldom 1+2) distal setae,

posteriorly with many (maximally about 30+30) setae, leaving a medial stripe

bare. Manubrium-dens junction with two low cushion-shaped ridges (Fig. 39c).

Dens anteriorly only distally with 1, i, 2 setae, posteriorly with a variable number

(about 6) of ill-defined humps and 7 setae. Mucro proportionally short, bidentate,

iamellate (Fig. 37h, 1).

Dorsally on abd, there is a rosette shaped like a head of lettuce. It consists of

strongly crisped integument that is relatively coarsely grained (granulation visible

at x 400). The hairs in this vicinity are longer and more slender and bent than the

normal body hairs (Fig. 39e, f).

Female genital orifice of two strongly protruding lips, each having 1 +1 normal

seta, i.e., almost as long as setae anterior and posterior to the genital orifice.

Spermatheca not seen.

Description of the male (Fig. 38).

The male is distinctly larger than the female: mean size of 18 random specimens

was 1.6 mm (s = 0.15). But the essential difference, very striking even at low

magnifications, is the possession by the males of a large number of strong yellow

spines on antenna, head, thorax, and abd ,_,. Moreover, the hair cover on abd, and

the expression of the rosette are different, and the antennae are aberrantly shaped.

Ant, is roughly normal in shape, differs only in the possession of one small,

slightly hook-like spine in lateral position. Ant, is somewhat thickened and has a

slightly oblique base, and also one lateral spine, which is a bit curved. Ant, is

distinctly thickened and has two spines of different length, as has ant,. Ant, is even

more inflated than ant,, and the antennal sockets are correspondingly enlarged.

The spines on head and body are essentially aligned along the lateral parts.

Although asymmetries and anomalies do occur (e.g. the vertical pair on the head

is often absent), the arrangement is essentially a very orderly one. The spines are

replaced on abd, by strong, straight macrochaetae (sometimes having a few

serrations).

The rosette on abd, is much less distinct in the male than in the female, and the

integument lacks the coarse texture occurring in that sex. The setae on abd, (and

to a lesser extent also those on abd,) in the discal area, i.e., close to the rosette, are

strongly protracted to a wavy thread, terminating in a tiny globule (Fig. 39d).

The male genital orifice is a large cone, encircled by about 20 setae; papilla

bearing 4+ 4 setae (Fig. 40a). Vas deferens visible in a few specimens only.

Discussion. Leaving aside sexual dimorphism, the new species comes close to

Clavisotoma Ellis, 1970, especially because of the hump structure of the dens.

However, the pair of setae antero-distally on the manubrium argues against such a

relationship. Otherwise, the resemblance to the littoral Proisotoma buddenbrocki

Strenzke, 1954, is rather striking. It is unfortunate that no submature males are

present in the material, making it impossible at present to say whether the

secondary sexual traits are obtained gradually or not.

W. N. ELLIS: Collembola from Central Crete 299

Fig. 39. Dimorphotoma porcellus n. sp. a, 9, frontal region of head in latero-dorsal aspect; b, palpus and

lobus externus; c, manubrium-dens junction; d, dorsal aspect of abd,_; of a male, showing chaetotaxy

(only some setae drawn) and the rosette; e, dorsal view of abd, of a female; f, rosette of a female in

lateral view

300 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Sexual dimorphism on the scale described here is not common in the

Isotomidae. A comparatively low degree of dimorphism occurs in Proisotoma

papillosa Stach, 1937, Hydroisotoma schaefferi (Krausbauer, 1898), and Austra-

lotomurus echidnus (Womersley, 1934). A really strong sexual dimorphism is known

to occur in only three instances (which are, however, not undisputed — cf. Goto,

1972): Guthriella Borner, 1906, Rhodanella Salmon, 1945, and Dimorphotoma

Grinbergs, 1975 (Dimorphiella Grinbergs, 1968, nec Valkanov, 1928). Guthriella is

much too specialized to be compared with the present species. Rhodanella, with

only the species R. minos (Denis, 1928), is considered by Goto (1972) a possible

ecomorph of Proisotoma stachi Goto, 1957, although the evidence he gives is not

very decisive, and there is no explanation as to why the ecomorphosis would affect

only one sex. However this may be, both the type of sexual differentiation (a pair .

of large horns on the male head, as well as a number of strong but not spiniform

body setae) and the female or submature male morphology exclude a close affinity

between Rhodanella minos and porcellus. Note that Denis originally placed minos in

Vertagopus.

Even the similarity to Dimorphotoma muriphila (Grinbergs, 1968) is not very .

strong. In that species the antennae are not affected by the sexual dimorphism,

and the rosette on abd, with its accompanying specialized setae is not present.

Moreover, in muriphila abd, is strikingly longer than abd,. Less important

differences with respect to muriphila concern the occurrence in this species of

spines in the male not only dorso-laterally but also dorso-medially, and also on

abd,. Finally, muriphila has a tridentate mucro. Still, although I cannot exclude the

possibility that by including porcellus in Dimorphotoma | am making the genus

polyphyletic, I think it better not to create a new genus for porcellus at present.

The specific name, which means a sucking-pig, refers not only to the plump

habitus but also to the roast pig for which Drosia, the type locality, is widely and

rightly renowned.

The species occurred very locally, and in enormous numbers. Very many

specimens were seen walking, apparently erratically, on the ground and on large

boulders. The collecting date was just after some days of heavy autumn rains,

which had thoroughly soaked the loamy subsoil. This and the fact that the material

seems to be composed exclusively of adults suggest that the specimens had passed

the period of summer drought in the subsoil, and had been driven to the surface by

the inundation of their summer resort. Two findings are relevant here: first a

comparatively large percentage (in the order of 10%) of the specimens were

infected by Sporozoa; and second, the genital area was conspicuously dirty, and I

had some difficulty in finding a specimen suitable for a drawing of the male genital

papilla. I should also mention that active testicular tissue, which is usually clearly

visible even in cleared specimens, could not be found, and the same holds for

spermathecae and a distinct vas deferens. In almost all specimens the intestine was

full (unidentifiable, mainly amorphous material). My suggestion is that, syn-

chronized as they were by the weather conditions, the specimens had not yet

passed through their moult to the sexual phase (c.f. e.g. Joosse & Veltkamp, 1970).

No specimens in pre-ecdysis condition were seen.

I have, of course, considered the possibility that D. porcellus might only be an

W. N. ELLIS: Collembola from Central Crete 301

ecomorph. This is not wholly inconceivable after Goto’s (1972) study and also

because the specimens show some integumentary structures often found in

| association with ecomorphosis. Moreover, if ecomorphosis were to occur in this

| species, it would be manifest during the summer drought, and presumably the

| specimens had not moulted since then. But I reject this possibility for the

following reasons:

(a) the specimens are adult;

(b) the intestine is usually filled;

females, all over the body in the males;

(d) the general morphology of the animals is very stable, anomalities virtually do

not occur; only the arrangement of the spines is somewhat variable, but

Isotomidae are usually not very stable in their chaetotaxy; the dental chaetotaxy is

very stable;

(e) eyes, mouth parts, and furca are very constant in shape, and do not present a

reduced aspect (only the mucro is a bit short);

(f) no “normal”, “female-type”’ males are found in the material;

(g) ecomorphosis due to parasitization seems excluded, since no difference could

be found in either sex between heavily parasitized specimens and animals that

were virtually free of Sporozoa.

Since care was taken to ensure random selection of both sexes for mounting on

slides, the sex ratio 9/4 = 1.2 is approximately correct.

Isotomiella minor (Schaffer, 1896)

Material: sample 4: 35 and 8 juv.; 7: 1 juv.; 29: 4 juv.

I. minor is, either primarily or secondarily, a cosmopolitan species. It is only

surprising that the species is so rare in Crete.

Isotoma notabilis Schaffer, 1896 (Fig. 40e)

Material: sample 4: 1 juv.; 5: 5 juv.; 6: 29 and 5 juv.; 19: 1 juv.; 21: 8 juv. + 20

ex. in alcohol; 24: 18 juv.; 25: 16 juv.; 29: 1 juv.; 35: 129 and 10 juv.; 37: 3 juv.; 49:

| juv.

An extremely common and widespread species.

Isotoma vaillanti Murphy, 1958

Material: sample 14: 8 juveniles.

Note: there can be no doubt that J. vaillanti is an ecomorph; but since it is

impossible to guess of which species, I continue provisionally to use this name.

After its description from mountainous Algeria, /. vaillanti was mentioned by

Cassagnau (1971a) from the Greek mainland, i.e., the southern part of the Pindos

range: Mt. Timfristos (Evritania).

W. N. ELLIS: Collembola from Central Crete 303

Isotoma olivacea Tullberg, 1871 s. auct. (a.o. Gisin, 1960) (Fig. 40 f)

Material (all immature): sample 23: 7; 31: 1; 41: 1.

The material is comparable to immatures of J. ‘‘olivacea’’ from western Europe.

Since Dr. A. Fjellberg is preparing a revision of this group, I limit myself to giving

a drawing of the antennal organ I (Fig. 40 f; juvenile specimen!).

Isotoma viridis Bourlet, 1839

Material: sample 11: 2 juv.; 19: 4 juv.; 23: 1 juv.; 24: 5 juv.; 26: 3 juv.; 32: 1 juv.;

34: 1 juv.; 36: 2 juv.; 39: 1 juv.; 40: 19 and 7 juv.; 44: 1 juv.

The high proportion of immatures in /. viridis and Isotomurus palustris is striking.

Isotomurus palustris (Muller, 1776)

Material: the present material comprises only immature specimens: sample 12:

Egadi: 09: 7221: 8; 25: 1; 26: 1: 29: 1: 35: 9:56: 4937: 13 409 15439194422;

45:1.

The larger specimens agree satisfactorily with the current interpretation of

palustris s. str.

Isotomurus spec.

Isotoma spec. Ellis, 1974.

Material: sample 26: 7 juv.

Discussion. The material in its unmistakable pigmentation agrees with the single

(also immature) specimen that I described briefly from Rhodes. My guess that this

specimen might be an ecomorph of an /sotomurus is corroborated by the fact that

the Kriti specimens have much longer and more differentiated setae. The

lasiotriches arrangement on abd „_, is, as far as I can establish 1, 3, 1. The labrum

is typical for the genus; mucronal seta absent; lateral flaps of ventral tube with

3+3 setae.

ONCOPODURIDAE

Oncopodura crassicornis Shoebotham, 1911 (Fig. 41a)

Material: sample 6: 25 ; 25: 19; 26: 19; 27: 29; 43: 19, lg and I juv.; 44: |

juv.; 45: 1 juv.

Note. The species has already been recorded from Rhodes (Ellis, 1974). The

structure of antennal organ I underlines the affinity of the Oncopoduridae to the

Isotomidae (Fig. 41 a).

304 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

TOMOCERIDAE

Tomocerus lamelligerus (Börner, 1903) (Fig. 41 b-g)

Material: Sample 25: 4 specimens, including at least 1 female and 1 male (the

other two may not be fully mature).

Description. Total length 2.2-3.0 mm. Pigmentation very weak, mainly on pleura

of th,, anterior margin of th,, coxae, and frontal region of head. Scales normal,

tinged dark brown.

Antennae 0.8-0.9 times body length, first two articles bearing scales. Eyes 6 on a

triangular black eye patch. Labral chaetotaxy typical: 4/5, 5, 4; labral margin with

4 recurved spines. Mouth parts seemingly normal, not dissected. “Prostheca” not

visible in transparency. Cephalic chaetotaxy seems unstable; anteriorly a group of.

about 6 setae, discally about 6+6 setae. Rear margin of head with a row of many

short spines.

Trochanteral organ of P, composed of one stiff seta on trochanter; one identical

seta basally on femur (Fig. 41 e). Feet covered with scales up to the tibiotarsi.

Tibiotarsal spines present, but difficult to count, because they are caducous and

intergrade between strong setae.

Unguis with strong pseudonychia, and on inner lamella 4 teeth plus a slightly

stronger basal tooth. Unguiculus without apical filament or outer tooth, but with a

small tooth on the anterior of the two inner lamellae only (Fig. 41 b). Tenent hair

strong, tubiform.

Abdomen and thorax with lasiotriches lacking accessory scales at the base.

Ventral tube with many setae and scales. Retinaculum quadridentate, without

scales and with only one seta on the corpus (Fig. 41 c).

Manubrium anteriorly with scales, laterally with a single row of strong setae,

posteriorly with two broad longitudinal bands of many setae. Dental spines (Fig.

41 g) in two specimens 3, 2/4, 1; in the adult female 4, 2/, 1, 3, 1, and in the male 5,

2/1, 1, 3, 1. The basal spines are arranged in an irregular double row (Fig. 41 d).

The spines are brown, striate, and complex. Mucro with one apical and one

ante-apical tooth, and two basal teeth, the outer one having an accessory tooth.

Intermediate teeth are absent. The two basal teeth each have a proximal lamella;

the ante-apical tooth gives rise to a very short and low outer lamella, as well as to a

conspicuously high, undulating inner lamella that stretches as far as the inner basal

tooth (Fig. 41 f). Dentes basally with one pair of inner modified scales, about half

as long as the accompanying normal scales.

Discussion. The species is well characterized by the combination of the

presence of strongly compound dental spines and a mucro without intermediate

teeth but a high lamella. The only species in the European fauna with these

characters is 7. lamelligerus (Borner, 1903), described from Calabria, and later

recorded from Bulgaria (Rusek, 1965, citing an inaccessible paper by Drenowski,

1937; the material was identified by Stach) and Yugoslavia (e.g. Cvijovic, 1972). A

similar mucro is described in T. terrestralis (Stach, 1922) which has the dental

spines simple, and the North American (Californian) species 7. teres Christiansen,

1964, which differs by having the dental spines essentially simple and the tenent

hair pointed or only weakly clavate.

W. N. ELLIS: Collembola from Central Crete 305

Fig. 41. Oncopodura crassicornis Shoebotham. a, ventral aspect of ant,. Tomocerus lamelligerus (Bòrner).

b, claw of P,; c, retinaculum; d, arrangement of dental spines; e, trochanter and basal part of femur of

P, with trochanteral organ; f, mucro; g, dental spines, inner view

306 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Since Borner’s description is exceedingly concise, and consequently the

identification of the Greek material was not beyond doubt, I compared the Cretan

specimens with a topotypical specimen, kindly loaned to me by Dr. R. Dallai,

from Sienna. This specimen, an immature collected in Palmi, Calabria on 27

March, 1972, was mounted, and the chaetotaxy could not be studied. However, all

other structural details agreed with the Greek specimens. The strong tooth on the

inner lamella of the unguiculus mentioned by Börner is less pronounced in the

specimen seen, much as in the Cretan examples.

CYPHODERIDAE

Cyphoderus albinus Nicolet, 1841

Material: 2 specimens from sample 49.

The species has a very large distributional area covering Europe.

Cyphoderus spec. cf. maroccanus (Delamare, 1948) (Fig. 42 a, b)

Material: sample 5: 1g and 1 presumably juvenile specimen.

The proportionally short mucro (1/7 times dens) places these specimens within

the group singled out by Delamare as Cyphoda.

In one specimen (the male) there are 5 outer and 5 inner scales on the dens, but

I am not completely certain about the number in the other specimen; in both

specimens the internal apical scale is at least twice as long as the exterior one. The

mucro has a vestigial subapical tooth (Fig. 42 a). The unguis has only one of the

basal teeth strongly developed. An unpaired tooth is lacking in most claws, but

was seen in one instance (Fig. 42 b).

The related species C. grassei (Cassagnau & Delamare, 1948), differs by the very

narrow basal tooth on the unguis, and in having a distinct tunica.

It is very hard to allocate the two specimens: the shape of the mucro and the

distributional evidence point towards identification as grassei, but the shape of the

claw suggests maroccanus.

TROGLOPEDETIDAE

Troglopedetes cretensis n. sp. (Fig. 42 c-e, 43 a-e)

Material: sample 35: 1; 38: 1; 44: 2; 45: 1; 46: 3; all specimens of unknown sex,

perhaps largely immature, except one 9, the holotype, from sample 46.

Description. Total length of largest specimen, the holotype, 1.0 mm; remaining

specimens 0.7-0.8 mm.

Colour white, no trace of pigment, scales in large specimens with a brown hue.

Habitus paronelloid, with the conspicuous long and stiff furca.

Antennae twice head diagonal. Ant. 1: 2: 3: 4 = 3.5: 6.5: 4.8: 10.0. Ant, divided

into two equal parts by a distinct suture. Ant,_, dorsally with some scales. Ant,

dorsally in basal position with 4 small spines, ventrally with 2 small sensillae and 3

W. N. ELLIS: Collembola from Central Crete 307

Fig. 42. Cyphoderus cf. maroccanus (Delamare). a, mucrodens; b, claw of P,. Troglopedetes cretensis n.

sp. c, cephalic chaetotaxy; d, diagram of body chaetotaxy; e, latero-distal hair cover of abd,

308 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Fig. 43. Troglopedetes cretensis n. sp. a, lasiotriches and accompanying structures of abd,; b, claw of P,;

c, lasiotriches and accompanying structures of abd,; d, mucro; e, lasiotriches and accompanying

structures of abd,. Pseudosinella paprivata n. sp. f, inner carina of unguis; g, claw of P,; h, labial

chaetotaxy; i, eye patches of three specimens at the same magnification (interference contrast

microscope), to show variability in number of ocelli

W. N. ELLIS: Collembola from Central Crete 309

sense hairs. Ant,_, with many smooth sense hairs of variable length, and with a

small number of sensillae in the form of a very short spine (type F of Christiansen,

1958); sensillae of type B — short sense hairs, apically dilated and cleft — not

found. No apical retractile papilla. Mouth parts normal for the family group.

Labral margin with (4?) globular thickenings that I could not analyse. Prelabral

setae 4/5, 5, 4. Labral chaetotaxy quite comparable to that of Lepidocyrtus:

M,M;rEl,L,; not only r but also 1, in the form of a short spine.

Ocelli absent. For cephalic chaetotaxy, see Fig. 42 c.

Feet without scales. Trochanteral organ composed of 9 smooth spines. Femora

and tibiae with plumose hairs and a small number of sensillae (?) of type F referred

to above. Tibiotarsus, with smooth supra-empodial hair; tenent hair distinctly

clavate. Unguiculus rather broad, untoothed, unguis with well-developed basal

teeth and one strong unpaired tooth (Fig. 43 b).

Ventral tube with a limited number of plumose setae at anterior and posterior

faces; anterior face not easily studied, posterior with 3+3 type F sensillae and at

least one pair of scales. Retinaculum quadridentate, corpus without scales, having

only one median hair.

Manubrium: dens: mucro= 10.0: 5.8: 1.7. Manubrium anteriorly and laterally

scaled, posteriorly with ciliate setae. Dens anteriorly with large scales — the distal

ones as long as the mucro — posteriorly with ciliate setae and postero-internally

with a row of at most 15 simple smooth spines. The spines are distally slightly

longer than basally; in the middle of the dens they are half as long as width of

dens. Mucro almost 5 times as long as maximally thick, with apical and subapical

teeth, and postero-basally 3 or 4 sawteeth (Fig. 43 d).

Body chaetotaxy oligochaetotic (Fig. 42 d). Each segment with one pair of

pseudopores. Lasiotriches: one pair on head, 2, 3, 3, on abd,_, showing

accompanying modified scales at their bases. Many microchaetae, a number of

which on abd, are conspicuously long (Fig. 42 e, 43a, c, e).

To give an impression of the variability of some characters, Table 7 shows some

data on the length of the specimens, the number of dental spines, and the number

of dorsal teeth on the mucro.

Discussion. The new species is very close to 7. cavernicola Delamare, 1944,

described in great detail from some caves in Portugal. Differences are: the mucro

is narrower in cavernicola with only two basal teeth (apparently constantly) and the

Table 7. Total length and numbers of dental spines and dorsal teeth of mucro in Troglopedetes cretensis

n. sp.

Length (mm) dental spines mucronal teeth

1.0 15 + 15 3+3

0.8 12 + 12 3+4

0.8 2412 3 +3

0.8 10 + 10 3+4

0.7 9 +12 3+4

0.7 9 +10 2 +3

0.7 8 +9 SE

0.6 12 + 12 3 +3

310 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

tenent hair in cavernicola is very weakly clavate. There are also differences in some

chaetotactic details although in interpreting Delamare’s drawing of the chaetotaxy

of cavernicola one must include the pseudopore, which was not known at that time.

The th, of cavernicola are stated to show 3-4 insertions on each side within the

groups of six setae; in cretensis this is 1+the pseudopore. In th; cavernicola

has 3+3 insertions within the groups of 3; in cretensis this is 0 + the pseudopore.

In abd, cavernicola has 4 strong macrochaetae lateral to the anterior lasiotriches; in

cretensis there are 5.

ENTOMOBRYIDAE

Heteromurus major (Moniez, 1889)

Material: sample 11: 1iex.; 13: 2iex. 219: Ixjuw.2522iex (13), 267 3exa(lauye

18); 27:4 ex. (29); 29:4. ex. (19,25); 31: 2 ex. (18.)53322 ex (le and 1 juv.); 38:

1 juv.; 43: 1 juv.

Discussion. The material is in good agreement with the specimens recorded

earlier by me (1974) from Rhodes. Labial chaetotaxy is again MMREIL, and this

very constantly. The smooth seta 1,, also drawn by Hüther (1970), is peculiar.

These results are somewhat at variance with the information given by Martynova,

Chelnokov & Rasulova (1974), who report a rather strong variability in the labial

chaetotaxy of southeastern Russian major (MMR-IL, MMREIl, MM/m ...l,

MMRELI) in which I, seems to be the only seta which is constantly smooth.

Heteromurus sexoculatus Brown, 1926, bona species

Material: sample 5: 1 ex.; 6: 1 ex.

Discussion. The good luck of finding two more specimens of H. sexoculatus gives

me the opportunity to revise my earlier opinion that sexoculatus is a variety of

major, since they showed two characters which could be confirmed in the

specimen from Rhodes.

The labial chaetotaxy is not completely stable, but either M,mz,rel,l, or

m,m,rel,l, (m, is about 2/3 the length of m,). In one specimen this occurs

asymmetrically; in another, seta r is symmetrically absent. However this may be,

the number of smooth setae is much higher than in H. major. In conjunction with

this the setae at the ventral face of the head are smooth or almost so, only

becoming gradually more serrate towards the proximal and lateral regions.

The posterior face of the manubrium bears some pairs of stiff erect and virtually

smooth setae, about 1.5 times the length of the normal serrate setae. The number

of these hairs is difficult to establish, since they tend to fall out easily, and the

number is perhaps variable; in any case, one pair is situated very proximally, and

another 3/4 of the distance from base of manubrium; in one specimen such a seta

was also observed on the dental base. Setae of this type are missing in major, but

are present in nitidus and related species. No smooth setae are present on the

tibiotarsi. Body chaetotaxy as in major.

W. N. ELLIS: Collembola from Central Crete 311

I asked Mr. P. N. Lawrence of the British Museum whether the types of

H. sexoculatus were still extant, and he informed me that the whereabouts of

Brown’s collection are unknown.

Heteromurus nitidus (Templeton, 1835)

Material: sample 9: | juv.; 24: 1 juv.; 38: 1 ad.

The species has been recorded throughout Europe.

Pseudosinella octopunctata Borner, 1901

Material: sample 4: 29 and 2g ; 5: 1 ex.; 6: lex; 10: 19; 11: 3 ex. (1 juv.); 12: 1

E10: 20: 19; 26 lers; 292 ex. (1@ ); 30: Tex; 3274 ex: (19); 36: 2iex.

(1g ); 42: 2 ex.; 45: 1 ex.; 48: lex.

The species has already been recorded from Rhodes (Ellis, 1974). Recently, Da

Gama (1973) recorded it from Turkey.

Pseudosinella albida (Stach, 1930)

Material: sample 9: 1 ad.; 25:2 9 and 1 juv.; 31:22 ;36:1¢.

Discussion. The material agrees with the description given by Stomp (1971), but

the pigmentation is much stronger, as is normal for Greek material (Stomp, 1972).

There is a contradiction between Stomp’s descriptions of 1971 and 1972 as to the

presence of seta s on abd,. In the present material seta s is indeed present, as

stated in the earlier publication. The species seems to be common in Greece (see

also Da Gama, 1973), and is also known from Spain and Italy.

Pseudosinella imparipunctata Gisin, 1953

Material: sample 4: 1 ; 11:19; 14: 1 ex.; 16: 1 ex.; 21: lex. 34: 19; 35: .Ixjuw.;

43:19.

Discussion. The material agrees with the literature in most currently available

characters: labial chaetotaxy MrELL, chaetotaxy of abd, pABQgq, seta s lacking

on abd,, body macrochaetae R111/10/030+2, no scales on ant,. The unguis is as

described by Gisin (1953), closely resembling that of alba (Packard, 1873).

However, the configuration of the eyes is not completely in agreement with the

original description. Most of the specimens show two distinct, well-pigmented

eyes, subequal and separated by slightly less than their diameter. In the two 9

mentioned above, the eyes on one side are much closer to each other, almost

touching (and again subequal). In the immature from sample 35, one side shows 3

subequal, well-spaced eyes, lying in a straight row.

P. imparipunctata was described from Switzerland, and has been recorded from

Spain (Selga, 1973), Portugal (Da Gama, 1961), Germany (Hüther, 1961), and

Bulgaria (Kosarov & Zonev, 1966).

312 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Pseudosinella paprivata n. sp. (Fig. 43 f-i, 44 a, b)

Material: sample 5: 39, 1g , 2 immature g , 7 specimens of unknown sex, partly

juv.; 9: 1 juv. (identification not definite, excluded from type-series); 27: 19, 4

juv.; 44: 2 juv. Holotype isag from sample 5.

Description. Total length 0.6 mm. Habitus normal. Scales thin. Some pigment

only in and around the poorly delimited eye patch.

Antenna 1.4 times head diagonal. Ant 1:2:3:4 = 3.8:5.2:5.9:10.0. No scales on

antenna.

Eye number-very difficult to establish (cf. Fig. 43 i). In samples 27 and 44, 3

pigmented corneae are visible; in sample 5 (and in the doubtful specimen from

sample 9) mostly 2+2, but sometimes virtually 1 +2 or 1 +1 ocelli can be counted. :

Since no other distinction could be made within the material, these differences, if

real, are not considered important. Labial chaetotaxy in all specimens except 29

and | juv. from sample 5: M-ELL; in the three specimens mentioned: MrELL.

Evidently, the reduction of seta r has led in most specimens to its total suppresion.

The row of setae anterior to the prelabial setae are also ciliate, though less than

the prelabial ones, just as in P. ciliata Ellis, 1974 (Fig. 43 h).

Feet without scales. Empodium lanceolate. Tenent hair feebly clavate. Unguis

with comparatively weak basal teeth, inserted rather high, and two comparatively

strong distal teeth (Fig. 43 f, g).

Abd, with a rather special chaetotaxy, formulated as: --Bqq. Not only are p anda

constantly absent, but also the exterior of the two microchaetae in front of the

lasiotriche is absent, and at its site there is a small scale (Fig. 44 a). Abd, without

seta s; seta a well ahead of e (Fig. 44 b). Arrangement of macrochaetae:

RO11/10/0101 +2.

Mucro with subequal teeth.

Discussion. The constant absence of both a and p on abd, places this species in

an isolated position among all European species described so far. It is generally

agreed that Pseudosinella is derived polyphyletically from Lepidocyrtus, largely

through some still extant stem species. In the present case, however, I do not know

of a Lepidocyrtus species from which this form could have been derived.

Lepidocyrtus lignorum (O. Fabricius, 1775) (Fig. 44 c, d)

Material: sample. 6: 1 ex.; 20: 1 ex.; 21: 1 juv.; 23: 1; 25: 1; 30: 3 (2 juv.); 31: 3

(19); 36: 1 juv.; 37: 14 ; 38: 4 (29); 45: 1 juv.; 50: 3 (1 juv.).

Discussion. The specimens are in good agreement with West European material

of this species, except that in some of the largest specimens the labial chaetotaxy

shows some (symmetrical) duplication, resulting in the presence of an accessory

seta M,’ and two setae R’, R” (Fig. 44 c). But most specimens show the normal

labial chaetotaxy M,M,REL,L,. Other characteristics of the species are: scales

present on femora and tibiae and on ant,_,, body chaetotaxy R001/00/0101 +3,

chaetotaxy of abd,: paBqg, seta s lacking on abd.

313

W. N. ELLIS: Collembola from Central Crete

Sui /

n q 4

O | no 7}

a, AD D

if / 3

SP ber a

Fig. 44. Pseudosinella paprivata n. sp. a, chaetotaxy of abd,; b, anterior lasiotriches of abd, with

associated microchaetae. Lepidocyrtus lignorum (O. Fabricius). c, labial chaetotaxy; d, ventral tube,

anterior view. Lepidocyrtus lignorum forma ? e, pigmentation of abd,_,. Lepidocyrtus serbicus Denis. f,

ventral tube, anterior view

314 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Lepidocyrtus lignorum forma? (Fig. 44 e)

Material: sample 25: 2 specimens.

Discussion. The specimens agree morphologically in all details with L. lignorum

(which occurred in the same sample). However, they show a pigmentation pattern

I do not remember ever having seen in lignorum: two broad spots on abd,, almost

filling the dorsum of that segment, leaving only a median uncoloured spot and two

smaller marks postero-laterally on abd,. Also pigmented are the frontal ocellus

and — weakly — the antennae. Eye patches black as usual.

It is interesting to regard this material in relation to the “lignorum var.?”

mentioned by Da Gama (1973) from the Peloponnesus and the Ionian islands.

Here, however, abd, and abd, are entirely pigmented, as well as the posterior |

corner of abd,. The relation between these two forms and L. instratus Handschin,

1924, from Switzerland and east-central Europe, merits closer study.

Lepidocyrtus serbicus Denis, 1936 (Fig. 44 f)

Material: sample 15: 1 ex.; 19: 1& ; 21: 19; 24: 29; 25: 13 ex. (at least 3 adults

and 19); 27: 8 ex. (at least 29 and 2g ); 35: 2 ex. (19); 37: 1 ex.; 44: lex.

Discussion. The material is in good agreement with the descriptions, but some

clearly immature specimens seem to lack seta M, on the labium. Otherwise, the

characters of serbicus are stable: body chaetotaxy R111/00/0101 +2; antennae and

feet without scales; labial chaetotaxy M,M,REL,L,; chaetotaxy abd,: p-B-q,; seta

s lacking on abd, (and seta e rather longer than the others).

Lepidocyrtus serbicus, described from Yugoslavia, has been recorded from

Poland (Szeptycki, 1967), and Romania (Gisin, 1965). Although some references

are ambiguous, on the European mainland serbicus seems to be usually caverni-

colous.

Seira graeca Ellis, 1966 (Fig. 45 a)

Seira pillichi graeca Ellis, 1966.

Seira ferrarii; Ellis, 1974.

Material: sample 38: 19. |

The study by Dallai (1973), unfortunately received only at the end of 1974,

convinced me of the importance of the chaetotactic details of th,, and forced me

to re-examine my material from central Greece and Rhodes. The posterior group

of macrochaetae on th, proved to consist of a tight group of 7+3+5 macrochaetae

(Fig. 45 a). Only in two not fully mature specimens from Rhodes was 6+3+5

found asymmetrically.

Dallai (1973) cites two differences between S. ferrarii Parona, 1888, and S.

sacchii Parisi, 1969: (1) ferrarii has 6+3+5, sacchii 7+3+5 macrochaetae

posteriorly on th,, and (2) ferrarii has 8, as against 10 macrochaetae sacchii has in

the transverse row between eye patch and frontal ocellus. S. graeca and ferrarii

differ only in th, chaetotaxy, and thus the only remaining difference between

graeca and sacchii would be the interocular macrochaetae. As a matter of fact, in

W. N. ELLIS: Collembola from Central Crete 315

5 I o |

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Fig. 45. Seira graeca Ellis. paratype. a, chaetotaxy of th,_,. Seira spec. b, chaetotaxy of th,; c, dorso-

median chaetotaxy of abd,

graeca this number is 8+8 or, almost as frequently, 8+9. The difference between

graeca and sacchii is thus very small indeed. S. sacchii is so far known only from Italy.

Seira spec. (Fig. 45 b, c)

The habitus of the living animal is unknown, but in chaetotactic details the

specimen is very close to S. dagamae Dallai, 1973, except for one clear feature: th,,

which has 14 paramedian macrochaetae in dagamae, has 15 in the present speci-

men; in particular, the group postero-internally from the pseudopore has one

macrochaeta more. I cannot evaluate this specimen at present (Fig. 45 b).

For convenience, I present below a practical key to the European Seira of

which the chaetotaxy is known.

1. Discal group of th, with 4 setae; anterior row on abd, simple ............ 2

— Discal group of th, with more than 4 macrochaetae; anterior row on abd,

COBE ee ARIE NEN MSI 7

Ps Raramedian:group ofiabd,; with Smacrochaetae: i... 3

— Paramedian group of abd, with 4 macrochaetae ....................... 5

316 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

3. Posterior sroup'ofith>;withilSimacrochacta Re RE 4

22 Posterior group of th, with 18-20'macrochaetae RO

Seer ERA Ica ii AIRSHIP ea rnd ts Var Fl hed mor cota Am a unnamed species (see p. 315)

5. Posterior field of th, with 14 macrochaetae ............ ferrarii Parona, 1888

—= Posterior tield;on th, witholSimacrochactae nnn 10E ea 6

6. Between eye patch and frontal ocellus, 8-9 macrochaetae .................

EEN EEEN ORR B RE le Me en sacchii Parisi, 1969

7. th, with postero-medial group (behind pseudopore) of 7-9 macrochaetae

u re MEL ara UPR do cS SITO AE OS RE Le lusitanica Da Gama, 1964

— th, with postero-medial group of 4 macrochaetae ........................

PIRRO SSR RITA EEN squamoornata (Schtscherbakow, 1898) |

— th, with postero-medial group of 3 machrochaetae .................... 8

Discalfield'of.th7,with:Wmacrochactae RR

RA a Rene pallidipes Reuter, 1895, sensu Loksa & Bogojevic, 1970

— Discalfield'of th, with LOM maerochaetae WEE Ae es eee 9

Anterior macrochaeta of intermediate group of abd, twice as far from

posterior seta in this group, as from its lateral neighbour ..................

SOUS ith au PR ARDA I Re RRL I een saxatilis Gisin & Da Gama, 1962

— Three timesithis.distanee®@. = e Ne dollfusi Carl, 1899

S. arenaria Da Gama, 1966, banyulensis Denis, 1924, italica (Cassagnau &

Delamare, 1953) and petrae Jacquemart, 1973, are considered synonyms of S.

ferrarii.

Entomobrya muscorum (Nicolet, 1841)

Material: sample 8: one adult specimen.

The ‘specimen is very pale, and the meagre pigmentation resembles most

closely that of E. pazaristei Denis, 1936. However, the extremely long antennae

of the specimen are more in favour of an identification as muscorum. The species

has been recorded from some central European countries, Madeira, and North

and South A merica (South, 1961).

Entomobrya handschini Stach, 1922

of unknown sex and 1 juv.; 26: 1 juv.; 27: 1 juv.; 38:4 9 and DER

This characteristic species, one of the few truly epigeic species encountered in

Crete, was described from Hungary, and has since been recorded from many

countries in south-eastern Europe and from Spain.

W. N. ELLIS: Collembola from Central Crete 317

Entomobrya multifasciata (Tullberg, 1871)

Material: sample 15: 3 adults; 22: 7 9, 2g, and I juv. + 7 ex. in alcohol; 27: |

juv.; 38:1 0,14, 1 juv., and 3 adults of unknown sex, + 7 ex. in alcohol.

This common European species has already been recorded from Rhodes.

Entomobrya cf. multifasciata (Tullberg, 1871)

Material: sample 23: 1 9, 1g and 2 specimens of unknown sex; 32: | ex.;

33:1g and3juv.;42:1g and 2 juv.; 49: 2 ex.(1juv.).

All these specimens are wholly or almost entirely depigmented; the few traces of

pigment and the few reliable morphological characters do not contradict identi-

fication as multifasciata.

NEELIDAE

Megalothorax incertus Borner, 1903

Material: sample 23: 1 ex:; 25: 2 ex.; 27: 1 ex.; 35: 1 ex.: 36: 2 ex.2'43°2 ex: 44:

Sex. 45: 2'ex.

The species has already been recorded from Rhodes. I am not completely

convinced as to the identification of the specimen from sample 23.

SMINTHURIDIDAE

Sminthurides malmgreni (Tullberg, 1876)

Material: sample 26: 1 9,3% ‚and 5 juv.

The species has been recorded from most European countries, North America,

and Japan; records from the southern Palaearctis include Portugal, Spain, Yugo-

slavia, Romania, N. Africa, and the Azores.

Sphaeridia pumilis (Krausbauer, 1898) sensu Massoud & Delamare

Deboutteville, 1964

Material: sample 6: 1 9; 7:2 9 andlg;10:4juv.; 17:14; 18:5 9,33 ,and9

juv.; 19:19;21:1g and2juv.; 23: 6 9 and 2 g, and about 120 ex. in alcohol; 24:

Bed? 7,25. 3795,26: IO 27-2" and 2 3°: 29: 1-9 32: 10'ex. in alcohol; 33:

(RSrand 3 3 ; 34:2 9; 35: 2 9; 36:4 anda 537: 19; 41: 1 g 5 42:2 9; 43:1 9;

ls; 49: 1 juv.

This common and widespread species has already been recorded from Rhodes.

SMINTHURIDAE

Arrhopalites spec.

Material: sample 4: 1 juv.; 25: 2 juv.; 43: 1 juv.; 44: 4 juv.

The material is too young to permit speculation about the species identification.

318 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Sminthurinus aureus (Lubbock, 1862)

S9,3g,and4juv.;43:1 9.

Discussion. The identification of isolated immatures (samples 4, 23, 24, and 36)

can of course only be conjectural. The species is widespread throughout Europe.

None of the specimens shows the “typical” yellow coloration; all are marked

with blue, usually well-delimited patches, which in some instances cover the

major part of the body. The general habitus is rather similar to that of S. reticulatus

Cassagnau, 1964, from the Pyrenees.

Sminthurinus spec. aureus group

Material: 3 immatures from sample 18.

Sminthurinus alpinus bisetosus n. ssp. (Fig. 46 a, b)

S. spec. near alpinus Gisin, 1953; Ellis, 1974.

Material: sample no. 970.010 from Rhodes, Lindos, 1 g and 1 juv.; from Kriti,

sample 19: 2 juv.; 21: 14 ; 23: 4 juv.; 24: 1 juv.; 27: 6 juv.; 3l: 13 543: 1x ;44: 1 9.

Holotype: the female from sample 44.

In the discussion on the two specimens from Rhodes, I noted as divergent from

S. alpinus Gisin, 1953, the two (instead of one) setae on the corpus tenaculi and the

number of setae in the eye patch (‘‘two, as seen in profile” — instead of one). Now

that I have more material, the latter observation seems rather dubious. Young

specimens, with the eye spot slightly transparent, seem to have only one seta in the

eye patch. This character, interesting though it may be, is of limited practical use.

The number of setae on the retinaculum is in all adults definitely two; doubt exists

in some immatures, where the two setae differ developmentally. The wart on ant,

is in the Greek material weakly trilobed, but this can also occur in material of

typical alpinus (Fig. 46 b). In all other characters, notably the distinct dental

chaetotaxy, the Greek material resembles alpinus, and I consider it most

appropriate to evaluate the single difference in the retinaculum as a subspecific

one.

After its description from Switzerland and Austria, alpinus was recorded from

the Tatra (Nosek, 1969), Bulgaria (Rusek, 1965), and India (Choudhuri & Roy,

1972). The record from India needs confirmation.

Stenognathellus cassagnaui Yosii, 1966 (Fig. 46 c)

S. denisi Cassagnau, 1953; Ellis, 1974.

Material: sample 27: 1 9; 36: 1 juv.; 45:24.

Discussion. Dr. J.-M. Betsch, upon inspecting my only female from Rhodes,

drew my attention to some distinctions with respect to true denisi: the middorsal

seta on abd,, which is deeply bifurcated in denisi, is simple in cassagnaui. More-

W. N. ELLIS: Collembola from Central Crete 319

over the dens has two latero-distal setae (anterior to the distal whorl) in denisi,

but only one in cassagnaui (Fig. 46 c).

The Greek material is in good agreement with the species of Yosii, described

from Nepal, except that the eye patch is well pigmented in the Greek material,

and ant, is perhaps slightly less inflated. The appearance of this inflation depends

strongly however, on the observation angle. Until now, the species had not been

found again since its description.

= >

IN |

N N

Sur

Fig. 46. Sminthurinus alpinus bisetosus n. ssp. a, mucrodens; b, wart on ant, Stenognathellus cassagnaui

Yosii. c, dens. Sminthurus nigromaculatus Tullberg. d, subcoxae | and 2 and coxa of P,; the seta marked

x on subcoxa 2 is absent in S. viridis. Ptenothrix italica Dallai. e, female postabdomen

320 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Sminthurus multipunctatus Schaffer, 1896

Material: sample 15: 1 9.

This species has been recorded from southern and eastern Europe.

Sminthurus flaviceps Tullberg, 1871

Material: sample 27: 1 juv.; 35: 1 juv.; 38: 3 juv.

This species, described from Sweden, has been recorded from Germany and

most southern and eastern European countries. The record from the Netherlands

(Buitendijk, 1941) is erroneous, as I was able to verify through examination of the

original material. |

Sminthurus nigromaculatus Tullberg, 1871 (Fig. 46 d)

Material: sample 15: 23 juv.; 21: 1 juv. (cf.); 26: 1 juv. (cf.); 35: 1 9; 36: 1 3 and

3 juv.; 38: 6 juv. + 23 specimens, presumably also immature, in alcohol; 46:1%.

Discussion. Many authors have been reluctant to accept nigromaculatus as a

good species, because of the minimal set of characters available to distinguish

it from viridis (Linnaeus, 1758): a preference for drier habitats, for pollen as food

(little or no plant epidermis is consumed), and the presence of two (sometimes

one) dark spots on the postabdomen. I think that I have now found a good

morphological character. Subcoxa 2 of P, in viridis has one rather thin hair;

nigromaculatus has the same, but also anteriorly another stouter hair (Fig. 46 d).

The latter is found in nigromaculatus in the Netherlands and Rhodes (my separa-

tion of viridis and nigromaculatus in material from Rhodes is thus corroborated)

and in the present nigromaculatus from Crete. The character holds for larger

immatures, but very young immatures of nigromaculatus, which usually already

have the spots on the postabdomen, show only the single hair of viridis — but in

such specimens the number of setae on the coxa of P, is 2, instead of 4, and the

setae on the dens are fewer in number.

Comparison of the situation in S. nigromaculatus with that in other Symphy-

pleona (Allacma, Neosminthurus, Lipothrix of the same subfamily, Bourletiella and

Deuterosminthurus of the Bourletiellinae, and even Dicyrtoma of the Dicyrtomidae)

shows that in the others subcoxa 2 of P, has only one seta, which, to judge from its

appearance and location, is homologous with the single seta in S. viridis. I

therefore consider the possession of an extra seta by S. nigromaculatus to be an

apomorphous character. It is a bit surprising, however, that this character is also

present in Caprainea echinata (Stach, 1930) and C. marginata (Schott, 1893).

Within the genus Sminthurus s. str., I found that S. guthriei Stach, 1920, shows

the plesiomorphous state, but S. multipunctatus Schaffer, 1896, and S. maculatus

Tömösvary, two species which are very close to nigromaculatus, possess the

apomorphous character state.

The stability of the chaetotaxy in the coxal region of P, is great. Both viridis

and nigromaculatus are very common in the Dutch fauna and generally in Europe.

W. N. ELLIS: Collembola from Central Crete 321

Among the large material available I found only one aberrant specimen, an

adult 9 of nigromaculatus from the Netherlands in which the extra seta was absent

on one side.

The specimens from Crete are all intensely mottled with blue pigment all over

the body. This also applies to the immatures, which excludes the possibility of a

gradual darkening of adult specimens by subcutaneous accumulation of waste

products. In general appearance they have no resemblance to the nigromaculatus

from Rhodes.

It thus seems that viridis and nigromaculatus are two well-separated species

which must have evolved in a comparatively remote past. S. viridis, the most

primitive of the two, has not evolved any further and remained monomorphous. S.

nigromaculatus, on the contrary, is in a state of evolutionary radiation and the

species comprises a number of forms, many of which doubtless have not yet

reached the level of species.

Only the refined methods of beta-taxonomical approach can clarify this com-

plex. Its recognizable members are ‘’maculatus’’, the pale, two-spotted western

European nigromaculatus, the dark marbled nigromaculatus from southern Europe,

and the single-spotted nigromaculatus described from Rhodes.

Caprainea echinata (Stach, 1930)

This Mediterranean species has already been mentioned from Rhodes.

Neosminthurus natalicus Ellis, 1974

Material: sample 24: 2 juv.; 25: 2 juv.; 35: 1g and 4 juv.; 44: 4 juv.

This apparently Mediterranean species was previously known from Ibiza and

Rhodes.

DICYRTOMIDAE

Dicyrtoma (Dicyrtoma) melitensis Stach, 1957

Material: sample 46: | immature specimen.

Discussion. The identification is made with some reservation, because the

unguicular filament of P, of both feet is distinctly knobbed. The dental chaeto-

taxy, however, points to melitensis. The species has been described from Malta

and was recorded from Stromboli (Altner, 1961), Sardinia (Parisi, 1969), and the

small Italian island of Pianosa (Dallai, 1969b).

Dicyrtomina (Calvatomina) cf. articulata Ellis, 1974

Material: sample 27: 6 juv.

The material unequivocally belongs to the subgenus Calvatomina Yosii, 1966, but

is too young to be attributed with certainty to articulata.

322 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Ptenothrix italica Dallai, 1973 (Fig. 46 e)

Material: sample 31: 1 9.

The single specimen is in good agreement with Dallai’s description of italica,

from southern Italy. Since Dallai did not have females, I add a drawing of the

female postabdomen. The anal appendages are comparatively thick (Fig. 46 e).

Although Ptenothrix cavicola Cassagnau & Delamare Deboutteville, 1955, de-

scribed from the Lebanon, is very close to italica, on the basis of the few dis-

tinctive characters available (pigmentation of ant,, presence of two instead of

three teeth on lateral carina of unguis) the Cretan specimen is to be attributed

to the western species.

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ésl. Spol. zool. 36 (4): 275— 279.

W. N. ELLIS: Collembola from Central Crete 325

———, 1973a. Neue Collembolen von den Uberschwemmungswiesen Süd-Mährens. — Vést. èsl. Spol.

zool. 37 (3): 183—194.

———, 1973b. Zur Collembolenfauna (Apterygota) der Praealpe Venete. — Boll. Mus. Civ. Stor. nat.

Venezia 24: 71—95.

———, 1974. Zur Taxonomie der Tullbergiinae (Apterygota: Collembola). — Vést. ésl. Spol. zool. 38

(1): 61— 70.

Salmon, J. T., 1962. New Collembola from 83 deg. South in Antarctica. — Trans. R. Soc. N.Z. [Zool.] 2

(18): 147—152.

———, ‘1964-65. An index to the Collembola. — Bull. R. Soc. N. Zeal. 7 (1—3): 1—651.

Selga, D., 1962a. Tres especies nuevas de colémbolos del Puerto de Navacerrada (Guadarrama). —

Publnes Inst. Biol. apl. Barcelona 33: 33—41.

———, 1962b. Proisotoma (Collembole) de los suelos del Guadarrama con descripcion de una nueva

especie del mismo género. — Boln R. Soc. esp. Hist. nat. [B] 60: 69—76.

———, 1963. Cuatro especies nuevas de Colémbolos de la Sierra de Guadarrama. — Publnes Inst.

Biol. apl. Barcelona 35: 83—96.

———, 1966. Descripciön y comentarios ecolögicos de cuatro nuevas especies de colémbolos. — Boln

R. Soc. esp. Hist. nat. [B] 64: 145—160.

———, 1973. Catalogo de los Colémbolos de la Peninsula Ibérica. — Graellsia 26: 133—284.

South, A., 1961. The taxonomy of the British species of Entomobrya (Collembola). — Trans. R. ent.

Soc. Lond. 113 (13): 387—416.

Stach, J., 1947. The apterygotan fauna of Poland in relation to the world-fauna of this group of insects.

Family: Isotomidae. — Acta monogr. Mus. Hist. nat. 1947: 1—488, pls. 1—53.

Stomp, N., 1968. Tetracanthella hygropetrica luxemburgensis n. ssp. de la région du „Gres de Luxem-

bourg” (Insecta, Collembola, Isotomidae). — Bull. Mus. natn. Hist. nat. [2] 40 (4): 734— 741.

———, 1971. Contribution à l’etude des Pseudosinella endogés. Espèces européennes de Pseudosinella

à 5+5 yeux. (Collembola, Entomobryidae). — Revue Ecol. Biol. Sol 8 (1): 173—188.

———, 1972. Deux nouvelles espèces de Pseudosinella endogés d’Europe. (Collemboles, Entomobryi-

des). — Revue suisse Zool. 79 (1): 279—286.

Szeptycki, A., 1967. Fauna of the springtails (Collembola) of the Ojców National Park in Poland. —

Acta zool. cracov. 12 (10): 219—280, pls. 17—22.

Thibaud, J. M., 1967. Description d’une espèce nouvelle de Collembole Acherontiella cassagnaui n. sp.

— Annls Spéléol. 22 (2): 393—400.

Yosii, R., 1960. Studies on the Collembolan genus Hypogastrura. — Am. Midl. Nat. 64 (2): 257—281.

———, 1965. On some Collembola of Japan and adjacent countries. — Contr. biol. Lab. Kyoto Univ.

19: 1-71.

———, 1966a. Collembola of Himalaya. — Coll. Arts Sci. Chiba Univ., Nat. Sci. Ser. 4 (4): 461—531.

———, 1966b. On some Collembola of Afghanistan, India and Ceylon, collected by the Kuphe-

expedition, 1960. — Res. Kyoto Univ. sc. Exp. Karakoram Hindukush 1955, 8: 333405.

—— —, 1972. Collembola from the alpine region of Mt. Poroshi in the Hidaka Mountains, Hokkaido.

__— Mem. natn. Sci. Mus. 5: 75—99.

Zivadinovié, J., 1972. Species of Collembola (fam. Poduridae, Onychiuridae and Isotomidae) as mem-

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diënjak srp. Akad. Nauk 25: 175—226, tab. 1—3.

326 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 8, 1976

Acheroxenylla nov. gen.

affinis Borner

afurcata Denis

albertinae n. sp.

albida Stach

albinus Nicolet

americanus Denis

anthrenoidea n. sp.

armata Nicolet

articulata Ellis

asigillata Borner

aureus Lubbock

bisetosus n. ssp.

cassagnaui Yosii

crassicornis Shoebotham

237-238

225-227

259-266

291, 292-296

311

306

284

275, 278-280

245

321

267-269

318

318, 319

318-319

303, 305

cretensis n. sp. (Acheroxenylla) 236, 238-242

cretensis n. sp. (Neanura)

cretensis n. sp. (Troglopedetes)

critica n. sp.

debilis Cassagnau

decipiens Steiner

dubius Krausbauer

echinata Stach

flaviceps Tullberg

gibbosa Bagnall

gisini Strenzke 243,

graeca Ellis

277-278

306-310

230-231

289, 290-292

266-267

269

321

320

245

245, 247-249

314-315

granulata Cassagnau & Delamare

Deboutteville

handschini Stach

hygropetrica Cassagnau

imparipunctata Gisin

incertus Borner

italica Rusek (Mesaphorura)

italica Dallai (Ptenothrix)

krausbaueri Borner

ksenemani Stach

lamelligerus Borner

274-276, 278

316

282, 283

311

317

231

3198322

231

283, 285-286

304-306

libanensis Cassagnau & Delamare

Deboutteville

lignorum O. Fabricius

linnaniemii Denis

longiseta Caroli

major Moniez

malmgreni Tullberg

manubrialis Tullberg

269-272

312-313

282

279, 280-281

310

317

249

marchicus Frenzel 285

maritima Tullberg 244

maroccanus Delamare Deboutteville

306, 307

melitensis Stach 321

minitaurus n. sp. 271, 272-274, 275

minor Schaffer 301

minuta Tullberg 292

mucronata n. sp. 271, 272-274

multifasciata Tullberg 3107

multipunctatus Schaffer 320

muscorum Nicolet 316

nanus Ellis 283, 285

natalicius Ellis 321

navacerradensis Selga 284-285

nigromaculatus Tullberg 319, 320-321

nitidus Templeton 311

notabilis Schaffer 301, 302

octopunctata Borner 311

olivacea Tullberg 302, 303

orientalis n. ssp. 252, 253-255, 256

ossica Cassagnau 266-267

palustris Muller 303

paprivata n. sp. 308, 312, 313

parvula Schaffer 259

parvulus Stach 282-284

ponticus Stach 286-287

porcellus n. sp. 295, 296-301, 302

prolatus Gisin 232-233

pseudoghidinii Dallai 233, 235, 236

pumilis Krausbauer 317

rhodia Ellis 242-244

schoetti Von Dalla Torre 292, 295

serbicus Denis 313, 314

sexoculatus Brown 310-311

stachianus Bagnall 235

succinea Gisin 244-245

tethyca n. sp. 246, 248, 249-251, 253

thermophilus Axelson 286

tricuspis Borner 228

triglenus n. sp. 287-288, 290

trisetosus Denis 282

vaillanti Murphy 301

vernalis Carl 253

viridis Bourlet 303

xenonis n. sp. 234, 235, 237

MUS. COMP, ESOL:

ED -TSE.Z

LIBRARY

DEEL 119 AFLEVERING 9 1976

| FEB4 1977

HARVARD

UNIVERSITY

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

. DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

WARREN T. ATYEO & PAUL C. PETERSON. — The feather mite genera Zumptia

Gaud & Mouchet and Parazumptia gen. nov. (Acarina, Analgoidea),

p.327—335, fig. 1—14.

Tijdschrift voor Entomologie, deel 119, afl. 9 Gepubliceerd 29-XII-1976

THE FEATHER MITE GENERA ZUMPTIA GAUD & MOUCHET

AND PARAZUMPTIA GEN. NOV. (ACARINA, ANALGOIDEA)')

WARRENT. ATYEO

Department of Entomology, University of Georgia, Athens, Georgia 30602

and

PAUL C. PETERSON

Department of Biological Sciences, Youngstown State University, Youngstown, Ohio 44555

With 14 figures

ABSTRACT

Parazumptia gallinulae gen. nov., spec. nov., is described from Gallinula chloropus (Rallidae), Luzon,

Philippine Islands, and India; the genus Zumptia Gaud & Mouchet, 1959, is redefined; Z.

dermoglyphoides Gaud & Mouchet, 1959, is illustrated, and Z. macclurei spec. nov. is described from

G. chloropus, Luzon.

The feather mite fauna of the avian family Rallidae is varied. Each genus of

Analginae (Analgidae) and Pterolichidae associated with these birds contains

many species. Conversely, mites assigned to the subfamily Xolalginae (Analgidae)

are highly modified and each named species has been the basis of a monobasic

genus. Some of these xolalgine species have a wide host range, others are known

from single collections or from single localities. Extremes of these associations are

exemplified by Gymnalloptes pallens (Trouessart & Neumann, 1888), known to

occur on eight species of Rallidae from Eurasia, Africa, and the Philippine Islands,

and by Zumptia dermoglyphoides Gaud & Mouchet, 1959, known only from the

type series.

In the present paper three taxa will be described, a new monobasic genus and

second species of Zumptia, all discovered in collections made by members of the

Migratory Animal Pathological Survey in Southeast Asia. Including the new taxa,

there are eight species assigned to seven genera of Xolalginae from the Rallidae;

the host-parasite associations, either previously recorded (Atyeo, 1974; Gaud,

1968, 1974) or new are:

Fulica atra (L., 1758), european coot

Analloptes megnini Trouessart, 1885: Morocco

Gymnalloptes pallens (Trouessart & Neumann, 1888): Morocco, India

Gallinula chloropus (L., 1758), common gallinule, moorhen

1) Research supported by the National Science Foundation (BMS75-03394) and the Research Council,

Youngstown State University.

3277

328 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 9, 1976

Gymnalloptes pallens (Trouessart & Neumann, 1888): France, Madagascar

Parazumptia gallinulae gen. nov., spec. nov.: Philippine Islands, India (new

records)

Zumptia macclurei spec. nov.: Philippine Islands (new record)

Limnocorax flavirostris (Swainson, 1837), black crake

Beaurcournuella loculosa Gaud, 1974: Africa(1 9)

Cryptosikya protalgoides Gaud, 1971: Cameroons

Gymnalloptes pallens (Trouessart & Neumann, 1888): Cameroons

Zumptia dermoglyphoides Gaud & Mouchet, 1959: Cameroons

Podica senegalensis (Vieillot, 1817)

Gymnalloptes pallens (Trouessart & Neumann, 1888): Cameroons

Porzana parva (Scopoli, 1769), little crake

Dogielacarus uncitibia Dubinin, 1949: France, U.S.S.R.

Gymnalloptes pallens (Trouessart & Neumann, 1888): Europe, Morocco, Iran

(new record)

Porzana fusca (L., 1766), ruddy crake

Analloptes megnini Trouessart, 1885: Philippine Islands (new record)

Gymnalloptes pallens (Trouessart & Neumann, 1888): Philippine Islands (new

record)

Rallus aquaticus (L., 1758), water rail

Analloptes megnini Trouessart, 1885: Europe

Beaucournuella loculosa Gaud, 1974: France

Gymnalloptes pallens (Trouessart & Neumann, 1888): France

Five of the above feather mites, Dogielacarus uncitibia, Cryptosikya protalgoides,

the two species of Zumptia, and Parazumptia gallinulae can be considered

having special modifications. Based on collections from over half of the species of

Rallidae, these mites have extremely limited host distributions, occur sporadically,

have a low rate of infestation, and probably maintain low populations on

individual birds. It could be concluded that each of these five mite species have a

limited and specific microhabitat and that each must encounter considerable

pressure when becoming established on a new host individual.

In the descriptions below, the terminology for chaetotaxal signatures follows

Atyeo & Gaud (1966). Type materials are deposited in the University of Georgia

(UGA), the British Museum (Natural History) (BMNH), the National Museum of

Natural History (NMNH), and the collection of J. Gaud (GAUD).

Family ANALGIDAE

Subfamily XOLALGINAE

Zumptia Gaud & Mouchet

Zumptia Gaud & Mouchet, 1959: 526; Gaud, 1968: 25; Atyeo, 1974: 479. Type species: Zumptia dermo-

glyphoides Gaud & Mouchet, 1959.

W.T. ATYEO & P. C. PETERSON: Zumptia and Parazumptia 329

Diagnosis. Xolalgine mites lacking setae sR on trochanters I, II; humeral setae

short, basally dilated, apically attenuated; subhumeral setae ventral to humerals,

short, setiform; dorsal shields reduced; long, narrow metapodosomal shields

lateral to hysterosomal shield; dorsal hysterosomal gland well developed (?);

hysterosomal setae d/ absent, /1 positioned posterolateral on propodosoma;

epimerites I Y-shaped; legs with four functional segments (femora and genua

fused); tarsi I, II with whorl of four setae near midlength (/a, ra, wa, aa). Males with

dentate adanal discs; well-defined posterior lobes with at least interlobar

membrane well developed; genital organ small, reflexed; genital region sur-

rounded by circular striae; ventral setae with one pair lateral and one pair

immediately posterior to genital region; genital discs anterolateral to genital

organ; legs III equal to or greater than legs IV. Females with characteristic

epigynium, i.e., anterior margin straight, posterior margin concave.

The two species of Zumptia are known from limited materials, Z. dermo-

glyphoides from the original series of five specimens and Z. macclurei spec. nov.

from three collections from the Philippine Islands and India. The poor condition

of the two specimens of Z. dermoglyphoides available for study precludes a

complete redefinition of the genus as certain structures are not visible, e.g., dorsal

hysterosomal glands which are not apparent but are probably present.

The interpretation of the tarsal chaetotaxy for Zumptia and Parazumptia

indicates the presence of setae normally found in the Acaridae but not present in

the Analgoidea, that is, setae aa on tarsi I and II. In the Analgoidea four ventral

and ventrolateral setae occur, they are /a, ra, wa, and s; in these instances, s is near

the apex on the midventral surface and occurs on legs I-IV. In the taxa under

consideration, the fourth seta is not considered to be seta s as it occurs only on

legs I and II and all setae occur in a whorl. Another explanation for this seta would

be that it is ba, but ba in known Analgoidea occurs on the dorsal surface either in

close proximity to @ / or in the interspace between @ / and 03.

Striation patterns have not been noticeably different in the analgoid taxa, but

between the species of Zumptia and Parazumptia, there are striking differences. On

the dorsal propodosoma in the area between the posterior margin of the

propodosomal shield and the sejugal suture and including the scapular setae,

Zumptia species have the striae forming a pattern of inverted V’s and U’s; in

Parazumptia, the propodosomal striae are horizontally directed and parallel. On the

propodosomal venter, Zumptia species have the striation pattern as a series of V’s

with the posterior terminations of the more lateral striae bent toward the meson;

in Parazumptia, the striae are parallel and the posterior terminations are bent

laterad or are continuous with hysterosomal striae.

The last major difference between the two genera relates to the pretarsal

structures (compare Fig. 6 and 11). The divided central plate of Zumptia is

characteristic of numerous xolalgine genera; the modifications of the Parazumptia

pretarsus are found in Cryptosikya and Dogielacarus (Atyeo, 1974).

- Zumptia dermoglyphoides Gaud & Mouchet (Fig. 1—4)

Zumptia dermoglyphoides Gaud & Mouchet, 1959: 526—7, Fig. 12A, B; Gaud, 1968: 26.

330 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 9, 1976

The species is known only from the type series consisting of two males, two

females, and one nymph. The type host, Limnocorax flavirostris, has been

repeatedly examined by Gaud and others, but to date, there has never been any

reported success in recollecting Z. dermoglyphoides. Gaud (1968) discusses the

possibility that the L. flavirostris specimens might have been misidentified, but he

150u

200

Fig. 1-4. Zumptia dermoglyphoides Gaud & Mouchet: ventral (1) and dorsal (2) aspects of male; ventral

(3) and dorsal (4) aspects of female. Setae: cl, central; cx3, coxal: d 2-5, / 1-5, dorsal and lateral

hysterosomals; 4, humeral; sh, subhumeral

W.T. ATYEO & P. C. PETERSON: Zumptia and Parazumptia 331

suggests that this is probably not the case as the other mites in the collection are

known ectoparasites of this bird species.

The species will not be formally redescribed as the study specimens are in poor

condition — our illustrations are in reality probable reconstructions as relates to

the hysterosomal shields.

Type data. From Limnocorax flavirostris (Swainson, 1837) (Rallidae): 3 holo-

type, 1 ¢,2 9 paratypes, Yaoundé, Cameroons, April, 1958, J. Mouchet. The type

series is in the collection of J. Gaud.

Zumptia macclurei spec. nov. (Fig. 5-9)

The males of Zumptia dermoglyphoides and Z. macclurei are easily separated; in

the former species, legs III and IV are subequal and in Z. macclurei, legs III are

much larger than legs IV.

Male (holotype). Length, including gnathosoma and terminal lobes 270 u, width

108 u. Idiosoma broad, gently tapering posterior to humeral setae to deeply cleft

terminus. Dorsal propodosoma with shield small, not extending to scapular; area

between posterior margin of shield and sejugal suture with inverted V and U

striations. Dorsal hysterosoma with setae d2-d4 internal to shield margins;

terminus with deep U-shaped cleft 37 u in height; membrane extending posterior

to setae /5. Ventral idiosoma with epimerites I Y-shaped, with surface fields; striae

obliquely directed and meeting at meson in V-configurations posterior to

epimerites I; humeral setae dagger-shaped, 46 u in length; subhumerals ventral to

humerals and setiform; setae c3 each on small plate posterior to genital organ;

adanal discs dentate with striae radiating from corolla. Legs 4-segmented (genua

and femora fused); setae absent on trochanters I, II; tarsi I, II with whorl of 4 setae

near midlength; setae s absent; tarsus IV with setae d, e as short spines inserted on

apicodorsal protuberance.

Female (paratype). Length, including gnathosoma 393 u, width 139 u. Idiosoma

elongate, parallel-sided, posteriorly truncated. Propodosoma with striae directed

dorsally and ventrally as in male. Hysterosoma with narrow, posteriorly truncated

shield bearing setae d2 - d4 at margins, ventral hysterosoma and legs as figured.

Type data. From Gallinula chloropus (L., 1758) (Rallidae): 4 holotype, 12 g , 19

Q paratypes, Dalton Pass, North Vizeaya, Luzon, Philippine Islands, February 18,

1967, U.S. Migratory Animal Pathological Survey team (NU 13,442). Holotype and

paratypes: UGA; paratypes: AMNH, BMNH, NMNH, GAUD.

Remarks. The new species is named for Dr. H. Elliott McClure, recently retired

from the Migratory Animal Pathological Survey, who over the course of many

years has sent us thousands of feather mite collections from Southeast Asia.

_ Parazumptia gen. nov.

Diagnosis. Xolalgine mites lacking setae sR on trochanters I, II; humeral setae

long, setiform; subhumeral setae anteroventral to humerals, setiform; propodo-

somal shield reduced; hysterosomal shield rather large; long, narrow metapodo-

somal shields lateral to hysterosomal shield; dorsal hysterosomal gland well

332 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 9, 1976

Fig. 5-9. Zumptia macclurei spec. nov.: ventral (5) and dorsal (7) aspects of male; enlarged pretarsus (6);

ventral (8) and dorsal (9) aspects of female. Setae: cl, central; cx3, coxal; h, humeral; 42, //, dorsal and

lateral hysterosomals; sh, subhumeral

W.T. ATYEO & P. C. PETERSON: Zumptia and Parazumptia 388

developed; two pairs of hysterosomal setae absent (d/, d2), ll positioned

posterolateral on propodosoma; epimerites I Y-shaped; legs with four functional

segments (femora and genua fused); tarsi I, II with whorl of four setae near

midlength. Males with dentate adanal discs; well-defined posterior lobes with

extra- and interlobar membranes, latter incised at midlength; genital organ small,

reflexed; genital region partially surrounded by well-defined pregenital apodeme

in which genital discs are incorporated; ventral setae as figured, with one pair

obviously associated with coxae IV (cx4); legs III, IV subequal in length, legs IV

slightly larger than III in diameter. Female similar to male in basic characteristics;

epigynium small, transverse, and not extending to level of genital discs.

Type species: Parazumptia gallinulae spec. nov.

Parazumptia gallinulae spec. nov. (Fig. 10-14)

This taxon is superficially similar to species of Zumptia in that the dorsal shields

are weakly developed, the legs are four-segmented, and the males have deeply

cleft idiosomata with each lobe bearing at least one large interlobar membrane.

Parazumptia gallinulae is easily distinguishable from the known species of Zumptia

by the long, setiform humeral setae; the same setae in Zumptia are short and

dagger-shaped.

Male (holotype). Length, including gnathosoma and terminal lobes 340 u, width

154 u. Idiosoma broad, gently tapering behind humeral setae to deeply cleft

terminus. Dorsal propodosoma with shield small, granulated, not extending to

scapular setae; area posterior to shield, inciuding insertions of scapular setae with

transverse striae with superimposed sclerotization. Dorsal hysterosoma with

weakly developed shield; two pairs of setae absent [dl and d2(?)]; narrow

sclerotized band lateral to dorsal gland opening; hysterosomal glands large;

terminus with deep V-shaped cleft 93 u in height; each lobe bearing membranes on

inner and outer margins. Ventral idiosoma with epimerites I Y-shaped, with stem

extending almost to sejugal region; striae vertically directed between epimerites I,

II and between epimerites II and ventral margin of scapular shield; subhumeral

setae fine, anteroventral to humeral setae; setae c/ and cx3 in transverse line; setae

c2 and a within apodemes connecting pregenital apodeme and terminus; setae c3

(cx4) near trochanter IV; genital discs incorporated in pregenital apodeme; adanal

discs dentate with prominent rays radiating from corolla. Legs four-segmented

(genua and femora fused); setae sR on trochanters I, II absent; tarsi I, II with

whorl of 4 setae near midlength (ra, la, wa, aa); setae s absent; tarsus IV with setae

d, e as short spines inserted on apicodorsal protuberance.

Female (paratype). Length, including gnathosoma 385 u, width 150 u. Idiosoma

with small, weakly developed shields, terminus truncated. Dorsal propodosoma

with shield as in male, but area posterior to shieid with striae as in other regions,

i.e., without superimposed sclerotization; other aspects except setal positions

similar to male. Ventral idiosoma with epimerites I, II similar to male; epigynium

small, not extending to genital discs; other features as figured.

Type data. From Gallinula chloropus (L., 1758) (Rallidae): 3 holotype, 93,7 9,

Dalton Pass, North Vizeaya, Luzon, Philippine Islands, February 18, 1967, U.S.

334 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 119, AFL. 9, 1976

Fig. 10-14. Parazumptia gallinulae gen. nov., spec. nov.: ventral (10) and dorsal (12) aspects of male;

enlarged pretarsus (11); ventral (13) and dorsal (14) aspects of female. Setae: c/, central; cx3, coxal: d3, _

II, dorsal and lateral hysterosomals; 4, humeral; sh, subhumeral

W.T. ATYEO & P. C. PETERSON: Zumptia and Parazumptia 335

Migratory Animal Pathological Survey team (MAPS) (NU 13,442); 2 g , same data

India, December 12, 1969, MAPS (UGA 6256). Holotype, paratypes: UGA;

paratypes: AMNH, NMNH.

Remarks. The same specimen of Gallinula chloropus harboured Zumptia macclurei

and Parazumptia gallinulae. The former mite species was not found on other

specimens of the moorhen infested with P. gallinulae.

LITERATURE CITED

Atyeo, W. T., 1974. Dogielacarus uncitibia Dubinin, 1949, redescribed and reassigned (Acarina: Anal-

goidea). — J. Kansas ent. Soc. 47: 478—482.

Atyeo, W. T., & J. Gaud, 1966. The chaetotaxy of sarcoptiform feather mites (Acarina: Analgoidea). —

J. Kansas ent. Soc. 39: 337—346.

Gaud, J., 1968. Acariens Sarcoptiformes plumicoles (Analgoidea) parasites sur les oiseaux Ralliformes

et Gruiformes d’Afrique. — Annls. Mus. r. Afr. cent. Ser. in8°, Zool. (164): 1—101.

———, 1974. Quelques espèces nouvelles de Sarcoptiformes plumicoles (Analgidae & Dermoglyp-

hidae) parasites d'oiseaux d'Europe. — Acarologia 15: 727—758.

Gaud, J., & J. Mouchet, 1959. Acariens plumicoles des oiseaux du Cameroun. — Annis. Parasit. hum.

comp. 34(4): 493—545.

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UITGEGEVEN DOOR

MUS. COMP. ZOOL,

LIBRARY

MAR 2 3 1977

HARVARD

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGINGNIVERSITY

* Een sterretje duidt een naam aan, welke nieuw is voor de wetenschap.

* An asterisk denotes a name new to science.

Uit het register zijn weggelaten de namen uit het artikel van Van Lith, welke reeds zijn opgenomen in

het eigen register (p. 122).

ACARI

Analloptes 327, 328

Beaucournuella 328

Cryptosikya 328, 329

dermoglyphoides 327, 328,

[329

Dogielacarus 328, 329

* gallinulae 328, 331, 335

glyphoides 328

Gymnalloptes 327, 328

loculosa 328

* macclurei 328, 329, 331,

[335

megnini 327, 328

pallens 327, 328

* Parazumptia 328, 329,

[331, 335

protalgoides 328

uncitibia 328

Zumptia 327, 328, 329,

(3311; 335

AVES

Fulica atra 327

Gallinula chloropus 327, 328,

(331, 333, 335

Limnocorax flavirostris 328,

[330

Podica senegalensis 328

Porzana fusca 328

Porzana parva 328

Rallus aquaticus 328

COLEOPTERA

Calathus 1, 12, 13

melanocephalus 12, 13

oblongopunctatus | et seq.

Pterostichus | et seq.

COLLEMBOLA

acauda 237

Acherontides 238

Acherontiella 237, 238

Acherontiellina 237, 238, 244

* Acheroxenylla 221,

[237-238

affinis 225 et seq.

afurcata 259 et seq.

alba 311

* albertinae 221, 291 et seq.

albida 311

albinus 306

Allacma 320

alpinus 318

americanus 284

anophthalamis 284

* anthrenoidea 221, 275,

À [278 et seq.

arenaria 316

armata 245

articulata 321

asigillata 267 et seq.

assimilis 247, 249

aterrima 251

atrata 237

aureus 318

banyulensis 316

bayeri 259

bipartita 227

Biscoia 238

* ssp. bisetosus (S. alpinus)

[221, 318, 319

boerneri (Pseudachorutes)

[274

boerneri (Xenylla) 237

bougisi 242, 244

Bourletiella 320

buddenbrocki 298

Calvatomina 321

cancellatus 233

capitata 247, 251

cassagnaui (Coloburella) 282

cassagnaui (Pratanurida) 274

cassagnaui (Stenognatellus)

[318-319

cavernicola 309, 310

cavicola 322

Ceratophysella 251

christianseni 247, 249

ciliata 312

Clavisotoma 294, 296, 298

conlatus 221, 232, 233

crassicornis 303, 305

crassus 272

* cretensis (Acheroxenylla)

[221, 236, 237 et seq.

* cretensis (Neanura) 221,

[277 et seq.

* cretensis (Troglopedetes)

[221, 306-310

* critica 221, 230 et seq.

Cryptopygus 292

Cyphoda 306

dagamae 315, 316

debilis 289 et seq.

decipiens 266, 267

denisi 318, 319

f. dentata (H. packardi) 251

denticulata 244, 245

Deuterosminthurus 320

Dicyrtoma 320

Dimorphiella 300

Dimorphotoma 300

dollfusi 316

domestica 316

dubius 269

dunarius 237

echidnus 300

echinata 320, 321

edinensis 237

engadinensis 244, 245

ferrarii 314, 316

flaviceps 320

geronensis 272

ghidinii 233

ghilarovi 259

gibbosa 245

gisini (Hypogastrura) 243,

[245, 247 et seq.

gisini (Onychiurus) 221, 232,

[233

gisini (Stenaphorura) 228

graeca 221, 224, 314 et seq.

grandis (Colonavis) 263

grandis (Frisea) 263

granulata 274-276, 278

grassei 306

guthriei 320

Guthriella 300

handschini 316

hankoi 294

hellenica 280, 282

hygropetrica 282, 283

hygrophila 232

hystrix 282

imparipunctata 311

incertus 317

insignis 290

instratus 314

Isotomurus 303

italica (Mesaphorura) 231,

[232

italica (Ptenothrix) 319, 322

italica (Seira) 316

krausbaueri 231, 232

ksenemani 283, 285-286

ladeiroi 261, 265, 266

lagrecai 261

lamelligerus 304 et seq.

Lepidocyrtus 309, 312

libanensis 269 et seq.

lignorum 312 et seq.

linnaniemii 282

Lipothrix 320

longiseta 279, 280-281

ssp. lusatica (N. tetroph-

[thalma) 278

lusitanica 316

ssp. luxemburgensis (T.

[hygropetrica) 282

macrotuberculata 251

maculatus 320, 321

major 310

malmgreni 317

manubrialis 249

marchicus 285

marginata 320

maritima 244

maroccanus 306, 307

ssp. matthesi (T. hygro-

[petrica) 282

melitensis 321

meridionalis 285

Mesaphorura 221, 255

Metaphorura 225, 228

Micranurida 274

* minitaurus 221, 255 et seq.

minor 301

minos 300

minuta 292

monophthalma 286

monticola 251

* mucronata 221, 271 et seq.

multifasciata 317

multipunctatus 320

muriphila 300

muscorum 316

nana 253, 255

nanus 283, 285

natalicius 321

navacerradensis 284-285

Neanura 274

Neosminthurus 320

nigromaculatus 319 et seq.

nitidus 310, 311

notabilis 301, 302

novemspina 228

occidentalis 245

octopunctata 311

Odontella 255, 257

olivacea 302, 303

Onychiurus 237

* ssp. orientalis (O. nana)

[221, 252 et seq.

ossica 266-267

packardi 251

pallidipes 316

palustris 303

papillosa 300

* paprivata 221, 308, 312,

[313

paradoxa 251

parvula 259

parvulus (Pseudachorutes)

[272

parvulus (Folsomides)

[282-284

pazaristei 316

penicula 285

petrae 316

phlegraea 280

pillichi 314

ponticus 286-287, 290

* porcellus 221, 295,

[296-301, 302

Pratanurida 274

pratensis (Pseudachorutes)

[272

ssp. pratensis (F. navacerra-

[dencis) 284

prolatus 221, 232-233

Propexenylla 237

Pseudacherontides 238

Pseudachorutes 274

pseudodiplophthalma 286

pseudoghidinii 233, 235, 236

pseudostachianus 235

pumilis 317

pusilla 228

ramicuspis 228

reticulatus 318

Rhodanella 300

ssp. rhodia (A. bougisi)

[242-244

sacchii 314, 315, 316

sahlbergi 251

salaymehi 286

salmoni 263

saxatilis 316

schaefferi 300

schoetti 292, 295

Seira 221, 315

sensibilis 232

serbicus 313, 314

sexoculatus 221, 310-311

Sminthurus 320

spinularius 237

squamoornata 316

stachi (Odontella) 255

stachi Bagnall (Onychiurus)

[235

stachi Denis (Onychiurus)

[235

stachi (Proisotoma) 300

stachianus 235

Stachorutes 274

Stenaphorura 228

subcrassus 272

sublamellifera 255

sublatus 221, 232, 233

succinea 244-245

sylvatica 231, 232

ssp. tatricola (N. tetrophthal-

[ma) 278

tenuisensillata 232

teres 304

terrestralis 304

* tethyca 221, 246, 248,

[249-251, 253

tetrophthalma 278

theeli 251

thermophilus 286, 287

Tremoisea 266

tricuspis 228

* triglenus 221, 287-288, 290

ssp. trilatus (O. prolatus) 221,

[232, 233

trioculatus 290

trisetosus 282

trybomi 251

tuberculata 282

tullbergi 251

vaillanti 301

vallvidrerensis 255

vernalis 253

Vertagopus 300

viatica 251

viridis (Sminthurus) 319,

[320, 321

viridis (Isotoma) 303

* xenonis 221, 234, 235, 237

Xenylla 221, 237, 238, 242,

[244

Xenyllina 238

Xenyllodes 259

yosiii 232

DIPTERA

angustatus 17

clypeatus 16, 17

fulviventris 16, 17

Hylemia 16

Melanostoma 15 et seq.

Melithreptus 16

mellinum 16, 17

Platycheirus 15 et seq.

scalare 17

scambus 16, 17

Sphaerophoria 16

variata 16

HYMENOPTERA

Acaelius 63, 64

Acampsis 37, 47, 65

alternipes 65

analis 67

Ancylocentrus 74

Aneurobracon 35

Apanteles 62, 63

Aphaereta 46

Aphidius 71

apicalis 74, 76

Apodesmia 62

Aridelus 50

Ascogaster 66

ater 74

auctus 55

Austrozele 68

bergi 61

Blacus 48, 67

Brachistes 68

Bracon 57, 58

Braunsia 65

brevicaudis 68

buccata 54

Bucculatriplex 45

campsolechiae 46

Cardiochiles 63

cembrae 71

Cenocoelius 67

Charmon 37, 72

Chelonus 65

Chenusa 61

Chrysopophtorus 50, 75, 76

clavator 60

Clinocentrus 46

Colastes 46

conterminus 75

compressa 56

Cosmophorus 71, 73

cremastobombyciae 62, 63

Cryptoxilos 50

deceptor 74, 75, 76

Dendrosoter 59

Doryctodes 59

drymoniae 61

Dyscoletes 48

Dyscritulus 49

Earinus 64

Elachistocentrum 68

Ephedrus 71

ervi 71

Eubazus 68, 69

Eubadizon 37

excavatus 60

extensor 72

falcator 62, 63

Foesteria 69

Ganychorus 67

Gnaptodon 60

Histeromerus 57, 58

Holdawayella 50

Hormius 46

Hybrizon 48, 54

Ichneutes 68

imperator 59

infumator 73

instabilis 66

irrorator 65, 66

japonensis 64

japonicus 63

klugi 71

laevigator 72

lapponicus 68

Leiophron 50, 74, 76

liparae 61

Macrocentrus 37, 51, 67

macroscapa 74

manni 55

matsumurai 65

Mesostoa 44, 56

Meteoridea 37, 47, 51, 64

Meteorus 50, 75

Microchelonus 65

Microctonus 75, 76

Microgaster 63

Microplitis 63

Microtypus 72, 73

Mirax 62, 63

monstrosum 77

mystacinus 57, 58

Neoneurus 55

nigripennis 68, 70

nitidulus 64

Oncophanes 46

Opius 62

Orgilus 72

Perilitus 50, 75, 76

Phanomeris 46

plagiator 71

Polemochartus 61

polyzonius 62

Praon 49

Proterops 68, 70

protuberans 59

Pseudodicrogenium 35, 77

pumilio 60

punctata 61

regius 71, 73

Rhyssalus 60

Rogas 45, 46, 60, 61

Ropalophorus 50

Sigalphus 37, 47, 65, 66

similis 68

Streblocera 50, 74

striatus 67

sulcatus 75

Syntretus 50, 75

takeuchii 63

Tanycarpa 61

Telengaia 45

Termitobracon 44

theretrae 63

thoracicus 67

tibialis 69

tosensis 65

unipunctator 61

urinator 57, 58

wesmaeli 72, 73

Wesmaelia 50

Ypsistocerus 44, 55

Zele 37, 50, 73

Zemiotes 37, 50, 74, 75

LEPIDOPTERA

abottii 179, 182

abyssinica (Melitaea) 175,

[182, 195

ssp. abyssinica (E. belemia)

[170

acamas 176, 184, 192

alceae 179, 182, 208

ssp. alcippus (D. chrycippus)

[172, 183, 184

alexanor 168

Anaphaeis 171, 184, 192

Anthea 192

Apharitis 176, 184, 192, 204

Appias 167

Araschnia 199

Argynnis 175

Aricia 192

aristaeus 207

asterope 175, 192

atalanta 174, 192

ausonia 170, 192, 193

australis 172

aurora 192

aurota 171, 184, 192, 204

Azanus 178, 192

balkanicus 177, 182

Bassaris 173

beckeri 169

belemia 170, 182, 192, 193

boeticus 192

Boloria 175

Borbo 181, 183, 197, 211

borbonica 181, 183, 197, 211

brassicae 168

brassicoides 168, 182, 183,

[195, 208

Brenthis 175

buchanani 176

buqueti 192

Byblia 192

Cacyreus 192

calais 170, 183, 192

c-album 199

callidice 169

Calopieris 192

canidia 168

Carcharodus 179, 182, 195

cardui 175, 191, 192, 205

Castalius 192

Catopsilia 171, 183, 184, 185,

[197

Celerio 217-219

celtis 173

Charaxes 173

charlonia 192

Chilades 177, 184, 192

chloridice 169

chrysippus 172, 184

chrysonome 171, 183, 192

cilissa 176

clarki 179, 182

Colias 171, 182, 192, 193

Colotis 170, 183, 184, 192,

[200

cramera 192

crashayi 192

cretosus 192

crocea 171, 172, 192

Cyclyrius 177, 185, 205

Cynthia 173, 175

Danaus 172, 183, 184, 197

daplidice 169, 182, 192, 193,

ne [210

davidis 169

demodocus 205

demoleus 168, 205

deota 168

deserticola 192

Deudorix 167, 176

Dichora 172

didyma 175, 192, 193

dispar 166

distorta 169, 182, 195

doris 180, 182, 208

dubernardi 169

electo 171, 172, 182

eleusis 192

Elphinstonia 170, 192

epargyros 176

erate 171, 172, 182, 193

eriphia 192

eris 192

Euchloe 170, 182, 183, 192,

[193

eulimene 192

hybr. euphaes 217, 218

euphorbiae 217-219

eupompe 192

evagore 170, 192, 200

evippe 192

excelsior 175, 182

falloui 170, 182, 183, 192, 193

fausta 170, 184

felix 176, 182, 195

fieldii 172

florella 171, 183, 184, 185,

[197

Freyeria 177, 178

galba 178, 179, 184, 192, 198

ssp. gambica (G. pumilio)

[181

Gegenes 180, 182, 192, 204

geron 180

gilletti 176

glauconome 169, 182, 191,

[192, 204

Graphium 167

halimede 192

hanningtoni 175, 182, 195,

[196

helice 169, 182, 208

Heliophorus 179

helle 166

Heodes 166

hierta 175

Hipparchia 175, 182, 207,

[208

hippophaes 217-219

hybr. hippophorbiae 217-219

hospiton 168

hottentota 180, 182

hyale 172

Hypolimnas 173, 184, 185

ilithyia 192

ssp. inconspicua (P. thrax)

181

indica 174

Iolaus 192

hybr. irene 219

Issoria 175, 182, 195, 196

jasius 173

jesous 178

karsandra 178, 207

kershawi 175

knysna 178, 192, 207

labdaca 172

laius 172, 177

Lampides 177, 192

Lasiommata 176, 182

lathonia 175

levana 199

liagore 192

Libythea 172

lilacinus 176

lingeus 192

livia 176, 183, 192

livornica 217, 219

Lycaena 166, 167, 178, 182,

[192, 195, 208

machaon 168, 192, 193, 199

maderakal 176, 182

mafa 179, 180, 198

mandersi 177

ssp. marnoana (C. erate) 171

mathias 181

maxima 176

melete 168

Melitaea 175, 182, 192, 193,

[195

ssp. meridionalis (P. brassi-

[coides) 168

minuscula 178

misippus 173, 184, 185

Mycalesis 167

myrmecophila 176, 184

naganum 168

narina 172

Nepheronia 192

Neptis 173, 185, 205

nigricans 169

nilus 176

niso 180

nostrodamus 180, 181, 182,

[192, 204, 209

ssp. nouna (C. evagore) 170

nursei 192

oenone 192

orbifer 179, 180, 183, 186,

[198, 201,211

orithya 175, 184

orus 179, 182

osthelderi 180

Palaeochrysophanus 166

Papilio 168, 192, 193, 199

Pararge 176, 195

hybr. pauli 217, 219

Pelopidas 181, 184

phisadia 171, 184, 192

phlaeas 166, 179, 182, 192,

[195, 208

phlomidis 180

Pieris 168, 182, 183, 192, 208

Pinacopteryx 192

pirithous 177, 183

plexippus 172, 197

plinius 177

Polygonia 199

Pontia 168, 182, 191, 192,

[193, 195, 204, 208, 210

Pontieuchloia 169

Precis 175, 184, 192

protodice 169

ssp. pseudohecate (C. electo)

[172

ssp. pseudophlaeas (L.

[phlaeas) 179, 208

pumilio 180, 181, 182, 211

rivularis 173

rapae 168, 192

rosaceus 177, 192

sappho 173

ssp. scotti (M. abyssinica)

[175

sebaldus 192

sertorius 179, 180, 183, 186,

[198, 201, 211

ssp. shima (L. phlaeas) 179,

[208

sisymbrii 169

smaragdifera 175, 182

Spialia 179, 182, 183, 186,

[198, 201, 208, 211

Synchloe 169

Syntarucus 177, 183

Tarucus 176, 182, 192

tewfiki 175, 182, 208

theophrastus 176, 192

Thersamonia 166

thrax 181, 184

trochilus 178

Vanessa 173, 191, 192, 205

hybr. vespertilioides 219

Virachola 176, 183, 192

virginiensis 175

ssp. vulcanica (V. indica) 174

webbianus 177

ssp. wissmanni (C. alceae)

[179

xuthus 168

Ypthima 175, 192

ssp. zelleri (B. borbonica)

[181

Zizeeria 178, 192, 207

ORTHOPTERA

Acheta 124, 139

Acinipe 142

Acrida 125, 153, 154

Acridella 149

Acridium 145, 150

Acrometopa 124, 125, 154

Acrotylus 125, 150 et seq.

aegyptium 124, 145

Aeolopus 150

affinis 124, 129

Aiolopus 125, 150

albifrons 124, 127

algericus 139

algirius 139

Anacridium 124, 145

anatolica 133

Arachnocephalus 124, 140

astyla 124, 130, 131, 154

barbarus 124, 144

bicolor (Acrida) 153

bicolor (Stenobothrus) 148

bimaculatus 124, 139

biroi 125, 148, 154

Bolivarius 124, 137, 155

bordigalensis 124, 139

brunneus 125, 148

caerulans 152

caerulescens 125, 151

Calliptamus 124, 144

Caloptenus 144

cavicola 154

chabrieri 130, 154

chopardi 139

Chorthippus 125, 148, 154

cicindeloides 124, 140

ssp. cinerascens (L. migra-

toria) 14

coerulans 152 a

concii 128, 154

conica 124, 143

Conocephalus 124, 126, 153.

ssp. cophtha (G. vulgaris)

[141

cretensis (Acromorpha) 124,

[125, 154

cretensis (Gryllomorpha)

[124, 140, 154

cretensis (Poecilimon) 124,

[126, 134, 154

cretica (Eupholidoptera)

[130, 131, 154

* ssp. cretica (P. grisea) 124,

[127, 153, 154

cylindrica 143

dalmatina 124, 140

danicus 149

decorus 125, 150

Decticus 124, 127

depressa 124, 141

discolor 124, 126, 153

Discoptila 124, 140, 154

distinguendus 126

Dociostaurus 125, 148

Dolichopoda 124, 138, 154

[et seq.

domesticus 124, 139

elegans 124, 137, 155

Epacromia 150

Ephippiger 134, 135

Ephippigera 137

escalerai 124, 129, 154

Eupholidoptera 124, 129, 130

[et seq.

ssp. exornatus (S. caerulans)

[153

fieberi 124, 142, 154

* forcipata 124, 129, 130,

(131, 153, 154

fuscum 126

* gemellata 124, 130, 134,

[136, 153, 154

germanica 124, 137

geticus 124, 154

giornae 124, 145

Glyptobothrus 125, 148

graeca 143

ssp. grandis (C. italicus) 144

gratiosa 152

grisea 124, 127, 129, 153

grylloides 143

Gryllomorpha 124, 140, 154

Gryllotalpa 124, 141

gryllotalpa 124, 141

Gryllus 124, 139, 149, 152

ssp. hebraeus (O. yersini) 143

Heteracris 124, 145, 153

Homorocoryphus 124, 126,

[153

idomenaei 137

incerta 124, 129

Incertana 124, 129

ssp. inficitus (A. insubricus)

[125, 151

inflatus 154 et seq.

insubricus 125, 151

intermedia 124, 129

italicus 124, 144

jonicus 126

laeta 150

* latens 124, 130, 134, 153,

[154

lilifolia 124, 125

lindbergi 124, 140, 154

lineola 145

Liogryllus 139

littoralis 124, 145, 153

Locusta 125, 149

longicornis 124, 143, 153, 154

longipes 125, 150

macropoda 125

maroccanus 125, 148

meridionalis 124, 141

Microscirtus 152

migratoria 125, 149

miniata 154

Mioscirtus 152

Modicogryllus 124, 139, 154

sp. monticola (P. grisea) 127

Morphacris 152

Myrmecophilus 124, 140

Myrmophilina 124, 140

nana 124, 125, 153

nasuta 125, 149, 153

naxia 155

nigrofasciatus 150

nitidulus 124, 126, 153

ochraceus 124, 140

Ochrilidia 124, 146, 154

Oecanthus 124, 141

Oedaleus 125, 150

Oedipoda 125, 149 et seq.

Omocestus 148, 154

Orchamus 124, 142, 154

Pachytilus 149

* pallipes 124, 130, 135, 153,

[154

Pamphagus 142

Paranocarodes 124, 142, 154

paraskevi 124, 138, 154 et seq.

Paratettix 124, 141

patruelis 125, 151

pellucens 124, 141

petraeus 148, 154

Pezotettix 124, 145

Phaneroptera 124, 125, 153

Pholidoptera 131

Platycleis 124, 127 et seq.,154

Platyphyma 145

Platypterna 146

Poecilimon 124, 126, 134,

[154

Porthetis 142

procera 149

pruinosa 124, 146

Pyrgomorpha 124, 143

raulinii 124, 142, 154

Rhacocleis 124, 137

roeweri 124, 138, 154

rubescens 152

Scintharista 152

Sepiana 124, 130, 153

sepium 124, 130, 153

servillea 125, 154

siculus (Calliptamus) 144

siculus (Steropleurus) 137

ssp. similis (H. littoralis) 145

ssp. sparsa (P. nana) 125

Sphingonotus 125, 152, 154

spinulosus 124, 138, 154

Stauroderus 148

Stauronotus 148

Stenobothrus 148

Steropleurus 137

strepens 125, 150

ssp. syrica (T. longicornis)

[143

Tartarogryllus 124, 139

Tessellana 127

Tetrix 124, 141

Tettigonia 124, 127

thalassinus 125, 150

Thamnotrizon 130

tibialis 124, 146, 154

ssp. transiens (P. grisea) 127,

[128

ssp. transjonica (T. longi-

[cornis) 143

Trigonidium 124, 140

Troglophilus 124, 138, 154

Tropidopola 124, 143, 153,

[154

Truxalis 125, 149

Tryxalis 149, 153

turrita 125, 153

Tylopsis 124, 125

ungarica 153

unguiculata 149, 153

Uromenus 124, 137, 155

variabilis 149

venusta 125, 152, 154

vestitus 124, 140

viridissima 124, 127

vulgaris 141

Xiphidion 124, 126, 153

yersini 124, 142, 154

PLANTAE

Alopecurus pratensis 16

Anthoxanthum odoratum 16

Artemisia dracunculus 16

Arundo donax 223

Astragalus 134

Betula 2

Bryum 224 ;

Capparis 222

Carpobothrus 224

Ceratonia siliqua 224

Convolvulus lanatus 180

Dactylis glomerata 16

Echallium 224

Echallium elaterium 223

Euphorbia cyparissias 217

Euphorbia polychroma 218

Festuca pratensis 16

Ficus 222

Fossombronia 223, 224

Helictotrichon pubescens 16

Hippophae rhamnoides

[217-219

Juncus 223

Juniperus 135

Molinia coerulea 16

Malva 224

Oxalis pes-caprae 223, 224

Phleum pratense 16

Pistacia lentiscus 223, 224

Plantago arenaria 16

Plantago lanceolata 15

[et seq.

Plantago media 16

Poa annua 16

Polygonum 179

Posidonia 224

Punica 222

Quercus coccifera 135, 222,

[223, 224

Quercus robur 2

Riccia 223

Rumex 179

Salicornia fruticosa 224

Sanguisorba minor 16

Sarcopoterium 223

Scirpus lacustris 16

Selaginella 224

Selaginella denticulata 223

Tamarix 224

Targionia 223

Tilia 189

Vitis 222

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Acme

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ERNST MAYR LIBRARY

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