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~~ DEEL 125 1982

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

Tijdschrift voor Entomologie, deel 125, 1982

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

BESTUUR (BOARD)

Woorzitten(Chaiman) Prenen R. H. Cobben

Vice-voorzitter (Vice-President) ................ L. H. M. Blommers

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AMA TOSS it vn SOE EE RE Rijksmuseum van Natuurlijke Historie,

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Afleveringen 1—3 verschenen 15.VII.1982

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ISSN 0040-7496

INHOUD VAN DEEL 125

Assem, J. van den , M. J. Gijswijt & B. K. Nübel. — Characteristics of courtship and mating

behaviour used as classificatory criteria in Eulophidae-Tetrastichinae (Hymeno-

ptera), with special reference to the genus Tetrastichus s.l. …

Belle, J. — A review of the genus Archaeogomphus Williamson (Odonata, Gomphidae) .........

Brock, J. P. — A systematic study of the genus Ophion in Britain (Hymenoptera, Ichneumoni-

CRE). FEET PON IA RR AR RENO ARI I O

Cobben, R. H. — The Hebrid fauna of the Ethiopian Kaffa Province, with considerations on spe-

Giesjsroupine(ebridaenHleteroptera MIR RR I TTT

Gijswijt, M. J., zie Assem, J. van den.

Kielland, J. — Revision of the genus Ypthima in the Ethiopian Region excluding Madagascar

(BepidopreraßSatyrıdae) IRE O E

Leereveld, H. — Anthecological relations between reputedly anemophilous flowers and Syrphid

flies. III. Worldwide survey of crop and intestine contents of certain anthophilous

NES 5 aes a EON

Nübel, B. K., zie Asser, J. van den.

Willemse, F. — A survey of the Greek species of Poecilimon Fischer (Orthoptera, Ensifera, Pha-

MELOPEer NAE RIE TO SII ee ee

155

ER aa

len

Hi:

DEEL-125 Ne EVE RANG: i 1982

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

R. H. Cossen. — The Hebrid fauna of the Ethiopian Kaffa Province, with consid-

erations on species grouping (Hebridae, Heteroptera), p. 1—24, figs. 1—59.

Tijdschrift voor Entomologie, deel 125, afl. 1 Gepubliceerd 15-VII-1982

D LH, ae Ba a Pa ND

Wf i aro i ip Li pr pe N n KL,

A 5 i

1 aa We, :

brik N'EN | î

} Pad, 1 hr

DIN ee TE A

rer, DADI

| pl

Ki Aer

) Me

;

med

; DO)

THE HEBRID FAUNA OF THE ETHIOPIAN KAFFA

PROVINCE, WITH CONSIDERATIONS ON SPECIES

GROUPING (HEBRIDAE, HETEROPTERA)

R. H. COBBEN

Department of Entomology, Agricultural University, Wageningen, the Netherlands

ABSTRACT

Nine species of “velvet surface bugs” (Hebridae) are recorded from the area around Jim-

ma, the capital of the Ethiopian Kaffa Province. Six new species are described. Four belong

to the genus Hebrus (H. bimaculatus, H. pseudopusillus, H. spinitibialis, H. gidshaensis).

The other two new species (H. bongaensis, H. niemeri) and Hebrus pelengei Poisson are

assigned to Hebrometra n. gen., a taxon restricted to the sprinkle zone of waterfalls. An-

other new Hebrometra species is described from Malawi (H. malawiensis) which is consid-

ered an ancestral member of the Hebrometra clade. The phylogenetic significance of anten-

nal, genital and external microcuticular characteristics of Hebrus and Hebrometra is dis-

cussed; arguments are given to abandon the current subgeneric splitting of Hebrus.

INTRODUCTION

Hebridae are very tiny (generally 1.5—2.5

mm), plumb-bodied predacious bugs living in

clumps of mosses and other short hygrophilous

vegetation or on bare, permanently moist soil

along the margins of pools, ponds, rivulets or in

marshes. The number of described species is ap-

proximately 150, but because of their size and

obscure way of life probably many more species

still have to be discovered. The family predomi-

nantly occurs in the warm regions, particularly

of the Old World. The Hebridae form an inter-

esting group in studies on the phylogeny of het-

eropterous families, since, as a representative of

the major group Gerromorpha, it contains a

number of archaic features (Cobben, 1968,

1978). Two of these original traits concern the

habitually damp soil-substrate required for sur-

vival, and the carnivorous way of feeding with

rasping-filing maxillary stylets (Andersen,

1979; Cobben, 1979).

The mense described in this paper was col-

lected near Jimma, the capital of the Kaffa Prov-

ince in the southwestern part of Ethiopia, dur-

ing a two month visit (Oct.-Nov.) in 1969. Up

!) Recently Andersen (1981), in a superb revision of

the family, recognizes seven genera (Hyrcanus,

Merragata, Lipogomphus, Timasius (with ten new

species, mostly from India), Neotimasius, Hebrus,

Hebrometra (see the present paper)).

to now some 40 hebrid spp. are known from

Africa (inclusive of Madagascar), of which four

occur in Ethiopia. The fact that I could sample

nine species, six of them new, in an area of

roughly 50 km radius around Jimma, demon-

strates that the Ethiopian hebrid fauna is com-

paratively rich and still poorly known. Apart

from the worldwide and largest genus Hebrus

Curtis, the family contains some genera with

only few species: Merragata White (Old and

New World), Hyrcanus Distant, Timasius Dis-

tant, and Timasiellus Lundblad (restricted to the

Oriental Region!). So far only representatives of

Hebrus have been recorded from Africa. In this

paper some typical hebrid species will be de-

scribed as new; four other species are recorded

which warrant a separate generic status on the

basis of some synapomorphic characteristics.

Before treating the species collected, some an-

tennal, genital and external micro-cuticular

characteristics will be discussed in view of their

importance for a future revision of the world-

wide genus Hebrus.

DISCUSSION OF CHARACTERS

The antennae

Most African species have been described by

the French specialist Poisson in a series of pa-

pers from 1934 onwards. In 1943 he first divid-

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

0.025

arr mg

Fig. 1. Hebrus gerardi; a, right fore wing of 4; b, hind tibia of & with row of incised spines; c—e, paramere of

specimen from Ethiopia; f—h, paramere of two specimens from Ivory Coast. Fig. 2. Left view of buccula of: a

Hebrus gerardi, b, Hebrus katompei and Hebrus pseudopusillus; c, Hebrus pusillus, Hebrus gidshaensis, and He-

brus spinitibialis; d, Hebrus bimaculatus. 3. Hebrus bimaculatus; a, b, paramere; c, hind tibia of d ; d, pronotum.

COBBEN: Ethiopian Hebrid fauna 3

ed the genus into the subgenera Hebrusella and

Hebrus s.s. The first subgenus has five distinct

antennal segments, whereas representatives of

the second seem to possess only four segments

when superficially viewed. The last segment in

this latter species group reveals a peculiar differ-

entiation half-way, rightly circumscribed by

Poisson (1955a) as “une zone de chitine souple

vaguement annelée sans noeud articulaire”. This

differentiation was already noted by Lundblad

(1933) (“mit einer falschen Gelenkverbin-

dung”). I adopt here the term “pseudoarticula-

tion” (Miyamoto, 1965) since it indeed has no

articulatory function. This point of weakness

enables the apex of the antennae to vibrate in air

currents and it is easily broken off during ultra-

sonic cleaning. The pseudoarticulation has an

uninterrupted continuation of setae, and is

merely perceptible in scanning-micrographs as

an area with weak, transverse striation (fig. 49).

Under transmitted light, there appears to be a

thin transparent outer cuticle, whereas the scle-

rotized inner cuticle is broken up by obliquely

spiralising strips (fig. 11b.).

Poisson (1955b) described Hebrus houti from

S. Africa having four simple antennal segments

without any sign of subdivision of the last one,

and placed it in a new subgenus Subbebrus. He

described it again as new subgenus in 1957 with

the note that the reduction of antennal segments

is undoubtedly of secondary nature. I doubt this

very much since four-segmented antennae occur

in the hebrid genera Hyrcanus and Merragata,

and in all other families of waterstriders. This

nodal number is found further in all other major

groups of Heteroptera with the exception of

most Pentatomidae (second segment divided)

and some reduviid taxa (many subdivisions ei-

ther of the 2nd or 4th segment) (Miller, 1956). It

is therefore more likely that the pseudoarticula-

tion in Hebridae is a transitional state towards

the 5-segmented condition (all larval instars of

the subgenus Hebrus lack the pseudoarticula-

tion!). A pseudojoint is also found in the hebrid

genus Timasius (Miyamoto, 1965) and even in

Madeovelia (Poisson, 1959) which is affiliated

with the Mesoveliidae (Cobben, 1968; Ander-

sen & Polhemus 1980). Outside the Hebridae, a

pseudoarticulation is also present in the Paraph-

rynoveliidae (Andersen, 1978), and in Hydro-

metridae (Heterocleptis, Andersen personal

comm.). Thus, there is evidence that this ten-

dency for duplication of the last segment

evolved more than once independently. Its pres-

ence or absence alone is then not reason enough

for subgeneric delimitation. The problem of

subgeneric divisions within Hebrus is further

complicated by three additional subgenera from

Africa and Madagascar: Paratimasius Poisson,

1952, Timasielloides Poisson, 1952, and Parati-

masiellus Poisson, 1956. These have also the an-

tennal construction of the subgenus Hebrus,

and moreover some other external features,

which on the basis of the descriptions seem too

weak and not discontinuous enough to warrant

subgeneric status.

Male genital structures

Realizing that the characters used for the

present subgeneric grouping of species could be

liable to parallelisms, I paid particular attention

to the male genitalia of the Ethiopian and Euro-

pean species, since in other families these struc-

tures have been proven to be of great value in

supraspecific ranking. The symmetrical para-

meres of Hebridae are already generally used

for the diagnoses of species, although Lundblad

(1933) has already stressed that their shape is

highly dependent on the angle under which it is

viewed. One picture from a fixed angle or var-

ious pictures of the same paramere (e.g. fig. 1f—

h) are therefore necessary for adequate compar-

isons between species. The three-dimensional

shape and vestiture of typıcal hebrid paramere

(Hebrus pusillus) ıs shown in the SEM micro-

photographs of figs. 54—56. The outer surface

of the hooked paramere is densely beset with

long hairs (fig. 54). The inner surface has a

group of three pegs and a perpendicular squa-

mous outgrowth. This latter protuberance bears

three sensillum-like structures (figs. 56, 57)

which obviously play a role when touching the

female ovipositor. Such microstructures deserve

attention in future comparisons between other

hebrid taxa.

The intromittent male organ of Hebridae has

never been studied for the purpose of unravel-

ling intrafamilial relationships, presumably be-

cause of its very tiny dimensions (the not inflat-

ed phallus measures only 0.1—0.2 mm). My

earlier experience when comparing genitalia in

most heteropterous families (see Cobben, 1978,

for the diversity within the families of water-

striders) concerned only Hebrus ruficeps from

Europe. It belongs to the subg. Hebrusella sen-

su Poisson with five fully developed antennal

segments. The phallus which can be inflated by

slight manual pressure on the abdomen, as in

most Gerromorpha, unfolds a very simple com-

position (fig. 22). Inflation is simply brought

4 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

== ne

=> ST

==>"

Fig. 4. Hebrus katompei; a, b, paramere; c, hind tibia of 6; d, hind tibia of ©. Fig. 5. Hebrus pusillus, speci-

mens from the Netherlands; a, b, paramere; c, front leg of d; d, hind tibia of d. Fig. 6. Hebrus pseudopusillus,

hind tibia of d. Fig. 7. Left fore wing of: a, Hebrus bimaculatus; b, Hebrus katompei and Hebrus pseudopusil-

lus; c, Hebrus spinitibialis (black area represents circumference of abdomen; similar Hebrus gidshaensis, but

smaller proximal white spot).

COBBEN: Ethiopian Hebrid fauna 5

about by blood pressure into the endosoma.

The ductus seminis is very thin and runs en-

tirely free through the simple basal articulatory

apparatus, phallotheca and endosoma. The lat-

ter is more than four times the length of the

phallotheca in which it is folded during rest. It is

bilaterally symmetrical and is divided in a long

proximal cylindrical conjunctiva and a distal ve-

sica with several pairs of swellings. Two lateral

crescent-shaped sclerites support the basis of a

pair of preapical flap-like extensions. These

sclerites are well-discernible in the phallus if the

entire endosoma, being out of function, is re-

tracted (vss in figs. 23, 24).

Some of the Ethiopian species, however, have

more complex internal endosomal equipment,

occurring also in the second northern-European

species H. pusillus, belonging in the subgenus

Hebrus sensu Poisson. Instead of just two sup-

porting sclerites there are two more sclerotized

thickenings leading each to an undulating apical

process. The greatest difference with the He-

brusella-type, however, are two sclerotized

threads extending from two invaginated lobes

with which they form a proximal loop (es in

figs. 24, 25). The SEM micrographs (figs. 58, 59)

show these struts very clearly after rupture of

the vesical membrane.

The function of the struts becomes clear

when pairs in copula are studied (figs. 28, 29;

the fixed material was kindly put at my disposal

by Mrs. Karin Heming-v. Battum). They appar-

ently serve as a more solid expansion of the ve-

sica, the loops being pressed against the ventral

wall of the female gynatrium. The result is that

the secondary gonopore of the phallus takes a

fixed position close to the gutter-shaped wall

thickening (gwth in fig. 29) which leads to the

entrance of the tubular spermatheca for receiv-

ing the sperm. Figs. 27b—d further show that

during copulation the male genital segments are

rotated through an angle of 90°, that only one

paramere is functioning as a clasper, and that

the phallus is inserted into the apical region of

the ovipositor.

The striking discrepancies in phallic struc-

tures mentioned above for H. (Hebrusella) rufi-

ceps and H. (Hebrus) pusillus could enhance

subgeneric differences. However, although the

pusillus phallic type seems to be restricted to the

subgenus Hebrus having the pseudoarticulatory

antennal joint, some of the Ethiopian spp. do

have the endosomal looped struts, while other

related spp. do not possess them. The Ethiopian

material further contains some species with a fa-

cies unlike typical Hebrus (described below as a

new genus), but with the antennal structure of

the subgenus Hebrus type. Nevertheless, their

endosomal structures are very simple and with-

out sclerotic struts.

Microsculpture of the dorsal integument

The shape of pronotum and scutellum, and

the distribution of deep cuticular pits on it, are

commonly used in the literature for the diag-

noses of species. Often it is not clear whether

the pictures given are from dry material or from

specimens in preservative fluid. In the former

case, some pits are easily overlooked. One or

two specimens of the species listed below were

studied with the scanning electron microscope.

The resulting pictures, made under proportion-

ally low magnification, were compared with

light optical observations of additional material

for allowance of generalizations to be made on

constant features by species. The purpose was

to analyse whether the micro-level might reveal

differences in species-group categories. As dis-

cussed above, the species grouped under the

Hebrusella- or Hebrus-type have different an-

tennae. The integument of head and thorax re-

veals the following three discrepancies between

both groups. The row of bare spots on the me-

dian line of the vertex, representing internal

muscle insertions of the pharynx pump, is prox-

imally marked by one deep pit (figs. 41, 50) in

all species of the Hebrus-type studied. In H. ge-

rardi (fig. 30) and in H. ruficeps, representing

the Hebrusella-type of species, this pit is lack-

ing. In the latter group, the pronotal pits bear a

central projection, mostly cylindrical (figs. 31—

35), whereas in the Hebrus-type species studied,

these pits are bordered with a fringe of microtri-

chia and lack a central projection (figs. 37, 43).

The mid-dorsal pronotal pits in the Hebrusella-

type (fig. 30) are lower in number than in the

Hebrus-type species (figs. 36, 47, 51). The func-

tion of such pits, which are often associated

with particular, probably sensillar structures in

other families (see Cobben, 1978: 159—164), is

not clear. The three species group dividing char-

acters should be compared in more species in

order to test their general validity.

The external cuticular surface of water strid-

ers is of a complex nature, predominantly serv-

ing as a water-proof coat (Andersen, 1977;

Cobben, 1978). The basic pattern of sculpture is

rather uniform in all hebrid species studied,

consisting of a thick layer of microtrichia, often

with intertangled apices (figs. 39, 44). Within

6 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

Fig. 8. Hebrus spinitibialis; a, b, hind tibia of 6, in lactic acid slide (a), dry (b); c—e, paramere; f, front leg of d.

Figs. 9—10. Hebrus gidshaensis; a, front leg of 3; b, hind tibia of 8; 10, apex of antenna, ventral, with the

patch of sensory pegs, occurring in all Ethiopian Hebrus species. Fig. 11. Hebrometra niemeri; a, antenna; b,

pseudoarticulation of last segment.

COBBEN: Ethiopian Hebrid fauna 7

this layer, so-called “peg-plates” (Andersen,

Le.) or sieve-pores (Cobben, l.c., possibly rep-

resenting outlets of epidermal glands) are dis-

tributed (figs. 38, 39, 43—45, 52, spherical rims

including a group of conical pegs). Other cuti-

cular extensions are mushroom-like, calyx-like,

shoe-lift-like and slender macrotrichia with

fluted surface (figs. 32, 39, 44, 52). Shoe-lift

structures with a slender form occur in the spe-

cies bimaculatus, pseudopusillus and katompei,

and with a tapering form in the species spimiti-

bialis and gidshaensis. They are absent in the

new genus, described below, and in the Hebru-

sella-type spp., H. gerardi and H. ruficeps. The

latter two species possess calyx-like structures

instead.

From the foregoing it seems provisionally

recommendable not to use the antennal and

phallus structures mentioned as the only subge-

neric criteria within the large group of other-

wise typically shaped Hebrus species. Another

reason is the apparent absence of clear ecologi-

cal differentiation between the species having

four, five or an intermediate number of antennal

segments. For convenience, however, I will in-

dicate in the descriptions of species given below

the stage of antennal development between

brackets: (Hebrus-type fourth segment not

completely subdivided) or Hebrusella-type;

with five segments).

SURVEY OF THE ETHIOPIAN SPECIES

The following four species have been record-

ed so far from Ethiopia.

H. alluaudi Poisson, 1943. Addis Ababa (Pois-

Sons 955: 154); Galla Tieea (Mancino

1961: 41); also known from Kenya.

H. mancini Poisson, 1955. Addis Ababa (Pois-

son, 1955a: 154-155) (= Hebrometra pel-

engei (Poisson, 1954), see this paper).

H. jeanneli Poisson, 1943. Gembi near Agaro,

1963 (Linnavuori, in litt.); also known from

Er Africas

H. violaceus somaliensis Poisson, 1953. Sululta,

1963 (Linnavuori, in litt.); known from So-

malia; the nominate form is widespread in

East and South Africa.

I collected in the habitats around Jimma nine

species. Six species belong to Hebrus, four of

which are described as new. Three other spe-

cies, two of which are new, are deviative

enough to erect a new genus for them.

HEBRUS Curtis, 1833

Hebrus (Hebrusella-type) gerardi Poisson,

1950

(figs. 1, 2a, 30—35)

General colour dark brown; vertex behind

ocelli, inner margin of eyes, collar and anterior

mid part of pronotum light brownish; anterior

half of endocorium strikingly white (fig. 1a);

membrane unicolorous dark fuscous in 6, with

four vague lighter spots in 2. Upper side with-

out distinct hairs, except for the three pairs of

long, white cephalic trichobothria and the light

hairs on the corial veins. Antennal formula (2d,

19): 4:4—4.3:5—6:4—4.5:4—5.5. Extremities

yellowish brown. Tibia 3 of & very slightly

curved; innerside with a row of spines extend-

ing from before the middle towards the apex,

the spines being incised on top (fig. 1b). Buccula

and paramere as figured (figs. 2a, 1c—e).

Length of 26 1.7—1.8, of 2 1.9 mm.

Material: 1 dg, 1 2, Jimma, Hippo Lake

27.x.1969; 1 3, idem, 28.x1.1969, on open wet

sandy soil with sparse short grasses.

This is apparently the rarest species in the

Kaffa province. The three specimens were col-

lected at one spot within an extensive marshy

area where three other Hebrus species were

abundant. I attribute this species provisionally

to H. gerardi Poisson, described from Zaire (1

d) in 1950. Poisson (1957b) recorded additional

material from the same region, and Linnavuori

(in litt.) from Nigeria and Sudan. The long row

of incised spines of the hind tibia is characteris-

tic, but the paramere and the antennal formula

presented in the papers of Poisson deviate

somewhat from the Ethiopian specimens. Pois-

son’s drawing of the wing shows hairs on the

endocorium, whereas in my material this wing

area is entirely glabrous. I collected in the Ivory

Coast (Bouaké, iv.1964) a series which con-

forms externally quite well with the Ethiopian

animals. The parameres, however, reveal differ-

ences in shape and placement of spines (fig. 1f—

h). Future studies of material from other parts

of Africa will be necessary for a right taxonomic

interpretation of the geographic differences

mentioned. Hoberlandt (1950) described H.

wygodzinskyi from Angola. This species is close

to H. gerardi, but the mid part of the hind tibia

is broadened, bearing a short row of pointed

spines. This condition being quite different in

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

H. gerardi, I am inclined to consider H.

wygodzinskyi a valid species rather than a sub-

species of H. gerardi, as was proposed by Pois-

son (1957).

Species rs

The following five species are externally very

much the same and close to the Palaearctic He-

brus pusillus and associates. The sequence of the

species treated below follows the increasing dif-

ferentiation of the fore and hind leg of the

males, which are undoubtedly related with the

copulatory act. The hind femur of H. pusillus is

strongly arched; the hind tibia is weakly sinuate

and bears a row of some nine long spiny hairs

on the medio-dorsal surface (fig. 5d). This setal

comb has not been mentioned in the literature

(e.g. Jordan, 1954; Stichel, 1955, Poisson, 1957).

Since H. pusillus also has been recorded from

North Africa, material identified as such should

be carefully restudied for the shape and pilosity

of the tibiae, and the presence of the phallic

struts.

The attribution of single females to their

proper male sex is another complication. Gener-

ally, species of which males have clearly arched

hind tibiae, the corresponding females have also

sinuate hind tibiae, though less marked. There

Fig. 12. Hebrometra n. gen.; a, H. bongaensis, © ; b, right hemielytron of H. pelengei; c, d, right hemielytron of

H. niemeri (c, specimen from Natal, d, from Tanzania, East Usambara Mts.).

COBBEN: Ethiopian Hebrid fauna 9

appears, however, no regularity in this sexual

difference in closely related species. For exam-

ple, the d of H. spinitibialis has conspicuously

arched hind legs, whereas the 2 has straight

hind tibiae. The hind tibia of the 9 of H. ka-

tompei is, on the contrary, sinuous, whereas the

male hind tibia is much less arched than in H.

spinitibialis. Such relations make the complete

definition of species, which often occur togeth-

er, very difficult. It took me some effort to

make the right conspecific combinations be-

tween males and females. The shape of the buc-

Figs. 13—15. Hebrometra, right paramere; a, dorsal, b, other view; 13a, b, H. bongaensis (arrow points to sen-

sillum); 14a—c, H. pelengei; 15a, b, H. niemeri (specimen from Natal). Fig. 16. Left profile of head; a, H.

bongaensis (arrow points to setiferous tubercle); b, H. niemeri; c, H. pelengei. Fig. 17. Pronotum of H. bong-

aensıs.

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

culae, not liable to sexual differences, is an inad-

missible help in this dilemma.

Another severe source of confusion is the an-

tennal formula. I compared my measurements

on known species with the data from the litera-

ture, and found many incongruences. In the for-

mulae given, the length of the internodal joints

is usually indicated separately. Because of the

small sizes, I think that it is more preferable to

take the length of each segment inclusive of the

basal internode and to measure the last segment

with the pseudojoint as a single unit. My calcu-

lation of the ratios of the segments of European

H. pusillus (mean of five specimens) resulted in

the formula: 7:4:7:10, whereas the recalculation

of Poisson’s (1943) formula reads: 4.4:4:6.2:10,

thus a considerable shorter first segment. The

formula given by Jordan (1954) is: 5:4:6:9.

Because of all the difficulties mentioned

above, it is clear that earlier descriptions of spe-

cies belonging in this complex may not give all

the essential information. I tried to make the

right nomenclatorial decisions on the species

listed below. The type material in the private

collection of Poisson being not available for

study, I had to rely mostly on his original de-

scriptions.

Hebrus (Hebrus-type) bimaculatus n.sp.

(figs. 2d, 3, 7a)

General facies of the pusillus group. Prono-

tum black with a weak shine of metallic blue.

Proximal half of endocorium milky white and

hairless (fig. 7a). Buccula extending ventrally

beyond the level of the ventral head lobe (fig.

2d). Paramere as drawn in fig. 3a, b. Antennal

formula (mean of 3 d and 2 2): 7.5—8:5.5—

6:6:11.5—13.5. Hind tibia of d and © straight

(fig. 3c). Length of 4 d: 2.4—2.5 mm, of 2 9:

25 2 6, mE

Holotype d : 70 km from Jimma on the route

to NddiS ba 28219609 MParatypes SOA

idem. At the bottom of deep narrow ravine in

sprinkle zone of waterfall, restricted to the

north side on loamy soil, which received the

heat of the afternoon sun. Holotype and para-

types in the author’s coll; paratype in the Mus-

éum National d'Histoire Naturelle, Paris.

The shape of the paramere is like that of H.

caeruleus Poisson, 1934, as figured in Poisson

(1943). This species (later spelled H. coeruleus)

is recorded from the Cape Province, S.W. Afri-

ca, Angola, Mozambique, Equatorial and East

Africa (Poisson, 1934, 1943, 1957; Hoberlandt,

1950; Linnavuori, 1971). I had the opportunity

to study the & holotype and one 2 paratype

Muséum National d’Histoire Naturelle, Paris.

The third antennal segment of H. caeruleus is

1.5 times the length of the second (1.6 x

according to Poisson, 1934), whereas these seg-

ments are subequal in 7. bimaculatus. The buc-

cula of H. caeruleus is rather of the pusillus-type

(fig. 2c), and the total length of about 2 mm is

smaller than in H. bimaculatus. The metallic co-

lour tends to blue in H. bimaculatus and to

green in H. caeruleus. It is possible that the two

specimens from Addis Ababa and Galla Lieca,

identified as H. alluaudi Poisson by Poisson

(1955) (Mancini, 1961), are identical with this or

the next species. In A. alluaudi the third anten-

nal segment should be longer than the first seg-

ment. Both specimens (in Museo Civico di Sto-

ria Naturale, Genoa) are females and lack the

antennal segments 3 and 4. In his table to the

African species, Poisson (1943) contrasted the

brilliant metallic colour of H. caeruleus against

the black colour with some violet reflection in

H. violaceus. H. bimaculatus n. sp. conforms in

this respect with H. violaceus and also with H.

violaceus somaliensis Poisson and H. vaillanti

Poisson, 1953, but unlike these species pos-

sesses straight hind femora. H. bimaculatus is

very close to H. pseudopusillus n.sp., with

which it was found together at the type locality.

Apart from the differences in the paramere and

buccula, the white endocorial base in H. pseu-

dopusillus is smaller and bears some minute

hairs (fig. 7b).

Hebrus (Hebrus-type) pseudopusillus n.sp.

(figs. 2d, 6, 7b, 36— 39)

Characterized by straight hind femur and

hind tibia, also in the male (fig. 6). Otherwise it

has the aspect of H. pusillus. The paramere is

like that of H. spinitibialis n.sp. (fig. Sc—e), and

resembles that of H. pusillus (fig. 5a, b), but

with the apical bristles not curled. In contrast to

H. pusillus, the penis lacks sclerotic looped

struts (see page 5). Antennal formula of five

specimens: 6.5—7:4.5—5:7:10—11.5 (not dif-

ferent from H. pusillus). Ground-colour black-

ish, corial area between the veins dark, legs pale

white with fuscous knees, apex of metanotum

incised (fig. 37). In H. pusillus: ground-colour

brown, area between wing veins light, legs uni-

colourous yellow brown, metanotal apex not in-

cised. The general aspect of A. psendopusillus,

straight hind tibia and shape of the paramere is

much like in H. campestris Linnavuori, 1971

(described as a subspecies of H. soudani Pois-

COBBEN: Ethiopian Hebrid fauna 11

son, but now considered a valid species, Linna-

vuori, 1980). In contrast, however, H. campes-

tris is about 0.2 mm smaller and it possesses the

looped penis struts (checked in 2 paratypes).

Length of 5 d: 2—2.3 mm, of 5 2: 2.2—2.4 mm

(A. pusillus is generally 0.2 mm smaller).

Material. Holotype d: Jimma, Hippo Lake,

27.x.1969. Paratypes: Idem, x and xi, 1969, 19

d, 16 ®; Jimma, waterfall, 5.x.1969, 2 2; Jim-

ma airport, 17.x1.1969, 4 dg, 2 2; Jimma, 30 km

en route to Addis Ababa, 23.x.1969, 3 6, 6 ®;

idem, 40 km en route to Addis Ababa,

6.x1.1969, 3 © ; idem, 70 km en route to Addis

Ababa, 23.x.1969, 8 6, 4 © ; Bonga, 29.x.1969,

1 d,2 2; Bonga waterfall, 6 2; Bonga, 8 km en

route to Jimma, 19.x1.1969, 1 2. Holotype and

paratypes in author’s coll.; paratypes in the coll.

of Linnavuori, the Africa Museum, Tervuren,

the Muséum National d’Histoire Naturelle, Pa-

ris, the Zoologisk Museum, Copenhagen, and

the Rijksmuseum van Natuurlijke Historie, Lei-

den.

This is the most common species in the region

investigated. It occurs in a wide variety of hab-

itats, muddy places without running water,

along rivulets, in a wet marl ditch and near the

sprinkle zone of waterfalls. The often simulta-

neous occurrence with other species of the same

Fig. 18. Paramere of Hebrometra pelengei, paratype. Fig. 19. Parameres of Hebrometra niemeri from Tanza-

nia; a—c, Usamba Mts.; d, e, Mt. Rungwe. Fig. 20. Hebrometra malawiensis; a, paramere; b, pseudoscutellum

(compared with that of other Hebrometra, c); d, buccula.

12 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

species group, such as H. katompei and H. spin-

itibialis, makes recognition of single females dif-

ficult. Females of H. pseudopusillus are distin-

guished from females of other species by the

following combination of characters: shape of

buccula (fid. 2b), wing-design (fig. 7b) and

straight hind tibia.

Hebrus (Hebrus-type) katompei Poisson, 1950

(figs. 2b, 4, 7b)

Specimens available from the surroundings of

Jimma are most probably conspecific with H.

katompei, described after 1 d from Zaire. The

curvature of the d hind tibia (fig. 4c) and the

paramere (fig. 4a, b) agree with the figure in

Poisson (1950). The antennal formula of 3 d

and 2 @ is: 6—7:5—6:6—8.5:10—11.5. The

recalculated formula of Poisson reveals a

proportionally smaller second segment

(5.9:4.1:5.9:9.8). The dorsal surface of the hind

tibia of the male has a narrow band of short

spines (fig. 4c), which is not figured by Poisson,

probably because he studied his material under

lower magnification. Otherwise, the species is

very close to H. pseudopusillus (the same type

of buccula, fig. 2b), but in contrast with this

species it possesses the phallic looped struts, as

in H. pusillus. The female of H. katompet is on-

ly recognizable as such on the basis of the buc-

cula (fig. 2b) and the weakly curved hind tibia

(fig. 4d); metanotum broadly, but shallowly in-

cised apically (fig. 40). Another species having

the hind tibia curved in the male sex and the

paramere more or less shaped as in H. katom-

pei, is H. mizae Hoberlandt, 1950, from An-

gola. The antennal formula, however, is devia-

tive (6.1:4:3.5:9.2), the pubescence more dense,

and the hind tibia straight in the 9.

Material: Jimma, Hippo Lake, x-x1, 1969, 15

6,3 23 Ione, Mon IED, 23 8,21 Le nn,

hood IZ 3 Gs Wma, wets,

5.x.1969, 3 d, 2 2; Bonga, 8 km in direction of

mie, 19441969, 2S, 1 2 Gin doe audrors

coll.).

Hebrus (Hebrus-type) spinitibialis n.sp.

(figs. 2c, 7c, 8, 41—46)

Close to A. pusillus, from which it differs in

the following respects as far as the male is con-

cerned: hind tibia conspicuously curved with a

medio-dorsal row of some twelve long spines

(figs. 47, 8a, b; note that the aspect appears

quite different when the leg is slide-mounted

(fig. 8a), or when viewed dry (fig. 8b)), hind tar-

sus with a group of long, extending setae; front

femur with six pegs, front tibia with a ventral

row of oblique spines (fig. 8f); paramere with

apical setae not curled (fig. 8c—e); penis with-

out looped sclerotic struts. Wings somewhat re-

duced, not covering the entire abdomen, mem-

brane with four light areas (fig. 7c). For cuticu-

lar differentiations, see the scanning

micrographs (figs. 41—45). In H. pusillus these

characters are: male hind tibia only weakly

curved, with nine less firm spines (fig. 5d), front

legs without pegs and spines (fig. 5c), paramere

with curled hairs (figs. 5a, b), penis with looped

sclerotic struts. Buccula of H. spinitibialis (fig.

2c) and straight hind tibia of female similar as in

H. pusillus. The antennal formula of 5 d and 5

2 is: 7:5—6:7—8:10—11; thus the length of the

second segment is proportionally longer (0.73 x

the length of segment 3) than in H. pusillus

(0.57). The dorsal pubescence in H. spinitibia-

lis is slightly more dense and longer than in H.

pusillus. The major part of the endocorium and

the area between the exocorial veins is light in

H. pusillus, whereas only a very small basal part

of the endocorium is whitish in A. spinitibialis

(fig. 7c).

Material. Holotype d: Jimma, Hippo Lake,

5.1.1969. Paratyıpes, idem 3 6, | en

27.x.1969, 1 3, 1 Os Jımma, 4310631557

waterfall, 5.x.1969, 2 d, 1 2; Jimma, 70 km en

route to Bonga, 15.x1.1969, 1 d; Jimma, 70 km

en route to Addis Ababa, 23.x.1969, 2 9;

Bonga, along rivulet on chalky loam, 29.x.1969,

10 d, 14 2; Bonga waterfall, 22.x1.1969, 5 6, 6

2 4,1 ©; Gidsha valley, 2400 m, 23.x1.1969, 1

3; Shappa, 7 km from Bonga, xi, 1969, 5 6, 6

2. Holotype and paratypes in the author’s coll.;

paratypes in the coll. of Linnavuori, the Africa

Museum, Tervuren, the Muséum National

d’Histoire Naturelle, Paris, the Zoologisk Mu-

seum, Copenhagen, and the Rijksmuseum van

Natuurlijke Historie, Leiden.

Hebrus (Hebrus-type) gidshaensis n.sp.

(figs. 2c, 7c, 9, 10, 47, 48)

Close to the preceding species, H. spinitibia-

lis, by general facies, wing reduction and wing

pattern (basal white spot of corium further re-

duced than depicted in fig. 7c), similar gula and

paramere, and straight hind tibia in the female.

The male is characterized by stronger curved

hind femur and tibia (fig. 47); consequently, the

hind leg has a remarkably arched appearance,

which obviously plays a role in holding the fe-

COBBEN: Ethiopian Hebrid fauna 13

Figs. 2124. Genital structures; 21. Hebrometra bongaensis, inflated phallus and genital segments; a, right

lateral view; b, endosoma, dorsal view; c, ventral view; 22, Hebrus ruficeps, inflated phallus; left, sinistral lateral

view; right, dorsal view; 23, Hebrus ruficeps, non-inflated phallus, dorsal view with left paramere; 24, Hebrus

pusillus, idem. (co, conjunctivum; ds, ductus seminis; es, endosomal strut; p, paramere; phb, phallobase; phth,

phallotheca; pro, proctiger; py, pygophor; rm, retractor muscle; vss, vesical sclerite).

Figs. 25, 27—29. Hebrus pusillus; 26, Hebrus ruficeps; 25, endosoma of inflated phallus, dorsal. 26, 27a, right

view of genital segments; 27b-d, abdominal apices of pairs in copula; d, d and © somewhat pulled apart to show

the torsion of the d capsule and the intrusion of the phallus apically in the © ovipositor. 25, inflated phallus,

right lateral view. 29, d, ® genital structures fixed in copula, 2 viewed from the right lateral side (note that the

d pygophore is rotated 180°). (ds, ductus seminis; es, endosomal strut; gwth, gutter-shaped wall thickening; gy,

gynatrium; ov, ovipositor; pro, proctiger; py, pygophore; sfc, secondary fecundation canal; sp, spermatheca; vs,

vesica; vss, vesical sclerite).

COBBEN: Ethiopian Hebrid fauna 15

male during copulation. The hind femur is lon-

ger (width of pronotum/femur length is 1.1,

against 1.56 in A. spimitibialis) and bears a row

of tubercles along its inner side (figs. 47, 48).

The row of conspicuous spines of the hind tibia

is longer, reaching nearly the apex (fig. 9b).

Front femur with about eight pegs, front tibia

with a row of blunt pegs (fig. 9a). Antennal for-

mula as in HH. spinitibialis. Despite the strong re-

semblance to the preceding species, H.

gidshaensis has looped struts within the endoso-

ma of the penis. Length of 5 d: 2.5—2.7 mm; of

5 9:2.5 mm (i.e. 0.1 mm larger than the preced-

ing species).

Material. Holotype dg: Gidsha valley,

23.x1.1969. Paratypes, idem, 7 d, 11 2. Holo-

type and paratypes in the author’s coll.; para-

types in the coll. of Linnavuori, the Africa Mu-

seum, Tervuren, the Muséum National d’His-

toire Naturelle, Paris, the Zoologisk Museum,

Copenhagen, and the Rijksmuseum van Na-

tuurlijke Historie, Leiden.

The Gidsha is a flat, open, wet, grassy valley

at an elevation of 2400 m halfway Bonga and

Shappa. The Hebrus species was collected along

a rivulet fed by a nearby spring. That it belongs

to a valid species, reproductively isolated from

its close relative H. spinitibialis, is strengthened

by the fact that the latter species also occurred

in the same habitat. Both species, H. gidshaensis

and H. spinitibialis, were found in closer asso-

ciation with running water than the other He-

brus species treated in the present paper.

Hebrometra n. gen.

Type species: Hebrometra bongaensis n. sp.

Description

Slender, parallel-sided body, three times as

long as wide, total length varying from 2.5—3.0

mm. Antennae and legs longer than in the genus

Hebrus. All constituting parts, inclusive of the

pretarsus, contribute to the greater length of the

legs. Third tibia 1.1—1.3X longer than the pro-

notal width. Hind femur and tibia straight in

both sexes. Ratio: width of head/length of first

antennal segment is: 1.4—1.5 (in Hebrus: 1.7—

2.5). Third antennal segment + 1.2 longer than

first segment (subequal in Hebrus). (The elong-

ate body shape and associated lengthening of

appendages is sometimes not so pronounced

(see p. 18), in which case metrical conditions are

intermediate with those of typical Hebrus). Last

segment of antenna with a pseudoarticulation

(fig. 11a, b), and with a cluster of preapical sen-

SR pegs ( as in Hebrus, fig. 10). General co-

lour dull blackish. Bucculae only weakly devel-

oped (fig. 16). Head with a seta (probably sen-

sory) inserted on tubercle anteriorly of eye (fig.

16, arrow), in addition to the three pairs of tri-

chobothria. Ratio pronotum width/pronotum

length: 1.5—1.6 (in Hebrus ca. 1.8); pronotum

with a more distinct anterior collar than in He-

brus. Paramere elongated, with median process

(fig. 13). Endosoma of phallus largely membra-

nous, with only two, tiny apical sclerites (fig.

21).

Other external and internal structures (repro-

ductive organs, metasternal scent gland, salivary

gland) not differing from typical Hebrus, al-

though the principal salivary gland of the type

species has a more elongated posterior lobe and

a seemingly unicellular anterior lobe.

I decided to erect a new genus for the three

species from Ethiopia listed below, on the basis

of three character states, which separate it from

the genus Hebrus:

1) body elongate with distinctly elongated

appendages, for ratios see above (in Hebrus the

body length is 2.1—2.5 times the greatest width

of the pronotum, and the hind tibia is subequal

to or shorter than the pronotal width). The

lengthening of Hebrometra refers mainly to the

abdomen, head and pronotum remaining more

or less of the original Hebrus-type. The three

species collected by me appear to be adapted to

the sprinkle zone under and beside waterfalls.

Label records of abundant material from other

parts of Africa all point to a similar type of ma-

dicolous habitat. I found the species only in a

narrow, shaded zone, walking on a thin layer of

water running over black rocks covered with

sparse moss and algal patches, often also on wet

vertical cliffs and dark gravel. The long-legged

condition and the longer antennae may be an

adaptation to this niche. Other hebrid species

mentioned in this paper, as well a Microvelia

spec. (Veliidae), lived very close to Hebrometra,

but sharply separated from it on stagnant water

or not-flooded substrate. In all localities in

Ethiopia, Hebrometra was accompanied by Sal-

dula jihafana Brown (Saldidae); sometimes both

insects were entirely intermixed.

2) Presence of a seta inserted on an elevation

in front of the eye. The same character occurs

also in the oriental hebrid genus Timasıns,

which has superficial resemblance with Hebro-

metra and which lives under similar conditions

(N. N. Andersen, pers. comm.).

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

3) Bucculae weakly developed (fig. 16). In all

other hebrids the bucculae seem to be well pro-

nounced (fig. 2).

4) Paramere elongate (figs. 13—19).

I consider these four characters derived in

comparison with their respective conditions in

Hebrus. Moreover, the adaptive shift towards

the very humid restrictive zone bordering wa-

terfalls warrant the distinction of a separate

clade. Provisionally, Hebrometra could readily

be evaluated as the sister group of Hebrus. In

considering cladistic relations of hebrid genera,

Andersen (1981) also suggests the possibility

that Hebrometra, in spite of its distinctive hab-

itus, could have arisen from some subordinate

group within Hebrus. The recent discovery of a

new Hebrometra species with one of the four

most distinctive generic characters only weakly

developed (see p. 18) favours this view. If the

Hebrometra clade is classified as separate genus,

as proposed here, the genus Hebrus then be-

comes a paraphyletic group. The fact that my

investigation of only four waterfalls, over a dis-

tance of about 40 mi. in the Kaffa province of

Ethiopia, yielded three species of the new ge-

nus, suggests that this particular habitat har-

bours more species in Africa.

Hebrometra bongaensis n.sp.

(figs. 12a, 13b, 16a, 17, 21, 49— 53)

Body, predominantly dull black, clothed with

short adpressed hairs. Head and pronotum of-

ten with an inconspicuous blue-metallic hue; for

cuticular details see scanning micrographs (figs.

50—53). Head with several erect hairs in addi-

tion to the trichobothria and the microsetae.

Buccula weakly developed (fig. 16a); rostrum

light-brown, apex reaching level of hind coxae.

Vertex behind eyes, collar of pronotum, aceta-

bulae, and sometimes also lateral edges of pro-

notum, yellow-brownish; velvet spot between

eye and posterior trichobothrium. Apex of tri-

angular metanotum weakly notched (fig. 51).

Forewings reaching wholly or nearly the apex

of abdomen, black, with grey-pruinose basis of

exo- and endocorium, endocorium neatly de-

marcated from membrane; whole surface of co-

rium regularly covered with short, golden-sil-

very hairs, adpressed in caudal direction; mem-

brane shining, irregularly blotched with black

and pruinose areas; a lateral oblique line and an

apical spot whitish. Extremities unicolorous

light brownish. Antenniferous tubercle promi-

nent; antennal formula: 9:7:11.5:13; segments

with short hair-dress, and some scattered, semi-

long, erect setae on 2—4. Legs slender and long,

with short pubescence; length in mm of femur,

tibia and tarsus (excluding pretarsus) of leg

1:20—21—7.5, of leg 2:24—25—8, of leg

3:30—36.5—11, respectively; cleaning comb of

tibia 1 with ca. 17 spines, of tibia 2 with ca. 9

spines, of tibia 3 poorly developed. Paramere

(fig. 13) with rounded apex and triangular medi-

an process, bearing a sensillum. The phallus in

artificially inflated condition is shown in fig.

21a—c. Length of 8 d: 2.7—3.0 mm, of 7 9:

2.9—3.0 mm.

Holotype d: Bonga reusel, 22.x1.1969.

Paratypes, idem, 9 8,5 2, 1 larva Voudemmiies

DAMON Wes, Wo Jensen), 6 8,5 2, 70 km

from Jimma en route to Addis Abeba,

23.x.1969; 1 d, 2 2, Sakka, 20 km South of Jim-

ma, 19.x.1969. Holotype and paratypes in the

author’s coll; paratypes in the coll. of Linnavu-

ori, the Africa Museum, Tervuren, the Zoolog-

isk Museum, Copenhagen, the Muséum Na-

tional d’Histoire Naturelle, Paris, and the

Rijksmuseum van Natuurlijke Historie, Leiden.

Hebrometra pelengei (Poisson, 1954)

(= H. mancinii Poisson, 1955, nov. syn.).

(figs. 12b, 14, 16c, 18)

Hebrus pelengei was originally described

from Zaire (Gorges de la Pelenge, 1.250—1.600

m, 20—23.vi.1947). Of the series of 38 speci-

mens I was able to study 1 d and 1 2 deposited

in the Smithsonian Institution, Washington, and

4 8 in the Tervuren Museum. Since Poisson did

not designate types, I selected 1 d as the holo-

type (deposited in Tervuren) and labelled the re-

maining specimens as paralectotypes. The spe-

cies is characterized by complete lack of hairs

on the endocorium. Examination of the type of

Hebrus mancinii from Addis-Abeba, Ethiopia

(in Museo Civico di Storia Naturale, Genua) did

not reveal specific differences with A. pelengez.

The drawings of the parameres of H. pelengei

(Poisson, 1954, fig. 4c) and A. mancinu (Pois-

son, 1955, fig. 4b, c) suggest some dissimilari-

ties, but these fall within the range of variability

which I found in conspecific material from lo-

calities in Zaire and Ethiopia (see figs. 14 and

18). I collected 1 d and 1 ® in Jimma,

18.x.1969; idem 1 6, 5.x.1969; idem, 1 ©,

1.x1.1969; 1 d, Sakka, 20 km south of Jimma,

19.x.1969 (simultaneous with H. bongaensis).

Some specimens have the endocorium entirely

hairless as in the type series of H. pelenget,

others possess a central tuft of a few transversal

silvery scales.

COBBEN: Ethiopian Hebrid fauna 1177

H. pelengei differs in the following respects

from H. bongaensis: Last antennal segment

1.4—1.5X the length of third segment (1.1—

1.2X in À. bongaensis). Bucculae less developed

(fig. 16c). Apex of metanotum brownish. Base

of endocorium white, remainder part brown

and not well demarcated from membrane, with-

out hairs or with only a few silvery hairs in the

centre (fig. 12b). Exocorium dark with ad-

pressed hairs on the veins and a strip of minute

silvery hairs on the area between the veins. Male

genital capsule shorter and with longer setae.

Paramere with nasiform median process of

varying shape (figs. 14, 18) without sensillum

on the median process.

Hebrometra niemeri n. sp.

(figs. 11, 12c, d, 15, 16b, 19)

Differing from the previous two species in the

shape of the bucculae (fig. 16b) and wing-pu-

bescence (fig. 12c). The basis of the endocorium

is white like in A. pelengei. The endocorium

bears a dense layer of transverse silvery setae;

the area between the exocorial veins also with

silvery setae, oriented under an angle different

from the half-long golden setae on the veins.

Paramere varying (fig. 19 a—e), mostly with

blunt apex and irregularly notched median pro-

cess without sensillum. The antennal formula

and tendency for more brownish areas on pro-

notum and metanotum is more like in H. pel-

enger. Length varying from 2.5—2.8 mm (6)

and 2.9—3.3.m (?).

It is almost certain that this species refers to

Hebrus mancinii f. panganii Poisson, 1963, al-

though I was unable to study the type material.

The holotype was said to be deposited in the

Muséum National d’Histoire Naturelle, Paris,

but I was kindly informed by dr. Carayon that

it is not available there. However, Poisson’s de-

scription clearly mentions the presence of the

silvery hair-layer on the endocorium. The type

material is from the same mountain (Usambara)

in Tanzania, from which now abundant recent

material is available. The reason that I introduce

here a new name is the following. Above it was

concluded that H. mancinti is a synonym of H.

pelengei. The Usambara material described as

forma panganii is clearly another species.

According to the International Code, 1961, arti-

cle 17, a name published as “variety” or “form”

may only remain available as species name, if it

was published before 1961.

I name the species Hebrometra niemeri in

| memory of my host in Jimma, Hugo Niemer

(WHO, med. doct.) who, with his daughter,

tragically perished in Malawi in 1976. The spe-

cies has a wide south-eastern distribution from

Ethiopia up to the southern edge of the Cape

Province. The great majority of the Tanzanian

specimens collected in the Usambara Mountains

at different altitudes are characterized by long

setae on the inner corial vein and along the

oblique margin of the membrane (fig. 12d);

third antennal segment two times the length of

segment two or slightly more; legs dark shaded;

membrane reaching the apex of the abdomen.

Another population from Tanzania, living on

Mt. Rungwe at a distance of some 700 km from

the Usambara Mts., has no such obvious long

setae on the corium, the third antennal segment

less than two times the length of the second seg-

ment, legs lighter coloured, and the membrane

somewhat reduced. The parameres in both pop-

ulations are not discriminative. In spite of the

rather constant differences mentioned I refrain

for the present from dividing up the species in

subspecies. A long series from Uluguru Mts.,

about half-way the two localities mentioned

above, is more or less intermediate in the char-

acters recorded above, suggesting clinal varia-

tion. Other material from Ethiopia, Malawi, and

S. Africa lacks the long corial setae, or has the

setae of intermediate length, whereas other

characters resemble those of the Usambara

specimens.

Material. Ethiopia: Holotype &, Bonga,

22.x1.1969, in author’s coll. Paratypes, idem, 2

3; Tanzania: East Usambara Mts., Sigi River,

750 m, 4 d, 4 2, numerous larvae, 15.v11.1980,

leg. M. Stoltze & N. Scharff (in Zoological Mu-

seum, Copenhagen); East Usambara Mts.

Amani, Dodwe Stream, 1000 m, 3 d, 1 larva,

~ 5.vii1.1979, leg. M. Stoltze (in Zool. Mus. Co-

penhagen); Amani, seeps near Sigi river, CL

VI ©, Ile 2, 3 lees Win IO les, |. IC

Polhemus (in Polhemus coll.; Cobben coll.);

idem) Sigaunivers) ©9996 one loan O79:

leg. J. T. Polhemus; idem, Dodwe Stream, CL

IE nor TEE PollaenuseZaire:

Kivu, Bulingwe (Runingo), Itombwe plane, Est

(pierre mouillée), 6 d, 10 9, 6.v1.1958, leg. G.

Marlier (in Mus. Tervuren); Uvira, Vallée de la

Miele (cocherssrursselanu) 3 G, ZS

26.v.1958, leg. G. Marlier (in Mus. Tervuren);

Uvira, Mulenga-Lemera, 5 ©, 19.1.1955, leg. G.

Marlier (in Mus. Tervuren). West Usambara

Mts., Bumbuli, 1300 m, 8 6, 6 ®, 3 larvae,

1.viii.1980, leg. M. Stoltze & N. Scharff (in

Zool. Mus. Copenhagen and Rijksmus. Nat.

18 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

Hist, Leiden); Mt. Rungwe SW. 1900 m, 13 d,

1 2,1 larva, 20.vii.1980, leg. M. Stoltze & N.

Scharff (in Zool. Mus. Copenhagen and

Rijksmus. Nat. Hist, Leiden); Mts. Uluguru,

Morning Side (sur roche ruisselante), 1600 m,

16 d, 27 2, 21—29.v1.1971, leg. L. Berger, N.

Leleup, J. Debecker (in Mus. Tervuren). S. Afrı-

ca: Natal, Cascades, Mont aux Sources, 1 d, 1

2, 5.11.1968, leg. J. Munting (coll. ACNC);

Natal, Gillits, Pinetown district 1 9, 21.x1.1963,

leg. B. & P. Stuckenberg; Cape Province, Gra-

hamstown, 1 ®, 1924, leg. M. L. Winslow;

any IL Le 22.001082, lez ar Io Mi, Camee

(coll. USNM); Rust in Vrede, Swartberg, nr.

Oudtshoorn, 1 2, 1.1.1927, leg. G. E. Hutchin-

son (coll. BMNH).

A NEW HEBROMETRA SPECIES FROM MALAWI

WITH A MORE HEBRUS-LIKE FACIES

Hebrometra malawiensis n.sp.

(fig. 20)

General facies like typical members of the ge-

nus Hebrometra (see p. 15 and fig. 12) but with-

out such pronounced elongated appendages.

Dull, black, with short pubescence and several

dark erect semilong hairs on head; pronotum

with weakly metallic blue shine. Buccula only

weakly developed (fig. 20d). Distal lateral edges

of pronotum, underside of pronotal collar, ac-

etabula and rostrum brownish. Antennae dark

brown with short pubescence; segment 2 with

some erect longer setae like on segments 3 and

4; last segment with pseudoarticulation; ratio of

segments of holotype (d) and paratype (@), re-

speetigely: TORO ande ZES7ENO Led

brownish, femora proximally yellowish brown;

third tibia basally very weakly sinuate in both

sexes; ratio length tibia 3 to maximal width of

pronotum 0.9—1.0. Pseudoscutellum (fig. 20b)

more clearly bordered and tapered than in other

Hebrometra spp. (fig. 20c). Forewing black, on-

ly extreme base of endocorium light; regular

golden pubescence on veins, silvery hair-cover

in between veins (much like in Hebrometra nie-

meri, fig. 12c). Paramere elongated, of the He-

brometra-type (fig. 20a). Phallus with only two

apical sclerites. Length and width of d: 2.1.

mm, 0.8 mm; of 2 2.4 mm, 0.98 mm.

Holotype d: Malawi, Zomba Plateau, CL

985, 1.111.1979, leg. J. T. Polhemus (in Polhemus

coll.). Paratypes, idem, 7 6, 7 © (also in au-

thor’s coll.).

Discussion: H. malawiensis is in several re-

spects intermediate between typical Hebrus spp.

and the three species of Hebrometra described

in this paper. The ratio width of head: length of

first antennal segment is 1.8 (1.7—2.5 in He-

brus, 1.4—1.5 in Hebrometra). The ratio length

of tibia 3: width of pronotum is 0.9—1.0 (0.8—

0.9 in Hebrus, 1.10—1.3 in Hebrometra). The

ratio pronotum width: pronotum length is 1.7—

1.8 (1.8—1.9 in Hebrus, 1.5—1.6 in Hebrome-

tra).

On the basis of the general facies, weakly de-

veloped buccula and shape of paramere, this

species is assigned to Hebrometra. The setifer-

ous tubercle in front of the eye is less pro-

nounced and the appendages are less elongated

than in typical Hebrometra. On the basis of this

character state, I conclude that H. malawiensis

is one of the first representatives of the Hebro-

metra clade in which the elongation of the legs

as a further adaptation to the sprinkle zone has

not yet been fully expressed (see p. 15).

ACKNOWLEDGEMENTS

The expedition to Ethiopia was made possible

by financial support of the Netherlands Foun-

dation for the Advancement of Tropical Re-

search (WOTRO). My thanks are due in partic-

ular to the family Niemer for housing and trans-

port facilities in and around Jimma, and to the

Bonga Mission for the hospitality received. The

following colleagues assisted in some way or

other with regard to the material treated in this

paper: N. M. Andersen (Copenhagen), J. Car-

ayon (Paris), R. C. Froeschner (Washington), L.

Hoberlandt (Prague), R. Linnavuori (Somerso-

ja, Finland), R. Poggi (Genoa), J. T. Polhemus

(Englewood), G. Schmitz (Tervuren). I am

greatly indebted to N. Andersen for valuable

comments on the manuscript. Mrs. Karin

Heming-v. Battum (Edmonton, Canada) sup- |

plied material of Hebrus pusillus, fixed in copu- |.

la. S. Henstra and F. Thiel of the Technical and

Physical Engineering Research Service, Wage-

ningen, made the photomicrographs with the

Jeol scanning electron-microscope.

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——, 1978. A new family of semiaquatic bugs for

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COBBEN: Ethiopian Hebrid fauna 19

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some species of the adjacent countries. 8. Addi-

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tilen und semiaquatilen Hemipteren von Sumatra,

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Civ. Nat. Genova 72: 31—46.

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20 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

Figs. 30—35. Hebrus gerardi, scanning electron micrographs of cuticular structures; 30, dorsal view of thorax

(80x); 31—35, detail of right-mid part of pronotum; 31 (300X), 32 (583X), 33 (1166x), 34 (1750x), 35

(5830x).

COBBEN: Ethiopian Hebrid fauna 21

Figs. 36— 41. Scanning electron micrographs; 36—39, Hebrus pseudopusillus; 36, dorsal view of thorax and

wing bases (58X); 37, metanotum (116%); 38, 39, detail of right-mid part of pronotum, 38 (583X), 39 (1750x);

40, Hebrus katompei, metanotum (116); 41, Hebrus spinitibialis, anterior part (32X).

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

Figs. 42—47. Scanning electron micrographs; 42—46, Hebrus spinitibialis; 42, metanotum (116X); 43—45, de-

tail of right-mid part of pronotum, 43 (250X), 44 (1750X), 45, sieve-pore (5830X), 46, setal comb of d hind

tibia (175%); 47, Hebrus gidshaensis, 3 hind leg (58x).

COBBEN: Ethiopian Hebrid fauna 23

Figs. 48—53. Scanning electron micrographs; 48, Hebrus gidshaensis, pegs on d hind femur (350X, see Fig.

47); 4953, Hebrometra bongaensis; 49, last antennal segment with striation of pseudojoint, arrow (350X); 50,

head (93x); 51, thorax (57); 52, endocorium (1750X); 53, microsetae on membrane (1750X).

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 1 (1982)

Figs. 54—59. Scanning electron micrographs; Hebrus pusillus, specimens from the Netherlands, male genitalia;

54, left view of genital capsule (175%); 55, posterior view of intromittent organ with left paramere (204X); 56,

inner surface of left paramere (583); 57, sensilla on inner outgrowth of paramere (5830x); 58, dextral side of

genital segments with artificially erected phallus, apex of vesica ruptured so that internal struts are exposed

(87%); 59, vesical struts (583X).

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INHOUD

H. LEEREvELD. — Anthecological relations between reputedly anemophilous flow-

ers and Syrphid flies. II]. Worldwide survey of crop and intestine contents of

certain anthophilous Syrphid flies, p. 25—35.

Tijdschrift voor Entomologie, deel 125, afl. 2 Gepubliceerd 15-VII-1982

AA ee I

ANTHECOLOGICAL RELATIONS BETWEEN REPUTEDLY

ANEMOPHILOUS FLOWERS AND SYRPHID FLIES.

III. WORLDWIDE SURVEY OF CROP AND INTESTINE

CONTENTS OF CERTAIN ANTHOPHILOUS SYRPHID FLIES

H. LEEREVELD

Hugo de Vries-laboratorium, Amsterdam, Netherlands

ABSTRACT

A worldwide survey of the pollen present in the digestive tract of about 160 syrphid flies

of the genera Melanostoma and Platycheirus and some related taxa confirms the preference

of most representatives of this group of Diptera for pollen of nominally anemophilous

plants such as Plantago, Typha, grasses and Cyperaceae. In many cases they had been feed-

ing on pollen of such plants exclusively. The results of this survey are discussed in connec-

tion with (especially some ecological) conditions and with the pollination syndromes of

such plants.

INTRODUCTION

Representatives of the genera Melanostoma

and Platycheirus (= “M-P group”) are effective

pollinators of Plantago lanceolata L. (Stelleman

& Meeuse, 1976, see pt. 1 of this series), and al-

so exhibit consistent feeding visits to grasses

and other anemophiles (Van der Goot & 'Gra-

bandt, 1970; Leereveld et al., 1976; and person-

al observations by the present author, Meeuse

and Stelleman; to be summarised by Stelleman,

in prep.). Grabandt (in an unpublished report)

and Holloway (1976, New Zealand) confirmed

the ambogamous nature of Plantago lanceolata,

already pointed out by several older workers,

by such gut content studies of flies of the M-P

group. Strikingly high percentages of pollen of

anemophiles, particularly of P. lanceolata, were

recorded, the first author also mentioning ap-

preciable quantities of grass pollen.

In a description of Platycheirus species occur-

ring in Norway, Nielsen (1971) reported,

among others, Plantago lanceolata as a source

of food of several species. Waitzbauer (1976) al-

so noticed a relation between representatives of

the M-P group and anemophilous plants when

he recorded insect visits of three species of

Platycheirus to Typha angustifolia in Austria.

More recently Mesler (1977) noted the frequent

feeding visits of the related Mesograpta margi-

natus to Plantago lanceolata in Michigan,

U.S.A. His descriptions of the feeding behav-

iour of this syrphid species resembles that ot

members of the M-P group studied in the Neth-

erlands and in Germany. The present study was

aimed at establishing a world-wide validity of

the already signalised anthecological relation

between members of the M-P group of hover

flies and anemophilous plant taxa in western

Europe. To this end the gut contents of M-P

syrphids hailing from various parts of the world

were analysed palynologically.

MATERIAL AND METHODS

The flies, if not collected by members of our

research team, were kindly provided by various

institutes and collectors in this country and

abroad. The total amount of flies at our disposal

was about 160 from 15 different countries.

Apart from M-P species from Europe material

was received from North America and New

Zealand. All specimens from New Zealand were

referred to Melanostoma fasciatum, a species

not known from Europe but resembling our

M. scalare very closely. Two specimens be-

longed to Platycheirus immaculatus, a rare spe-

cies in the Netherlands. Of the related genus

Pyrophaena 10 Norwegian specimens of

P. granditarsa were studied. The 54 localities

are not only widely scattered over the world,

but are also situated at different altitudes (from

sea level to about 1500 m alt.).

Table 1 shows the number of localities per

26 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 2 (1982)

Table 1. Countries, number of localities and number of specimens used for analysis of

P y

gut contents. (Dutch specimens refer to samples; extensive data in Van der Goot &

Grabandt (1970).

Country no. of no. of no. of specimens

specimens localities per locality

Canada 5 3 3,2X1

Czechoslovakia 4 2 3,1

England 43 1 43

Finland 9 2 5,4

France 1 1 1

Italy 6 3 3x1

Netherlands 3 3 3x1

New Zealand 15 2 9,6

Norway 44 11 11, 9, 2X5, 4, 3, 2X2, 3X1

Roumania 8 4 4,2, 2X1

Spain 2 2 2x1

Sweden 15 10 4,3,2,7X1

U.S.A. 2 1 2

U.S.S.R. 19 8 7,6,6X1

Yugoslavia 1 1 1

country and the number of specimens per local-

ity. It appears from this table that the large ma-

jority of the specimens came from England,

Scandinavia and Russia, which (owing to the lo-

cal floral composition) may cause a certain bias

when conclusions are drawn from the analyses

of the gut contents. The number of specimens

from Canada and the USA is unfortunately

rather scanty. The preparation was carried out

as follows. |

Every individual specimen was treated in an

almost boiling 10% KOH solution for about 5

min. in order to soften the outer chitin skeleton

for further manipulation. If the boiling period is

extended the fly may come apart and pollen es-

capes from the digestive tract, so that excessive

boiling must be avoided. Since many specimens

had been mounted on needles and the holes in

their bodies might cause a leakage of pollen dur-

ing the boiling, it was deemed recommendable

to boil needle-mounted flies needle and all. Still,

during excessive boiling some pollen may es-

cape through the perforations in the body

alongside the needle. Another complication is

the chemical reaction between KOH and the

metal of the needle often leading to the forma-

tion of ferric hydroxide which may precipitate

in the form of flakes, thus obscuring the micro-

scopical observation. Also for this reason the

treatment with hot KOH solution must be kept

as short as possible. To the KOH solution with

the specimen a few drops of a solution of safra-

nin in distilled water are added. The safranin

stains the exine, which is indispensable for the

subsequent microscopical identification. After

the insect has been rinsed under a strong jet of

distilled water (so as to rinse away pollen grains

adhering to the body), it can be dissected in a

simple way on a glass slide under a binocular

microscope. By means of fine needles and

tweezers the outer skeleton is ruptured in order

to expose the digestive tract. When excessive

amounts of pollen were present (a swollen ab-

domen of a fly is indicative!) two slides were

prepared, one of the anterior part of the tract (in

head and thorax) and one of the posterior part

(in the abdomen). In this way at least some indi-

cation may be acquired of a possible change in

the feeding behaviour in a fairly short time-

span. A second advantage is a practical one: a

very large quantity of pollen in one slide is not

sufficiently surveyable. Finally the pollen pre-

paration is mounted in glycerin jelly and the

cover glass is sealed off on all sides by means of

paraffin wax.

Since our palynological department has an ex-

tensive standard pollen collection of slides pre-

pared by means of the customary acetolysis

process of Erdtman, it would have been an ob-

vious choice if the same treatment would have

been applied to the pollen extracted from flies,

but in view of a desirable minimum of loss of

pollen and a preferably least complicated proce-

dure, the above-mentioned simple method of

preparation was followed.

The pollen identifications were primarily

based on Erdtman et al. (1961). Pollen of Plan-

taginaceae, irrespective of the country of origin,

LEEREVELD: Anthecological relations 3 27

was referred to four types: the P. coronopus, the

P. lanceolata, the P. major and the P. maritima

types. In addition the slide collection was con-

sulted and also a small reference collection of

pollen of P. lanceolata, P. media, P. major,

P. coronopus, and P. maritima especially made

for the purpose and prepared in the same way as

the flies by treating the whole anthers of these

plants in KOH, etc. This collection, when used

for comparison, already indicated that the pol-

len grain diameters reported in the above-cited

Scandinavian pollen flora are of very little use;

the variation in size is particularly great in

P. lanceolata, even in pollen extracted from a

single anther. Pollen of grasses and Cyperaceae

was not further identified, firstly because this

would require an appreciable experience (which

was lacking) and secondly because there was no

suitable reference collection available. During

the identification the pollen of reputedly or pos-

sibly anemophilous forms such as species of Ty-

pha and Salix were separately recorded.

The counts were performed as follows: if the

number of grains did not exceed 500 they were

all idenufied as far as possible, but if it exceeded

500, a homogeneous portion of the slide was se-

lected at a magnification of about 400X and

three separate counts of 100 grains were made

of which the mean value was taken.

RESULTS

Some specimens studied did not have any pol-

len in their digestive tract. This may be attrib-

utable to three causes, viz., (1) a catch before

anthesis of the food plants had taken place (not

very likely), (2) a poor or diseased condition of

the fly (not so obvious either), or (3) consump-

tion of nectar only (this was noticed by personal

observations of particularly Platycheirus scuta-

tus, while nectar consumption of M-P syrphids

has been also reported by other workers).

The majority of the specimens studied yielded

substantial amounts of pollen, as already men-

tioned, but the effect of the digestive juices on -

the grains, especially on the ectexine sculpture,

renders well-digested pollen unrecognisable

owing to a reduced affinity to the stain, to the

corrosion and abrasion of the surface, shrinkage

and shrivelling of the grains, etc. The latter

damage is partly caused by dehydration and by

the mechanical action of compression and peri-

staltic movements of the gut.

Fortunately there are usually enough better

preserved, identifiable grains to get some idea of

the diet. An additional complication is that the

staining of the contents of the digestive tract

starts at the oral end and not or hardly from the

anal end, so that the pollen grains consumed lat-

est are usually also the best stained. Occasional-

ly the staining was poor because the throat was

blocked during the boiling process. Grass pollen

seems to have a lower affinity to the safranin

dye than dicotyledonous pollen, and pollen

grains of Cyperaceae have hardly any, but, even

when not stained, are conspicuous by their own

brownish colour. Among the flies there were

some caught at the beginning of this century.

Without exception the pollen in their digestive

tract had undergone a process of aging which

gave them the appearance of fresh pollen sub-

jected to the standard Erdtman process of ac-

etolysis. This may be a way to procure naturally

aged pollen of certain species.

The results of the analyses are shown in table

2 in which four categories are distinguished,

three consisting of pollen of easily identifiable

anemophilous forms, and the fourth including

the remainder. If it was deemed to be relevant

(and possible!) to provide some additional in-

formation concerning other anemophiles, the

Plantago types other than the P. lanceolata

type, and the relative amounts of pollen types

present; these are separately entered in the

fourth column. Although the specimens were

always rinsed to wash away any pollen attached

to the outer surface of the body, some may have

got into the slide, but at the worst the number,

as far as can be ascertained, does not exceed 5

grains per fly and is, therefore, negligible. When

only very few identifiable pollen grains were

present in the slides the absolute number (and

no percentage) is given, while the “+” symbol

means that the category is ila only spo-

radically.

From previous studies of our research group

(see parts I and II of this series) the feeding hab-

its of the pollenconsuming members of the M-P

type are dependent on (a) the local habitat and

stands of vegetation, (b) the time of the day

(consumption mainly taking place before 10.00

hrs or 11.00 hrs) and (c) the weather. Such data

were not available for the specimens sent to us,

but one may infer that especially the first factor

is important because the diet sometimes clearly

reflects the only available sources (see under

“Discussion”).

After excessive feeding in the early morning

pollen is usually present in the animal body for

several hours after the last consumption has tak-

en place and animals caught later in the day still

Table 2. Results of analyses of gut contents of syrphid flies of the M-P group. Explanation in text. |

Abbreviations used: tricolp.: tricolporate (dicotyledonous pollen type; verruc(ulate), psil(ate), fenestr(ate), |

retic(ulate), percolp(ate), scabr(ate): Compos. — Ligul. = Compositae Liguliflorae. Records in parentheses: |

duplicate slides.

Slide no. and

country of

origin

® PAL 7, Netherl.

PAL 23, Italy

(PAL 24, Italy

PAL 31, Italy

PBL 21, Roumania

PBL 25, Roumania

PBL 26, Roumania

PBL 70, England

(PBL 71

PBL 75, England

PBL 78, idem

PBL 84, idem

PBL 89, idem

PBL 94, idem

PBL 99, idem

PCL 7, idem

PCL 8, idem

PBL 79, idem

PBL 81, idem

PBL 82, idem

PCL 1, idem

PCL 6, idem

dg PAL 14, Netherl.

PAL 56, USSR

PAL 72, Norway

PAL 79, Norway

PAL 82, Norway

PBL 52, Sweden

PBL 83, England

PBL 87, England

PCL 11, England

PBL 74, England

PBL 95, England

® PAL 9, Netherl.

PAL 17, Czechosl.

PAL 21, Yugosl.

PAL 25, Spain

(PAL 26, Spain

PAL 51, USSR

(PAL 52, USSR

PAL 55, USSR

PAL 75, Norway

PBL 30, USA

PBL 31, USA

TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 2 (1982)

pollen composition recorded

Plantago

lanceolata

type

Melanostoma mellinum

grasses

100%

100%)

100%

67%

314

100%

100%)

100%

98%

62%

94%

88%

89%

100%

95%

86%

94% 6%

62% 38%

100%

96%

98%

4% 96%

10% 90%

100%

18% 80%

30% 70%)

100%

100%)

100%

28% 48%

95%

100%

Cyperaceae

other ones

14% Tiliaceae, 14% Pteridophyta

5% other kinds

7 other ones, 1 prob. Caryophyllac.

P. major type (= P. media): +

tricolp.-reticulate: +

2% tricolp.-echinate

38% Compos.-Ligul.

6% Compos.-Ligul.

12% dicotyl., 2 types

11% mainly tricolp.-retic.

5% fenestrate

(prob. Compos.-Ligul.)

14%, mainly Rosaceae

4% tricolp.-verrucul.

1% tricolp.-echinate,

1% tricolp.-reticul.

24% mainly of one type

5% fenestrate

(prob. Compos.-Ligul.)

LEEREVELD: Anthecological relations 3 29

Table 2 (cont)

Slide no. and pollen composition recorded

pot grasses Plantago Cyperaceae other ones

origin ] Di

anceolata

type

(Melanostoma mellinum, cont.)

BEBA 27, Spain 95% 5% of one type (rosaceous)

PAL 81, Norway 67

9 PAI 32, Finland 100%

(PAL 33, Finland 100%)

PAL 34, Finland 100%

(PAL 35, Finland 100%)

PAL 36, Finland 100%

PAL 53, USSR 30% 70%

(PAL 54, USSR 100%)

PAL 73, Norway 100%

(PAL 74, Norway 100%) Pinus (a single grain)

PBL 53, Sweden 85% 15%, 2 Picea, 1 prob. Caryophyllac.

8: PAL 57, USSR 90% 10% triporate-psilate,

one Pinus grain

PBL 34, Sweden 100% triporate-psilate: +

other types: +

98% Rosaceae

(Potentilla anserina subtype,

2% P. erecta subtype),

other types: +

E225 Italy 32% 68% dicotyledonous,

tricolp.-scabrate

PAL 77, Norway 45% 55%, mainly of one type

(PAL 78, Norway do.) (do.)

PBL 22, Roumania 18% prob. 82% Caryophyllac.

PBL 23, Roumania 88% P. major type,

10% dicot. tricolp.-reticulate,

2% other types

PAL 30, Italy 93% P. major type, 7% Rosac.

PAL 58, USSR 100% P. major type (= P. media)

PBL 24, Roumania 104 tricolp.-scabr.,

4 P. major type (= P. media),

other types: 3

6 PAL 1, France 100%

PAL 76, Norway 100% fenestrate type

PAL 80, Norway mainly Ericaceae and Rosaceae,

Caryophyllac.: +

2 PBL 35, Sweden contained no pollen

Melanostoma scalare

2 PBL 76, England 100%

PBL 85, England 100% Ranunculaceae: +

PBL 88, England 100% tricolp.-echinate: +

PBL 90, England 100%

PCL 3, England 100% 3h

(Melanostoma scalare, cont.)

PEReT 5, England 100%

PCL 12, England 100% triporate-psilate: +

d PBL 68, England 100% dicots: +

PBL 72, England 100%

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 2 (1982)

Table 2 (cont.)

Slide no. and pollen composition recorded

Bren, oi grasses Plantago Cyperaceae other ones

5 lanceolata

type

PBL 86, England 100%

PBL 92, England 100% dicots: +

2 PBL 77, England 93% 44 grains tricolp.-echin.,

3 other ones

PBL 91, England 91% 9%

PBL 98, England 88% 12% mainly Ranunculaceae

PCL 4, England 96% 4% Rosaceae

PCL 10, England 79% 21% tricolp.-reticulate

3 PBL 67, England 90% 10% tricolp.-echinate

PBL 96, England 94% 6% tricolp.-echinate

2 PBL 80, England 100%

PBL 93, England nearly 100% Rosaceae

PCL 2, England 23% 77% tricolp.-reticulate

3 PBL 69, England sl 100% Compos.-Ligul.,

other types: +

1 indeterminate

d PAL 83, Norway 100% Rosaceae

PAL 84, Norway 100% Compos.-Ligul.

PBL 40, Sweden 100% mainly 2 types of dicots

Melanostoma fasciatum

® PBL 51, New Zeal. 100%

3 PBL 43, New Zeal. 100%

PBL 44, New Zeal. 100%

PBL 45, New Zeal. + 100% +

PBL 46, New Zeal. 77% 33% mainly 4 types of dicots

PBL 47, New Zeal. 65% 35% do.

PBL 48, New Zeal. 100%

PBL 50, New Zeal. 100%

PBL 49, New Zeal. esophagus 5% 80% Malvaceae,

15% Compos.-Ligul.

gut: 6% 89% Compos.-Ligul.,

5% Malvaceae, other types: +

2 PBL 59, New Zeal. 100% de

PBL 61, New Zeal. 81% 19%

PBL 64, New Zeal. 91% 9%

d PBL 62, New Zeal. 84% 1% 15%

PBL 63, New Zeal. 100% +

2 PBL 60, New Zeal. 100%

Platycheirus clypeatus

2 PAL 19, Czechosl. 1 2 not identifiable

(PAL 20, Czechosl. 6 3) (2 not identifiable,

4 of different types)

PAL 28, Italy 100%

PAL 29, Italy 100%

PAL 40, Finland 100%

PAL 46, Canada 100%

(PAL 47 100%)

LEEREVELD: Anthecological relations 3 31

Table 2 (cont.)

ET CORET Pre Mer ENE eri LIT di SEO Ri a

Slide no. and pollen composition recorded

country of grasses Plantago Cyperaceae other ones

ra lanceolata

type

EE EN me | lia i ASA

PBL 3, Norway 100% + triporate-scabrate

PBL 4, Norway 100%

PBL 27, Roumania 100% +

PBL 28, Roumania 100%

d PAL 99, Norway 87% 12% Rumex-like, 1% differ.

(PAL 100, Norway do.) (do.)

PBL 1, Norway 130 4 Rumex-like, 4 different

PBL 5, Norway 2 1 1 tricolpor.-reticul.

PBL 6, Norway 100% do.

PBL 7, Norway 100%

PAL 18, Czechosl. 100%

PBL 8, Norway 100%

® PAL 37, Finland 100%

(PAL 38, Finland 100%)

6 PBL 33, Sweden 100%

PBL 54, Sweden 11% 86% 3%

® PAL 39, Finland 72% Comp. Ligul.,

28% Melampyrum

PBL 2, Norway + 100% dicotyl, pericolp.-scabrate,

other types: +

é PBL 41, Sweden 47% 4% 49%, mainly 2 tricolp.

dicot types

PAL 45, Canada contained no pollen

PBL 38, Sweden contained no pollen

Platycheirus angustatus

2 PBL 73, England 100%

PAL 50, Canada 100% 1 Pinus

Platycheirus scutatus

8 PAL 85, Norway 1 100% of 2 tricolp. types

PAL 86, Norway 100% of 3 tricolp. types

PBL 32, Sweden contained no pollen

Platycheirus albimanus

ES PAL 159, USSR 100%, mainly 2 types

PAL 62, USSR 10 13, 3 types

PAL 64, USSR 100% fenestrate

PAL 97, Norway + 100% tricolp.-scabrate

PAL 98, Norway ca. 50 fenestrates

PBL 36, Sweden 100% mainly Caryophyllac.

and tricolp. types

PBL 37, Sweden 100%

PBL 42, Sweden + various types + Typha latifolia

3 PAL 60, USSR mainly Ericaceae + various tricolp.

PAL 61, USSR 100%

PAL 63, USSR 100% various

PAL 65, USSR 3 24 tricolp.-psil., 15 indeterminate,

1 different

PAL 95, Norway mainly tricolp., fenestrate,

and tricolp.-psilate types

PAL 96, Norway various types

Table 2 (cont)

Slide no. and

country of

origin

PBL

39, Sweden

13, Netherl.

87, Norway

88, Norway

90, Norway

66, USSR

70, USSR

71, USSR

89, Norway

67, USSR

68, USSR

69, USSR

91, Norway

92, Norway

93, Norway

94, Norway

42, Finland

43, Finland

9, England

58, Sweden

44, Finland

48, Canada

49, North Am.

56, Sweden

57, Sweden

41, Finland

55, Sweden

12, Norway

13, Norway

14, Norway

16, Norway

9, Norway

10, Norway

11, Norway

15, Norway

17, Norway

18, Norway

19, Norway

20, Norway

pollen composition recorded

TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 2 (1982)

grasses Plantago Cyperaceae

lanceolata

type

Platycheirus fulviventris

93% 4%

Platycheirus immarginatus

93% 7%

Stra 100%

Platycheirus peltatus

71%

37%

Platycheirus scambus

100%

100%)

100%

(Platycheirus scambus, cont.)

100%

ar 78%

100%

28% 72%

1 17%

8% 22%

Pyrophaena granditarsa

100%

100%)

90%

100%

other ones

mainly Caryophyllac.,

also tricolp. types

3% prob. Caryophyllac.

12% Caryophyllac., 17% other types

100%

97% various

66%

50% Rumex-like, rest one type

100%

100% Liliaceae

mainly Rosaceae,

also Typha latifolia

mainly Rosaceae

100%

22% Rumex sp., Secale +

Pteridophyta: 1

83% Rumex sp., Tilia +, 1 Pinus

70% Pteridophyta, various +

10% trilete

+ Chenop., + different

+ tricolp.-scabrate

100%, mainly one type

100%)

100%, various types

100%, in two types

100%, in three types

100%, mainly in 2 types

100% Rosaceae

100% in two types

LEEREVELD: Anthecological relations 3 33

yield quantities of recognisable pollen. That

weather conditions were adverse when the flies

were caught is hardly likely because as a rule

entomologists do not go out on field trips to

catch sun-loving insects when the conditions are

unfavourable.

The table does not only show confirmations

but also deviations from the general consump-

tion pattern of a certain syrphid species. It also

gives some indication of the local habitat at a

certain date whenever diverse species are avail-

able from the same site and date. The recon-

struction of the biotope from the feeding habits

of a number of sympatrically occurring hover

flies may eventually provide us with a useful re-

cord of the consumptive behaviour of a certain

syrphid species of the M-P group in that bi-

otope and enables the comparison of that specif-

ic behaviour with the general tendencies. Such a

systematic study has not been carried out as yet

but seems highly promising.

DISCUSSION AND CONCLUSIONS

The usefulness of a pollen vector is to a large

extent dependent on the faithfulness of this vec-

tor. Generally speaking the efficiency, i.e., the

chance of an efficient transfer of pollen, in-

creases as the faithfulness is greater. The con-

sumption of pollen of anemophiles or apparent

anemophiles by syrphid flies is only benificial to

a plant species if a previous visit was to a flower

or inflorescence of a specimen of the same spe-

cies. This is a salient point in the present investi-

gation because the relatively high degree of

faithfulness of the syrphid flies of the M-P

group, as established by direct observations and

resulting in effective pollen transfers (Stelleman

& Meeuse, 1976), is assessable by the composi-

tion of the consumed quantities of pollen. The

presence of practically only one kind of pollen

| suggests a strongly monolectic feeding behav-

iour, and even if fair amounts of only two or

three kinds of pollen is consumed, i.e., only a

few plant taxa are selectively being visited, the

| efficiency of the flies as pollinators is not greatly

| reduced. However, since the groups distin-

| guished may contain several species it would be

an exaggeration, not to say misleading, if the in-

sect is called faithful to a single taxon because it

| is faithful to a type of plant and not necessarily

to only one species. It is quite clear that many

| individuals or even species are faithful to a cer-

tain type. Observations in the field of the be-

haviour of a single fly suggest that in many situ-

| ations the animal visits only Plantago, or only

the inflorescences of a single grass species (e.g.,

one of the few or the only one in anthesis at the

time of feeding), or only a single genus of the

Cyperaceae (such as Rhynchospora), or nothing

but Typha. Knowledge of the conditions ob-

taining at the locality (time of the year, floral

composition at the site, time of the day, avail-

ability and relative frequency of the potential

sources of pollen food in the habitat, etc.) may

permit the conclusion of a high degree of faith-

fulness, but this is information usually not avail-

able when flies collected without extensive field

notes are being studied. Bearing this in mind,

one nevertheless arrives at a tentative evaluation

of the results of the analyses as shown in table 2.

It seems as if the behaviour is inconsistent

and the table rather meaningless, but a perusal

of table 2 suggests that ambiguous evaluations

may be explained by the local habitat. Melanos-

toma mellinum and other species studied in an

environment teeming with Plantago, grasses,

etc. seem to have feeding preferences, but it

stands to reason that in certain vegetation types

or in a certain season Plantago pollen is not

available. Unpublished records of visits of M-P

syrphids by P. Stelleman in heathland and

moorland areas in the Netherlands in the month

of August show that the only opulent sources of

pollen available, viz., Molinia coerulea and

sometimes Rhynchospora, are visited. In spring

Typha may be eaten by flies already active at

that time and living near the water containing

the stands of Typha because no other source

may be present at that site at that time. Waitz-

bauer (1976) recorded Platycheirus clypeatus,

P. fulviventris, and P. perpallidus as occurring

in large numbers on the male inflorescences of

Typha from the end of May till the middle of

June. Presumably other good sources of pollen

were not available at the site at the time (or the

producers had not come into flower yet).

Taking all this into account one may safely

conclude that at least a majority of the taxa of

the M-P group of syrphids occurring in Europe,

America, and New Zealand exhibits a feeding

behaviour similar to that observed in several

sites in Europe (especially in Holland) and are

faithful to a certain source of food, sometimes

Plantago, sometimes grasses, sometimes Cype-

raceae (or combinations of these). The flies are

obviously capable of discriminating between the

plants growing in their habitat and of selecting

their pollen sources. An earlier study of gut

contents of syrphids of the M-P group including

Pyrophaena by Van der Goot & Grabandt

34 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 2 (1982)

(1970) had already shown the feeding prefer-

ences of these flies. The present study confirms

the deviating consumption pattern of certain

species, and suggests that Platycheirus albima-

nus and especially P. scutatus are exceptional.

P. scutatus often feeds on nectar alone (personal

observations by our Amsterdam research group

in Hessia, Germany) and also differs in its be-

haviour in that it is actively feeding till late af-

ternoon whereas most members of the M-P

group are only active from dawn to 10.00 hrs or

11.00 hrs in the morning.

Pollen of grasses and/or Plantago was found

in specimens from Europe, N. America and N.

Zealand. Pollen of Cyperaceae was only present

in appreciable quantities in specimens from

Scandinavia, the adjoining part of the USSR,

and Canada. This is no doubt attributable to the

locally great abundance of cyperaceous forms in

open arctic and subarctic vegetation types

(heathlands, tundra, etc.).

In other parts of the world more or less local

stands of Cyperaceae may attract representa-

tives of the M-P group at least when other pol-

len sources are not abundant. Carex and Rbyn-

chospora are visited in, e.g., the Netherlands in

special habitats (moorland, fens, etc.).

When this paper had been submitted for pub-

lication, a large number of syrphids of the M-P

group (about 180) became available for dissec-

tion. The results confirm the general picture,

viz. that all over the world the diet of such flies

consists very often of pollen of nominal anemo-

philes (Cyperaceae, Gramineae, Plantago lan-

ceolata, P. maritima, P. media), even in such re-

mote areas as Madagascar (Melanostoma annu-

lipes: 100% Cyperaceae pollen, almost certainly

of a Cyperus spec. of the C. papyrus group).

Since an effective transfer of pollen of Planta-

go by M-P flies has been demonstrated (Stelle-

man & Meeuse, 1976) the presence of apprecia-

ble quantities of Plantago pollen in the digestive

tract of such syrphids may be taken as clearly

indicative of an effective pollination with the

flies as the animal vector. SEM photomicro-

graphs (Stelleman, 1978) have shown that grass

pollen may also become attached to the body of

M-P syrphids and it is tentatively accepted by

the present author that the consumption of

grass pollen is indicative of a possible effective

pollination of some grasses by these flies. The

same probably holds for pollen of Cyperaceae,

although so far pollen grains of Cyperaceae

have not been found adhering to insect bodies,

but this is under investigation.

By assuming that the presence of appreciable |

quantities of ingested pollen in the digestive |

tract of a species of syrphid is sufficient proof of |

a possible effective pollen transfer by the species |

in question, the final possible, tentative conclu- |

sion can be drawn that plant forms with a more |

or less manifest anemophilous pollination syn- |

drome are not infrequently pollinated by such |

flies. This plant-insect interaction is found in |

many parts of the world, so that most species of |

syrphid flies belonging to the M-P group are ap- |

parently specialised feeders. Several questions |

remain unanswered, such as the possible role of |

“Pollenkitt” in the adhesion of pollen grains to |

the insect body, and the relative importance of a

possible anemophilous pollen transfer in respect

of the effective entomophilous pollination. The

study of the topic is continued by our research

group in various directions.

ÄCKNOWLEDGEMENTS

The author is much indebted to the Dept. of

Palynology, H. de Vries-Laboratorium, espe-

cially to A. Smit (late of that Dept.), to Dr. B.

van Geel and to Mr E. Beglinger for their help

with the identifications (S. and v.G.) and techni-

cal assistance (Mr B.).

Without the contribution of material from

many parts of the world this study would not

have been possible. The consignments of speci-

mens and information received from the follow-

ing persons are gratefully acknowledged: V.

Brädeszu (Bucarest, Roumania), S. Gaunitz

(Växjö, Sweden), V.S. van der Goot and Mr R.

A. Grabandt (Amsterdam), Dr. B. A. Holloway

(Auckland, New Zealand), B. J. D. Meeuse

(Seattle, U.S.A), Dr. T. R. Nielsen (Sandnes,

Norway), Dr. G. V. Price (Ascot, England), and

Dr. V. Richter (Leningrad, U.S.S.R.).

The encouragement from and critical dis-

cussions with Prof. A. D. J. Meeuse and Dr. P.

Stelleman is much appreciated. Prof. Meeuse al-

so took care of the vetting and the English

translation of the original draught.

REFERENCES

Erdtman, G., B. Berglund & J. Praglowski, 1961. An

introduction to a Scandinavian pollen flora. —

Stockholm.

Goot, V. S. van der, & R. A. Grabandt, 1970. Some

species of the genera Melanostoma, Platycheirus

and Pyrophaena (Diptera, Syrphidae) and their re-

lation to flowers. — Entom. Ber. 30: 135—143.

Holloway, B. A., 1976. Pollen-feeding in hover-flies

(Diptera: Syrphidae). — N. Zeal. J. Zool. 3: 339—

350.

LEEREVELD: Anthecological relations 3 35

Leereveld, H., A. D. J. Meeuse & P. Stelleman, 1976.

Anthecological relations between reputedly ane-

mophilous flowers and syrphid flies. II. Plantago

media L. — Acta Bot. Neerl. 25: 205—211.

Mesler, M. R., 1977. Notes on the floral biology of

Plantago in Michigan. — Michigan Botanist 16:

73-83.

Nielsen, T. R., 1971. Syrphidae (Dipt.) from Jaeren,

Norway, I. With description of two new species.

— Norsk ent. Tidsskr. 18: 53—73.

Stelleman, P., & A. D. J. Meeuse, 1976. Anthecologi-

cal relations between reputedly anemophilous

flowers and syrphid flies. I. The possible role of

syrphid flies as pollinators of Plantago. —

Tijdschr. Entom. 119 (2): 15—34.

Stelleman, P., 1978. Applications of scanning electron

microscopy in anthecology. — Acta Bot. Neerl.

27: 333—340.

Waitzbauer, W., 1976. Die Insektenfauna männlicher

Blütenstände von Typha angustifolia. — Zool.

Anz., Jena, 196: 9—15.

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J. BELLE. — A review of the genus Archaeogomphus Williamson (Odonata, Gom-

phidae), p. 37—56, figs. 1—31.

Tijdschrift voor Entomologie, deel 125, afl. 3 Gepubliceerd 15-VII-1982

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A REVIEW OF THE GENUS ARCHAEOGOMPHUS

WILLIAMSON (ODONATA, GOMPHIDAE)

JEAN BELLE

Onder de Beumkes 35, Velp, The Netherlands

ABSTRACT

The genus Archaeogomphus Williamson is reviewed. Both sexes of A. densus sp. n. (male

holotype: Santa Catarina, Brazil) and the hitherto unknown male of A. infans (Ris) are de-

scribed and illustrated. The female from Misiones, Argentina referred to A. infans by Ris is

conspecific with the new species A. densus. New distributional records of Archaeogomphus

species are given and a key to the five known species is constructed. Generic characters,

structures of specific value, mutual mating adaptations, immature stages, habitats, behav-

iour in the field, and distribution are discussed. The manner of grasping of the female by the

male during copulation is indicated. A historical survey of the genus is provided as well as a

full bibliography on the subject.

CONTENTS

page

inroducronee ti td a äh: 37

ENcknovzledsements, ... odes coe Ae sno 38

Sliecomcel SUR ee à - RN 38

ihelsenus Archaeogomphus-.................. 39

The species of Archaeogomphus .............. 40

Efay LO MES pete st ATEN NIEREN, 42

AVIS ORIG) EC RES A ARSA RR Te 42

PA hamatus (Williamson). 3.11. 20. asten 44

grata Williamson) 2.2.2: id die. 44

À, ADS NES ME ER RAR EE 45

FAMMICHSUSISPEC TON, foe net ne sales 45

Aimoteonitheimmaturestagesı i. ii... att. 47

Habitat and behaviour inthe field ............ 47

Georraphieidistribution RS E n 48

Beierenees ep eee toa ae ini TI BOI a, 49

“I named a Gomphine genus Archaeogomphus and

Tillyard took one look and said it was the most highly

specialized Gomphine he ever saw. Had it turned up

as a fossil what would it have been? God only knows

— but I think it would have been whatever they

needed to fit in with something else. At the present

time when systematists are nearly extinct, I think

some good generic monographs dealing with what is

observable would be a fine thing.”

E. B. Williamson in a letter to J. G. Needham

Dated February 18, 1928 (Entomological News, 62)

INTRODUCTION

The incentive to start this study was the re-

ceipt of a new species of Archaeogomphus Wil-

liamson, 1919, from the collection of Mr. Carl

Cook (Center, USA) in December, 1977 and

the discovery of a (hitherto undescribed) male

of A. infans (Ris, 1913) in the collection of René

Martin, Paris, in the same month. Later on,

specimens of the new species were received

from Prof. Dr. Gerhard Jurzitza (Karlsruhe,

BRD), Prof. Dr. Angelo B. M. Machado (Belo

Horizonte, Brazil), and Prof. Dr. Minter J.

Westfall, Jr. (Gainesville, USA). Besides, the

Argentine specimen of Archaeogomphus re-

ferred to A. infans by Ris (1913) proved to be

identical with the new species as well.

Earlier I had obtained specimens of A. hama-

tus (Williamson, 1918) and A. nanus Needham,

1944, from the Rijksmuseum van Natuurlijke

Historie at Leyden (the Netherlands), speci-

mens of A.furcatus Williamson, 1923, and

A. nanus from the Museo de Biologia at Caracas

(Venezuela), and a specimen of A. hamatus col-

lected in Surinam by my eldest son. Paratypes

of A. hamatus in the Smithsonian Institution at

Washington, D.C. (USA), one of the Kartabo

males of A. hamatus in the Academy of Natural

Sciences of Philadelphia (USA), and some of the

paratypes of A. hamatus and A. furcatus as well

as a specimen of A. nanus in the Museum of Zo-

ology at Ann Arbor (USA) were available for

study. Additionally the material of A. nanus

previously assembled in Surinam (see Belle,

1970, 1972, and 1977) was re-examined. Finally,

Mr. and Mrs. George H. Beatty (Lemont,

USA), Dr. Oliver S. Flint, Jr. (Washington,

D.C., USA) and Dr. Dennis R. Paulson (Seattle,

38 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

USA) provided me with material of A. furcatus

collected during their explorations in Central-

America.

The present opportunity is used to review the

genus Archaeogomphus and publish a key to the

five known species. |

All figures of structural details illustrating

this paper are reproductions of original camera

lucida drawings (the details were completed by

free hand), except for fig. 21, which is a repro-

duction of an illustration published by William- _

son (1923), while the figures repesenting thorac-

ic colour patterns are diagrammatic, all drawn

over the same outline.

ÄCKNOWLEDGEMENTS

The following lists the names of the institu-

tions and individuals from whom material was

received on loan or as a gift; the names are pre-

ceded by the abbreviations used in the text. I

wish to express my gratitude to all those who

have assisted me whole-heartedly.

AC — Author’s collection.

ANSP — Academy of Natural Sciences of

Philadelphia; Dr. Daniel Otte.

GE — Collection Cook, Center; Mr.

Carl Cook.

@] — Collection Jurzitza, Karlsruhe;

Prof. Dr. Gerhard Jurzitza.

CM — Collection Machado, Belo Hori-

zonte; Prof. Dr. Angelo B. M.

\ Machado.

(CIP — Collection Paulson, Seattle; Dr.

Dennis R. Paulson.

CUI — Cornell University, Ithaca; Prof.

Dr. L. L. Pechuman.

FEM — Frost Entomological Museum,

Lemont; Mr. and Mrs. George H.

Beatty.

FSC — Florida State Collection of Ar-

thropods, Gainesville; Prof. Dr.

Minter J. Westfall, Jr.

MBC — Museo de Biologia, Caracas; Dr.

Janis Racenis (+).

ML — Rijksmuseum van Natuurlijke

Histoire, Leyden; Dr. P. H. van

Doesburg.

MNP — Muséum National d'Histoire Na-

turelle, Paris; Dr. Jean Legrand.

MNHW — National Museum of Natural His-

tory, Washington, D.C.; Dr. Oli-

ver S. Flint, Jr.

MZM — Museum of Zoology, Ann Arbor;

Dr. Irving J. Cantrall.

SMF — Natur-Museum und Forschungs-

Institut Senckenberg, Frankfurt

am Main; Dr. Heinz Schröder.

— Zoologisches Staatsinstitut und

Zoologisches Museum, Hamburg;

Prof. Dr. H. Strümpel.

ZMH

HISTORICAL SURVEY

1903. Needham published a figure of the right

pair of wings of a shrivelled teneral male

of Archaeogomphus from Sao Paulo, Bra-

zil, under Agriogomphus sp.

1909. Ris published a figure of the (transposed)

right pair of wings of a female of Archae-

ogomphus from Espirito Santo, Brazil,

under Agriogomphus sp.

1913. Ris described Archaeogomphus infans

under the generic name Agriogomphus on

the basis of two females. The holotype is

the Brazilian female of which he had

published a figure of the right pair of

wings in 1909. The description was illus-

trated with a diagram of the colour de-

sign of the female holotype. The other fe-

male, referred to the same species, is

from Misiones, Argentina.

1918. Williamson gave a description of Archae-

ogomphus hamatus under the generic

name Agriogomphus on the basis of 13

males and 9 females from Colombia. Im-

portant details of generic and specific val-

ue were elucidated by figures, and photo-

graphs of the left pair of wings of the

male and the female were published. In

the same paper he included a description —

with an illustration of a Colombian fe:

male of Archaeogomphus under Agrio-

gomphus species but refrained from nam-

ing it since the corresponding male was

unknown.

1919. Williamson created the new genus Ar-

chaeogomphus with Agriogomphus ham-

atus as the type-species to contain his

two Agriogomphus species of 1918 and

those of Needham (1903) and Ris (1913).

He placed Archaeogomphus in the group

of genera of the legion Gomphus of Selys

having numerous intermedian cross-veins

in the wings.

Williamson placed Archaeogomphus in

the Epigomphus series of his classification

of the family Gomphidae.

1923. Williamson described and illustrated Ar-

chaeogomphus furcatus on the basis of a

male and a female from Venezuela, and

the female from Colombia which he had

1920.

BELLE: The genus Archaeogomphus 39

described under Agriogomphus species in

1918.

Needham described and illustrated the

larva of an Archaeogomphus species on

the basis of a single male exuviae from

Santa Catarina, Brazil. In the same paper

he also gave 13 features of Archaeogom-

phus in his Verification Table for the

Neotropical Gomphidae.

Fraser compared the penile organ of Ar-

chaeogomphus hamatus with that of the

gomphids pertaining to other genera.

Tillyard and Fraser placed Archaeogom-

phus in the subfamily Gomphinae.

Needham described and illustrated Ar-

chaeogomphus nanus on the basis of two

males and a female from Surinam. He re-

ferred the male of his supposed Agrio-

gomphus sp. of 1903 to Archaeogomphus

hamatus.

Calvert recorded Archaeogomphus ham-

atus from Guyana.

Belle described and illustrated the larva

of Archaeogomphus nanus on the basis of

two exuviae, one of which belonging to a

reared male, and also published photo-

graphs of the exuviae and of the right pair

of wings of the male and female of Ar-

chaeogomphus nanus.

1979. Belle placed Archaeogomphus in the sub-

family Epigomphinae.

1940.

1944.

1948.

1970.

THE GENUS ARCHAEOGOMPHUS

Archaeogomphus Williamson, 1919a: 2—5; 1920: 7—

8; 1923: 1, 5—8. Gaige, 1934: 16—17. Tillyard &

Fraser, 1938—1940: 372. Fraser, 1940: 547.

Needham, 1940: 382, 386—389. Fraser, 1943:

161. Klots, 1944: 3. Needham, 1944: 172—173,

175, 179—181, 183. Calvert, 1948: 68. Anony-

mous, 1951: 305. Schmidt, 1951: 130. Fraser,

1957: 93. Belle, 1970: 1, 5. St. Quentin, 1967: 327;

1973: 355, 359. Paulson, 1977: 178. Belle, 1979:

45.

The type-species of Archaeogomphus is Ag-

riogomphus hamatus Williamson, 1918, as ex-

plicitly stated by Williamson (1919a) when he

established the genus.

Characters of the genus

An amended definition of the genus Archaeo-

gomphus was given by Williamson (1923) after

the description of A. furcatus.

In the venation of the wings Archaeogomphus

is at once recognizable from the other Neotrop-

ical gomphid genera having but a single row of

cells beyond the triangle in all wings, by the

much more open character and the lack of a bas-

al subcostal cross-vein. Other striking charac-

ters in the wings of Archaeogomphus are: Sub-

triangles, supratriangles and triangles open; two

or more intermedian cross-veins in hind wings;

middle fork oblique forward; vein M2 arising

about half a cell-width or more beyond subno-

dus in fore wings, generally closer to subnodus

in the hind wings; anal area one cell wide in fore

wings, two cells wide in hind wings; second pri-

mary antenodal cross-vein normally the fifth

antenodal cross-vein; one cubito-anal cross-

vein in addition to inner side of subtriangle; no

anal triangle in hind wings of male.

The venational features used by St. Quentin

(1973) for the recognition of the genus Archaeo-

gomphus seem to be partly derived from the fig-

ures and photographs of the wings published by

Needham (1903), Ris (1909), Williamson

(1918b) and Belle (1970). All these illustrations

show the sectors of the arculus connected at

their origin but in fact they are separated at their

origin by a distance that is equal to or greater

than the thickness of either sector.

Needham (1903), after studying the wings of

his male of Archaeogomphus, said: “We may re-

mark, in passing, that the Gomphinae [= Gom-

phidae], having attained a fair measure of ad-

vantageous vein arrangement, and a fair (or, in

the strongest species, even a superior) degree of

vein differentiation, seem to have turned to a

specialization of another sort. The imagoes are

on the wing less constantly than other Odonata

and fly for a shorter period. During this time

their chief concern is with reproduction, hence

we find the external genitalia highly speciali

zed”. The male of Archaeogomphus is unique in

the form of the tenth abdominal segment. Wil-

liamson (1919a) stated it as follows: “This has

the dorsum armed basally on either side with a

strong interno-posteriorly directed hook and

produced apically in a long snout-like projec-

tion which is more than twice as long as the ru-

dimentary, flap-like, superior appendages; . . .”.

As to how the dorsal hooks may function in

copulation, opinions were given by Williamson

(1918b) and Walker (in Williamson, 1918b).

Curiously enough, Needham (1944), despite his

statement of 1903 cited above, considered the

dorsal hooks functionless or merely decorative.

The knowledge of the males of A. infans and

A. densus spec. nov. and the discovery of a pair

of submedian spines at the superior margin of

40 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

the occipital foramen of the corresponding te-

males enable me to answer the question of the

manner of the grasping of the female by the

male during copulation. The males of A. infans

and A. densus spec. nov. exhibit an inferior

groove at either lateral side of the base of the

snout-like projection of the tenth abdominal

segment, an adaptation that is in perfect confor-

mity with the following method of grasping

during copulation. In seizing the female the

apex of the snout-like projection is pushed

against the superior margin of the occipital for-

amen; the denticles along the inferior margin of

the apex of the snout-like projection prevent a

slip backward. The spines at the superior mar-

gin of the occipital foramen rest in the inferior

grooves at the base of the snout-like projection.

The superior caudal appendages are pressed

against the rear of the head. The dorsum of the

tenth abdominal segment is placed on the pro-

thorax of the female with the dorsal hooks

snapped behind the posterior margin of the pro-

thorax.

The male of A. infans possesses minute denti-

cles at the inner side of the tip of the dorsal

hooks in order to prevent slip on the posterior

margin of the prothorax, and the rear of the

head of the paratype female of A. furcatus ex-

hibits two distinct mating marks against which

the apices of the male superior appendages may

be applied during pairing. Both characters are in

agreement with the copulatory position of the

male tenth abdominal segment and superior

caudal appendages.

The postero-lateral projections (horns) on the

occiput of the female of some species are not of

essential value in this method of grasping, but

serve only as lateral guards to give the male

tenth abdominal segment additional support.

Mr. Kiyoski Inoue, Osaka, who studied a

male of A. nanus, wrote in a letter (dated

6.x11.1975) to me: “I think these hooks fit the

structures on female prothorax in tandem, if it is

true, a transition phenomenon from Zygoptera

holding prothorax to Anisoptera holding head is

observed here”.

When Williamson (1919a) established the ge-

nus Archaeogomphus he only knew the male of

one species and the female of two species of this

genus. After describing A. furcatus in 1923 he

gave an amended definition of the genus but ne-

glected the generically important penile organ.

All species of Archaeogomphus possess a penis

of the same complicated structure that is typical

of the genus and that emphasizes the isolated

position of Archaeogomphus. A detailed de-

scription and a figure of the penile organ were

published by Williamson (1919a), Fraser (1940),

and Needham (1944).

The femoral hair and armature are typical of

the genus. The second and third femora have a

conspicuous row of arranged bristles along the

outer and inner sides, and between these rows,

on the ventral side, are scattered small spines,

which are twice to three times longer in the fe-

males than in the males. The first femora have

an arrangement of bristles only along the inner

side, while in both sexes the spines are equal in

length.

Also the shape of the vulvar lamina is typical

of Archaeogomphus. The vulvar lamina consists

of a broad short base and two long, slender,

hardly diverging and acutely pointed branches.

The colour pattern of the pterothorax is very

similar in the two sexes of each species but the

pale colours are clearer in the male than in the

corresponding female.

The wings are often slightly amber-coloured

on the basal half but for the rest they are clear.

THE SPECIES OF ARCHAEOGOMPHUS

General remarks

With the description of A. densus spec. nov.

in the present paper the total number of species

belonging to the genus is brought up to five.

These species, together with their type locality,

sex of the holotype and type location are chro-

nologically listed in table 1.

Distinct differences of specific value exist in

the venation of the wings, the position and rela-

tive length of the pterostigma, the colour design

of the body, the hood of the penial peduncle,

the vulvar lamina, the occiput of the female, and

the conformation of the superior margin of the

occipital foramen of the female.

Table 2 shows the features of the wings of the

diverse species listed in order of the date of de-

scription. The reticulation of the wings of

A. furcatus exhibits the slightest density, that of

A. densus the greatest (hence the specific name

densus). Although A. nanus is the smallest rep-

resentative of the genus its wings are more

densely veined than those of A. hamatus and

A. furcatus. The number of antenodal cross-

veins in the wings of the male of A. infans is re-

markably low, possibly due to individual varia-

tion (only one male available).

In all species, except in A. densus, the trian-

gles are normally four-sided. In A. densus, the

BELLE: The genus Archaeogomphus

Table 1. Chronologic list of the species ot Archaeogomphus, with type locality, sex of the holotype, and type

location

Species Type locality Holotype Type location

infans (Ris, 1913) Brazil (Espirito Santo) Q ZMH

hamatus (Williamson, 1918) Colombia (Fundación) d MZM

furcatus Williamson, 1923 Venezuela (Bejuma) d MZM

nanus Needham, 1944 Surinam (Litani River) 6) GEN

densus spec. nov. Brazil (Nova Teutonia) 3 MZM

Table 2. Features in the wings of Archaeogomphus species

Species 1 2 3 + 5 6 7 8 9 10

Cell-r Anten. Anten. Postn. Postn. Cellsa. Cellsa. Cellsp. R.length Length

MiM2 ınfw inhw infw inhw Culfw Culhw Cu2hw n-ptfw of hw

infans 3 2 10 8 5—7 5—6 1—2 2—3 3—4 3 19.5

hamatus 3 2 10—11 8—9 +46 +—6 1—2 1—2 0—2 3 16—17.5

furcatus 3 2 10—12 9 4—6 5—6 1 1—2 0—3 3 18—18.5

nanus È 2 10—12 9 5—7 4—6 1—2 1—3 1—2 3 13.7—16

densusd 3 jee ert remo id Te, RENE RER Dod as 17-19

infans 2 2 11 9 7 7 3 25,06 3 21.5

hamatus © 2 11 z 5—7 6—7 1—2 2—3 1—3 3—3.5 18—20

furcatus 2 2—3 10-12 9—10 5-8 5—8 1—2 1—5 2—6 2.7—3.3 19—21.5

nanus 2 2 10-12 9 7—8 6—8 1—2 2—4 3—5 3 16—18

densus 2 3 11-13 911 6-7 6-8 6-9 49 6—7 2.3—2.5 20-21

For ease of reference the abbreviated multiple column-headings are to be taken as follows:

. Antenodal cross-veins of first series in fore wing.

. Antenodal cross-veins of first series in hind wing.

. Postnodal cross-veins in fore wing.

. Postnodal cross-veins in hind wing.

COND UI R WN mm

wing.

. Cell-rows near marginal row of cells, between M1 and M2.

. Cells on anterior side of Cul which do not reach M4, in fore wing.

. Cells on anterior side of Cul which do not reach M4, in hind wing.

. Cells posterior to Cu2 and distal to postanal cells which do not reach the posterior wing margin, in hind

9. Relative length of distance between nodus and pterostigma in fore wing, the costal edge of the pterostigma

being taken as the standard for comparison.

10. Length of hind wing in millimeters.

All numbers in the table are approximate and sometimes derived from a single specimen.

triangle is always three-sided in the hind wings

and sometimes four-sided in the fore wings.

The lateral dilatations of the eighth abdomi-

nal segment of the male of A. infans and A. den-

sus are well-developed and widen broadly back-

wards, those of A. hamatus, A. furcatus and

A. nanus are very slightly developed and a little

prolonged in posterior direction.

The posterior margin of the tenth abdominal

segment of the male of A. infans and A. densus

is denticulated at the level of the base of the su-

perior caudal appendages and along the ventral

side of the segment, that of A. hamatus, A. fur-

catus and A. nanus is entirely denticulated (ex-

cluding the snout-like projection).

In A. infans and A. densus, the superior mar-

gin of the occipital foramen of the female bears

a pair of submedian spines, and in correlation

with these spines the snout-like projection of

the tenth abdominal segment of the correspond-

ing males has a pair of inferior grooves in which

the spines fit during copulation. The superior

margin of the occipital foramen of the female ot

A. hamatus has a pair of submedian lobes,

whereas that of the female of A. furcatus and

A.nanus has no prominences. The snout-like

projection of the tenth abdominal segment ot

the male of these three species exhibits no ma-

ting adaptation in the shape of a pair of grooves.

The rear of the occiput of the female of

42 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

A. furcatus has a pair of long, posteriorly di-

rected horns, that of the female of A. hamatus a

small prominence at either lateral end, whereas

in the females of other species it is smooth, lack-

ing any projecting angles whatever. The rear of

the occiput of the male of A. furcatus has a small

knob at either lateral end, that of the male of

other species has no prominences of any kind.

Classification

The species of the genus Archaeogomphus can

be conveniently divided into two well-defined

groups, VIZ.:

(1) The infans group, comprising the two

species A. infans and A. densus spec. nov. The

species of this group have the five veins co-

alesced at the posterior angle of the triangle in

the hind wings. The lateral dilatations of the

eighth abdominal segment of the males are well-

developed and widen broadly in posterior direc-

tion. The posterior margin of the tenth abdomi-

nal segment of the males is denticulated at the

level of the base of the superior caudal append-

ages and along the ventral side of the segment.

The females have a pair of submedian spines at

the superior margin of the occipital foramen.

(2) The hamatus group, comprising the three

species A. hamatus, A. furcatus and A. nanus.

The species of this group have the five veins

normally not coalesced at the posterior angle of

the triangle in the hind wings but separated into

two groups, three anterior and two posterior.

The lateral dilatations of the eighth abdominal

segment of the males are narrow. The posterior

margin of the tenth abdominal segment of the

males is entirely denticulated (excluding the

snout-like projection). The superior margin of

the occipital foramen of the females lacks

spines.

KEY TO THE SPECIES OF Archaeogomphus

. Males

— Females

2. Posterior margin of base of hind wings

hardivanenlaten er ern nanus

— Posterior margin of base of hind wings

Steonelyaaneulaten ee 3

3. Lateral dilatations of eighth abdominal seg-

ment well-developed, strongly widened

posterad and extending to nearly half the

lengthiof segment ee 4

— Lateral dilatations of eighth abdominal seg-

ment Narrow; Sne NE 5

4. Wings with three rows of cells near margin-

—

al row, between M1 and M2...... densus

— Wings with two rows of cells near marginal

row, between M1 and M2........ infans

5. Ventral margin of penial peduncle, in poste-

rior view, broad, concave and with a shal-

low median notch hamatus

— Ventral margin of penial peduncle, in poste-

rior view, deeply excised V-shaped ........

furcatus

6. Rear of occiput with a pair of long, posteri-

orly directed spines (see fig. 8).... furcatus

— Rear of occiput with small prominences or

without any prominences.............. 7

7. Rear of occiput with a small prominence at

either lateral end (see fig. 7) .... hamatus

— Rear of occiput smooth, lacking any pro-

jecting angles whatever eenen

8. Wings with three rows of cells near margin-

al row, between M1 and M2...... densus

— Wings with two rows of cells near marginal

row, between M1 and M2.............. 9

9. Superior margin of occipital foramen with a

pair of submedian spines ........ infans

— Superior margin of occipital foramen with-

out spines nanus

Treatment of the species

The species are treated chronologically. For

each species are given a list of synonyms, the

material studied (in addition to those mentioned

in my former papers of 1970, 1972 and 1977),

and a description or descriptive notes.

Archaeogomphus infans (Ris, 1913)

(figs. 3, 10, 15, 19, 2326, 31)

Agriogomphus spec., Needham, 1903: 738, fig. 27.

Ris, 1909: 11, fig. 1. Agriogomphus infans Ris,

1913, 72—73, fig. 11 (©). Williamson, 1918b:

15—17; 1919a: 5; 1923: 1.

Archaeogomphus infans; Williamson, 1923: 7—8. Fra-

ser, 1943: 161. Needham, 1944: 176. St. Quentin,

1973: 359. Paulson, 1977: 180.

Material. — Brazil: Espirito Santo, 20.v.1898,

1 2 (holotype), J. Michaelis leg. (ZMH); Espiri-

to Santo, 1 d (MNP).

This species was described in a paper dealing

with Argentine Odonata. As explicitly stated by

Ris (1913) the type is the female from Espirito

Santo. However, the locality data of the type as

given by Ris prove to be incomplete. The label

attached to the pin of the type indicates that the

female has been taken in Espirito Santo, Brazil,

on 20.v.1898 by J. Michaelis. Since the female

BELLE: The genus Archaeogomphus 43

from Misiones, Argentina, referred to the same

species by Ris, is conspecific with my new spe-

cies A. densus described hereafter, and since the

distance from Espirito Santo to Argentina is

about 1000 miles, the occurrence of A. infans in

Argentina is subject to doubt.

The female holotype of A. infans has a sub-

quadrangular depression in the swelling behind

each compound eye. The distance between the

two depressions, however, is much too great for

a possible correlation with the grasping organs

of the corresponding male. In my opinion the

depressions are injuries owing to the fact that

the head was turned one-fourth around with its

rear surface to the side when the specimen was

originally stored in an envelope (the pterotho-

rax of the type is partly and the abdomen

largely flattened).

The colour pattern of the present male resem-

bles greatly that of the female holotype, but the

dark interpleural and metapleural stripes are not

connected with each other. These stripes are

largely confluent above the spiracle in the holo-

type female. The wings of the male are some-

what less densely veined than those of the fe-

male holotype, a character likewise found in the

two sexes of other congeners. The number of

antenodal cross-veins in the present male is only

10 in each fore wing and only 8 in each hind

wing; in Archaeogomphus normally 11 in the

fore wing and normally 9 in the hind wing.

The male of Archaeogomphus of which

Needham (1903) published the right pair of

wings may be A. infans as already suggested by

Williamson (1923). The venation agrees in hav-

ing the five veins at the posterior angle of the

triangle practically coalesced in the hind wing,

and in having this wing with four cells posterior

to Cu2 which do not reach the posterior wing

margin (in the male of A. hamatus, at the most

two cells posterior to Cu2 which do not reach

the posterior margin, cf. table 2). Also the anal

margin of the hind wing is more resembling that

of A. infans than that of A. hamatus. It is not to

be forgotten that Needham’s figure of the wings

is a drawing made after boiled and unrolled

wings of a very teneral male (cf. Williamson,

1918b: 1, footnote 5; Needham, 1944: 175) and

that due to these conditions certain deviations in

the form of the wings and the position of the

veins are probable. Finally, the place of capture

(Sao Paulo, Brazil) is in conformity with the

type locality of A. infans (Espirito Santo, Bra-

zil). On the distribution map (fig. 31) the sym-

bol referring to the locality of the male from Säo

Paulo is marked by a question mark.

Description of male (hitherto undescribed;

pterothorax partly crushed). — Total length, 35

mm; abdomen, 27 mm; hind wing, 19.5 mm;

costal edge of pterostigma in fore wing, 2.0 mm.

Face pale grey-green, the frons somewhat

darker. Lateral sides of labrum and margin of

facial lobes pale yellow. Free border of labrum

fringed with brown-yellow hairs. Frons round-

ed on edge, its superior surface grey-green in

front, becoming darker at base. Vertex as in ho-

lotype female, dark brown, the transverse ridge

becoming bulbous behind each lateral ocellus.

Occiput dark brown, rounded on edge, without

crest line but with hairs. Swollen area of head

behind each compound eye dark brown, sparse-

ly covered with long brown hairs. Tempora yel-

low. Spines at superior margin of occipital for-

amen closer to each other and smaller than in fe-

male holotype. Labium and adjacent mouth

parts pale green.

Prothorax largely brown. Front margin of an-

terior lobe brownish yellow. Swollen rear mar-

gin of middle lobe green. Posterior lobe pale

yellow but brown in middle.

Pterothorax dark brown with pale green

markings; its colour pattern shaped as shown in

fig. 3.

Wings clear but subcostal and cubito-anal in-

terspaces very slightly brownish yellow tinged.

Venation of wings dark brown, including fron-

tal margin of costa. Pterostigma dark brown,

surmounting 2! cells. Intermedian cross-veins

45/3—3 in fore and hind wings, respectively.

Base of hind wings strongly angled. Other ve-

national features given in table 2.

Femora brown, the outer side with a yellow-

ish brown band along anterior row of spines.

Tibiae brown, the outer side yellow. First tibiae

without lamina tibialis. Tarsi and claws brown,

but second joint of tarsi yellow.

Abdomen predominantly yellow-brown on

segments 1 to 7, dark brown on segments 8 and

9, and brown on segment 10, including caudal

appendages. Segment 1 brown above. Segment 2

grey-green along dorsal anterior margin and on

auricles. Segments 3 to 6 becoming dark brown

backwards, being dark brown ringed, respecti-

vely on half, two-fifths, one-third, and one-

fourth the way along each segment. Segment 7

without such a ring. Lateral dilatations of seg-

ment 8 broadly widening in posterior direction

44 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

and extending to a point about half the way

along segment 9. Lateral dilatations of segment

9 much narrower than those of segment 8, equal

in width, more or less inflexed against sternum,

and prolonged backwards. Dorsal side of seg-

ment 10 convex except for dorsal side of snout-

like projection which is concave.

Archaeogomphus hamatus (Williamson, 1918)

Grass 7 Wil, 16, 20, Sil)

Agriogomphus hamatus Wiliamson, 1918b: 4-11, pl.

1 figs. 1-12, pl. 2 figs. 14-15 (d, 2). Gaige,

1934: 17. Belyshew & Haritonov, 1977: 85 (nam-

atus = lapsus calami pro hamatus).

Archaeogomphus hamatus; Williamson, 1919a: 1—2,

5, PIL lime, 28 19190 43 19203 73 1255 1-5, A

9, pl. 1 fig. 4. Calvert, 1935: 10. Fraser, 1940: 543,

pl. 3 fig. 11; 1943: 161-162. Needham, 1944:

175—177, 179. Calvert, 1948: 49, 67—68. St.

Quentin, 1973: 359.

Material. — Brazil: Goiás, Rio Paraim,

29.x.1977, 1 2, Keiko leg. (CM). — Colombia:

Dept. Magdalena, Fundación, 10.1.1917, 1 ó

(paratype) (ML); 14.1.1917, 1 d, 1 2 (para-

types) (MZM); 14—17.1.1917, 1 d, 1 2 (para-

types) (MNHW), all paratypes J. H. & E. B.

Williamson leg. — Guyana: Bartica District,

Kartabo, no date, 1 & (ANSP). — Surinam:

Marowijne District, Albina, vin.1973, 1 2, J. J.

Belle leg. (AC).

Williamson (1918b) gave a figure of the vul-

var lamina with the parallel slender divisons.

The females recorded here have the tips ot these

divisions divergent (fig. 11). The female from

Surinam is somewhat larger and stouter than the

other two females; its measurements are: total

length, 33 mm; abdomen, 25 mm; hind wing, 19

mm. Those of the other females are: total

length, 30 mm; abdomen, 22.5 mm; hind wing,

18 mm (paratype) and 18.5 mm (Rio Paraim).

The distance between the nodus and the ptero-

stigma in the fore wing of the female from Suri-

nam, however, is three and a half times the

length of the costal edge of the pterostigma; it is

only three times in the other females. The fe-

male from Rio Paraim is somewhat paler than

the other females and it has better developed

first pale antehumeral stripes.

The female of A. hamatus is readily recogniz-

able from that of the other congeners by the

conformation of the occiput the rear margin of

which possesses a small prominence at either

lateral side.

The tenth abdominal segment (including the

snout-like projection) of the male of A. hamatus

is concave on the dorsal side.

The numerals for the number of cross-veins |

and cells as stated by Williamson, 1918b: 9) are

not correct in all respects. Some of the misstate-

ments were already corrected by him (William-

son, 1923: 4). The photograph of the left pair of

wings of the male of A. hamatus (Williamson,

1918b: fig. 14) shows only 10 and 8 antenodal

cross-veins in the fore and hind wings, respecti-

vely. These numerals are 11 and 9, respectively,

in the original description. Further, in the wings

figured, there are only 4 postnodal cross-veins

in the fore wing as well as in the hind wing;

according to the original description 5 or 6.

Archaeogomphus furcatus Williamson, 1923

(figs. 1,4,8—9, 12, 21,31)

Agriogomphus sp. Williamson, 1918b: 17—18, pl. 1

fig. 13 (9).

Archaeogomphus furcatus Williamson, 1923: 2—9, pl.

1 figs. 1—3 (6). Gaige, 1934: 17. Needham, 1944:

176. Calvert, 1948: 68. St. Quentin, 1973: 359.

Material. — Colombia: Dept. Antioquia,

Cristalina, 14.11.1917, 1 2 (paratype), J. H. & E.

B. Williamson leg. (MZM). — Costa Rica:

Prov. Guanacaste, Rio Santa Rosa (3.7 mi E. of

Las Canas), 25.v11.1967, 1 2; Rio Piedras, N. of

Bagaces, 27.vi1.1967, 1 ©, both O. S. Flint, Jr. &

M. A. Oruz B. leg. (MNHW); Rio Tempisque

(11.3 mi W. of Liberia), 9.viii.1964, 1 reared d

+ exuviae, F. G. Thompson leg.; Hacienda Ta-

boga, 9.x11.1966, 1 ©, D. R. Paulson leg.; Río

Corobici (3.4 mi NW. of Las Canas), 12.11.1967,

2 larvae, M. L. Paulson leg.; Río Santa Rosa (3.5

mi N. of Las Canas, 300°), 26.vii.1967, 1 2, M.

L. Paulson leg.; Prov. Heredía, Río Puerto Vie-

jo (1.5 mi S. of Puerto Viejo), 14.1v.1967, 1

reared 2 + exuviae, 9 larvae, D. R. & M. L.

Paulson leg. (CP). — Mexico: Oaxaca, Rio

Choapan (16.6 km S. of Tuxtepec), 27.v.1962, 1

di, Ga HH & An E° Beatty leer (AO) 3 @hiapass

Rio Despoblado (8.7 mi NW. of Huixtla),

31.vii.1965, 1 reared d + exuviae, 1 reared 2 +

exuviae, 2 larvae; Rio El Mosquito (16.2 mi

Rio Cuil (1.92 mi NW of Einsam)

23.vi.1966, 1 larva, all D. R. Paulson leg. (CP).

— Venezuela: Tächira, La Fria, 1.iv.1958, 1 ©,

J. Racenis leg. (MBC).

The occiput of the female of A. furcatus is pe-

culiar by the two long, posteriorly directed

BELLE: The genus Archaeogomphus 45

spines. On the contrary, the occiput of the cor-

responding male has at the most a very small

knob at either lateral end.

The rear of the head of the female paratype

mentioned above exhibits two distinct postgenal

cicatrices on which apparently the tips of the

male superior caudal appendages were applied

during copulation. The other females here re-

corded (as well as the females of the other con-

geners) have no, or no distinct copulation

marks.

The single (reared) female from Mexico re-

sembles those from continental South America

but the females from Costa Rica are different in

some respects. Most striking is the development

of the occipital spines which in the females from

Costa Rica are much stouter and longer. The

wings of the females from Costa Rica have also

a denser reticulation. They have generally one

extra cell or two extra cells (near the marginal

row) for a third row, between M1 and M2. For

the Costa Rican females the numbers in the col-

umns of table 2 are 23, 11—12, 9—10, 7—8,

6-8, 1-2, 3—5, 3—6, 3.1—3.3, 20—21.5, re-

spectively; for the Mexican female Fraserand

the South American females these numbers are

2, 1011,79, 5—6, 5—6, 1, 1—2, 2—3, 2.7—3,

19—20, respectively. However, no difference of

any importance was found between the single

(reared) male from Costa Rica and the males

from Mexico and South America. The confor-

mation of the hind lobe of the accessory genita-

lia is very similar and the wings have no denser

reticulation. Also no differences were found be-

tween the larvae from Costa Rica and Mexico.

Dr. Alice F. Beatty informed me (letter dated

25.v1.1979) that the Mexican male from Tuxte-

pec is one of a series of 13 d and 1 © collected

on 27 and 29.v.1962 at the same locality. An-

other male was taken west of Cosamaloapan de

Carpio in Veracruz by Harold White. Many lar-

vae were also assembled from three localities in

the Pacific coastal region of Chiapas on 18—

23.11.1963, and some of these larvae were reared

(see immature stages). A detailed study of the

Mexican material of A. furcatus by Mr. and

Mrs. Beatty is in progress.

Archaeogomphus nanus Needham, 1944

(figs. 5, 13, 17, 22, 31)

Archaeogomphus nanus Needham, 1944: 171, 176—

180, pl. 14 figs. 1a-d (d, ®). Belle, 1970: 1, 5—7,

figs. 1—5, pl. 1a-b, 2b (larva); 1972: 217—218. St.

Quentin, 1973: 359. Belle, 1977: 290.

Material. — Surinam: Upper Litani River,

18.vii.1939, 1 d (FSC); Marowijne District, La-

wa River, Benzdorp; 18.1x.1960, 1 8, 1 2, J.

Belle leg. (ML). — Venezuela: Bolivar, Caicara,

3.1.1957, 1 2, J. Racenis leg. (MBC); Bolivar, El

Dorado-Brasil (km 67), 27.v11.1966, 1 9, Camp-

os leg. (AC). — Brazil: Mato Grosso, Rio Tapi-

rapé, Porto Velho, 23.1.1963, 1 2, Celia Pinhei-

ras leg. (ML); Território do Roraima, Surumu

(NW. of Depòsito, 4° 14’ N., 60° 55’ W.),

ix.1966, 1 6, Moacir Alvarenga leg. (MZM).

The male from the Upper Litani River be-

longed to the gomphid material sent to Need-

ham (1944) for description by Dr. D. C.

Geijskes. The cellophane envelope in which the

specimen has been stored contains a label “Ar-

chaeogomphus nanus Paratype Det. by J. G.

Needham” but the male has no official standing

since it has not been cited in the original de-

scription. The male has the five veins coalesced

at the posterior angle of the triangle in the hind

wings. A. nanus has these veins normally sepa-

rated into two groups, three anterior and two

posterior.

The Brazilian male from Surumu differs in

size and coloration with my males from Suri-

nam. It is a smaller specimen with the mea-

surements: total length, 27 mm; abdomen, 20.9

mm; hind wing, 13.7 mm. The males from Suri-

nam have the (average) measurements: total

length, 29 mm; abdomen, 22.5 mm; hind wing,

15.5 mm. The light colours of the pterothorax

are light grey; they are tawny yellow in the

males from Surinam.

The Venezuelan female from El Dorado-Bra-

sil has the basal half of the hind wings strikingly

brown-tinged. The trigonal interspace in the

hind wings of this female starts with an extra

initial cell at the posterior angle of the triangle.

The vulvar lamina of A. nanus has not been

figured before. Needham’s (1944) depiction of

the vulvar lamina (his fig. te, pl. 14) fits that of

Cyanogomphus demerarae Selys (his Ebegom-

phus strumens) whereas that of fig. 4d (pl. 14)

fits that of Agriogomphus sylvicola Selys.

Archaeogomphus densus spec. nov.

(es 6 AUS 2751)

Agriogomphus infans Ris, 1913: 72—73 (® from Ar-

gentina).

Archaeogomphus infans; Fraser, 1947: 432. Paulson,

WTS N75?

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

Material. — Argentina: Misiones, 14.1v.1909,

1 ® (paratype), Joergensen leg. (SMF, no.

15232). — Brazil: Minas Gerais, Lagoa Santa,

Cimpinho 211097) SN Cd rzitzanles:

(CJ); same locality data, 2 d (AC); same locali-

ty, 21.11.1979, 1 & and 21.iv.1979, 1 d, both C.

Mascarenhas leg. (CM); Minas Gerais, Serra do

Cipó, Santana do Rio Acho, Soberbo stream

(19° 20’ S., 43° 38’ W.), 14.1.1975 (6 p. m.), 1 ©

(paratype), Angelo B. M. Machado leg. (CM);

same locality, 29.11.1975 (1 p. m.), 1 © (para-

type), Paulo A. R. Machado leg. (AC); Santa

Catarina, Nova Teutonia, 3.1.1941, 1 & (holo-

type) (MZM); same locality, 4.1.1941, 1 ® (par-

atype) (MZM); same locality, 11.1949, 1 © (allo-

type) (CC), F. Plaumann leg.

In the venation of the wings this species dif-

fers markedly from all other congeners by the

following two characters: (1) there are three

rows of cells near the marginal row, between

M1 and M2; two in other congeners; (2) the dis-

tal portion of the wings beyond the pterostigma

is more prominent and, correlated with this

character, the pterostigma is nearer to the nodus

than in other congeners; the distance from no-

dus to pterostigma is 2.3 to 2.5 times the length

of the costal edge of the pterostigma, 3 to 3.5

times in other congeners.

In the colour pattern of the pterothorax it dif-

fers from all other congeners in having the first

pale antehumeral stripes isolated.

The nearest relative of the new species is

A. infans. It agrees with A. infans in the co-

alescence of the five veins at the posterior angle

of the triangle in the hind wings. The female

agrees with A. infans in having the superior

margin of the occipital foramen armed with a

pair of submedian spines but the male differs

from this species by the lack of these spines.

Male (holotype; colours of thorax and abdo-

men obscured; abdomen broken between seg-

ments 5 and 6). — Total length, 33 mm; abdo-

men, 25 mm; hind wing, 19 mm; greatest width

of hind wing, 5.5 mm; costal edge of ptero-

stigma in fore wing, 2.0 mm.

Face grey-green, the frons darker but the

frontal ridge paler. Lateral margins of labrum

and facial lobes pale yellow. Free border of la-

brum fringed with brown-yellow hairs. Vertex

dark brown, with well-developed transverse

ridge behind lateral ocelli. Occiput brown,

rounded on edge, without crest line but with

hairs. Anterior part of occiput slightly elevated

at either lateral end. Swollen area of head be-

hind each compound eye dark brown and

sparsely covered with brown hairs. Tempora

yellow. Labium and adjacent mouth parts pale

green.

Prothorax dark brown but anterior border of

first lobe brown-yellow.

Pterothorax dark brown with pale yellowish

brown markings; its colour pattern shaped as

shown in fig. 28.

Legs brown but outer side of tibiae and outer

side of second joint of tarsi yellow. First tibiae

without lamina tibialis.

Wings slightly brown tinged on basal half.

Venation of wings dark brown but frontal mar-

gin of costa yellow. Pterostigma brown, sur-

mounting 2'%2—3'% cells. Intermedian cross-

veins 4—4/2—3 in fore and hind wings, re-

spectively. Base of hind wings strongly angled.

Other venational features as shown in verifica-

tion table 2.

Abdomen brown, the segments 3 to 6 darker

at apex. Lateral dilatations of eighth abdominal

segment well-developed, broadly widening

backward and extending to about one-third the

length of segment 9. Dorsal side of segment 10

basally convex but concave on snout-like pro-

jection. Inner side of tip of dorsal hooks of seg-

ment 10 without minute denticles.

Female (allotype; broken in several pieces but

complete). — Total length, 31 mm; abdomen,

23 mm; hind wing, 20 mm; greatest width of

hind wing, 6 mm; costal edge of pterostigma in

fore wing, 2.4 mm.

Head similar to that of male holotype, but su-

perior margin of occipital foramen with a pair of

submedian spines diverging downward. Pale

markings of pterothorax less extended than in

holotype. Colour pattern of pterothorax shaped

as shown in fig. 6. Legs as in holotype, but

spines of femora longer and less in number. Ab-

domen dark brown. Vulvar lamina extending to

a point about one-third of the way along venter

of segment 10. Apical segments 7, 8, 9, and 10

approximately in ratio 35:28:16:10, with the

caudal appendages 6 on the same scale.

Wings slightly brown tinged, especially on

basal half. Venation in right pair of wings

shaped as shown in fig. 2. Pterostigma sur-

mounting 3—3V2 cells. Intermedian cross-veins

4—4/2—3 in fore and hind wings, respectively.

The males from Lagoa Santa are smaller than

the type; the measurements of the smallest one

are: total length, 30 mm; abdomen, 22 mm;

BELLE: The genus Archaeogomphus 47

hind wing, 17 mm. Further, the dark colours are

darker and pale colours are paler than in the ho-

lotype (the males were put in aceton, whereas

the colour design of the type is obscured due to

post mortem changes). As a result the abdomen

of the males from Lagoa Santa is distinctly an-

nulated on the middle segments (broad black

rings covering joinings of segments and a nar-

rower black ring just before middle of each).

The face, the first pale antehumeral stripes, and

the pale colours of the abdominal segments 3 to

7 of these males are greenish white. The abdo-

minal segments 8, 9 and 10, however, are yellow

except for the nodules which are black. The

lateral dilatations of the abdominal segments 8

and 9 of some males from Lagoa Santa are in-

flexed against the sternum of these segments.

The body coloration of the Argentine female

is largely obliterated due to post mortem

changes. The slender divisions of the vulvar

lamina of this female are more bent from each

other than in the other females.

The variation in the number of antenodal

cross-veins in A. densus is as follows: antenodal

cross-veins in fore wing, male 11 (81%) or 10

(19%), female 11 (80%), 12 (10%) or 13 (10%);

antenodal cross-veins in hind wing, male 9

(75%) or 8 (25%), female 9 (90%) or 11 (10%).

Remarks: (1) The allotype female of A. den-

sus, while in my possession as a loan, had a fall

that broke off the tip of the right fore wing and

left hind wing, after drawings had been made of

the right pair of wings. The wing tips, inclosed

in a cellophane triangular envelope, have been

added to the otherwise complete specimen. (2)

Mr. Carl Cook wrote me (in a letter dated

13.11.1978), that the allotype of A. densus in his

collection at present, will, together with the

whole of his collection, be deposited in the Na-

tional Museum of Natural History, Washing-

ton, D.C.

A NOTE ON THE IMMATURE STAGES

The larva of Archaeogomphus is peculiar by

the libelluline-like appearance owing to its thin

skin, its slender legs, its very large compound

eyes, and its broad lateral labial lobes which are

concave. The tarsi are 2—2—2 jointed, they are

2—2—3 jointed in the larvae of other Neotropi-

_ cal Gomphidae.

Needham (1940) described the exuviae of Ar-

chaeogomphus from Nova Teutonia, Santa Cat-

arina, Brazil. The cast-off skin possibly belongs

to A. densus since this species is known from

the same locality.

Schmidt (1951) studied the sclerotizations in

the lateral body wall of the abdomen of an Ar-

chaeogomphus larva from Nova Teutonia, Santa

‘Catarina. This larva may also belong to A. den-

sus.

Belle (1970) described comparatively the exu-

viae of A. nanus. The cast-off skin belongs to a

reared individual collected in Surinam by Dr. D.

C. Geijskes.

During their explorations in Central-Ameri-

ca, Mr. and Mrs. Beatty as well as Dr. Paulson

collected many larvae of A. furcatus and suc-

ceeded in rearing some imagines from them. A

description of the larva of A. furcatus will be

given by Mr. and Mrs. Beatty.

HABITAT AND BEHAVIOUR IN THE FIELD

There is some environmental and behavioural

information available on Archaeogomphus spe-

cies. Williamson (1918a, 1918b, 1923) described

the localities and circumstances in which he had

collected his Archaeogomphus species in Co-

lombia and Venezuela. As regards the behav-

iour in the field of A. hamatus, Williamson

(1918b) cited the following: “The gomphines

were usually resting on twig tips or similar

perches from six inches to two and one-half feet

high, lower locations being preferred. No dif-

ference was detected in the actions of the sexes.

They were not wary, but flushed, the flight was

followed with difficulty and the individual was

usually lost”.

Dr. Geijskes provided the triangular envelope

in which the unofficial paratype of A. nanus has

been stored with the field note that the speci-

men was taken “in bushes”. This is in agreement

with my experiences in the field with regard to

this species. The favourable environment of

A. nanus is obviously the tropical rain forest. I

found the species in numbers during my collect

ing trips in the interior of Surinam, but nearly

always along the banks of the rivers and larger

streams and only by way of exception along the

banks of the small creeks. Its behaviour in the

field is not typical of gomphids in general.

When searching for specimens of A. nanus I ap-

proached carefully the overhanging bushes and

smaller trees along the banks of the streams in a

small wooden boat masterly navigated on the

swiftly flowing water by a Red Indian. The

males were generally found hidden in these

overhanging bushes and trees, and when dis-

48 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

turbed they flew between the branches over

short distances from one branch to another or

took refuge into the bushes. This circumstance

made it difficult, if not impossible, to net them

and will explain that, although many of these

males were seen, only a relatively small number

of them could be captured. Females were also

often taken by me on the river banks along the

footpaths immediately behind the overhanging

small trees and bushes. Larval cast-off skins can

be found close to the water edge, attached

against the roots or twigs of the bank vegeta-

tion. Dr. Geijskes told me, that, when exploring

the shores of an island of the Marowijne River

in Surinam, he saw a number of specimens

(males?) of A. nanus perching together on a

bare twig “like a row of birds”. This was a most

curious sight.

Dr. Paulson, in a letter (dated 30.1x.1980) to

me, wrote about A. furcatus in Costa Rica:

“They are devilishly hard to find, and I have

never seen a male on territory. One adult female

was collected near, but not at, a fair-sized rocky

river (Rio Santa Rosa) with wooded banks; ac-

tually she was on a twig in dry forest 30’ from

the river and 8’ above the ground. The other

adult (Taboga) was taken about 100’ from a

rather slow, muddy river through moist forest,

also 8’ above the ground on a flat leaf. It was

sunny, time 11:00. Her eyes were dull red over

gray, which seemed an unusual eye color for a

gomphid. Other rivers at which I have taken

them (larvae) include the Corobici, which is

fairly wide and quite rocky, and the Puerto Vie-

jo, which is also large but entirely sandy. The

larvae live in pools with sand and detritus, along

with Phyllogomphoides, Phyllocycla and other

gomphids.” And about the Archaeogomphus in

South Mexico he said in the same letter: “I have

not taken larvae in small streams, although I be-

lieve one or more of the rivers in Chiapas was

no more than 5 m in width”.

Apparently A. furcatus prefers the more open

rivers and larger streams, as A. nanus does in

Surinam. However, this behaviour is not typical

of Archaeogomphus species in general as clearly

appears from the experiences with A. densus. In

a letter (dated 10.iv.1978) to me, Prof. Machado

wrote about the females of A. densus collected

by himself and his son Paulo: “My specimen

was found perching on a dead branch about 2

meters high, inside the small stream called “So-

berbo” close to the point where it ends in the

Cipo River. Paulo’s specimen was found in the

bushes about 10 meters from this stream. My

specimen was collected late in the afternoon

(about 6 p. m.) and Paulo’s at 1 p. m. Although

the place is called “Serra do Cipó” (Cipó moun-

tain) actually it is not on the “Serra” but on the

Cipó River Valley close to the “Serra” at the

municipality of Santana do Rio Acho, a poor

village which you will probably not find on the

map. The exact position of the area is 19° 20’

Lat. and 43° 38’ Long. at about 70 kilometers

North of Belo Horizonte”. Concerning the

males of A. densus collected by himself and

Prof. Jurzitza he wrote in a letter (dated

16.x.1979): “The first males were discovered by

Gerhard Jurzitza about 70 kilometers from the

place where the females were collected. They

were perched in the grass at the banks of a

stream 1—2 meters wide, tributary of the Rio

das Velhas close to the village of Campinho, 10

kilometers from the town of Lagoa Santa. The

vegetation of the region has been described in

detail by the Danish botanist Eugenio Warming

(Warming, J. E. B. — 1892 — Lagoa Santa: Et

bildrag til den biologiske plantegeografi. Det.

Kongel. Danske Videnske. Selskabs. Skrifter. R.

6B: 153—488). It is a kind of savanna known as

“cerrado” with gallery forests along the

streams. However, most forest has been de-

stroyed and no vestige of it exists nowadays

along the stream where the Archaeogomphus

were collected.”

GEOGRAPHICAL DISTRIBUTION

The genus Archaeogomphus is primarily a

South American group but occur also in Cen-

tral-America (fig. 31). The species belonging to

this genus are relatively rare, and due to the

scantiness of the material available a prediction

cannot or can hardly be made as to the occur-

rence of a certain species in a given region.

I. The fans group. A. infans was described

from Espirito Santo, Brazil. Its range apparently

covers a part of the south-eastern coastal border

of Brazil. Sao Paulo lies in this region and this

circumstance corroborates my supposition that

Needham’s (1903) male of Archaeogomphus be-

longs really to this species. A. densus, described

from Nova Teutonia (Santa Catarina), Brazil,

but also known from Lagoa Santa and the near-

by Serra do Cipó (Minas Gerais), Brazil, and

Misiones, Argentina, apparently occurs in the

south-eastern part of Brazil and the north-east-

ern part of Argentina.

II. The hamatus group. A. hamatus was de-

scribed from Fundación, Colombia. Calvert

(1948) recorded the species from Kartabo,

BELLE: The genus Archaeogomphus 49

Guyana. The discovery of A. hamatus in Albi-

na, Surinam, and at the Rio Paraim (Goiás),

Brazil, extends the range of this species further

eastwards and considerably more southwards.

Although no record is known from Venezuela

the occurrence of A. hamatus along the entire

northern coastal region of continental South

America is probable and even its occurrence in

the southern part of Central-America can be ex-

pected. A. furcatus was described from Bejuma,

Venezuela, and Cristalina, Colombia. The new

locality Táchira in Venezuela is situated about

half-way between Bejuma and Cristalina. The

surprising discovery of the species in Costa Rica

(Guanacaste and Heredia) and South Mexico

(Veracruz, west of Cosamaloapan de Carpio;

Oaxaca, south of Tuxtepec; and Chiapas, coast-

al region of the Sierra Madre) extends its range

considerably towards the north. A prediction

can be made as to the occurrence of the species

in the other Central-American countries. A. na-

nus was described from the upper part of the Li-

tani River, Surinam. The present records from

Bolivar, Venezuela, and the Rio Tapirapé, Bra-

zil, extends its range considerably to the West

and South. Obviously the species occurs not

alone in the Amazone basin but also along a

large part of the northern coastal region of

South America.

Using the zoogeographical division of South

America after Sclater & Sclater, 1899 (cf. Rapo-

port, 1968: 68) we can say that the members of

the infans group occur in the Sud-brasileno re-

gion whereas the members of the hamatus

group occur in the Colombiana and Amazónica

regions.

REFERENCES

Anonymous, 1951. Letter of E. B. Williamson to J. G.

Needham dated February 18, 1928. Ent. News 62:

304— 305.

Belle, J., 1970. Studies on South American Gomphi-

dae (Odonata) with special reference to the species

from Surinam. Stud. Fauna Suriname 11: 1—158;

pls. 1—21.

| Belle, J., 1972. Further studies on South American

Gomphidae (Odonata). Tijdschr. Ent. 115: 217—

240.

Belle, J., 1977. Some gomphine material from Suri-

nam, preserved in the Leyden Museum of Natural

History, wıth a note on the larva of Desmogom-

phus tigrivensis Williamson (Anisoptera: Gom-

phidae). Odonatologica 6 (4): 289—292.

Belle, J., 1979. An attempt at the subfamily classifica-

tion of the Gomphidae, based on some new inter-

pretations of wing venation (Anisoptera). Odona-

tologica 8: 43—46.

Belyshew, B. F. & A. Y. Haritonov, 1977. Determiner

of dragonflies (genera of Boreal faunistical king-

dom and some contiguous territories, species of

the USSR fauna). Publishing House “Nauka”, Si-

berian Branch, Novosibirsk, 1977: 398 pp.

Calvert, P. P., 1935. Edward Bruce Williamson. Ent.

News 46: 1—13; pl. 1.

Calvert, P. P., 1948. Odonata (dragonflies) of Karta-

bo, Bartica District, British Guiana. Zoologica,

N.Y. 33: 47—87; pls. 1-2.

Fraser, F. C., 1940. A comparative study of the penes

of the family Gomphidae (order Odonata). Trans.

R. ent. Soc. Lond. (A) 90: 541—550; pls. 1—6.

Fraser, F. C, 1943. Notes on the genera Agriogom-

phus Selys and Ischnogomphus Williamson with

the description of the male of Agriogomphus syl-

vicola Selys (Odonata). Proc. R. ent. Soc. Lond.

(B) 12: 161166.

Fraser, F. C., 1947. The Odonata of the Argentine

Republic I. Acta zool. lilloana 4: 427—461.

Fraser, F. C., 1957. A reclassification of the order

Odonata. Roy. zool. Soc. N. S. W., Sydney. 133

pp.; 1 pl.

Gaige, F. M., 1934. Edward Bruce Williamson 1877—

1933. Rpt. Dir. Mich. Mus. Zool. 1932—33: 7—

24;1 pl.

Klots, E. B., 1944. Notes on the Gomphidae (Odona-

ta) with descriptions of new species. Am. Mus.

Novit. 1259: 1—11.

Needham, J. G., 1903. A genealogic study of dragon-

fly wing venation. Proc. U. S. natn. Mus. 26:

703—764; pls. 3334.

Needham, J. G., 1940. Studies on Neotropical gom-

phine dragonflies. Trans. Am. ent. Soc. 65: 363—

394; pls. 20—22.

Needham, J. G., 1944. Further studies on Neotropical

gomphine dragonflies. Trans. Am. ent. Soc. 69:

171—224; pls. 14—16.

Paulson, D. R., 1977. Odonata. In: S. H. Hurlbert,

Ed., Biota acuatica de Sudamérica Austral: 170—

184. San Diego St. Univ.

Rapoport, E. H., 1968. Algunos Problemas bi-

ogeográficos del nuevo mundo con especial re-

ferencia a la región neotropical. Biol. Amér. aust.

4:53—110.

Ris, F., 1909. Libellulinen in Coll. Zool. de Selys

Longchamps 1: 1—120. Illus. Brussels, Hayez,

Impr. des Academies.

Ris, F., 1913. Neuer Beitrag zur Kenntnis der Odona-

tenfauna von Argentina. Mem. Soc. ent. Belg. 22:

55—102.

Schmidt, E., 1951. Sclerotizations in the lateral body

wall of the nymphal abdomen in Odonata. Ent.

News 62: 125— 134.

St. Quentin, D., 1967. Entwicklungstendenzen im

Flügelgeäder der Odonaten. Entom. Abb. Mus.

Tierk. Dresden 32 (18):311—339.

St. Quentin, D., 1973. Die Gomphidenfauna Südame-

rikas. Annln naturh. Mus. Wien 77: 335— 363.

Tillyard, R. J. & F. C. Fraser, 1938— 1940. A reclassı-

fication of the order Odonata. Based on some new

interpretations of the venation of the dragonfly

50 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

wing. Aust. Zool, 9: 125—169; 195—221; 359—

396.

Williamson, E. B., 19182. A collecting trip to Colom-

bia, South America. Misc. Publs Mus. Zool. Univ.

Mich. No. 3, February, 1918: 1—24.

Williamson, E. B., 1918b. A new species of Agrio-

gomphus (Odonata). Occ. Pap. Mus. Zool. Univ.

Mich. 59: 1—18; pls. 1-2.

Williamson, E. B., 1919a. Archaeogomphus, a new

genus of dragon-flies (Odonata). Occ. Pap. Mus.

Zool. Univ. Mich. 63: 1—8; pl. 1.

Williamson, E. B., 1919b. Notes on species of the ge-

nus Heteragrion Selys with descriptions of new

species (Odonata). Occ. Pap. Mus. Zool. Univ.

Mich. 68: 1—88; pls. 1-12.

Williamson, E. B., 1920. A new gomphine genus from

British Guiana with a note on the classification of

the subfamily (order Odonata). Occ. Pap. Mus.

Zool. Univ. Mich. 80: 1—12; pl. 1.

Williamson, E. B., 1923. A new species of Archaeo-

gomphus (Odonata). Occ. Pap. Mus. Zool. Univ.

Mich. 134: 1—10; pl. 1.

BELLE: The genus Archaeogomphus

VE TS

[OA

Cu2 Cul

Wilhamson (para-

2 shaded): 1, A. furcatus

air of wings of female (cells mentioned in table

Figs. 1-2. Right p

c. nov. (allotype), Brazil.

type), Colombia; 2, A. densus spe

52 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

Figs. 3—6. Diagram of pterothorax: 3, A. infans (Ris) d, Brazil; 4, A. furcatus Williamson © (paratype), Co-

lombia; 5, A. nanus Needham d, Surinam; 6, A.densus spec. nov. ® (allotype), Brazil. Figs. 7—8. Occiput of

female, dorsal view: 7, A. hamatus (Williamson), Surinam; 8, A. furcatus Williamson (paratype), Colombia.

Fig. 9. Archaeogomphus furcatus Williamson © (paratype), Colombia. Superior part of rear of head, showing

postgenal cicatrices (pgc).

BELLE: The genus Archaeogomphus 53

Cig

19 20

Figs. 10—14. Vulvar lamina: 10, A. infans (Ris), Brazil; 11, A. hamatus (Williamson), Surinam; 12, A. furcatus

Williamson (paratype), Colombia; 13, A. nanus Needham, Surinam; 14, A. densus n. sp. (allotype), Brazil.

Figs. 15— 18. Superior margin of occipital foramen of female; 15, A. infans (Ris) (holotype), Brazil; 16, A. ha-

matus (Williamson), Surinam; 17, A. nanus Needham, Surinam; 18, A. densus n. sp. (allotype), Brazil. Figs. 19—

22. Outline of penial peduncle, posterior view: 19, A. infans (Ris), Brazil; 20, A. hamatus (Williamson) (para-

type), Colombia; 21, A. furcatus Williamson (holotype), Venezuela (After Williamson, 1923); 22, A. nanus

Needham, Surinam.

54 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

Figs. 23—26. Archaeogomphus infans (Ris) 3, Brazil: 23, apical segments of abdomen, left lateral view; 24,

ninth and tenth abdominal segments, dorsal view; 25, accessory genitalia, right lateral view; 26, right pair of

wings (cells mentioned in table 2 shaded).

BELLE: The genus Archaeogomphus | 55

Figs. 27—30. Archaeogomphus densus n. sp., holotype 4, Brazil: 27, apical segments of abdomen, left lateral

view; 28, diagram of pterothorax; 29, accessory genitalia, right lateral view; 30, right pair of wings (cells men-

tioned in table 2, shaded).

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 3 (1982)

hamatus

furcatus

nanus

infans

densus

Fig. 31. Distribution of the species of Archaeogomphus Williamson.

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J. P. Brock. — A systematic study of the genus Ophion in Britain (Hymenoptera,

Ichneumonidae), p. 57—97, figs. 1—52, graphs 19.

Tijdschrift voor Entomologie, deel 125, afl. 4 Gepubliceerd 31-XII-1982

A SYSTEMATIC STUDY OF THE GENUS OPHION IN

BRITAIN (HYMENOPTERA, ICHNEUMONIDAE)

TERSBROGR

Imperial College, London, U.K.

ABSTRACT

A taxonomic revision of the British species of Ophion is presented, including study of

strategic type material. Fourteen species are recognised, two of which are new. Bobs

and(distributionalldata are summarised as far as the existing data allow. A full discussion of

criteria of species recognition is given, and some problems which suggest future experi-

mental work are outlined.

CONTENTS

TERN SEO ea ANT War. elende We 57

Nomie tae eee Es DNA UE. et 58

Ereimowledsements RI. LME oe. kee. es 58

Characters used in definition of species ... 59

{Uriteriaforspeciesilimits Austen. ue 62

EBEHERIEIIASDOSISU. NE AIR ER US Mal 63

ESeyavonspeciesinis ont vat ear Min 63

BY:SCENGARICIPArte. ARE N Lol MARIE 74

Eimmanylolhostspeeiesm.i.. Wa. es doi 95

CLUS ee e a 96

Appendix (notes on Platophion) ......... 96

EKAIN) a ENNE 96

INTRODUCTION

The species of Ophion are familiar testaceous

Ichneumonids of generally nocturnal habits,

frequently taken in light traps. With the excep-

tion of two distinctive species, they present un-

usual taxonomic difficulty — not only due to

the close relationship of species one to another,

but also owing to the quite exceptional variabili-

ty of those structural characters which can be

used to define species limits. Ophion species are

parasites of (mainly) lepidopterous larvae

(usually Noctuidae), but determinations of the

species have been so spurious in the past that a

large proportion of the published host records

requires further confirmation.

By the time Morley (1915) came to mono-

graph the Briush species of the genus, a number

of species had been proposed by Ratzeburg

(1848), Thomson (1888), Kriechbaumer (1879)

and Brauns (1889), to add to three “traditional

species” of Linnaeus (1758), Fabricius (1798)

and Gravenhorst (1829). Most of the later spe-

cies were inadequately distinguished from /u-

teus of Linnaeus, certain of han actually being

based upon abnormal colour or structural vari-

ants of otherwise “anonymous” species. Morley

himself (loc. cit.) described several new species

from the British fauna; certain of these were

quite valid, although others have been shown to

be synonyms of earlier species (see Gauld,

1976). Other keys to European species of the

same period (notably that of Schmiedeknecht,

1908) tended to be compilatory in nature and

added comparatively little to an understanding

of the luteus complex. It is to Morley’s credit

that he correctly dismissed a number of charac-

ters used by authors as being unstable within a

species, but at the same time, he was unable to

find more than a very few new features on

which to base his own diagnoses.

With the doubtful exception of Schmiede-

knecht’s revision of his own earlier key (1936),

no real advance was made over Morley until

comparatively recently. However, J. F. Perkins

of the British Museum (Natural History) had

arranged the national collection according to his

own assessment of both old and new taxonomic

characters, supplemented by study of important

type material during the 1930’s. Perkins also

drew up a manuscript key to the British species

of Ophion and related genera, which work has

been used by several subsequent workers, in-

cluding the present author. The manuscript was

an incomplete study, since its author was work-

ing more intensively on his revisions of Ichneu-

moninae and Pimplinae; the key remained ten-

tative in nature, since Perkins did not include a

detailed study of infraspecific variation, nor did

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

he use all characters which have now come to

light. Nevertheless, his work did constitute a

great advance over the keys of Morley and

Schmiedeknecht.

More recently, Gauld (1973, 1976, 1978) has

published several papers on the British species

of Ophion. He was able to confirm many of the

synonyms indicated by the Perkins manuscript

and, to some extent, to reassess some of the

characters used to define species. Gauld also

drew attention to some characters which had

not previously been used to define Ophion spe-

cies. At a later stage, Gauld (1978) produced a

key to the species of the difficult /uteus-mocsa-

ryi complex. Oosterbroek (1978), partially fol-

lowing Gauld (loc. cit.), has written an account

of the Dutch species, which includes several

fresh taxonomic observations. The key given by

Gauld now allows identification of a much

larger proportion of material than could possi-

bly be gained from the combined works of

Morley and Schmiedeknecht, but it will still be

found that a dangerously large percentage of

specimens will present great difficulty in identi-

fication with this key. This fact is due to two

reasons:

(1) Gauld relies on the Perkins manuscript key

for definition of species. As a consequence,

many tentative observations which Perkins

made and which were far from being at all fully

worked out, are now used by Gauld in much

the same taxonomic context as Perkins sug-

gested using them;

(2) Gauld apparently greatly underestimates

the variability of many of the characters which

he uses; this includes both Perkins characters

and certain of those discovered by Gauld him-

self.

In trying to solve the problems of species def-

inition in the Ophion luteus complex, I have

found myself adopting a very flexible concept of

species limits, since I find that nearly all avail-

able characters present an alarming degree of

variation — even compared to other Ichneumo-

nid genera which I have studied. I have entirely

omitted several characters of authors, which ap-

pear to be much too variable to be of any value.

I have also placed several others (including

some of those studied by Perkins and Gauld) in

a more confirmatory than diagnostic context

than has hitherto been proposed. In addition, I

have brought forward a number of new charac-

ters which greatly assist in defining species lim-

its. The key and descriptions given here stress

features which I have found to be less variable,

supported by other somewhat more plastic varı-

ables which nevertheless help confirm species

diagnosis. I have usually quoted several charac-

ters simultaneously in the key, since this prac-

tice makes allowance for variants which may

lack one or more of the usual defining features

of a species. This accounts for the complexity of

the key, but this difficulty is inevitable if accu-

rate determination is to be procured.

NOMENCLATURE

Owing to the inadequacy of many published

descriptions, along with the consequent misde-

termination of much Ophion material by past

authors, it has not been possible to present a

complete study of synonymy. I have followed

what I believe to be the best solution, in dealing

principally with the work of more recent au-

thors, with original descriptions, and to some

extent with identifications of workers for which

material has actually been made available for re-

examination. Several type specimens have ap-

parently been lost, and the possibility remains

that further research could well bring to light

some data which might lead to subsequent ad-

justment in nomenclature.

Generic synonymy was treated by Townes

(1971), and is not repeated here.

ACKNOWLEDGEMENTS

For loan of specimens, I am grateful to: M.

Fitton, British Museum (Natural History)

(BMNH); I. Gauld, Commonwealth Institute

of Entomology, London; A. Brindle, University

Museum, Manchester (MU); P. Oosterbroek,

Instituut voor Taxonomische Zoologie,

Amsterdam; J. Aubert, Laboratoire d’Evolution

des Etres Organisés, Paris; E. Königsmann,

Zoologisches Museum, Berlin, J. Papp, Termes-

zettudomanyi Muzeum, Budapest; E. Diller,

Zool. Staatssammlung, Munich.

In addition, many specimens were loaned or

donated by private collectors. The reared

material of M. R. Shaw (Reading), and the light

trap catches sent by the late L. W. Siggs (Min-

stead, New Forest, Hants.), were of quite ex-

ceptional value. Other material (mostly taken at

light) was given to me by: M. C. Birch, K.

Stewart, C. O’Toole, M. R. Shaw, J. St. E. Car-

dew, R. B. Angus, I. Lansbury, M. J. Smith, N.

Rae-Jones, and M. P. Hassel. Most of this

material is now in the Brock collection (BK),

Ulster Museum, Belfast.

I am indebted to the Science Research Coun-

cil (U.K.) for a grant in support of Ph. D. work

Brock: Ophion in Britain 59

(which included the present study) at Imperial

College, London. R. G. Davies of that institu-

tion helped with statistical treatment of some of

the data and also read through the manuscript,

making helpful suggestions towards im-

provement.

CHARACTERS USED IN DEFINITION OF SPECIES

Head characters

1. The ocellar-ocular interspace, i.e., the dis-

tance between a posterior ocellus and the near-

est point on the compound eye, compared with

the interocellar space (that between the posteri-

or ocelli). Species of the luteus complex fall into

two sections, one in which the posterior ocelli

are nearly or quite in contact with the eyes (e.g.

fig. 40) and a second group of species in which

there is a distinct, often large ocellar-ocular in-

terspace (see figs. 11, 25). Gauld (1973) terms

these species groups the luteus and mocsaryi

groups, respectively, at first freely following the

Perkins (MS) definition, later (1978) attempting

a modification of this. Unfortunately, intraspe-

cific variation does not allow diagnosis of the

two species groups as defined by Gauld. In

practice, males of two mocsaryi group species

very often have a distinct ocellar-ocular inter-

space, which is not smaller than that found in

many specimens in the other species group. In-

deed, it is quite frequently larger.

2. The distance between the posterior ocellus

and occipital carina — here compared to the

maximum (usually apical) width of the first fla-

gellar segment, also with the diameter of a pos-

terior ocellus. These characters seem correlated

with the last, but have not previously been used

in the taxonomy of Ophion.

3. The length of the temple: the maximum

length of the temple as seen in lateral view, here

compared to the maximum transverse diameter

of the compound eye.

4. The stemmaticum: this is the raised area

bearing the three ocelli. In Ophion it is bounded

by sulci of varying definition. The posterior sul-

cus (see fig. 51) lies behind the posterior ocelli.

The lateral sulcus lies between the posterior

ocellus and the compound eye and runs for-

wards to the front ocellus (see fig. 33). The pos-

terior and lateral sulci of the stemmaticum may

or may not connect, since the former frequently

runs directly into the margin of the compound

eye. Sometimes the connection to the eye is

through a stemmatico-ocular line, which is quite

distinct from the sulci bounding the stemmati-

cum (see fig. 28). The degree of definition of

these sulci has been compared one to another,

also with that of the middorsal sulcus, which

lies between the posterior ocelli on the middor-

sal line of the head. — Although variable within

species limits, these features are useful in defin-

ing species and supraspecific groups within

Ophion. They have not been used previously in

the taxonomy of the genus and are here defined

for the first time.

5. The occipital carina. This may be rounded,

squared, angled or dipped centrally on the ver-

tex. It is a variable character which nevertheless

remains useful in defining species of Ophion.

Although much used in Ichneumonid taxonomy

generally, the form of the occipital carina has

not been indicated in previous studies of the

present genus.

6. The gena: comparison of genal length with

the width of the base of the mandible is a con-

ventional character. It was used by Perkins in

his manuscript key to Ophion, followed and ex-

panded upon by Gauld. Less precise references

to the form of the gena will be found in earlier

keys to European Ophion species.

7. The postgena and genal inflection: the

length of the postgena is here defined as the dis-

tance between the mid point of the genal inflec-

tion and the nearest point on the compound eye.

The genal inflection is that length of hyposto-

mal carina lying between the origin of the latter

at the mandible base and its junction with the

genal carina. These dimensions are further clar-

ified by fig. 26, and are so defined for the first

time. I have compared the length of the post-

gena with that of the genal inflection, the latter

in turn with the width of the mandible base.

8. The compound eyes in frontal view: these

may be approximately parallel, divergent (fur-

ther apart towards mouth than towards vertex),

or convergent.

9. Facioclypeal area: delimited above by the

line of the antennal sockets, laterally by the

margin of the compound eyes, below by the

lower edge of the clypeus. The shape of this area

is significant and obviously correlated with the

last mentioned character. Neither has been used

previously in definition of Ophion species.

10. The mandibles: of particular significance

is the degree of puncturing present on the ven-

tral flange, along the more medial region. Also,

the gape — here defined as the angle between

the two teeth. The internal angles of the teeth lie

in the excised triangular area where the teeth

coalesce, and vary in the extent to which they

are defined (compare figs. 14 and 37). Finally,

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

the basal width of the top edge of the mandible

varies in comparison to the maximum width of a

tooth (see figs. 45 and 46). Several of these char-

acters are here defined for the first time, none

have been used previously in the taxonomy of

the genus. Morley (1915) refers to the acute or

obtuse nature of the apical teeth of the mandi-

bles, but this feature is too variable within spe-

cies to be of any real use in definition of taxa.

11. Sculpture of epistoma and orbits: I have

compared the distribution of epistomal punc-

tures with those of the facial orbits. This is a

variable character, but useful in defining certain

species.

12. Maxillary palpi: the last segment is gener-

ally distinctly longer than the penultimate, but

in certain species these may be of more or less

equal length.

The antennae

13. First segment of flagellum: length/width

ratios given refer to the maximum (usually api-

cal) width. Gauld (1978) uses the 10th segment

in this context, but I do not find the distal seg-

ments any less variable than the proximal. In

any case, the limits given in Gauld’s couplet

seven (loc. cit.) are subject to much overlap in

actual practice.

14. Number of flagellar segments: this is

subject to much variation within a species and

has consequently been little used by authors. I

have used this character within a more statistical

concept of species limits, the numerical nature

of variation being naturally suited to this kind

of analysis. Gauld follows Perkins in using the

number of flagellar segments to define minutus

and scutellaris. With regard to the difficult /4-

teus — mocsaryi complex, Oosterbroek (1978)

tentatively suggests limits of variation to certain

species — limits which must be extended

through study of additional material in the pre-

sent work.

15. Sensilla of first flagellar segment: Gauld

(1978) uses this character to define both species

and supraspecific groups to some extent. I have

not found the limits given by Gauld reliable in

practice. Certain species which normally pos-

sess sensilla over most of the surface of the first

flagellar segment show variation to the extent

where they may be nearly or quite absent. Con-

sequently, I have omitted this character from

the present study. I shall mention in addition,

that certain species which seem particularly well

defined with reference to this character are

known from only a small number of specimens.

As a general rule, I think it unwise to place too

much weight on a character which is seen to be

subject to much variation in other species for

which suitably large samples have been available

for study of intraspecific variation.

The thorax

16. Sculpture: of particular use is the distri-

bution of punctures on the mesopleura, also the

form of microsculpture in that region. I have

usually only drawn attention to divergence

from the normal condition of moderate punctu-

ration and weak microsculpture in descriptions.

17. Colour: most species are largely testa-

ceous in colour. Gauld’s key (1978) identifies

only minutus and obscuratus (together with the

rare forticornis) as having yellow lines on the

mesonotum (see fig. 6b), but obscuratus fre-

quently lacks these markings and there are other

species in which the mesonotum is sometimes

vittate (e.g., parvulus and luteus, also longige-

na).

18. The epicnemium: I have used several new

characters from this region, some of which re-

quire definition. The sternal angles lie ventrally,

behind the front coxae, the pleurosternal angles

at the point where the pleural epicnemium sud-

denly turns into a more vertical plane. The ap-

proximately horizontal portion of the epicnemi-

um lying in the pleurosternal region, in front of

the pleurosternal angle, I have termed the lower

sector of the (pleural) epicnemium. The upper

sector is that portion of the pleural epicnemium

which lies between the pleurosternal angle and

that point at which the epicnemium turns for-

wards to meet the front edge of the mesopleu-

ron. These structures are shown in fig. 2. Char-

acters of particular importance are: (a) shape

and angulation of the pleurosternal angle (fig. 2,

SP); (b) the degree to which the sternal and

pleurosternal angles are aligned along a trans-

verse plane (compare figs. 36 and 49); (c) the ra-

tio of lengths of lower and upper sectors of the

pleural epicnemium (see figs. 2, X/Y, 49, 52).

19. Scutellar carinae: in certain species these

are more or less frequently well defined. In

Gauld’s key (1978) the stability of this character

has been overestimated. Thomson (1888) used

the scutellar carinae in his treatment of Ophion,

while Morley (1915) correctly assigns a measure

of instability to the character. It is nevertheless

useful (within limits) for definition of species

within the genus.

BrocK: Ophion in Britain 61

The wings

20. Distribution of trichiae; Perkins (MS)

was first to realise the significance of this char-

acter, particularly with reference to the distri-

bution of trichiae on the submedian cell of the

forewing. In attempting a reassessment of the

reliability of Perkins’ observations with refer-

ence to infraspecific variation, I find I have been

gradually led to the belief that, while the trichia-

tion of the submedian cell remains a useful con-

firmatory character for a fair proportion of

material studied, it is less than reliable when

used as a key character. I have regularly met

with sizeable population samples in which a

species normally having an evenly trichiose sub-

median cell has the trichiae more sparsely dis-

tributed above than below — and vice versa.

This breaks down the apparent division be-

tween luteus and those species related to longi-

gena, especially since more buccate headed ex-

amples of luteus are liable to overlap with the

longigena subgroup, not only in head charac-

ters, but also in the distribution of trichiae on

the submedian cell. Furthermore, the parallel

subdivision of the mocsaryi group on the basis

of a similar character dichotomy, is prone to a

very wide area of overlap. Further investigation

of the distribution of trichiae on the forewing

however, has led to the discovery of a rather

more stable character concerning the degree to

which these trichiae extend onto the area imme-

diately beneath the prestigma. The distribution

of trichiae in this region is compared with that

found elsewhere on the wing; the bare area be-

neath the stigma (substigmal speculum) may ex-

tend beneath the prestigma (compare figs. 39

and 48). Gauld’s (1978) use of the trichiation of

the submedian cell follows the Perkins manu-

script and is unreliable for reasons discussed

| above.

21. The nervellus: in conventional nomencla-

ture, the nervellus of the hind wing may be bro-

ken at, above or below the centre. This is a very

unstable character in Ophion (see remarks on

costatus Ratzeburg).

22. Radius of forewing: the radius may be

slightly thickened towards the base in certain

species. Perkins (MS) drew attention to the non-

sinuate radius of parvulus. This sinuation occurs

on the second abscissa (see figs. 38a/b).

Venational terminology is that used by past au-

thors working with the Ichneumonidae, rather

than the Comstock-Needham system.

Legs

23. Hind leg: the length/width relationships

of the hind femur may be very variable within a

species, but a trend towards more or less elon-

gate shape can definitely be idenufied in several

species. I do not use actual length: width ratios

in descriptions, as these are both variable and

difficult to estimate in normal taxonomic prac-

tice. Fig. 30 illustrates the usual range of shapes

found in the hind femur of Ophion species.

Gauld (1978) was the first author to draw atten-

tion to variation in the form of the hind tro-

chantellus in Ophion. He showed that luteus

and its near allies have a more elongate hind tro-

chantellus than other species. I find I can fully

confirm Gauld’s observations for all but one

Ophion species which, due to an unusual vari-

ability in respect of this character, will not run

correctly in that author’s key. In addition to the

form of the hind femur and trochantellus, I have

tound the shape of the hind coxa useful in defin-

ing one species.

24. The tibiae: there is useful interspecific

variation in the degree of development of the

tibial spinules, especially those of the hind ubia

(see fig. 41). This character has not been used

previously in Ophion; it is used in a largely

“confirmatory” rather than diagnostic context

herein. Perkins, in his manuscript key to

Ophion, made use of the relative lengths of the

apical spurs of the middle tibia in defining cer-

tain species. Gauld (1978) follows Perkins, but

without critical reappraisal of the variability of

the character (see remarks under O. luteus).

The propodeum

25. Past authors have made reference to the

greater or lesser development of the carinae

defining the area superomedia (see fig. 1). This

is a variable character, though nonetheless valid,

providing other features are taken into consid-

eration at the same time. Of rather more use as a

taxonomic character is the degree of devel-

opment of the carina lateralis in relation to that

of the other, more dorsal propodeal carinae (see

figs. 43 and 50). This is an easy character to ap-

preciate and it is rather less variable within a

species than other propodeal features. Curious-

ly enough, it has been overlooked by previous

authors who have worked on Ophion.

The abdomen

26. The most useful character I have found

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

on the abdomen relates to the relative position

of the petiolar spiracles and the ventral mem-

brane of the same segment. This simple charac-

ter has also been overlooked by authors (see

figs. 35 and 43). — Morley (1915) refers to the

degree of basal constriction of the third tergite

as a species level character in Ophion, but I have

not found this at all stable within a species. — A

preliminary study of the male genital sclerites

was also made. Some potentially useful charac-

ters were found, but it would not have been

possible to obtain sufficient material for dissec-

tion with some of the less common species.

Since infraspecific variation could not therefore

be fully assessed, genitalic characters have been

left aside in this revision. It is often quite

wrongly assumed that these structures possess

some special property whereby they are ex-

cepted from the necessity of consideration of

stability; personal experience suggests that this

is far from being the case.

CRITERIA OF SPECIES LIMITS

In any taxonomically difficult species group,

there is bound to be controversy surrounding

the selection of criteria for species recognition.

From a practical as well as theoretical stand-

point, these difficulties are likely to present

their greatest obstacle within what can be

termed close species pairs. In purely pheneuc

terms, a close species pair consists of any given

species, together with that other from which it

is most difficult to separate morphologically

From a biological viewpoint, we shall be con-

cerned with questions relating to whether or not

such segregates do represent species isolates, or

whether they may simply reflect some disjunct

aspect of infraspecific variation. In Ji

there are several close species pairs within

which statistical analysis is very desirable for a

logical, objective answer to be made to such

questions. The actual analysis proceeds along

several inter-related lines of inquiry:

(1) Statistical aspects of morphological isola-

tion. Where statistical rather than absolute dif-

ferences exist between members of a close spe-

cies pair, some form of comparison of character

state frequency distributions is obviously desir-

able. Two points must be made here: (a) where

sexual dimorphism or any other factor affects

the shape of the distributions, a non- parametric

test should replace the usual parametric equiva-

lent for significance testing; (b) assignation of

identity to specimens used for investigating sus-

pect bimodality in continuous variables must be

based entirely upon characters other than that

being analysed (otherwise, an element of circu-

lar reasoning will enter into our species con-

cept!).

I have adopted the Mann-Whitney test in

place of the more familiar “t” test for compari-

son of frequency distributions. Such distribu-

tions rarely seem to follow a near-normal form.

For the most part, I have been concerned with

comparison of the number of flagellar segments

between members of a close species pair, since

this character possesses a large variance, which

is also easily compartmentalised on a direct nu-

merical scale. Specimens from which these data

were drawn had previously been identified on

characters other than flagellar segment number.

An expected result for isolating members of a

close species pair would be based on significant

bimodality of two approximately symmetrical

distributions for number of flagellar segments.

Areas of actual overlap would be biologically

acceptable, owing to the fact that initial species

diagnosis was not based upon that same charac-

fers

(2) Behavioural criteria. There are two very

important aspects of behaviour which bear di-

rectly on the question of species identity: (a) the

temporal frequency distribution (based upon

ee of capture for as large a sample as possi-

ble); (b) host preference. Expected correlations

for true species isolates would be based on the

hypothesis that closely related species would be

likely to exhibit behavioural divergence in these

two niche-related parameters. In parasites, it

might be expected that selected hosts appear at

different seasons for two related parasite species

— or that two synchronously appearing para-

sites avoid competition by modifying their host

preferences. Where two morphologically dis-

tinct species appear at the same time, attacking

the same hosts, we may have reason to suspect

that these are not biologically distinct units.

Areas of difficulty.

There are several complicating factors which

render the foregoing criteria less than perfect in

reality. Dimorphism (and polymorphism) may

occur not only in relation to obvious sexual dif-

ferences, but also in correlation with environ-

mental factors. Where the host range of a spe-

cies contains a disjunct size distribution, the

parasite may also exhibit disjunct variation. En-

vironmental influences acting upon species with

an extended flight period may also be reflected

in morphological variation in parasites. Mem-

BROCK: Opbion in Britain 63

bers of a close species pair could therefore be

forms of a single species which has been sub-

jected to differing environmental influences sur-

rounding embryogenesis.

In attempting to find a solution to one prob-

lem of the kind just mentioned, I have looked

for evidence arising from the regression of fla-

gellar segment number on wing length. This is a

special rather than general technique, which is

explained in detail at a later point.

A further problem lies in the possibility that

biologically distinct species need not exhibit

morphological differences. It is possible that

certain bivoltine parasite species really represent

two biologically isolated forms, each with a

complete annual life cycle. This will come to

light only when (for example) “spring brood”

parasite larvae produce adults the following

spring instead of appearing at the normal “se-

cond brood” time. However, this could also be

interpreted as a single parasite species with dif-

ferent developmental periods dependent upon

host size or upon other environmental factors.

This is a real, rather than imaginary problem

with certain Ophion species. Hopeful solutions

to these difficulties are presented following for-

mal description of the species concerned. In one

case, a small scale behavioural experiment was

conducted in order to support the interpretation

of other sources. The relative scarcity of rearing

records and utter lack of breeding experiments

with Ophion obviously contribute greatly to

some of the major problems discussed herein.

GENERIC DIAGNOSIS

The genus Ophion can be recognised by the

recent keys of Townes (1971), also Gauld

(1973). Gauld also discusses recognition of Pla-

tophion as a genus distinct from Ophion, the

former segregate not having been recognised by

Townes (loc. cit.). Most of the characters given

by Gauld for separation of Platophion may oc-

cur in Ophion s. str., but there still remains

some evidence to suggest that these genera are

best retained as separate entities (see also Oos-

terbroek, 1978). The genus Platophion is not

treated in the present study. I have appended

short notes on species discrimination in Plato-

phion (Appendix).

KEY TO THE SPECIES

1. Small species (wing length at most 11 mm);

nervellus distinctly reclivous above (fig. 3);

body frequently yellow marked (cf. fig. 6b),

never black patterned; flagellum with 40—

49 segments; propodeum with reduced ca-

rination, at least in region anterior to the

posterior transverse carina (cf. fig. 5); radi-

us somewhat thickened towards junction

with stigma (fig. 4); clypeus up to more

than twice wider than high. (Widely dis-

tributed, often common)... minutus Kr.

Wing length usually over 11 mm, nervellus

rarely distinctly reclivous, excepting when

body distinctly black patterned, or when

there are less than 50 flagellar segments;

propodeum rarely lacking carinae anterior

to posterior transcarina (cf. figs. 15, 50); ra-

dius not, or scarcely thickened towards

stigma (fig. 38); clypeus usually much less

thanitwiceiwiderthantie he 2

. Body with distinctive black pattern (cf.

fig. 6a); thorax with large punctures; wings

orange tinted; nervellus tending to be recli-

vous above, radius slightly thickened to-

wards stigma. (Widely distributed, seldom

COMMON) EEE ventricosus Grav.

Head and thorax not black marked; thorax

with superficial puncturation; wings gener-

ally hyaline; nervellus rarely reclivous

above, radius not noticeably thickened to-

wand sistem aten ALG ee EERE. 3

. Antennal flagellum rarely with less than 64

segments; hind femur very slender (cf.

fig. 30a), hind coxae elongate — appearing

smaller to slightly larger in area than the

lateral area of the propodeum, latter often

with no keel connecting the carina lateralis

to the spiracle (fig. 8); mid tibial spurs of

nearly equal length; punctures of epistoma

tending to be further apart than those of fa-

cial orbits (fig. 9); ocellar-ocular interspace

from almost absent in females to 0.65 X in-

terocellar space in some males, diameter of

hind ocellus often as much as twice distance

between ocellus and occipital carina; meso-

notum often dark suffused. (Common spe-

cies of early spring) ....... scutellaris Ths.

Flagellum usually with less than 64 seg-

ments; hind femur less slender (figs. 30b-

d); hind coxae slightly to very much larger

in area than the lateral area of the propo-

deum, carina lateralis generally connected

to spiracle by a keel (fig. 43); mid tibial

spurs usually of very unequal length; punc-

tures of epistoma often as close or even

closer together than those of orbits (e.g.

fig. 21); ocelli various, hind ocelli frequent-

ly smaller in relation to distance to occipital

carina; mesonotum only dark suffused

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Fig. 1. Propodeum, dorsal (generalised), showing terminology of carinae and areae: cl, carina lateralis; ap, area

petiolaris; ab, area basalis; as, area superomedia; ad, area dentiparis. Fig. 2. Mesosternopleuron, antero-lateral,

showing terminology of the epicnemial carina. sp, sternopleural angle; s, sternal angle; x, upper sector; y, lower

sector. Fig. 3—5. Ophion minutus Kriechbaumer; 3, nervellus of hind wing; 4, fore-wing — stigma and base of

radius; 5, propodeum, dorsal (maximum development of dorsal carinae). Fig. 6. Head, thorax and propodeum,

showing patterning in: (a) Ophion ventricosus Gravenhorst, (b) O. obscuratus Fabricius. Fig. 7. Ophion scutel-

laris Thomson, lateral carinae (c) of mesoscutellum.

BROCK: Ophion in Britain 65

when yellow markings are also present

(re, GID) eg veen PRE 4

INTO TESI ats N pe 5

Femaless Aa O EEN 6

. Ocellar-ocular interspace at least wider

than the posterior sulcus of the stemmati-

cum (fig. 33) — very often in range 0.25 to

1.0 X interocellar space (see figs. 11, 18,

23); posterior sulcus of stemmaticum most

usually joining lateral sulcus without first

touching compound eye, although it may

connect to latter by means of a stemmatico-

ocular line (fig. 28), especially when the

ocellar-ocular interspace is long. ........ 7

Posterior ocelli actually touching eyes

(figs. 47, 51), or separated from them by

width of the posterior sulcus of the stem-

maticum only — in which case the latter

touches the compound eye, often without

connecting to the lateral sulcus (which may

be weak or absent) (fig. 40); stemmatico —

ocular line rarely in evidence .......... 22

. Ocellar-ocular interspace wider than poste-

rior sulcus of stemmaticum, (fig. 33), very

often in range 0.2 to around 0.5 X interocel-

lar space (figs. 11, 20, 23); posterior sulcus

of stemmaticum very often connecting with

lateral sulcus — whether or not former

jomseompeundreye(ch. mg, 33)... 7

Posterior ocellus touching compound eyes,

or separated from them only by the width

of the posterior sulcus of the stemmaticum

— latter often running into back of eye

without connecting to lateral sulcus of

stemmaticum, which latter may be weak or

alsem (es. 10.472,51) SI SOR 22

. Ocellar-ocular interspace often more than

0.25 X interocellar space (figs. 11, 20, 25);

vertex generally weakly to sharply angled

centrally (figs. 20, 23, 47, 51) and/or hind

trochantellus greater or equal in length to

its minimum width in dorsal view (e.g., fig.

10a); body often yellow patterned (cf.

fig. 6b); carina lateralis of propodeum often

weaker than the dorsal carinae or even ab-

sent (fig. 16). — Mandible often with dis-

tinet puncturing on ventral flange, fre-

quently in combination with acute gape and

weak internal angles to teeth (fig. 14); radi-

us usually sinuate towards base (fig. 38a);

scutellum rarely carinate when first flagellar

segment is much more than 3.0 X longer

than broad

Ocellar-ocular interspace never greater

than about 0.25 X interocellar space; occip-

10.

il.

ital carına usually squared, dipped or

rounded centrally on vertex (figs. 11, 33,

40) and/or hind trochantellus shorter than

its minimum width in dorsal view

(fig. 10b); body rarely yellow patterned

above; carina lateralis more usually about as

strong as the other dorsal carinae (fig. 43).

— Mandible with at most a few fine punc-

tures on ventral flange, the gape obtuse to

rectangular, internal angles to teeth sharply

defined (fig. 37); radius quite often evenly

curved towards base (fig. 38b); one species

frequently has the scutellum carinate

(fig. 7), and the first flagellar segment well

over 3.0 X longerthan broad .......... 22

. Body without yellow patterning, stigma not.

yellowsat apen me (eee ER 9

Body yellow patterned (cf. fig. 6b) — or

stigma with conspicuous yellow apex... 14

MATE ONS ENE MENSE RS 10

Hemales AR KOTA NOOO SIEDO 11

Head less buccate — i.e. agreeing with at

least two of the following: ocellar-ocular

interspace less than 0.4 X interocellar

space; distance between posterior ocellus

and occipital carina less than 0.8 X diameter

of a posterior ocellus and less than 2.0 x

maximum width of first flagellar segment

(dorsal head characters, figs. 11a, 19, 23);

gena less than 0.45 X width of base of man-

dible (figs. 9, 12, 21). — Flagellum with at

least 51 segments, temple always shorter

daam eye lengde mg, la) sossen 14

Head more buccate — ocellar-ocular inter-

space at least 0.4 X interocellar space; dis-

tance between posterior ocellus and occipi-

tal carina at least 0.8 x diameter of ocellus,

usually not less than 2.0 X maximum width

of first flagellar segment (except when latter

is only around 3.0 X longer than broad)

(figs. 11b, 25, 28); gena at least 0.45 Xx

width of base of mandible (figs. 13, 26, 29).

— Flagellum often with less than 51 seg-

ments, temple about as long or even longer

doen eve (res 25,27, 28) oso 12

Head less buccate — ı.e., ocellar-ocular in-

terspace usually much less than 0.4 X inter-

ocellar space, distance between posterior

ocellus and occipital carina usually less than

0.8 x diameter of a posterior ocellus and up

to c. 1.3 X maximum width of first flagellar

segment (fig. lla); gena at most 0.4 xX

width of mandible base (figs. 9, 12, 21). —

Flagellum with at least 51 segments.

Temple always distinctly shorter than eye

66 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Figs. 8,9. Ophion scutellaris Thomson; 8, propodeum, lateral, al = lateral area, c = hind coxa; 9, head, frontal

view. Fig. 10. Hind trochantelli, a) elongate form, b) short condition. Figs. 11—14. Ophion luteus (Linnaeus);

11, heads, dorsal, (a) and indicating range of variation (b); 12, head, lateral; 13, ditto — showing longer genal

condition; 14, mandible, mandibular gape indicated by arrow, f = mandibular flange.

12.

13:

14.

Brock: Ophion in Britain 67

(Ge, Ue) 14

Head more buccate — ı.e., ocellar-ocular

interspace at least 0.4 X interocellar space,

distance between posterior ocellus and oc-

cipital carina at least 0.8 X ocellar diameter

and greater than or equal to 1.3 X maxi-

mum width of first flagellar segment (ex-

cept when latter is only around 3.0 X lon-

ger than broad) (figs. 27, 28); gena 0.35 to

0.65 X width of base of mandible (figs.

12,29). — Flagellum often with less than 51

segments. Temple up to as long as eye (fig.

27)

Occipital carina rounded, squared or

dipped centrally (sometimes weakly angled)

(figs. 11, 33, 40); hind trochantellus as long

or longer than its minimum width in dorsal

view (fig. 10a); mandible usually with an

acute gape, weak internal angles to the

teeth, the flange distinctly punctate medial-

(e, EE LEA Lit 14

Occipital carina weakly to sharply angled

centrally figs. 19, 20); hind trochantellus of-

ten shorter than its minimum width in dor-

sal view (fig. 10b); mandible with a rectan-

gular or obtuse gape, sharp internal angles

and more or less impunctate flange (fig.

Dn 13

Temple shorter than eye length; flagellum

with at least 51 segments; stigma usually

yellow at apex when flagellum has less than

57 segments; hind trochantellus more

usually as long as its minimum width in

JOIE Ie MIO) nno 16

Temple at least equal to eye length, and/or

flagellum with less than 51 segments; stig-

ma never yellow at apex; hind trochantellus

always shorter than its minimum width in

densalkvrewa(tie lb) Sangen 17

Occipital carina usually rounded, squared

or dipped centrally, sometimes weakly an-

gled (fig. 11, 33, 40); mandibular gape

usually acute, with weak (or no) internal

angles at base of teeth (fig. 14); epistomal

punctures often similar to or of sparser dis-

tribution than those of facial orbits; lateral

carina of propodeum generally absent or

more weakly defined than the dorsal cari-

nae (fig. 16, also 50); mesopleura with

weakly sculptured interspaces between the

punctures. — First flagellar segment often

distinctly greater than 4.0 X longer than

broad (fig. 11); postgena greater than or

equal to length of genal inflection (cf. figs.

12, 13); stigma rarely paler at apex, body

15.

16.

rarely yellow patterned. Extremely vari-

able, especially in genal length, interocellar

space and other head characters (Abun-

dant) luteus L.

Occipital carina weakly to very sharp-an-

gled centrally (figs. 20, 23, see also 47, 51);

mandibular gape obtuse to rectangular, in-

ternal angles of teeth sharp (fig. 37); punc-

tures of epistoma similar to, or of closer dis-

tribution than those of orbits (fig. 21);

lateral carina of propodeum sometimes

complete and strong; mesopleural inter-

spaces more roughly sculptured on average.

Hind trochantellus sometimes much short-

er than minimum width (fig. 10b). — First

flagellar segment less than 3.0 to around 4.0

X longer than broad (sometimes a little lon-

ger) (figs. 19, 20, 23); postgena usually in

range of less than, or equal to length of gen-

al inflection; stigma often yellow at apex

(typically in combination with yellow pat-

tennediDody 1260) e 15

Flagellum with less than 50 segments; dis-

tance between posterior ocellus and occipi-

tal carina around 2.0 or more X maximum

width of first flagellar segment (fig. 18);

third abdominal tergite not more than twice

as broad apically as at base, in lateral view

(fig. 17a); the central flagellar segments

tending to be less elongate on average;

Mesonotum “vittate” (fig. 6b). (Rare, sand

dunes) forticornis Mrly.

Flagellum with at least 51 segments (usually

more); distance between posterior ocellus

and occipital carina much less than 2.0 x

maximum width of first flagellar segment

(figs. 19, 23); third tergite up to 3.0 X wider

at apex than at base (fig. 17b); central fla-

gellar segments tending to be more elongate

in yellow marked examples .......... 16

Body usually richly yellow patterned (fig.

6b), stigma rarely lacking yellow apex when

other markings are absent; first flagellar

segment usually over 3.0 X longer than

broad when there are 57 or more flagellar

segments (fig. 19); sternopleural angle of

epicnemium obtuse to (sometimes) weakly

acute (fig. 22); hind trochantellus Sy

about as long as its minimum width in dor-

sal view (cf. fig. 10a); general microsculp-

ture finer (see fig. 22); hind femur more

slender on average (figs. 30a/b, sometimes

c); scutellum not carinate laterally; propo-

deum sometimes with area basalis, but rare-

ly with area dentipara or closed superome-

68 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Figs. 15— 16. Ophion luteus (Linnaeus); 15, propodeum, dorsal; 16, propodeum, lateral. Fig. 17. (a) Ophion

forticornis Morley, abdominal tergites 1—3, lateral; (b) Ophion obscuratus Fabricius, the same. Fig. 18. Ophion

forticornis Morley, head, dorsal (central flagellar segments inset). Figs. 19—22. Ophion obscuratus Fabricius;

19, head, postero-dorsal view (central flagellar segments inset); 20, head, dorsal; 21, head, frontal view; 22,

mesothorax, slightly rotated ventral view — sternopleural angle of epicnemium arrowed. Fig. 23. Ophion cras-

sicornis sp. nov. head, dorsal (central flagellar segments inset).

Brock: Ophion in Britain 69

Fig. 24. Ophion crassicornis sp. nov., mesothorax, slightly rotated ventral view — sternopleural angle of epicne-

mium arrowed. Fig. 25—27. Ophion longigena Thomson; 25, head, dorsal (male); 26, head, lateral — x, postge-

nal length, y, length of genal inflection; 27, head, dorsal (female). Figs. 28—29. Ophion brevicornis Morley; 28,

head, dorsal (female) (proximal and central, flagellar segments inset), sol, stemmatico-ocular line; 29, head,

lateral (female). Fig. 30. Range of form of hind femur in Ophion species. Figs. 31—32. Ophion perkinsi sp.

nov.; 31, head, dorsal; 32, head, lateral. Fig. 33. Ophion pteridis Kriechbaumer, head, dorsal — Iss = lateral

sulcus of stemmaticum.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

dia (see fig. 1). Polymorphic. (Abundant,

March to January, with peaks in spring and

autumn) obscuratus F.

Body not yellow patterned, stigma lacking

pale apex; first flagellar segment from less

than 3.0, to (sometimes) 4.0 X longer than

broad (fig. 23); flagellum with at least 57

segments; sternopleural angle of epicnemi-

um usually sharply acute angled (fig. 24);

hind trochantellus often distinctly shorter

than its minimum width in dorsal view (ap-

proaching fig. 10b); general microsculpture

rougher (see fig. 24); hind femur thicker on

average (figs. 30c, sometimes b); scutellum

often with lateral carinae (fig. 7); propo-

deum lacking area basalis, but often with

dentiparal carinae and closed area supero-

media. (Uncommon, early to mid-summer)

rin. Ge ee crassicornis sp. nov.

MAES te re 18

ÉÉMAleS EN RC PR ee: 20

18. Temple around 1.3 X eye length, very

strongly expanded behind eyes in dorsal

view; ocellar-ocular interspace over 0.5,

to about 1.0 X interocellar space; distance

between posterior ocellus and occipital ca-

rina 2.5 — 3.0 X maximum thickness of

first flagellar segment (latter about 4.0 X

longer than broad) and 0.8 to 2.0 X diame-

ter of a posterior ocellus (head, dorsal fig.

25); postgena around 4.0, to nearly 6.0 X

length of genal inflection, latter distinctly

less than 0.4 X width of mandible base;

gena 0.8 to 1.0 X base of mandible (head,

lateral fig. 26); eyes strongly divergent be-

neath in frontal view; flagellum with 48 to

56 segments (small sample). (Very rare, per-

haps southern; early summer)

oo e ER eee longigena Ths.

Temple not or scarcely longer than eye, less

expanded behind; ocellar-ocular inter-

space not greater than 0.8 X interocellar

space; distance between posterior ocellus

and occipital carina not greater than 2.0 X

maximum thickness of first flagellar seg-

ment (latter sometimes only around 3.0 X

longer than broad) and at most 1.7 X

diameter of posterior ocellus (heads, dorsal

figs. 28, 31); postgena not more than 2.0 X

length of genal inflection; latter not less

than 0.4 X width of mandible base; gena of-

ten less than 0.8 X latter dimension (heads,

lateral figs. 29, 32); eyes at most slightly di-

vergent; flagellum most often with less than

S0lsesments |: EC AEN TIC)

19. Petiolar membrane approximately opposite

20.

21°

line of spiracles (cf. fig. 43); first flagellar

segment up to 3.4 X longer than broad or

less; temple about equal to eye length and

up to 1.5 X length of first segment of flagel-

lum (fig. 28); scutellum at most weakly ca-

rinate. (Rare) brevicornis Morl.

Petiolar membrane far behind line of spira-

cles (cf. fig. 35); first flagellar segment

around 4.0, to about 5.0 X longer than

broad; temple at most a little longer than

first segment of flagellum (fig. 31); scutel-

lum generally with strong lateral carinae

(cf. fig. 7). (Rare, but widespread in

Britain) perkinsi sp. nov.

Temple fully as long as eye; distance be-

tween posterior ocellus and occipital carina

up to 1.6 X diameter of a posterior ocellus

(fig. 25); genal inflection less than 0.45 X

width of mandible base; postgena at least

around 4.0 X length of genal inflection (fig.

26); abdomen sometimes broadly black

across apex; flagellum with 48 to 56 seg-

ments (small sample!). Rare, perhaps south-

ern longigena Ths.

Temple not quite as long as eye; posterior

ocellus to occipital carina up to 1.3 X ocel-

lar diameter (figs. 28, 31); genal inflection

not less than 0.45 X width of base of man-

dible; postgena less than 2.0 X genal inflec-

tion (fig. 32); abdomen sometimes darkened

ventrally, but not determinately black api-

cally; flagellum usually with less than 50

Sepments se EEE 21

Petiolar membrane approximately in line

with the spiracles (cf. fig. 43); first flagellar

segment up to about 3.4 x longer than

broad; temple slightly shorter than eye and

up to 1.5 X length of first flagellar segment:

distance between posterior ocellus and oc-

cipital carina at most 1.8 X maximum width

of first flagellar segment; central flagellar

segments subquadrate, the antenna unusu-

ally short and thick (head, dorsal, fig. 28);

gena greater than or equal to 0.5 X width of

base of mandible. (Rare)

Petiolar membrane emitted well behind line

of spiracles (cf. fig. 35); first flagellar seg-

ment in region of 4.0 to 5.0 x longer than

broad; temple much shorter than eye and

about same length as first flagellar segment,

or a little longer; distance between posteri-

or ocellus and occipital carina 1.5 to 2.0 X

maximum width of first flagellar segment;

22.

23.

Brock: Ophion in Britain ZA

central flagellar segments elongate, the an-

tenna more slender in appearance (head,

dorsal, fig. 31); gena less than or equal to

0.5 x width of mandible base (fig. 32).

(Rare, but widespread in Britain). .........

ode EE perkinsi sp. nov.

Mandible more elongate on average, usually

with a distinctly punctate flange, often with

an acute gape combined with weak or ab-

sent internal angles to the teeth (fig. 14);

hind trochantellus usually at least as long as

its minimum width in dorsal view (fig. 10a),

sometimes shorter; carina lateralis of pro-

podeum tending to be absent or weaker

than the more dorsal carinae (fig. 16) when

vertex is not angled at the occipital carina

(e.g. figs. 11, 33, 40); body very often yel-

low patterned (fig. 6b) including apex of

stigma; scutellum rarely carinate (excepting

when the first flagellar segment is only

around 3.0 X longer than broad); radius al-

ways sinuate towards base (cf. fig. 38a) 14

Mandible shorter and wider on average, the

flange with at most a few minute punctures

medially, the gape rectangular to obtuse, in-

ternal angles of teeth sharply defined (fig.

37); hind trochantellus always shorter than

its minimum width in dorsal view (fig. 10

b); carina lateralis usually about as strong as

the dorsal propodeal carinae when vertex

not angled centrally on occipital carina (fig.

43); body (including stigmal apex) rarely

with distinctive yellow patterning; scutel-

lum often carinate (fig. 7), in which case

first flagellar segment much more elongate

(e.g. figs. 34, 40); radius quite often not or

scarcely sinuate towards base (fig. 38b) 23

First flagellar segment usually at least 3.4 X

as long as broad (figs. 33, 40); carina latera-

lis usually about as strong as the dorsal pro-

podeal carinae (fig. 43); pleurosternal an-

gles of epicnemium far anterior to sternal

angles, excepting when former are very ob-

tuse angled (figs. 36, 42); substigmal specu-

lum more or less extending beneath the pre-

stigma (fig. 39) or trichiae in this region

much longer than width of adjacent cross-

vein (fig. 34); top edge of mandible basally

narrower than width of a tooth (fig. 45) —

except when the edge is rounded into front

of mandible. — Scutellum often carinate

(fig. 7); radius often not sinuate towards

DASS JO ELA IR eN ete 24

First flagellar segment often less than 3.0 X

longer than broad, rarely more than 3.3 X

245

23.

(tigs. 47, 51); carina lateralis very often ab-

sent or weaker than the dorsal propodeal

carınae (fig. 50); pleurosternal angles of

epicnemium very often nearly in line with

sternal angles, not broadly obtuse, often

sharp (figs. 49, 52); substigmal speculum

not extending beneath prestigma, the tri-

chiae in this region at most marginally lon-

ger than the adjacent crossvein ıs wide (fig.

48); top edge of mandible basally about

same width as a tooth, edge sharply defined

from front of mandible (fig. 46). Scutellum

rarely with trace of carinae; radius always

sinuate towards base (cf. fig. 38a)...... 25

Scutellum often carinate (fig. 7); radius sin-

uate towards base of second abscissa (fig.

38a); membrane of petiolar segment well

behind line of spiracles (fig. 35); number of

flagellar segments rarely less than value of:

1.19 X wing length (mm) + 36.45 (to near-

est whole number); pleurosternal angles of

epicnemium rectilinear to somewhat ob-

tuse, lower sector of epicnemium usually

not more than about 0.5 X upper sector

(fig. 36); lateral sulci of stemmaticum about

as deep, or deeper than the middorsal sulcus

(fig. 33). — Colour more reddish, micros-

culpture coarser; area superomedia usually

open posteriorly and petiolar carinae less

complete (cf. fig. 15); facioclypeal area

more quadrate on average. (Common and

widespread) pteridis Kr.

Scutellum rarely with trace of carinae; radi-

us often evenly curved towards base of sec-

ond abscissa (fig. 38b); membrane ot petio-

lar segment approximately opposite the spi-

racles (fig. 43); number of flagellar

segments rarely more than value ot: 1.13 X

wing length (mm) + 37.5 (nearest whole

number); pleurosternal angles of epicnemi-

um usually rounded and broadly obtuse,

lower sector of pleural epicnemium gener-

ally nearer 0.6 X upper sector (fig. 42);

lateral sulcus of stemmaticum usually

weaker than middorsal sulcus, frequently

obsolete or absent (fig. 40). Colour more

testaceous, frequently with yellow mark-

ings (sometimes as in fig. 6b); microsculp-

ture weak; area superomedia more often

closed, petiolar carinae more complete (cf.

fig. 1); facioclypeal area more transverse on

average. (Common, apparently less so

Northwards) parvulus Kr.

Flagellum with 50-58 segments; nervellus

broken at, or below centre; occipital carına

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

ssp

Figs. 34—37. Ophion pteridis Kriechbaumer; 34, fore wing, trichiation of disco-cubital cell — ps = prestigma,

ssp — substigmal speculum; 35, first abdominal segment, lateral (relative positions of spiracle and sternal mem-

brane indicated); 36, mesothorax, slightly rotated ventral view — broken lines indicating degree of opposition of

epicnemial angulations; 37, mandible, frontal view — gape arrowed, 14 = internal angle of tooth, f = flange.

Fig. 38. Fore wing, stigma and radius; (a) Ophion pteridis Kriechbaumer, (b) Ophion parvulus Kriechbaumer.

Radial sinuation arrowed in (a). Figs. 39—40. Ophion parvulus Kriechbaumer; 39, fore wing, trichiation of dis-

co-cubital cell; 40, head, dorsal (variation in shape of first flagellar segment indicated, inset), mds = mid-dorsal

sulcus. Fig. 41. Hind tibiae, central region, showing spinulation, in (a) Ophion luteus (Linnaeus) and (b) Ophion

parvulus Kriechbaumer.

BROCK: Opbion in Britain 73

51 =

Figs. 42—45. Ophion parvulus Kriechbaumer; 42, mesothorax, slightly rotated ventral view, lengths of lower

and upper sectors of epicnemial carina arrowed; 43, propodeum and first gastral tergite, lateral. Juxta-position

of spiracle and sternal membrane indicated; 44, head, sub-ventral view of left half, showing relative lengths of

genal inflection and postgena; 45, mandibles, lateral view — x = basal width of top edge, y = (maximum) width

of (dorsal) tooth. Fig. 46—50. Ophion mocsaryi Brauns; 46, mandibles, lateral view; 47, head, dorsal; 48, tri-

chiation of disco-cubital cell; 49, mesothorax, rotated ventral view, opposition of epicnemial angles indicated;

50, propodeum and hind coxa, sinistro-posterior view, (a), spiracular shape, (b) (inset), ditto in O. costatus

Ratzeburg. Figs. 51—52. Ophion costatus Ratzeburg; 51, head, dorsal — pss = posterior sulcus of stemmati-

cum; 52, mesothorax, rotated, ventral view.

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

more angled centrally, distance between

posterior ocellus and occipital carina usual-

ly greater (never less) than maximum width

of first flagellar segment and usually around

0.7 X ocellar diameter; ocellar interspace

often wider than maximum width of first

flagellar segment, the latter 2.4 to about 3.5

X longer than broad; posterior sulcus of

stemmaticum moderately impressed (head,

dorsal fig. 47); lower sector of pleural epic-

nemium usually less than 0.6 X length of

upper sector (fig. 49); propodeum usually

nearly completely areolated, but with the

carina lateralis absent to weaker than dorsal

carinae in most specimens; propodeal spira-

cle more ovoid (fig. 50a); last two segments

of maxillary palpus moderately to strongly

unequal in length. (Widespread, sometimes

common) mocsaryi Brauns

— Flagellum with 56-63 segments; nervellus

frequently broken above centre; occipital

carina rounded to weakly angled; posterior

ocellus to occipital carina usually less, sel-

dom greater than width of first flagellar

segment and usually less than 0.7 X ocellar

diameter: ocellar interspace often narrower

than first flagellar segment, latter shorter on

average (often much less than 3.0 X longer

than broad); posterior sulcus of stemmati-

cum very deeply impressed (head, dorsal

fig. 51); lower sector of epicnemium usually

nearer 0.6 X upper sector (fig. 52); propo-

deum with or without a strong carina

lateralis (cf. figs. 1 & 43); propodeal spira-

cle more linear (fig. 50b); last two segments

of maxillary palpus equal, to moderately

unequal in length. (Widespread, seldom

common) costatus Ratzburg

SYSTEMATIC PART

Ophion

Fabricius

Ophion Fabricius, 1798, Entomologica Systematica,

suppl.: 210, 235. Species: Type /chneumon luteus

Linnaeus, designated by Curtis, 1835.

Ophion minutus Kriechbaumer

(figs. 3—5)

Ophion minutus Kriechbaumer, 1879: 105 (lectotype

male, Munich). Schmiedeknecht, 1908. Morley,

1915. Perkins, 19-- (MS). Townes et al., 1965 (in

part). Gauld, 1973, 1976, 1978. Oosterbroek,

1978.

Ophion parvulus Kriechbaumer; Morley, 1915

(misidentification).

Ocellar - ocular interspace 0.2 to 0.5 times in-

terocellar space; posterior ocellus to occipital

carina a little greater to nearly twice longer than

first flagellar segment and a little narrower to

distinctly wider than diameter of a posterior

ocellus; maximum length of temple much short-

er than greatest transverse diameter of com-

pound eye, subequal to much longer than first

flagellar segment; posterior sulci of stemmati-

cum deep, tending to join middorsal sulcus be-

hind the posterior ocelli; lateral sulcus of stem-

maticum generally absent; occipital carina

usually squared or dipped centrally on vertex.

Gena from less than 0.4 to about 0.6 times

width of mandible base; postgena distinctly lon-

ger than genal inflection, latter 0.4 to 0.7 times

width of mandible base. Sculpture of epistoma

not markedly different from that of facial or-

bits; last two segments of maxillary palpus

about equal in length. First flagellar segment 3.5

to over 4.0 times longer than its maximum

width. Range: 40 to 49 segments.

Mesonotum with fine punctures; body fre-

quently yellow patterned, especially with “vit-

tate” mesonotum; pleurosternal angle of epic-

nemium very strongly obtuse; mesopleural

punctures sparse, interspaces not strongly mi-

crosculptured.

Wings with submedian cell somewhat evenly

trichose, nervellus of hind wing distinctly recli-

vous above; substigmal speculum not extending

beneath prestigma; radius somewhat thickened

towards base, wide angled with top edge of

wing. Forewing length: 8—11 mm.

Hind femur rather thick, trochantellus short,

tibial spinules moderate; mid tibial spurs

strongly unequal.

Propodeum varying from near lack of carinae

to a condition where only those anterior to the

posterior transverse carinae are reduced. Carina

lateralis generally in evidence.

Petiolar membrane just behind line of petiolar

spiracles; gaster sometimes darkened at apex.

Variation. — The only really significant vari-

ation lies in the extent of the yellow markings,

which may be very profuse or quite absent.

Gauld (1978) states that no transverse carinae

occur on the propodeum, but these are fre-

quently present — with a maximum devel-

opment as indicated in the figure (fig. 5).

Remarks. — Morley (1915) used the name

parvulus Kr. for non-yellow marked examples

of minutus. Townes et al. (1965) synonymises

these two species, presumably as a result of the

Brock: Ophion in Britain 75

discovery that authors had followed Morley;

Perkins had examined the Kriechbaumer type

and found his parvulus to be quite different

from that of the Morley interpretation (MS). See

also remarks under parvulus Kr. in the present

work.

There should be no difficulty in recognising

the present species from the key.

Biology. — A parasite of Geometrid moths,

the adults of which fly during the winter month

(Erannis species). The parasite is on the wings

during May and June. It is attracted to light, and

is quite frequently collected during the daylight

hours. The chief habitat seems to be deciduous

(and particularly Oak dominated) woodland,

but includes suburban gardens.

Material examined. — 10 2, Wytham, Berks,

28.v.—9.vi.1967, 68, M.V. trap (M. C. Birch); 1

2, Low Moss Plantation, Glasgow, 1.v1.1966; 1

2, Cadder Wilderness, Glasgow, 23.v.1966; 2

d, Wytham, Berks, M.V. trap, 28.5.1968; 1 d,

Pinner, Middlesex, 13.vi.1975, at light; 1 6,

Low Moss Plantation, Glasgow, 19.v.1966 (all

coll. J. P. Brock, unless otherwise stated); 1 d,

Minstead, New Forest, 22.v.1976, M.V. trap (L.

W. Siggs) (BMNH); 1 d, 1 9, Byfleet, SR, 26—

29.v.1949 & 9—11.v.1947 (R. B. Benson)

(BMNH); 2 6,1 ©, Bricket Wood, HT, 31.v.—

5.vi.1951 (R. B. Benson) (BMNH); 1 6, Min-

stead, New Forest, M.V. trap, 22.v.1966 (L. W.

Ses) MB MINE) 27° 8, Whetstone, HT,

12.v.1961 (MV) & 30.v.1959 (P. H. Ward)

(BMNH); 1 d, Blean Wood, Kent, 20.v.1973

(BMNH); 1 6, Abbots Wood, Sussex,

29.5.1932 (H. Donisthorpe) (BMNH); 1 2,

Claygate, Surrey, 4.vi.1952 (D. M. S. Perkins)

(BMNH). Reared material: “H. progemmaria,

New Forest”, 19.vi.1916 (Lyle) (BMNH); “A.

aurantiaria, about 7.v.1902” (Morley coll.)

(BMNH). At the time of writing, I have a few

live cocoons of a small Ophion which can only

be this species, from Erannis defoliaria

(Clerck). These were from host larvae on Oak,

May 1979, Copse Wood, Ruislip, nr. London.

Ophion ventricosus Gravenhorst

(fig. 6a)

Ophion ventricosus Gravenhorst, 1829: 702. Morley,

1915. Gauld, 1978.

Ophion impressus Thunberg, 1822: 262 (pre-occ).

Perkins, 19-- (MS). Gauld, 1973, 1976. Ooster-

broek, 1978.

Ocellar-ocular interspace 0.6 to 1.0 times in-

terocellar space; posterior ocellus to occipital

carina 1.5 to 2.0 times maximum width of first

flagellar segment and up to nearly 2.0 times

diameter of a posterior ocellus; maximum

length of temple up to a little less than greatest

transverse diameter of eye, distinctly longer

than first segment of flagellum. Stemmaticum

with deep posterior sulcus, no lateral sulcus.

Middorsal sulcus deep. Occipital carina round-

ed, squared or dipped on vertex. Gena 0.4 to 0.6

times width of mandible base; postgena 1.3 to

3.0 times length of genal inflection, latter 0.3 to

about 0.6 times width of mandible base. Episto-

mal punctures sparser than those of facial orbits.

Last two segments of maxillary palpus unequal

in length. Dorsal edge of mandible basally wider

than a tooth. First flagellar segment from 2.5 to

around 3.6 times longer than maximum width.

Flagellum with 48 to about 55 segments (mean

of 53).

Mesonotum with large deep puncturation,

deep reddish rather than usual testaceus colour;

pleurosternal angle of epicnemium very obtuse

and weakly defined. Mesopleural punctures

large and deep. Scutellum sometimes with

lateral carinae defined. Thoracic pattern fig. 6a.

Submedian cell of forewing with evenly dis-

tributed trichiae; nervellus slightly reclivous

above; substigmal speculum not extending be-

neath prestigma; radius slightly thickened to-

wards base. Wings somewhat orange-tinted.

Forewing length: 8— 14mm.

Hind femur rather stout, tibial spinules mod-

erate, hind trochantellus short; mid tibial spurs

distinctly unequal.

Propodeum with most of the carinae general-

ly in evidence, including carina lateralis. The

costulae are discrete from the usual transcarina

in this region. The propodeum is usually

marked with black.

Abdomen generally darkened towards apex;

membrane of petiolar segment well behind line

of spiracles.

Variation. — There is no really significant

variation in this species, other than that involv-

ing the greater or lesser extent of the black

markings of the head, thorax, propodeum, ab-

domen and legs.

Remarks. — This species is unlikely to be

mistaken for any other European member ot the

genus.

Biology. — W. Watson has reared this species

of Ophion through the Geometrid Apocheima

pilosaria (D. & S.). Habitats of captured adults

seem usually to be deciduous woodland. It is a

widely distributed species, of rather sporadic

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

occurrence. Records extend to Loch Lomond-

side in southern Scotland, but Ophion ventrico-

sus appears to be absent from many localities

which contain large populations of the above

host.

Material examined. — 1 d, Garth, Loch Lo-

mond, Scotland, 21.vi.1964, M.V. trap (K.

Stewart) (BK); 2 &, 3 ©, Bricket Wood, Herts.,

27.v1.1948, 20.v.1947, 23.v.1939, 4.v1.1951 (R.

B. Benson) (BMNH); 1 d, New Forest,

8.v.1960 (J. F. & R. A. Perkins) (BMNH); 1 dg,

Rammamere Heath, BX, 31.vi.1950 (R. B. Ben-

son) (BMNH); 1 d, Woodwalton Fen, Hunts.,

12—14.v.1958 (R. B. Benson) (BMNH); 1 dg,

Wenlock Edge, Shropshire, 24.v1.1958 (J. F.

Perkins) (BMNH); 1 &, Leamington Spa, WW.,

10.vii1.1954 (M. Bibikoff) (BMNH); 1 d, Ox-

dog Surrey Zi (ip JE Persians)

(BMNH); 1 d, Byfleet, Surrey, 9-11.v.1947 (R.

B. Benson) (BMNH); ex. Apocheima pilosaria

(D. & S.): 1 2,2 à, host coll. v/vi.1975, adult

emerged vi.1976 (W. A. Watson) (Shaw coll.);

further British records can be found in Morley

(1915).

Ophion scutellaris Thomson

(figs. 7—9)

Ophion scutellaris Thomson, 1888: 1192 (lectotype,

Lund). Perkins, 19-- (MS). Gauld, 1973, 1976,

1978. Oosterbroek, 1978.

Ophion longicornis Brauns, 1889. Schmiedeknecht,

1908. Morley, 1915.

Ophion stigmaticus Morley, 1915.

Ocellar-ocular interspace from less than 0.2

(females) to about 0.65 (males) times interocel-

lar space; posterior ocellus to occipital carina

1.0 to 1.6 times width of first flagellar segment

and 0.5 to around 1.0 times diameter of a poste-

rior ocellus. Maximum temple length much

shorter than greatest transverse diameter of eye

and 0.7 to 1.0 times length of first flagellar seg-

ment. Sulci of stemmaticum complete, lateral

sulcus often deeper than middorsal sulcus. Man-

dibular flange subglabrous, gape not acute, in-

ternal angles sharp. Surface of mandible notice-

ably deplanate. Occipital carina generally

rounded or squared, not angled centrally. Gena

around 0.4 (female) to nearly 0.8 (male) times

width of mandible base; postgena 0.6 to 1.8

times genal inflection, latter generally around

0.7 times width of mandible base. Eyes conver-

gent in frontal view; facioclypeal area subqua-

drate to transverse; epistoma tending to be

more sparsely punctured than orbits; last two

segments of maxillary palpus unequal. Anten-

nae unusually elongate; first flagellar segment

3.6 to over 4.0 times longer than broad. 62 to 72

flagellar segments.

Thorax with fine puncturation, weak micro-

sculpture; mesonotum often with darkened

areas; pleurosternal angle of epicnemium sharp,

rectilinear to obtuse, not aligned with sternal

angles. Scutellum often with strong lateral cari-

nae.

Submedian cell of forewing somewhat evenly

trichose (variable); nervellus not reclivous

above; substigmal speculum not clearly pro-

duced beneath prestigma; radius sinuate to-

wards base. Forewing length: 14— 18 mm.

Hind coxa and femur elongate, former small

in relation to lateral area of propodeum. Tro-

chantellus short. Tibial spinules only rudimen-

tary. Mid tibial spurs of only slightly unequal

length (1: 0.8 or more).

Propodeum with carina lateralis, lacking

complete area superomedia and dentiparae (and

sometimes the carina lateralis). Keel between

latter and spiracle often absent.

Petiolar membrane far behind line of spira-

cles.

Variation. — The keel joining the lateral cari-

na to the propodeal spiracle was found to be

present in roughly thirty percent of specimens

and its absence cannot therefore be used as a di-

agnostic feature of the species as indicated by

Gauld (1978). The other characters given by

that author are more reliable and are those

brought forward by Perkins, in his manuscript

key to Ophion (but see obscuratus flagellar vari-

ation!).

Remarks. — Perkins found that specimens

placed under the name scutellaris Ths. in the

Morley collection were in fact referable to pteri-

dis Kr. He also recorded that Morley had used

the name longicornis Brauns for the present spe-

cies. This synonymy was later checked and pub-

lished by Gauld (1973, 1976). Perkins also indi-

cated that Morley’s own O. stigmaticus was a

form of scutellaris Ths. (see Gauld 1973, 1976).

There should be no difficulty in identifying

this species, as it is on the wing well before any

other with which it might be confused. Early

specimens of obscuratus may however be taken

along with it; these will generally be recognised

by their yellow markings or by the shorter fla-

gellum of the latter species. Two of the paratype

specimens of stigmaticus that were redetermined

by Gauld (loc. cit.) as scutellaris are in fact refer-

BROCK: Opbion in Britain 77

able to obscuratus. À further two paratypes

from the Morley stigmaticus material, stated by

Gauld to be luteus (sensu Perkins — Gauld) are

males of parvulus and obscuratus respectively.

Biology. — Hosts are overwintering Noctuid

larvae which reach near-maturity in spring.

Normally, they appear to feed on grasses or

other low plants of open ground (meadows,

commons, heaths, and woodland clearings). The

parasite is often seen flying in sunshine in such

habitats, and is probably common in most suit-

able localities throughout Britain.

Material examined. — 3 2, 8.v.1965,

15.1v.1964, 10.1v.1963, Minstead, New Forest,

M.V. trap (L. W. Siggs) (BMNH); 3 d, 3 9,

same data as above (13.1v.1974—ıv.1975) (BK);

2 2, Freshfield, Lancs., 9.1v.1964 (C. O'Toole)

(BK); 3 2, Wytham Wood, Berks., M.V. trap

(M. C. Birch) (BK); 3 2, Wytham, Berks., M.V.

trap, v.1967/68 (BK); 1 2, Milngavie, Glasgow,

11.v.1966 (common in this locality) (BK); no

data (? Lancs.) (C. O’Toole) (BK); several males

observed at Shabbington Wood/Waterperry

Wood, near Oxford, 1968/69 (BK); 2 4,2 2,

Minstead, New Forest, M.V. trap, 2.11.—

27.1v.1966 (L. W. Siggs) (BMNH); 1 d, Oxen-

bourne Down, Hants., 1.1v.1973, M.V. trap (G.

Else) (BMNH); 1 ?, Dartford Heath, Kent, 1—

Peel son Re ee Ford) BMNEN; 1d i 2,

Glen Brittle, Skye, W. Ross, 20—21.1v.1962 (L.

Blozesch)E. BMNENE 2) 65 VArkley (AS

ASSET Glo wart) (BMNH); 1 95

Kent’s Oak, Romsey, Hants., 6.1v.1969, at light

(C. R. Vardy) (BMNH); 1 6, Wimbledon

Common, Surrey, iv.1958 (J. A. & D. J. Clark)

(BMNH); reared through “T. fimbria” and

wNexamtvosraphas, 1909, 1910 (Lyle)

(BMNH); also Hayling Island, SH., ex Aporo-

phyla nigra (Sperring) (BMNH).

Ophion luteus (Linnaeus)

(figs. 11—16, 41a, graphs 1, 1a, 5)

Ichneumon luteus Linnaeus, 1758 (holotype male,

London; Linn. coll.). Morley, 1915. Perkins, 19--

(MS). Gauld, 1973, 1976, 1978. Oosterbroek,

1978.

Ophion distans Thomson, 1888. Morley, 1915.

Ophion ?slaviceki Kriechbaumer, 1892: 233 (lecto-

type male, Munich). Perkins, 19-- (MS). Gauld,

1976, 1978.

Ophion calcaratus Morley, 1915.

Ocellar-ocular interspace usually in range

0.2—0.6 times interocellar space (sometimes

much shorter); posterior ocellus to occipital

carina 1.3 to 2.0 times maximum width of first

flagellar segment and from 0.6 to over 1.0 times

diameter of posterior ocellus; temple with maxi-

mum length distinctly shorter than greatest

transverse diameter of eye, usually 1.0 to 1.3

times length of first flagellar segment; sulci of

stemmaticum more or less complete, sometimes

with a stemmatico-ocular line to eye; occipital

carina usually rounded or squared, but

sometimes weakly angled centrally. Gena

around 0.4 to as much as 0.9 times width of

mandible base; postgena 1.0 to 1.8 times length

of genal inflection, latter 0.7 to 0.8 times width

of mandible base. Eyes weakly divergent in

frontal view; facioclypeal area moderately to

strongly transverse; mandible elongate, usually

with acute gape, weak to absent internal angles

to the teeth, the flange with distinct medial

punctures. Punctures of epistoma similar to or

of sparser distribution, than those of orbits; last

two segments of maxillary palpus unequal in

length. First segment of flagellum usually

around 4.0 times longer than broad (may be

much shorter, or even more elongate than this);

usual range of flagellar segments, 52—59 (mean

54 in male, 56 in female): see also remarks be-

low.

Thorax with shallow puncturation and weak

microsculpture; mesonotum sometimes with

weak yellow “vittae”; pleurosternal angle of ep-

icnemium usually obtuse, not aligned to sternal

angles. Lower sector of pleural epicnemium up

to about 0.5 times upper sector. Scutellum rare-

ly with trace of carinae.

Submedian cell of forewing sparsely trichose

above, but sometimes evenly trichose. Substig-

mal speculum more or less extending beneath

prestigma; stigma not yellow apically; radius

sinuate towards base. Forewing length: 10—18

mm.

Hind femur moderately elongate, trochantel-

lus as long or longer than minimum width.

Hind tibia with strong spinules. Mid tibial spurs

moderately to very strongly unequal in length.

Carina lateralis absent to weaker than dorsal

propodeal carinae. Superomedia and dentipara

incomplete.

Membrane of petiolar segment near line of

spiracles.

Variation. — An exceedingly variable species,

especially in head characters. There is wide

overlap with the longigena subgroup in more

buccate headed individuals, but these can be dis-

tinguished on characters given in the key. Gen-

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

erally speaking, luteus is characterised by the

form of the mandible, lack of scutellar carinae,

elongate hind trochantellus and non-angulate

vertex. Since all of these characters are subject

to variation and exception, any doubtful speci-

mens should be compared point by point with

the description. Confusion with obscuratus is

possible, since some forms of luteus may be yel-

low marked (and obscuratus not). Overlap with

related species is however, not too frequent and

the key has been constructed to deal with most

variants which will be encountered. Specimens

with unusually small ocellar-ocular interspace

may be confused with parvulus, but these can be

eliminated on the form of the hind trochantel-

lus.

Sexual dimorphism is quite marked in this

species, the greater extremes of variation in

most characters being attained most frequently

by males.

Remarks. — Thomson separated his O. dis-

tans on the basis of characters which are cer-

tainly unreliable and which occur in any combi-

nation within contiguous populations of luteus.

The subsequent history of distans is as follows:

(1) Perkins found that the Linnaean type of

luteus was in fact of the distans form; since he

regarded this as a good species, Perkins used the

first available name, slaviceki Kr. for luteus

auctt. nec L. and placed distans as a synonym of

luteus L. (unpubl. MS).

(2) Gauld (1973, 1976, 1978) published the

Perkins findings, at the same time re-examining

the types. Unfortunately, he did not look fully

into the question of infraspecific variation in

these insects. As a result, the name slaviceki Kr.

has gained currency for the O. luteus of au-

thors.

(3) Following analysis of characters used to

separate luteus and “distans”, together with

study of material named by Perkins in the Brit-

ish Museum collections, I came to the conclu-

sion that luteus and slaviceki were simply

variants of a single species.

(4) I have seen the type of slaviceki Kr., but

am not satisfied that the specimen so designated

agrees with the description given by Kriech-

baumer (1892). It is a typical specimen of luteus,

lacking the unusual features of slaviceki as given

by the describer (loc. cit.).

(5) Oosterbroek (1978) places slaviceki Kr.

sensu Gauld as a synonym of luteus L. He is in

agreement with the present author concerning

the unreliability of characters used to split /4-

teus.

Gauld (1976) discusses the synonymy of

O. calcaratus Morley with slaviceki Kr. —

Gauld.

The identity of the true slaviceki remains an

open question, unless it can be shown that the

lectotype was the specimen from which Kriech-

baumer drew his original description.

Specimens of luteus sensu Perkins in the Na-

tional collection are smaller on average and have

fewer flagellar segments than normal luteus

specimens. For the most part, they also have

earlier dates of capture than the latter. From a

regression curve of wing length on flagellar seg-

ment number (graph 5 and 1a), these individuals

Cal

(ei

(©)

i=)

distans

MAY JUNE JULY AUG, SEPT.

Graph 1. Temporal frequency distribution for

Ophion luteus (L.).

20 1

(©)

[ea]

Fu .

distans

poe

mld

46 48 51 55 60

NUMBER OF FLAGELLAR SEGMENTS

Graph 1a. Flagellar segments frequency distribution

for Ophion luteus (L.).

would seem to be a simple downwards extrapo-

lation of the usual /uteus (sensu lato) trend —

very possibly representing an occasional, sea-

sonal, host-correlated dimorphism in the species

— as observed for the related obscuratus.

The only /uteus sensu stricto material named

Brock: Ophion in Britain 79

by Gauld that I have seen, were the two already

referred to under scutellaris: these were misde-

terminations of two other species.

Biology. — Apart from the rearing of some

luteus sensu Perkins-Gauld through the dune-

land Noctuid Ochropleura (Agrotis sens. auctt.)

praecox (L.), I have been unable to trace any au-

thentic rearing of this species. This is remarka-

ble, since luteus is the commonest species of the

genus, and is found in almost any kind of hab-

itat. Circumstantial evidence would seem to

suggest that the host is some Noctuid larva of

such frequent occurrence that it is seldom

reared by Lepidopterists! From the distans rear-

ings, this could well be an overwintering larva

— but the temporal frequency distribution of

adult luteus is very similar to that of pteridis

(which attacks hosts which overwinter as pu-

pae, not larvae).

Material examined. — 36 6,55 ©, Minstead,

New Forest, M.V. trap, vii/vili/ix.1963—75 (L.

W. Siggs) (BMNH/BK); 5 ®, Whetstone,

23.vii—5.x.1959 (P. H. Ward) (BMNH); 1 9,

Sudbury, 4.1x.1925 (P. Harwood) (BMNH); 1

2, Romsey, Kent, 17—21.1x.1963 (C. R. Vardy)

(BMNH); 1 ©, Beckenham, Kent, 14.v111.1963

(R. I. Vane-Wright) (BMNH); 1 d, Whetstone,

25.vii.1959 (P. H. Ward) (BMNH); 2 ©, Wood-

chester Park, Gloucs., ix.1972 (BK); 3 ®, Soli-

hull, Birmingham, 17—18.v1.1969 (M. R.

Shaw) (BK); 8 d, Solihull, Birmingham, M.V.

trap, 17—24.v11.1969 & 29.v11.1973 (M. R.

Shaw) (BK); 1 d, Cornwall, 4—10.v111.1974 (J.

St. E. Cardew) (BK); 2 4, Sanderstead, Surrey,

22.viii.1950 (R. W. Crosskey) (BMNH); 1 ©,

parvulus 49 mocsaryi 55

luteus-/ \

pteridis 54_/ A

FREQUENCY

British Museum, 12.1x.1925 (J. Waterston)

(BMNH); 1 6, Hayes, Kent, 18.v11.1963 (A.

Banks) (BMNH); 1 6, Colchester, NE,

17.viu.1952 (D. De Havilland) (BMNH); 1 6,

Keston, WK., 11.1x.1949 (A. E. Wood)

(BMNH); 1 ©, Kent's Oak, Romsey, Hants., at

light, 7.x.1972 (C. R. Vardy) (BMNH); 1 9,

Beckenham, Kent, 30.viii.1966 (R. I. Vane-

Wright); 1 ®, Tooting Common, Surrey, at

light, 10.1x.1924 (O. G. Heath) (BMNH); 1 ©,

Botley Wood, Hants., 9.x.1974 (G. R. Else)

(BMNH); 1 ©, nr. Stroud, Glos., 23.1x.1960 (A.

C. Pont) (BMNH); 1 2, Arkley, Hants., M.V.

trap, 22.vili.1954 (Sir L. Wakely) (BMNH); 1

®, Digby, Lincs., 20.1x.1958, at light, 20.1x.1958

(C. R. Vardy) (BMNH); 1 ®, Juniper Hall,

Boxhill, Surrey, 18.1x.1963 (R. G. Adams)

(BMNH); 1 ©, Parlock, 16.vi1.1956 (D. M. S.

Perkins).

Material of luteus-distans in coll. BMNH: 2

g, Elveden, Suffolk, 2.vi.1911 (Nurse); 1 d,

Milton, Cambs., 28.v.1924 (Benson); 1 d, Gair-

loch, Scotland, 9.vi.1958 (Benson); 2 d, 1 9,

Monk’s Soham, 31.v.1909, reared (? host) (Mor-

ley); 1 8, 12.viii.1906, no other data (Morley); 1

®, no data (Stephens coll.); 1 ®, “Clutten”

(Harwood); 4 6,1 2, Hunstanton, 27.v.1918, ?

3.v.1918, 25.v.1918 (Lyle).

Ophion obscuratus Fabricius

(figs. 6b, 17b, 19—22, graphs 3, 4a—b)

Ophion obscuratus Fabricius, 1798: 237 (lectotype

male, Copenhagen). Perkins, 19-- (MS). Gauld,

1973, 1976, 1978. Oosterbroek, 1978.

Ophion obscurus Fabricius, 1804. Schmiedeknecht,

1908. Morley, 1915.

crassicornis 61

costatus …

obscuratus 58

NUMBER OF FLAGELLAR SEGMENTS

Graph 2. Flagellar segments frequency distribution for seven critical Ophion species (relative frequency values).

80 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Ocellar-ocular interspace less than 0.2 to

(sometimes) 0.6 times interocellar space; poste-

rior ocellus to occipital carina 1.0 to 1.7 times

maximum width of flagellum 1, 0.5 to 0.7 times

diameter of posterior ocellus. Temple with max-

imum length always distinctly shorter than

greatest transverse diameter of eye, subequal to

length of first segment of flagellum; stemmati-

cum with complete sulci, lateral sulcus often

weak or erased. Stemmatico-ocular line

sometimes present. Occipital carina weakly to

sharply angled centrally. Gena 0.3 to rather

more than 0.4 times width of mandible base;

postgena less than, or equal to length of genal

inflection, latter 0.7 to 1.0 times width of man-

dible base. Mandible with rectangular to obtuse

gape, sharp internal angles to the teeth, flange

generally with distinct punctures medially.

Epistomal punctures similar to, or closer to-

gether than on orbits. Last two segments of

maxillary palpus unequal in length. Dorsal edge

of mandible as wide as a tooth. First flagellar

segment approximately 3.0 to a little over 4.0

times longer than broad. Usual range of 54—62

flagellar segments (mean of about 58). See re-

marks on variation below.

Mesonotum not strongly punctate, often vit-

tate. Yellow markings often spreading to pleu-

ron, propodeum, abdomen. Pleurosternal angle

of epicnemium often obtuse, rarely acute, usual-

ly somewhat rounded. Not aligned with sternal

angles. Mesopleural punctures moderately

close, with distinctly microsculptured back-

ground. Scutellum without distinct lateral cari-

nae.

Forewing with evenly distributed trichiae on

submedian cell; substigmal speculum more or

less extending beneath prestigma; stigma usual-

ly yellow at apex; radius sinuate towards base.

F.wg.: 10—18 mm.

Hind femur quite slender, trochantellus

around same length as its minimum width in

dorsal view. Tibial spinules quite strongly de-

veloped. Mid tibial spurs moderately unequal in

length, sometimes strongly so.

Propodeum sometimes with area basalis de-

fined, rarely with trace of dentiparal carinae; su-

peromedia incomplete, carina lateralis rather

weak, usually less developed than more dorsal

carinae.

Membrane of sternite of petiolar segment ap-

proximately in line with spiracles. Tergite 3 up

to 3.0 times wider at apex than at base.

Variation and biology. — The unusually wide

FREQU: END-MONTHLY TOTALS

JAN. FEB. MAR. APR.

T T T T T T ass

MAY JUNE JULY AUG.SEPT. OCT. NOV. DEC.

Graph 3. Temporal frequency distribution for

Ophion obscuratus F. O. crassicornis sp. nov. shown

for smaller sample.

spectrum of variation found with regard to

structure and colour in this species points to a

seasonal (and probably at least partially host-

correlated) polymorphism. This is reflected for

example, by the frequency distribution of fla-

gellar segment number (graph 4a), and by the

strongly trimodal frequency distribution for

wing length in a large, quasi-random sample

(graph 4b). The three morphs can be roughly

categorised as follows:

(1) Spring form — wing length 13—15/16

mm, 54—60 flagellar segments. Ocellar-ocular

interspace usually in the range: 0.2 to 0.4 X in-

terocellar space. Distance between posterior

ocellus and occipital carına much greater than

width of fırst flagellar segment; testaceous with

moderately distinct yellow markings. Hosts me-

dium-sized Noctuidae, frequenting more open

habitats and woodland clearings. Flies both in

sunshine and at night, common. Feb. to June.

(2) Autumnal “giants” — wing length

15/16—18 mm, 56—68 flagellar segments; ocel-

lar-ocular interspace usually 0.1 to 0.3 X intero-

cellar space; posterior ocellus to occipital carına

not much greater than width of flg. 1. Colour

tending towards infuscate-testaceous, with very

vivid yellow markings. Hosts unknown. Abun-

dant, particularly so in gardens in both urban

and suburban localities. Most usually attracted

to light, rather than being captured in daylight.

August to February, including the winter

months.

(3) Autumnal “dwarfs” — wing length 10—

13 mm, 51—58 flagellar segments; posterior

ocellus to eye, 0.35—0.6 X interocellar space.

Posterior ocellus to occipital carina 1.25—1.7 X

width of flg. 1; colour reddish-testaceous, most

often with weak, or no yellow patterning (ex-

cepting usually, the stigmal apex). Hosts small

Brock: Ophion in Britain 81

FREQUENCY

49 54 58 63 68

NUMBER OF FLAGELLAR SEGMENTS

Graph 4a. Flagellar segment frequency distribution

for Ophion obscuratus F.

Noctuidae on heathland. Sporadic, but ot

widespread distribution; sometimes common.

August to February.

Winter records for the “autumnal” morphs

do not refer to hibernating individuals, since

both sexes have been taken on the wing during

December and January. A long, staggered emer-

gence period is therefore likely for these “late”

forms. From the available host data, it seems

12) =|

> 6

iS)

[2]

[se]

o + T T T T T T Ar

WING LENGTH (mm)

Graph 4b. Wing length frequency distribution for

Ophion obscuratus F. (Continuous line: 3 morphs

separately; broken line: combining all morphs.)

very likely that host choice is strongly corre-

lated with structural polymorphism in this spe-

cies. The stable spring population attacks its

hosts early in the year, but adults do not emerge

from spring cocoons until the following year.

This has been shown to be the case from my

own experience, independently confirmed by

scutellaris

G |

= __- Obscuratus

jen

(©)

lan

8 = DI T T T T T

42 45 50 55 60 68 72

Statistical significance,

species

forticornis

longigena gp.

parvulus

luteus

pteridis

NUMBER OF FLAGELLAR SEGMENTS

correlation/regression analyses

P levels species P levels

<0.05 crassicornis <0.01

<0.001 obscuratus (a) < 0.001

<0.001 obscuratus (b) N.S.

<0.001 scutellaris N.S.

<0.001 mocsaryi/costatus < 0.001

Graph 5. Regression of flagellar segment number with wing length for the Ophion luteus complex.

82 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

M. R. Shaw (pers. comm.). Several facts do

however, seem to support the view that biologi-

cally distinct species are probably not responsi-

ble:

(a) From a regression analysis of wing length

on number of flagellar segments taken across all

forms of obscuratus (graph 5), it was shown that

these two characters are directly correlated

(probably as a simple growth function) for small

to medium-sized individuals, irrespective of

other characteristics (including times of appear-

ance). Unusually large individuals show no such

relationship, flagellar segment number varying

quite independently of wing length. A similar

analysis was done for most other Ophion spe-

cies, the results showing that the form of direct

linear relationship of the two variables studied is

the usual one taken by a graph of wing length

plotted with flagellar segment number. This was

not the case with scutellaris, however. I inter-

pret this as follows: flagellar segment number

and wing length are directly correlated with

each other due to the fact that both are correla-

tions of a simple growth function. In unusually

large individuals (or in species such as scutellaris

in which there is always an unusually large

number of flagellar segments) there is a relaxing

of the canalizing selection normally acting to

hold genetic or environmentally induced varia-

tion of flagellar segment number within tolera-

ble limits, so that such variation is allowed a

certain degree of expression independently of

growth function relationships. This fits very

well with the widely known fact that greater

variation is observed in multiple-compartmen-

talised structures following a serially homolo-

gous form. Smaller “autumnal giant” obscuratus

forms merge with larger spring forms, in their

agreement with expected regression in the two

characters studied. In contrast, members of the

pteridis-parvulus species pair show very widely

displaced regression lines for the same variables.

Thus, there is at least some indication of poly-

morphism rather than speciation between

morphs of the present species.

(b) Since the dwarf form of obscuratus seems

to be the best defined morph, some further in-

vestigatión is necessary in this area. This morph

also fails to produce large enough forms for any

example to fit into the “ grey area” of the regres-

sion analysis. Since rearing records of this

morph were from Lycophotia varia, and since

the form appears always to be collected in local-

ities frequented by this moth, it seemed appro-

priate to test whether or not females collected

from heaths would show any preference be-

tween hosts offered in captivity. Two females of

this form were taken at Chobham Common in

October, 1979 and offered larvae of L. varia,

M. impura (certainly one host of the spring

brood of ‚obscuratus) — together with two other

“remote” Noctuids: Anarta myrtilli and Apa-

mea sp.; a Tenthredinid larva was also offered

as a “control”. The result was that oviposition

was attempted many times in all species of Noc-

tuid larvae, apparently without preference. The

sawfly larvae were completely ignored. This

does not prove that such non-preference neces-

sarily happens in the wild state, nor that devel-

opment of parasites can take place in all at-

tacked hosts. Of rather more significance from

the point of view of interpreting variation in the

parasite species, is the fact that, (a) parasite

adults showed no apparent preference for larvae

at different stages in their development, at-

tempting oviposition in third to final instar

hosts of several species; (b) known hosts of

O. obscuratus are peculiar for the fact that they

themselves possess a staggered growth period.

Both small and large larvae of L. varia were

found simultaneously in the autumn, while

M. impura larvae representing widely different

developmental stages can be found in spring.

Thus, it seems highly likely that the apparently

host-correlated polymorphism of the parasite is

related to this characteristic of the host species.

J. F. Perkins (MS) thought it possible that the

dwarf form at least could be specifically distinct,

while Gauld (1978) does not take this morph at

all into account in his key to the British Ophion

species. This form also appears to have some

points in common with the O. subarcticus of

Hellen, also kevoensis (see Jussila, 1976).

Material examined. — 13 6, 9 2, Minstead,

New Forest, 1963—75 (L. W. Siggs) (BMNH)

(BK); 7 d, Bingley, St. Ives — data incomplete

(J. Wood) (MU); 5 d, Lichfield — data incom-

plete (Carr) (MU); 1 2, Formby Pt, Lanes.,

16.x1.1950, ex Triphaena comes, emerged

4.11.1951 (K. C. Greenwood) (MU); 1 d, Dela-

mere, Cheshire, 19.x.1924 (H. Britten) (MU); 1

d, Skirwith, Cumbs., 19.ix.1926 (H. Britten)

(MU); 1 @, Chat Moss, ances, 8541933050

Britten) (MU); 1 4, Witherslack, 16.v.1925, no

other data (MU); 1 6, Longdendale, Cheshire,

16.x.1932 (H. Britten) (MU); 1 &, Hale, Ches-

hire, 21.v.1922 (H. R. P. Collett) (MU); 1 d, 2

2, Staley, Cheshire, 22.x.1939 (H. Britten)

(MU); 2 d, Wytham, Berks., M.V. trap (M. C.

BROCK: Opbion in Britain 83

Birch) (BK); 1 &, Rutland, 2.v.1974 (J. St. E.

Cardew) (BK); 1 6, Waterperry Common, nr.

Oxford, 9.v.1968 (BK); 1 6, Wytham, Berks.,

M.V. trap, 2.v1.1969 (BK); 1 d, Low Moss

Plantation, Glasgow, 30.1v.1966 (BK); 1 ?, Ox-

ford, 22.x.1968 (R. B. Angus) (BK); 1 9, Ox-

ford, 25.x.1967 (I. Lansbury) (BK); 1 ©, Ruislip

Common, nr. London, ex Mythimna impura

larvae, vi.1975, emerged 11.1976 (BK); 7 d, 5

©, Whetstone, Herts., 27.1v.—10.v.1959, 7—

12.x.59, 5.v.60 (P. H. Ward) (BMNH); 1 &,

. Esher, Surrey, x.1945 (BMNH); 1 d, Worcester

Park, Surrey, at light, 28.11.1960 (R. D. Eady)

(BMINED MS 1 ® Beckenham, Kent,

19.x.1966 (R. I. Vane-Wright) (BMNH); 1 d,

Sinam Common, Hants., 18.11.1972 (coccoon),

Adult 191972 (G. R: Else) (BMNE) 1. 3;

Arkley, HT., 23.11.1954 (T. G. Howarth)

(BMNH); 1 ?, Weybridge, Surrey, v.1951 (E.

Taylor) (BMNH); 1 2, Banstead, Surrey, at

light, 27.3.1951 (S. Wakely) (BMNH); 1 9,

Hurt Wood, Surrey, 12.v.1954 (R. B. Pike)

(BMNH); 1 ®, Causfield, Fareham, Hants.,

v.1971, M.V. trap (BMNH); 1 2, Aviemore,

Inverness-shire, 2—4.v1.1934 (R. B. & J. E.

Benson); 1 2, Bricket Wood, Herts., 23.v.1959

(R. B. Benson) (BMNH); 1 d, Digby, Lincs., at

light, 20.1x.1958 (C. R. Vardy) (BMNH); 1 2,

Ivinghoe, BX., 13.v.1954 (R. B. Benson)

(BMNH); 1 d, Earl’s Court, London, 23.1.1952

(Lang) (BMNH); 1 ®, 1 6, Chelsfield,

20.x.1950 (A. M. Swain) (BMNH); 1 g, Parks

Fence, Oxford, 30.x.1926 (BMNH); 3 6,9 ©,

New Forest, SH., “ex strigula” (L. porphyrea),

DR 27451811909 Lyle) it (BMNED); 2. 2,

Ainsdale, Lancs., ex Agrochola or Xanthia sp.,

host larvae coll. 5.vi.1975, parasite larvae

emerged from host prepupa, vi.1975, adults

emerged 2.iv. & 6.1v.1976 (M. R. Shaw coll); 1

2, Lyndhurst, New Forest, SH., ex Noctua in-

terjecta, host larva 19.v.1975, parasite larva

from host prepupa, c.16.vi.1975, adult emerged

11.1v.1976 (M. R. Shaw coll.); 5 specimens ex

Leucanıa impura, 10—12.1v.1907, “Brit.Is.” (P.

Harwood) (BMNH); reared from Amathes aga-

thina, Poole, Dorset, 24.11.—5.1v.1914 (Lyle)

(BMNH); from “7. fimbria”, New Forest,

Hants., 24.11.1918, ZII ıv.1903,

16.11.1915 (Lyle) (BMNH); from “N. neglec-

ta”, 14.1v.1904 (Lyle) (BMNH); from N. baja,

New Forest, Hants., 9.11.1913 (Lyle) (BMNH);

from N. augur, Shibden, Hx., 13.1v.1923 (Lyle)

(BMNH). In addition, six males and two fe-

males were taken at Chobham Common in Sur-

rey, 10—20.x.1979. Several others were ob-

served flying in sunshine at the same time; these

were all “autumnal dwarfs”.

Ophion forticornis Morley

(figs. 17a, 18)

Ophion forticornis Morley, 1915: 270-271 (lecto-

type, London). Perkins, 19-- (MS). Gauld, 1973,

1976, 1978. Oosterbroek, 1978.

Ophion? baueri Habermehl, 1930: 114.

Ocellar-ocular interspace less than 0.4 (fe-

male) to over 0.6 (male) times interocellar space;

posterior ocellus to occipital carina around

twice maxımum width of first flagellar segment,

0.8 times to about equal to diameter of posterior

ocellus; temple distinctly shorter than greatest

transverse diameter of eye and subequal to

length of first flagellar segment; stemmaticum

with mid-dorsal sulcus stronger than lateral sul-

cus; stemmatico-ocular line indistinct. Occipital

carina sharply angled centrally. Gena from 0.35

(female) to nearly 0.8 (male) times width of

mandible base; postgena a little less to a little

greater than length of genal inflection. Latter

over two thirds width of mandible base. Facio-

clypeal area more or less transverse; mandible as

obscuratus; epistomal sculpture also as obscura-

tus; last two segments of maxillary palpus un-

equal; first flagellar segment 4.0—4.5 times lon-

ger than broad, 43 to 49 flagellar segments

(small sample). Antenna appearing rather short

and stout, with subquadrate central segments.

Mesonotum weakly to moderately strongly

punctate; colour similar to obscuratus, but no

non-yellow marked specimens seen. Pleuro-

sternal angle of epicnemium subrectangular,

more nearly aligned to sternal angles than in

obscuratus. Mesopleural punctures about as far

apart as their diameter. Scutellum not carinate.

Wing characters as obscuratus, including yel-

low stigmal apex. Forewing length: 1014 mm.

Hind femur moderately slender, trochantellus

about as long as wide, tibial spinules weak. Mid

tibial spurs moderately to only slightly unequal

in length.

Propodeum with dorsal carinae more strong-

ly raised than normal, dentiparal and superome-

dial carinae tending to be more nearly complete

than in obscuratus.

Membrane of first sternite near line of spira-

cles. Third abdominal tergite not more than

twice as broad at apex than at base, in lateral

view — abdomen thus appearing more slender

than in obscuratus (more so in Dutch than in

British material however).

84 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Variation. — With the exception of the num-

ber of flagellar segments, the characters given

by Gauld (1973, 1978), after Perkins (MS), lie

within the range of variation found in O. obscu-

ratus. In fact, the shape of the flagellar segments

and mesonotal puncturation are unstable char-

acters in both species. There is also overlap in

the development of the propodeal carinae; the

latter character was indicated by Oosterbroek

(1978) as an additional feature of forticornis. I

have also discovered overlap of this kind with

regard to the additional characters which I have

used to attempt clear distinction between obscu-

ratus and the present species.

Remarks. — Oosterbroek (1978) is probably

correct in placing bauer: Habermehl as a syno-

nym ot forticornis Morley, although neither he

nor I has been able to locate the type of baueri.

The arguments for and against retention of

forticornis as a distinct species are as follows:

(1) Overlapping character states appear not

to occur together (in combination) in individual

variants of either obscuratus or forticornis.

(2) The range of variation (with particular

reference to the number of flagellar segments) in

a combined obscuratus — forticornis would be

very significantly greater than one would expect

within a single Ophion species, from study of

variational range in other species of the genus.

Against:

(3) There is no absolute structural gap be-

tween obscuratus and forticornis, other than that

exhibited by the number of segments in the fla-

gellum. This latter feature could be due to small

sample of forticornis available.

(4) O. forticornis could conceivably be a

duneland ecotype of obscuratus. In support of

this would be the somewhat parallel variants of

crassicornis and luteus which I have seen from

similar localities; on the other hand, typical

obscuratus forms can also be taken amongst

sand dunes.

Biology. — I believe there is a specimen in

the Hope Department of Entomology, Oxford,

reared through Ochropleura (Agrotis auctt.)

praecox (L.). The species flies in early summer,

frequenting sand dunes (also the habitat of the

host record species cited above). Rare.

Material examined. — Morley’s original

material came from Felixstowe, Suffolk, on sand

dunes. — Holotype ®, 6.v.1897, paratype 9,

7.v.1900 (BMNH); I have seen one other Brit-

ish specimen, in the Hope Department Collec-

tions (Dale), reared from “Agrotis praecox” (see

above).

The following material from Holland was

sent by P. Oosterbroek, whom I contacted at

the helpful suggestion of I. D. Gauld: 1 2, Ven-

lo, 1877 (Oudemans coll.); 1 &, Venlo (Oude-

mans); 1 d, Putten, 14.v.1913; 1 &, Hilversum,

11.v.1893 (Oudemans); 1 ©, Assel, 5.6.1970 (J.

B. Wolschrijn).

Ophion crassicornis spec. nov.

(figs. 23, 24, graph 3)

Ophion “species W”; Perkins (MS).

Ophion “species X”; Gauld, 1978 (in part).

Ocellar-ocular interspace 0.2 to 0.5 times in-

terocellar space; posterior ocellus to occipital

carina 1.0 to 1.4 times maximum width of first

flagellar segment, 0.65 to 0.8 times diameter of

posterior ocellus. Maximum length of temple

distinctly shorter than greatest transverse

diameter of eye, usually noticeably longer than

first segment of flagellum; stemmaticum with

usual sulci, lateral sulcus tending to be weaker

than middorsal sulcus. Stemmatico-ocular line

sometimes present, may be represented by zone

of rough sculpture. Occipital carina usually

sharply angled centrally. Gena 0.3 to 0.4 times

width of mandible base; postgena 0.65 to 1.3

times length of genal inflection, latter 0.6 to 1.0

times width of mandible base. Eyes somewhat

convergent towards mouth in frontal view; fa-

cioclypeal area subquadrate; mandible with rec-

tangular to obtuse gape, sharp internal angles to

teeth, flange generally with distinct medial

puncturation. Epistoma and orbits both closely

punctate; last two segments of maxillary palpus

unequal. First flagellar segment from less than

3.0 x longer than broad, to nearly 4.0 times

(usually nearer lower end of ratio) usual range

of 59 to 62 flagellar segments (mean around 60):

exceptionally with as few as 57.

Mesonotum with fine puncturation; colour

tending towards darker testaceous, without yel-

low markings. Pleurosternal angle of epicnemi-

um generally sharply acute angled, produced

backwards to nearly lie in line with sternal an-

gles. Mesopleura closely punctured with dis-

tinct microsculpture between them. Scutellum

usually carinate.

Forewing evenly trichose over submedian

cell; substigmal speculum not clearly extending

beneath prestigma. Forewing 12—18 mm.

Hind femur relatively stout to moderately

Brock: Ophion in Britain 85

slender, the trochantellus much shorter than, to

about equal in length to its minimum width in

dorsal view; tibial spinules weaker than in /u-

teus; mid tibial spurs moderately unequal in

length.

Propodeum lacking area basalis, but often

with closed area superomedia and dentiparal

carinae indicated; carina lateralis usually weak.

Membrane subopposite spiracles of petiole.

Abdomen often darkened ventrally.

Variation. — I have examined some unusual

specimens of this species from northern locali-

ties. These differ from typical crassicornis as fol-

lows: temple shorter on average, first flagellar

segment rather more elongate, nervellus broken

at or below centre (tending to be broken at or

above centre in typical specimens); noticeably

smaller.

Although I have little doubt that these repre-

sent a form of crassicornis, I exclude them from

designation as paratype material. It is possible

(though uncertain as yet) that these specimens

typify a form found in more open habitats, such

as sand dunes or moorland. Again, they may

simply represent a trend in northern popula-

tions generally.

Remarks. — This species was discovered by

Perkins, who also suggested the name proposed

for it. In his manuscript, Perkins placed crassi-

cornis close to longigena. It is the same as “sp.

X” of Gauld (1978) and “sp. W.” of Perkins

(MS). Gauld (loc. cit.) follows Perkins in placing

the present species near longigena, but segre-

gates within the /uteus-longigena section will

not separate satisfactorily on the characters

given in Gauld’s key. In particular, the trichia-

tion of the submedian cell (as used by Perkins

and Gauld) is unstable (cf. luteus) and the hind

trochantellus of crassicornis is unusually variable

(sometimes agreeing with /uteus, alternately

with longigena, following Gauld’s diagnosis).

While admitting that certain similarities do

exist between the present species and members

of the longigena group, I find that there is more

agreement with obscuratus: some non-yellow

marked forms of the latter species require care-

ful study if confusion with crassicornis is to be

avoided. This however applies mainly to the

dwarfed autumnal morph of obscuratus, which

can generally be separated by the smaller num-

ber of flagellar segments.

Biology. — I have seen one specimen reput-

edly reared through the Noctuid Aporophyla

nigra (Haw.). The normal flight period of the

parasite seems rather later than would be ex-

pected if this was the usual host. Ophion crassi-

cornis seems to be increasing in numbers at

about the same time as the mid-summer decline

in obscuratus (see graph 3).

Gauld (1978) found his “species X” to be a

common one, while my own experience sug-

gests that crassicornis is of rather sporadic oc-

currence. Most probably, Gauld’s view was at

least partly based on the large number of speci-

mens placed under the Perkins manuscript name

in the national collection. A closer examination

of these during 1979 has shown that four differ-

ent species have been confused under that title.

These included the “true” crassicornis, the

“unknown” males of brevicornis Morl., and a

Scandinavian species related to the latter. Al-

though it now seems that Ophion crassicornis is

rather a rare species, the existing records do

suggest quite a wide distribution. Gauld also

sent a specimen of “sp. X” to J. Aubert, who in-

tended describing and naming the species on the

basis of this and one other specimen. Through

the very helpful cooperation of these workers, I

was allowed to examine these insects, with a

view to including them as paratypes of crassicor-

nis. Both however, were examples of Ophion

brevicornis Morley, thus apparently confirming

> cc

the aggregate nature of Gauld’s “sp. X”.

Material examined. — Holotype d, Badbury

Rings, Dorset, 4—6.v11.1955 (I. H. H. Yarrow)

(BMNH); Paratypes. Minstead/Linford, New

Forest, Hants., M.V. trap, 2.v1.1975, 30.v1.75,

19—25.v1.75, 6.vu.75, 7—13.v11.755 (L. W.

Siggs) (BK); 6 6, 5 2, Minstead, New Forest,

M.V. trap, 6.v1.1963, 3.v1.1965, 6.v1.1964, 31.v.

& 5.v1.1963, 3.v1.1964, 7.vi.1964, 3.v.1964,

13.v1.1965, 17.v1.1963, 14.v11.1963 (L. W. Siggs)

(BMNH); 2 dg, Gritnam, New Forest, Hants.,

M.V. trap, 4.v1.1976, 28.v.1976 (BK, private

coll.); reared: Hayling Island, SH., 13.vi.1961,

ex Aporophyla nigra (A. H. Sperring) (BMNH).

Notre paratypesa) wlio Sicile MSc pesta!

17.iv.1965, 450 m (K. M. Guichard) (BMNH);

5 Oy Spain iv ONIN Beal (ERS MEE

Cardew) (BK); 1 6, Bolton Perry (VC64),

23.v.1943 (W. G. Blamely) (MU); 1 d, Stanton

Sand Pits (VC61), 14.vi.1943 (W. D. Hincks)

(MU); 1 à, Leeds Lime Hills (VC64), 28.v.1944

(J. Wood) (MU); 1 6, Keighley Dist. (VC63),

5.vi.1943 (J. Wood) (MU); 1 6, Branley Moor,

Cheshire, 13.v1.1926 (A. K. Lawson) (MU).

86 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Ophion longigena Thomson

(figs. 25—27)

Ophion longigena Thomson, 1888: 92 (lectotype,

Lund). Morley, 1915. Gauld, 1978. Oosterbroek,

1978.

Ophion costatus Ratzeburg; Brauns, 1889 (? misiden-

ufication). Perkins, 19-- (MS) (in part). Gauld,

1973, 1976.

Ocellar-ocular interspace around 0.5 (female)

to 1.0 times interocellar space (male); posterior

ocellus to occipital carina 1.8 to 3.0 times maxi-

mum thickness of first flagellar segment, 0.8 to

2.0 times diameter of posterior ocellus; temple

with maximum length just greater than, to 1.3

times greatest transverse diameter of eye and

around 1.0 to 1.5 times length of first flagellar

segment. Stemmaticum with complete sulci,

stemmatico-ocular line often represented. Oc-

cipital carina squared or rounded centrally.

Gena 0.4 to 1.0 times width of mandible base;

postgena around 4.0, to nearly 6.0 times length

of genal inflection, latter 0.3 to 0.4 times width

of mandible base. Facioclypeal area transverse;

mandible with rectangular to obtuse gape, dis-

tinct internal angles to the teeth, the flange in-

distinctly punctured medially. Epistomal punc-

tures tending to be closer than those of orbits;

last two segments of maxillary palpus unequal.

First flagellar segment distinctly over 3.0, to

around 4.0 times longer than broad. Flagellum

with 48 to 56 segments in sample studied.

Thorax with moderate puncturation; meso-

notum sometimes with trace of yellow “vittae”;

abdomen sometimes black apically (female).

Pleurosternal angle of epicnemium more or less

rectangular, nearly aligned with sternal angles;

lower sector of pleural epicnemium 0.5 to 0.6

times upper; scutellum with distinct lateral cari-

nae.

Submedian cell of forewing with evenly dis-

tributed setae; substigmal speculum not extend-

ing beneath prestigma. Forewing 12—15 mm.

Hind femur relatively stout, trochantellus

shorter than its minimum width in dorsal view.

Tibial spinules quite strong. Mid tibial spurs of

moderately unequal length.

Propodeum tending towards complete areola-

tion.

Position of membrane of petiolar segment

somewhat variable.

Variation. — It is difficult to assess fully the

range in variation of this species, due to the

small amount of material available for study.

Sexual dimorphism seems considerable for head

characters, the lower limits for dimensions be-

longing more to the female than the male end of

the range observed. The female may have the

apical abdominal tergites determinately black.

Characters quoted by Gauld (1978) relating to

venational differences between longigena and

brevicornis are unreliable in practice. I am also

not satisfied that Gauld’s use of the form of the

front tarsus is not subject to overlap (in any

case, a wide overlap does exist with another

species not recognised by Gauld).

Remarks. — The name costatus Ratzeburg

has been used for this species (Perkins, MS;

Gauld, 1973, 1976) and was suggested as a pos-

sible synonym by Morley (1915). Interpretation

of the name costatus Ratzeburg is discussed

elsewhere in the present study.

Biology. — The few available rearing records

for this species are from Cucullia (Noctuidae).

The parasites are on the wing in early summer,

during which time the host larvae are presum-

ably in early instar. The species is excessively

rare in occurrence, perhaps of southern distri-

bution only in Britain.

Material examined. — 1 d, Spurn (VC61),

14.v11.1952 (W. D. Hincks) (MU); 1 6, Priors

Marston, Warwicks. 26.vi.1943 (D. Allen)

(MU); 1 9, Isle of Wight, Freshwater, ? 111.1971

(MU); d, no locality data, “? 11.vi.1944, ex C.

roule of 1942” (Sir L. Wakely) (BMNH);

2 3, “Colchester, Harwood” (BMNH): 6, ex

Harwood coll., no data (BMNH): d, Wood-

walton Fen, Hunts, vii.1923 (P. Harwood)

(BMNH); dg, Corfe Castle, Dorset, 4.vii.1901,

ex C. chamonillae (Bankes coll.) (BMNH); ®,

“S wold, 20.vii.01” (C. Morley) (BMNH).

Ophion brevicornis Morley

(figs. 28, 29)

Ophion brevicornis Morley, 1915: 274 (holotype,

London). Gauld, 1976, 1978 (in part). Ooster-

broek, 1978 (not examined).

Ophion costatus Ratzeburg; Brauns, Perkins, 19--

(MS) (in part).

Ophion “species X” Gauld, 1978 (in part).

Ocellar-ocular interspace 0.4 (female) to

about 0.6 (male) times interocellar space; poste-

rior ocellus to occipital carina 1.3 to 1.6 times

maximum width of first flagellar segment; dis-

tinctly less than, to 1.4 times diameter of poste-

rior ocellus. Maximum length of temple just

shorter than greatest transverse diameter of eye

(female) — about equal in male; temple 1.25—

1.5 times length of first flagellar segment. Sulci

Brock: Opbion in Britain 87

of stemmaticum — posteriors very deep and

broad, laterals weak to moderate; stemmatico-

occular line represented by zone of rough sculp-

ture over a groove like depression. Occipital

carina angled centrally (may be triangulate).

Gena 0.5 to 0.8 times width of mandible base;

postgena from shorter than (female) to nearly

twice length of genal inflection (male). Latter

0.6 to 0.7 times width of mandible base. Facio-

clypeal area strongly transverse; mandible rec-

tangular to obtuse in gape, internal angles sharp,

flange with a few minute punctures only. Epis-

toma and orbits with similar puncturation; last

two segments of maxillary palpus strongly un-

equal. First flagellar segment distinctly less than

3.5 times longer than maximum width. Flagel-

lum with 45 to 50 segments (small sample!).

Thorax rather closely punctured, with vari-

able microsculpture; rather dark testaceous in

colour; position of epicnemial angles rather

variable, lower sector of pleural epicnemium 0.5

to more than 0.6 times upper. Scutellum not, or

weakly carinate laterally.

Submedian cell of forewing with trichia even-

ly distributed; substigmal speculum not extend-

ing beneath prestigma; radius weakly, or not

sinuate towards base. Fore wing: 10.5—13 mm

in length.

Hind femur short and stout, trochantellus

shorter than its minimum width in dorsal view;

tibial spinules weak or moderate; mid tibial

spurs of moderately unequal length.

Propodeum tending towards complete areola-

tion, but lateral carina may be absent and super-

omedia incomplete.

Membrane of petiolar segment approximately

opposite line of spiracles.

Variation. — Without a larger material, it is

difficult to assess infraspecific variation.

Remarks. — The male paratype referred to

by Gauld (1976) is a specimen of O. perkinsi.

J. F. Perkins placed brevicornis as a probable

synonym of longigena Ths. (MS, and in BMNH

collection), but this has been rejected by Gauld

(1976, 1978). See also remarks under longigena.

Oosterbroek (1978) places the present species

close to parvulus Kr., but has clearly not had

the opportunity to compare other longigena

group material. From that author’s observa-

tions, brevicornis would appear to be commoner

in Holland than it is in Britain.

Biology. — Oosterbroek (loc. cit.) cites Cos-

mia trapezina (Linnaeus) as a host. From the

few British records of capture, a Noctuid host

maturing in early summer would seem to be a

likely target. If Cosmia is the usual host, then

this may be a good biological distinction from

longigena Ths.

Material examined. — 2 ©, Pinner, Middle-

sex, garden M.V. trap, vi.1977 (BK, private

coll.); 1 36, Wenlock Edge, Shropshire,

24.v1.1958 (J. F. Perkins) (BMNH); 3 ©, Col-

chester, at light, 1—3.vu.1936 (C. B. Williams)

(BMNH); 2, no data (Harwood) (BMNH); d,

2, Höör District, Sweden, 17—21.vi.1938 (J. F.

Perkins) (BMNH); ©, Ideford, “Ens 32

21.vi.1942 (J. F. Perkins) (BMNH); 2 2, no

data (BMNH); ®, Darenth, Kent (coll. Cham-

pion) (BMNH); 2 ®, Minstead, New Forest,

Hants, 21—31.vu.1963 (L. W. Siggs) (BMNH).

The Morley type female was taken at Bentley

Woods, Suffolk, 23.vi.1899, by E. Platten (see

Gauld, 1976) (BMNH). Just prior to comple-

tion of this work, J. Aubert sent me two addi-

tional specimens: 1 d, Ideford, 7.v1.1942 (J. F.

Perkins); 1 2, Maujolia Neucliatie, 22.vi.1947

(J. Aubert) (see O. crassicornis).

Ophion perkinsi spec. nov.

(figs. 31, 32)

Ophion brevicornis Morley, 1915 (in part). Gauld,

1976 (in part).

Ophion “species X” Perkins, 19-- (MS).

Ocellar-ocular interspace 0.4 to 0.8 times in-

terocellar space; posterior ocellus to occipital

carina 1.5 to 2.0 times maximum width of first

flagellar segment, just less than, to about 1.25

times the diameter of a posterior ocellus; maxi-

mum length of temple less than greatest trans-

verse diameter of eye; same dimension a little

less to a little more than length of first segment

of flagellum. Lateral sulci of stemmaticum weak

to absent, stemmatico-ocular line weak to quite

deeply impressed. Occipital carina distinctly an-

gled centrally. Gena 0.35—0.5 (female), up to

1.0 times width of mandible base (male). Post-

gena 0.8 to nearly 2.0 times length of genal int-

lection, latter less than 0.5 to 0.65 times width

of mandible base. Facioclypeal area distinctly

transverse; mandible with rectangular to obtuse

gape, distinct internal angles to teeth, flange

with indistinct medial punctures only. Punc-

tures of epistoma and orbits of variable distribu-

tion; last two segments of maxillary palpus dis-

tinctly unequal; first segment of flagellum

around 4.0, to about 5.0 times longer than maxi-

mum width. Flagellum with 45 to 50 segments,

the central ones elongate.

88 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Thorax with close puncturation, with notice-

able microsculpture on pleura. Epicnemial an-

gles not opposed, the sterno-pleural angle sharp

and more or less rectangular. Lower sector of

pleural epicnemium tending to be around 0.5

times upper sector. Scutellum with prominent

lateral carinae.

Submedian cell of forewing with trichiae

tending to be more closely spaced on lower

third of cell than above this area; substigmal

speculum not clearly extending beneath prestig-

ma; radius sinuate towards base. Forewing

length: 10.5—12.5 mm.

Hind femur not slender, trochantellus shorter

than its minimum width in dorsal view. Tibial

spinules not strong. Mid tibial spurs distinctly

unequal in length.

Propodeum with carina lateralis and area su-

peromedia tending to be complete, external

dentiparal carinae obsolete.

Membrane of petiolar segment far behind line

of the spiracles. Abdomen tending to be dark-

ened ventrally.

Variation. — It is likely that when more

material is seen, some extension may be re-

quired to the range of variation given here. In

the key, I have attempted to stress characters

which have been found stable within reasonable

limits in other species groups where more

material has been available for analysis. For this

reason, some differences between species in the

longigena group which may be quite reliable,

have been omitted from the key. A darkening of

the abdomen (both ventrally and apically) was

the only really conspicuous component of varia-

tion found in some specimens of perkinsi.

Remarks. — This species was discovered by

Perkins and is the “sp. X” of his manuscript key

to Ophion. He placed his “sp. X” with slaviceki

(see luteus L.) and distinguished the species on

characters which in reality, mostly lie within the

variation of luteus L. For several years, I had

two specimens of perkinsi placed as indet. nr.

brevicornis Morley. Eventually, I was able to

align these with material placed separately by

Perkins in the BM collections; although these

specimens were not labelled as such, it seemed

that they must be the material upon which Per-

kins had based his “sp. X”. Since that date, these

specimens have been amalgamated with “slavi-

ceki” in the national collection; they were re-

cently reinstated by the present author (1978).

Ophion perkinsi is not represented in the key

given by Gauld (1978), nor by Oosterbroek

(1978) in his study of the Dutch species. The

scutellar carinae and short hind trochantellus

width generally distinguish it from forms of lu-

teus with more buccate head; the antennal

structure will separate perkinsi from brevicornis,

to which latter species it is obviously closely re-

lated.

Biology. — Nothing is known regarding host

preferences of the present species. Habitats fre-

quented by it are as diverse as deciduous wood-

land and fen country. Its British distribution ex-

tends into Scotland, but it is rarely collected.

The species flies during early summer.

Material examined. — Holotype 2, Clandon,

5.vii.1953 (D. M. S. & J. F. Perkins) (BMNH);

Paratypes, 1 2, West Suffolk, 1920 (P. Har-

wood) (BMNH); 3 2, Woodwalton, Hunts.,

vii.1923 (P. Harwood) (BMNH); 2 6, Wood-

walton, Hunts., vu.1923 (P. Harwood)

(BMNH); 1 d, Dartmoor, 26.v11.1935 (R. C.

L. Perkins); 1 6, Killin, Perthshire, 22—

30.v1.1932 (R. B. Benson) (BMNH); 1 d, Wy-

tham Wood, Berks, 3.vii.1968, M.V. trap (BK);

1 4, Sandyhills, Lanarks., 23.viii.1966 (K. Stew-

art) (BK). As already stated, the Morley male

paratype of brevicornis is a specimen of the pre-

sent species (Surrey, 1900, coll. Sparke)

(BMNH).

Ophion pteridis Kriechbaumer

(figs. 33—38a, graphs 6, 6a, 9)

Ophion pteridis Kriechbaumer, 1879 (lectotype fe-

male, Munich). Perkins, 19-- (MS). Gauld, 1976,

1978. Oosterbroek, 1978.

Ophion scutellaris Thomson; Morley, 1915 (misiden-

tification).

Ophion pteridis Brauns; Gauld, 1973 (lapsus).

Ocellar-ocular interspace absent, to (excep-

tionally) 0.3 times interocellar space; posterior

ocellus to occipital carina 1.2—1.5 times maxi-

mum width of first flagellar segment, 0.5—0.85

times diameter of a posterior ocellus. Temple

much shorter than greatest transverse diameter

of eye, at least marginally shorter than first flag-

ellar segment. Sulci of stemmaticum complete,

posterior sulcus tending to run into back of eye,

lateral sulcus strong (usually deeper than mid-

dorsal sulcus). Occipital carina usually squared,

rounded or dipped (sometimes clearly angled)

centrally. Gena less than 0.3 to (some males) 0.7

times width of mandible base; postgena usually

less than or equal to length of genal inflection,

latter 0.7—0.8 times width of mandible base

Eyes somewhat convergent beneath in frontal

Brock: Ophion in Britain 89

view; facioclypeal area subquadrate to higher

than broad; mandible not long, gape rectangular

to obtuse, internal angles sharp, flange largely

impunctate. Punctures of epistoma and orbits of

similar distribution; last two segments of maxil-

lary palpus unequal; dorsal edge of mandible

about same width as a tooth, rounded into front

of mandible. First flagellar segment 3.5 (excep-

tionally only 3.0) to over 4.0 times longer than

broad. Flagellum with usual range of 53—57

segments (mean around 55): exceptionally, up

to 59 present.

Thorax with distinct coriaceous microsculp-

ture between punctures (especially on meso-

pleura); colour rather reddish; epicnemial an-

gles not aligned, pleurosternal angle sharp, more

. or less rectangular, lower sector of pleural epic-

nemium usually less than or equal to 0.5 times

upper. Scutellum usually carinate.

Submedian cell with sparse, elongate, evenly

distributed trichiae. Substigmal speculum not

extending beneath prestigma, trichiae in latter

region unusually elongate. Radius sinuate to-

wards base. Length 10—17 mm.

Hind femur quite slender, trochantellus

shorter than its minimum width in dorsal view.

Tibial spinules weak. Mid tibial spurs of un-

equal length (both elongate).

Propodeum with carina lateralis; superome-

dia and petiolar carinae rarely complete.

Membrane of petiolar segment well behind

line of spiracles.

Variation. — Males overlapping with the /#-

teus group with respect to the ocellar-ocular in-

terspace are not uncommon. The scutellar cari-

nae also, are frequently weak or absent. There is

also variation in the development of the propo-

deal carinae, which may range from consider-

able reduction to a condition of nearly complete

areolation; specimens approaching the latter ex-

treme are rare. Again, the darker colouration

and rougher microsculpture which tend to dis-

tinguish this from the next species, are subject

to quite frequent exception. Where difficulty 1s

experienced in separating variants of pteridis

and parvulus (or luteus from the former), a

point by point comparison with the descriptions

(bearing in mind the given variation) will gener-

ally produce a satisfactory result. I do not find

that Gauld (1978) has allowed sufficiently for

infraspecific variation in attempting to distin-

guish pteridis and parvulus.

Remarks. — Perkins’ views on the identity of

this species have been confirmed with reference

to the type. In his original description, Kriech-

baumer refers to a conspicuous dark and light

patterning, a feature absent in most specimens I

have studied — but certainly evident in the

type. I have seen a few British specimens which

do have this colouration, which I suspect is an

artefact produced either by a killing agent or by

decay of internal tissues.

Biology. — O. pteridis has been reared quite

regularly through night feeding Noctuid larvae

on seed pods of Campanula (Hadena). Other,

possibly regular hosts include Ceramica pisi

(Linnaeus) and Diataraxıa oleracea (Linnaeus).

These all pass the winter in the pupa state, and

for the most part, are not arboreal in habit.

They are apparently fairly mature larvae during

the flight period of the parasites.

Material examined. — Lectotype ® (Mu-

nich); 7 d, 50 ®, Minstead, New Forest,

Hants., M.V. trap, vii.—1x.1963—1975 (Siggs)

(BMNH); 1 d, 1 ©, Tiverton, vii.-ix.1954 (F.

H. Lyon) (BMNH); 1 6, Eltham, Surrey,

30.vii.1892 (MU); 1 d, Ogden Clough, Lancs.,

ex M. pisi, host 13.1x.1936, emerged 30.v.1937

(A. Brindle) (MU); 1 ©, Timperley, Cheshire,

121950 (EIER Galla) (MIO): 1 ® 1866,

Cheshire, 20.vin.1932 (H. R. Collett) (MU); 1

9, Framdley House, Cheshire, 3.1x.1946 (A. W.

Boyd) (MU); “Lichfield, Carr” (MU); 1 d,

Burnley, Lancs., 26.v.1923 (W. H. Clutten)

(MU); 1 6, Staley, Cheshire, 14.vu1.1945 (H.

N. Michaelis) (MU); 1 6, Burnley, Lancs.,

27.v.1923 (W. G. Clutten) (MU); 1 9, South-

port, 18.v.1929 (W. G. Clutten) (MU); 1 d,

Solihull, Birmingham, 17.viu.1969, M.V. trap

(M. R. Shaw) (BK); 1 d, Speke, Liverpool,

12.viii.1972 (BK); Marston, Oxford, 9.vin.1970

(BK); 1 6, Weymouth, Dorset, 16.vin.1974 (M.

J. Smith) (BK); 1 6, Richmond, Surrey,

31.vii.1971 (BK); 3 ©, Solihull, Birmingham,

viii.1969, M.V. trap (M. R. Shaw) (BK); 1 ?,

Mugdock, Glasgow, 11.1x.1971 (BK); 1 2,

Sandyhills, Lanarks., 23.vii.1966 (K. Stewart)

(BK); 1 2, Lambhill, Glasgow, 8.1x.1974 (BK);

1 2, Possil Marsh, Glasgow, 4.1x.1971 (BK); 1°

3,1 2, Pinner, Middlesex, garden M.V. trap,

vi.1977 (BK); 1 d, Claygate, Surrey, 2.v111.1958

(D. M. S. Perkins) (BMNH); 1 ©, Southamp-

ton, at light, 28.viii.1954 (W. Tampion)

(BMNH); 1 ?, Crapstone, Devon, 30.vı11.1968

(BMNH); 1 d, 1 9, Colthrop, 1906, ex D. cap-

sincola (Lyle coll.) (BMNH); 3 d, Exebridge,

NS., 15.viii.1958 (D. H. Harvey) (BMNH); 1

3, Digby, Lincs., at light, 18.vii1.1958 (C. R.

90 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Vardy) (BMNH); ? sex (damaged), Seaford, ex

D. cucubali, emerged 3.vi.1943 (Sir L. Wakely)

(BMNH); 1 à, Black Mountains, Powis, Wales,

ex Ceramica pisi, host larva coll. 3.viii.1975,

parasite emerged from prepupa, viii.1975, adult

parasite, 23.vii.1976 (S. J. Hopton) (Shaw coll.);

1 d, Taunton, ex C. psst, host larva 15.v111.1974,

paras. larva emerged ? vin.1974, adult

26.v1.1975 (K. Noble) (Shaw coll); 1 9, East

Didsbury, Manchester, ex D. oleracea, host lar-

va 15.1x.1974, adult paras. emerged 20.vii.1975

(M. R. Shaw); 1 d, 1 ©, Chislehurst, Kent, ex

D. oleracea, host larvae 15.x.1972, emerged

JOY, Dam lS (Stay colle I Bil 25

host larvae 26.vu.1973, parasite larva left host

prepupammer iele pars adults) davies &

13.vi.1974, ex H. bicruris (M. R. Shaw coll.); 1

g, Solihull, Warwicks., ex ? Mamestra brassi-

can host cols vil paras larva

c.15.1x.1973, adult 18.vii.1974 (M. R. Shaw

coll.); 1 d, 1 ?, East Didsbury, Manchester,

host 8.1x.1977, paras. larva 12.x.1977, adults

26.vu. & 9.viii.1978 (M. R. Shaw); ex D. capsin-

cola, 1.v1.1902 (Morley) (BMNH).

Ophion parvulus Kriechbaumer

(figs. 38b—40, 41b—45, graphs 6, 6a, 8, 9)

Ophion parvulus Kriechbaumer, 1879 (lectotype

male, Munich). Perkins, 19-- (MS: BMNH).

Gauld, 1976, 1978. Oosterbroek, 1978.

Ophion “species Y” Perkins, 19-- (MS: Oxford).

Ophion parvulus Brauns; Gauld, 1973 (lapsus).

Ocellar-ocular interspace absent to 0.3 times

interocellar space, longest in males; posterior

ocellus to occipital carina 1.2—2.0 times maxi-

mum width of first flagellar segment, 0.65—0.8

times diameter of a posterior ocellus. Temple

much shorter than eye, less than to distinctly

longer than first flagellar segment. Posterior

sulcus of stemmaticum tending to terminate at

margin of eye, lateral sulcus weaker than mid-

dorsal sulcus, frequently absent. Occipital cari-

na usually rounded, squared or dipped centrally

(though sometimes angled). Gena 0.3—0.75

times width of mandible base (longest in males);

postgena usually longer than or equal to length

of genal inflection, latter 0.35 to 0.7 times width

of mandible base. Eyes more or less parallel in

frontal view; facioclypeal area subquadrate to

wider than high; mandible with rectangular to

obtuse gape, sharp internal angles to the teeth,

flange indistinctly punctured medially. Punc-

tures Of epistoma similar, to somewhat closer

than those of orbits. Last two segments of max-

illary palpus unequal. Dorsal edge of mandible

narrower than a tooth, sharply defined from

front. First flagellar segment sometimes as short

as 3.0 times, usually around 4.0 to 5.0 times lon-

ger than broad. Usual range of 47—53 flagellar

segments (mean around 51); holotype with only

45 flagellars.

Thorax with moderate punctures, micro-

sculpture rather weak throughout, colour testa-

ceous, pleura often yellow marked above

(mesonotum rarely with yellow “vittae”); epic-

nemial angles remote, to more or less in line;

pleurosternal angle rounded, often distinctly

obtuse. Lower sector of pleural epicnemium 0.5

to 0.8 times upper sector. Scutellum rarely with

a trace of lateral carinae.

Submedian cell of forewing with trichiae

short, often sparser dorsally than below. Sub-

stigmal speculum generally produced beneath

prestigma. Radius evenly curved — to quite

deeply sinuate towards base.

Hind femur quite slender, trochantellus

shorter than minimum width in dorsal view;

tibial spinules weak, spurs of mid tibia not

elongate, of moderately unequal length.

Propodeum usually with carina lateralis about

as strong as dorsal carinae; superomedia and pe-

tiolar carinae usually incomplete, though less so

than in pteridis.

Membrane of petiolar segment subopposite

spiracles.

Variation. — Forms with a distinct ocellar-

ocular interspace are by no means rare, especial-

ly amongst males. These can readily be distin-

guished from luteus group species from the key.

Care must be taken however, to avoid misiden-

tification of yellow patterned individuals as

obscuratus, since the mesonotum may be “vitta-

te” and the stigmal apex yellow in exceptional

specimens. Much greater difficulty will be expe-

rienced in separating parvulus variants with

short first flagellar segment from mocsaryı, es-

pecially since the distribution of the trichiae on

the submedian cell (Gauld, 1978) is unstable; in

addition, the substigmal speculum to some ex-

tent, along with the carination of the propo-

deum, are subject to variation. Where ditticulty

is met, a point by point comparison of descrip-

tions will usually produce an acceptable solu-

tion. Confusion with pteridis is also quite likely

to occur, especially since there is a recurring

form of parvulus with distinctly sinuate radius.

When attempting to separate these from forms

of pteridis which converge on parvulus (e.g., in

Brock: Ophion in Britain

Graph 6. Temporal frequen-

cy distributions for Ophion

parvulus Kriechb. and O.

pteridis Kriechb.

FREQUENCY: HALF MONTHLY TOTALS

MAY

lack of scutellar carinae, etc.), special attention

should be paid to the shape of the facioclypeal

area, the longer postgena and temple ot parvu-

lus males and the position of the petiolar mem-

brane.

Remarks. — Perkins originally referred to

this species as “sp. Y” (MS), later suggesting the

name parvulus Kr. I have confirmed this syno-

nymy with reference to the type. The latter is an

abnormally small specimen of the species and it

lacks the darkened mesonotum mentioned by

Kriechbaumer (1879). It is otherwise in

agreement with the original description and

with that given in the present study.

Biology. — Hosts are Noctuidae which over-

winter as pupae. In early summer, Orthosia spe-

cies seem to be selected, while the autumn

brood has been reared through Euplexia and

Acronicta. The biological significance of this ap-

parent bivoltinism is unclear, since cocoons

from spring Orthosia appear to produce parasite

adults the following year (see remarks under

obscuratus). As unusual characteristic is the

habit of cocoon-forming within the ruptured

pupal remains of the host.

Common and widespread, perhaps less fre-

quent in the north.

Material examined. — Lectotype d (Mu-

nich); 34 d, 41 ®, Minstead, New Forest,

Hants., M.V. trap, vi—x1.1963—75 (L. W.

Siggs) (BK) (BMNH); 3 ©, Tiverton, vu.—

1954 (F, H. Lyon) (BMNH); 2 6, “O.

Winch. Hill, S. Hants.”, 4.vin.1954 (A. H. Sper-

ring) (BMNH); 1 2, Manchester, ex O. gothica,

JUNE

T T T T

JULY AUG. SEPT.

8.vi.1948 (L. Nathan) (MU); 1 6, Malham

Warns 25.21.1958. VEDA H nck) MU) MINCE

Frandley Hs., Cheshire, 11.v11.1957 (Boyd)

(MU); 1 ©, Manchester, ex gothica, 2.v1.1945

(MU); 1 ©, Doncaster, 14.vi.1921 (Clutten)

(MIU): 65 Keeler Dise, 2801925. (I:

Wood) (MD) 1 ©, Renan, Guns,

18531119537 (We la Davidson) (MIO) LE

ichfield, Carr” (MU); 1 ®, Delamere, Cheshire,

22.ix.1951 (H. N. Michaelis) (MU); 1 2, Burn-

ley, ex E.lucipara, 2.vii.1918 (Clutten)

(BMNH); 1 6, Chat Moss, Lancs., ex A. lepori-

na, 1.viii.1937 (L. Nathan) (MU); 3 d, Devon,

M.V. trap, 10.vii.1973 (M. R. Shaw) (BK); 1 d,

Stevenston, Ayrshire, 15.viu.1965 (BK); 1 6,

Milton Lockhart, Lanarks, vii.1965, ex coccoon

under bark, probably Acronicta sp. (BK); 13 ó,

Wytham, Berks., M.V. trap, 3.vil.—22.1x.1968

(J. P. Brock, M. C. Birch, N. Rae-Jones) (BK);

2 6, “Oxford” (M. P. Hassel) (BK); 1 d, Corn-

wall 4—10.vin.1974 (J. St. E. Cardew) (BK); 1

3, Solihull, WKS., 18.7.1969 (M. R. Shaw)

(BMNH); 1 6, Woodchester Park, Glos.,

ix.1972, M.V. trap (BK); 9 2, Wytham, Berks.,

28.v.-22.ix.1966 (BK); 1 9, “Oxford” (M. P.

Hassel) (BK); 1 4,3 ©, Pinner, Middx., garden

M.V. trap, vi.1977 (BK, private coll.); 3 d, 1 On

Colchester, 1945 (Harwood) (BMNH); 2 ?,

Arkley, 13.1x.1954 (T. G. Howarth) (BMNH);

1 2, Giggleswick, Yorkshire, 21.vui.1964 (E.

Burtt) (BMNH); 1 ©, Killin, Perthshire, vi.1932

(K. G. Blair) (BMNH); 1 d, Old Winchester,

Hants., 28.viii.1954 (A. H. Sperring) (BMNH);

1 ©, Bramshaw Wood, New Forest, Hants., on

birch, 20.vi.1964 (L. W. Siggs) (BMNH); 1 ?,

92 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Wimbledon, Surrey, 28.v.1957 (D. J. Clark)

(BMNH); 1 6, Windsor Forest, Berks., v.1953

(H. Donisthorpe) (BMNH); 1 (damaged) Bent-

ley, NE, ex C. coryli (Harwood) (BMNH).

Discussion of species criteria in the pteridis-

parvulus species pair

A Mann-Whitney test on the frequency dis-

tributions of flagellar segment number in these

two species shows significant median separation

for each segregate: P < 0.001 (graphs 2, 6a).

The bimodality of the parvulus curve (absent in

pteridis) may reflect an underlying genetic dif-

ference which has not received notice.

Pal

S _ pteridis

= N

= \

T Tu T T T T T T

49 50 51 5253 54 55 59

NUMBER OF FLAGELLAR SEGMENTS

Graph 6a. Flagellar segments frequency distributions

for Ophion parvulus Kriechb. and O. pteridis

Kriechb.

The temporal distributions of parvulus and

pteridis show a wide overlap, but pteridis is uni-

voltine, parvulus bivoltine. Again, pteridis

reaches a peak in numbers during the inter-gen-

eration depression of parvulus (graph 6).

From earlier remarks, it is clear that breeding

experiments to determine the biological nature

of bivoltinism in the last mentioned species

would be very desirable. Ecological separation

of pteridis and parvulus is decisive, but the latter

could possibly conceal two biologically distinct,

yet structurally identical species: circumstantial

evidence from biometric analysis of similar phe-

nomena in obscuratus however, seems to sug-

gest that a single species can be a “facultative bi-

voltine” unit.

Ophion mocsaryi Brauns

(figs. 46—50a, graphs 7—9)

Ophion mocsaryi Brauns, 1889: 89 (holotype female,

Budapest). Schmiedeknecht, 1908. Morley, 1915.

Perkins, 19-- (MS) (in part). Gauld, 1973, 1976,

1978 (in part). Oosterbroek, 1978.

Ocellar-ocular interspace absent, or very

short; posterior ocellus to occipital carina usual-

ly more than, sometimes equal to maximum

width of first flagellar segment, at least 0.5 and

usually around 0.7 times diameter of a posterior

ocellus; ocellar interspace greater than or equal

to maximum width of first flagellar segment.

Temple far shorter than eye, 1.0 to 1.25 times

length of first flagellar segment. Posterior sulcus

of stemmaticum deeply impressed, ending at

eye; lateral sulcı of stemmaticum weakly devel-

oped. Vertex with occipital carina sharply an-

gled centrally. Gena 0.2 to nearly 0.4 times

width of mandible base; postgena distinctly less,

to distinetly greater than length of genal inflec-

tion. Latter around two thirds of width of man-

dible base. Facioclypeal area subquadrate; man-

dible with rectangular to obtuse gape, sharp in-

ternal angles, Tse impunctate flange.

Epistomal and orbital puncturation similar. Last

two segments of maxillary palpus slightly to

strongly unequal in length. Dorsal edge of man-

dible as wide as a tooth, sharply offset from

front. First flagellar segment 2.4 to 3.5 times

longer than maximum width. Flagellum with

range of 50 to 58 segments (mean about 54).

Thorax moderately punctured, testaceous. Pleu-

rosternal angle of epicnemium sharp, often rec-

tangular, nearly aligned with sternal angles.

Lower sector of pleural epicnemium less than,

to equal to 0.6 times upper sector. Scutellum not

carinate.

Forewing with submedian cell having short,

evenly spaced trichiation. Nervellus broken at

or below the centre. Substigmal speculum not

produced into area beneath prestigma, that area

with short trichiae. Radius weakly to strongly

sinuate towards base. Forewing: 11.5—15.5

mm.

Hind femur moderately slender, trochantellus

shorter than minimum width. Tibial spinules

moderate. Mid ubial spurs unequal.

Propodeum usually lacking distinct carina

lateralis, dorsal carinae often complete and

strongly raised; spiracle more ovoid.

Membrane of petiolar segment approximately

in line with the spiracles.

Variation. — This species has generally been

characterised by authors, through the presence

of a complete area superomedia on the propo-

deum. This is not always in evidence, but is nev-

ertheless more frequently strongly developed

than in the majority of specimens of pteridis or

parvulus. Some variants of mocsaryi are liable to

be confused with parvulus and difficulty will al-

so be experienced in separating the present from

BROCK: Opbion in Britain 93

the next species. These problems are discussed

fully under parvulus and costatus.

Remarks. — Specimens identified as mocsaryi

in collections will generally be found to include

material of costatus Ratzeburg. These were not

distinguished by Perkins, either in his manu-

script key, nor in the British Museum collec-

tions. The type of mocsaryi has been seen and

the interpretation of authors confirmed.

Gauld (1978) does not allow for infraspecific

variation in attempting to distinguish the mocsa-

ryi — costatus species pair from the related pte-

ridis — parvulus pair. Females are particularly

likely to overlap on characters relating to the

tenth flagellar segment, as defined by that au-

thor. The characters used by Oosterbroek

(1978), partially after Gauld, are likewise unre-

liable for separation of species in the mocsaryı

group.

Biology. — This species has been reared

through night feeding Noctuid larvae maturing

during early summer on low shrubs. These in-

clude Triphaena and Orthostinae. Ophion moc-

saryi is widely distributed and sporadically

common in Britain, possibly more rare in Scot-

land (no records as yet).

Material examined. — Holotype ®, Buda-

pest, 1879, ex Cosmia ambusta (Prague); Para-

openden le Or same. data-as holotype,.d,

“19.1.1879”, ©, “19.iii.1879” (Berlin); 21 3,

Stark Ponds, Kingsteynton, 24.v.—8.v1.1942 (J.

F. Perkins) (BMNH); 6 6, 21 ©, Minstead,

NenamborestesHants:, NEVA traps 0 18.v.—

26.vii.19634—5 (L. W. Siggs) (BMNH); 1 d,

1 ®, Minstead, New Forest, M.V., 25.v. &

26.v.1975 (Siggs) (BK); 1 d, Formby Point,

Lancs., ex 7. fimbria, host 16.x1.1950, parasite

emerged 16.1v.1951 (K. C. Greenwood) (MU);

2 6, “Lichfield, Carr, 1920” (MU); 1 d, Fresh-

field, Lancs., 10.vi.1957 (W. D. Hincks) (MU);

? sex (damaged): Middlewood, Cheshire,

6.vi.1920 (H. Britten) (MU); 1 d, Formby

Moss, Lancs., ex T. fimbria, host 17.x1.1950,

emerged 23.1v.1951 (K. C. Greenwood) (MU);

1 2, Ainsdale, Lancs., swept from Salix repens

at night, 25.vi.1974 (M. R. Shaw) (BK); 1 9,

Wytham, Berks., M.V. trap, vu.1968 (M. C.

Birch) (BK); 2 2, Wytham, Berks., M.V.,

9.vi.1969, 27.vi.1968 (BK); 2 ©, “Oxford” (M.

P. Hassel) (BK); 1 d, Minstead, New Forest,

M.V., 28.v.1976 (L. W. Siggs) (BK); 1 d, New-

ton Abbot, SD., 1.vi.1941 (J. F. Perkins)

(BMNH); 1 ©, Tiverton, ND. vi.1955 (F. H.

Lyon) (BMNH); 4 6, Starks Ponds, King’s

Steynton, 24.v.—8.vi.1941 (J. F. Perkins)

(BMNH); 2 2, Minstead, New Forest, M.V.,

9.v1.1962 (L. W. Siggs) (BMNH); 1 2, Monk’s

Soham, Suffolk, with coccoon, 4.vi.1909 (Mor-

ley) (BMNH); 1 d, Ainsdale, Lancs., ex ? Xan-

thia icteritia on Salix repens, host larva coll.

5.vi.1975, parasite larva from host prepupa

v1.75, paras. adult 11.1v.1976 (M. R. Shaw coll.);

1 2, Ainsdale, Lancs., ex Agrochola or Xanthia

sp., host larva on S. repens 5.vi.1975, parasite

larva from host prepupa 16.vi.75, adult emerged

15.v.1976 (M. R. Shaw coll.); damaged speci-

men — St. Annes’, 1910, ex Orthosza lota (Clut-

ten) (BMNH).

Ophion costatus Ratzeburg

(figs. 50b—52, graphs 7—9)

Ophion costatus Ratzeburg, 1848: 79 (holotype de-

stroyed). Schmiedeknecht, 1908: 1438. Gauld,

1978. Oosterbroek, 1978.

Ophion mocsaryi Brauns; Perkins, 19-- (MS) (in part).

Gauld, 1978 (in part).

Ocellar-ocular interspace absent, or very

short; posterior ocellus to occipital carina less

than or equal to maximum width of first flagel-

lar segment, less than 0.5, to 0.7 times diameter

of posterior ocellus; ocellar interspace most

usually narrower than maximum width of first

segment of flagellum. Temple much shorter

than eye, 0.9 to 1.25 times length of first flagel-

lar segment. Posterior sulcus of stemmaticum

forming very deep grooves behind posterior

ocelli, ending at eye; lateral sulcus weak. Occi-

pital carina with rounded, weak angulation cen-

trally. Gena around 0.2 to nearly 0.4 times

width of mandible base; postgena distinctly less,

to distinctly greater than length of genal inflec-

tion — latter about two thirds width of mandi-

ble base. Facioclypeal area subquadrate. Mandi-

ble as in mocsaryi. Epistoma and orbits with fine

puncturation. Last segment of maxillary palpus

of same length, or slightly longer than penulu-

mate segment. First flagellar segment 2.3 to

nearly 3.0 times longer than maximum width

(shorter and broader on average than in mocsa-

ryi). Flagellum with usual range: 56 to 63 seg-

ments (mean about 61).

Thorax moderately punctured, tending to-

wards darker, more reddish testaceous. Pleuros-

ternal angle of epicnemium sharp, not obtuse —

more or less aligned with sternal angles; lower

sector of pleural epicnemiun around 0.6 X up-

per sector. Scutellum not carinate.

Wings similar to mocsaryi, but hind wing

with the nervellus intercepted above centre in

94 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

about 30% of specimens. Forewing: 14—17

mm.

Leg characters as in mocsaryi.

Propodeum often lacking distinct carina

lateralis, the dorsal carinae frequently complete

and often very strongly raised; propodeal spira-

cle more linear than in mocsaryı.

Abdomen as in mocsaryi.

Variation. — Due to overlap with the preced-

ing species with regard to the number of flagel-

lar segments and the interception of the nervel-

lus, it is likely that difficulty will be met with in

attempting to separate the present species from

mocsaryi. For this reason, all other characters

must be examined for reliable identification of

costatus.

Remarks. — The history of this species is

somewhat tortuous and can perhaps be summa-

rised as follows:

(1) The original description of costatus

Ratzeburg could apply to more than one species

of Ophion.

(2) The holotype of costatus no longer exists.

(3) Perkins used the name costatus for longi-

gena Ths. on the basis of his study of the Brauns

collection, since Brauns (1889) was first reviser

of costatus Ratz. (Perkins, pers. comm., 1978).

(4) Brauns himself doubted whether his own

species was the same as that described by Ratze-

burg. Furthermore, there now exists no material

of costatus Ratz.-Brauns in the Brauns collec-

tion.

(5) Schmiedeknecht (1908) identified costatus

Ratz. as a form close to mocsaryi Brauns, differ-

ing in characters which could only refer to

variants of the species for which I have adopted

the name costatus Ratz.

(6) Gauld (1973) published the Perkins use of

costatus (1.e., for longigena Ths.).

(7) Gauld (1978) now uses the name costatus

in the same context as did Schmiedeknecht,

without comment as to his change of opinion on

the Perkins interpretation. The characters given

by Gauld for separation of costatus and mocsa-

ryi follow the Schmiedeknecht diagnosis for the

most part, and will only separate about 30% of

material. _

It will be found, in some British collections,

that the present species has quite often been

misidentified as longicornis Brauns (see scutella-

ris Thomson). Post-1974, a number of speci-

mens in the British Museum and Manchester

Museum collections had been identified as cos-

tatus Ratzeburg, by the present author.

Biology. — Ratzeburg’s type of costatus was

reared through Acronicta aceris (Linnaeus). I

have come across no further material reared

through this host, for any Ophion species. M.

R. Shaw has reared costatus (as here recognised)

through Cucullia (Noctuidae). The host range

may then include both arboreal and other Noc-

tuid larvae which pass the winter in the pupa

stage. The true identity of Ratzeburg’s species

however, remains uncertain.

The species is seldom taken in any numbers,

but is widely distributed in England. As yet,

there are no Scottish records.

mocsanyi

costatus

FREQU: HALF MONTHLY TOTALS

JUNE JULY AUG.

Graph 7. Temporal frequency distributions for

Ophion mocsaryi Brauns and O. costatus Ratzeburg.

Material examined. — 6 d, Minstead, New

Forest, Hants., M.V. trap 31.v.1964, 4.v1.1964,

25.v1.1963, 25.v1.1965, 8.vi.1965, 2 ©, same

data, 4.vi.1964, 7.vii.1964 (L. W. Siggs)

(BMNH); 1 d, Hale, Cheshire, 16.vii.1923 (H.

RAP. Collett) (MU); MIN nen

Cheshire, 23.vii.1932 (H. R. P. Collett) (MU); 7

3, 3 2, Wytham Wood, Berks., M.V. trap,

4.v1.-12.v11.1968, 6.vi.1967; 2.vi.1969, 22.vi-

.1971 TP Brock, M: € Birch, MERS hm)

(BK); 2 2, Woodchester Park, Glos., 7.vii.1973

(M. R. Shaw); 2 d, Hereford, Craswall, 23—

24.v1.1973 (BMNH); 1 6, Reichenau district,

Semmeringgebiet, Lower Austria, 26.v.1957 (R.

B. Benson) (BMNH); 1 ©, Oxford, ex Cucullia

verbasci, host coll. 3.vii.1970, parasite adult

emerged: 28.v.1971 (M. R. Shaw collection); 2

3, Kirton, Suffolk, viii.1979 (M. J. Smith).

Discussion of species criteria in the mocsaryi —

costatus species pair

A Mann-Whitney analysis for comparison of

frequency distributions of flagellar segment

number showed significant separation; SND =

Brock: Ophion in Britain 95

À costatus

arvul

parvu US 65 |

FREQUENCY

NUMBER OF FLAGELLAR SEGMENTS

Graph 8. Flagellar segments frequency distributions

for Ophion parvulus Kriechb., O. mocsaryi Brauns

and O. costatus Ratzeburg.

7.03; P < 0.001 (see graph 8). A curve combin-

ing the two species is quadrimodal, preserving

the sexual bimodality apparent in the individual

distributions, plus the overall bimodality of the

species segregates.

In terms of overall resemblance, these two

species are closer than any other species pair in

Ophion. However, the actual zone of overlap in

flagellar segment number is distinctly less than

that found in parvulus/pteridis (compare graphs

6a and 8). Furthermore, the overall range for

this same character in a combined mocsaryi —

costatus aggregate is of even greater magnitude

than a similar combination of pteridis — parvu-

lus composition. The possibility of one extreme-

ly variable species may perhaps be ruled out on

the absence of any threshold effect in the regres-

sion curve (graph 5), as was observed for the

polymorphic obscuratus. On the other hand,

costatus is very nearly an extension of the moc-

saryı curve on a simple linear basis. The rather

small sample size for costatus inevitably makes

for difficulty in arriving at an absolutely final

assessment of the said data.

parvulus + pteridis

mocsaryi

+ costatus iN

FREQUENCY

NUMBER OF FLAGELLAR SEGMENTS

Graph 9. Combined flagellar segments frequency dis-

tributions for Ophion parvulus/pteridis and O. moc-

saryi/costatus.

Coming now to the question of the temporal

frequency distributions of the two species in

question (see graph 7), it is found that costatus

tends to be on the increase during the late de-

cline in mocsaryi numbers. This fits well with

the phasing of the host larvae, those of the first

species deriving from early summer moths, at a

time when mocsaryi hosts are mature larvae.

From the rather male dominated sex ratio in

costatus, it could be supposed that this was real-

ly a male trend from mocsaryi — but these

males would be appearing in the population at

the same time as the female sex was disappear-

ing! This seems biologically unlikely, and the

apparent predominance of males in costatus

could be an effect of small sample size: I have

also seen material obviously drawn from a heav-

ily male dominated population of “normal”

mocsaryı. In solving this anomaly, I have drawn

particularly on the extremely valuable material

collected at Minstead, New Forest on a year by

year basis, by the late L. W. Siggs.

A final comment on the possibility of a host

correlated dimorphism underlying the costatus-

mocsaryi species pair comes from the fact that

mocsaryi specimens reared through the large

Noctuid Noctua do not converge on costatus

form, as compared to the Orthosiinae rearings.

O. mocsaryi and costatus remain the most

closely related species pair amongst British

Ophion species. Investigations concerned with

establishing whether or not these segregates are

really biologically distinct should not only deal

with a larger and more widely distributed set of

population samples, but must also bear in mind

the probable biological criteria already dis-

cussed. It is particularly important that the defi-

nition of segregates given herein be taken fully

into account, in respect of apparent interme-

diate forms. The characters given by Gauld (loc.

cit.) certainly break down with an examination

of infraspecific variation, even with the relative-

ly small sample sizes already available. It per-

haps goes without saying, that breeding experi-

ments are much to be desired in resolving out-

standing problems.

SUMMARY OF HOST SPECIES FOR OPHION

(VERIFIED RECORDS ONLY)

minutus

Agriopis (Erannis auctt., ptm.)

A. aurantiaria (Hubn.)

A. marginaria (F.) (prob. plus E. defoliaria)

ventricosus

96 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 4 (1982)

Apocheima pilosaria (D. & S.) (Phigalia pedaria F.

of auctt.)

scutellaris

Lycophotia porphyrea (D. & S.) (L. varia, of

auctt.)

Noctua (Triphaena, auctt.) fimbriata (Schreber)

Xestia (Amathes, auctt.) xanthographa (Schiff.)

Aporophyla nigra (Haw.)

luteus

Ochropleura (Agrotis, auctt.) praecox (L.)

(usual host not known)

obscuratus

Acronicta cuspis (Hubn.)

Lycophotia porphyrea (D. & S.)

Noctua (Triphaena, auctt.) comes (Hubn.)

N. fimbriata (Schreber)

N. interjecta (Hubn.)

Mythimna impura (Hubn.) (formerly Leucania)

M. ferrago (F.)

Xestia (Amathes, auctt.) agathina (Duponch.)

X. castanea (Esper)

X. baja (Schiff.)

Graphiphora augur (F.).

Agrochola or Xanthia sp. indet.

forticornis

Ochropleura (Agrotis, auctt.) praecox (L.)

longigena

Cucullia chamomillae (Schiff.)

C. ?scrophularıae (Capieux)

brevicornis

Cosmia trapezina (L.)

perkinsi

unknown

crassicornis

Aporophyla nigra (Haw.)

parvulus

Orthosia cruda (Schiff.)

O. gothica (L.)

Euplexia lucipara (L.)

Acronicta leporina (L.)

Acronicta sp. indet.

Ipimorpha retusa (L.)

Cosma trapezina (L.)

Colocasia coryli (L.)

pteridis

Ceramica pisi (L.)

Hadena bicruris (Hufn.)

H. rivularis (F.)

Diataraxia oleracea (L.)

?Apamea unanimis (Hubn.)

mocsaryi

Agrochola lota (Clerck)

Noctua (Triphaena, auctt.) fimbriata (Schreber)

Atethmia ambusta (F.)

Xanthia or Agrochola sp. indet.

costatus

Acronicta aceris (L.) . . . lost type!

Cucullia verbasci (L.)

CONCLUSIONS

Comparative studies show much infraspecific

variation in Ophion species, which may greatly

overstep species-to-species differentials (see

curve 2, for example). Members of the luteus

complex are extremely closely related and diffi-

cult to define, other than in a context of highly

polythetic character distribution. The question

of temporal speciation cannot be ruled out for

obscuratus and parvulus, nor is it yet entirely

certain that mocsaryi and costatus are real bi-

ological species isolates. Breeding experiments

with the last two forms (also with luteus, obscu-

ratus and parvulus) will be necessary in order to

fully resolve some of the questions which can-

not be fully answered through morphological

investigations alone.

APPENDIX: NOTES ON PLATOPHION

Platophion species are at once recognisable

through their lack of any occipital carina; they

appear to be parasites of Thyatiridae (BMNH

records, confirmed M. R. Shaw, pers. comm.).

Both generic and species diagnoses have been

affected by the small number of available speci-

mens. Preliminary study suggests that the two

species can be distinguished thus:

ocellaris (Ulbricht)

First flagellar segment less than 3.5 X longer

than broad; flagellum with at least 50 segments;

paler, propodeum more weakly areolated.

areolaris (Brauns)

First flagellar segment at least 3.5 x longer than

broad; flagellum with less than 50 segments;

darker, propodeum more distinctly areolated.

As stated by Oosterbroek (1978), the charac-

ters given by Gauld (1973) are not reliable for

separation of these two species. While I have

been able to gain some knowledge of infraspe-

cific variation within ocellaris, it is quite likely

that discovery of further areolaris material will

further modify our concept of species differ-

ences within Platophion. Oosterbroek (loc. cit.)

reasonably questions the validity of the two

species here treated.

REFERENCES

Brauns, S., 1889. Die Ophioniden. — Arch. Ver.

Freunde Naturg. Mecklenb. 43: 58—72.

Curtis, J., 1835. British Entomology, 3. Hymeno-

ptera. — London.

Fabricius, J. C., 1798. Supplementum Entomologica

Systematica. — Halfniae.

1804. Systema piezatorum. — Brunsvigae.

Gauld, I. D., 1973. Notes on the British Ophionini

(Hym., Ichneumonidae) including a provisional

key to species. — Entomologist’s Gaz. 24: 55—

65.

Brock: Ophion in Britain 97

— 1976. Notes on the British Ophioninae (Hym.,

Ichneumonidae). Part 3. The identity of the spe-

cies described by Morley, 1915 and Thomson,

1888 and a checklist of British species. — Ento-

mologist’s Gaz. 27: 113—117.

— 1978. Notes on the British Ophioninae (Hym.,

Ichneumonidae). Part 4. A revised key to the spe-

cies of the genus Ophion Fabricius. — Entomo-

logist’s Gaz. 29: 145—149.

Gravenhorst, J. L. C., 1829. Ichneumonologia Euro-

paea, 3. — Breslau.

Habermehl, H., 1930. Neue und wenig bekannte pal-

aarktische Ichneumoniden. — Konowia 9: 109—

117.

Jusilla, R., 1976. Contribution to the knowledge of

the Norwegian fauna of Ichneumonidae

(Hymenoptera parasitica). — Norwegian Journ.

Ent. 23: 97—120.

Kriechbaumer, J., 1879. Ophion parvulus n. sp. and

O. minutus n. sp. — Ent. Nachr. 1879: 104—106.

1892. Ophion slaviceki n. sp. — Ibid. 1892: 233.

Linnaeus, C. von, 1758. Systema Naturae, ed. x. —

Holmiae.

Morley, C., 1915. Ichneumons of Gt. Britain, 5. —

London.

Oosterbroek, P., 1978. Dutch Ophionini (Hym., Ich-

neumonidae, Ophioninae). — Ent. Ber. 38: 103—

112.

Perkins, J. F. Undated. Manuscript (BMNH).

Ratzeburg, J. T. C., 1848. Die Ichneumonen der

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Schmiedeknecht, O., 1908. Opuscula Ichneumonolo-

gica 4: 1434— 1449. — Blankenburg i Thuringen.

Schmiedeknecht, O., 1936. Idem (Supplements) 25:

Sl. :

Thomson, C. G., 1888. Ofversigt af de i Sverige funna

arter af Ophion och Paniscus. — Opusc. Ent. 12:

1185—1201.

Thunberg, C. P., 1822. Ichneumonidea, Insecta

Hymenoptera, illustrata. — Mém. Acad. Imp. Sci.

St. Pétersbourg 8: 249—281.

Townes, H., 1971. The Genera of Ichneumonidae. —

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Inst. 5.

PDEELM25 AFLEVERING 5 1982

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

va JS. COMP. ZOOL

LIBRARY

JAN 3 1 1983

HARVARD

UNIVERSITY

INHOUD

J. KieLLanD. — Revision of the genus Ypthima in the Ethiopian Region excluding

Madagascar (Lepidoptera, Satyridae), pp. 99—154, figs. 1—185, pls. 18.

Tijdschrift voor Entomologie, deel 125, afl. 5 Gepubliceerd 31-XII-1982

REVISION OF THE GENUS YPTHIMA IN THE ETHIOPIAN

REGION EXCLUDING MADAGASCAR (LEPIDOPTERA,

SATYRIDAE)

JAN KIELLAND

4916 Boröy, Norway

ABSTRACT

The Afrotropical species of the genus Ypthima are revised. All known species and sub-

species are described, including the following new taxa: Y. condamini, Y. jacksoni, Y. lam-

to, Y. vuattouxi, Y. yatta, Y. antennata cornesi, Y. condamini nigeriae and Y. pupillaris

obscurata. The female of Y. diplommata Overlaet is described for the first time. Keys are

provided based on the male and female genitalia. Photographs of the butterflies and free-

hand drawings of the male and female genitalia are given, as well as distribution maps of all

known species and subspecies.

INTRODUCTION

The genus Ypthima was erected and de-

scribed by Hübner (1818). Ypthima huebneri

(Kirby, 1871) (an Indian species), was desig-

nated type-species by the Commission. Since

Elwes & Edwards (1893) revised the genus, no

attempt has been made to give a complete sur-

vey of the Ypthima species occurring on the Af-

rican continent. Other students have dealt with

species from certain parts of Africa only, mainly

from relatively well accessible areas. This has

led to many misidentifications and a confusing

synonymy.

In this paper an attempt is made to give an

up-to-date account of the genus Ypthima occur-

ring on the African continent. Work was long

hampered and delayed due to the difficulty in

obtaining enough material of some of the spe-

cies from various parts of Africa. To get a com-

paratively clear picture of the distribution of the

various species, it has been necessary to write to

many private collectors and institutions for loan

or exchange of material. To start with, the au-

thor’s private collection from Tanzania and the

large collection in the National Museum, Nai-

robi, were studied and arranged. It was soon ap-

parent that this was by no means enough, so re-

quests were made for type material and photo-

graphs from various museums in Europe and

South Africa. They were generously supplied

and did help to a great extent. Later, large col-

lections from the Ivory Coast revealed species

not yet described and many other interesting

facts. Finally, a study was made of the extensive

Ypthima collections in the British Museum

(Nat. Hist.) and this revealed further interesting

facts and helped to clear up some difficulties.

As many of the Ypthima species are extreme-

ly variable during the seasons and often external

characters alone are of little use in identifica-

tion, extensive dissections were carried out and

a key was made, mainly based on the genitalia

of both males and females. Apart from the geni-

talia, also antennae, labial palps and fore legs

were examined.

In spite of generous help from institutions

and private persons, further study is needed on

this group, particularly with regard to distribu-

tion and subspeciation. There are many parts of

Africa from where it is not easy to obtain

enough material.

ACKNOWLEDGEMENTS

The writer is greatly indebted to Mr. M. Clif-

ton of the National Museum, Nairobi, for loan

of specimens and free access to the Museum’s

collections, for undertaking photographing and

for sound advice; to the keeper of Entomology

and staff of the British Museum (Nat. Hist.),

particularly to Mr. T. G. Howarth, Mr. R. I.

Vane-Wright and Mr. P. R. Ackery for dissect-

ing types, for presenting the writer with photo-

copies of literature, loan of specimens, free ac-

cess to the museum’s collections and valuable

information and advice; to Dr. M. Condamin,

I.F.A.N., Dakar for presenting 40 specimens of

100 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Ypthima, photographs and the loan of 560 spec-

imens from the Ivory Coast and another large

collection from the Central African Republic

and for lots of valuable correspondence; to Mr.

L. Berger, Tervuren Museum, Belgium, for loan

of type material; to Dr. H. J. Hannemann, Ber-

lin Museum, for loan of specimens and to Dr. L.

Vari, Transvaal Museum, South Africa, for loan

of specimens and for permission to examine the

museum’s collections on my visit to South Afri-

ca. My thanks are also due to Mr. N. H.

Cornes, N.S.P.R. Institute, Lagos, Nigeria, for

the loan of Nigerian specimens and to Dr. M. B.

Usher, York University, for the opportunity to

examine his Ypthima material from Ghana, for

providing me with the localities, and for valu-

able advice on the manuscript; also the follow-

ing persons who have supplied me with speci-

mens: Father Maessen, Ghana; Dr. Mc Cleery,

Zomba, Malawi (now in England); Mr. S. C.

Collins, Nairobi, Kenya; Mr. W. H. Henning,

Johannesburg, South Africa. I also wish to

thank Dr. A. Lillehammer and Mr. M. Opheim

of Töyen Museum, Oslo, for facilities in the use

of a room and equipment for study while I was

on a visit to Norway in 1969; to Mr. R. Mehl,

Töyen Museum, for photographing some of the

specimens. Finally, thanks are due to Mr. T. G.

Howarth for reading and correcting my manu-

script and to Mr. R. I. Vane-Wright, Mr. R. L.

Smiles and Mr. M. B. Usher for reading the

manuscript and making suggestions.

ABBREVIATIONS USED IN THE TEXT

BM British Museum (Nat. Hist.), Lon-

don

BMG Berlin Museum, D.D.R.

DIG discoidal cell

d.s.f. dry season form

f.w. fore wing

h.w. hind wing

IFAN Institute Fondamental d’Afrique

Noire, Dakar

int. f. intermediate form

NMN National Museum, Nairobi

NSPRI N.S.P.R. Institute, Lagos, Nigeria

spec. nova new species

stat. nov. new status

TMB Tervuren Museum, Belgium

TMSA Transvaal Museum, South Africa

w.s.f. wet season form

The preparation numbers followed by an A

or K are genitalia preparations deposited in the

British Museum; all numbers without letters are

in the author’s collection and some in the Na-

tional Museum, Nairobi.

CHARACTERIZATION OF THE GENUS

External characters. — The fore legs of the

males are very minute and usually much shorter

than in the female, whose fore legs are much

better developed. In the female of Y. recta,

however, they are much reduced and not much

larger than in the male. The wings are rather

uniformly brown except in Y. albida; they have

two swollen veins in the basal area and there are

no androconial hair tufts on the wings. The an-

tennae are shorter than half the length of the

fore wing, except in the West African race of

Y. antennata. The club is usually narrow and

long, not much thicker than the shaft. The only

species here dealt with that has a thick club is

Y. antennata. The underside of the club is dou-

ble-grooved on each joint.

Male genitalia. — An important character in

the male genitalia of Ypthima is the shape of the

valva; the length of the valva in relation to the

length of the aedeagus and/or to the combined

length of the tegumen-uncus, the width and

general shape of the uncus are also of taxonomic

value, as is the vinculum, which may be nearly

straight as in Y. condamini, or strongly bent as

in Y. asterope.

At least in the genus Ypthima it is advisable

to study the genitalia unmounted, to make it

possible to compare species both in dorsal (or

ventral if preferred) and lateral aspect. Personal-

ly I prefer the dorsal aspect with regard to Yp-

thima. Certain important features can only be

seen in dorsal or ventral view, particularly the

width of the tegumen and the uncus, the posi-

tion of apophyses angulares which in certain

species are turned interiorly (towards each oth-

er). There are no subunci in Ypthima.

Female genitalia. — The taxonomically most

important characters in the female genitalia are

apparently found in the genital plates, one ante-

rior and one posterior to the ostium. They are

variable within the same species, but only to a

limited extent (no. 865, with a folded ant. plate,

but with external characters exactly like a nor-

mal Y. asterope, may constitute another species,

but is more probably an aberrant specimen of

Y. asterope.) The key is based to a large extent

on these characters. The nature of sclerotization

of the plates is also of major importance. The

shape and depth of the wall encircling the sinus

vaginalis is of taxonomic value and also to a cer-

tain extent, the presence or absence ot sclero-

KIELLAND: Ethiopian Ypthima 101

tized ribs of the anterior wall, but in some spe-

cies, as in Y.asterope, the wall is strongly

ribbed in some specimens, and not ribbed at all

in others. In view of this variability within a

species, this feature is not included in the key.

Other parts of taxonomic value are the length

and width of the ductus bursae and the size and

shape of the corpus bursae. There is no signum

in Ypthima.

EcoLoGY

The adult Ypthima is a typical inhabitant of

open grassland and woodland. A few species

however prefer moister habitats. Y. albida in-

habits forest margins and swampy places near

rivers and along forest edges. Y. doleta also pre-

fers humid areas and does not occur in more ar-

id country. In Tanzania, Y. doleta only occurs

in the humid Bukoba District in the north-west

corner. Y. asterope inhabits the more arid coun-

tries of Africa, such as Southern Africa, the dry-

er parts of Kenya to Somalia, parts of Ethiopia,

Sudan and west to Senegal. It seldom penetrates

into the more humid areas of tropical Africa,

and in Asia it occurs from the Arabian peninsula

to Lebanon, Israel and India. Personally I have

found both males and females of Y. antennata

concentrated in rocky areas of sandstone for-

mations, often settling on the rocks.

The flight of all Ypthima species is weak and

erratic and they tend to fly close to the ground

between grasses, frequently settling on the

ground or on low vegetation. As a rule, they do

not visit flowers, nor are they attracted by fer-

mented juices.

Discussion

At least with regard to Ypthima, the writer

has come to the conclusion that the comparative

length of the two last joints of the antennal club

is of much less taxonomic value than previously

thought. Generally the two last joints vary a

great deal within the same species, even within

the same locality, as shown with regard to

Y. impura and Y. pupillaris. The length and

thickness of the club in Y. antennata is also

very variable, to such a degree, that I at first

presumed that two species were involved. Spec-

imens of Y. antennata in Rhodesia to central

Tanzania normally, but not always, develop a

shorter and thicker antennal club than those

taken further north, from northern Tanzania to

Sudan. The genitalia, however, show no differ-

ence in either sex.

It has been found that the numerous aber-

rants described have little or no value as such.

For the greater part, they constitute seasonal

forms and also variations within the same sea-

son. Some even are distinct species, having

nothing to do with the species they were sup-

posed to be aberrants of. Many earlier writers

attached too much importance to the number

and size of the ocellar spots. They are in fact ex-

tremely variable within the same species.

The antenna-wing ratio is measured for each

species. However, I do not find it to be of too

much importance. It is extremely difficult to

measure the length of the antenna accurately,

because it is in most instances more or less

curved. Even when straightened out with alco-

hol, they quickly curve back to the original po-

sition and it is difficult to get it measured in

ume. A slight error may make a fairly big differ-

ence in the ratio. Species with only a little dif-

ference of the antennal length compared to the

wing can hardly be distinguished from each oth-

er in this manner and this applies to most of the

Ypthima dealt with here. The ratios also vary in

different habitats. An example of this is Y. an-

tennata, with a much higher ratio for the Nige-

rian race than for the aggregate from the eastern

part of Africa. Ratios for Y. impura impura and

Y. pupillaris pupillaris from the Ivory Coast are

higher than for the eastern races. Curiously

enough, it seems that the aggregates from west-

ern Africa exhibit higher ratios than do speci-

mens of the same species from Fast Africa. It

probably has something to do with the environ-

ment. (Y. doleta, however, shows no variation

in the ratio.) One might therefore presume that

this difference, constant for each habitat, sug-

gests separate species. At least for Y. impura

and Y. pupillaris, this is out of the question, as

both exhibit characteristic and constant genitalic

characters in both sexes and races.

As stated under Y. asterope, the specimens

from the Nairobi area with the more slender

valva, may prove to be distinct from Y. aste-

rope. However, the external characters show no

difference whatsoever, and too few specimens

of this form are available, and the female is not

yet known. The specimens of Y. asterope from

South Africa and Rhodesia differ somewhat

from specimens taken in Kenya to Sudan in the

underside h. w. discal band. The status of Y. as-

terope asterope and Y. asterope hereroica is not

quite clear. Apparently these two races do have

a common boundary and even overlap in one

area. In Tanzania Y. asterope asterope, as far as

I know, does only occur in the Tanga Region

102

(one male in the BM, taken by T. H. E. Jack-

son). The previous records proved to be wrong.

I have not seen any specimens from Malawi and

Zambia and no material of Y. asterope is avail-

able from Zaire. On the other hand, it turns up

again in Gambia and Senegal and Northern Ni-

geria. It occurs further north from here and

probably also across Africa, just south of the Sa-

hara to Uganda and Kenya. However, further

south there is a blank area straight across Africa

and Y. asterope does not occur again before we

get to Angola and the southern part of Africa.

In fact, Y. asterope is less common and wide-

spread in Africa than earlier believed. As to the

related species Y. condamini, Y. rbodesiana and

Y. congoana, these all occur within the “blank”

area. Y. condamini has a wide, although scat-

tered distribution from South Africa to Senegal

and Uganda. Y. rbodesiana is more restricted to

the eastern part of Africa, from Rhodesia to Ke-

nya and Katanga in Zaire. While the distribu-

tion areas of these two species considerably

overlap the area of Y. asterope, Y. congoana has

hitherto only been taken in the “blank” area,

being mainly restricted to Katanga.

The discovery of Y. antennata on the plateau

in Nigeria proves that this species is more

widely distributed than was previously thought.

The Nigerian specimens, however, constitute a

separate race. It is very likely that Y. antennata

occurs in many more areas of Africa, as this spe-

cies, in spite of its very wide club, is frequently

confused with and often referred to Y. asterope.

KEY TO THE CONTINENTAL AFRICAN SPECIES OF

Ypthima, BASED ON THE MALE GENITALIA

DVI ES M RICE PR EP EE, 2

= OWN INES DOWN AIN N 4

2. Upperside markings clearly defined ..... 3

Upperside markings faint or greatly re-

Ace: MARMEREN CITI albida uniformis

3. Upperside wings bluish white; underside

pale albida occidentalis

Upperside wings whitish; underside rather

dark albida albida

4. Valva, distance from base to ventral bend

less than half the total length of valva (in

CPAP POW EGNOS) EE A 6

Valva, distance from base to bend more

than half the length of valva; dorsal aspect

of uncus gradually tapering to a narrow dis-

tal portion’: Je RE eee 5

5. Apex of valva thick, knob-like...... doleta

— Apexofvalvaslender ........ diplommata

. Uncus as under 5; aedeagus sinuous or

TIJDSCHRIFT VOOR ENTOMOLOGIE,

10.

112,

15°

14.

DEEL 125, AFL. 5 (1982)

rather straight, not evenly curved from the

open basal portion, except in condamini.. 7

Uncus, basal portion flattened, not gradual-

ly tapering to the narrow part, see fig. 75;

aedeagus evenly curved and gradually ta-

pering 20

. Aedeagus usually very sinuous, except in

condamıni; distal portion of valva more or

less flattened dorsad and usually with sharp

inner edge 12

Aedeagus straighter; valva not flattened

distad and without sharp inner edge ..... 8

. Aedeagus as long as, or longer than valva;

valva elbowed as normal 10

Aedeagus shorter than valva, distal half

sharply narrowed ventrally; valva narrow

and only a little elbowed

. Valva narrow and straight; aedeagus much

shorter than valva, ratio 0.75; h.w. upper-

side discal and submarginal bands promi-

nent pulchra

Valva, narrow portion sinuous; aedeagus a

little shorter than valva, ratio 0.87; h.w. up-

perside discal and submarginal bands

obscured praestans

Antennal club short and thick; valva, ven-

tral curve even and gradual, distance from

base to curve equal to, or a little longer than

distal poro 1 A antennata

Antennal club thin; valva, ventral curve

acute, distance from base to curve much

shorter than distallportiow =a ee Sent 11

Valva as long as tegumen-uncus... jacksoni

Valva much longer than tegumen-uncus....

eee. vuattouxi

Accessory plate placed at a sharp angle to

the longitudinal section of valva; aedeagus

longer than valva 13

Accessory plate almost perpendicular to the

longitudinal axis; aedeagus shorter than,

equal to or longer than valva .......... 16

Vinculum long, almost straight; valva, nar-

row distal portion short and robust,

somewhat swollen distad of the ventral

bend see ties 430 00 Or condamını

Vinculum moderately or strongly curved;

narrow distal portion of valva not robust

E Dese so eo 2 14

Vinculum moderately curved; inner edge of

valva, narrow distal portion very sharp

from apex to near wide portion; diaphrag-

ma unsclerotized; appendix angulare bent

INCELIOLI VARE RIOT simplicia

Vinculum strongly curved; inner edge of

valva moderately sharp, for a short distance

15.

16.

18.

Dil

22%

25:

26.

KieLLAND: Ethiopian Ypthima 103

EE I O IT 15

Diaphragma with a triangular sclerotiza-

tion; appendix angulare straight. ..........

EEE HAR. asterope (atypic)

Diaphragma without sclerotization; appen-

dix angulare bent interiorly … simplicia

Valva robust; apex and accessory plate

OEE Di EEE 17

Valva much more slender, distal portion

slender, width of accessory plate not as

bueadtasapextonvalvaliin. neten. DT. 19

Valva dorsal margin twice the length of

SAGE LL A asterope asterope

Valva dorsal margin 1.5 times the length of

Bremen: late ls ab ac 18

Aedeagus as long as valva; upperside f.w.

ocellar area usually sharply defined ........

den els ee EEE asterope hereroica

Aedeagus much longer than valva; upper-

side f.w. ocellar area obscured … yatta

. Uncus rather broad; aedeagus as long as, or

a little shorter than valva....... rhodesiana

Uncus narrow; aedeagus as long as or a

little longer than valva.......... congoana

. Uncus wide from base to near apex, to nar-

row portion; valva, basal portion very

wide, narrow portion very slender ..... 26

Uncus and valvanotasabove.......... 21

Uncus, narrow portion strongly curved;

valva long, slender and rather evenly ta-

pering; aedeagus longand very thin .... 25

Uncus and valva normal; aedeagus short 22

Valva, base of narrow distal portion thick-

ened, see fig. 61, thus robust part twice as

lougasthe thintportion.. 1.4 lamto

Narrow portion of valva not thickened

BES ee EE 23

. Diaphragma with a triangular sclerotiza-

tion; antenna-wing ratio 0.36.......... 24

Diaphragma without sclerotization; anten-

HA Wwinorato 044 er ae es a recta

. Basal end of aedeagus widened and straight

COMPRI Eri ci. granulosa (inland form)

end of aedeagus rounded, not

widened 0... granulosa (coastal form)

H.w., upperside discal bands clearly de-

BE eee cece ec. impura impura

H.w., discal bands not clearly defined......

impura paupera

Narrow distal portion of valva slender, as

long as, or longer than dorsal length of

broad portion; inner side of f.w. ocellar

area above, clearly defined; h.w. discal

markings distinct ...... pupillaris pupillaris

Narrow distal portion of valva less slender,

. Antennal

. Anterior plate simple,

. Sinus vaginalis deep;

in most cases shorter than dorsal length of

broad portion; inner side of f.w. ocellar

area not clearly defined; h.w. markings

obseuredes ads sole pupillaris obscurata

KEY TO THE CONTINENTAL AFRICAN SPECIES OF

Ypthima BASED ON THE FEMALE GENITALIA

1. Anterior plate heavily sclerotized; posterior

plate with posterior part sclerotized, the

frontal part formed like an unsclerotized

tonsuelike lobe me Sr DER À

Anterior plate round and unsclerotized;

posterior plate as above; edge of anterior

platesbinelyasenra er ER congoana

Both plates unsclerotized; sometimes ante-

rıor plate weakly sclerotized ........... 8

Differing from above................. 10

. Anterior plate small, rather narrow; poste-

Lior plate wider andishontem ARRE e 3

Anterior plate large and wide ........... 7

. Sclerotized part of posterior plate rather

narrow, usually not bipectinate distad; its

anterior lobe small and pointing ventrad

Sclerotized part ot posterior plate large and

bipectinate distad; anterior lobe as above;

bursa very long and narrow ......... yatta

Sclerotized part of posterior plate large and

broad; anterior lobe tongue-like and point-

ing more posteriorly; ductus wide and

shortabursanwid ef ea hate 4

. Apex of posterior plate bipectinate; under-

side of wings rather pale; discal line of h.w.

evenlyacurvede APCE Meer rhodesiana

Apex of posterior plate rounded with or

without apical emargination; underside of

wings dark; h.w., discal line irregular ......

iL simplicia

club abrupt and thick; bursa rath-

er large; sinus vaginalis moderately deep;

posterior apophyses rather narrow ........

frutti uan tas pd cr antennata

Antennal club gradual, narrow; bursa

smaller; sinus vaginalis wide and shallow;

posterior apophyses rectangularly shaped 6

bilobed, with or

without spines at margin; anterior wall of

sinus vaginalis evenly rounded; posterior

platswuariable ae ese asterope

Anterior plate strongly folded; anterior

wall of sinus vaginalis prolonged anteriorly

EE RUE. de asterope (atypic)

anterior plate not

harp! ione 0301305: Ki AA. granulosa

Sinus vaginalis wide and shallow; anterior

plate almost circular, covered with spiny

104 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

hars oe Rito. SE, condamini

8. Posterior plate sack-like, wider than ante-

rior plate; both plates unsclerotized ..... 9

— Anterior plate wider than posterior plate,

strongly bilobed, sometimes slightly sclero-

Zedd AR MR ARI, doleta

9. 2Wnssiwhtten ANSE eea albida

Nine brown nn ur ee jacksoni

10. Anterior plate sclerotized, large and wide,

simple, usually widest in the middle .... 13

— Anterior plate tongue-like, parallel-sided;

distal end of posterior plate bent over the

ant. plate; sinus vaginalis shallow .. pulchra

— Anterior plate complex, folded, sinus vagi-

nalisdéepet eee bene saleen 11

11. Both anterior and posterior plates complex

ane.tolded un. sonen ee 12

— Only the anterior plate folded; posterior

Platelonsandmarrows re impura

12. Anterior part of posterior plate bilobed ....

des re Sep pa ee pupillaris

— Anterior part of posterior plate evenly

founded hare ey Base a praestans

13. Posterior plate unsclerotized, not bilobed ..

SARO GRA RES RATER RE Pinar EEN 14

ete A vuattouxi

14. Posterior plate wide at base with long,

tongue-like frontal lobe, edges even; sinus

vaginalis deep; lateral sclerites wide. .... 15

— Posterior plate widest in the middle, folded,

CASES EVEN ANO diplommata

15. Anterior plate variable, edge uneven and

serrated all the way; posterior plate very

wide basad with a moderately long anterior

lobe; bursa small; fore legs only a little

loca daam im WHO TINE 5.2... recta

— Anterior plate evenly edged and serrated

only at the distal end; posterior plate less

wide at base with a longer anterior lobe;

bursa longer; fore legs much better devel-

pei mine male 0.0 000000 lamto

DESCRIPTION OF THE Ypthima SPECIES

Ypthima asterope (Klug)

This species, which is distributed throughout

most of south-west Asia and Africa excluding a

belt crossing the central part, has been the cul-

prit of many misidentifications. In earlier times

when dissection was not generally done, au-

thors like Strand in particular, described many

aberrations which in most cases were forms be-

longing to other species, particularly Y. granu-

losa. Unfortunately, Strand’s types were de-

stroyed during the 1939—45 war and the de-

scriptions alone are not sufficient for sure

identification. However, it is fairly certain that

his Y. asterope ab. interrupta is the w.s.f. of

Butler’s Y. granulosa and his ab. uniocellata the

d.s.f. Strand (1909: 23) points out that the un-

derside h.w. has six ocelli which is not at all

normal for Y. asterope; all the Y. asterope I

have examined, have from two to four ocellae.

However, six ocelli is common in Y. granulosa

and the d.s.f. has minute spots as in Strand’s

uniocellata.

Externally Y. asterope can usually be sepa-

rated from the closely related species Y. yatta

sp. nova by the paler subapical ocellar area of

the f.w. and 32 antennal joints against 29 in

Y. yatta.

The male genitalia of Y. asterope are general-

ly thought to be easily recognisable by the up-

turned, truncate apex of the valva and squarely

placed, distal accessory plate. This however

could very well lead to mistakes, as there turns

out to be several species with more or less simi-

larly shaped valva. There is a type of Y. asterope

with the accessory plates of valva placed at a

rather sharp angle; it occurs in the Nairobi area

of Kenya and its status is rather dubious (geni-

talia figured and described below). The external

characters are almost exactly identical with

those of Y. asterope and due to lack of material,

I will not make any attempt to describe it here

as a species. Y. yatta has a similarly shaped val-

va, but more robust. There are a few more spe-

cies with valves approaching those of Y. aste-

rope, viz. Y. congoana, Y. condamini, Y. lamto

and to a certain extent Y. rhodesiana. Y. lamto,

however, has a differently shaped uncus. But

none of these have the truncate apex with

squarely placed accessory plate as in normal

Y. asterope, and the female genitalia differ.

Ypthima asterope asterope (Klug)

(figs. 1—5, 12—19, 100—102, 137145, pl. 1

figs. 1—8)

Hipparchia asterope Klug, 1832, pl. 29 figs. 11—14

(Aden).

Ypthima asterope ab. binucholata Strand, 1909: 112

(Sadani, Tanzania).

Ypthima asterope ab. biocelligera Strand, 1909: 112

(Sadani, Tanzania).

External characters (pl. 1 figs. 1—8). — An-

tenna male and female 32 to 33 joints, shaft

black, white ringed above and below; club grad-

ual, but distinct, smooth, 12—13 joints, nar-

KIELLAND: Ethiopian Ypthima 105

Ant v sv

Du bu

Figs. 13. Male genitalia of Ypthima asterope Klug. 1, dorsal aspect; 2, lateral aspect; 3, aedeagus (Apx ang,

appendix angulare; Ae, aedeagus; Ax p, accessory plate, terminal process of valva; C p, costal process; Df, dia-

phragma; Fa, phallobase; Jx, juxta; Sa, saccus; Tg, tegumen; Un, uncus; V, valva, clasper; Ve, vesica; Vin, vincu-

lum; V n p, valva narrow portion; V w p, valva wide portion). Fig. 4. Female genitalia of Ypthima asterope

Klug, ventral aspect (Ant v sv, anterior wall of sinus vaginalis; A pl, anterior genital plate; Apo po, posterior

apophyse; A p pl, anterior part of posterior genital plate; C, colliculum; Corp bu, corpus bursae, bursa; Du bu,

ductus bursae; L sc, lateral sclerites; O bu, ostium bursae; Pap a, papilla analis, anal lobes, ovipositor lobes; P pl,

posterior genital plate; S v, sinus vaginalis, genital chamber).

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

rowly and finely double grooved, last joint as

long as penultimate; palpi second joint curved,

twice as long as last joint.

Male. — Upperside wings fuscous brown

(faded in old specimens); f.w. subapical ocellar

area clearly defined, rounded and usually much

paler than the ground colour, with a well de-

fined, dark brown outer border, a somewhat in-

distinct inner border line; the submarginal area

is paler than the ground colours a rather wide,

dark margin; cilia basally pale, then brown and

white tipped. H.w. with an ocellar spot in 2,

sometimes also in 1b (w.s.f.); there is no clear

discal line, a faint submarginal line, incurved in

4 and 5 and a dark marginal line; cilia as in the

f.w., but slightly paler. Underside ground col-

our whitish and striation rather spaced, giving a

pale appearance, particularly in the h.w.; f.w.

ocellar area roundish, reaching middle Da space

2; a thin, black marginal line; h.w. a double or

single spot in 1b, one in 2 and in most cases one

in 6. The spots of the w.s.f. are larger than those

of the d.s.f.; there is an uneven discal line,

which generally is clearer in the d.s.f., excurved

at vein 5; there is a submarginal line, almost

straight from the tornal spot to vein 3, thence

wavy and indistinct to near apex; a black mar-

ginal line. Length of f.w., 15 to 17 mm, antenna-

wing ratio 0.43.

Female. — Ground colour only slightly paler

than in the male; the ocellar area as in the male;

sometimes a faint indication of a h.w. discal line.

Length of f.w., 16 to 18.5 mm, antenna-wing ra-

uo 0.39.

Male genitalia (figs. 1—3, 12—19). — Prep.

nos. 393, 908, 923, 713A, 2275—77 from Soma-

lia; 394, 597, 598, 862, 2281, 2284 from Kenya;

1842K, 2267, 2268 from Sudan; 715 from Sene-

gal; 730A, 762A, 2047 from Angola; 726A from

Mozambique; 696A from Ethiopia; 1846K from

Arabia; 1848K from Lebanon; 1854K from Na-

tal, S. Africa; 1915 from Rhodesia; 2266 from

Amani, Tanzania. Dorsal aspect: Tegumen

wide, as long as uncus, proximal margin evenly

curved; valva, distal portion rectangularly

shaped, flattened dorsally, forming a sharp in-

ner (lateral) edge; outer margin evenly curved

from base to the discal rectangular part; acces-

sory plate upturned, at almost 90° angle to the

longitudinal axis of valva. Lateral aspect: Vincu-

lum short, strongly curved; valva robust, the

narrow distal portion thicker than usual in Yp-

thima, but much slenderer and longer than in

Y. yatta; distal portion truncate, upturned; ae-

deagus as long as valva.

Female genitalia (figs. 4, 100—102, 137—

145). — Prep. nos. 559, 6925) 9075) 909559228

711A, 714A, 715A, 1840K from Somalia; 712

from Senegal; 865, 716A, 717A, 1864, 1875

from Kenya; 706A, 707A from Ethiopia; 1845K

from Arabia; 1847K, 1946K, 1949K from Leba-

non; 1947K from Trans-Jordan; 1948K from

W. Aden. Anal lobes triangular; sinus vaginalis

wide and shallow, evenly rounded; edge of an-

terior wall usually wavy; inside of the wall with

more or less sclerotized longitudinal ribs, a rec-

tangular frontal sclerotized area; anterior plate

small, longer than wide, bilobed distally; poste-

rior plate consisting of an unsclerotized frontal

tongue-like protuberance and a basal, wider,

but rather narrow, sclerotized portion, tapering

on both sides (this portion is rather variable).

The lateral sclerites are U-shaped; bursa

obliquely ovoid to pear-shaped. The oblique-

ness is obscured if bursa is turned 90°. In speci-

men no. 873 from Kiambitti Hill, Embu, Kenya,

the anterior plate is trilobed apically, the middle

lobe small and the frontal part of the anterior

wall unsclerotized, but otherwise corresponds

with normal Y. asterope.

As stated before, there is a form of Y. aste-

rope in the Nairobi area, Kenya, with genitalia

somewhat different from typical Y. asterope. It

can be described as follows:

Prep. nos. 598, 686A, 687A. Dorsal aspect:

The basal margin of tegumen with three conca-

vities; apart from the dorsal one there is one on

each side, making the outline wavy; valva more

slender, outer margin less curved, distal portion

more rounded and less square looking, although

somewhat flattened dorsad and the inner edge is

as sharp as normal; the apical accessory plate

placed more at an angle, about 70°. Lateral as-

pect: Uncus a little longer than tegumen, the

lateral emarginations of the basal part of tegu-

men discernible; valva more slender and the

narrow portion longer than normal; aedeagus

longer than valva, sinuous, with a blunt apex;

diaphragma with a slightly sclerotized patch

ventrad of anus.

Specimens belonging to this type, have so far

only been taken in the Nairobi area to Yatta in

Kenya and may very well constitute a distinct

species, but too little material is available to as-

certain this. The external characters are exactly

like those of asterope.

Note. — Strand’s Y. asterope ab. binucholata

and ab. biocelligera are almost certainly not

Y. asterope, but probably Y. granulosa. The

type series of ab. binucholata may constitute

KIELLAND: Ethiopian Ypthima 107

„ asterope asterope Klug

. asterope hereroica Grünberg

- congoana Overlaet

. yatta species nova

- antennata antennata van Son

. antennata cornesi subspecies nova

„ Wuattouxi species nova

- jacksoni species nova

Fig. 7. Distribution of Ypthıma species.

more than one species.

Habitat. — Rather arıd bush country.

Distribution (fig. 5). — Most of S.W. Asıa

and Africa excluding the more humid parts of

Central Africa and excluding S.W. Africa to

Cape Province. I have seen no records from

Zaire to S. Nigeria and Ghana, nor from Zam-

bia, Malawi, Botswana, Burundi, Ruanda,

Uganda and Mozambique except the southern-

most part of Mozambique. It certainly occurs in

Botswana, northern part of Uganda, further

north in Mozambique and possibly also in S.

Zambia and Malawi. From several of these

- rhodesiana Carcasson

Y. condamini condamini species nova

. condamini nigeriae subspecies nova

- simplicia Butler

- recta Overlaet

y. granulosa Butler

Y. lamto species nova

Fig. 8. Distribution of Ypthima species.

countries I have not seen very much material. In

spite of heavy collecting ın Tanzanıa I have nev-

er taken Y. asterope there. The only record ıs a

single male from Amani, Usambara taken by

IAE jackson:

Ypthima asterope hereroica Van Son

(figs. 5, 38—41, pl. 2 figs. 1, 2)

Ypthima asterope hereroica Van Son 1955: 156, fig.

165.

Ypthima asterope ab. hereroica; Grünberg 1910: 101

(Damaraland).

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

¢. pupillaris pupillaris Butler

. pupillaris obscurata subspecies nova

. impura impura Elwes & Edwards

Y. impura paupera Ungemach

Fig. 9. Distribution of Ypthima species.

External characters (pl. 2 figs. 1—2). — An-

tenna as in Y. asterope asterope; pattern of the

wings similar to *Y. asterope asterope, but un-

derside somewhat darker and slightly more un-

evenly striated giving a mottled appearance; un- -

derside h.w. discal band usually more roundly

excurved at vein 5 than in specimens from Natal

and Transvaal. Van Son (1955: 155) states that

in the nominate race the discal line “is strongly

angled outwards on vein M2”. This applies best

to specimens in eastern South Africa to Rhode-

sia, but the aggregates from northern Africa and

S.W. Asia do not as a rule have a strongly an-

gled discal band. In fact, a long series from So-

malia in the B.M. have all a curved discal band

as in ssp. hereroica. The aggregate from eastern

South Africa could possibly constitute a sepa-

rate race, but due to the variability of Y. aste-

rope as a whole, it is very difficult to point out

constant characters to separate the South Afri-

can aggregate from the nominate race.

Male genitalia (figs. 38—41). — Prep. nos.

1919—21, 1843K, 2044-46, 2369—70 from

S.W. Africa. Valva is slightly more robust than

in Y. asterope asterope, almost as in Y. yatta sp.

n.; the upturned apical portion usually shorter,

but in prep. 1843K this was not noticeably

shorter and it apparently varies; aedeagus ap-

proximately as long as valva; saccus short and

wide in dorsal aspect; the diaphragma with a

small triangularly shaped, thinly sclerotized

area below the anal opening. Van Son (1955:

156) says that there is no sclerotization. The

specimens I have examined had sclerotization,

O Y. praestans Dverlaet

% Y- diplommata Overlaet

Fig. 10. Distribution of Ypthima species.

but others could well be without.

Female genitalia. — Prep. nos. 1922, 1923

from S.W. Africa; 1918 from Cape Province.

Two preparations have normal genitalia as in

Y. asterope asterope, while no. 1918 has an aty-

pic posterior plate with bilobed posterior por-

tion.

Habitat. — Dry, open country.

Distribution (fig. 5). — Cape Province to

S.W. Africa and probably S. Angola.

Ypthima yatta spec. nova

(figs. 5, 20—23, 134, 135, pl. 2 figs. 36)

Differs from Ypthima asterope Klug and its

subspecies in the slightly broader wings, the less

distinct ocellar area, due to the colour ap-

proaching that of the rest of the wing; submar-

ginal border not paler than the rest of the wing;

the genitalia much more robust than in Y. aste-

rope asterope, but only a little more than in ssp.

hereroica Grünberg.

External characters (pl. 2 figs. 3—6). — An-

tenna 29 joints, (in Y. asterope asterope and

Y. asterope hereroica 32 joints); shaft white

ringed, dark brown below, blackish above; club

gradual, but rather thick as in Y. asterope, 10

joints, the 11th going over into the shaft, under-

side double grooved in full length of the joints,

end joint slightly longer than penultimate; palpi

fawn, second joint twice as long as third joint.

Male. — Upperside wings evenly brown (old

specimens); f.w. ocellar area hardly paler than

the ground colour, faintly indicated; a very thin

and faint, brown marginal line; cilia plain

KIELLAND: Ethiopian Ypthima 109

brown, (in Y. asterope basally whitish, then

brown and pale tipped); h.w. with one spot in 2,

an unclear submarginal line and a fine, dark

margin; cilia a little paler than that of the f.w.

Underside darker and more uniformly and

closely striated than in Y. asterope; h.w. discal

band shaped as in Y. asterope; a double spot in

ib, one in 2 and one in 6; the outer margin an-

gled at vein 4, rounded in Y. asterope. Length of

f.w. 16 to 17 mm, antenna-wing ratio 0.40.

Female. — As in the male with rather

obscured inner part of the ocellar area, but a

little larger with broader wings; ground colour

paler. Underside as in the male with uniform,

close striation; slightly paler. Length of f.w.

16.3 to 17.5 mm.

Male genitalia (figs. 20—23). — Prep. nos.

517, 863, 864 from Kenya; 704A, 712A from

Ethiopia. Dorsal aspect: Rather similar to Y. as-

terope hereroica Grünb., but more robust; distal

part of valva very wide and short. Lateral as-

pect: Tegumen-uncus as in Y. asterope; valva

more robust; the narrow portion very thick,

particularly the proximal half; aedeagus much

longer than valva, sinuous as in Y. asterope, bas-

al open portion (phallobase) constricted in the

middle and thick at base, apical portion bent

ventrad, the tip upturned, tapering to a point;

diaphragma without sclerotization.

Female genitalia (figs. 134— 135). — Prep.

nos. 741, 720A, 734A, 1954K from Ethiopia. Si-

nus vaginalis rather wide and shallow; anterior

wall ribbed, with an oblong frontal sclerotiza-

tion; lateral sclerites broad, V-formed; anterior

plate very small, crumpled, almost like that of

Y. simplicia, but sclerotized; posterior plate

large, bipectinate distad, not quite symmetrical;

the sclerotized part large; a smaller, unsclero-

tized frontal part; anal lobes large, without apo-

physe; bursa very long and narrow, tapering

proximad, with no distinct junction with duc-

tus, quite distinct from other Ypthima species

dealt with here.

Habitat. — Not known to author, but proba-

bly similar to that of Y. asterope, in rather arid

country.

Distribution (fig. 5). — Thika in Kenya and

south-central Ethiopia.

Holotype d: Kenya, Thika, Yatta, 3.v.1942,

N. Mitton. Allotype 2: Ethiopia, Sheikh Hus-

sein, Upp. Schobehli R., 20.ix.1894, Don Smith.

Paratypes: Kenya, Thika, Yatta, 3.v.1942, N.

Mitton, 1 d; same data, but 13.v.1952, R. H.

Carcasson, 1 d; Ethiopia, Arussi Galla, Daroli,

13.111.1901, C. V. Erlanger, 2 d ; Ethiopia, Gen-

@ >. albida albida Butler

WM Y. albida uniformis Bartel

@ :. 6: occidentalis Bartel

X x. pulchra Overlaet

Fig. 11. Distribution of Ypthima species.

ale river, 16.1v.1901, C. V. Erlanger, 1 d; Gur-

ra, Dagaje, 4—5.1v.1901, C. V. Erlanger, 1 ®;S.

and S. E. Ethiopia, 1909—130, R. E. Drake-

Brookman, 1 ®. Holotype and two male para-

types in the NMN; allotype and six paratypes in

the BM.

Ypthima congoana Overlaet stat. nova

(figs. 5, 24—27, 103, 146, pl. 8 figs. 1, 2, 4, 5)

Ypthima asterope congoana Overlaet, 1955b: 85, figs.

12a, 12b (Upemba, Katanga).

A male and a female paratype were kindly

lent to me by Dr. L. Berger from Musée Royal

de L’Afrique Centrale, Tervuren, Belgium. In

all, two paratypes and one additional specimen

from B.M. have been examined.

Y. congoana can safely be raised to specific

status. The male and female genitalia differ from

those of Y. asterope. The three specimens exam-

ined are larger than any of the Y. asterope of the

corresponding sex I have seen.

External characters (pl. 8 figs. 1—2, 4—5). —

Male. Antenna with 32 joints, brown, speckled

and ringed white; club gradual, distinct and

long, consisting of 14 joints; (the antennal club.

of the paratype is aberrant with a serrated club

and abnormally long end joint); second palpal

joint three times as long as the end joint. Upper-

side f.w. ground colour dark brown; ocellar

area and costa paler and clearly defined; the area

from the hind margin, nearly up to vein 2 slight-

ly paler than the ground colour. This is not pro-

nounced in the second male. There is a dark

110 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

18 ze

Figs. 12—27. Male genitalia of Ypthima species. 12—15, Y. asterope asterope Klug, typic; 12, dorsal aspect; 13,

lateral aspect; 14—15, aedeagus in dorsal and lateral aspect. 16—19, Y. asterope asterope Klug, atypic; 16, dorsal

aspect; 17, lateral aspect; 18—19, aedeagus in dorsal and lateral aspect. 20—23, Y. yatta species nova, Kenya;

20, dorsal aspect; 21, lateral aspect; 22—23, aedeagus in dorsal and lateral aspect. 24—27, Y. congoana Overlaet,

paratype, Shifumanzi, Katanga; 24, dorsal aspect; 25, lateral aspect; 26—27, aedeagus in dorsal and lateral as-

pect.

KIELLAND: Ethiopian Ypthima ili

marginal line; cilia, greyish white with a brown

median line; h.w. ground colour as the f.w., but

evenly distributed over the entire wing, perhaps

slightly paler in the submarginal and the discal

section; there is a faint discal and a clear sub-

marginal line; an eye spot in 2. Underside of the

wings greyish-white, closely and evenly striated

brown; f.w. basal area darker in the paratype

(not in the second specimen); the pale ocellar

area clearly defined, the inner border thick, the

outer thin; a fine dark margin; h.w. evenly

striated, an indistinct discal band angled distad

between vein 4 and 5, almost in a straight line

from area 4 to the hind margin and from area 4

to costa; a double ocellar spot in 1b, one in 2

and one in 6; those in 2 and 6 of equal size, an

indistinct submarginal band and a fine marginal

line. Length of f.w. 17 mm in the paratype, 17.5

in the other male, antenna-wing ratio 0.47;

higher than in Y. asterope.

Female. — Antenna with 35 joints; club 12

joints, the last a little longer than penultimate,

conical. Upperside ground colour paler than in

the male; f.w. brown and with no darker basal

and median area; the ocellar area very clearly

defined, surrounded by a dark brown border; a

fine dark marginal line; cilia white with a thick,

brown median line; h.w. with a faint discal line

shining through from beneath (almost as in

Y. rhodesiana); an irregular submarginal line,

commencing almost at tornus, but turning

slightly interiorly at vein 2; an ocellar spot in 2.

Underside greyish, paler than in the male. (Both

males are w.s.f., the female d.s.f.) F.w. ocellar

spot and its area as in the male; the striation

brown, in the h.w. nearly black, but very fine; a

very fine, brown outer marginal line, followed

by a thicker black line; the two lines separated

by a white line; cilia as upperside; h.w. with

three tiny eye-spots, a double in 1b, one in 2

and one in 6; the two marginal lines not so clear

as in the f.w. (seen through a lens); the discal

band as in the male, but more sinuate. Length of

f.w. 19 mm, antenna-wing ratio 0.45.

Male genitalia (figs. 2427). — Prep. nos.

850 and 1825K. Dorsal aspect: Valva more slen-

der than the atypical form of Y. asterope from

the Nairobi area and much more slender than

normal Y. asterope; the distal end and processes

as in Y. rhodesiana, but uncus narrower; appen-

dix angularis of tegumen sharply bent interiorly

at midway from its base. Lateral aspect: Tegu-

men a little longer than uncus, in Y. asterope

shorter or equal; valva distal half narrow, only

slightly flattened dorsad close to the apical pro-

cess; basal end short, with a dorsal bulge.

Female genitalia (figs. 103, 146). — Prep. no.

851. Genital chamber deep, square anteriorly, in

Y. asterope shallow and rounded, no sclerotiza-

tion of the frontal area; anterior plate totally

different from that of Y. asterope, rounded,

rather large and broad with finely serrated edge,

roughish surface, poorly sclerotized; posterior

plate of the same type as in Y. asterope, with a

posterior, sclerotized broader part and an ante-

rior tongue-like unsclerotized lobe.

Habitat. — Not known to author, but proba-

bly same as that of Y. rhodesiana.

Distribution (fig. 5). — Katanga in Zaire.

Ypthima rhodesiana Carcasson

(figs. 6, 52—55, 105, 147—151, pl. 4 figs. 1—6)

Ypthima rhodesiana Carcasson, 1961: 14, fig. 11

(Choma, Zambia).

Related to Y.asterope, Y.congoana and

Y. condamini. Differs in the underside h.w. dis-

cal band showing through on the upperside; the

discal line much more evenly curved, almost

parallel to the outer margin.

External characters (pl. 4 figs. 1—6). — An-

tenna male 34 joints, female 30 to 31 joints;

shaft white below, blackish brown, sprinkled

with white and white ringed above; palpi sec-

ond joint two and a half times the length of the

end joint.

Wet season form. Male. — Upperside ground

colour variable, from pale to dark brown; f.w.

ocellar area clearly defined, paler than the

ground colour, with a dark border line all

around, outdrawn towards tornus, but not ex-

tended over vein 2; submarginal area paler than

the ground colour, margin darker; the median

area a little paler in some specimens; cilia brown

with a darker median line and paler tips; cilia of

the h.w. almost white in the distal half; h.w.

with a discal line, very clearly seen from be-

neath; a submarginal line, only slightly incurved

in space 4 to 5; a marginal dark brown line. Un-

derside ground colour pale gray, with even,

brown striation; there is a blackish brown mar-

ginal line in both wing pairs; h.w. with a promi-

nent discal line, almost evenly curved, but a

little irregular from near vein 1b to vein 5,

where it makes a gentle curve basad and with a

small bend before reaching costa; the whole line

is almost parallel to the outer margin; there is a

less distinct submarginal line; ocellar spots in

1b, 2 and 6 are small and mostly without pupils;

the spot in 2 nearly always with yellow ring, the

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Figs. 28—41. Male genitalia of Ypthima species. 28—33, Y. simplicia Butler, Largido (Longido?), Tanzania; 28,

dorsal aspect; 29, lateral aspect; 30—31, aedeagus of prep. no. 1832 in dorsal and lateral aspect; 32, aedeagus of

prep. no. 1831 in lateral aspect; 33, juxta. 34—37, Y. simplicia Butler, syntype, Ethiopia; 34, dorsal aspect; 35,

lateral aspect; 36, aedeagus, lateral aspect; 37, juxta. 38—41, Y. asterope hereroica Grünberg, S.W. Africa; 38,

dorsal aspect; 39, lateral aspect; 40, aedeagus, lateral aspect; 41, juxta.

KIELLAND: Ethiopian Ypthima 113

ones in 1b and 6 often without yellow ring and

very small. Length of f.w. 15.8 to 18.5 mm, an-

tenna-wing ratio 0.47.

Female. — Ground colour and markings as in

the male; wings slightly broader; h.w. tornal an-

gle more rounded, angled in the male. Length of

f.w. 17 to 18.5 mm, antenna-wing ratio 0.44.

The d.s.f. exhibits very little difference from

the w.s.f. The ocellar spots are reduced to tiny

black points, often difficult to detect with the

naked eye; the upperside ground colour is

sometimes a little paler, particularly the median

area in the f.w.

Male genitalia (figs. 52—55). — Prep. nos. 12,

154160190104 429, 43234, 436, 441, 544,

546, 549— 50, 552, 2007 from Mpanda district,

Tanzania; 868 from Kenya; 742A, 758A, 1839K

from Rhodesia. Dorsal aspect: Uncus distal

portion rather robust and blunt; accessory

plates placed at the apex of valva, distal portion

only slightly flattened as in Y. congoana.

Lateral aspect: Uncus shorter than tegumen;

vinculum strongly bent as that of Y. asterope;

valva basal half rather robust, more than the

length of the distal narrow portion; aedeagus as

long as, or slightly shorter than valva.

Female genitalia (figs. 105, 147—151). —

Prep. nos. 443, 449, 450 from Mpanda, Tanza-

nia. Sinus vaginalis evenly rounded, wide, but

not very deep, edge of anterior wall slightly

wavy; lateral sclerites U-shaped; anterior plate

small, bladelike, rounded, covered with fine tu-

bercles, emarginated distad; posterior plate con-

sists of a wide, cup-like and bipectinate heavily

sclerotized posterior portion and a narrower,

unsclerotized anterior part, formed as a tongue-

like lobe; ductus wide and short; bursa rather

large and longitudinally oval.

Note. — The male genitalia of Y. rhodesiana

and those of Y. congoana are closely related,

but the female genitalia show distinct and con-

stant differenses from Y. congoana in the ante-

rior plate. Moreover, Y. congoana is a very local

species.

Habitat. — Brachystegia woodland. Often

flying together with Y. condamini sp. n.

Distribution (fig. 6). — From southern part

of Kenya to Tanzania, Zambia, Rhodesia and

south Zaire.

Ypthima simplicia Butler

(figs. 6, 28—37, 127, 128, 173178, pl. 2 figs. 7,

8, pl. 3 figs. 1—6)

Ypthima simplicia Butler, 1876: 481 (Ethiopia).

In the BM collection there are three males

corresponding with Butler’s Ypthima simplicia.

One is labelled by him and is from the type lo-

cality. The two others are from Godman-Salvin

collection, 1902—116, labelled “Abyssinia”. A

fourth specimen, also labelled by Butler and

named Y. simplicia, belongs to Y. impura. This

one and the other one labelled by Butler had

previously been selected as syntypes of Y. sim-

plicia (unpublished). However, as the first three

specimens correspond best with Butler’s de-

scription, I reject the one that belongs to Y. im-

pura as syntype and select the other specimen

labelled by Butler as lectotype. I have dissected

the lectotype myself; the two other specimens

had been dissected before and mounted dry, but

they are good enough to be properly examined

and undoubtedly they belong to the same spe-

cies as the specimen which I have selected as

lectotype of Y. simplicia. Any female type can-

not be traced.

Description of the lectotype. External charac-

ters (pl. 2, figs. 7—8). — Antenna missing. Up-

perside ground colour greyish-brown; probably

dark brown in fresh specimens; f.w. ocellar area

almost as dark as the ground colour, encircled

by a darker line; ocellar area slightly prolonged

towards tornus; a pale yellow ring around the

black, blue pupilled ocelli; a dark marginal

stripe; cilia of both wings pale grey with a

brown middle line and paler tips; h.w. uniform-

ly brown without eye spots. (The two other

specimens have an ocellar spot in area 2. This

and the paler ocellar area of the f.w. are the only

differences between the type and the two other

specimens.) There is a fine, dark marginal line as

in the f.w. Underside greyish white with

somewhat uneven, brown striation; f.w. ocellar

area as the rest of the wing and encircled by a

brown ring slightly prolonged towards tornus;

there is a clear, black marginal line; h.w. with

two eye spots, one in area 2 and one in 6, almost

of the same size; a faint discal line, more visible

near costa as a blotch; a black marginal line as in

the f.w.

Genitalia (figs. 34—37). — Prep. no. 736A.

Tegumen-uncus resembling that of Y. rhodesia-

na, rather long and evenly tapering; valva slen-

der, outer half of distal narrow portion gradual-

ly widening; accessory plate turned in and

placed at an angle, approximately at 70° to the

longitudinal axis of valva; aedeagus longer than

valva; distal portion slender; apophyse angulare

bent interiorly.

114 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

4 x

Dtm EN D m

NEN > arti

NN ER

y erg

Figs. 42—45. Male genitalia of Y. condamını species nova. 42, dorsal aspect; 43, lateral aspect; 4445, aedea-

gus, dorsal and lateral aspect. Fig. 46. Female genitalia of Y. condamini species nova.

In the BM collection there is a series from

Ethiopia which corresponds with Y. simplicia,

and I have examined rather fresh specimens col-

lected in Ethiopia by S. C. Collins. As many of

these specimens are in better condition and

much fresher than the lectotype, it seems appro-

priate to describe them here.

External characters (pl. 3 figs. 1—4). — An-

tenna 29 joints, last joint as long as, or slightly

longer than penultimate, creamy on the under-

side, brown on the upperside; palpi, last joint

two and a half times the length of second joint.

Wet season form. Male. — Upperside ground

colour sooty brown in fresh specimens, other-

whise like lectotype. H.w. with an ocellar eye

spot in 2; a dark submarginal line, incurved be-

tween vein 3 and 5; a fine dark marginal line.

Underside ground colour of both wings rather

dark grey with dark brown, evenly distributed

striation, making the underside look darker than

usual in the Ypthima here dealt with, almost as

in Y. recta. The discal line in the h.w. is irregu-

lar and more or less indistinct; a black marginal

line which is indistinct and patchy. Length of

f.w. 17 mm, antenna-wing ratio 0.44.

Female. — Larger than the male; upperside

ground colour slightly paler; ocellar area a little

larger, otherwise the same markings. Underside

of specimens from the same season as dark as in

the males and with same markings, but ocellar

spots smaller and in some there is also a double

spot in 1b. Length of f.w. 17.8 to 20.5 mm, an-

tenna-wing ratio 0.38.

Dry season form. Male. — Upperside as the

w.s.f. Underside f.w. differs little from the

w.s.f., but h.w. usually with better developed

markings and with much smaller ocellar spots;

the discal line sharply produced at vein 4 and 5,

incurved between 2 and 4, somewhat similar to

Y. asterope from Natal. Length of f.w. 15.6 to

16.3 mm.

Female. — Little different from the w.s.f.

Length of f.w. 15.5 to 16.5 mm.

Male genitalia (figs. 28—33). — Prep. nos.

697A—701A, 710A, 1741 from Ethiopia;

1822K, 1831K, 2271K from N. Tanzania;

1844K, 2273K, 2283 from Kenya. Dorsal as-

pect: Somewhat resembling an atypic form of Y.

asterope, but accessory plate of valva strongly

inclined, quite variable, but usually about 45° to

50° to the longitudinal axis of valva; fultura U-

formed. Lateral aspect: Uncus as long as, or

longer than tegumen; narrow portion of valva

longer than wide portion; inner edge of narrow

part very sharp, extending from inner edge of

accessory plate almost the whole length of the

narrow portion; aedeagus as long as, or longer

than valva, straight dorsad, but distal half nar-

rowed ventrally; vinculum moderately curved,

less than in Y. asterope; apophyse angularis bent

interiorly. The accessory plate of the valva of

the lectotype is less strongly inclined and the

KIELLAND: Ethiopian Ypthima 115

sharpness ot the inner edge of the narrow part

of valva less pronounced and less extended.

However, the specimens examined vary quite a

lot with regard to these features.

Female genitalia (figs. 127—128, 173—178).

— Prep. nos. 703A, 708A, 709A, 1743, 1832K,

1953K from Ethiopia; 2274 from Kenya; 2272K

from N. Tanzania. Sinus vaginalis rather wide

and deep with a rounded, frontal sclerotization;

lateral sclerites widely U-shaped, broadening

laterally; anterior plate heavily dentate laterally,

bilobed distally and with a rather crumpled-

looking middle section; posterior plate with a

frontal, unsclerotized part as usual and a poste-

rior, lightly sclerotized part, shaped somewhat

as in Y. asterope, but more heavily bilobed; anal

lobes with small apophyses and uneven edges of

9th tergite; bursa obliquely ovoid.

Habitat. — Montane grassland and bush,

according to S. C. Collins while collecting in

Ethiopia. I have myself taken it in similar hab-

itat at Ngong hill, near Nairobi in Kenya.

Note. — Butler (1888: 59) records Y. simpli-

cia from Wadelai, Equatorial Africa, but this is

doubtful and possibly a misidentification. Speci-

mens not available.

The specimens from Longido and W. Kili-

manjaro, Tanzania, and from Kenya differ

somewhat from the Ethiopian specimens; with

paler and more unevenly striated undersides and

clearly defined discal and submarginal lines. The

Kenya-Tanzania aggregate may belong to a sep-

arate race, but a very long series of specimens

from both seasons is needed for description.

The Tanzania-Kenya specimens are d.s.f.

Distribution (fig. 6). — Central and South

Ethiopia, South Kenya, North Tanzania, and a

few records from Sudan.

Ypthima condamini spec. nova

This new species is quite common in Mpanda,

Kigoma and Ufipa in Tanzania and has also

been collected by the author about 20 miles

north of Tabora, Tanzania. In the NMN and

BM there are specimens from Kenya and Ugan-

da and in the BM also from Malawi, Zambia, a

record from West Zaire and two from Angola,

all of which belong to the nominate race. In the

Y. asterope collection sent to the writer on loan

by Dr. Vari (TMSA) there was one male Y. con-

damini condamini taken and dissected by Van

Son from Wolkberg farm, Lotaba Drift, Trans-

vaal.

Apparently the species is widely distributed

in Africa and locally common, but it has often

been confused with the similar Y. asterope. The

West African specimens are different from

those of other parts of Africa and are distin-

guished as a separate subspecies.

Y. condamini frequently flies together with

Y. rhodesiana, but can be separated from it by

the underside h.w. discal line which is angled

distad in space 4 to 5; in Y. rhodesiana the discal

line is evenly rounded and almost parallel to the

outer margin. The only certain means of sepa-

rating it from Y. asterope is by examining the

genitalia. This can be done just by brushing

aside some abdominal hairs to expose the valva

which differs from that of Y. asterope in the ac-

cessory plate placed at an angle and the valva it-

self with a dorsal convexity in the middle. This

feature is apparent in only one more species, Y.

lamto.

Ypthima condamini condamini subspec. nova

(figs. 6, 42—46, 166, 167, pl. 4 figs. 7—14)

External characters (pl. 4 figs. 7—14). — An-

tenna 33 joints, shaft brown, with a whitish,

longitudinal stripe; club gradual, only slightly

thicker than the shaft, thinner than the club of

Y. asterope, underside of each joint with large,

but shallow double grooves; the last joint longer

than the penultimate; palpi sprinkled with black

and white hairs, second joint two and a half

times as long as the last joint.

Wet season form. — Male. Upperside wings

fuscous brown, a little paler in the apical half;

f.w. ocellar area slightly paler than the ground

colour, bordered by a clearly defined, dark line,

most prominent distad; margin dark; cilia

brown, finely white checkered; h.w., a spot in 2,

a submarginal band, excurved in 4 and 5. Un-

derside ground colour whitish grey with brown

striation; f.w. ocellar area with a U-formed

brown border, thickest below the eye-spot (the

ocellar area varies individually in size and

form); h.w. with one ocellar spot in 2 and a

smaller one in 6; in one specimen from Sibweza,

Mpanda, Tanzania, the spots are equal in size;

there is a faint submarginal line and sometimes a

discal and subbasal band. Length of f.w. 15.8 to

17 mm, antenna-wing ratio 0.44.

Female. Considerably larger than the male,

with more rounded wings; slightly paler ground

colour; f.w. ocellar area more clearly defined

and paler. Length of f.w. 18.6 mm, antenna-

wing ratio 0.39.

Dry season form. — Upperside similar to that

of the w.s.f., except that the males have no spots

in the h.w.; 50% of the females have a spot in

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

space 2, the rest none. Underside h.w. very dif-

ferent from that of the w.s.f.; eye spots reduced

to tiny black points, male visible without a

lens; there is a faint subbasal line and a promi-

nent discal band; the discal band is clearer in the

female; the submarginal line is hardly clearer

than in the w.s.f. Length of f.w. male 15.5 to

16.5 mm, female 18.5 mm.

Male genitalia (figs. 42—45). — Prep. nos. 1,

lil, 17% 18, 24, 25, 229, 2890, 491, ZON Did, Dro

820, 821, 823, 1322, 1509, 1666, 1929, 2006

from Tanzania; 861, 2278K from Uganda;

722A, 723A from Malawı; 693A, 725A from

Sudan; 731A from Angola; 1415, 1423 from W.

Zaire. Dorsal aspect: Tegumen-uncus narrow-

ing rather evenly to the blunt distal apex of un-

cus; the uncus is more solid than in Y. asterope

and Y. yatta; valva robust, the narrow portion

swung out and the apical accessory plate placed

at a sharp angle, 45°. In Y. asterope the narrow

portion continues the convex bend of the first

part of valva and the shape becomes quite dif-

ferent from that of Y. condamini. Lateral as-

pect: Vinculum straighter than in other conti-

nental African Ypthima (in Y. asterope the vin-

culum is much shorter and very bent); uncus

rather straight and robust, the narrow portion

much shorter than the proximal, broad part,

about one quarter of the length of valva; valva

robust, distal narrow portion wider than nor-

mal, with a dorsal convexity, the shape close to

valva of Y. lamto sp. n.; aedeagus longer than

valva; saccus rather narrow.

Female genitalia (figs. 46, 166—167). — Prep.

nos. 447, 448, 822, 824, 825, 1321 from Tanza-

nia; 870 from Uganda; 694A, 695A from Sudan;

724A, 1838K from Malawi; 1827K from Zam-

bia; 718A from Kenya; 729A from Angola.

Very unlike Y. asterope. Anterior wall of sinus

vaginalis reinforced with strong, longitudinal

ribs and an anterior sclerotized plate with even

edges; the rest of the wall’s margin is wavy;

there is a large, rounded and hairy anterior plate

with a slight emargination at the distal end; a

thinly sclerotized posterior plate, consisting of

two parts, an anterior, unsclerotized tongue-

like lobe and a slightly sclerotized and strongly

wavy posterior plate. This is entirely fused to

the likewise strongly wavy lateral sclerites, thus,

with these forming a wavy wall or ribbon which

is quite unlike the situation found in related spe-

cies; ostium small; ductus short and bursa rath-

er small, slightly obliquely shaped.

Habitat. — Brachystegia woodlands of higher

and medium altitudes. In Tanzania found up to

2000 m as at Ufipa highlands and at about 700

m at Ruaha near Iringa. The flight is low and

skipping between grasses.

Note. — In the d.s.f. there are two types of

“wing shapes, both in the male and the female.

The commonest is the narrow type, but often

there is also a type with considerably broader

wings. The genitalia are constant in all and the

two forms have been taken in the same habitats,

so there is no possibility of subspeciation. The

difference in the wing shape has also been men-

tioned by Overlaet in his description of Ypthi-

ma (asterope) congoana. It seems that the nar-

row wing type is confined to the d.s.f.

Distribution (fig. 6). — From North Ethiopia

and South Sudan to East Africa, Zambia, Mo-

zambique and Transvaal, and some records

from West Zaire and North Angola.

Holotype d: Tanzania, Mpanda, Sibweza,

16.11.1970, J. Kielland; Allotype 9: Same data

and collector, 4.1.1970; Paratypes: Same, but

31.vu.1961, 12.1.1970, 6.1.1970 and 28.11.1969, 5

d; same, but Kanindi (Rukwa Valley), 5.x.1970,

2 6; Kigoma, Mgambo, 9.v1.1968, 1 6; Mpan-

da, Mbaya, 23.1x.1970, 1 2; Mpanda, Sibweza,

30.vi.1968, 1 ®; Mpanda, Katuma, 2.11.1967, 1

SEM panda Ikakay 71x 1978, Saar

land; Uganda, Karamoja, Labwor Hills, Aremu,

111.1952, T. H. E. Jackson, 1 25 sameybur West

Madi, Metu, vi.1954, 1 ©. Holotype and allo-

type in NMN; paratypes to NMN, BM and J.

Kielland collection.

Ypthima condamini nigeriae subspec. nova

(fig. 6, pl. 4 figs. 15—18)

External characters (pl. 4 figs. 15—18). —

Dry season form. Male. More pointed f.w. and

smaller than both the dry and wet season forms

of the nominate race; the upperside ground col-

our is considerably paler. Both the pointed

wings and ground colour resemble the paler

forms of Y. rhodesiana. The ocellar area is very

distinct and bordered by a dark line; h.w. in

some specimens with, and others without ocel-

lar spot in area 2. Underside pale as in Y. conda-

mini condamini; h.w. discal, subbasal and sub-

marginal lines very sharply defined; the subba-

sal line more parallel to the discal line than in Y.

condamini condamini, making an outward,

sharp bend in the cell near the base of vein 2,

corresponding with the bend in the discal line.

This subbasal line is evenly rounded in the nom-

inate race, when present. Length of f.w. 14.9 to

15.4 mm, antenna-wing ratio 0.44.

Female. Upperside as in the male, but larger;

KrerLAND: Ethiopian Ypthima 117

Figs. 4762. Male genitalia of Ypthima species. 47—51, Y. antennata van Son; 47, dorsal de en

aspect; 49, aedeagus, prep. no. 536; 50, valva, prep. no. 875; 51,valva, prep. no. 858, Sudan.52 55, é A en

na Carcasson; 52, dorsal aspect; 53, lateral aspect; 54—55, aedeagus, dorsal and lateral aspect. 56—5 id ua

touxi species nova, paratype, Lamto, Ivory Coast; 56, lateral aspect; 57, dorsal aspect. Las Ye iss DE ue

cies nova, holotype, Mtwapa Creek, Mombasa, Kenya; 58, dorsal aspect; 59, lateral spas GO 2, Y. lamto

species nova, Lamto, Ivory Coast; 60, dorsal aspect; 61, lateral aspect; 62, aedeagus, lateral aspect.

118 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

underside a little more suffused, with subbasal

line less sharp. Length of f.w. 17.2 mm, anten-

na-wing ratio 0.40.

Wet season form. Male. Wings usually slight-

ly more rounded than in the d.s.f.; underside

paler. Length of f.w. 16 mm, antenna-wing ratio

0.43.

Distribution (fig. 6). — From Nigeria to

Ghana, Upper Volta and Senegal.

Holotype d: Nigeria, Farniso near Kano,

BADOO, Buena Cet, Albogpe 2:

Same data and collector. Paratypes: Same, but

20) gravel 25 Degene IID, 2 ©, esus Niger,

Borgu, Yelwa, 2.vi1.1899, Wilson, 1 d, w.s.t.;

Senegal, Badi, Niokolo Koba, 22.x1.1959 and

15—25.vii.1955, Mission IFAN, 2 8; same, but

18.1.1965, 1 ©. Holotype, allotype and three

paratypes in the B.M. collection; two males and

one female paratypes in IFAN, Dakar.

Ypthima antennata Van Son

Antenna longer than in other continental Af-

rican species and recognizable by the abrupt,

short club; the club is of variable length, in

some specimens almost circular, in others ob-

long, deeply grooved on the underside; palpi:

third joint long and thin, second joint only one

and a half times the length of the third joint.

Ypthima antennata antennata Van Son

(figs. 7, 47—51, 104, 152—155, pl. 5 figs. 1—4)

Ypthıma antennata Van Son, 1955: 156, pls. 5—6 figs.

86, 87 (Munnik, N. Transvaal).

External characters (pl. 5 figs. 1—4). — An-

tenna male 31 to 37 joints, female 32 joints,

black and white ringed. Wet season form. Male.

Wings, upperside uniformly dark greyish

brown in fresh specimens; f.w. ocellar area

slightly paler than the ground colour, but with a

distinct, darker border all the way around, in

most specimens not reaching vein 2; margin

finely black bordered; cilia greyish with a dark

median stripe; h.w. with a very faint, uneven

submarginal band close to the margin, excurved

in 4 and 5; some specimens without any trace of

eye spots, others with one spot in 2. Underside

ground colour whitish grey with uneven, brown

striation; f.w. basal portion a little paler than the

rest of the wing; the ocellar border distinct;

h.w. with two ocellar spots of even size, in 2

and 6, sometimes also one in 1b; a subbasal

band, thickened and protruding distad as a W in

4 and 5; a submarginal band, rather far from the

margin; no lines between this and the margin,

only normal striation.

Dry season form. Very little difference from

the w.s.f.; underside eye spots in the w.s.f. only

slightly larger. Length of f.w. 14.3 to 17 mm,

antenna-wing ratio 0.47.

Female. Antennal club more oblong than that

of the male. Underside wings slightly paler than

in the male; f.w. subapical eye spot larger, more

rounded and clearly defined, reaching vein 2;

h.w. markings as in the male, some without and

others with an ocellar spot in 2. Underside f.w.

ocellar area and its border as on the upperside;

h.w. markings as in the male. Length of f.w. 18

to 19 mm, antenna-wing ratio 0.45.

Male genitalia (figs. 47—51). — Prep. nos. 9,

346, 537, 783 from N. iIlanzania: 97089870556!

547, 555, 881 from W. Tanzania; 869, 1965

from Uganda; 858, 859, 751A from Sudan;

1865, 1866, 1912 from Kenya. Dorsal aspect:

Tegumen rather wide at base, narrowing to-

wards its junction with uncus, which is of nor-

mal width at base and basal half, then narrowing

to a slender distal portion; appendix angularis of

tegumen bent interiad; valva strongly curved in-

teriad in an even line from base to apex; a small

apical process. Lateral aspect: Dorsal outline of

tegumen concave in the middle; uncus slender

from base to apex; valva slender, broadest near

base, tapering to the sharply upturned distal

portion; aedeagus as long as valva, the basal

third thin, the middle section thick, then ta-

pering to a sharp, upturned point.

Female genitalia (figs. 104, 152—155). —

Prep. nos. 560 from Uganda; 1197, 1826K from

Malawi; 561, 784, 880 from Tanzania. Confu-

singly similar to Y. asterope. Sinus vaginalis

rather shallow, but slightly wider than in Y. as-

terope; the anterior wall evenly and broadly U-

- shaped; anterior plate small, longer than broad,

bilobed distally, but the shape slightly variable;

in the specimens from W. Nile, Uganda, its dis-

tal part is broadest, in the specimen from Sero-

nera, Tanzania, no. 784, it is parallel-sided; pos-

terior plate usually wider than that of Y. aster-

ope, but in asterope it also varies in width and

depth; like in Y. asterope there is a central, un-

sclerotized lobe; the lateral sclerites are U-

shaped and connected to the posterior plate;

ductus moderately long and wide; bursa slightly

larger than in Y. asterope.

Habitat. — Mainly open grassland with scat-

tered trees. Frequently taken in rocky places of

sandstone formations, settling on rocks.

KieLLAND: Ethiopian Ypthima 119

Figs. 63—78. Male genitalia of Ypthima species. 63—67, Y. recta Overlaet; 63—64, dorsal and lateral aspect of

prep. no. 849, paratype, Kafakumba, Katanga; 65—67, dorsal and lateral aspect, and aedeagus of prep. no. 724,

Lamto, Ivory Coast. 68—69, Y. granulosa Butler, atypic, Kakamega, Kenya; 68, dorsal aspect; 69, lateral aspect.

70—74, Y. granulosa Butler, coastal type, prep. no. 852, Kakamega, Kenya; 70, dorsal aspect; 71, lateral aspect;

72—73, aedeagus, dorsal and lateral aspect; 74, distal end of valva, prep. no. 1413, Kitwi, Kenya. 75—78, Y.

granulosa Butler, inland type, prep. no. 853, Kakamega, Kenya; 75, dorsal aspect; 76, lateral aspect; 77—78,

aedeagus, dorsal and lateral aspect.

120 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Distribution (fig. 7). — Widely distributed

over most of the eastern part of Africa; from

Transvaal in South Africa to Rhodesia, Mozam-

bique, Central Zambia, western and northern

part of Tanzania to West Kenya, Uganda and

South Sudan.

Ypthima antennata cornesi subspec. nova

(fig. 7)

Amongst a small collection kindly sent to me

for examination by Mr. M. A. Cornes, Lagos,

Nigeria, a male and a female proved to belong

to a distinct race of Y. antennata. While exam-

ining the collection of Ypthima in the BM, a few

more specimens from Nigeria were found.

External characters. — Antenna male 31

joints, female 30 joints; shaft and club as in Y.

antennata antennata.

Wet season form. Male. Differs from the

nominate race in the f.w. subapical ocellar area,

which is less rounded and more drawn out to-

wards tornus; tornus itself more sharply angled

and the outer margin stands almost perpendicu-

lar upon the inner margin (the outer margin is

much more sloping in ssp. antennata). H.w.

outer margin less convex, nearly straight; f.w.

submarginal line closer to the ocellar ring. Un-

derside, wings evenly striated; in ssp. antennata

the striation is very uneven in both seasonal

forms and the h.w. bands are much more clearly

indicated. The underside of ssp. cornest is very

close to that of Y. asterope; the eye-spots in 2

and 6 are very large and there is a small spot in

1b. Length of f.w. 14.8 mm, antenna-wing ratio

0.53; much higher than in the nominate race.

Female. Paler than the male and with more

rounded wings and larger eye-spots on the f.w.

Otherwise it differs from the nominate race as

does the male. Length of f.w. 15.5 mm, anten-

na-wing ratio 0.49.

Male and female genitalia. — As in ssp. an-

tennata.

Distribution (fig. 7). — Nigeria.

Holotype d: North Nigeria, Benue Plateau,

Jos, June 1968, M. A. Cornes. Allotype ®:

Same data and collector.

Notes. — I have the pleasure in dedicating

this new race to its collector, Mr. M. A. Cornes

of the NSPR Institute. Lagos, Nigeria. Holo-

type and allotype to NSPRI collection

While this paper was in press, additional re-

cords were received from Mr. Usher, York, of

the occurrence of this subspecies in Ghana (see

fig. 7).

Ypthima jacksoni spec. nova

(te VASE EE pl 3100829)

External characters (pl. 3 figs. 5— 8). — An-

tenna 29 joints; upperside shaft brown, sides

greyish-white and underside brown and grey

checkered; club gradual, distinct; the underside

of each joint with two narrow grooves, the end.

joint as long as penultimate; second palpal joint

one and a quarter times as long as the third

joint.

Male. Upperside, wings greyish-brown (fresh

specimens are most certainly darker); ground

colour and markings hardly distinguishable

from Y. asterope; the ocellar area clearly de-

fined and slightly paler than the ground colour;

costa striated; the section between the yellow

ring and submarginal line is paler; submarginal

line (ocellar border) narrow, dark brown; mar-

gin darker than the submarginal area; cilia

brown; the cilia of Y. asterope are basally whit-

ish, then brown and tipped white; h.w. uni-

formly greyish-brown; a small ocellar spot in 2;

a faint submarginal line and a dark margin. Un-

derside f.w. distinguishable from Y. asterope in

the entire area being almost uniformly striated;

the usually dark anal area, common in Ypthıma

species, is grey like the ground and with fine,

brown striation; h.w. paler and uniformly

striated with brown; without subbasal and dis-

cal lines; a very fine, mottled, dark brown mar-

gin; ocellar spots small, but entire; there is a

double pupilled spot in 1b, one in 2 and one in

6. Length of f.w. 14 to 14.7 mm, antenna-wing

ratio 0.45.

Female. Ground colour a little paler than in

the male; wings more rounded and larger. Up-

perside f.w. with a large ocellar area, larger than

in the male; a brown border surrounding the

area as in the male, but its proximal part near

costa forming a nearly 90° angle; costa striated

as in the male; h.w. with a small ocellar spot in 2

and a tiny one in 1b; a clear submarginal line,

rather irregular; a dark brown margin; cilia

brownish grey with a distinct, median line. Un-

derside f.w. striated as in the male, but the

brown border of the ocellar area as on the up-

perside, except the part parallel to costa which is

very thin and indistinct; a black marginal line;

h.w. not paler than the f.w.; there is a clearly

defined discal line almost parallel to the margin;

a less clear subbasal line; both lines fairly evenly

curved; near apex there is a blotch which is

placed on edge, more clearly seen on the right

wing; there is a submarginal line which looks

KIELLAND: Ethiopian Ypthima 121

uneven due to the striation; a double spot in 1b,

one in 2, 5 and 6. Length of f.w. 16 mm, anten-

na-wing ratio 0.38.

Male genitalia (figs. 58—59). — Prep. nos.

534, 596, 705A, 738A, 739A, 2270. Dorsal as-

pect. Tegumen-uncus gradually tapering to-

wards the distal end of the uncus; valva slender,

sinuate; apical accessory plate small, situated on

the innerside, pointing up and placed 45° to the

longitudinal axis of valva. Lateral aspect. Uncus

a little longer than tegumen; vinculum only

slightly bent; saccus short; valva furnished with

a large costal process; basal part of valva short

and broad, narrow portion very slender with

distal end upturned; aedeagus very robust, lon-

ger than valva, with the distal end blunt and up-

turned.

Female genitalia (figs. 125—126). — Prep.

nos. 735A, 2269. Peculiar; lateral sclerites short

with weak sclerotization; genital plates unscle-

rotized; anterior plate small, rounded and den-

tate along the edges; posterior plate formed like

a rounded sack; sinus vaginalis with a small,

oval frontal sclerotization; anal lobes compara-

tively large, without apophyses.

Habitat. — Judging from distribution, both

humid and dryer woodland formations.

Distribution (fig. 7). — Kenya; coast near

Mombasa, Teita; Ethiopia at Kata Mane river,

Darolli Arussi Galla.

Holotype gd: Kenya, Mombasa, Mtwapa

Creek, iv.1950, E. Pinhey; Allotype 2: Ethio-

pia, Kata Mane Riv, 27—31.111.1901, C. V. Er-

langer; Paratypes: Kenya, Samburu, 12.x.1911,

IEEE jackson, 18; British Central Africa,

Masongaleni, 26.1x.1911, no name, 1 d; Kata

Mane, 27.10.1901, C. V. Erlanger, 1 d: Ethio-

pia, 1909—130, B. E. Drake-Brookman, 1 d.

Holotype to NMN; allotype and paratypes to

I have the pleasure in naming this species af-

ter one of its collectors, the late Mr. T. H. E.

Jackson.

Ypthima vuattouxi spec. nova

(figs. 7, 56, 57, 110, pl. 5 figs. 5—8)

Four males and one female of this species

from Lamto, Ivory Coast, were sent to the au-

thor on loan by Dr. M. Condamin, IFAN, Da-

kar. Later I found several males and females in

the BM collection from Gambia, Nigeria and

Cameroun.

Diagnosis. — Closely related to Y. jacksoni.

Genitalia differ in valva being more evenly bent

and not sinuate; basal wide portion differently

shaped and uncus thinner. Differs from Y. lam-

to in upperside ocellar area being without, or

nearly without striation. Underside f.w. tornal

area not, or slightly darkened; female genitalia

differ strikingly, as shown in the figure.

External characters (pl. 5 figs. 5—8). — An-

tenna 33 joints, brown above, whitish below,

faintly ringed with white on the shaft; club

gradual, but clearly defined, consisting of 10

joints, 11th going over into the shaft; the under-

side of the joints double-grooved, the last as

long as the penultimate; palpi second joint near-

ly twice as long as the last joint, curved, fawn,

dorsal hairs mainly white.

Male. Upperside wings brown, slightly

speckled with grey, particularly the area around

the h.w. ocellar spot in 2, as seen through a lens;

f.w. ocellar area distinctly bordered with dark

brown; the ocellar and submarginal area paler

than the ground colour; in one paratype the ex-

tension of the black central area intrudes into

the pale yellow ring, forming a tiny eye spot

with trace of a blue pupil in space 3 (this is only

an individual variation); the ocellar area is fairly

evenly rounded with slight extension towards

tornus; cilia greyish with a median, brownish

stripe on both wings; h.w. with an ocellar spot

in 2 and a small, single pupilled spot in 1b; in

some specimens this is lacking or very faint; a

submarginal, dark band and a fine marginal dark

border as in the f.w.; the submarginal band is

slightly excurved in 4 and 5. Underside grey,

finely striated with brown in both wings; f.w.

ocellar area evenly striated as the rest of the

wing, bordered with brown, open towards cos-

ta, the dark border area extended towards tor-

nus; one large subapical ocellus, in one paratype

also a small, black pupilled one in 2; h.w. with

one double pupilled spot in 1b, one in 2 and one

in 6. Length of f.w. 14.5 mm, antenna-wing ra-

tio 0.46.

Female. Upperside rather similar to a small Y.

asterope; larger than the male; antenna 32

joints, end joint shorter than penultimate;

otherwhise like that of the male. Upperside

wings brown, slightly paler than in the male;

f.w. ocellar area larger and paler; costa granu-

lated; h.w. with one ocellar spot in 2; a faint,

dark submarginal band, slightly widened in 4.

Underside similar to that of the male, or slightly

more greyish and f.w. ocellar area a little larger;

h.w. submarginal band widened, slightly dark-

ened and excurved in 4; cilia greyish with a

prominent, brown median line. Length of f.w.

16 mm, antenna-wing ratio 0.42.

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Figs. 79—90. Male genitalia of Ypthima species. 79—82, Y. impura Elwes & Edwards; 79, dorsal aspect; 80,

lateral aspect; 81—82, aedeagus, dorsal and lateral aspect. 83—86, Y. pupillaris Butler, Nigeria; 83, dorsal as-

pect; 84, lateral aspect; 85—86, dorsal and lateral aspect of aedeagus. 87, Y. diplommata Overlaet, holotype,

Kafakumba, Katanga. 88, Y. praestans Overlaet, holotype, Tshiole, Sankuru, Zaire. 89—90, Y. doleta Kirby; 89,

dorsal aspect; 90, lateral aspect.

KIELLAND: Ethiopian Ypthima 123

Male genitalia (figs. 56—57). — Prep. nos.

697, 1233, 1234, (986 Condamin prep.) from

Ivory Coast; 1836K from Cameroun; 1841K

from Nigeria. Closest to Y. jackson: sp. n. Dor-

sal aspect: Tegumen wide basally, narrowing

towards uncus which tapers evenly into a long,

narrow apical portion; difters in this aspect

from that of Y. jacksoni; valva very long, slen-

der and evenly curved (not sinuate) from base

towards apex; just before apex it makes an out-

ward, sharp bend, angled on the innerside; the

small apical accessory plate is placed at an angle

of about 45° to the longitudinal axis of valva.

Lateral aspect: dorsal outline of tegumen evenly

curved, the portion above appendices angulares

narrowly constricted; uncus very slender; valva

much longer than the combined length of tegu-

men-uncus; the thicker basal part rather nar-

row, shorter than the length of the very slender

distal portion, aedeagus very long, as long as, or

longer than valva; the basal open part very slen-

der, abruptly thickening about 1/5 distance

from extreme base; the rest robust, almost par-

allel, except for the distal upcurved point; sac-

cus longer than in Y. jacksoni.

Female genitalia (fig. 110). — Prep. no. 987,

Condamin prep. from Ivory Coast. Sinus vagi-

nalis wide, not very deep; anterior plate very

wide and large, slightly emarginated at its distal

border, one and a quarter times as wide as the

posterior plate, which also is large, as wide as

long and trilobed distally; both plates are rather

weakly sclerotized. In Condamin’s preparation

it looks like the posterior plate consists of two

portions, the one described above and an ante-

rior, unsclerotized, tongue-like process, as in

most species of Ypthima. The ductus and bursa

are not visible; the anal lobes are rectangular

and evenly outlined as those of Y. asterope.

Habitat. — Savanna.

Distribution (fig. 7). — Lamto, Ivory Coast;

Ghana, Nigeria and Cameroun (records re-

ceived while this paper was in press).

Holotype d : Ivory Coast, Lamto (Toumodi),

8.11.1967, R. Vuattoux. Allotype 9 : Same locali-

ty, 12.1v.1965, S. Kwamé. Paratypes: 3 d from

same locality, 11.1.1968, R. Vuattoux;

25.11.1965, 29.11.1965, S. Kwamé.

Note. — The specimens are all typical w.s.

forms.

Ypthima lamto spec. nova

(figs. 8, 60—62, 136, 163—165, pl. 5 figs. 9—

14)

Dr. Condamin, IFAN, Dakar, kindly pre-

sented to the author some specimens from Ivory

Coast belonging to this species and later sent a

large series for examination. Later a male from

Cameroun was presented by Mr. J. Plantrou.

External characters (pl. 5 figs. 9—14). — An-

tenna 34 joints, (in the related Y. granulosa Btl.

30 joints), greyish with a dark, brown-grey

checkered line dorsally, ending at the last joint;

shaft rather thick, very gradually thickening in-

to the club which is broadest four joints from

the apex; the club is orange, except for the dor-

sal stripe; underside is double-grooved for near-

ly the full length of the joints; palpi second joint

two and a half times the length of the third joint,

rather straight.

Male. Upperside strikingly similar to Y. gran-

ulosa, particularly to the coastal form; f.w. fus-

cous brown, ocellar area outdrawn towards tor-

nus and thickly striated with dark brown, clear-

ly dark bordered on either side; submarginal

area slightly lightened with grey, a double mar-

ginal line, the outer dark brown, the inner less

clear; the subtornal area of same dark brown as

the submarginal line; cilia greyish brown with a

darker median line; the double pupilled sub-

apical ocelli with brownish-yellow ring. In

some males the ocellar area is of paler striation

and of larger extent towards margin and tornus.

Normally the ocellar striation reaches the mid-

dle of area 2 and the submarginal line is situated

in the middle between margin and the yellow

ocellar ring; in some specimens the ocellar stria-

tion reaches vein 2 and beyond; h.w. ground

colour as in the f.w.; the only markings are the

single ocellar spot in 2, a dark submarginal line,

excurved in 4 and 5 and a less clear, irregular

median line; the submargin is slightly paler; cilia

as in the f.w. Underside grey with dark, olive-

brown striation, almost as in Y. pupillaris; f.w.

ocellar area bordered with dark brown which

continues in the subtornal area, reaching tornus;

both wings with a black margin, bordered inside

by a line of the same shade as the greyish

ground colour; cilia grey with a dark median

line; h.w. with an uneven, dark submarginal

line, incurved and widened in 4 and 5; a fairly

clear, but uneven median line and an indistinct

subbasal line; four ocellar spots, two in 1b, one

in 2 and one in 6; the one in 2 a little larger, but

sometimes the one in 2 and 6 are equal and in

others the ocellus in 6 is larger, the two in 1b are

a little smaller. Length of f.w. 14.5 to 16 mm,

antenna-wing ratio 0.36.

Female. Ground colour paler than that of the

male, the whole surface more or less striated;

124 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

the f.w. ocellar area paler, striation more

spaced, extending further towards tornus; the

submarginal bands in both wings, the h.w. me-

dian band and the brown ocellar ring more

clearly defined; h.w. discal area slightly paler

than the basal area and striation clearly defined;

submarginal area pale and striated; underside

marked and striated as in the male, but striation

more spaced, giving it a much paler appearance.

Length of f.w. 16 to 17.5 mm, antenna- wing ra-

tio 0.37. Slightly higher ratio in the female than

in the male, which is unusual.

Dry season form. There is no typical d.s.f. in

the numerous specimens examined from Lamto.

Amongst the paratypes three males and one fe-

male approach this form with very small eye

spots on the h.w. underside, but are otherwise

similar to the w.s.f. I was informed by Dr. Con-

damin that Lamto is situated close to the rain

forest area with very short dry season and is,

therefore, unlikely to produce typical d.s.

forms.

Male genitalia (figs. 60—62). — Prep. nos.

EO 695, G99; 729, 1205), 1247, 1240, 161);

1276, 1278 from Lamto, Ivory Coast; 1726

from Cameroun. The whole appearance of the

genitalia similar to that of Y. recta. Tegumen

and uncus also resembling those of Y. granulo-

sa. Dorsal aspect: The basal part of uncus rather

flattened dorso-ventrally and the appearance

wide in dorsal view; valva sinuate, with the dis-

tal part of narrow portion widened and outer

margin making a strong outward bend; distal

part of inner margin straight. Lateral aspect:

The narrow portion of uncus laterally com-

pressed and forming a V in cross section; short-

er than the narrow portion of the uncus in Y.

recta; appendices angulares of tegumen bent,

with apices converging towards each other; val-

va robust, two slight concavities ventrad, one

on each side of the median bend, dorsally con-

vex at junction of thick and narrow portion; ae-

deagus shorter than valva; diaphragma unscle-

rotized.

eme somme (es. 196 ID) —

Prep. nos. 691, 695, 1241, 1252754, 1265 67,

1277 from Lamto, Ivory Coast. Sinus vaginalis

wide and rather deep; anterior plate large and

broad, distally dentate with a shallow median

emargination; posterior plate unsclerotized,

consisting of a broader posterior part and a

tongue- like anterior section; ductus short and

ide. bursa small, obliquely Sean Daped

Habitat. — Savanna country, according to

Condamin

Distribution (fig. 8). — Lamto and Tiassale,

Ivory Coast, and Mbalmayo, Cameroun.

Holotype d : Ivory Coast, Lamto (Toumodi),

29.v1.1965, S. Kwamé. Allotype 2: Same, but,

13.vii.1967, R. Vuattoux. Paratypes: w.s.f. 20

3, same locality, but 20.vi.1964, 21.vi.1965,

26.v1.1964, 12.1v.1965, 18.11.1965, 20.v1.1965,

29.01.1965, 174911.1965, 2710119651 9A111819658

16.v11.1965, 10.v11.1965, 27.11.1964, 4.viu.1965,

17.viu.1965, S. Kwame. 13 2, same locality, but

23.11.1965, 27.11.1965, 25.11.1965, 29.11.1965,

2.v.1965, 29.v1.1965, 17.vu.1965, 11.vii1.1965, S.

Kwamé; 4.x.1967, R. Vuattoux; 19.11.1964, M.

Condamin; 2.1v.1962, C. Bigot. Paratypes ap-

proaching to d.s.f.: 3 d, same, 8.1.1968, R.

Vuattoux; 7.1.1946, M. Condamin; 25.111.1965,

S. Kwamé; 1 2 same, but 7.11.1964, M. Conda-

min. Holotype, allotype and the bulk of the par-

atypes to be returned to IFAN. Other paratypes

in NMN and J. Kielland collection.

Ypthima recta Overlaet

(figs. 8, 63—67, 121—124, pl. 5 figs. 15—18, pl.

8 figs. 3, 6)

1955: 87, figs.

Ypthima recta Overlaet, 25a, 25b

(Upemba, Zaire).

This species may easily be mistaken for Y.

granulosa, but the underside is usually darker. It

also resembles dark forms of Y. simplicia in ex-

ternal characters.

External characters (pl. 5 figs. 15—18, pl. 8

figs. 3, 6). — Antenna 36 joints, shaft and club

pale brown; shaft white ringed dorsally, club

gradual, end joint longer than penultimate; sec-

ond palpal joint two and a half times the length

of the last joint, nearly straight, first joint equals

the last joint in length.

Male. Upperside, wings dark brown (fresh

specimens); f.w. ocellar area striated, only

slightly paler than the ground colour, extended

towards tornus to vein 1b; there is a double

marginal line, rather wide apart and a submar-

ginal line midway between the ocellar spot and

margin, in some specimens slightly closer to the

margin; h.w. with an ocellus in 2, a discal band

outwardly dentate in area 3 and excurved in 5, a

submarginal line, incurved between vein 3 and 6

and a double, marginal line as in the f.w. Under-

side striated all over as in Y. granulosa; the f.w.

ocellar area clearly indicated by a darker border,

in a few specimens indistinct, thickest on the in-

ner side; the area is of the same shade and stria-

tion as the rest of the wing; there is a double

marginal line, in some specimens the inner line

KIELLAND: Ethiopian Ypthima 125

17227

ro,

ZZZ

Figs. 91—102. Male (91—99) and female (100—102) genitalia of Ypthima species. 91—93, Y. pulchra Overlaet,

paratype, Kafakumba, Katanga; 91, dorsal aspect; 92—93, aedeagus, dorsal and lateral aspect (prep. P. Muteshi).

94— 96, Y. albida Butler, Kibondo, Tanzania; 94, dorsal aspect; 95, lateral aspect; 96, aedeagus, lateral aspect.

97—99, Y. albida uniformis Bartel, Zaire; 97, dorsal aspect; 98, lateral aspect; 99, aedeagus, lateral aspect. 100—

102, Y. asterope Klug; 100, Somalia, typic; 101, Kiambitu Hill, Kenya, somewhat atypic; 102, Yatta, Thika, Ke-

nya, highly atypic, particularly the folded anterior plate, which is flattened in the drawing.

126 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

is indistinct due to the striation; h.w. with a

double spot in 1b, in some specimens also a spot

in 5, situated proximad of a line drawn between

the spot in 2 and 6; a discal band as on the up-

perside and an irregular subbasal line which is

indistinct in many specimens; an indistinct sub-

marginal line and marginal lines as in the f.w.

Length of f.w. 17.2 to 17.7 mm, antenna-wing

ratio 0.44.

Female. Larger than the male; wings slightly

more rounded; upperside ground colour a little

paler; f.w. ocellar area wider, a little paler,

reaching close to the hind margin; otherwise

markings as in the male. Length of f.w. 18.7

mm, antenna-wing ratio 0.36.

Male genitalia (figs. 63—67). — Prep. nos.

849 from Katanga, Zaire; 1698, 1699, 1703,

1704, 1706, 1708, 1710, 1711 from Ngara, Tan-

zania; 2003, 2042, 2285, 2286 from Angola;

1876 from Endebes, Kenya. Similar to Y. gran-

ulosa, particularly the coastal form. Dorsal as-

pect: Tegumen medium broad, uncus broad in

basal half, then abruptly narrowing to the thin

apical portion; valva long, slender and sinuous,

apical portion broadening and turned out; the

process is wide and prominent, placed at about

70° to the longitudinal axis of valva, sloping in-

teriad. Lateral aspect: Appendices angulares of

tegumen turned interiad; valva broadest one

third from base, tapering towards base and to-

wards the apical narrow portion which is long

and slender, slightly upturned at apex; aedeagus

shorter than valva.

Female genitalia (figs. 121—124). — Prep.

nos. 387 from Elizabethville, Zaire; 860 no date

label; 1701, 1702 from Ngara, Tanzania. Geni-

tal chamber deep, squarely formed; lateral scle-

rites extremely broad and tapering to a point

proximad; anterior wall with a sclerotized fron-

tal area, broad dorsad, tapering ventrad; ante-

rior plate wide and short with serrate margin in

paratype, but the plate is variable, although al-

ways uneven or serrate and wide, sometimes

very wide; in no. 860 trilobed apically; posteri-

or plate tongue-like, sclerotized and not clearly

seen, very wide at base (the shape of the posteri-

or plate is constant); ductus wide and short;

bursa small, obliquely ovoid.

A single male from Lamto, Ivory Coast pre-

sented to the author by Dr. M. Condamin, dif-

fers so much from typical Y. recta that it de-

serves to be remarked upon. The upperside

sooty brown; f.w. ocellar area entirely without

striation and the whole insect extremely like Y.

impura Elw. & Edw. and Y. pupillaris Bel; the

only external distinction may be the submarg-

inal dark line which is situated closer to the

margin than to the middle of the distance from

margin to the ocellar yellow ring; in Y. impura

and Y. pupillaris the line is either closer to the

eye spot, or placed in the middle. Underside

with strongly indicated ocellar dark border on

the f.w., reaching 1b; h.w. with a subbasal, an

inner discal, a faint outer discal and a submar-

ginal band; there is a double marginal line, the

inner blackish; the ocellar spots are very small, a

double one in 1b, one tiny spot in 2 and one in

6. Genitalia as in Y. recta.

Note. — It might be suggested that the coast-

al form of Y. granulosa is identical with Y. rec-

ta. However, this is out of the question as the

female genitalia of the two differ considerably.

Moreover, the fore leg of the female in Y. recta

is much reduced; tarsi one-jointed, fusiform;

tibia bent in the middle, as long as the tarsi;

femora thicker, but only a little longer than the

tibia. In Y. granulosa the female fore leg is well

developed as usual.

Habitat. — Woodland and open grassland of

rather humid climate. Often at margins of ever-

green forests.

Distribution. — See fig. 8.

Ypthima granulosa Butler

(figs. 8, 68—78, 106—109, 160—162, pl. 6 figs.

1—8)

Ypthima granulosa Butler, 1883: 101 (Victoria Nyan-

za, Tanzania).

Ypthima cataracta Van Son, 1955: 154, pls. 5, 6, fig.

81 (Rhodesia), syn. nov.

Ypthima asterope ab. inocellata Strand, 1909: 112

(Uluna-Langenburg).

Ypthima asterope ab. triocellata Strand, 1909: 112

(Langenburg, N. Nyassa).

Ypthima asterope ab. biocellata Strand, 1909: 112

(Dar es Salaam, Tanzania).

Ypthima asterope ab. interrupta Strand, 1909: 113

(Dar es Salaam).

Ypthima asterope ab. uniocellata Strand, 1909: 113

(Dar es Salaam).

General remarks. — As mentioned before,

the types of Strand’s aberrations of Y. asterope

asterope were destroyed during the war of

19391945. To identify specimens from de-

scriptions only, is very difficult, due to the vari-

ation of Ypthima within a given species and this

makes it difficult to analyse Strand’s descrip-

tions with certainty. However, due to the cir-

cumstances stated below, we must presume that

the above aberrations do not belong to Y. aster-

ope, but are forms of Y. granulosa, common in

KIELLAND: Ethiopian Ypthima 127

coastal areas of East Africa and also occurring

in several places inland. Personally, I have col-

lected hundreds of Ypthıma in the coastal areas

of Tanzania and examined other collectors’

specimens, all from areas mentioned by Strand

regarding Y. asterope. All of these turned out to

be Y. granulosa and Y. impura. Y. asterope ab.

biocelligera and Y. asterope ab. binucholata

probably also belong to Y. granulosa, but are

vaguer and Strand’s series could consist of other

species as well and should not be included here.

Strand (1909: 113) says about ab. interrupta

“Hinterflügel unten mit 6 ocellen, ----”. This

corresponds with a form of Y. granzlosa which

also occurs at Dar es Salaam (the type locality ot

Y. interrupta). It has small ocellar spots on the

h.w. underside; even in the wet season the spots

are small, although quite perfect with rings and

pupils. The d.s.f. has from four to six minute

black spots. Strand’s ab. #niocellata of Y. aster-

ope corresponds well with the d.s.f. of Y. gran-

ulosa. He says: “Hinterflügel unten mit 5 bis 6

winzig kleinen Ocellen (die in den Feldern 4

und 5 zum Teil ganz rudimentàr und die innere

der beiden am Analwinkel gelegene Ocellen nur

durch ein schwarzes Pünktchen angedeutet.)”

This specimen was also taken at Dar es Salaam.

Y. asterope ab. inocellata is another d.s.f. of Y.

granulosa of the inland with darker underside.

Y. asterope ab. triocellata and Y. asterope ab. bi-

ocellata are also forms of Y. granulosa with two

or more ocellar spots on the upperside h.w. Thıs

is often the case with the atypical form of Y.

granulosa, described below. Further, Van Son

(1955: 163) mentions that he had a species from

Mozambique which he correctly thought was

Y. granulosa (in fact the coastal form). He also

says that he has seen genitalia drawings made by

Elwes & Edwards of a species from Zanzibar

which also resembles Y. granulosa. It may

‚therefore seem curious that Van Son did not

recognize his species Y. cataracta as Y. granulo-

sa. There is a specimen in NMN from Dondo,

Mozambique resembling Strand’s interrupta

and van Son’s description of the specimens he

saw from Mozambique. His description of the

genitalia also resembles that of the coastal form

of Y. granulosa, which differs somewhat from

the typical, or inland form (figs. 70—74). The

two forms of Y. granulosa, differing both in

genitalia and size and also often in facies, are in-

deed confusing and it led me to think that they

were two separate species, but extensive dissec-

tions of both my own and NMN material re-

vealed that there were intermediates and al-

though the female genitalia varied to some ex-

tent, they did not vary in accordance with other

characters. Sometimes this is also the case with

the male. Therefore, it must be presumed that

the two forms belong to the same species. They

cannot be separate races, as the two fly together

in most areas. The larger form, though, is by far

the commonest along the coastal belt, inland to

Morogoro, while the smaller form is much more

common further inland.

External characters (pl. 6 figs. 1—8). — An-

tenna short, 28 to 30 joints; shaft brown, white

ringed, end joint much longer than penultimate;

club long and distinct; palpi very long, brown

and white hairs, sprinkled with white.

Typical form. Mainly inland. — Wet season

form. Male. Upperside wings fuscous brown;

the subapical ocellar area triangularly shaped,

not as clearly defined as in Y. asterope Btl.,

more like Y. yatta, but to a varying degree;

faintly striated, more or less drawn out towards

tornus as in Y. lamto; the submarginal area with

same palish shade as the distal half of the ocellar

area; the size of the ocellar spot is variable, but

with few exceptions smaller than in Y. asterope.

H. w. mostly with an eye-spot in 2, but in some

specimens there is no trace of any spots; there is

a faint indication of a discal and a submarginal

band; margin as in the f.w.; cilia in both wings

brown with a darker median line. Underside

f.w. very variable; in some specimens the ocellar

area is rather clearly defined by a dark line

which in some specimens is thin, in others

thicker, but in some specimens the area is more

or less obscured, particularly on the distal side;

in most, the apex is much paler than in the rest

of the wing, due to a fine striation, which is also

extended to the rest of the wing, but more scat-

tered; however, in others, the basal part, up to

the end of the cell, has the same degree of stria-

tion and shade as the apex, in a few, almost the

entire wing, but the area between the eye-spot

and the hind margin is always darker and less

striated; there is a blackish marginal line; the

h.w. is finely granulated; there is a very uneven

discal band, sharply protruding distad in area 3

to 4 and in some specimens also in 5; the rest of

the band is very wavy; there is a likewise un-

even, but generally fainter submarginal band.

The variation of the unevenness seems to corre-

spond with the size of the ocellar spot in space

2; in specimens with a large spot, the submar-

ginal line in this area is pushed closer to the

margin; there is a marginal line as in the f.w.;

normally there are four ocellar spots, two in tb,

128 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

109

Figs. 103— 115. Female genitalia of Ypthima species. 103, Y. congoana Overlaet, paratype, Elisabethville,

Zaire; bursa missing. 104, Y. antennata van Son, Serengeti, Tanzania. 105, Y. rhodesiana Carcasson. 106—109,

Y. granulosa Butler; 106, prep. no. 1012, Tabora, Tanzania; 107, prep. no. 460, Kenya; 108, prep. no. 871, Kivu,

Zaire, posterior plate removed; 109, posterior plate of prep. no. 871. 110, Y. vwattouxi species nova, allotype,

Lamto, Ivory Coast (prep. Condamin). 111, Y. albida Butler. 112—113, Y. doleta Kirby; 112, Abijan, Ivory

Coast; 113, Karera Sango Bay, Masaka, Uganda. 114-115, Y. pulchra Overlaet, paratype, Landoa, Lualu River,

Katanga; 115, anterior and posterior plates in lateral aspects.

KrELLAND: Ethiopian Ypthima 129

7 NE

\ /

one in 2 and one in 6; the size of the ocellar

spots is very variable even in specimens from

the same season; in some they are tiny, while in

others quite large, particularly in 2 and 6; there

are a few specimens with only one small spot in

2 and some with a spot in 5 and others in 3 as

well, all together six spots; cilia as on the upper-

side. Length of f.w. 15.5 to 17.7 mm, antenna-

wing ratio 0.36. There is one male from Kenya

with ratio 0.40. Otherwise the highest measured

is 0.38 and the lowest 0.35.

Female. Upperside paler than in the male and

with more widely spaced striation; ocellar area

paler and larger and more clearly defined; h.w.

130 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

as in the male, but slightly paler. Underside

much paler and with more widely spaced stria-

tion than in the male; otherwise similar. The un-

derside of the female has an affinity to that of

female Y. recta, but the specimens are usually

smaller. Length of f.w. 17 mm, antenna-wing

ratio 0.35.

Dry season form. This varies very little from

the w.s.f. in East Africa; the underside h.w.

ocellar spots are always small, sometimes tiny

or lacking, except for a tiny spot in space 2. Van

Son (1955) describes the d.s.f. from Rhodesia as

being slightly different from those occurring in

Fast Africa. In specimens from Rhodesia there

is no trace of any eye-spots on the h.w. This is

probably due to the somewhat more arid cli-

mate in Rhodesia so that d.s.f. becomes more

extreme.

Atypical form. Mainly coastal. — Fore legs,

antenna and palpi as in the normal form; f.w.

ocellar area paler and more clearly defined, less

difference between male and female; h.w. discal

and submarginal bands rather prominent. Un-

derside generally paler with more widely spaced

striation, as in the females; the specimens are

slightly larger on the average.

Male genitalia (figs. 75—78) (typical). —

Ba mos. 192, 377, 307, BD, 87%, B33, ZE,

1697, 1700, 1705, 1707, 2289 from Tanzania;

375, 1914 from Rhodesia; 385, 398, 853—55,

925, 927, 1413, 1738 from Kenya; 386 from

Uganda; 1414 from Burundi; 2000, 2001, 2037

from Botswana; 2004 from Angola. Dorsal as-

pect: Basal portion of uncus wide, wider than in

Y. recta; the narrow portion comparatively

short, almost as in Y. lamto; valva narrow por-

tion more or less curved interiad and distal por-

tion pointing away, but not as much as in the

atypical form; accessory plates situated distad

and generally not extended along the inner mar-

gin of valva; aedeagus short, less than two third

of the length of valva, constricted at the distal

end of the open, basal portion; the extreme base

straight (seldom rounded). Lateral aspect: Dor-

sal outline of tegumen-uncus rather straight;

wide portion of valva shorter and not so robust

as in Y. lamto; narrow portion sharply pointed

upwards and much longer than in Y. lamto; a

slight concavity on each side of the ventral

bend; diaphragma with a rectangularly shaped

nica ventrad of anus; saccus fairly long

and narrow; basal portion of aedeagus sharply

bent : ventrad.

Female genitalia (figs. 106—109, 160—162).

— Prep. nos. 460, 1483, 1667—69 from Tanza-

nia (coastal); 921 from Uganda; 1012—15 from

Singida, Tanzania; 1412 from Malawi; 1917

from Rhodesia. Sinus vaginalis deep, a sclero-

tized anterior section of the anterior wall, which

is finely dentate along the edge; anterior plate

very large, broad and deeply incised at the distal

middle, with a rounded lobe on each side; pos-

terior plate consists of two parts, the foremost

almost unsclerotized, tongue-like and pointed

distad; the posterior section sclerotized, more

or less incised at the distal middle; this part of

the genitalia varies as shown in the figures. The

lateral sclerites U-shaped, rather thick at base,

gradually tapering towards the proximal ends;

ductus fairly wide, rather short; bursa oval and

straight; colliculum large and oval.

Male genitalia (figs. 68—69) (atypical). —

Prep. nos. 219, 342, 344, 924, 148486 from

Tanzania; 920 from Mozambique; 871 from Ki-

vu, Zaire; 874 from Zambia; 852 from Kenya;

371, 376, 926, 929, 930; 932 frome sande

seems necessary also to describe the atypical

form of the male genitalia. Tegumen-uncus lon-

ger than normal; uncus narrower, distal portion

longer; valva narrow portion strongly curved

interiad, with distal end pointing away; the ac-

cessory plate in most specimens extended along

the inner edge of valva; aedeagus not con-

stricted and open portion evenly rounded.

Lateral aspect: Valva long and sinuate; aedeagus

sligthly longer than normal.

Intermediates between typical and atypical

forms occur.

Habitat. — Open, deciduous woodlands.

Distribution (fig. 8). — East Africa to Mala-

wi, Zambia, Rhodesia and northern Botswana

and a few records from the eastern part of Zaire.

Ypthima pupillaris Butler

and

Y. impura Elwes & Edwards

These two species are in most cases extremely

similar, and before I attempt to describe them, I

would like to make the following remarks. Both

are represented by two races. The nominate race

of Y. pupillaris occurs in South Africa and the

western part of Africa north to Senegal and Y.

pupillaris obscurata in eastern Africa. Y. impura

impura is West African and its race paupera oc-

curs from South Africa to East Africa and

Ethiopia. Previous authors have often confused

the two species with each other. This confusion

is, I think, due to the very great variability of Y.

impura, both in size, markings and coloration.

It is quite interesting to notice that Y. impura

KIELLAND: Ethiopian Ypthima 131

Figs. 116—126. Female genitalia of Ypthima species. 116—118, Y. pupillaris Butler, Ivory Coast; 116, sinus va-

ginalis flattened down, with genital plates and lateral sclerites; 117, left anal lobe; 118, bursa with ductus. 119—

120, Y. impura Elwes & Edwards, Ivory Coast; 119, ventral aspect; 120, right anal lobe in lateral aspect. 121—

124, Y. recta Overlaet; 121, prep. no. 860; 122, prep. no. 1382; 125, posterior plate detached; 124, prep. no. 887,

paratype, Elisabethville, Zaire, part of genitalia missing. 125—126, Y. jacksoni species nova, allotype, Kata Mane

River, Ethiopia; 125, sinus vaginalis flattened down; 126, right anal lobe.

132

TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

KIELLAND: Ethiopian Ypthima 133

from the eastern part of Africa resembles Y. pu-

pillaris from that same area, while the W. Afri-

can form resembles Y. pupillaris from that part

of Africa.

The W. African race of Y. pupillaris can easily

be distinguished from the E. African form of Y.

impura on external characters, but the E. Afri-

can races of the two species are difficult to sepa-

rate with certainty, without exposing the geni-

talia, and the W. African races hardly at all; they

resemble each other perfectly and I cannot point

to any particular character which is distinct for

each species.

I have before me 62 males of Y. pupillaris pu-

pillaris and 220 males of Y. impura impura from

Lamto, Ivory Coast, and it is not possible to de-

termine individual specimens with certainty,

without examining the genitalia. The two spe-

cies are both very variable and at the same time

very much alike. But when having the two large

series beside each other, it is noticeable that on

average, Y. pupillaris has more clearly defined

markings on the upperside of the wings. All the

62 Y. pupillaris, representatives of the two sea-

sons, have clear h.w. submarginal and discal

bands. This is very variable in Y. impura, but in

most specimens the markings are more

obscured.

Fortunately, the two species can easily be

separated on the genitalia by brushing aside

some hairs of the tip of the abdomen and exam-

ining with a good hand lens. The wide uncus of

Y. pupillaris and the narrow, sharply bent uncus

and long, slender valva of Y. impura cannot be

mistaken, and they are also quite distinct from

all other species here dealt with. In Y. impura

from Ivory Coast I have never seen very large

h.w. underside ocellar spots as often occurs in

Y. pupillaris from that part of Africa. The gen-

eral appearance when large series are examined,

is the paler underside of Y. impura. The differ-

ence between dry and wet season forms of Y.

impura impura is more pronounced than in Y.

pupillaris pupillaris.

Ypthima pupillaris Butler

For distinguishing characters of this species

see above.

Ypthima pupillaris pupillaris Butler

(figs. 9, 83—86, 116—118, 171, pl. 6 figs. 9—

12)

Ypthima pupillaris Butler, 1888: 59 (River Dangu,

Ganyese Tambu, Eq. Afr.)

Ypthima gazana van Son, 1955: 151, pl. 5, figs. 79, 80,

gen. figs. 169, 170. Syn nov.

Ypthima pupillaris ab. depupillata Strand, 1909: 113

(Loma).

Ypthima pupillaris ab. macrocellata Strand, 1913: 150

(Cameroun).

Rhodesian specimens described as Y. gazana

by van Son (1955: 151) correspond with Y. pu-

pillaris pupillaris Butler from Ivory Coast. The

pronounced markings above of specimens from

W. Africa and Rhodesia are identical and much

clearer than in E. African specimens; on an av-

erage they are also smaller. These characteristics

are constant in all the specimens I have exam-

ined, in both seasonal forms from W. Africa and

Rhodesia, and different from E. African aggre-

gates.

Strand’s Y. pupillaris ab. depupillata is de-

scribed from a single worn female from Loma,

with no other locality name; with large ocellar

spots and no pupils in spots 2 and 6 of the h.w.

underside (these, however, may have been

rubbed off, if small). Whether the specimen be-

longs to Y. pupillaris or to Y. impura is a ques-

tion which must remain unanswered, as the

specimen most certainly was destroyed together

with most of the other Ypthima types in Berlin,

during the 1939—45 war. Y. pupillaris ab. ma-

crocellata from Cameroun, could be Y. pupilla-

ris, but also possibly Y. impura. This cannot be

verified as the type is lost. Strand says “Das

Exemplar zeichnet zich durch grosse Ocellen

der Unterseite der Flügel aus”.

External characters (pl. 6 figs. 9—12). — An-

tenna male 35, female 36 joints; shaft brown and

Figs. 127—135. Female genitalia of Ypthima species. 127—128, Y. simplicia Butler, Ethiopia; 127, prep. no.

1743, with sinus vaginalis detached and flattened down; 128, posterior plate, prep. no. 707A. 129—130, Y. di-

plommata Overlaet, Upper Longwe, Bungo, Angola; 129, genitalia with posterior plate removed; 130, posterior

plate. 131—133, Y. praestans Overlaet, Katanga; 131, genitalia with anterior plate and bursa removed, posterior

plate straightened; 132, lateral sclerites with anterior plate, ductus and bursa; 133, posterior plate in natural po-

sition. 134—135, Y. yatta species nova, Sheik Hussein, Upper Shebeli River, Ethiopia; 134, sinus vaginalis with

genital plates; 135, right anal lobe.

134 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

white ringed above, white below, thicker white

line in the female; club gradual, double grooved

on the underside for full length of the joints,

end joint as long as, or longer than penultimate;

Van Son (1955) says that in South African speci-

mens the end joint is much longer. Palpi: second

joint more than twice the length of the terminal

joint.

Wet season form. Male. Upperside, ground

colour of the wings brown (rather faded speci-

mens); f.w. inner side of the ocellar area rather

obscured in most specimens by the ground col-

our invading the area up to the ocellar dark

ring; outer dark brown border of ocellar area

very pronounced; a paler submarginal border

and two clearly defined marginal lines, the inner

line thickest, like Y. impura; h.w. with rather

prominent discal and submarginal bands,

somewhat resembling Y. doleta; two margınal

lines; one or two ocellar spots in 1b and one in

2; in some the spot in 1b is lacking, or extremely

faint. Underside f.w. ground colour greyish,

with dense olive-brown striation; ocellar area

clearly defined by the dark brown border; in

some specimens this dark border is merging

with the brown eye-ring, with hardly any stria-

tion in between; in others, there is striated space

in between them; the brown border is generally

drawn out towards tornus, sometimes nearly

reaching the hind margin of the wing; there are

clearly defined brown bands on the h.w. sub-

marginal and discal areas, a double marginal

line, the inner thickest, almost black; a double

ocellar spot in 1b, one in 2 and one in 6; the size

of the eye spots is very variable. Length of f.w.

16.3 to 19.3 mm, average 17.7 mm antenna-

wing ratio 0.43.

Female. Similar to the male, but the wings

more rounded; underside as the male with large

ocellar spots, those in 6 and 2 of the h.w. large

and of equal size, the tornal spot much smaller.

Length of f.w. 16 to 20.8 mm, average 18.5 to

19 mm, antenna-wing ratio 0.41.

Note. Apart from the Rhodesian specimens,

the material examined comes from humid parts

of W. Africa where typical d.s.f. are not likely

to occur.

Male genitalia (figs. 8386). — Prep. nos.

708, 727, 1239 from Ivory Coast; 1352, 1823

from Nigeria; 390 from Rhodesia. Dorsal

aspect: Uncus very wide and dorso-ventrally

flattened from base to apex, unlike all other Yp-

thima of the continental African group. Lateral

aspect: Saccus rather large, robust; valva, basal

two third very broad, squarish, with dorsal

margin almost parallel to ventral margin,

abruptly bent ventrad to form the narrow distal

part; the apical process placed dorsad, pointing

up and at 90° to the longitudinal axis of valva;

aedeagus as long as valva, the basal one quarter

(phallobase) narrow and turned ventrad, the rest

evenly tapering to a rather blunt, upturned tip.

Female genitalia (figs. 116—118, 171). —

Prep. nos. 701, 709, 1269, 1270, 1272 from Ivo-

ry Coast; 888 from Rhodesia. Anterior wall of

sinus vaginalis flattened down in the figure. The

anterior plate very wavy, one large fold on each

side of the raised frontal portion, which consists

of two rounded lobes separated by a shallow

emargination, protruding as a fold; the frontal

portion of the posterior plate consisting of two

rather pointed lobes, separated by a V-formed

indentation, its lateral folds strongly bent back;

the lateral sclerites wide nearly all the way, V-

shaped; anterior wall of sinus vaginalis with a

tongue-like frontal sclerotization; ductus long,

bursa kidney-shaped.

Habitat. — Woodland and grassland.

Distribution (fig. 9). — Ivory Coast to Nige-

ria, W. Zaire and Rhodesia.

Ypthima pupillaris obscurata subspec. nova

(figs. 9, 170, 172, pl. 6 figs. 13—18)

External characters (pl. 6 figs 13—18). —

Male. Differs from the nominate race in the

more evenly coloured fuscous brown upperside,

with only very slight indications of h.w. bands;

inner side of the f.w. ocellar area more obscured

by the dark brown ground colour invading the

area up to the dark ocellar ring. Underside, h.w.

marginal lines less clearly defined; particularly

in the d.s.f. the lines are almost obscured in the

striation and hardly discernable. The specimens

are larger than in ssp. pupillaris, even the d.s.f. is

a little larger than the W. African race. Length

of f.w., w.s.f. 20.8 mm, d.s.f. 18.7 mm, on the

average. Antenna-wing ratio 0.40 (lower than in

Y. pupillaris pupillaris).

Female. Slightly paler than the male; wings

more rounded; very faint indication of h.w. dis-

cal band; underside as in the male, with very

large eye-spots in the w.s.f., which are reduced

to small black dots in the d.s.f., the f.w. apical

eye-spot is also smaller in the d.s.f. The female

differs little from the nominate race. Length ot

f.w. 19 to 20.5 mm.

Male genitalia. — Prep. nos. 20, 57, 122, 223,

228, 391, 721, 866, 542 from Tanzania. Same as

Y. pupillaris pupillaris; the narrow distal portion

of valva often a little shorter.

KIELLAND: Ethiopian Ypthıma 135

Female genitalia (figs. 170, 172). — Prep. nos.

ONM O5 1397 from Tanzanıa. As

the nominate race. The posterior plate varies a

little, as shown in the figures. This variation

seems to occur in both races.

Habitat. — Rare at lower altitudes with

woodland, but locally common in montane

grassland above 1500 m.

Distribution (fig. 9). — Eastern Africa.

Holotype 6: Tanzania, Mpanda, Ngondo,

1500 m, 14.1.1970, J. Kielland. Allotype © : Tan-

zania, Kigoma, Lubalizi, 1200 m, 7.11.1970, J.

Kielland. Paratypes: Mpanda, Sitwe, 1700 m,

20.v.1970, 24.x.1958, 2 d; Mpanda, Kampisa,

1500 m, 24.x1.1968, 25.x1.1962, 2 d; Kigoma,

Mweze, 1700 m, 11.vi.1968, 2 d; Mpanda,

Ngondo, 1500 m, 14.1.1970, 2 d; J. Kielland;

Mufindi, November 1958, R. H. Carcasson, 4

d; Njombe, xii 1968, C. Maccleary, 2 6; Kigo-

ma, Mt. Kungwe, ix.1954, 1 d; Uganda, An-

kole, Kalinzu, x1.1961, 1 © ; Acholi, Madi Opei,

MAP MIE E] ackson 1) 92 Holoty,pe and

allotype in the NMN, paratypes in NMN, BM

and J. Kielland collection.

Ypthima impura Elwes & Edwards

For distinguishing characters, see above.

Ypthima impura impura Elwes & Edwards

(figs. 9, 79— 82, 119, 120, 156—159, pl. 7 figs. 1,

Ypthima impura Elwes & Edwards, 1893: 23 (An-

gola).

Elwes & Edwards (1893) said about this spe-

cies and Y. doleta that they are very close in ex-

ternal characters, particularly the underside. Al-

though the W. African aggregates may resemble

Y. doleta to a certain extent, in my opinion the

external characters are closer to Y. pupillaris

and as stated before, almost identical. But the

genitalia of all these species are very unlike each

other in both sexes.

External characters (pl. 7 figs. 1—2). — An-

tenna male 37 joints, female 36 tot 37 joints; end

joint > = < RL shaft black and white

ringed above, speckled with white below; palpi

black and white hairy; second joint twice the

length of the last joint.

Wet season form. Male. Upperside, ground

colour dark greyish brown; f.w. ocellar area a

little paler than the ground colour, usually

clearly indicated by a dark brown border; there

is a double, prominent, brown marginal line; the

h.w. with a thick and sharply defined, dark

brown discal band; a prominent submarginal

line; a double marginal line and a brown cilia

with a darker brown median line; there is an

ocellar spot in 2 and in some specimens there is

also one spot in 1b. Underside, f.w. with clearly

defined ocellar area; a brown, double marginal

line. In all specimens from Ivory Coast, Central

Congo and Ghana, the size of the h.w. eye-

spots tend to correspond with those of the in-

termediate w.s.f. of Y. impura paupera; none

are very big, as often happens in the extreme

w.s.f. of ssp. paupera; also the uneven striation

and prominent bands correspond.

Female. A little paler and larger than the

male, with broader wings.

Dry season form. The only difference from

the w.s.f. is the ocellar spots, which are reduced

to tiny points, often hardly discernable. None

of the specimens has the pale underside of the

d.s.f. of ssp. paupera. Length of f.w., w.s.f. 6 16

io 18.5 mn 2 175 co 19.5 mom clot, 6 165%

19 mm, © 18 to 20.5 mm, antenna-wing ratio d

0.43, 2 0.41.

Male genitalia (figs. 79—82). — Prep. nos.

403 from Angola; 704, 706, 710, 711, 726, 1236,

1237, 1242-46, 1255, 1259, 1264, 1268 from

Ivory Coast; 1226 from Ghana; 1355 from Ni-

geria. Dorsal aspect: Tegumen and base ot un-

cus wide, then abruptly tapering to a long and

narrow distal portion; distal end ot valva trun-

cate, with a wide accessory plate, extending

along the inner edge (this is normal, but in a few

specimens the plate is situated distad and in

many specimens the plate is interrupted.)

Lateral aspect: Tegumen normal, but uncus di-

rected upwards with the distal end strongly

curved ventrad; valva long and comparatively

narrow over the whole length, gradually ta-

pering; aedeagus longer than tegumen-uncus,

but shorter than valva.

Female genitalia (figs. 119120, 156—159).

— Pago, mos, MOO PSP 1256—58,

1260, 1271—73, 1275, 1279—82, 1284 from

Ivory Coast; 402 from Zaire. Sinus vaginalis

large and deep; the anterior wall with a frontal

sclerotization; anterior plate variable, normally

with three distal lobes, the middle largest,

rounded, some specimens with, others ition

an emargination in the middle; the posterior

plate is also variable, but is always very long and

narrow, with a Sunded apex in specimens ex-

amined from East and West Africa, but accord-

ing to van Son (1955), with a spear-shaped distal

end in specimens from South Africa; the lateral

sclerites V-shaped; ductus rather short; bursa

136

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFI

170 171

169

B 0

175

… 5 (1982)

172

KieLLAND: Ethiopian Ypthima

ovoid, with entrance of ductus centrally placed.

Van Son states that the bursa is “pyriform” (also

shown in his figure 172). None of the many

specimens I have examined from other parts of

Africa correspond with this statement; they

were all ovoid.

Distribution (fig. 9). — Ivory Coast to Nige-

ria and scattered records from Cameroun, Cen-

tral Congo and Angola.

Ypthima impura paupera Ungemach, comb

nov.

(fig. 9, pl. 7 figs. 3—8)

Ypthima pupillaris paupera Ungemach, 1932: 51

(Ethiopia).

Ypthima pupillaris paupera f. hiemis Ungemach,

1932: 51 (Ethiopia).

Ypthima impura f. badhami Van Son, 1955: 150, pls.

56, figs. 77, 78 (Rhodesia), d.s.f.

Ungemach (1932) described paupera as a sub-

species of Y. pupillaris with the form hiemis

from Ethiopia. Photos and genitalia drawings

sent to me by Dr. Condamin clearly show that

this is the E. African race of Y. impura, repre-

sented by wet and dry season forms; paupera

being the w.s.f. and hiemis the d.s.f. Specimens

of Y. impura paupera from Ethiopia are darker

than specimens from E. Africa.

Ssp. paupera is a very variable race and a de-

scription of just one form is very inadequate for

an identification. The underside h.w. eye-spots

vary from tiny black dots in the extreme d.s.f. to

very large spots in the extreme w.s.f.; the un-

derside coloration and markings are also ex-

tremely variable. The extreme d.s.f. has a very

pale underside, with rather thin and widely

spaced striation, but prominent discal and medi-

an lines; this probably had led to the confusion

with certain forms of Y. asterope. The interme-

diate form has slightly bigger eye-spots; the

submarginal and discal bands of the h.w. are

137

broader, the striation closer, giving it a darker

appearance; there is a w.s.f. with large eye-

spots, but otherwise with very little markings

except for the even, olive-brown striation, rath-

er similar to and easily misidentified with cer-

tain forms of Y. pupillaris; individual speci-

mens, two females from Endebes, Kenya, have

the underside very heavily striated, with pro-

nounced h.w. submarginal and discal bands,

with olive-brown blotches. Also the wing

length varies greatly, as well as the length of the

last antennal joint.

Van Son (1955) writes about Y. impura that

the w.s.f. is generally bigger than the d.s.f. In

Tanzania the form of the dry season appears to

be the largest. Further he states that the anten-

nal end joint equals the penultimate in length.

By examining numerous specimens, both from

Fast and West Africa, I have found that the last

joint > = < penultimate.

Due to the very great variability of this spe-

cies, it is difficult to make a proper description

of it and only the characters of the genitalia can

with certainty separate it from other species.

External characters (pl. 7 figs. 3—8). — An-

tenna 34 to 37 joints, shaft black and white

ringed above, speckled with white below; club

gradual; palpi white and black hairy, second

joint twice the length of the last joint.

Wet season form. Male. Upperside ground

colour dark greyish brown; f.w. ocellar area in-

distinct due to the ground colour invading it.

However, it is rather more clearly indicated

than in Y. pupillaris obscurata. The submargin is

slightly paler than the ground colour, separated

from the ocellar area by a dark brown line; in

some specimens the outer section of the ocellar

area is a little paler than the inner part, but in

others, there is hardly any difference; h.w. dis-

cal band is hardly discernible in some speci-

mens; submarginal and marginal lines and cilia

as in Y. impura; there is an ocellar spot in 2, in

Figs. 136—178. Female genitalia of Ypthima species. 136, Y. lamto species nova, Lamto, Ivory Coast. 137—

138, variations in genital plates; 137—141, anterior plates of Y. asterope Klug; 137—139, normal shapes, East-

ern Africa to Somalia; 140, atypic, Kiambitti Hill, Kenya; 141, atypic, Tiaroye, Senegal. 142—145, posterior

plates of Y. asterope Klug; 142, corresponding to 137; 143 to 138; 144 to 134; 145 to 141. 146, Y. congoana

Overlaet, anterior plate. 147—151, Y. rhodesiana Carcasson; 147—148, anterior plates; 149— 151, posterior

plates. 152—155, Y. antennata van Son; 152—153, anterior plates; 154—155, posterior plates (152 and 154, Se-

rengeti, Tanzania; 153 and 155, White Nile, Uganda). 156—159, Y. impura Elwes & Edwards, posterior plates;

156-157, common forms from East and West Africa; 158, South Africa; 159, uncommon form. 160-162, Y.

granulosa Butler, posterior plates. 163—165, Y. lamto species nova, anterior plates. 166—167, Y. condamini

species nova, anterior plates, always covered with spiny hairs. 168—169, Y. doleta Kirby, anterior plates. 170—

172, Y. pupillaris Butler, posterior plates; 170, 172, ssp. obscurata subspecies nova, Tanzania; 171, ssp. pupillaris

Butler, Ivory Coast. 173—178, Y. simplicia; 173—175, posterior plates; 176, anterior plate of 173; 177, 178,

strongly aberrant form of anterior plate corresponding to 175 (177, frontal aspect; 178, flattened down).

138 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

some specimens there is also one spot in 1b. Un-

derside extremely similar to that of Y. pupillaris

obscurata with olive-brown striation; h.w. with

a large spot in 2, one in 6 and a smaller, double

pupilled one in 1b.

Female. The ocellar spot in the f.w. more

rounded; the yellow ring paler; wings more

rounded; underside ocellar spots a little larger;

otherwise similar to the male.

Early wet season form (intermediate form).

Both sexes differ from the extreme w.s.f in paler

and better defined ocellar areas, a more promi-

nent underside h.w. discal band, generally

thicker submarginal bands and a paler ground

colour; the striation is seldom olive tinged and

the specimens are generally casier to separate

from Y. pupillaris. This form is typical for west

Tanzania and occurs mostly during the first part

of the rainy season.

Dry season form. Upperside dark greyish

brown, the f.w. ocellar area clearly defined,

more or less similar to the intermediate form;

the underside is much paler than in the other

forms, due to the much more widely spaced,

blackish striation; the h.w. discal line clearly de-

fined, the submarginal and marginal lines hardly

discernible, particularly in the h.w. Females

with both wings much more rounded than in

the male, but otherwise similar.

The pale form reminds one of the underside

of the d.s.f. Y. praestans. Length of f.w. Inter-

mediate f. gd 16.5—19.5 mm, 2 18—20.5 mm.

Wess, © 1620 nom, © 12215 rain, Det,

& 175—205 mm, 20221 5imm Antenna.

wing ratio d 0.41, ® 0.39 (lower than in the

nominate form).

Male and female genitalia: As in Y. impura

impura.

Habitat. — Deciduous woodland, particu-

larly Brachystegia woodlands, and open hab-

itats.

Distribution (fig. 9). — Ethiopia and East Af-

rica to Zambia, Rhodesia, Transvaal and Natal

and southern and probably eastern part of

Zaire.

Ypthima pulchra Overlaet

(figs. 11, 91—93, 114, 115, pl. 8 figs. 7, 8, 10,

11)

Ypthima pulchra Overlaet, 1954: 41, fig. 1a—b (Ka-

tanga, Zaire).

À paratype male and a female were kindly

sent on loan to the author by Mr. Berger, TMB

The description is as follows.

External characters (pl. 8 figs. 7—8, 10-11).

— Antenna 34 joints, shaft black and white

ringed, club short, broader than in Y. praestans

and Y. doleta, terminal joint longer than penul-

timate; palpi second joint very straight, two and

a half times the length of the last joint.

Male. Upperside ground colour dark brown;

f.w. with prominent, striated ocellar area; sub-

marginal area as the ocellar area, divided by a

thick, dark brown submarginal line; inner bor-

der of ocellar area close to eye-spot sharply de-

fined, as in Y. diplommata and Y. doleta; a thick

brown margin; cilia white at base, then brown

with slightly paler tips; h.w. heavily marked, as

in Y. doleta, by very irregular discal and sub-

marginal bands; the submarginal band widened

and incurved in space 4 and 5; a larger eye-spot

in 2 and a smaller one in 6; marginal line as in

the f.w.; cilia a little paler. Underside f.w. with a

dark, curved median line, the basal area inside

this line paler and striated; the median section

up to the ocellar area darker brown; apex and

subapex pale and striated; a marginal line, thin-

ner than the upperside one; h.w. with promi-

nent median, discal and submarginal bands; the

submarginal bands making a wide bend into the

discal area, between vein 3 to middle of space 5,

its angle at vein 4; two ocellar spots in 1b, one

in 2 and one in 6; the h.w. is paler than the dark

area of the f.w.; the marginal line as in the f.w.

Length of f.w. 18.4 mm, antenna-wing ratio

0.42.

Female. Upperside ground colour as in the

male; the ocellar and submarginal areas paler;

there is also some striation basad from the ocel-

lar dark border; marginal line and cilia as in the

male; h.w. as in the male, the light area a little

paler; there is also an ocellar spot at tornus and

a tiny black dot near it. Underside f.w. ground

colour whitish grey, with very uneven striation,

thickest in the subtornal area; apex olive, with

no dark line between this and the ocellar area;

h.w. ground colour whitish grey, with very

scattered, small brown specks, becoming a little

closer towards the basal area; a median and a

discal band; the area between these, greenish

yellow or olive; a submarginal line with a large

olive blotch; eye-spots as in the male. Length of

f.w. 20 mm, antenna-wing ratio 0.40.

Both male and female are d.s.f.

Male genitalia (figs. 91—93). — No. NMN

92. Dissected by Mr. P. N. Muteshy. Dorsal

aspect: Tegumen very wide and short; uncus

wide basally and narrowing down to a thin dis-

tal portion; valva long and rather narrow and

KIELLAND: Ethiopian Ypthima 139

sinuate, distal end with accessory plate bent in-

teriad; aedeagus sinuate. Lateral aspect: Length

of tegumen-uncus only two third of valva which

is long and rather narrow, ventrad only slightly

elbowed before midway; outer narrow portion

very slender; aedeagus as long as tegumen-un-

cus, but much shorter than valva, distal end

curved upwards.

Female genitalia (figs. 114-115). — No.

NMN 93. Dissected by Mr. P. N. Muteshy. Si-

nus vaginalis wide, but rather shallow; margin

of anterior wall slightly wavy, with a rounded

frontal sclerotization, reaching the margin; an-

terior plate rather long, evenly rounded distally;

posterior plate also long, but a little wider and

distal part curved ventrad; the apex evenly

rounded; lateral sclerites U-shaped and rather

close together. The rest of the abdomen was

damaged and the bursa was lacking.

Habitat. — Not known to author, but proba-

bly deciduous formations.

Distribution (fig. 11). — Known from cen-

tral and western Katanga in Zaire.

Ypthima praestans Overlaet

(figs. 10, 88, 131—133, pl. 7 figs. 9—12, pl. 8

fig. 12)

Ypthima praestans Overlaet, 1954: 43, fig. 2 (Sanku-

ru).

The holotype male was sent to me on loan

from the TMB by Mr. Berger. Other specimens

were examined at the BM.

External characters (pl. 7 figs. 9—12, pl. 8.

figs. 9, 12). — Antenna male 33 joints, female 37

joints, shaft black and white ringed, club thin,

gradual, upperside outer half yellow, the inner

half black, 12 joints, underside of each joint

double grooved over one quarter of its length,

last joint longer than penultimate; palpi second

joint two and a half times the length of the last

joint.

Male. Upperside, wings dark sooty brown;

f.w. ocellar area pale, striated, extended towards

tornus to vein 1b, surrounded by a dark band

almost reaching the hind margin near tornus

and almost touching the rather wide, but un-

clear marginal line; h.w. with slightly paler dis-

cal area and with the discal and submarginal

bands obscured; five ocellar spots, a double spot

in 1b, one larger in 2, small ones in 4 and 5 anda

bigger one in 6. Underside, wings almost as in

some forms of Y. impura paupera; but there are

also spots in spaces 4 and 5 as on the upperside;

a median line in both fore and hind wings; h.w.

with a discal and a submarginal band, which is

broken at vein 4 and hardly discernible between

vein 4 and tornus; the marginal line is very in-

distinct. Length of f.w. 19 mm, antenna-wing

ratio 0.39.

Female (d.s.f.). Upperside, ground colour

brown to pale brown in some specimens; f.w.

ocellar area large, striated, paler than the ground

colour and extended nearly to the hind margin

at vein 1b, indicated by a brown line; there is a

double marginal line; the area between the mar-

ginal line and the ocellar border of the same

shade and striation as the ocellar area; h.w. with

more or less clearly marked discal line; a sub-

marginal line parallel to the margin over the

whole length; a double marginal line, the inner

thickest; there is an ocellar spot in 2 and a dou-

ble spot in 1b. The spot in 1b is in some speci-

mens very indistinct; in some specimens there is

also a faint dot in 6; the narrow strip between

the two marginal lines and the submarginal line

is paler than the ground colour; the discal area

is, in some specimens, paler than the ground

colour, in others hardly paler. The specimens

look very similar to the female ot Y. impura on

the upperside, the underside like the d.s.f. of

both Y. impura and Y. pupillaris, except that the

discal line is straighter. Underside with paler or,

in other specimens, darker brown striation; f.w.

ocellar area clearly indicated by a brown line,

prolonged to vein 1b, somewhat irregular on the

underside in area 1 and 2; there is a double mar-

ginal line, the outer one very thin; cilia whitish

with a brown, median line; h.w. with a double

black spot in 1b, another dot in 2 and one in 6,

no clear indication of a pupil; there is a clearly

visible discal line, straighter than normal in Yp-

thima; a subbasal line, parallel to the discal one,

which is indistinct in some specimens; a sub-

marginal line and a double marginal line, the in-

ner of these is, in some specimens, indistinct; ci-

lia as in the f.w. Length of f.w. 19 to 21.6 mm,

antenna-wing ratio 0.38.

Male genitalia (fig. 88). — Prep. nos. 753A,

756A, 759A. Dorsal aspect: Tegumen-uncus

slender; tegumen a little shorter than uncus;

valva sinuate, widely bent and slender, accesso-

ry plates placed at the apex pointing almost

straight ahead; fultura thin, widely U-shaped;

saccus large. Lateral aspect: Tegumen nearly

rectangularly shaped; its lateral processes long

and narrow and turned anteriad; valva long and

slender, narrow portion very slender and distal

part bent ventrad; dorsal process large; aedea-

gus longer than tegumen-uncus, but much

140 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Figs. MEZZI Ypthimomorpha itonia (Hewitson), male genitalia; 179, lateral aspect; 180—181, variations of

distal part of valva. 182, Ypthimomorpha itonia (Hewitson), female genitalia. Figs. 183—184. Mashuna upemba

DET female genitalia; 183, ventral aspect; 184, lateral aspect. Fig. 185. Mashuna mashuna Trimen, female

genitalia.

KIELLAND: Ethiopian Ypthima 141

shorter than valva, distal half narrow, basal part

broad.

Female genitalia (figs. 131—133). — Prep.

nos. 757A, 1828K, 1829K, 2038, 2039, 2041. Si-

nus vaginalis deep; anterior and posterior plates

large and rather complex, not unlike those of Y.

pupillaris; lateral sclerites very wide in front, at

the base produced, forming a wide lobe which

looks like an extra fold of the anterior plate; the

anterior plate is bilobed and turned in at the

edges; the posterior plate which looks rather

complexly folded with two lateral lobes, com-

pletely changes its shape when straightened, as

shown in the figures; ductus narrow, bursa

small, almost circle-shaped.

Habitat. — Not known to author, but proba-

bly deciduous tree formations.

Distribution (fig. 10). — Recorded from Ka-

tanga in Zaire, Angola and West Zambia.

Ypthima doleta Kirby

(figs. 10, 89, 90, 112, 113, 168, 169, pl. 7 figs.

13—16)

Ypthima doleta Kirby, 1880: 335 (Sierra Leone).

Ypthima doleta ab. tripunctata Strand, 1909: 114

(Cameroun).

Ypthima doleta ab. quadripunctata Strand, 1909: 114

(Cameroun).

Ypthima doleta ab. quinquepunctata Strand, 1909:

114 (Cameroun).

Ypthima doleta ab. septemocellata Strand, 1909: 114

(Cameroun).

Ypthima doleta ab. strigata Strand, 1913: 150.

Ypthima doleta ab. defecta Strand, 1913: 150 (Uelle-

burg).

Ypthima doleta ab. asteropina Strand, 1913: 150

(Alen) (perhaps another species)

Ypthima doleta ab. dschangensis Strand, 1914: 153

(Cameroun).

As usual, Strand’s original descriptions of the

aberrations refer to variations of the ocellar

spots only.

External characters (pl. 7 figs. 13—16). —

Antenna male 34 joints, female 33 joints; shaft

dark brown, white ringed above, a longitudinal,

white stripe below; club thin, brown above, last

Joint as long as penultimate in most specimens,

longer in others, as in some specimens from

Uganda (NMN) and from Cameroun (IFAN);

underside of the club with two small, circular

grooves anteriad of each joint; palpi second

joint only slightly curved, two and a half times

as long as the last joint.

Male. Rather similar to some forms of Y.

praestans and Y. pulchra. Upperside wings

brown; f.w. ocellar area large, with a dark

brown border, almost reaching the hind margin;

ocellar and median areas, costa and submargin,

except apex, striated; margin brown bordered;

cilia brown with paler tips; h.w. with dark

brown discal and submarginal bands; the discal

line sharply excurved at vein 5, the submarginal

line incurved at vein 4 and in most specimens

more or less thickened between vein 3 and 6;

the discal and submarginal lines converge and

join each other close to the spot in space 1b,

near tornus, enclosing a paler, striated postdis-

cal area; there is a double ocellus in 1b, one in 2,

one in 3 and a poorly developed one in 6; the

one in 2 is the largest and placed closer to the

base of the wing than the ocellus in space 3 and

is in line with the second spot in 1b and the spot

in 6. Underside somewhat resembling Y. impu-

ra impura from West Africa, but generally paler

and often more evenly striated; there is a medi-

an line; the brown discal and submarginal lines

coinciding with those of the upperside, are, in

some specimens, broken up by the whitish

ground colour; ocellar spots as on upperside,

but the one in 6 is normally developed and of

the same size as the spot in 2. Length of f.w.

18.5 to 20 mm, antenna-wing ratio 0.42.

Female. Similar to the male, but the wings

much more rounded and rather paler; upperside

f.w. subapical eye-spot larger, the ocellar brown

border variable in thickness; striation of the f.w.

more extended basad than in the male, but the

extent of the striation varies. Underside as in the

male. Length of f.w. 19 to 22 mm, antenna-wing

ratio 0.39—0.40.

Male genitalia (figs. 89—90). — Prep. nos.

7201238 won Lory Coast. 12212225; 1349,

2005 from Ghana; 389, 2287 from Zaire; 877,

884, 885 from Uganda; 1724, 1725 from Cam-

eroun; 2002 from Angola. Dorsal aspect: Tegu-

men wide at base; uncus gradually tapering to a

narrow distal portion; valva strongly and evenly

curved from base to near apex; here the distal

portion is thickened and the apex is turned out-

ward; accessory plates placed at a sharp angle.

Lateral aspect: Tegumen nearly straight dorsad,

basal part straight, rectangular; vinculum rather

short and strongly curved; uncus as long as te-

gumen, in the dorsal middle rather hunched;

valva robust, narrow portion short and strongly

thickened distad; aedeagus shorter than valva,

rather robust and evenly curved.

Female genitalia (figs. 112—113, 168—169).

— Prep. nos. 718, 719, 1274 from Ivory Coast;

888, 1010 from Uganda; 1432 from Bukoba,

142 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

Tanzania. Sinus vaginalis large and deep, margin

even, anterior sclerotization narrow, reaching

the margin, distal part serrated; anterior plate

variable, but always bilobed distally and with an

anterior, smaller part, in some specimens bi-

lobed and in others single, more or less weekly

sclerotized; posterior plate unsclerotized with a

tongue-like distal part; ductus wide; bursa rath-

er small, slightly oblong.

Variation in Ghana and Ivory Coast. — All

specimens examined from Ghana and Ivory

Coast differ from aggregates from other parts of

Africa as follows. Male. F.w. subapical ocellar

area darker due to close striation; the area itself

smaller and its brown border wider and more

pronounced; the ocellar spot larger and the yel-

low ring brighter; h.w. median and submarginal

bands also more pronounced. Underside, f.w.

ocellar spot larger and brighter; the ocellar area

more heavily bordered in brown; h.w. markings

better developed.

In the female the f.w. ocellar spot generally

larger and on upper and underside ocellar area

much smaller than in aggregates from other

parts of Africa.

Specimens from the type locality of Sierra

Leone are similar to those from central and east-

ern Africa and I do not consider it advisable to

name the Ghana-Ivory Coast form as a distinct

race.

Habitat. — Flying in rather humid habitats of

farmland, forest margins and glades.

Distribution (fig. 10). — From Sierra Leone

through Ivory Coast and Ghana, Togo, Nigeria

to Zaire and Angola and from there to Uganda

and N.W. Tanzania. There are also a few re-

cords from S. Sudan.

Ypthima diplommata Overlaet

(figs. 10, 87, 129, 130, pl. 7 figs. 17, 18, pl. 8

figs. 13, 14)

Ypthıma diplommata Overlaet, 1954: 44, fig. 3 (Kafa-

kumba, Katanga).

The holotype (male, d.s.f.) was sent to the au-

thor on loan by Mr. Berger and a brief descrip-

tion follows.

External characters (pl. 7 figs. 17—18 pl. 8

figs. 13, 14). — Underside, wings ground colour

and markings as in Y. pulchra with perhaps a

paler tornal area of the f.w.; a large spot in 2,

one in 3 and a black spot in 6. Underside f.w.

differs in the paler subtornal area with a well de-

fined ocellar brown border; the yellow ocellar

ring brighter; h.w. ground colour whitish with

scattered brown striation, heavily marked

brown in the median area from costa to about

middle of the cell between the median and discal

line; a marginal brown blotch from apex to vein

4; there are two small ocellar spots in 1b, one in

2 and one in 6. Length of f.w. 18 mm.

A w.s.f. male from Zambia, sent by Dr. Pin- —

hey, had the following characters. Antenna 34

joints, end joint longer than penultimate; there

are small oval, double grooves on the underside

of each joint of the club; second palpal joint

twice the length of the end joint. Upperside

h.w. ocellar spots in space 2 and 3. Underside

wings darker than the d.s.f., similar to Y. doleta

with striation on h.w. uneven; there is a double

pupilled spot in 1b, a larger one in 2, a smaller in

3 and a large spot in 6.

Two females which must belong to this spe-

cies were found in the BM collection among

specimens of Y. doleta and Y. praestans. The fe-

male has remained undescribed and is here de-

scribed for the first time. Upperside almost as in

Y. doleta, but with only one ocellar spot in the

h.w. in area 2. Antenna 29 joints; palpi second

joint less than twice the length of the third joint.

Underside somewhat similar to a large female

Y. granulosa; on the h.w. there are the usual

ocellar spots as in Y. doleta, a double one in 1b,

one in 2, one in 5 and in 6. Length of f.w. 18 to

18.2 mm, antenna-wing ratio 0.39.

Male genitalia (fig. 87). — Prep. Overlaet and

prep. nos 761A, 2407 from Zambia. Dorsal

aspect: Tegumen-uncus rather long and slender;

uncus tapering to a narrow, distal portion; valva

narrow distal portion bent interiad; accessory

plate robust and furnished with 5 to 7 strong

teeth. Lateral aspect: Tegumen nearly rectangu-

lar-shaped as in Y. praestans and Y. doleta, but

appendices angulares much broader; lower half

of vinculum wide; valva, narrow portion much

shorter, about one third of wide portion; aedea-

gus as long as, or shorter than valva.

Female genitalia (figs. 129—130). — Prep.

nos. 754A no data label; 2040 from Angola. Si-

nus vaginalis wide and deep, squarely shaped;

lateral sclerites broad; anterior plate large, sim-

ple, bilobed distally with rather wavy edge; pos-

terior plate peculiar, roundly fan-shaped with

longitudinal ridges, unsclerotized; anal lobes

with short, blunt apophyses; ductus very short;

bursa small, nearly circular.

Habitat. — Probably similar to that of Y.

praestans.

Distribution (fig. 10). — Katanga, W. Zambia

and Angola.

KieLLAND: Ethiopian Ypthima 143

Ypthima albida Butler

This species is distinguished from all other

known Ypthima by the silvery white ground

colour of the upperside of both wing pairs.

Ypthima albida albida Butler

(figs. 11, 94—96, 111)

Ypthima albida Butler, 1888: 59 (Foda, Equatorial

Africa).

Ypthima albida argentata Bartel, 1905: 134 (Niaru-

gungu, Ruanda). Syn. nov.

Ypthima albida ab. conradsi Strand, 1909: 113 (Uke-

rewe Isl., L. Victoria).

The race argentata Bartel described from

Ruanda, is a typical East African form, similar

to specimens examined from Uganda and Kenya

and specimens from Kibondo and Bukoba,

N.W. Tanzania. A specimen of Y. albida argen-

tata from the type locality in Ruanda, was kind-

ly sent to me on loan by Dr. Hannemann of the

BMG. Photos of the type of Y. albida, of speci-

mens from the type locality of argentata and al-

so of Y. albida uniformis Bartel, were sent to

writer from the BM. It is difficult to determine

from the photos alone and Butler’s type of Y.

albida is rather worn, but I do not think there is

much difference between ssp. albida and speci-

mens from Ruanda, or other parts of East Afri-

ca. There is individual variation with regard to

the width of the marginal dark bands, and size

and number of the h.w. ocellar spots, which is

normal amongst Ypthima.

The f.w. subapical ocellar spot in the albida

type, looks rather obscured, as Butler himself

stated, but this, I think, is due to fading and

rubbing. Aurivillius (1914) says that argentata

only differs from Y. albida albida in the upper-

side h.w. having two eye-spots instead of only

one in Y. albida albida, but as stated before, the

number of eye-spots is very variable in Ypthi-

ma. From this we can conclude that Y. albida

argentata Bartel is a synonym of Y. albida albi-

da Butler.

External characters. — Antenna male and fe-

male 35 joints, brown and white ringed dorsal-

ly, white sprinkled ventrally; club thin and very

gradual, end joint a little longer than penulu-

mate, distal half of each joint double grooved;

palpi second joint two and a half times as long

as the third joint.

Male. Upperside, wings silvery, or whitish;

f.w. with a normal subapical eye spot, with the

ocellar area bordered only on the outside by a

brown submarginal band, which in some speci-

mens fades out near tornus, in others at vein 2;

from the eye spot to the base of the wings the

ground colour is not interrupted; there is a

brown margin which is more or less fused into

the submarginal band; there is also more or less

brown dusting, or striation between the sub-

margin and the eye-spot; costa dusted with

brown; h.w. with margin striated with brown;

in some specimens the striation includes and

obscures the ocellar spots and a submarginal

line which is situated rather close to the margin;

in others the brown submarginal line is clearly

defined and with very little dusting between this

and the margin, which is finely bordered with

brown; cilia brown with slightly paler tps;

there is always a prominent ocellar spot in 2,

mostly also one in 1b, which is more or less

obscured in the dark border; there is often a

third, small spot in 3 and sometimes a plain,

dark spot in 6. Underside dark brown, in some

specimens almost blackish brown, with fine

whitish striation; f.w. with distinct ocellar area,

bordered on the innerside by a median band;

h.w. with very irregular, olive tinted median

and submarginal bands; the submarginal band is

strongly widened in 4 and 5 to a blotch, reach-

ing the margin; a double pupilled spot in 1b,

one in 2 and one in 6. On the underside the

spots are nearly equal in size and there is no

spot in 3, even in specimens with a spot in that

area on the upperside. Length of f.w. 19 to 20

mm, antenna-wing ratio 0.39.

Female. Upperside, wings striated with

brown, particularly the f.w.; f.w. with a more or

less clearly defined, dark inner border to the

ocellar area; the distal quarter of the h.w. gener-

ally more heavily striated than the rest of the

wing, but in some specimens the whole surface

is rather evenly striated. Underside as that of

the male, but the olive blotches and bands more

pronounced. Length of f.w. 20 to 22 mm, anten-

na-wing ratio 0.38.

Male genitalia (figs. 94—96). — Prep. no. 456

from Kibondo, Tanzania; 889 from Fort Portal,

Uganda; 890 from Kaimosi, Kenya; 941 from

Kalinzu, Uganda; 942, 944 from Bugoma and

Katera, Uganda. Dorsal aspect: Tegumen and

uncus short and wide; distal narrow portion of

uncus very short; valva strongly, but evenly

curved interiad, its accessory plate with 4 or 5

teeth. Lateral aspect: Uncus tapering to a sharp

point; vinculum straight; saccus robust, wide;

valva rather robust and strongly hunched dor-

sad; the narrow portion with a ventral concavi-

ty near its juncture with the wide portion; ae-

144 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

deagus basal, open part narrow, the rest wide

and rather straight.

Female genitalia (fig. 111). — Prep. nos. 892,

1009 from Katera Sango Bay, Uganda. Anal

lobes triangular as in Y. asterope; sinus vaginalis

moderate, rather shallow; anterior plate unscle-

rotized, squarish, with uneven edges, distal end

with two shallow emarginations; posterior plate

unsclerotized, short and wide, with an anterior,

central lobe, which is rounded distad; ductus

wide, moderately long; bursa small, obliquely

oblong.

Habitat. — Forest margins and swampy riv-

ersides.

Distribution (fig. 11). — Uganda to W. Ke-

nya and N.W. Tanzania, Burundi and eastern

part of Zaire.

Ypthima albida uniformis Bartel

(figs. 11, 97—99)

Ypthima albida uniformis Bartel, 1905: 134 (Central

Africa).

Ypthima albida uniformis ab. pseudalbida Dutrane,

1945: 94 (Kamitunga).

Ypthima albida uniformis ab. impunctata Dufrane,

1945: 94 (Kamitunga).

External characters. — In Bartel’s description

of this race, he stated that there are no upperside

markings except for some dark dusting at the

basal part of costa and at apex of the f.w. This

corresponds with the photos before me of speci-

mens from Kivu in Zaire, but the two f.w. sub-

apical pupils are present and in one specimen,

also a faint yellow ring around each pupil; the

h.w. has only one ocellar spot in 2. The under-

side resembles that of the nominate race, but

with more obscured h.w. median band and f.w.

ocellar area. Length of f.w. (Bartel) 20 mm.

Female. Upperside, white ground colour less

dusted with brown than in other races; the f.w.

subapical ocellar spot well developed, but the

yellow and outer brown rings unclear; there is a

spot in 2 and a smaller one in 3 of the h.w. Un-

derside as in the male, but white striation better

developed and the f.w. subapical ocellar spot

larger.

Male genitalia (figs. 97—99). — Prep. no

14.50. As nominate race.

Female genitalia. — Prep. no. 1449. As nomi-

nate race.

Distribution. — See fig. 11.

Ypthima albida occidentalis Bartel

(fig. 11)

Ypthima albida occidentalis Bartel, 1905: 134 (North

Cameroun).

Ypthima albida ab. argentoides Strand, 1914: 152

(Dschang, Cameroun).

I have not seen the type of this race, but three

males which apparently belong here, were kind-

ly sent to me on loan, by Mr. M. A. Cornes,

NSPR Institute, Lagos, Nigeria. The specimens

had been taken by R.G.T. Leger, at Obudu Pla-

teau, Northern Nigeria, January 1972. The de-

scription of these is as follows.

External characters. — Male. Antenna as in

ssp. albida, but the dorsal side finely sprinkled

with white; palpi as in the nominate race. Up-

perside wings, ground colour slightly darker

than in ssp. albida; the brown dusting of the

margins darker, but this may be due to the com-

paratively recent capture of the Nigerian speci-

mens, so they have had no time to fade yet. The

f.w. submarginal band reaches tornus and there

is very little white dusting between this and the

margin; the h.w. eye spot in 3 is larger than in

the other races. Length of f.w. 18.5 mm, anten-

na-wing ratio 0.39.

Female. Not known to me, but described by

Bartel (1905) as follows: “. . . unterscheidet sich

von d dadurch, dass die Grundfarbe weisslich,

sehr stark durch dunkle Atome verdüstert ist.

Die gelbe Umrandung des Augenfleckes ist

breiter als beim d.”

Male genitalia as in Y. albida albida.

Note. — Strand’s ab. argentoides may belong

to this race, as the locality suggests. As the type

is lost, nothing can be proved, but Strand’s

statement that the underside is more sprinkled

with white than in ab. conradsi and argentata

(which are forms of Y. albida albida), suggests

that argentoides belongs to ssp. occidentalis.

The equal size of the h.w. upperside ocellar

spots is also an indication.

Distribution (fig. 11). — Cameroun to the

northern part of Nigeria.

Two GENERA SEPARATED FROM YPTHIMA BY

VAN SON

Ypthimomorpha Van Son, 1955

Type species: Ypthima itonia Hewitson, 1865

KieLLAND: Ethiopian Ypthima 145

Ypthimomorpha itonia (Hewitson)

(figs. 179—182)

Ypthima itonia Hewitson, 1865: 287.

Ypthima itonia ab. hoeneli Holland, 1896: 744 (East

Africa).

Ypthima itonia ab. quadriocellata Strand, 1909: 113

(Tanzania, Majıta).

Ypthima itonia ab. microocellata Strand, 1909: 113

(Tanzania, Ukerewe).

Ypthima itonia ab. pluripupillata Strand, 1909: 114

(Central Africa (Zaire)).

Ypthima itonia ab. subocellata Strand, 1909: 114

(Songea, Ungoni in Tanzania).

Ypthima itonia ab. pluriocellata Strand, 1913: 150

(Cameroun, Assoko Jaundestation-Simekoa).

This species is widespread in Africa in savan-

na and savanna-like habitats, particularly in

moist and swampy places. It is variable in shade

of colour, in size and number of ocellar spots,

and many aberrations have been described,

which in reality are of very little value as such.

No subspecific characters can be pointed out

and variation seems to be influenced by season

and environment. One male from Cameroun,

examined by the writer at the NMN, fitted

Strand’s description of Ypthima itonia ab. pluri-

pupillata, with three pupils in the f.w. subapical

ocellar spot. The genital preparation showed a

slight difference in the distal part of valva. In

Ethiopia very large specimens have been taken

in numbers. The genitalia of these correspond

with Y. ıtonıa.

When Van Son separated this species from

Ypthima, one important factor was not men-

tioned. The uncus is loosely attached to the te-

gumen by a membrane and this easily breaks off

during preparation. This character does not oc-

cur in the Ypthima dealt with here. As a matter

of fact, two Thailand species, Ypthima baldus

Fabr. and Y. fasciata Hew. have the same char-

acters of the genitalia as Y. itonia, with regard

to the loose uncus as well as to the valvae. Male

genitalia, figs. 179—181.

Female genitalia, fig. 182.

Mashuna Van Son

Mashuna Van Son, (July) 1955: 159, figs. 178a, b, c

and 179.

Type species: Ypthima mashuna Trimen, 1895.

Ypthimorpha Overlaet, 1955a: 23 (published in De-

cember).

Mashuna mashuna (Trimen)

(fig. 185)

Ypthima mashuna Trimen, 1895: 181, pl. 5, fig. 1

(Mashonaland).

Described from Mashonaland in Rhodesia.

Female genitalia, fig. 185. A very local species,

frequenting swampy areas from Rhodesia to

Natal and Bihe in Angola.

Mashuna upemba (Overlaet)

(figs. 183, 184)

Ypthimorpha upemba Overlaet, 1955a: 23, fig. 16a—c.

Described from Mukana, Buye-Bala, Mubale

and Lusinga at Upemba Park in Katanga. One

male and one female of this species were taken

by the author, July 1967, flying over a swampy

patch just north of Mbisi forest, 2000 m, in Ufi-

pa, Tanzania. March 1978 another five males

and one female were taken in grassy glades of

the Mbisi forest, 2200 m.

Female genitalia, figs. 183—184.

Distribution. — A very local species fre-

quenting swampy areas in Katanga, West Tan-

zania and N.E. Angola.

REFERENCES

Aurivillius, P.O.C., 1914. Rhopalocera. In: Seitz, Die

Gross-Schmetterlinge der Erde, 13: 113—119, pl.

29. Stuttgart.

Bartel, M., 1905. Neue Aethiopische Rhopalocera des

Kgl. Museums für Naturkunde in Berlin. — Nov.

Zool. 12: 134.

Butler, A.G., 1876. On a collection of Lepidoptera

from Abyssinia. — Ann. Mag. Nat. Hist. (4) 18:

481.

——, 1883. On some Lepidoptera from the Victoria

Nyanza. — Ann. Mag. Nat. Hist. (5) 12: 101—

102.

——, 1888. On Lepidoptera from Equatorial Africa.

— Proc. Zool. Soc. Lond. 1888: 59—60.

Carcasson, R. H., 1961. New and little-known Afri-

can Lepidoptera. — Coryndon Mem. Mus., Occ.

Papers 7: 14-15.

Dufrane, A., 1945. Lépidoptères du Kivu. — Bull.

Annls. Soc. r. Ent. Belg. 81:94.

Elwes, J. J. & J. Edwards, 1893. A revision of the Ge-

nus Ypthima with especial reference to the charac-

ters of the male genitalia. — Trans. Ent. Soc.

Lond. 1893: 23—24.

Grünberg, K., 1910. Lepidoptera. In: L. Schultze,

Zoologische und anthropologische Ergebnisse

einer Forschungsreise in Südafrika. —Denkschr.

med.-naturw. Ges. Jena 16: 101.

Hewitson, W. C., 1865. A monograph of the Ypthi-

ma, with descriptions of two new genera of diur-

nal Lepidoptera. — Trans. ent. Soc. Lond. (3) 2:

287.

Holland, W., 1896. List of the Lepidoptera collected

in East Africa, 1894, by Mr. William Chanler and

Leutmand. — Proc. U.S. Nat. Mus. 18: 744.

Hübner, J., 1818. Zutrage zur Sammlung exotischer

Schmetterlinge, bestehend in Bekundigung ein-

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982)

zelner Fliegmuster neuer und rarer nichteuro-

päischer Gattungen. — Augsburg.

Kirby, W. F., 1880. Remarks on New or Interesting

Lepidoptera. — Scient. Proc. Roy Dubl. Soc. 2:

335.

ing, I 1851882, msec, Im: Co GC, lilhrenlberg,

Symbolae Physicae, seu Icones et descriptiones

Corpum Naturalium novorum aut minus cognito-

rum, quae ex initeribus per Libyam, Aegyptum,

Nibiam, Dongalam, Syriam, Arabiam et Habessi-

niam. Pars Zoologia. Symb. Phys., pl. 29 figs.

11—14. — Berlin.

Overlaet, F. G, 1954. Satyrides Africains nouveaux.

— Lambillionea 54: 41—44.

——, 1955a. Exploration du Parc National de l’U-

pemba. — Miss. de Witte 27: 22—24.

— — 1955b. Addendum. Exploration Parc National

de L’Upemba. — Miss. de Witte 27: 85—88.

Strand, E., 1909. Zur Kenntniss afrikanischer Ypthi-

ma-Arten auf Grund des im Berliner Museums

vorhandenes Materials. — Ent. Zeitschr., Stutt.

24: 112—114.

——, 1913. Zoologische Ergebnisse der Expedition

G. Tessmann etc. Lepidoptera. — Arch. Natur-

gesch. 79A (7): 150—151.

, 1914. Lepidoptera aus Kamerun. — Arch. Na-

turgesch. 80A (2): 153.

Trimen, R., 1895. On some new species of butterflies

from tropical and extratropical South Africa. —

Trans. ent. Soc. Lond. 1895: 181.

Ungemach, H., 1932. Contribution à l’étude des Lépi-

doptères d’Abyssinie. — Mem. Soc. Sci. Nat. Ma-

roc 1932: 51.

Van Son, G., 1955. The Butterflies of Southern Africa.

Vol. 2. — Transvaal Museum, Pretoria.

147

Ethiopian Ypthima

KIELLAND

Pri

as)

LO)

{an}

—

lar

(ci

{os}

iss}

Sì

DIE

DES

—6, d, Natal; 7—

Figs. 1—8. Upper and undersides of Ypthima aste

Lebanon);

9, d.s.f., Beirut (

148 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982) |De, 2 7

Figs. 1—8. Upper and undersides of Ypthima species. 1—2, Y. asterope hereroica Grünberg, 6, w.s.f., S.W.

Africa; 3—4, Y. yatta species nova, 4, paratype, Ganale River (Ethiopia); 5—6, Y. yatta species nova, 9, allo-

type, Sheik Hussein, Upper Schebehli River (Ethiopia); 7—8, Y. simplicia Butler, w.s.f., Bole Valley (W. Etho-

pia).

PL:3 KIELLAND: Ethiopian Ypthima 149

Figs. 1—8. Upper and undersides ot Ypthima species. 1—2, Y. simplicia Butler, 2, w.s.f., Bole Valley (W.

Ethiopia); 3—4, Y. simplicia Butler, 3, d.s.f., Arussi Galla (Ethiopia); 5—6, Y. jacksoni species nova, d, para-

150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982) PL. 4

Figs. 1—18. Upper and undersides of Ypthima species. 1—6, Y. rhodesiana Carcasson, Mpanda, Sibweza (Tan-

Va; /——6, 8, des RuahaNataBark(lanzana) Ss long Mpanda, Rukwa (Tanzania); 11—12, d, w.s.f.,

PLD KIELLAND: Ethiopian Ypthima 151

Figs. 1—18. Upper and undersides of Ypthima species. 1—4, Y. antennata antennata van Son; 1—2, 6, w.s.t.,

w.s.f., Mbalmayu (Cameroun); 13—14, @, w.s.f., Lamto (Ivory Coast); 15—18, Y. recta Overlaet; 15—16, d,

Rulenge, Ngara (Tanzania); 17—18, 2, Ntungamu, Ngara (Tanzania).

152 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 5 (1982) Pr.6

Figs. 1—18. Upper and undersides of Ypthima species. 1—8, Y. granulosa Butler; 1—2, d, inland type, 20 miles

North of Tabora (Tanzania); 3—4, d, coastal type, d.s.f., Morogoro (Tanzania); 5—6, d, coastal type, w.s.f.,

Kenya coast; 7—8, ®, coastal type, w.s.f., Mikumi (Tanzania); 9-12, Y. pupillaris pupillaris Butler, Lamto

(Ivory Coast); 9—10, d, w.s.f.; 11—12, 9, d.s.f.; 13—18, Y. pupillaris obscurata subspecies nova; 13-14, 6,

d.s.f., Ngara (Tanzania); 15—16, ®, w.s.f., Mpanda, Kampisa (Tanzania); 17—18, 9, d.s.f., Kigoma, Mahale.

(Tanzania).

Pits 7/

KIELLAND: Ethiopian Ypthima

155

Figs. 1—18. Upper and undersides of Ypthima species. 1—2, Y. impura impura Elwes & Edwards, 6, d.s.t.,

Lamto (Ivory Coast); 3—8, Y. impura paupera Ungemach; 3—4, 4, w.s.f., Kigoma, Ngombe (Tanzania); 5—6,

2, d.s.f., Mpanda (Tanzania); 7, 2, w.s.f. Endebes (Kenya); 8, d, w.s.f., Kigoma, Ngombe (Tanzania); 9—12,

Y. praestans Overlaet, 2 ; 9—10, w.s.f., North of Santa Gomba (Angola); 11—12, d.s.f., Chianga (Angola); 13—

16, Y. doleta Kirby; 13—14, d, Likpe Mate (Ghana); 15—16, 2, Mbalmayu (Cameroun); 17-18, Y. diplom-

mata Overlaet, ?, Northeast of Negola (Angola).

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 5 (1982) PL. 8 |

Figs. 1—14. Upper and undersides of types of Ypthima species. 1—2, 4—5, Y. congoana Overlaet, paratypes;

pulchra Overlaet, paratypes; 7—8, d, Kafakumba; 10—11, 2, Sandoa; 9, 12, Y. praestans Overlaet, d, holo-

type, Sankuru; 13—14, Y. diplommata Overlaet, 3, holotype, Kafakumba.

DEEL 125 AFLEVERING 6 1982

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

MUS. COMP. ZOOL

LIRRARY

JAN 3 1 1983

HARVARD

UNIVERSITY

INHOUD

FER WILLEMSE. — A survey of the Greek species of Poecilimon Fischer (Orhto-

ptera, Ensifera, Phaneropterinae), p. 155—203, figs. 1—272, maps 1A.

| Tijdschrift voor Entomologie, deel 125, afl. 6 Gepubliceerd 31-X11-1982

A SURVEY OF THE GREEK SPECIES OF POECILIMON

FISCHER (ORTHOPTERA, ENSIFERA, PHANEROPTERINAE)

FER WILLEMSE

Laurastraat 67, Eygelshoven, The Netherlands

ABSTRACT

Thirty-four species of Poeclimon are recorded from Greece. Evidence was found that the

variation of some taxa is considerably larger than was understood previously. As a result

some taxa are synonymized: P. albanicus Ramme, 1933, with P. jonicus (Fieber, 1853), P.

veluchianus Ramme, 1933, with P. chopardi Ramme, 1933, P. nitidus Werner, 1932, with P.

thessalicus Brunner von Wattenwyl, 1891, and P. ebneri peristericus Karaman, 1961, with P.

ebneri Ramme, 1933. Reviewed diagnoses are given of P. jonicus (Fieber, 1853), P. chopardi

Ramme, 1933, P. zimmer: Ramme, 1933, and P. hoelzeli Harz, 1966. P. pindos sp.n., P. kli-

suriensis sp.n. and P. jonicus lobulatus ssp.n. are described from the Greek mainland and P.

ikariensis sp.n. from the Aegean island of Ikaria. First records for Europe and Greece are

given of P. anatolicus Ramme, 1933, and for Greece of P. hoelzeli Harz, 1966, P. pancici

Karaman, 1958, and P. miramae Ramme, 1933. Full distributional data and maps are in-

cluded.

INTRODUCTION

During the examination of material of Poecili-

mon it became apparent that our knowledge of

the taxonomy and distribution of the Greek

species is unsatisfactory. Evidence was found

that the variation in some species is considera-

bly larger than was understood previously,

which induced a renewed evaluation of the di-

agnostic characters of some taxa. Opportunity

is taken here to give a survey of the systematics

and faunistics of all Greek species of the genus.

MATERIAL AND ACKNOWLEDGEMENTS

The material on which this study is mainly

based consists of 4,500 specimens representing

about 30 species and preserved in the author’s

collection (CW). Most of this has been collected

in Greece by the author in the years 1963 to

1979, always between the Ist of July and the

30th of August. Full label data (i.e. dates of cap-

ture and names of collectors) are given only of

type-specimens and specimens not collected by

the author. In addition 200 specimens were bor-

rowed from the institutions listed below (abbre-

viations are given in parentheses): Naturhisto-

risches Museum, Wien (NMW); Instituut voor

Taxonomische Zoölogie, Amsterdam (ITZ); Is-

tituto Policattedra di Biologia Animale, Catania

(IBA); Natuurhistorisch Museum, Maastricht

(NMM); Zoologische Staatssammlung, Mün-

155

chen (ZSM); British Museum (Natural Histo-

ry), London (BMNH).

For the loan of material thanks are due to A.

Kaltenbach, Vienna; G. Kruseman, Amster-

dam; M. La Greca and A. Messina, Catania; W.

Schacht, München; D. Ragge and Mrs. L. Pit-

kin, London.

The collecting of material by the following

persons is gratefully acknowledged here: M. C.

& G. Kruseman, A. C. & W. N. Ellis and J.

Duffels, all Amsterdam; L. Blommers, Rhenen;

B. van Aartsen, ’t Harde; J. Smid & F. Smid-

Elbers, Arnhem; L. Willemse and J. Tilmans,

Eygelshoven (the previous all in the Nether-

lands); A. Malicky, Lunz; M. Dethier and N.

Doneux-Stiernet, Chavannes/R.

All measurements are given in mm.

SYSTEMATIC PART

Poecilimon Fischer, 1853

Poecilimon Fischer, 1853: 225. Brunner von Watten-

wyl, 1878: 36; 1882: 257; 11891: 24. Ramme, 1933:

497. Bei-Bienko, 1954: 250. Harz, 1969: 91.

Eupoecilimon Tarbinsku, 1932: 183. Ramme, 1951:

336.

Type-species: Poecilimon superbus Fischer,

18553

Diagnosis. — A general diagnosis of the ge-

156 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

nus can be found in the above-mentioned litera-

ture.

Distribution. — The genus ranges from Sici-

ly, Italy and southern Austria through the Bal-

kans, Turkey, the southern part of the U.S.S.R.

to the Caucasus, extending eastward as far as Si-

biria, Kazakhstan and the highlands bordering

Central Asia, from Altai to Tien Shan, and

southward as far as Iran and Israel.

Remarks. — About 100 species have been de-

scribed. In their revision of the genus, Ramme

(1933) and Beî-Bienko (1954) emphasized the

diagnostic value of a large number of characters.

In the course of the present study it came out

that the amount of variation was not always suf-

ficiently known and some features seem to have

been weighted too excessively. This is here

shown in the cases of P. chopardi, P. zimmeri

and P. jonicus. An explanation of the high vari-

ability of some species may be found in the iso-

lation of the diverse populations, especially of

those occurring in the high mountains. in these

cases we are apparently dealing with vicariant

populations in which the characters change in a

sometimes remarkable degree. The confusion in

the systematics of some taxa is mainly due to

the problem of a correct evaluation of the rate

and stability of the differences, which may be

difficult or impossible to understand, when re-

lated forms cannot be sufficiently studied.

In the present study the following species and

subspecies are recorded from Greece:

P. ornatus (Schmidt, 1849)

P. pancici Karaman, 1958

P. hoelzeli Harz, 1966 (stat. nov.)

P. pindos sp.n.

P. gracilis (Fieber, 1853)

P. obesus Brunner von Wattenwyl, 1878

P. beieri Ramme, 1933

P. nobilis Brunner von Wattenwyl, 1878

P. thoracicus (Fieber, 1853)

P. laevissimus (Fischer, 1853)

P. jonicus jonicus (Fieber, 1853) (= P. albanicus

Ramme, 1933)

P. jonicus lobulatus ssp.n.

P. werneri Ramme, 1933

P. tessellatus (Fischer, 1853)

P. macedonicus Ramme, 1926

P. brunneri (Frivaldsky, 1867)

P. pergamicus Brunner von Wattenwyl, 1891

P. cretensis Werner, 1903

P. ikariensis sp.n.

P. syriacus Brunner von Wattenwyl, 1891

ebneri Ramme, 1933 (= P. ebneri peristericus

Karaman, 1961)

klisuriensis sp.n.

zwicki Ramme, 1939

orbelicus Panic, 1883

miramae Ramme, 1933

anatolicus Ramme, 1933

chopardi Ramme, 1933 (= P. veluchianus

Ramme, 1933)

zimmeri Ramme, 1933

. thessalicus Brunner von Wattenwyl, 1891

(= P. nitidus Werner, 1932)

. propinquus Brunner von Wattenwyl, 1878

. aegaeus Werner, 1932

P. mytilensis Werner, 1932

P. deplanatus Brunner von Wattenwyl, 1891

P. sanctipauli Brunner von Wattenwyl, 1878

P. hamatus Brunner von Wattenwyl, 1878.

The occurrence in Greece of P. geoktschaicus

Shchelkanovtsev, 1910, should be omitted. P.

hadjisarandou Werner, 1938, is considered a

species incertae sedis.

SIDE PRESI

Poecilimon ornatus (Schmidt, 1849)

(figs. 1-9, 199, 200, map 1)

Ephippigera ornata Schmidt, 1849: 184 (type-locality:

S. Karnten).

Poecilimon ornatus; Ebner, 1910: 411. Werner,

19336: 199. Ramme, 19356 MORE

fig. 1, pl. 8) fig. 1, pl. 11) fies de Me MR

Bienko, 1954: 270. Harz, 1969: 119, figs. 32, 268,

270, 271, 319, 322. Willemse, 1977: 52.

Barbitistes fieberi Fieber, 1853: 185 (type-locality:

Triest).

Material studied. — Austria: Hochobir, Potschula-

Sattel, 1500 m, viii.1940, Ramme, 1 6; vii.1957, Kal-

Italy: Trentino, Bordala, 1200 m, 18.viii.1946, A.

Galvagni, 1 d.

Yugoslavia: Juliske Alpe, Komna, 10.1x.1919, S. Se-

liskar, 1 2; Vipava, 17.vi.1963, C. A. W. Jeekel, 1 d,

1 2:7 Ücka, 70011100) ra 21er Amman,

nice, 1300 m, 1 d; Mlinista, 1200 m, 1 d; Bare near

Gorazde, 800 m, 4 6; between Bare and Gorazde, 900

m, 5 6, 1 89s Zelengoray) Suyeska 015 OR AGE

Dragos Sedlo, Nat.Park Sutjeska, 1000 m, 10 d, 8 9;

Biogradsko Jezero, 1400 m, 18.vii.1958, Deeleman, 1

3; between Bukovica and Savnik, 1300 m, 1 6; Cakor

Pass, 1700 m, 1 d; Pesurici, W of Visegrad, 1200 m, 1

d; Prevalac near Bresovica, 1300—1550 m, 13 d, 2

2; Hajla Planina, between Drelje and Pepié, 700—

1400 m & Bregu i Brahimages, 1650 m, 3 &; Mukos

Pass, Babuna R., 1040 m, 2 d, 4 2; Stepanci, Babuna

R., 640 m, 1 ®; 3 km W of Izvor, 400 m, 1 2; Buko-

vik, N of Kicevo, 900 m, 3 d; Staroec, S of Kitevo,

i |

5 6

WILLEMSE: Greek Poecilimon 157

3 4

7 8 9

Figs. 1—9. Poecilimon ornatus (Schmidt), dorsal view of left male cercus. 1, Kärnten, Hochobir; 2, Trentino,

Bordala; 3, Montenegro, Bukovica-Savnik; 4, Serbia, Pesurici; 5, Makedonija, Mukos Pass; 6, Makedhonia, Pi-

sodhéri; 7, Makedhonia, Mt. Vérmion; 8, Makedhonia, Velvendös-Katafiyio; 9, Ipiros, Mt. Tzoumerka.

1000 m, 1 d; Krusje near Resen, 1000 m, 1 d; be-

tween Trepjca and Otesevo, 1500 m, 28 d, 20 2; Go-

lesnica Planina, am Pepelak, 1800 m, vii.1936, Knip-

per, 1 d.

Greece: 2 km E of Alona, 1000 m, 1 6; 4 km E of

Pisodhéri, 1600—1700 M, 37 d, 14 ®; Vérnon Range,

Mt. Bela Voda, 1700-1971 m, 12 6, 8 2; Mt. Vérnon,

6 km SW of Dhrosopiyi, 1200 m, 13 d, 2 2; Mt. Vér-

mion, W of Näousa, EOS refuge “Tria Piyádhia”,

1850 m, 8.vili.1973, La Greca, 6 d (IBA); Mt. Piéria,

2000 m, 7.v111.1973, La Greca, 5 d, 4 2 (IBA); Mt.

Piéria, between Velvendós and Katafiyio, 1400 m, 1

3,1 2; Mt. Piéria, 10 km NW of Fteri, 1500 m, 1 9; :

Gävros, N of Kastoriá, 1000 m, 3 d; Eptakhörı, 1180

m, 10 d, 4 ®; Samarina, 1600 m, 2 d; Mt. Smólikas

above A. Paraskevi, 800—2000 m, 27 6, 4 ®; Mt.

Smólikas, between Foúrka and Samarína, 1500 m, 10

3: Métsovon, 3 km E & 3 km N, 1300—1400 m, 27

d, 12 2; Mt. Tzoumerka above Katarraktis, 1700—

2100 m, 12 8, 13 9 (all CW).

Diagnosis. — See Ramme (1933), Beî-Bienko

(1954) and Harz (1969).

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Variation. — Some variation of the male cer-

cus is noteworthy (figs. 1—9). Basal fold of

lower ovipositor valve (figs. 199, 200) always la-

melliform, protruding horizontally and slightly

arched upward anteriorly, strongly and roundly

impressed above and forming with the gonang-

ulum a deep and round pit. Size and coloration

highly variable. Specimens from Yugoslav and

Greek Makedhonia are usually more varico-

loured than those from the rest of the range. In-

tegument of male abdomen invariably shiny.

Distribution. — The species has a wide range,

which covers S Austria, NE Italy, W & C Yu-

goslavia, NW Greek Makedhonía, Albania and

the northern half of the Píndhos range.

Greek localities. — Makedhonia: — (“Mazedo-

nien”) (Ebner, 1910; Ramme, 1933; Beî-Bienko,

1954; Harz, 1969); Florina: Mt. Kaimakchalan

(Werner, 1933a); 2 km E of Alona, 1000 m; 4 km E of

Pisodhéri, 1600—1700 m; Mt. Bela Voda, 1700—1971

m; 6 km SW of Dhrosopiyí, 1200 m (all Willemse,

1977); Imathia: Mt. Vérmion, W of Náousa, 1350 m

(Willemse, 1977); Kozani: Mt. Vérmion, 1850 m; Mt.

Piéria, 2000 m; between Velvendós and Katafiyio,

1400 m (Willemse, 1977); Piéria: 10 km NW of Ftéri,

1500 m (Willemse, 1977); Kastoriá: Gavros, 1000 m;

Eptakhöri, 1180 m (both Willemse, 1977); Grevená:

Samarína, 1600 m.

Ipiros: Ioannina: between Foúrka and Samarína,

1500 m; Mt. Smólikas above A. Paraskeví, 800—2000

m; Métsovon, 1300—1400 m; Arta: Mt. Tzoumérka

above Katarraktis, 1700—2100 m (all Willemse,

1977). (Map 1).

Remarks. — In Greece the species may occur

together with several other members of the ge-

nus, e.g. P. hoelzeli, P. gracilis, P. jonicus joni-

cus, P. jonicus lobulatus, P. ebneri, P. chopardi,

P. zimmeri, P. thessalicus.

Poecilimon pancici Karaman, 1958

(fig. 10, map 1)

Poecilimon pancici Karaman, 1958: 36, figs.

(type-locality: Gabres bei Kumanovo).

1969: 122.

1—7

Harz,

Material studied. — Greece: Mt. Pangaion above

Akrovoúnion, 1250 m, 11 d,4 2 (CW).

Diagnosis. — See original description.

Variation. — The Greek material is rather

uniform, except for the male pronotum, which

may be more or less raised posteriorly.

Distribution. — Hitherto known only from

the type-locality Gabreë in N Yugoslav Make-

donija. The Greek locality indicates that the

range extends considerably more southeast-

ward.

Greek locality. — Makedhonía: Kavala: Mt. Pang-

aion, 1250 m. (Map 1.)

Remarks. — The above specimens agree com-

pletely with the description of P. pancıcı. The

male cercus (fig. 10) and subgenital plate, the

abdominal terminalia and the pronotum in both

sexes resemble P. ornatus. The fastigium of the

vertex, however, resembles more that of P. affı-

nis, being as narrow as — or often slightly nar-

rower than — the greatest width of the scape,

with parallel margins. Whether P. panaici repre-

sents a valid species is an open question.

The Greek specimens were found in a forest

clearing near a mountain refuge, mainly on ferns

and together with P. orbelicus and P. anatolicus.

Poecilimon hoelzeli Harz, 1966

(figs. 11—14, 128—130, 184— 186, 201, 202,

map 1)

Poecilimon ornatus; Ramme, 1951: 93

Rupa).

Poecilimon ornatus hoelzeli Harz, 1966: 23, figs. 10—

14 (type-locality: Mala Rupa); 1969: 120, fig. 349.

Harz & Kaltenbach, 1976: 331.

Poecilimon sp. aff. ornatus; Willemse,

1977: 52.

(only Mala

1974: 352;

type, labelled: Mala Rupa 1200 m, 20.vu.1917, Poecı-

limon ornatus Schm. det. W. Ramme, Poecilimon or-

natus hoelzeli Harz det. Kurt Harz, Poecilimon hoel-

zeli Harz K. Harz Sept. 1977, type-labels (ZSM).

Greece: Mt. Olimbos, W of Litókhoron, 1750 m,

2.vui.1965, 1 d, 1 2? & Spiliós Agápitos, 2400 m,

Mt. Olimbos, Refuge A between Priòni and Míukas,

1800—2400 m, 10 6,3 2 & 1000-2400 m, 5 6,2 £

& Refuge B above Sparmós, 1800—2300 m, 55 d, 37

9; Leptokaria-Karia, 500—1000 m, 7 6; Mt. Piéria

NW of Ftéri, 1500 m, 3 6; Ftéri, 1000 m, 1 6; 10 km

SW of Dheskäti, 700 m, 1 6; Khrisomiléa, 900—1200

m, 2 d (all CW).

Diagnosis. — Differs from P. ornatus as fol-

lows. Male (figs. 184, 185) of slightly smaller

size; cercus (figs. 11—14) shorter, more robust

over a longer distance, and more strongly in-

curved apically; integument of abdominal ter-

gites dull, and yellowish bands narrower and

more widely separated. Female (fig. 186) slight

ly smaller; basal fold of lower ovipositor valve

(figs. 201, 202) with a large, knob-shaped pro-

cess dorso-anteriorly.

WILLEMSE: Greek Poecilimon 159

19 20 21

Figs. 10—23. Poecilimon species, dorsal view of left male cercus. 10, P. pancıcı Karaman, Mt. Pangaion; ii,

P. hoelzeli Harz; 11, Mt. Piéria, Katafiyio; 12, Mt. Olimbos, Refuge B; 13, Leptokaria-Karia; 14, Khrisomiléa;

15, 16, P. pindos sp. n.; 15, Mt. Timfi; 16, Mt. Mavrovoúni; 17, 18, P. gracilis (Fieber) (Pisodhéri); 19— 22,

P. obesus Brunner von Wattenwyl; 19, Pos; 20, Kapsia; 21, Khrisovitsi; 22, Koutsélioh; 23, P. beter: Ramme

(holotype).

160 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Measurements: body d 21.0—31.0, 9 21.0—

29.0; pronotum d 6.0—8.3, 9 7.2—8.1; elytron

& 42-50, 2 0222 1: mel remuer & 16.5—

21.0, 2 17.2—19.1; ovipositor 14.0—16.0

Variation. — This species is as variable as P.

ornatus.

Pronotum and male cercus as in figs. 11—14,

128130:

Knob-shaped processus of basal fold of lower

ovipositor valve always present and but slightly

variable.

Distribution. — Ranges from the type-locali-

ty Mt. Kozuf (“Mala Rupa”), at the border of

Yugoslav and Greek Makedhonia, southward to

Mt. Piéria and Mt. Ólimbos in S Greek Maked-

honía and extreme NE Thessalia; westward

along S Greek Makedhonía and N Thessalia, ex-

tending into the eastern slopes of the Píndhos

range.

Greek localities. — Makedhonia: Kozani: Mt.

Piéria above Katafiyio, 1600—2190 m; Piéria: Mt.

Piéria, 10 km NW of Ftéri, 1500 m; Ftéri, 1000 m;

Mt. Olimbos above Litökhoron, 1000—2400 m; be-

tween Leptokariá and Karıa, 500—1000 m; Grevená:

10 km SW of Dheskäti, 700 m.

Thessalia: Larisa: Mt. Olimbos above Sparmós,

1800—2300 m; Trikkala: Khrisomiléa, 900—1200 m

(partly Willemse, 1974, 1977). (Map 1.)

Remarks. — Although much resembling P.

ornatus, the stability and importance of the dis-

tinctive characters justify its taxonomic recogni-

tion. Besides, the sympatric occurrence of P.

hoelzeli and P. ornatus forms evidence of the

specific status of both taxa, as was Karaman’s

opinion (Harz & Kaltenbach, 1976: 331).

The species was found sparsely on ferns and

low shrubs of undergrowth and clearings of

woodland and, more abundantly, above the tim-

berline on diverse herbaceous plants with pref-

erence for thistles and stinging nettle. The type-

locality is the Yugoslav part of Mt. Kozuf, for-

merly named Mala Rupa (Doflein, 1921: 96—

118, figs.).

The species may occur together with P. orna-

tus, P. thessalicus, P. jonicus lobulatus and P.

chopardi.

Poecilimon pindos sp.n.

(figs. 15, 16, 131133, 187189, 203, 204,

map l)

belled: Hellas, Pindos Ori, Tymfi above Papikon,

1800—2000 m, 1.vu1.1971, F. Willemse c.s.; para-

types: as holotype, 31 d, 24 2; Mt. Mavrovouni, N

of Metsovon, 1900—2100 m, 3.viii.1971, F. Willemse

CS 10,22 (ACN)

Diagnosis. — Male (fig. 187, 188). Size medi-

um. Integument moderately shiny to dull. Fasti-

gium of vertex sloping, margins parallel, from

slightly more than half to almost as wide as

greatest width of scape. Pronotum (figs. 131—

133) with transverse sulcus cutting median line

before middle of pronotal length; metazona

strongly widening posteriorly, moderately

dome-shaped, raised above tegmina; hind mar-

gin varying between very slightly convex and

distinctly emarginate. Elytra well visible, apical

margin reaching from just beyond fore margin

to just beyond hind margin of second tergite.

Hind margin of all tergites straight. Hind femur

with few ventral spinules. Cercus (figs. 15, 16)

short, robust, apical half regularly and strongly

incurved, outer side near apex gradually nar-

rowing into slightly recurved and downcurved

apical tooth. Subgenital plate relatively short

and wide, hind margin straight, postero-lateral

edges slightly protruding, ventral side with

weak median keel.

General colour green to yellowish green. An-

tennae not at all annulated. Pronotal dorsum, on

either side, with a yellow lateral streak, in meta-

zona bordered medially with rusty brown. Ely-

tra yellowish brown, stridulatory area darker

brown. Abdomen with pair of well separated

yellowish dorsal bands which are often bor-

dered with black along fore margin of tergites,

forming small triangular dots both medially and

laterally. Tip of cercus blackish. Legs not or

finely spotted rusty brown.

Female (fig. 189). Pronotum almost cylindri-

cal, metazona scarcely widened and raised pos-

teriorly. Elytra overlapping dorsally, either just

visible and reaching fore margin of first tergite

or completely covered by pronotum. Basal fold

of lower ovipositor valve (figs. 203, 204) stout,

often slightly inflated anteriorly, moderately

protruding horizontally and strongly arched

upward anteriorly, slightly impressed above and

forming with the gonangulum a shallow round

pit.

Coloration green, more or less brown-

spotted. Lateral pronotal streaks as in male but

narrower and less conspicuous. Abdomen

green, except in one specimen with yellowish

bands as in male, though less sharply defined.

Measurements: body d 18.0—22.5, 9 16.5—

23.0; pronotum d 5.8—7.1, 2 6.2—7.1; elytron

d 4.5—4.8, ® 0.1—1.0; hind femur d 15.2—

WILLEMSE: Greek Poecilimon 161

16.1, 2 15.1—18.2; ovipositor 11.3—13.1.

Variation. — Shape of male pronotum vari-

able (figs. 131—133), but male cercus (figs. 15—

16) and other characters vary only slightly. The

specimens from Mt. Mavrovoúni agree com-

pletely with those of the type-locality.

Distribution. — Hitherto known only from

the northern part of the Pindhos range.

Greek localities. — Ipiros: loánnina: Mt.

Timfi above Papingon, 1800—2000 m; Mt.

Mavrovounı near Metsovon, 1900—2100 m.

(Map 1.)

Remarks. — Readily distinguished from oth-

er species of the genus except P. affinis (Fri-

valdsky, 1867). In his study of the latter species,

Karaman (1974) recognized several subspecies.

Fortunately I have a rich material (183 d, 48 9,

CW) of P. Di. before me belonging to these

ARR and moreover specimens from Bos-

nia-Hercegovina, central Serbia, Montenegro

and Albania. The new species differs from P. af-

finis ( s.1.) in smaller measurements, less shiny

integument, and especially in the size and shape

of the male cercus. Throughout the range ot P.

affinis (s.1.) the male cercus is uniformly much

longer de less incurved apically. It would be

incongruous to arrange the new taxon under P.

affinis as now understood.

The type-locality is the stony slopes sur-

rounding a mountain refuge above the village of

Pápingon, where it occurs abundantly on herba-

ceous plants and low shrubs, together with P.

zimmeri. Near Métsovon it was found together

| with P. ornatus, P. jonicus lobulatus and P. cho-

| pardi.

Poecilimon gracilis (Fieber, 1853)

(figs. 17, 18, 134, 135, 205, 206, map 1)

Barbitistes gracilis Fieber, 1853: 261 (type-locality: Il-

lyrien).

Poecilimon gracilis; Ramme, 1933: 515, pl. 6 fig. 5, pl.

8 fig. 6, pl. 11 fig. 5. Beî-Bienko, 1954: 277. Harz,

1969: 123, figs. 280, 324, 363—365. Willemse,

19772 53%

2 Poecilimon mavrovi; Karaman, 1958: 38, figs. 8—

12. Harz, 1969: 123.

Material studied. — Austria: Obir, Hoffmannsalpe,

14.viii.1927, 1240 m, R. Ebner, 1 2. — Yugoslavia:

Donje Bare, Nat. Park Sutjeska, 1500 m, 17 d, 11 ?;

Kosmet; Drelje-Pepié, 700—1400 m, 1 d; Mavrovi

Anovi near Gostivar, 1100 m, 1 d; Mt. Perister, 1400

m, 31 d, 10 2. — Greece: Vérnon range, Mt. Bela

100271000 19710033028, 252: 4 kin E ot Pr

sodhéri, 1600—1700 m, 54 6, 11 2; Mt. Vernon, 6

km SW of Dhrosopiyi, 1200 m, 3 6,2 9 (all CW).

Diagnosis. — See Ramme (1933), Bei-Bienko

(1954) and Harz (1969).

Variation. — The shape of the pronotum

(figs. 134—135), subgenital plate and cercus

(figs. 17, 18) of the male, and the coloration in

both sexes, are highly variable. The number of

ventral spinules on the hind femur varies from

several to none and may differ even at either leg

of one specimen. The basal fold of the lower

ovipositor valve (figs. 205, 206) is less variable.

Distribution. — The species ranges from S

Austria, W. Yugoslavia, Albania, Yugoslav

Makedonija to the extreme northwestern part of

Greek Makedhonia.

Greek localities. — Makedhonia: Flórina: Vérnon

range: 4 km E of Pisodhéri, 1600—1700 m; Mt. Bela

Voda, 1700— 1971 m; 6 km SW of Dhrosopiyi, 1200

m (all Willemse, 1977). (Map 1.)

Remarks. — Part of the material listed above

agrees with the descriptions of P. mavrovi. The

characters of this taxon, however, seem to coin-

cide with those seen in certain populations of P.

gracilis. The question whether both taxa are

synonymous cannot be settled here.

In Greece the species was found together

with P. ornatus, P. jonicus jonicus, P. ebneri and

P. chopardt.

Poecilimon obesus Brunner von Wattenwyl,

1878

(figs. 19— 22, 136—138, 207, 208, map 1)

Poecilimon obesus Brunner von Wattenwyl, 1878: 38

(type-localities: Epirus and Parnass). Werner,

1929: 481. Ramme, 1933: 513, pl. 6 fig. 2, pl. 8

es > pl ill tie, So 1 ims, 3, deo, Werner,

1934: 324; 1938: 167. Beî-Bienko, 1954: 271.

Harz, 1969: 118, figs. 269, 345 — 348.

Material studied. — Greece: 5 km S of Kalávrita,

750 m, 12.viii.1975, J. Duffels, 1 & (ITZ); Päos, 500

ay Il da asta 750 on, 2 6, il 933 kan Je of Khrisovit-

si, 1200 m, 1 8; Koutseliön, 650 m, 1 à (all CW).

Diagnosis. — See the description in Brunner

von Wattenwyl (1878), Ramme (1933) and

Harz (1969). The description of the female was

incomplete. The elytra of the female widely

overlap, extending beyond hind margin of pro-

notum and reaching first donne tergite.

Basal fold of lower ovipositor valve (figs. 207,

208) stout, protruding downward and weakly

arched upward anteriorly, latero-dorsal surface

weakly impressed, forming with the gonangu-

lum a shallow concavity.

Variation. — The fastigium of the vertex

162 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

varies from slightly wider than to twice as wide

as the greatest diameter of the scape. Some vari-

ation of the male pronotum and cercus, as in

figs. 19—22, 136—138.

Distribution. — Known to occur in the W

and S part of the Greek mainland and the Pel-

opOnnisos.

Localities. — Ipiros: — (“Epirus”) (Brunner von

Wattenwyl, 1878; Werner, 1929; Ramme, 1933;

Harz, 1969); Ioannina: Koutselión, 650 m; Arta: Arta

Ramme, 1933).

Central Greece: Voiotia: Mt. Parnassós (“Parnass”)

(Brunner von Wattenwyl, 1878; Ramme, 1933; Harz,

1969).

Pelopónnisos: Korinthia: between Kórinthos and

Arkh. Kórinthos (“zwischen Paläokorinth und Akro-

korinth”) (Werner, 1934); Argolís: Mycenae (Ramme,

1933; Werner, 1938); Akhaia: Kalávrita (Ramme,

1933); 5 km S of Kalávrita, 750 m; Mt. Panakhaïkón

(“Voidea”) (Werner, 1929); Páos, 500 m; Arkadhía:

Khrisovitsi, 1200 m; Kápsia, 750 m. (Map 1.)

Remarks. — This species is an early one,

adults having been collected in May and June

(Werner, 1929, 1934, 1938). Apparently it dis-

appears soon afterwards as our specimens taken

in July were already dead or almost so. In the

Pelopónnisos we found it together with P. tes-

sellatus. From Mt. Parnassós also P. chopardi is

known.

Poecilimon beieri Ramme, 1933

(figs. 23, 209, 210, map 1)

Poecilimon beieri Ramme, 1933: 514, pl. 6 fig. 4, pl. 8

fig. 5, pl. 11 fig. 4 (type-locality: Insel Meganisi).

Ebner, 1954: 553. Harz, 1969: ;120, figs. 272, 320,

350.

Material studied. — d holotype, labelled: I. Mega- :

nisi, 27.v.1932, Beier, auf Disteln, Poecilimon beieri

Ramme Ramme det., Typus; ® allotype, labelled:

Umg. Levkas, 18—22.v.1933, Beier, Poecilimon beieri

Ramme Ramme det., Typus (both NMW).

Diagnosis. — See the original description.

Elytra of female widely overlapping. Basal fold

of lower ovipositor valve (figs. 209, 210) re-

minding of P. obesus. Cercus of holotype as in

ire 23%,

Variation. — Insufficiently known.

Distribution. — Known only from the Ionian

islands of Meganisi and Levkás.

Localities. — Ionian Is.: Meganisi and Levkas (both

Ramme, 1933: Ebner, 1954; Harz, 1969). (Map 1.)

Remarks. — The description is based on two

males and one female only, which differ from P.

obesus merely in the width of the fastigium of

the vertex. More material is needed to establish

Its true status.

Poecilimon nobilis Brunner von Wattenwyl,

1878

(figs. 24—26, 182, 183, 211, 212, map 1)

Poecilimon nobilis Brunner von Wattenwyl, 1878: 50

(type-localities: Athen; Taygetes), Werner, 1933a:

401, Ramme, 1933: 513, pl. 6 fig. 3, pl. 8 fig. 4, pl.

12 figs. 4, 4a, Werner, 1937b: 146, Bei-Bienko,

1954: 269, fig. 154, Harz, 1969: 118, figs. 267,

342—344.

Poecilimon holtzi Werner, 1902: 116, 2 figs. (type-lo-

cality: Kambos); 1929: 481; 1933a: 401.

Material studied. — Morea, Kambos, v.1901,

Holtz, 1 6 (topotype of P. holtzi) (NMM); Sparti-

Kalamai, 1200 m, 2 d 2 2; Mt. Taiyetos, Tópitza,

1200 m, £ & above Tópitza, 1700— 2000 m, 19 & 20

2; Mt. Párnon above Vamvakou, 1500—1935 m, 2 ó;

Mt. Aroánia above Kalávrita, 1700—2200 m, 1 2; Mt.

Panakhaïkón above Zástova, 1000—1650 m, 1 d; Mt.

Mt. Mainalon above Kardhará, 1600—1800 m, 35 d 8

?;3 km W & 5 km E of Khrisovitsi, 1200 m, 1 6 7 9:

Diagnosis. — See the descriptions in Ramme

(1933), Bei-Bienko (1954) and Harz (1969).

Variation. — Hind margin of last abdominal

tergite of male extending arcuately to a various

degree (figs. 182, 183). This character is less

conspicuous or not at all developed in the fe-

male. Variation of the male cercus as in figs.

24-26. Basal fold of lower ovipositor valve

(figs. 211, 212) scarcely variable. Quite vari-col-

oured specimens may occur. A few males and

females before me have a general reddish brown

colour instead of green.

Distribution. — Confined to the Pelopónni-

sos and the southeastern part of Central Greece.

Localities. — Central Greece: Attiki: Athinai

(“Athen”) (Brunner von Wattenwyl, 1878; Ramme,

1933).

Pelopónnisos: Korinthía: Arkh. Kórinthos (“Ebene

von Paläokorinth”) (Werner, 1933a); Akhaia: Mt

Aroänıa above Kalávrita, 1700—2200 m; Mt. Panak-

haïkón above Zástova, 1000—1650 m; Mt. Eríman-

thos above Kaléntzi, 1700—2200 m; Arkadhia: Mt.

Mainalon above Kardhará, 1600—1800 m; 3 km W &

5 km E of Khrisovitsi, 1200 m; Messinia: Bassae, 1000

m; Kámbos (Werner, 1902; Ramme, 1933; Beî-Bien-

ko, 1954); Exokhöri (“Xechori”) (Werner, 1937b);

Lakonia: between Sparti and Kalamai, 1200 m; Topit-

sa, 1200 m; Mt. Taïyetos (Brunner von Wattenwyl,

1878; Ramme, 1933), above Tópitsa, 1700—2200 m;

WILLEMSE: Greek Poecilimon 163

24 25 26 27 28

29 30 31 32 33 34

Figs. 24-40, Poecilimon species, dorsal view of left male cercus. 24—26, P. nobilis Brunner von Wattenwyl; 24,

Mt. Mainalon; 25, Mt. Taiyetos; 26, Mt. Erimanthos; 27, 28, P. thoracicus (Fieber); 27, Túkheron; 28, Amórion;

29, 30, P. laevissimus (Fischer); 29, Kallithéa; 30, Ano Exänthia; 31—40, P. jonicus jonicus (Fieber); 31, Dhassia;

32, 33, Gazätika; 34, Perithia; 35, Asfika; 36-37, Mt. Smélikas; 38, Albania, Korab (holotype P. albanicus); 39,

Dhrosopiyi; 40, Mt. Perister.

Mt. Párnon above Vamvakoú, 1500—1935 m. (Map

1.)

Remarks. — With P. obesus and P. beieri it

shares the shape of the basal structure of the

lower ovipositor valve. It was found on diverse

low shrubs and herbaceous plants both in

woodland and above the timberline and may oc-

cur together with ?. tessellatus, P. zimmer and

P. laevissimus.

Poecilimon thoracicus (Fieber, 1853)

(figs. 27, 28, 213, 214, map 1)

Barbitistes thoracicus Fieber, 1853: 176 (type-locality:

erroneously Sicilien).

Poecilimon thoracicus; Ramme, 1933: 523: pl. 6 fig.

164 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

2, pl. 8 fig. 13, pl. 11 fig. 11, pl. 12 fig. 8. Beî-

Bienko, 1954: 295, fig. 171. Harz, 1969: 124, figs.

284, 330, 366—368.

Material studied. — Romania: Domogled, 1600—

Donje Bare, Nat. Park Sutjeska, 1500 m, 12 d, 21 9;

near Dzigolj, 1400 m, 5 d,6 ®; Pesurici, W of Vise-

grad, 1200 m, 10 d, 12 ®; Hajla Planina, Drelje-Pe-

pic, 700—1400 m & above Pepié, 1100— 1500 m, 24

d, 7 2; Prevalac, near Brezovica, 3 3; Mt. Bistra,

d,6 2; Ljuboten, N of Tetovo, 1000 m, 9 d,6 9;

Vratnica, N of Tetovo, 2 ó.

Albania: Pastrik, 26.viu.1918, Ebner, 1 d. —

Greece: Amörion, 40 m, 1 à, 1 2; Tukherén, 40 m, 1

3,3 2 (all CW).

Diagnosis. — See the descriptions in Ramme

(1933), Bei-Bienko (1954) and Harz (1969).

Variation. — Throughout its range the varia-

tion is remarkably slight. Male cercus and basal

fold of lower ovipositor valve as in figs. 27, 28,

2155214;

Distribution. — From Romania and Yugosla-

via to Albania and Bulgaria, extending into N

Greece.

Greek localities. — Thráki: Évros: Tukherón, 40

m; Amorion, 40 m. Makedhonia: (Bei-Bienko, 1954;

Harz, 1969).

Ipiros: (Harz, 1969).

Ionian Is.: Kérkira (“Corfu”) (Bei-Bienko, 1954).

(Map 1.)

Remarks. — Confirmation of its occurrence

in the area extending from western Greek

Thraki to Ípiros is needed. In Amörion (Thráki)

the species was found together with P. zwick:.

Poecilimon laevissimus (Fischer, 1853)

(tas. 29, 30, 215, 216, map 2)

Odontura laevissima Fischer, 1853: 225, pl. 12 figs. 5,

5*, 5a (type-locality: Messina, Sicilia).

Poecilimon laevissimus; Ramme, 1933: 533, pl. 6 fig.

21, PL 25, PL ih ine, 19), jal 12 tie, 168 19998

46. Ebner, 1954: 553. Harz, 1969: 137, figs. 288,

326, 420—423. Willemse, 1977: 53.

Material studied. — Greece: Levkás, Áno Exánthia,

2.v1.1977, A. Malicky, 3 &; Kallithéa, S of Lidhor-

ikion, C. Greece, 100 m, 18.vi.1979, J. Smid & F.

Smid-Elbers, 1 6, 1 2; Ilia, Kallithéa, 400 m, 9 g, 6

9; Bassae, 1100 m, 11 8, 10 ®; Karitaina, 500 m, 1 ®;

Kambos, 400—700 m, 3 d, 2 9: Tópitsa, 1200 m, 8

d,10 9 (all CW).

Diagnosis. — See the description in Ramme

(1933, 1939) and Harz (1969). The basal fold of

the lower ovipositor valve (figs. 215, 216) was

not yet described: strong, well protruding hori-

zontally and strongly arched upward anteriorly,

broadly and strongly impressed from above and

forming with the gonangulum a large deep

round pit. Female subgenital plate divided by a

deep and wide i impression separating a pair of

bulbous, roughly semi-circular lobes.

Variation. — The Greek material is compara-

tively uniform. The male cercus (figs. 29, 30)

varies slightly in length. Colour of female rang-

ing from uniform green to as distinctly variegat-

ed as in male. Rusty brown dorsal spots of first

and second abdominal tergites of male invari-

ably present.

Distribution. — Known from Sicilia, some

Ionian islands, Central Greece and the western

part of the Pelopónnisos.

Greek localities. — Ionian Is.: Meganísi (Ramme,

1933; Ebner, 1954; Harz, 1969); Levkas: Nidhrion

(“Nidri”) (Ebner, 1954; Harz, 1969); Ano Exanthia

690 m (Willemse, 1977); Zakinthos: 2 km N of Zákin-

thos (“Zante”) (Ramme, 1939; Harz, 1969); S of A.

Nikélaos, 140 m. Central Greece: Fékis: Kallithéa,

100 m. Pelopónnisos: Ilia: Kallithéa, 400 m; Bassae,

1100 m; Arkadhía: Karítaina, 500 m; Messinía:

Kambos, 400—700 m; Lakonia: Töpitsa, 1200 m (all

Willemse, 1977). (Map 2.)

Remarks. — Ramme found the species on As-

tragalus (Zäkinthos). We collected it on various

low shrubs, e.g. Rubus, Quercus. In Bassae and

Tópitsa (both in the Pelopónnisos) the species

occurred together with P. nobilis.

Poecilimon jonicus Fieber, 1853

Among our rich material two forms could be

recognized, differing in the shape of the basal

fold of the lower ovipositor valve. One of these

forms agrees with both P. jonicus and P. albani-

cus, which therefore are considered conspecific,

while the other is described below as a new sub-

species.

Poecilimon jonicus jonicus (Fieber, 1853)

(figs. 31-40, 139—144, 170-172, 217, 218,

map 2)

Barbitistes jonicus Fieber, 1853:

Corfu).

Poecilimon ionicus; Brunner von Wattenwyl, 1878: 49

(partim). Harz, 1969: 131, figs. 278, 325, 395—

397 (not Pelopónnisos).

Poecilimon jonicus; Brunner von Wattenwyl, 1882:

260 (partim). Ramme, 1933: 529, pl. 8 fig. 21, pl.

175 (type-locality:

WILLEMSE: Greek Poecilimon 165

11 fig. 17, pl. 12 fig. 12 (partim). Beî-Bienko,

1954: 294 (not Pelopónnisos).

Poecilimon albanicus Ramme, 1933: 531, pl. 6 fig. 19,

pl. 8 fig. 23, pl. 12 fig. 14 (type-locality: Korab,

Albania). Karaman, 1958: 41, figs. 18—21. Harz,

1969: 130, figs. 294, 386— 389. Syn. nov.

Material studied. — Albania: Alban. Exp. 1918,

Korab, 23—31.vii., Poecilimon albanicus Rme Ramme

cilimon albanicus) (NMW).

Yugoslavia: Makedonija: Staroec, S of Kicevo, 1000

m, 11 6, 6 2; Krusje near Resen, 1000 m, 4 d, 3 9;

10 km E of Resen, 1100 m, 7 8,6 ®; Trepjca-Otese-

vo, 1100 m, 16 d, 12 2; Mt. Perister, 1100 m & 1400

m, 19 d, 13 9 & 1800—2000 m, 31.vu.1965, Ent.

Greece: Mt. Vérmion, refuge Tria Pigádhia, W of

Näousa, 1350 m, 1 6, 2 2; Seli, near refuge Mt. Vér-

mion, 1600 m, 1 ® (all CW); Kozani, Mt. Vermion,

1200 m, 58 d, 40 2; Mt. Vérnon, Polipótamos, 1100

HG PE Eptakshóri, 1180 m, 18 6, 9 9; Lithia,

5 2; Mt. Smölikas above A. Paraskevi, 800—2100 m,

37 à, 22 2; Elevtherön, 1000 m, 7 6, 2 2; Kónitsa-

Elevtheron, 1200 m, 2 d; Aristi-Papingon, 450 m, 9

3,6 2; Pápingon, 1000 m, 14 d 2 ®; Kalpákion, 400

types of P. jonicus): Dhassia, 5 km SE of Korakiana,

16—30.v.1971, B. v. Aartsen, 1 d; Perithia, 450 m, 8

6,5 2; Lake Antinioti, Om, 3 2; Tsaki, 10 m, 4 6,3

9; Livadhı plain, 10 m, 1 6, 1 2; Láfkion, 100 m, 1

2 (all CW).

Diagnosis. — Male. Size medium to small. In-

tegument shiny. Pronotum (figs. 139—144)

long, not at all saddle-shaped, not or scarcely

widened posteriorly, metazona very slightly

raised and scarcely bulbously inflated, lower

margin of lateral lobe from widely rounded to

scarcely S-shaped posteriorly. Elytra extending

beyond hind margin of pronotum, reaching

middle of first tergite or slightly longer. Cercus

(figs. 31—40) slender, slightly narrowing distal-

ly, sometimes with a slight pre-apical widening;

distal half more or less evenly incurved, cylin-

drical but at inner side of incurvation slightly

flattened, outer side of apex roundly narrowing

into a simple acute tip or tooth. Subgenital plate

(figs. 170—172) longer than wide, with an ob-

tuse median and a pair of lateral keels, margins

converging posteriorly toward narrow, trans-

verse or weakly concave hind margin, postero-

lateral edges slightly or not produced.

General colour yellowish, yellowish-green or

green, more or less spotted with dark brown or

black. Antennae finely annulated. Pronotal dor-

sum of general colour, or often more or less rus-

ty brown with on either side a yellow lateral

line. Elytron of general colour, stridulatory part

sometimes darker brown. Abdomen with more

or less developed black median band, composed

of a pair of black dots along fcre margin of ter-

gites, the dots extending posteriorly without

reaching hind margin of tergites and always sep-

arated from each other by a yellow median line.

Lateral side of abdomen more or less spotted

black, without forming a true black lateral band.

Cercus brown, often blackish or dark brown

apically. Lower outer keels of all femora black,

upper side of fore and middle femora and upper

and outer side of hind femur often with a black

line.

Female. Integument less shiny than in male,

though not dull. Pronotum almost cylindrical,

lower margin of lateral lobe from straight to

weakly rounded posteriorly, not S-shaped. Ely-

tra widely overlapping, usually extending well

beyond hind margin of pronotum. Basal fold of

lower ovipositor valve (figs. 217, 218) thin, la-

melliform, distinctly protruding horizontally,

arched upward anteriorly and not or scarcely

inflated, roundly impressed from above and

forming with the gonangulum a distinct round

pit.

Coloration as in male, more often less bright-

ly so or even completely uniform. Ovipositor in

rather varicoloured specimens with a black dor-

so-lateral spot.

Measurements: body & 13.5—20.0, 2 13.0—

22.0; pronotum d 4.3—5.7, 2 5.0—6.3; elytron

6 ji, 2 06-17 inc eme @ 15{0—

18.6, 2 16.1—19.1; ovipositor 8.4—10.2.

Variation. — A highly variable species. Male

cerci, including those of topotypical specimens,

often not widened preapically (figs. 31—40).

Size and shape of pronotum and subgenital plate

of male (figs. 139—144, 170—172) rather vari-

able. Specimens from the western part of the

range viz. from the Ionian island of Kérkira and

the lowlands of Ipiros are large and brightly

coloured, while those from the eastern part are

smaller and less conspicuously coloured.

Distribution. — The range of the nominate

subspecies extends from Dalmatia, Montenegro,

Albania and SW Yugoslav Makedonija to NW

Greek Makedhonia, the northern part of the

Pindhos range and Ipiros, and reaches the Ioni-

an island of Kérkira.

Greek localities. — Makedhonia: Imathia: Mt. Vér-

mion, mountain refuge of Séli, 1600 m; Mt. Vérmion,

166 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

41 42 43

45°

CEL

51 52 53 54

Figs. 41—54. Poecilimon species, dorsal view of left male cercus. 41—44, P. jonicus lobulatus ssp. n.; 41, Mt.

Mavrovouni; 42, Khrisomiléa; 43, Ramia-Livadhion; 44, Niköpolis; 45, P. werneri Ramme (Lekhaina) (for

comparison 45a, P. jonicus jonicus (Fieber), Gazátika); 46—49, P. tessellatus (Fischer); 46, Ipsous; 47, Mt. Aroa-

nia; 48, Mt. Maínalon; 49, Mt. Panakhaikön; 50—52, P. macedonicus Ramme; 50, Litékhoron; 51, 52, Mt. Kho-

lomón; 53, 54, P. brunneri (Frivaldsky); 53, Titov Veles; 54, P. Pletvar.

refuge Tria Pigadhia, W ot Náousa, 1350 m; Kozanı:

Mt. Vermion, 1850 m; Flörina: 2 km E of Alona, 1000

m; Mt. Vérnon, 6 km W of Dhrosopiyi, 1200 m &

olipötamos, 1100 m; Kastoria: Lithia, along Lake

Kastoria, 700 m; Eptakhòri, 1180 m.

Ipiros: — (“Epirus”) (Brunner von Wattenwyl,

1878, 1882; Ramme, 1933; Bei-Bienko, 1954; Harz,

1969); loánnina: Fourka, 1400 m; Mt. Smólikas above

A. Paraskeví, 800—2100 m; Elevtherón & between

Elevtherön and Könitsa, 1000 & 1400 m; between

Aristi and Pápingon, 450 m; Pápingon, 1000 m;

Kalpákion, 400 m; Asfáka, 500 m.

Ionıan Is.: Kérkira: — (“Corfu”) (Fieber, 1853;

Brunner von Wattenwyl, 1878, 1882; Ramme, 1933;

Bei-Bienko, 1954; Harz, 1969); Dhassia; Perithía, 450

m; Lake Antinioti, 0 m; Petália, 500 m; Láfkion, 100

m; Livadhi plain, 10 m; Tsakı, 10 m; Gazätika, 20 m.

(Map 2.)

Remarks. — Specimens from the mountains

agree with P. albanicus, those from the island of

Kérkira with P. jonicus, while the rest of the

material is intermediate. Presumably P. jonicus

is a highly variable species and P. albanicus

merely represents the mountain form of the for-

WILLEMSE: Greek Poecilimon

mer. I consider both taxa conspecific and pro-

pose to synonymize P. albanicus Ramme, 1933,

| with P. jonicus (Fieber, 1853). From the original

| description of P. ghighii Salfi, 1937, it seems

probable that this is another synonym of P. jo-

nicus. This cannot be confirmed in the absence

of the types.

Few records from the Pelopónnisos under the

name P. jonicus (Ramme, 1933; Harz, 1969) are

assigned in this study to P. tessellatus.

The species lives from the lowlands up to

above the timberline. We found it, often abun-

dantly, in diverse habitats, e.g. hedges, open

grassland, woodland clearings and lush mead-

ows. It occurs on low bushes and herbaceous

plants, such as thistles, ferns, stinging nettle,

Verbascum, Sambucus, Rubus and many others.

In Greece it may occur together with other

members of the genus e.g. P. ornatus, P. gracilis,

P. chopardi, P. ebneri.

Poecilimon jonicus lobulatus subsp. nov.

(figs. 4144, 145—150, 190—192, 219—222,

map 2)

Material studied. — ® holotype, d allotype, la-

belled: Hellas, Ep. Dhodhénis, 8 km N of Kaléntzion,

950 m, 22.vii.1976, F. & L. Willemse & J. Tilmans

(CW); paratypes: Aristi, 700 m, 2.vi.1973, M. La

Greca, 2 6, 1 ® (IBA); Mt. Mitsikéli above Ling-

iádhes, 850—1650 m, 3.viii.1978, F. Willemse, 1 à, 2

2: Métsovon, 3 km E, 1300 m, 25 d, 11 2 &6 kmE,

1500 m, 5 6, 3 2, 9.vili.1966, F. Willemse c.s. & J.

Scherpbier, & 5—10 km N, 1600 m, 3.vin.1971, F.

Willemse c.s, 31 6, 15 2 & 3 km N, 1400 m,

11.vii.1976, F. & L. Willemse & J. Tilmans, 14 d, 13

2 (all CW); Métsovon, 1600 m, 1.v11.1973, M. La

von, 1600—2100 m, 3.viii.1971, F. Willemse c.s., 11

d,6 2; Khrisomiléa, 900—1200 m, 10.v11.1976, 24 d,

25.v1.1976, 1 3; Miléa-Katarrákus, 750 m,

25.vu.1976, 2 d, 2 2; Mt. Tsoumérka above Ka-

Rámia-Livádhion, 500 m, 25.vii.1976, 21 d, 15 ®;

L. Willemse & J. Tilmans) (all CW).

Diagnosis. — Differs from the nominate sub-

species in the female only: basal fold ot lower

ovipositor valve bulbously inflated anteriorly,

forming a large round processus (figs. 190—

192, 219—222).

17.0— 22.5; pronotum d 4.2—6.3, 2 4.5—7.2;

elytron d 1.1—2.5, ® 0.1—1.6; hind femur d

14.1—19.8, 2 14.1—21.0; ovipositor 8.2—11.0.

Variation. — Most of the characters, e.g. size,

167

coloration, length of elytra in both sexes, shape

of the pronotum (figs. 145—150), subgenital

plate and cercus (figs. 41—44) of the male, are

as variable as in the nominate subspecies. Large

and brightly coloured specimens are from the

lowlands, while those from the high mountains

are smaller and less brightly coloured. The pro-

cessus of the basal fold of the lower ovipositor

valve presents some clinal variation, being con-

spicuously more developed in southern popula-

tions than in northern ones (figs. 219—222).

Distribution. — The range of the subspecies

adjoins that of the nominate subspecies and cov-

ers the central part of the Pindhos range, ex-

tending into S. Ipiros.

Localities. — Ipiros: Ioannina: Aristi, 700 m; Mt.

Mitsikéli above Lingiádhes, 850—1650 m; surround-

ings of Métsovon, 1300—1600 m; Mt. Mavrovoúni

near Métsovon, 1600—2100 m; 8 km N of Kalént-

zion, 950 m; Arta: between Miléa and Sgára, 600 m;

between Miléa and Katarraktis, 750 m; Mt. Tsoumér-

ka above Katarraktis, 850—2100 m; between Ramia

and Livadhion, 500 m; Preveza: Nikópolis, 900—

1200 m.

Thessalia:

(Map 2.)

Trikkala: Khrisomiléa, 900—1200 m.

Remarks. — While the female may be readily

distinguished from both the nominate subspe-

cies and other species resembling P. jonicus, the

males of both subspecies are indistinguishable.

P. jonicus lobulatus occurs from the lowlands

up to above the timberline in similar habitats as

the nominate form. It may occur together with

P. chopardi, P. zimmeri, P. ornatus, P. pindos

and P. hoelzelı.

Poecilimon werneri Ramme, 1933

(figs. 45, 151, 152, 223, 224, map 2)

Poecilimon werneri ne 1933: 530, pl. 6 fig. 18,

pl. 7 fig. 7, pl. 8 fig. 22, pl. 11 fig. 18 (type-locali-

ty: AGiion), ÉD MES wes, 205 —

414.

(CW).

Diagnosis. — See the original description.

The basal fold of the lower ovipositor valve,

hitherto not described, is thin, lamelliform, pro-

truding horizontally, slightly arched upward

anteriorly, weakly impressed from above and

forming with the gonangulum a shallow con-

cavity (figs. 223, 224).

Variation. — Insufficiently known by lack of

material. Male pronotum variable, as shown in

168 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

figs. 151—152. The ovipositor of the female

studied lacks a black spot and is unicoloured.

Distribution. — Known only from the type-

locality, Agrinion in western central Greece and

from Lekhaína, western Pelopónnisos. (Map 2.)

Remarks. — This species differs from P. joni-

cus in the male cercus and the basal fold of the

lower ovipositor valve. While the cercus in P.

jonicus is slightly flattened at the inner side of

the incurvation (fig. 45a), in P. werneri it is not

at all flattened but perfectly cylindrical (fig. 45).

Basal fold of lower ovipositor valve in both sub-

species of P. jonicus distinctly more impressed

from above than in P. werneri, in P. jonicus lo-

bulatus moreover with a large anterior pro-

cessus. The distinction between P. werneri and

P. tessellatus refers to the same characters, and

also to the pronotum, which is appreciably

more inflated posteriorly in the latter.

We found this species on a hedge bordering

cultivated land along the highway just north of

the village of Lekhaina.

Poecilimon tessellatus (Fischer, 1853)

(figs. 46—49, 153—155, 225, 226, map 2)

Odontura tessellata Fischer, 1853: 227, pl. 12 figs. 7,

7a-b (type-locality: erroneously Ticino, Helvetia).

Poecilimon tessellatus; Ramme, 1933: 575. Harz,

1964: 447, figs. 11—17; 1969: 136, figs. 290-291,

415—417.

Poecilimon walter: Werner, 1937b: 145, figs. 3—4

(type-locality: Mykenä); 1938: 166, fig. 3.

Poecilimon jonicus; Werner, 1927: 428. Ramme, 1933:

529 (partım).

Poecilimon spec.; Ebner, 1912: 109.

Material studied. — Greece: Attika, 1858—1862,

Heldreich, 1 &; Poros I., 1870, Heldreich, 1 d; Mt.

Aroanıa above Kalávrita, 1700—2000 m, 21 6,7 ®;

Mt. Panakhaikön above Romanoü, 1000 m & above

Zástova-Psarthrí, 1000—1650 m, 4 d, 2 2; Ipsoús,

800 m, 4 6d, 29; 5 km E & 3 km W & 12 km W of

above Kardharás, 1250—1981 m, 49 d, 36 ® (all

CW); Käpsia, 12.vii.1976, J. P. Duffels, 1 3, 29

(ITZ); Mt. Parnon above Kastánitsa, 1500—1800 m, 5

8, 4 2 (CW); 13 km E of Langadhá, 1000 m,

15.v11.1976, J. P. Duffels, 8 4,4 2 (ITZ).

Diagnosis. — See the descriptions by Harz

(1964, 1969).

Variation. — Size and coloration are highly

variable. The usually solid black pigmentation

of the abdomen may be distinctly less devel-

oped, recalling P. jonicus. The black lateral

bands of the abdomen, however, are invariably

present. Male pronotum and cercus varying, as

in figs. 46—49, 153—155. Basal fold of lower

ovipositor valve (figs. 225, 226) similar to that

of nominate P. jonicus and hardly variable.

Distribution. — The range of the species cov-

ers the Peloponnisos, including some offshore

islands of Argolis, and extends into Atukí and

the Saronic islands.

Localities. — Central Greece: Attiki: — Saronic

Is.: Aiyina (Harz, 1964; 1969).

Pelopónnisos: Argolis: Mikinai (“Mykenä”) (?

Ebner, 1912; Ramme, 1933; Werner, 1937b, 1938;

Harz, 1964, 1969); Póros I. (Harz, 1964, 1969); Spét-

sai I. (Harz, 1964, 1969); Akhaía: Mt. Aroanıa above

Kalávrita, 1700—2000 m; Mt. Panakhaikén above

Romanoú & between Zástova and Psarthrí, 1000—

1650 m; Ilia: Olimpia (? Ebner, 1912); Arkadhía: Vit-

ina (Werner, 1927; Ramme, 1933); Ipsoús, 800 m;

surroundings of Khrisovitsi, 1100—1200 m; Kapsia;

Mt. Párnon above Kastánitsa, 1500—1800 m; Lako-

nia: 13 km E of Langádha, 1000 m. (Map 2).

Remarks. — Near P. jonicus, P. werneri, P.

superbus (Fischer, 1853) and some other species.

Readily distinguished by the pronotum (figs.

153—155), which is more saddle-shaped and

more bulbously inflated posteriorly, the lower

margin of the lateral lobe being more S-shaped

than in any of the allied species. In P. tessellatus

the elytra are slightly shorter than in P. jonicus

and the unmodified basal fold of the lower ovi-

positor valve is quite distinct from P. jonicus lo-

bulatus. The male cerci in P. tessellatus and P.

jonicus are much the same: slightly flattened at

the inner side of the incurvation instead of per-

fectly cylindrical as in P. werneri.

Previous records of P. jonicus from the Pel-

opónnisos are here referred to P. tessellatus. The

occurrence of the former species in this area is

not probable since its range does not seem to

extend south of Ipiros.

We found the species below the timberline

often on Quercus (phrygana habitat) and above

timberline on various herbaceous plants, often

together with P. nobilis and P. zimmeri.

Poecilimon macedonicus Ramme, 1926

(figs. 50—52, 227, 228, map 3)

Poecilimon macedonicus Ramme, 1926: 280, figs. 3b,

4b (type-locality: Plaus bei Hodowa, Yugoslav

Makedonija); 1933: 527, pl. 8 fig. 17. Weidner,

1950: 181 Beî-Bienko, 1954: 299, fig. 173. Harz,

1969: 135, figs. 283, 406-409.

Poecılimon fussi; Berland & Chopard, 1922: 167.

Ramme, 1951: 95.

Poecilimon elegans; Uvarov, 1923: 147.

WILLEMSE: Greek Poecilimon 169

Material studied. — Yugoslavia: Makedonya: Ko-

zuf Planina (Konsko), 800 m, 7.vin.1966, M. Kara-

man, 1 d, 1 2; Teovo, 22 km SW of Titov Veles,

26.vii.1965, Ent. Exc. Zool. Mus. Amsterdam, 2 6;

Suvodol, 18 km ENE of Bitola, 700-1000 m,

30.vii.1966, Ent. Exc. Zool. Mus. Amsterdam, 2 d, 4

(Babuna R.), 400m, 1 4,1 2; Izvor, 800m, 12.

Greece: Makedhonia: Mt. Kholomón, 700—1000

m, 18 d, 11 2; Palaiókastro, 600 m, 5 6, 3 2; Lak-

hands, 200 m, 4 6, 9 2; Litékhoron, 20 m, 2 d; Thes-

salia: 20 km NW of Elassén, 10 6, 10 ®; Kallithéa,

600 m, 8 6, 4 & (all CW); 5 km S of Elassón, 600 m,

5.viii.1975, J. Duffels, 1 3 (ITZ).

Diagnosis. — See the descriptions in Ramme

(1926—1933), Bei-Bienko (1954) and Harz

(1969).

Variation. — The specimens at hand are

slightly variable. The male cercus varies as in

figs. 50 —52. The basal fold of the lower ovipo-

sitor valve (figs. 227, 228) is uniform, being

thin, lamelliform, well protruding horizontally,

it is strongly arched upward anteriorly, with a

shell-shaped impression, forming with the go-

nangulum a deep ovoid pit which, as seen in

lateral view, is partly covered by the recurved

outer margin of the fold.

Distribution. — The range extends from SW

Bulgaria (?) through S Yugoslav Makedonija to

the northern and central parts of Greek Maked-

honia, and southward into the extreme north-

eastern part of Thessalia.

Greek localities. — Makedhonia: Sérrai: Strimo-

niké (“Strimonikon”) (Weidner, 1951); Khalkidhiki:

Mt. Khoomón, 700—1000 m; Palaiókastro, 600 m;

Thessaloniki: Lakhanás, 200 m; Thessaloniki (“Salo-

niki”) (Ramme, 1951); 6 km N of Thessaloniki

(“Lembet”) (Uvarov, 1923; Ramme, 1933; Bei-Bien-

ko, 1954) & “Happy Valley” (Uvarov, 1923); Ak.

Még. Karambournoú (“Karaburun”) (Uvarov, 1923);

Lankadhas (“Langaza plain”) (Uvarov, 1923); Kilkís:

about 20 km S of Kilkis (“Deve Kran”) (Uvarov,

1923); Pélla: Edhessa (“Vodena”) (Ramme, 1926);

Florina: Florina (Berland & Chopard, 1922; Ramme,

1926); Piéria: Litokhoron (Harz, 1969), 20 m.

Thessalia: Larisa: 20 km N of Elasson; 5 km S of

Elassón, 600 m; Kallithéa, 600 m. (Map 3).

Remarks. — A well defined species. Basal

fold of lower ovipositor valve recalling P. brun-

neri, but the pit in the latter slightly smaller,

more circular and more widely open laterally.

Ramme’s record (1951) of P. fussi from Thessa-

loniki is far beyond the known range of that

species and more probably refers to P. macedo-

nicus.

P. macedonicus occurs in the lowlands and

was found on diverse low shrubs and herba-

ceous plants, in forest clearings as well as open

country.

Poecilimon brunneri (Frivaldsky, 1867)

(figs. 53, 54, 229, 230, map 3)

Odontura brunneri Frivaldsky, 1867: 94, pl. 2 figs. 3,

3a—3d (type-localities: Fehertemplon (= Bela

Crkva), NE Yugoslavia and Mehadia, SW Roma-

nia).

Poecilimon brunneri; Berland & Chopard, 1922: 167

Ramme, 1933: 527, pl. 8 fig. 8, pl. 11 fi. 4. Werner,

1933a: 405; 1934: 324, fig. 3; 1937a: 108; 1938:

167. Bei-Bienko, 1954: 305, fig. 178. Kaltenbach,

1965: 469. Harz, 1969: 134, figs. 282, 401— 405.

Poecilimon berlandi Uvarov, 1923: 148, fig. 1 (type-

locality: Vakoufkeuy, NE of Florina), Ramme,

1926: 281.

Poecilimon lemnoticus Werner, 1932: 3 (type-locality:

Lemnos).

Material studied. — Yugoslavia: Makedonija: Izvor

(Babuna R.), 2 km N & 8 km E & 3 km W, 400 m, 5

36,21 2,7 km NE of Titov Veles, 250 m, 2 @; P. Plet-

var near Prilep, 900 m, 9 6, 12 2 (all CW); 20 km W.

of Skoplje, 30.vi.1975, J. Duffels (ITZ).

Greece: Dhidhimotikón, 40 m, 1 2 (CW).

Diagnosis. — See the descriptions in Uvarov

(1923, as P. berlandi), Ramme (1933), Bei-Bien-

ko (1954) and Harz (1969).

Variation. — The specimens before me cover

only a small part of the known range and show

but little variation. The male cercus varies as in

figs. 53, 54. Basal fold of lower ovipositor valve

(figs. 229, 230) much as in P. macedonicus but

differing consistently in the groove formed be-

tween the fold and the gonangulum, which is

smaller, round instead of elongate, and more

widely open laterally.

Distribution. — Widely distributed from the

southern Ukraina through Romania, Bulgaria

and European Turkey to Yugoslavia, Albania

and N Greece, including some northern Aegean

islands.

Greek localities. — Thraki: Evros: Dhidhimotikén,

40 m; Kavalla: Krinidhes (Kaltenbach, 1965).

Makedhonia: Florina: Vakoufkeuy, NE of Florina

(Berland & Chopard, 1922; Uvarov, 1923; Ramme,

1926).

eee Is.: Thásos (Werner, 1938); Samothrakı

(Werner, 1934).

Eastern Aegean Is.: Limnos (“Lemnos”) (Werner,

1932; Ramme, 1933; Werner, 1933a, 1937a). (Map 3.)

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

58 59 60 61

Figs. 55—74. Poecilimon species, dorsal view of left male cercus. 55—57, P. cretensis Werner; 55, 56, Mt. Idhi;

57, Mt. Levka; 58, 59, P. ikariensis sp. n.; 58, holotype; 59, paratype; 60—62, P. ebneri Ramme (Mt. Smélikas);

63, P. klısuriensis sp. n. (holotype, 63a tip from behind); 64, 65, P. zwicki Ramme; 64, Amorion; 65, Bulgaria;

66—68, P. orbelicus Pancic; 66, Bulgaria; 67, 68, Mt. Pangaion; 69, 70, P. miramae Ramme (Elasokhórion);

71-74, P. anatolicus Ramme; 71, 72, Kesan; 73, 74, Mt. Pangaion.

Poecilimon pergamicus

Brunner von Wattenwyl, 1891

Poecilimon pergamicus Brunner von Wattenwyl,

1891: 29 (type-locality: Pergamon, Asia Minor).

Ramme, 1933: 528, pl. 6 fig. 16, pl. 7 fig. 6, pl. 8

fig. 19, pl. 11 fig. 15. Beî-Bienko, 1954: 312. Harz,

1969: 129, figs. 289, 382— 385.

? Poecilimon pergamicus; Werner, 1933a: 402.

Diagnosis. — See the descriptions in Ramme

(1933) and Harz (1969).

Distribution. — Known only from the type-

locality, Pergamon, western Turkey.

Remarks. — Werner (1933a) recorded three

females from the eastern Aegean island of

Lésvos (“Mytilene”) but was not certain of his

identification.

WILLEMSE: Greek Poecilimon 171

Poecilimon cretensis Werner, 1903

(figs. 55—57, 173, 174, 231, 232, map 3)

Poecilimon jonicus cretensis Werner, 1903: 67, fig.

(type-locality: Rethymno, Kreta).

Poecilimon cretensis; Ramme, 1927: 186; 1933: 538,

pe 55 ple lt tig. 27, pl. 12 fig. 24. Harz,

1969: 137, figs. 292, 293, 323, 418, 419. Willemse

& Kruseman, 1967: 126.

Poecilimon distinguendus Kuthy, 1907: 554 (type-lo-

calities: Antrum Jovis, Mons Ida & Insula Dhia,

Creta).

Material studied. — Kríti: Mt. Idhi, between Kolita

and Psilorítis, 1700—2100 m, 93 6, 43 2; Mt. Léfka,

Linoséli above Xiloskaló, 1800 m, 2 d; Frángo

Kástello, 5.v.1973; W. Gravestein, 1 &; Phaistós,

23-26. V.1972, M. C. & G. Kruseman, 4 6;

Knossos, 17—21.v.1972, M. C. & G. Kruseman, 1 d

(all CW).

Diagnosis. — See the descriptions in Werner

(1903), Ramme (1927, 1933) and Harz (1969).

Basal fold of lower ovipositor valve (figs. 231,

232) moderately thick, strongly arched upward

anteriorly, protruding downward and forming

with the gonangulum a shallow concavity.

Variation. — Variation of the subgenital plate

and cercus of male as in figs. 55—57, 173, 174.

The coloration varies from completely green to

strongly black dorsally.

Distribution. — Known from the islands of

Kríu and Náxos.

Localities. — Kikladhes: Náxos (Ramme, 1927).

Kriti: a full list of localities can be found in Wil-

lemse & Kruseman (1976). (Map 3.)

Poecilimon ikariensis sp. n.

(figs. 58, 59, 156, 157, 175, 1976, 193—195, 233,

234, map 3)

Poecilimon hamatus; Werner, 1934: 323, fig. 2c (only

Ikaria).

2 paratypes, labelled: Hellas, Ikaria, Pass between

Aghios Kirykos and Karavostamon, 20.vi.1977, M. C.

& G. Kruseman (ITZ).

Diagnosis. — Male (figs. 193, 194). Size

small. Integument glossy. Fastigium of vertex

narrower than half width of scape, extending

anteriorly, not sloping with frons, lateral mar-

gins slightly convergent anteriorly towards

shallowly grooved apex. Pronotum (figs. 156,

157) short, not saddle-shaped, not widening

posteriorly, not or scarcely raised but bulbously

inflated posteriorly; fore and hind margins al-

most straight, lower margin of lateral lobe S-

shaped. Elytra short, extending just beyond

hind margin of pronotum apical margin not

reaching beyond halfway first tergite. Abdomi-

nal tergites without particulars. Cercus (figs. 58,

59) long, extending far beyond hind margin of

subgenital plate, slender, apical half cylindrical,

apical third strongly incurved, apex obtusely

rounded with superimposed small dorsal crest

which is somewhat serrate with a single or few

small denticles. Subgenital plate (figs. 175, 1976)

short, wide, apical part Hie hind margin

transverse with lateral edges triangularly ex-

tending posteriorly. Femora comparatively

short and robust, unarined.

General colour yellowish white, heavily var-

iegated with black. Occiput black. Pronotum

black with lower and fore margins and a pair of

dorsal spots behind transverse sulcus yellowish

white. Elytra black, anterior margins broadly

bordered with ivory white. Abdomen with a

median and, on either side, a lateral band com-

posed of black spots which do not reach com-

pletely hind margin of tergites. Last tergites,

supra-anal plate, cerci and hind margin of sub-

genital plate completely black or almost so.

Legs more or less variegated with black and

dark brown, hind femur often with series of

transverse black stripes.

Female (fig. 195). Integument shiny. Prono-

tum short, metazona scarcely bulbously inflated

posteriorly. Elytra reaching hind margin of pro-

notum or slightly longer, touching or overlap-

ping each other dorsally. Ovipositor short. Bas-

al fold of lower ovipositor valve (figs. 223, 224)

large, strong, well protruding horizontally,

slightly arched upward anteriorly, outer margin

sinuate dividing the fold into a pair of rounded

lobes, the anterior one slightly bent upward and,

forming with the gonangulum a large concavity.

Coloration more uniformly greenish except

elytra, which are black, the fore margins broad-

ly bordered with ivory-white.

Measurements: body d 15.0—17.0, 2 14.5—

13.0, 2 14.5—14.6; ovipositor 7.0—8.0.

Variation. — Shape of pronotum (figs. 156,

157), subgenital plate (figs. 175, 1976) and cer-

cus (figs. 58, 59) of the few males at hand but

slightly variable.

Distribution. — Known only from the type-

series from the Eastern Aegean island of Ikaría.

(Map 3.)

Remarks. — Readily recognizable by the

172 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 75— 93. P. chopardi Ramme, dorsal view of left male cercus. 75, Mt. Kaimakchalan; 76, Kozani; 77, Nea-

polis; 78, Mt. Smólikas; 79, Mt. Mavrovoúni; 80—82, Khrisomilea; 83, 84, Mt. Karava; 85—88, Mt. Tim-

fristós; 89, 90, 15 km W of Karpenision; 91, Timfristos village; 92, Ipatı; 93, Paliokhòri.

shape of the male cercus and the basal fold of

the ovipositor. It is the only member of the ge-

nus known from Ikaria.

Poecilimon syriacus Brunner von Wattenwyl,

1891

(figs. 235, 336)

Poecilimon syriacus Brunner von Wattenwyl, 1891: 29

(type-localities: Syria (Beirut, Hierosolyma) &

Bitlis in Asia minore). Werner, 1901: 288.

Ramme, 1933: 551, pl. 6 fig. 40, pl. 7 fig. 18, pl. 10

fig. 50, pl. 11 fig. 40. Beî-Bienko, 1954: 314. Harz,

1969: 146, figs. 303, 338, 452—455.

Material studied. — Samos, between Marathókam-

bos and Ayios Theödhori, 11.v1.1977, M. C. & G.

Kruseman, 1 @ (ITZ).

Diagnosis. — See the descriptions in Ramme

(1933), Bei-Bienko (1954) and Harz (1969).

Distribution. — A wide-ranging species,

reaching from Israel and the Lebanon to E and S

Turkey. Samos?

Remarks. — Recorded by Werner (1901)

from the eastern Aegean island of Samos, but

this record has never been confirmed. The fe-

male before me agrees with the description of P.

syriacus, but its identification remains uncertain

WILLEMSE: Greek Poecilimon 173

94 95 96 97 98

99 100 101 102 103

104 105 106 107 108

109

110 111 112 113

Figs. 94—113. P. zimmeri Ramme, dorsal view of left male cercus. 94, 95, Mt. Timti; 96, Mt. Tómaros; 97, Mt.

Tzoumerka; 98, 99, Mt. Panaitolikón; 100, 101, Mt. Vardhousia; 102, 103, Mt. Oíu; 104, Mt. Giona; 105, 106,

Mt. Parnassés: 107, Mt. Aroánia; 108, 109, Mt. Mainalon; 110, 111, Mt. Erimanthos; 112, 113, Mt. Panak-

haïkén.

174 TIJpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL.

by lack of an associated male. The basal fold of

the lower ovipositor valve in the female is illus-

trated (figs. 235, 236).

Poecilimon ebneri Ramme, 1933

(figs. 60—62, 177, 237, 238, map 3)

Poecilimon ebneri Ramme, 1933: 542, pl. 6 fig. 30, pl.

9 fig. 38, pl. 11 fig. 32 (type-locality: Pashtrik, Al-

banien). Harz, 1969: 141, figs. 298, 432—435.

Willemse, 1977: 53.

Poecilimon ebneri peristericus Karaman, 1961: 41,

figs. 10—14. Syn. nov.

Material studied. — Yugoslavia: Mt. Pelister, west-

ern slopes, 31.vii.1965, Ent. Exc. Zoöl. Mus. Amster-

dam, 1 6 (ITZ).

Greece: Kozani, Mt. Vérmion, 1850 m, 8.v111.1973,

na, 1600—1700 m, 3 d, 6 2; Mt. Bela Voda, 1700—

1971 m, 20 d, 14 ®; Mt. Smólikas above A. Paras-

Diagnosis. — See the descriptions in Ramme

(1933), Karaman (1961) and Harz (1969).

Variation. — Body size rather variable. The

male cercus varies as in figs. 60—62. Apical part

of male subgenital plate invariably long (fig.

177). Basal fold of lower ovipositor valve small

and strongly compressed downward, scarcely

forming any concavity with the gonangulum

(figs. 237, 238). General colour from green to

yellowish. Most specimens are uniformly col-

oured, some are slightly varicoloured.

Distribution. — Known from Albania (type-

locality), Mt. Perister in extreme SW Yugoslav

Makedonija, and from some mountains in NW

Greece.

Greek localities. — Makedhonia: Kozáni: Mt. Vér-

mion, 1850 m; Flérina: Mt. Bela Voda, 1700—1971

m; 4 km E of Pisodhéri, 1600— 1700 m (Willemse,

1977).

Ipiros: Ioannina: Mt. Smélikas above A. Paraskevi,

2000— 2300 m (Willemse, 1977). (Map 3.)

Remarks. — The distinctive features of P. eb-

neri peristericus correspond with the variability

shown by Greek material of P. ebneri. I propose

to synonymize both taxa.

The species lives on diverse plants covering

stony ground above the timberline. In Greece it

may occur together with P. ornatus, P. gracilis,

P. jonicus jonicus and P. chopardi.

Poecilimon klisuriensis sp. n.

(figs. 63, 63a, 158, 178, 196—198, 239—240,

map 3)

6 (1982)

paratypes, labelled: Hellas, N. Kastorıas, Klısoura,

1150 m, 31.vii.1976, F. & L. Willemse & J. Tilmans

(CW).

Diagnosis. — Male (figs. 196—197). Size

moderate. Integument slightly shiny. Fastigium

of vertex about half as wide as greatest width of

scape. Pronotum scarcely saddle-shaped, scar-

cely widening posteriorly, metazona slightly

raised, hind margin straight, lower margin of

lateral lobe anteriorly and widely

rounded posteriorly (fig. 158). Elytra well visi-

ble, apical margin radiati hind margin of first

tergite. Cercus (figs. 63, 63a) robust, base wide,

strongly conical, apical fourth obtuse-angularly

incurved and slender, inner and outer margins

crested toward apex and terminating into a

slightly downcurved acute tip. Subgenital plate

(fig. 178) slender, apical part quite elongate,

hind margin straight, ventral side apically with

weak median keel.

General colour yellowish green, markedly

spotted with dark brown. Pronotal metazona

with reddish brown median line and, on either

side, with a reddish brown streak. Stridulatory

area of elytron partly dark brown. Tip of cercus

black.

Female (fig. 198). Pronotum cylindrical. Ely-

tra small, lateral, widely separated from each

other, completely covered a pronotum. Basal

fold of lower ovipositor valve (figs. 239, 240)

robust, swollen, posterior part lobe-shaped and

extending strongly downward, forming with

the gonangulum a narrow small pit, which

opens laterally. Base of lower ovipositor valve

with lower margin thick and lateral side

wrinkled. Coloration as in male.

Measurements: body d 18.0, ® 15.2—17.5;

pronotum d 5.1, 2 5.0—5.5; elytron d 1.5, ?

0.0—0.0; hind femur & 12.9, 2 15.8—17.2; ovi-

positor 8.9—9.1

Distribution. — The species is known only

from the type-locality in NW Greek Makedho-

nia at the highest point of a pass between the

villages of Korisós and Lékhovo, at the border

between the districts of Kastoria and Flórina.

(Map 3.)

Remarks. — The species comes near P. ebne-

ri, from which it differs in the male cercus

which is more robust with the tip not bidentate;

the male subgenital plate is longer and the lobes

of the basal fold of the lower ovipositor valve

are also different. It differs from P. orbelicus in

the same features and the pronotum, which is

WILLEMSE: Greek Poecilimon 175

WERE

114

120

126

125

127°

Figs. 114-127. Poecilimon species, dorsal view of left male cercus. 114121, P. thessalicus Brunner von Wat-

tenwyl; 114, Mt. Pilion; 115, 116, Mt. Ossa; 117, Leptokaria-Karia; 118, 119, Mt. Olimbos; 120, Fteri; 121, Mt.

Piéria; 122—124, P. propinquus Brunner von Wattenwyl; 122, 123, Mt. Dírfis, 12

2a tip from behind; 124, Akr.

Soúnion; 125, 126, P. sanctipauli Brunner von Wattenwyl; 125, Ephesus, 125a tip from behind; 126, Lindhos;

127, P. hamatus Brunner von Wattenwyl; Lindhos, 127a, up from behind.

shorter and less raised in the latter. There ıs also

some resemblance with P. zwicki. In the latter

species, however, the male cercus is more slen-

der and its tip differently shaped, the pronotum

is narrower, the male subgenital plate shorter,

the female elytra are not placed laterally but

overlap and the basal fold of the ovipositor is

quite different.

The few specimens were found on the flowers

of herbaceous plants along the road.

Poecilimon zwicki Ramme, 1939

(figs. 64, 65, 179, 180, 241, 242, map 3)

Poecilimon zwicki Ramme, 1939: 47, fig. 2 (type-lo-

cality: Bansko, Piringebirge, Bulgarien). Kalten-

bach, 1965: 469. Harz, 1969: 134, figs. 327, 398—

400.

Material studied. — Bulgaria: Goce Deltev,

Greece: Amörion, 40 m, 1 d; Stavroupolis, 100 m,

176 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

128 129 130

131 132 133

136

134 135

137 138

Figs. 128—138. Poecilimon species, lateral view of male pronotum. 128—130, P. hoelzeli Harz; 128, Mt. Piérias

129, Mt. Olimbos; 130, Khrisomiléa; 131—133, P. pindos sp. n.; 131, 132, Mt. Timfi; 133, Mt. Mavrovotni;

134, 135, P. gracilis (Fieber), Mt. Bela Voda; 136—138, P. obesus Brunner von Wattenwyl; 136, Kalavrita; 137,

Kapsia; 138, Khrisovitsi.

1 2; Mt. Falakrón above Vólax, 1250 m, 1 6; Mt.

Vrondoús, 800 m,9 8,7 9 (all CW).

Diagnosis. — See the descriptions in Ramme

(1939) and Harz (1969).

Variation. — Male subgenital plate and cercus

variable as shown in figs. 64, 65, 179, 180. Basal

fold of lower ovipositor valve thin, lamelliform,

small, moderately protruding horizontally,

scarcely arched upward anteriorly, scarcely im-

pressed trom above, forming with the gonangu-

lum an elongate weak impression (figs. 241,

242).

Distribution. — The known range covers S

Bulgaria, E Greek Makedhonia and Greek

Thraki.

Greek localities. — Thraki: Evros: Amórion, 40 m;

Xanthi: Stavroúpolis, 100 m.

Makedhonía: Kavalla: Kalamika; Batis; Krinidhes;

WILLEMSE: Greek Poecilimon 177

Dhiastávrosis; Avramilia; Drama: Drama (all Kalten-

bach, 1965); Mt. Falakrón, 1250 m; Sérrai: Mt. Vron-

dots, 800 m. (Map 3.)

Remarks. — The Greek specimens were

found on ferns and diverse herbaceous plants of

woodland and open land. In Greece the species

may occur together with P. orbelicus and P. tho-

racicus.

Poecilimon orbelicus Panéic, 1883

(figs. 66—68, 243, 244, map 3)

Poecilimon orbelicus Panäic, 1883: 163 (type-locality:

Mt. Rilo, Bulgaria). Ramme, 1933: 547, pl. 6 tig.

sep Mio, pls 9 tig. 45, pl. 11 fie. 37, plal2

fig. 28. Bei-Bienko, 1954: 325, fig. 188. Harz,

1969: 146, figs. 317, 335, 456 —458.

Poecilimon bulgaricus Brunner von Wattenwyl, 1891:

30 (type-locality: Mt. Rilo, Bulgaria).

Material studied. — Bulgaria: Pirin Mts., (in Rus-

Greece: Mt. Vrondous, 1400 m, 7 4, 3 2; Mt. Fa-

Pangaion above Akrovoúnion, 1250—1900 m, 22 d,

28 © (all CW).

Diagnosis. — See the descriptions in Ramme

(1933), Beî-Bienko (1954) and Harz (1969).

Variation. — Male cercus as shown in text-

figs. 66—68.

Distribution. — Known from SW Bulgaria

and adjacent parts of Greek Makedhonia.

Greek localities. — Makedhonia: Thessaloniki

(“Saloniki”) (Bei-Bienko, 1954); Sérrai: Mt. Vron-

dots, 1400 m; Drama: Mt. Falakrén, 1250—1900 m;

Kavalla: Mt. Pangaíon, 1250—1900 m. (Map 3.)

Remarks. — The basal fold of the lower ovi-

positor valve (figs. 243, 244) resembles that of

P. ebneri but is more compressed downward

posteriorly, and the concavity between the fold

and the gonangulum is deeper. Bei-Bienko’s re-

cord (1954) is based on a single male labelled

“Saloniki”. He supposed that the precise locali-

ty is further north. The species may occur in

Greece together with P. pancici, P. anatolicus

and P. zwicki. We found it on the higher parts

of the mountains up to above the timberline.

Poecilimon miramae Ramme, 1933

(figs. 69, 70, 181, map 3)

Poecilimon miramae Ramme, 1933: 563, pl. 6 fig. 46,

pl. 10 fig. 58, (type-locality: Kilia, European Tur-

key). Bei-Bienko, 1954: 339. Harz, 1969: 159,

figs. 305, 336, 510—513.

Material studied. — Greece: Evros, Elasokhórion,

180 m, 3 d (CW).

Diagnosis. — See the descriptions in Ramme

(1933), Beî-Bienko (1954) and Harz (1969).

Variation. — Insufficiently known by lack of

material. Male subgenital plate and cercus as in

text-figs. 69, 70, 181.

Distribution. — The range covers NW Tur-

key, SE Bulgaria and E Greek Thráki. The oc-

currence in Greece was not yet known.

Evros: Elasokhérion,

Greek locality. — Thraki:

180 m. (Map 3.)

Remarks. — The Greek specimens were

found sparsely on Rubus, at the riverside of the

Erithropos.

Poecilimon anatolicus Ramme, 1933

(figs. 71—74, 245, 246, map 4)

Barbitistes flavescens; Fieber, 1853: 175.

Poecilimon flavescens; Brunner von Wattenwyl, 1878:

42; 1882: 266.

Poealimon anatolicus Ramme, 1933: 555, pl. 6 fig. 42,

pl. 7 fig. 20, pl. 10 fig. 53, pl. 11 fig. 43, pl. 12 fig.

33 (type-locality: Bursa, Anatolia). Bei-Bienko,

1954: 333.

Eupoecilimon anatolicus; Ramme, 1951: 336, figs.

93a—c, 94.

Material studied. — Turkey: Kesan, 60 m,

15.vii.1979, L. Willemse, 11 d, 10 2; Greece: Mt.

Pangaion above Akrovoúnion, 1250 m, 16 d, 6 9 (all

CW).

Diagnosis. — See the descriptions in Ramme

(1933, 1951) and Beî-Bienko (1954). Basal fold

of lower ovipositor valve (figs. 245, 246) strong,

swollen, its posterior part lobe-shaped and well

protruding horizontally, its anterior part

strongly compressed downward and arched up-

ward, longitudinally impressed above, forming

an elongate, dark brown concavity with the go-

nangulum, which is directed anteriorly.

Variation. — The male cercus is known to be

rather variable (figs. 71— 74).

Distribution. — Hitherto known only from

the Bursa area of extreme NW Anatolia. Its oc-

currence in the European part of Turkey and E

Greek Makedhonía was unknown and extends

the range considerably westward.

Greek locality. — Makedhonia: Kaválla: Mt. Pang-

aion, 1250 m. (Map 4.)

Remarks. — Both sexes are characterized by

178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

139 140

143 144

147

155

141

142

145 L ) 146

Figs. 139—157. Poecilimon species, lateral view of male pronotum. 139—144, P. jonicus jonicus (Fieber); 139,

Petalia; 140, 141, Perithia; 142, 143, Aristi-Papingon; 144, Trepjca-Otsesevo; 145—150, P. jonicus lobulatus ssp.

n.; 145, 146, Métsovon; 147, 148, Khrisomilea; 149, Kaléntzion; 150, Niköpolis; 151, 152, P. werneri Ramme

(Lekhaina); 153—155, P. tessellatus (Fischer); 153, Attiki; 154, Mt. Mainalon; 155, Mt. Panakhaikon; 156, 157,

P. ikarıensis sp. n.; 156, holotype; 157, paratype.

colour, shape of male cercus and of basal fold ot

the lower ovipositor valve. The shape of the lat-

ter is similar to that of P. chopardi and P. thes-

salicus.

Hitherto unknown from Europe. The Greek

specimens were found in a forest clearing on

ferns and other herbaceous plants, together with

P. orbelicus and P. pancici.

Poecilimon chopardi Ramme, 1933

(figs. 75— 93, 159—162, 247—252, map 4)

Poecilimon flavescens; Berland & Chopard, 1922: 167.

Poecilimon chopardi Ramme, 1933: 548, pl. 6 fig. 38,

pl. 9 fig. 47 (type-locality: S. de Monastir (“Bito-

la”) entre Bukova et Holéven). Beî-Bienko, 1954:

329. Karaman, 1962: 1 pag. (®). Harz, 1969: 145,

figs. 450—451.

Poecilimon veluchianus Ramme, 1933: 549, pl. 6 fig.

39, pl. 10 fig. 48, pl. 11 fig. 38, pl. 12 fig. 30 (type-

locality: Veluchi gebirge, 1800—2000 m). Werner,

1933b: 190 (as veluchiensis). Harz, 1969: 150, figs.

318, 468—473. Syn. nov.

Material studied. — Yugoslavia: Makedonija, Kaj-

Greece: 2 km E of Alona, 1000 m, 2 2; 12 km W of

Florina, 1000 m, 1 6; 6 km SW of Dhrosopiyi, 1200

m, 13 d, 4 ®; Polipótamon, 1100 m, 2 6, 1 ® (all

CW); 18 km SE of Kozani, 3.vii.1975, J. P. Duffels, 2

3,8 2 (ITZ); Velvendos, 250 m, 1 2; Dhrepano, 700

m, 2 ®; Neápolis, 800 m, 1 ®; Eptakhori, 1190 m, 1

WILLEMSE: Greek Poecilimon 179

158 159

162 163

166 167

164

160 161

165

168 169

Figs. 158—169. Poecilimon species, lateral view of male pronotum. 158, P. klisuriensis sp. n. (holotype); 159—

162, P. chopardi Ramme; 159—160, Paraskevi; 161, 162, Mt. Timfristos; 163—166, P. zimmer: Ramme; 163,

164, Mt. Parnassös; 165, 166, Mt. Mainalon; 167—169, P. thessalicus Brunner von Wattenwyl; 167, Mt. Ossa;

168, Mt. Pilion; 169, Mt. Olimbos.

6,1 2; Grevena, 10 km SW of Dheskäti, 700 m, 19

6, 18 2; Paraskevi near Dheskati, 600 m, 33 d, 27 9;

Kranıa Dheskatis, 600 m, 1 6; Trigon (Trikkala), 800

m, 1 dg; Kalabaka, Metéora, 14.v1.1979, J. Smid & F.

Smid-Elbers, 1 36; Khrisomiléa, 900—1200 m, 38 dg,

2; Mt. Karáva, 2000 m, 5.viii.1973, La Greca, 6 d, 5

800—2000 m, 47 d, 13 2; Métsovon, 3 km N & 3 km

CW); Métsovon, 1500 m, 1.v111.1973, La Greca, 1 d,

1 2 (IBA); Mt. Mavrovotni near Métsovon, 1800—

2100 m, 24 d, 10 2; Mt. Timfristós above Karpen-

luchianus) (both CW); 15 km W ot Karpenision, 1100

m, 8.vu.1975, J. Duffels, 26 3, 26 2 (ITZ); village of

Timfristós, 1100 m, 4 6, 1 2 (CW); Ipati- Neokh-

Orion, 800 m, 7.vii.1975, J. Duffels, 1 6 (ITZ); Pal-

aiokhori, 400 m, 3 d, 4 2 (CW).

Diagnosis. — Male. Size moderate. Integu-

ment uniformly dull. Width of fastigium of ver-

tex from slightly less to distinctly more than

half the greatest width of scape. Pronotum (figs.

159—162) slightly saddle-shaped, a little widen-

ing posteriorly, metazona more or less raised

and scarcely inflated, hind margin straight or

weakly incised, lower margin of lateral lobe

straight or almost so anteriorly and widely con-

vex posteriorly. Elytra well visible, apical mar-

gin reaching from almost hind margin of first to

beyond middle of second tergite. Cercus (figs.

75—93) from slender to robust, on proximal

half slightly conical, in distal half about cylin-

drical and gradually incurved; apex not

widened, slightly compressed dorso-ventrally,

sometimes even flattened above, terminating in-

to an apical tooth which stands between mid-

dorsal and the side, inner and outer margins

more or less serrate or provided with two to six

small teeth over a variable distance. Subgenital

plate short, lateral margins straight or sinuate,

converging posteriorly, hind margin narrow,

straight or slightly concave or with V-shaped

emargination, postero-lateral edges whether or

not produced.

General colour yellowish green, straw-yellow

or brown, usually distinctly spotted with dark

brown. Antennae finely annulated. Pronotal

dorsum on either side with a yellowish or

creamy-white streak which, in the metazona, is

bordered medially with rusty or reddish brown.

Elytra yellowish brown, with or without a pre-

apical dark spot, stridulatory area sometimes

dark brown. Abdomen of general colour; dor-

sum often paler brown or yellowish and

sometimes provided with a median dark brown

or blackish band, and on either side with an ill-

180

177

di di

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

173 174 175

179 180 181

Figs. 170—181. Poecilimon species, ventral view of male subgenital plate. 170—172, P. jonicus jonicus (Fieber);

170, Dhassia; 171, 172, Gazátika; 173, 174, P. cretensis Werner (Mt. Ídhi); 175, 176, P. ikariensis sp. n. (para-

types); 177, P. aba Ramme (Mt. Smólikas); 178, P. klisuriensis sp. n. (holotype); 179, 180, P. zwickı Ramme;

179, Amorion; 180, Bulgaria; 181, P. miramae Ramme (Elasokhorion). Figs. 182, 183. Poecilimon nobilis Brun-

ner von Wattenwyl, hind margin of last abdominal tergite of male; 182, Mt. Maînalon; 183, Mt. Taiyetos.

defined lateral band of similar colour; median

band composed of a pair of spots along fore

margin of tergites, extending and narrowing

posteriorly, usually without reaching hind mar-

gin of tergites and separated at middle by a yel-

lowish line. Cercus with tip more or less black.

Female. — Pronotum cylindrical, metazona

sometimes scarcely widened and raised posteri-

orly. Elytra completely covered by pronotum,

placed laterad and widely separated from each

other. Basal fold of lower ovipositor valve (figs.

247-252) swollen, strong, protruding down-

ward, arched upward anteriorly, longitudinally

pressa and forming with gonangulum an

elongate concavity directed antero-ventrally.

Coloration as in male, though usually more uni-

form.

16.0—18.8; pronotum d 3.8—5.2, 9 4.3—6.8,

13.1—16.4, ©).

Variation. — The species is highly variable.

The male cercus shows some geographical vari-

ation. Throughout the northern part of the

range (figs. 75—79) the apical tooth of the cer-

cus is almost invariably located in the middle of

the width of the apex, and both the length of the

inner and outer apical margins and their number

of teeth are about equal to each other. Through-

out the southern part of the range (figs. 80—93),

however, the apical tooth of the cercus is situ-

ated more laterally and at the same time the

length and the number of teeth of the inner api-

cal margin are increased while those of the outer

margin are decreased. The basal fold of the low-

er ovipositor valve is but slightly variable (figs.

VIII)

Distribution. — The range extends from the

extreme SW part of Yugoslav Makedonija

through W Greek Makedhonia and N Thessalia

to the Pindhos range, covering the eastern and

central mountains from the Albanian border as

far south as Karpenísion, and reaching eastward

the lowlands and hills at the northern sides of

WILLEMSE: Greek Poecilimon 181

Mt. Oîti and Mt. Parnassós in central Greece.

Greek localities. — Makedhonía: Kozäni: 18 km SE

of Kozäni; Velvendos; Dhrépano, 700 m; Neàpolis,

800 m; Flórina: Flórina (Berland & Chopard, 1922); 2

km E of Álona, 1000 m; 12 km W of Flórina, 1000 m;

6 km SW of Dhrosopiyi, 1200 m; Polipétamon, 1100

m; Kastoria: Eptakhóri, 1190 m; Grevená: 10 km SW

of Dheskáti, 700 m; Paraskeví near Dheskati, 600 m.

Thessalia: Larissa: Krania Dheskätis, 600 m;

Trikkala: Kalabáka; Trigón, 800 m; Khrisomiléa,

900-1200 m; Pertoúli; Mt. Karáva, 2000 m.

[piros: Ioannina: Mt. Smólikas above A. Paraskevi,

800-2000 m; Métsovon, 1500 m; 3 km N & 3 km E

& 5—10 km N of Métsovon, 1300—1600 m; Mt.

Mavrovoúni near Metsovon, 1800— 2100 m.

Central Greece: Evritania: Mt. Timfristös (“Velu-

chi”) (Ramme, 1933; Werner, 1933b), above Karpen-

ision, 1700-2100 m; 15 km W of Karpenision, 1100

m; Fthiötis: Timfristös village, 1100 m; between fpati

and Neokhörion, 800 m; Bolas: Palaiokhóri, 400 m.

(Map 4.)

Remarks. — During examination of the many

specimens listed above it became apparent that

characters of P. chopardi and P. veluchianus

usually considered distinctive are completely

bridged by variation. Therefore, these taxa

should be considered conspecific. Both were de-

scribed simultaneously in the same work

(Ramme, 1933). The description of P. chopardi

is preceding (pag. 548) that of P. veluchianus

(pag. 549). As none of these names have any

special significance, I select P. chopardi and

propose to synonymize P. veluchianus with it.

Though highly variable, the species is well

defined by the dull integument, the gradual and

never angulate incurvation of the male cercus,

and the small elytra and shape of the basal fold

of the female ovipositor. The species resembles

much P. zimmeri and P. thessalicus. The distinc-

tion from the former is discussed below.

P. thessalicus differs from P. chopardi in the

broader pronotum which is less saddle-shaped

and more widened behind, and also in the inte-

gument, which is conspicuously glossy instead

of dull.

Hitherto the species was known only from

scanty material from Yugoslav Makedonija

(Berland & Chopard, 1922; Ramme, 1933; Ka-

raman, 1962), a male from Florina (Berland &

Chopard, 1922), and some material from Mt.

Timfristós (Ramme, 1933; Werner, 1933b).

The species was found on diverse herbaceous

plants, from the lowlands up to above the tim-

berline. In Greece it may occur together with

P. ornatus, P. hoelzeli, P. pindos, P. gracilis,

P. jonicus jonicus, P. jonicus lobulatus and P. eb-

neri.

Poecilimon zimmeri Ramme, 1933

(figs. 94—113, 163—166, 253—258, map 4)

Poecilimon propinquus; Brunner von Wattenwyl,

1878: 44 (only Parnass).

Poecilimon thessalicus; Brunner von Wattenwyl,

1891:30 (partim, only Parnassus).

Poecilimon zimmeri Ramme, 1933: 547, pl. 6 fig. 37,

pl. 9 fig. 46, pl. 12 fig. 29 (type-locality: Parnass,

M.-Griechenland). Bei-Bienko, 1954: 326, fig.

189. Harz, 1969: 147, figs. 311, 459—463.

Material studied. — Greece: Mt. Timfi above Pá-

pingon, 1800—2100 m, 64 d, 37 2; Mt. Tomaros

Tzoumérka above Katarraktis, 1700—2100 m, 7 d;

Mt. Panaitolikón above Prousós, Xerovoúni, 1400—

1650 m, 11 3, 14 2; Mt. Oiti above Ipati, near refuge

& Korifin & 10 km NW of Pavlianí, 1750—2135 m,

98 4, 68 2; Mt. Vardhoúsia above Mousounitsa,

1600—2000 m, 41 d, 29 9; Lilaia, 29.v.1976, A. Ma-

licky, 1 6 (all CW); Mt. Giona, 1900 m, 13.vin.1973,

La Greca, 2 6, 3 2; Mt. Parnassós, 12.vin.1973,

Arákhova & plateau N of Arákhova & below summit,

1200—1900 m, 35 d, 65 2 (topotypes); Mt. Pannak-

haikön above Zästova, 1000—1800 m, 28 d, 29 9;

Mt. Aroánia above Kalávrita, 1700—2000 m, 5 d, 3

2: Mt. Erimanthos above Kaléntzi, 1700— 2000 m, 8

d, 9 ®; Mt. Mainalon above Kardhara, 1550—1981

m, 52 6, 43 ® (all CW).

Diagnosis. — Differs from P. chopardi as fol-

lows.

Male. — Cercus with apical incurvation

stronger, more abrupt, almost rectangular.

(Apex ranging from similar to P. chopardi to of-

ten slightly widened (figs. 94—113).) Colour

pattern of abdomen often more distinct.

Female. — Basal fold of lower ovipositor

valve (figs. 253—258) strong, well protruding

horizontally, often lobe-shaped and not strong-

ly bent downward anteriorly, shallowly and

roundly impressed above and with gonangulum

forming a pit with a lateral opening.

Variation. — Throughout its range the incur-

vation of the male cercus is invariably strong

and angulate. The length and width of the apical

part and the armature of the tip, however, are

quite variable (figs. 94—113). Also the variation

of the male pronotum (figs. 163-166) and the

basal fold of the ovipositor (figs. 253—258) is

noteworthy.

Distribution. — The species has a remarkable

distribution: from Mt. Timfi, the northwestern

part of the Pindhos range, southward along the

182 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 184— 186. Poecilimon hoelzeli Harz; 184, 185, &, Mt. Olimbos, Refuge B; 186, 2, idem.

western mountains of this range, through the

mountains of central Greece, south of Karpen-

ision, it reaches eastward Mt. Parnassós and

southward covers the Pelopónnisos.

Localities. — Ipiros: Ioannina: Mt. Timfi above Pa-

pingon, 1800— 2100 m; Mt. Tomaros above Va-

riádhes, 1200—1600 m; Arta: Mt. Tzoumérka above

Katarraktis, 1700—2100 m.

Central Greece: Evritania: Mt. Panaitolikön above

Prousos, 1400—1650 m; Fókis: Mt. Oîti, 1750— 2135

m; Mt. Vardhousia above Mousounitsa, 1600—2000

WILLEMSE: Greek Poecilimon 183

REN

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

192

Figs. 190—192. Poecilimon jonicus lobulatus ssp. n.; 190, 191, é paratype, Khrisomiléa; 192, ?, Metsovon.

m; Mt. Giona; Lilaia; Voiotia: Mt. Parnassós (“Par-

nassus”, Brunner von Wattenwyl, 1891) (“Parnass”,

Brunner von Wattenwyl, 1878; Ramme, 1933; Harz,

1969), 1200—2100 m.

Pelopönnisos: Akhaïa: Mt. Panakhaikon above

Zástova, 1000—1800 m; Mt. Aroanıa above Kalávrita,

1700—2000 m; Mt. Erimanthos above Kaléntzi,

1700-2000 m; Arkadhia: Mt. Mainalon above

Kardhara, 1550—1981 m. (Map 4.)

Remarks. — P.zimmeri and P. chopardi

probably represent vicariant species. Transitio-

nal forms have not been found.

Hitherto known only from the type-locality.

The record from Istanbul, Turkey (Ramme,

1933, Harz, 1969) is quite improbable.

The habitat is similar to that of P. chopardı,

and it was found together with P. pindos, P. no-

bilis, P. jonicus lobulatus and P. tessellatus.

Poecilimon thessalicus Brunner von

Wattenwyl, 1891

(figs. 114—121, 167—169, 259— 262, map 4)

Poecilimon thessalicus Brunner von Wattenwyl, 1891:

30 (partim, not Parnassus) (type-locality: Montes

WILLEMSE: Greek Poecilimon 185

Ossa in Thessalia). Ramme, 1933: 550, pl. 7 tig.

17, pl 10 fe 45 go) AU Sir SE Werner, 19332:

402; 1938: 166. Harz, 1964: 450, figs. 20—21;

1969: 150, figs. 474— 478. Willemse, 1974: 352;

19700353

Poecilimon nitidus Werner, 1932: 2 (type-locality:

Skopelos, N. Sporaden). Harz, 1964: 449, figs.

18—19; 1969: 151, figs. 302, 304, 340, 479—480.

Syn. nov.

Material studied. — 5—8 km S of Sérvia, 600 m,

900—1500 m, 7 6, 3 2; Velvendés-Katafiyio, 950

14001700 m, 1 8; 10 km NW of Ftéri, 1500 m, 14

3, 10 2; Ftéri, 1000 m, 1 d; Mt. Olimbos, refuge A,

Mt. Olimbos, 1750—1950 m, 10.viii.1973, La Greca,

4 d, 3 2 (IBA); Leptokaria-Karia, 500—1000 m, 48

3, 29 2; Mt. Olimbos above Sparmös, near refuge B,

1800—2300 m, 56 d, 59 2; Kallithéa, 600 m, 1.9; 4

km W of Elassón, 300 m, 1 2; Dhimitra-Anatoli, 400

m, 1 9; Mt. Ossa above Anatoli, 1250—1400 m, 68

d, 26 2 (topotypes of P. thessalicus); Mt. Pilion

above Portariá, 700—1200 m, 2 & & between Portariä

Diagnosis. — See Ramme (1933).

Variation. — The species is highly variable.

Hind margin of pronotum varying from widely

rounded to emarginate. Shape of pronotum in

lateral view slightly variable (figs. 167—169).

Uniformly coloured specimens occur frequent-

ly. Black spots of male elytra may be absent. In-

tegument, also of female pronotum and abdo-

men, always markedly shiny. Apex of male cer-

cus comparatively uniform. The incurvation,

however, varies somewhat geographically, e.g.,

in specimens from the northern part of the

range it is more angular than in those from the

southern part (figs. 114—121). Basal fold of

lower ovipositor valve (figs. 259—262) strong,

protruding horizontally, arched upward ante-

riorly and invariably longitudinally impressed

above, forming an elongate groove which is di-

rected anteriorly. The posterior part of the fold

is variable and may be developed into a lobe of

variable size.

Distribution. — The range covers E Thessa-

lia, adjacent parts of Greek Makedhonia and the

Northern Sporádhes.

Localities. — Makedhonía: Kozáni: 5—8 km S of

Sérvia, 600 m (Willemse, 1977); Polimilos, 900—1500

m; between Velvendös and Katafiyio, 950—1400 m;

Mt. Pieria above Katafiyio, 1400—1700 (both Wil-

lemse, 1977); Pieria: Fteri, 1000 m; 10 km NW of

Fteri; Mt. Olimbos above Litókhoron, 1750—2400

m; between Leptokariá and Kariá, 500—1000 m (all

Willemse, 1977).

Thessalia: Larisa: Mt. Olimbos above Sparmós,

1800—2300 m (Willemse, 1974, 1977); Kallithéa, 600

m, 4 km W of Elassön, 300 m (both Willemse, 1977);

Lárisa (Ramme, 1933; Harz, 1969); stony hill “Kasa-

baliotiko” of Mt. Ossa area (Werner, 1933a); between

Dhimitra and Anatolí, 400 m (Willemse, 1977); Mt.

Ossa (Brunner von Wattenwyl, 1891; Ramme, 1933;

Harz, 1969) above Anatolí, 1250—1400 m (Willemse,

1977); Ludrissa (Ramme, 1933; Harz, 1969); Magni-

sia: Mt. Pílion above Portariá, 700—1200 m (Wil-

lemse, 1977) & between Portaria and Zagora, 900 m.

Northern Sporádhes: Skiathos (Werner, 1938);

Skópelos (Werner, 1932; Ramme, 1933; Harz, 1964,

1969). (Map 4.)

Remarks. — P.nitidus was synonymized

with P. thessalicus by ıts author (Werner, 1933a)

and by Ramme (1933), but re-established by

Harz (1964). The variation of P. thessalicus,

however, completely includes the characters of

P. nitidus, so that I propose to synonymize both

taxa again.

The species strongly resembles P. chopardı

and differs from the latter in the shiny integu-

ment and in the less raised metazona of the pro-

notum. The distinction between P. thessalicus

and P. zimmeri is based on the same features

and quite often on the male cercus and basal

fold of the ovipositor as well.

P. thessalicus lives from the lowlands up to

above the timberline and was found on a wide

range of plants, sometimes extremely abundant

on thistles or stinging nettles. It may occur to-

gether with P. ornatus, P. hoelzeli and P. mace-

donicus.

Poecilimon propinquus Brunner von

Wattenwyl, 1878

(figs. 122—124, 263—266, map 4)

Poecilimon propinguus Brunner von Wattenwyl,

1878: 44 (partım, not Syra & Parnass) (type-

locality: Athen); 1882: 267 (partım, not Syra).

Werner, 1902: 116; 1927: 428. Ramme, 1933: 553,

pl. 6 fig. 41, pl. 10 fig. 52, pl. 11 fig. 42, pl. 12 fig.

32. Werner, 1933a: 402; 1933b: 190; 1934: 323,

ng, 12 1997/09 1085 1937/58 1453 10395166, Ar

1969: 154, figs. 310, 341, 487—493.

Material studied. — Mt. Dírfis above Steni, 1100—

1745 m, 5 &,5 2: Mi Pins, 1000 tm, 231973, A

Malicky, 1 9; cap Soúnion, 10—12.1v.1979, M. De-

thier & N. Doneux-Stiernet, 1 d,1 2 (all CW).

Diagnosis. — See the descriptions by Ramme

(1933) and Harz (1969).

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 193— 195. Poecilimon ikariensis sp. n.; 193, 194, d holotype; 195, £ allotype.

Variauon. — This species is known to be

quite variable. The cercus of the males before

me varies as in figs. 122—124. Further variation

has been figured by Ramme (1933). The basal

fold of the lower ovipositor valve (figs. 263—

266) is strong, protruding vertically downward,

widely arched upward anteriorly, vertically im-

pressed in the middle and forming with the go-

nangulum an elongate vertical groove, which

may cause a narrow or wide incision of the out-

WILLEMSE: Greek Poecilimon 187

198

Figs. 196—198. Poecilimon klisuriensis sp. n.; 196, 197, d holotype; 198, 2 allotype.

er margin of the fold and form a pair of lobes. (Ramme, 1933; Werner, 1937a; Harz, 1969); Kimi

Distribution. — Confined to Evvoia, Attiki, (“Kymi”) (Werner, 1938; Harz, 1969); Sténi (Werner,

the adjacent NE part of the Pelopónnisos, and en ee Stent, roca al To ud

: thinai (“Athen” runner von attenwyl, 1878,

somie of the offshore islands. 1882: Harz, 1969); Voula (“Wula”) (Werner, 1934,

3 Pankräti (Werner, 1934); Mt. Imittös (“Hymettos”)

Waele Ere bou — (“Eu “@Wemere190249 1927-5 Rara me 1933: emer, 1933a,

boea”) (Werner, 1933b); Khalkís (“Chalkıs”) 1937b); Mt. Párnis, 1000 m; Akr. Sounion; Makronisi

188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

I. (Werner, 1937b); Saronic Is.:

(Werner, 1937b, 1938; Harz, 1969).

Pelopónnisos: Korinthía: between Palaiokörinthos

and Akrokorinthos (Werner, 1934); Argolis: Galatas

(Werner, 1938; Harz, 1969); Spétsai I. (Werner,

1937b). (Map 4.)

Aiyina (“Aegina”)

Remarks. — The Síros (“Syra”) record in the

original description refers to P. aegaeus, the

Parnassós (“Parnass”) one to P. zimmert.

Poecilimon aegaeus Werner, 1932

(map 4)

Poecilimon propinguus; Brunner von Wattenwyl,

1878: 44 (only Syra); 1882: 267 (idem).

Poecilimon aegaeus Werner, 1932: 2 (type-locality:

Mykonos); 1933a: 403, fig. 1. Ramme, 1933: 552,

PL 7 ine, 19, joll, Io nes, Sil, Sila, pl, IN me. Al, poll

12 fig. 32. Harz, 1969: 151, figs. 337, 481—496.

Diagnosis. — See the descriptions in Ramme

(1933) and Harz (1969).

Distribution. — Known only from some is-

lands of the Kiklädhes and Northern Sporädhes.

Localities. — Kikládhes: Mikonos (“Mykonos”)

(Werner, 1932, 1933a; Ramme, 1933; Harz, 1969);

Siros (“Syra”) (Brunner von Wattenwyl, 1878, 1882;

Ramme, 1933; Harz, 1969); Tínos (Ramme, 1933;

Harz, 1969).

Northern Sporadhes:

1933). (Map 4.)

Skíros,

Linarıa (Ramme,

Poecilimon mytilensis Werner, 1932

(map 4)

Poecilimon mytilensis Werner, 1932: 3 (type-locality:

Mytilene). Ramme, 1933: 537, pl. 6 fig. 24, pl. 9

fig. 29, PL 11 fig. 23, pl. 12 fig. 20. Werner, 1933a:

404, tig. 2; 1934: 324. Harz, 1969: 144, figs. 300,

5891 441445.

Poecilimon brevicauda Werner, 1932: 3 (type-locali-

ty: Lemnos). Ramme, 1933: 537 (as synonym).

Diagnosis. — See Ramme (1933) and Harz

(1969).

Distribution. — Known only from the Ae-

gean islands of Lésvos and ? Limnos.

Localities. — Eastern Aegean Is.: Lésvos (“Mytile-

ne”) (Werner, 1932; Ramme, 1933; Werner, 1934;

Harz, 1969), Lebétimnos Mts. (Werner, 1934);

Limnos (“Lemnos”) (Werner, 1932; Ramme, 1933).

(Map 4.)

Remarks. — P. brevicauda is known only

from the female type which is considered to be

lost (Ramme, 1933). Additional material is

wanted to establish its synonymy with P. myti-

lensis.

Poecilimon deplanatus Brunner von

Wattenwyl, 1891

(map 4)

Poecilimon deplanatus Brunner von Wattenwyl, 1891:

27 (type-locality: Kós). Ramme, 1933: 538, pl. 9

fig. 32, pl. 11 fig. 26, pl. 12 fig. 23. Werner, 1936:

11. Harz, 1969: 144, figs. 299, 446 —449.

Distribution. — Known only from some is-

lands in the Dhodhekánisos.

Localities. — Dhodhekanisos: Kös (Brunner von

Wattenwyl, 1891; Ramme 1933; Harz, 1969); Käsos

(Ramme, 1933; Harz, 1969); Karpathos (Ramme,

1933; Harz, 1969), Mt. Lastos (“Lastrosgebirge”)

(Werner, 1936). (Map 4.)

Poecilimon sanctipauli Brunner von

Wattenwyl, 1878

(figs. 125, 126, 267— 270, map 4)

Poecilimon sanctipauli Brunner von Wattenwyl, 1878:

40, figs. 2a—d (type-localities: Ephesus; Smyrna;

Rhodos). Giglio-Tos, 1914: 3. Ramme, 1933: 535,

pl. 7 fig. 10, pl. tig. 27 p iil wie, il. jell, 12 ive.

21. Werner, 1933b: 190; 1936: 10. Jannone, 1936;

145. Bei-Bienko, 1954: 288, fig. 159. Harz, 1969:

144, figs. 436—440.

sus, v.1901, Werner, 1 6 (both OW)E A Greece:

Rödhos, Lindhos, 17—18.iv.1970, A. C. & W. N. El-

lis,6 8,6 2 (ITZ; CW).

Diagnosis. — See the descriptions by Brunner

von Wattenwyl (1878), Ramme (1933), Bei-

Bienko (1954) and Harz (1969).

Variation. — The armature of the cerci of the

available males differs slightly from the descrip-

tions in presenting a separately placed pre-api-

cal spine (figs. 125, 126). The basal fold of the

lower ovipositor valve is also somewhat vari-

able, though always placed transversely, bul-

bously inflated in the middle and forming with

the gonangulum an elongate vertical groove

(figs. 267270).

Distribution. — This species is known from

SW Turkey and some offshore Aegean islands,

from Samos to Ródhos.

Greek localities. — Eastern Aegean Is.: Samos,

Marathokambos (Werner, 1933b; Harz, 1969); Dhod-

hekanisos: Kos (Ramme, 1933; Harz, 1969), Mt.

Dikeo (Werner, 1936); Ródhos (Brunner von Watten-

wyl, 1878; Ramme, 1933; Harz, 1969), “Cannamat”

& “Phileremos” & Monolithos (Werner, 1936),

Lindhos (Jannone, 1936), Ayios Isidhoros (Giglio-

Tos, 1914). Kalimnos: Potea (Werner, 1936). (Map 4.)

WILLEMSE: Greek Poecilimon 189

Figs. 199-206. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon species; 199, 200, P. ornatus (Schmidt) (Eptakhorion); 201, 202, P. hoelzeli Harz (Mt. Olimbos, Ret-

uge B); 203, 204, P. pindos sp. n. (Mt. Timfi); 205, 206, P. gracilis (Fieber) (Mt. Bela Voda).

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Poecilimon

x pancici

O ornatus

O hoelzeli

x pindos

A gracilis

A obesus

XK beieri

® nobilis

4 thoracicus

Map 1. Distribution of Poecilimon species.

Poecilimon hamatus Brunner von Watten wyl,

1878

bites. 127, 1274, 271, 277, imap 4)

Poecilimon hamatus Brunner von Wattenwyl, 1878:

41 (type-localities: Rhodos; Tireh bei Smyrna).

Werner, 1901: 286. Giglio-Tos, 1914: 3. Werner,

1933b: 190. Ramme, 1933: 538, pl. 9 fig. Sil, fol, Mi

fig. 25. Werner, 1934: 323, fig. 2a, b (not Ikaria);

1936: 11. Jannone, 1936: 144. Bei-Bienko, 1954: .

317. Harz, 1969: 140, figs. 281, 328, 424—426.

Material studied. — Rédhos, Lindhos, 19.1v.1970,

A.C. & W. N. Ellis, 1 5, 1 2 (ITZ); Naxos, Sifonas,

26.v.1977, A. Malicky, 1 6,1 2 (CW).

Diagnosis. — See Brunner von Wattenwyl

(1878), Ramme (1933), Bei-Bienko (1954) and

Harz (1969).

Variation. — The male cercus is known to be

variable, that of the Ródhos male before me is as

in text-figs. 127, 127a. Also the basal fold of the

lower ovipositor valve is variable. A figure of

this structure of the Ródhos female at hand is

given (figs. 271, 272).

Distribution. — Known from the Izmir dis-

trict, W Turkey, from several eastern Aegean is-

lands, from Lésvos to Ródhos, and from the is-

land of Naxos, the Kikladhes.

Greek localities. — Eastern Aegean Is.: Lésvos

(“Mytilini”), Mt. Lebétimnos (“Lepetymnosgebirge”)

(Werner, 1934); Samos (Ramme, 1933), Kierki near

Marathókambos (Werner, 1933b).

Dhodhekanisos: Nisiros (Nysiros (Ramme, 1933);

Rédhos (Brunner von Wattenwyl, 1878; Werner,

1901; Ramme, 1933; Harz, 1969), Mt. Attaviros

(“Mt. Attairo”) (Werner, 1936), Lindhos (Jannone,

1936), Ayios Isidhoros (Giglio-Tos, 1914).

Kikladhes: Naxos, Sifonas. (Map 4.)

Remarks. — Werner’s records (1934) from

the island of Foúrnoi need confirmation.

WILLEMSE: Greek Poecilimon 191

Poecilimon

O laevissimus

@ jonicus jonicus

O jonicus lobulatus

B werneri

* tessellatus

Map 2. Distribution of Poecilimon species.

DOUBTFUL LITERATURE RECORDS

Poecilimon hadjisarandou

Werner, 1938

Poecilimon hadjisarandou Werner, 1938: 167, fig. 4

(type-locality: Taygetos?). Harz, 1969: 140.

Remarks. — Known only from the male type

which, according to the description, is depos-

ited in the Athens Museum. During my visit to

this museum (June 1974), the specimen could

not be traced. The type-locality, as noted by the

author, is doubtful. Due to the insufficient de-

scription and lack of any further material, the

identity of this taxon is not clear.

Poecilimon geoktschaicus Shchelkanovtsev,

1910

Poecilimon bosphoricus; Uvarov, 1923: 148.

Poecilimon geoktschaicus; Ramme, 1933: 559, 562, pl.

6 fig. 47 pl. 10 figs. 59—60, pl. 11 tig. 48; 1939:

49. Beî-Bienko, 1954: 343, 353.

(CW). Greece: Macedonia, Lembet, June 1916, M.

Burr, Poecilimon bosphoricus Br.W.? det. Uvarov,

Poecilimon bidens geoktschaicus Shch Ramme der,

“Fundort vermutlich falsch, wahrscheinlich Geok-Ta-

pa”, Ramme, 1 d (BMNH).

Remarks. — These records refer to material

from Lembet and “Happy Valley”, both near

Thessaloniki. The male before me, labelled

Lembet, agrees completely with the Russian

material. Both Ramme (1939) and Beî-Bienko

(1954) assume that the locality labels have been

exchanged with Burr’s material from the Cauca-

sus.

Poecilimon sp. Ebner, 1954

Poecilimon sp.; Ebner, 1954: 554.

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 20/— 214. Lateral and ventral view ot the basal told ot the dorsal margin ot the lower ovipositor valve

in Poecilimon species: 207, 208, P. obesus Brunner von Wattenwyl (Kapsia); 209, 210, P. beter: Ramme (allo-

type); 211, 212, P. nobilis Brunner von Wattenwyl (Khrisovitsi); 213, 214, P. thoracicus (Fieber) (Tukheron).

WILLEMSE: Greek Poecilimon 193

Poecilimon

X orbelicus

@ macedonicus Ù

O brunneri

* cretensis ®

A ikariensis

A ebneri

A klisuriensis

Im miramae

Map 3. Distribution of Poecilimon species.

Remark. == Ihe record refers to a female

from Arta, Ipıros.

Poecilimon sp. Ebner, 1912

Poecilimon sp.; Ebner, 1912: 111.

Remarks. — Refers to juvenile specimens

from the island of Kithira (= Cerigo), off the

shore of the southern Pelopönnisos. Hitherto

no other record of the genus from this ısland ıs

known.

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Brunner von Wattenwyl, C., 1878. Monographie der

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——, 1882. Prodromus der europäischen Ortho-

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——, 1891. Additamenta zur Monographie der Pha-

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Doflein, Fr., 1921. Mazedonien: i-viii, 1—592, figs. —

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Ebner, R., 1912. Zur Kenntnis der Orthopterenfauna

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168—176, 184-188, 201—207, 232—238, 252 —

258.

Fischer, L. H., 1853. Orthoptera Europaea: ı-xx, 1—

194 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Poecilimon

anatolicus

chopardi

zimmeri

thessalicus

propinquus

aegaeus

mytilensis

deplanatus

* sanctipauli

4 hamatus

«EX Sooex

Map 4. Distribution of Poecilimon species.

454, figs. — Engelmann, Leipzig.

Frivaldszky, J., 1867. Monographia Orthopterorum

Hungariae: 1—201, figs. — Eggenberger, Pest.

Giglio-Tos, E., 1914. Escurzioni zoologiche del Dr.

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Harz, K., 1964. Orthopterologische Beiträge V. —

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——, 1966. Neues von europäischen Orthopteren. —

Mitt. deutsch. Ent. Ges. 25: 21—24, figs.

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Harz, K. & A. Kaltenbach, 1976. Die Orthopteren

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Jannone, G., 1936. Nuovi contributi alla conoscenza

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Karaman, M., 1958. Neue Orthopteren-Arten aus

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——, 1961. Beitrag zur Kenntnis der Orthopteren

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Ramme, W., 1926. Neue und wenig bekannte euro-

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WILLEMSE: Greek Poecilimon 195

liens und Kreta. — Eos 3: 111—200, figs.

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der Orthopteren von Südost-Europa und Vorder-

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Schmidt, F. I., 1849. (Beschreibung der Ephippigera

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(9): 184.

Tarbinskii, S. P., 1932. Materialy k poznaniyu prya-

mokrylykh nasekomykh SSSR. Izv. leningradsk.

— Inst. Bor’by s Vreditelyami Sel’skogo 1 Lesno-

go Khozyaistva 2: 181—205, figs.

Uvarov, B., 1923. A list of Orthoptera of Macedonia

with zoogeographical remarks. (In: M. Burr, B. P.

Campbell & B. Uvarov, A contribution to our

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Verh. zool.-bot. Ges. Wien 1903: 65—69; figs.

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——, 1929. Zoologische Forschungsreise nach den Jo-

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——, 1932. Neue griechische Orthopteren aus dem

Gebiet des Aegaeischen Meeres. I. — Anz. Akad.

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——, 1933a. Ueber Orthopteren aus Ost-Griechen-

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— Mitt. zool. Mus. Berl. 18: 395—415, figs.

, 1933b. Ergebnisse einer zoologischen Studien-

und Sammelreise nach den Inseln des Aegaeischen

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Wien (1) 142: 185—204.

——, 1934. Dritter Beitrag zur Kenntnis der Tierwelt

der Aegaeischen Inseln. — Sber. Akad. Wiss.

Wien (1) 143: 313—337, figs.

, 1936. Ergebnisse einer zoologischen For-

schungsreise nach dem Dodekanes, unternommen

von Kustos Dr. Otto Wettstein. 1. Orthopteren

aus dem Dodekanes. — Sber. Akad. Wiss. Wien

(1) 145: 9—16.

— — 1937a. Ergebnisse der vierten zoologischen

Forschungsreise in die Aegaeis (1936). — Sber.

Akad. Wiss. Wien (1) 146: 89— 118, figs.

——, 1937b. Beiträge zur Kenntnis der Tierwelt des

Peloponnes, der Inseln Kythira und Euboea sowie

der kleinen Inseln im Saronischen Golf. — Sber.

Akad. Wiss. Wien (1) 146: 135—153, figs.

— — 1938. Ergebnisse der achten zoologischen

Forschungsreise nach Griechenland (Euboea, Ti-

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Wien (1) 147: 157—173, figs.

Willemse, F., 1974. A new subspecies of Paranocara-

‘ cris bulgaricus (Ebn. & Dren.) from Greece

(Orthoptera, Acridoidea, Pamphaginae). — Biolo-

gıa gallo-hellenica 5: 351—352, figs.

— — 1977. Interesting distribution records of

Orthoptera from the Greek mainland and some

neighbouring islands. — Ent. Ber., Amst. 37: 52—

59, map.

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of Crete. — Tijdschr. Ent. 119: 123—164, figs,

map.

196 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 215— 222. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon species: 215, 216, P. laevissimus (Fischer) (Ano Exanthia); 217, 218, P. jonicus jonicus (Fieber)

(Dhrosopiyi); 219— 222, P. jonicus lobulatus ssp. n.; 219, 220, Métsovon; 221, 222, Niképolis.

WILLEMSE: Greek Poecilimon 197

Figs. 223—230. Lateral and ventral view ot the basal told ot the dorsal margin ot the lower ovipositor valve in

Poecilimon species: 223, 224, P. werneri Ramme (Lekhaina); 225, 226, P. tessellatus (Fischer) (Khrisovitsi); 227,

228, P. macedonicus Ramme (Elassón); 229, 230, P. brunneri (Frivaldsky) (Izvor).

198 TITDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 231—238. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon species: 231, 232, P. cretensis Werner (Mt. Idhi); 233, 234, P. ikariensis sp. n. (allotype); 235, 236,

P. syriacus Brunner von Wattenwyl (Samos); 237, 238, P. ebneri Ramme (Mt. Bela Voda).

WILLEMSE: Greek Poecilimon 199

Figs. 239—246. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon species: 239, 240, P. klisuriensis sp. n. (allotype); 241, 242, P. zwicki Ramme (Stavroúpolis); 243,

244, P. orbelicus Pancic (Bulgaria); 245, 246, P. anatolicus Ramme (Mt. Pangaion).

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 247—254. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon species: 247—252, P. chopardi Ramme; 247, 248, Paraskevi; 249, 250, Mt. Timfristós; 251, 252, Pal-

aiokhóri; 253, 254, P. zimmeri Ramme (Mt. Parnassós).

WILLEMSE: Greek Poecilimon 201

Figs. 255—262. Lateral and ventral view of the basal fold of the dorsal margin ot the lower ovipositor valve in

Poecilimon species: 255—258, P. zimmeri Ramme; 255, 256, Mt. Panakhaikön; 257, 258, Mt. Panaitolikon;

259—262, P. thessalicus Brunner von Wattenwyl; 259, 260, Mt. Ossa; 261, 262 Leptokaria-Karia.

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 6 (1982)

Figs. 263—270. Lateral and ventral view of the basal fold of the dorsal margin ot the lower ovipositor valve in

Poecilimon species: 263—266, P. propinquus Brunner von Wattenwyl; 263, 264, Mt. Dirfis; 265, 266, Mt.

Parnis; 267—270, P. sanctipauli Brunner von Wattenwyl; 267, 268, Smyrna; 269, 270, Lindhos.

WILLEMSE: Greek Poecilimon 203

Figs. 271, 272. Lateral and ventral view of the basal fold of the dorsal margin of the lower ovipositor valve in

Poecilimon hamatus Brunner von Wattenwyl (Lindhos).

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DÒ DEEL 125 AFLEVERING 7 1982

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

MOLOG, N

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S E 2 È UBRARY

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ON È DI VERSITY

INHOUD

J. VAN DEN Assem, M. J. Gijswijr & Bircit K. NùseL. — Characteristics of

courtship and mating behaviour used as classificatory criteria in Eulophidae-

Tetrastichinae (Hymenoptera), with special reference to the genus Tetrastichus

s.l., p. 205— 220, figs. 1—13.

Tijdschrift voor Entomologie, deel 125, afl. 7 Gepubliceerd 31-XII-1982

CHARACTERISTICS OF COURTSHIP AND MATING

BEHAVIOUR USED AS CLASSIFICATORY CRITERIA IN

EULOPHIDAE-TETRASTICHINAE (HYMENOPTERA), WITH

SPECIAL REFERENCE TO THE GENUS TETRASTICHUS sl.

J. VAN DEN ASSEM!), M. J. GIJSWIJT?) and BIRGIT K. NUBEL!)

ABSTRACT

The usefulness of behavioural characters for taxonomic work was tested in a comparative

study of courtship and mating behaviour of Chalcidoidea-Tetrastichinae. In particular,

species-groups of the large genus Tetrastichus were scrutinized. Similarities and differences

are discussed. Transformations between character-states are found which seem to reflect

developments in the evolution of this group of wasps. Earlier, parallel developments were

traced in other families of Chalcidoidea (e.g. in Pteromalidae, Encyrtidae).

Within the Eulophidae, the subfamily Tetra-

stichinae represents a morphologically well-de-

fined entity. However, the present subdivision

of this taxon is unsatisfactory. The current prac-

tice (Boucek, 1977) is to distinguish 17 genera

(exclusive of several poorly known Girault gen-

era), most of which with a single or a few spe-

cies. Amongst this group Tetrastichus stands

out with well over 200 species in Europe alone

(and a comparable number in North America,

ignoring other areas). Tetrastichus is subdivided

in species groups and a number of species solae,

but the general feeling is that it does not repre-

sent a natural entity. Very probably some of the

species groups should be raised to generic rank.

Graham (1961) realized the necessity to split up

Aprostocetus (= Tetrastichus), but he expressed

as his opinion “that such subdivision will have

to be based upon new combinations of charac-

ters, those hitherto employed being unreliable

for generic diagnosis”.

It is at this point that we venture to present a

type of character not employed so far in this

group. Earlier (Van den Assem & Povel (1973);

Van den Assem (1974)) characteristics of

courtship behaviour as taxonomic criteria were

used in a few groups of Pteromalidae and we

presumed that in the Tetrastichinae courtship

could also offer a set of valuable characters for

defining entities above the level of species.

Authors’ addresses: 1) Dept. of Zoology, University of

Leiden, and ?) Agrobiological Laboratory Duphar BV,

*s-Graveland, both The Netherlands.

205

For various reasons the Tetrastichinae seem

to represent a favourable group for a compara-

tive study of courtship behaviour. Recently, a

number of entomologists has become interested

in this group and, most important, are preparing

revisions of several genera of Tetrastichinae.

These revisions will offer good opportunities

for a comparison of arrangements of species and

groups of species (and assignment of generic-

status) on morphological criteria with one based

on characteristics of display behaviour.

There were additional reasons for choosing

the Tetrastichinae. Preliminary observations led

us to suppose that in this group, and in the

whole complex of Eulophidae for that matter,

evolution of courtship had proceeded parallel to

developments traced in other groups of Chalci-

doidea, viz., Pteromalidae and Encyrtidae. Te-

trastichus especially is extremely rich in clusters

of closely related species. Comparison of dis-

plays may give an idea of processes involved in

species radiation.

We thought it possible to collect a good many

species within a reasonable period of time and,

judging by our experience with Pteromalidae,

sa behaviour did not need complicated

arrangements before it could be observed in the

laboratory. As to this latter point, our expecta-

tions have not completely been realized. We

have collected 39 species of Tetrastichus and

about a dozen of Melittobia (a review of our Te-

trastichus material is given in the Appendix), but

some of them never courted, not even when we

patiently waited for hours. It is impossible to

206 TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

understand why this should be so. Presumably

one or more environmental factors were amiss.

For example, the males of 7. asparagi did not

court conspecific females in a bare glass vial, or

only briefly, with long intervals. Adding a small

quantity of dry sand made the males court im-

mediately, and removal of the sand made them

stop again. Adding sand to other non-courting

species did not have the desired effect.

With some species we obtained only very few

specimens and sometimes a male had died be-

fore one or more females had emerged. In yet

other species males proved to be very rare or

absent. Thelytoky is presumed to be the rule in

those species.

CHARACTERISTICS OF COURTSHIP BEHAVIOUR

SHARED BY ALL TETRASTICHINAE

A number of characteristics of the courtship

display are shared by all species of Tetrastichi-

nae. These general aspects are reviewed below.

More specific properties of displays will be dealt

with in the following sections.

Prior to the onset of courtship proper, the

male mounts the female and takes up a

courtship position on top of her. The precise

placement of the male’s front tarsi varies be-

tween species (on the thorax, neck, back of

head, or face of the female), but the position al-

most never varies within a species (7. lasiocera

was the only exception).

The position of the male’s head relative to the

female’s changes periodically during the perfor-

mance of his display, and accordingly we distin-

guish “high”-phase episodes and “low”-phase

Qs)

Fig. 1. Schematic representation of the positions of

male relative to female during the “high”- and “low”-

phase episodes; a, b, high, resp. low from aside; c, d,

high, resp. low positions of heads from the front.

episodes; see fig. 1 for a schematic representa-

tion. Usually these changes are very conspicu-

ous (except in a few species where it is less so,

and only rarely high phases appear to be en-

tirely absent). The high-phase position of the

male is achieved primarily by stretching his

front legs, his front tarsi remaining in the same

position throughout. As a consequence his head

comes further from the female’s than during the

low phase (when the male has bent his front legs

to some extent). In the low phase a contact be-

tween heads or their appendages (antennae,

mouth parts) is a frequent occurrence.

There is a second alternation, synchronous to

the high-and-low one: in the low phase the

male’s entire body is situated more to the front

(in several species rather extremely so) and in

the high phase more to the rear, relative to the

female. These shifts are also due to the

movement of the male’s legs, while his front tar-

si remain in the same position.

A sequence of male courtship consists of a

repetition of identical bouts of movements. In

some species bouts of one type of movements

alternate with bouts of another; in other species

successive bouts may be separated by intervals

which seem to be “rest” periods.

They can be presented schematically as fol-

lows:

low: bout — high:

high: rest, etc.,

or,

low: bout a > high: bout b — low: bout a

— high: bout b, etc.

In this respect the majority of Melittobia spe-

cies are the single exception we know of: here a

sequence of courtship is not confined to the

mere repetition of similar elements but new el-

ements come in underway (Van den Assem,

1975).

A courting male may move several limbs in

fixed patterns. The details of the movements are

specific but all species perform movements with

the antennae and almost all vibrate wings. In

cases where we have recorded these vibrations

on tape, we obtained specific sound patterns

(Van den Assem & Putters, 1980). In some gen-

era leg movements are involved, but in Tetrasti-

chus such cases seem to be rare.

When sexually receptive, the female raises her

abdomen to expose the genital orifice. At the

same time she lowers herself to the substrate

and stretches her head to the front with the an-

tennae pointing upwards. This posture holds for

all Tetrastichinae. Females may become recep-

rest — low: bout —

VAN DEN ASSEM, Gijswijr & NÜBEL: Courtship and mating behaviour 207

tive at certain points of a male’s display, e.g., al-

ways at the end of a low phase. Since the male

repeats similar motor patterns periodically, the

onset of female receptivity cannot be predicted

accurately beforehand. Melittobia females are

exceptional again: they will indicate receptivity

at only one single point, following the male’s fi-

nale (Van den Assem, 1975).

Following copulation a male may either walk

away from the female, or he may resume

courtship from the position characteristic for

the species. Post-copulatory sequences were ob-

served in some species. The motor patterns in-

volved in post-copulatory displays are identical

to those of pre-copulatory displays; post-copu-

latory sequences are usually rather short.

CHARACTERISTICS OF COURTSHIP DISPLAYS OF

TETRASTICHUS SPECIES GROUPS

Within the genus Tetrastichus there are sever-

al morphologically recognisable species groups.

With a number of such groups we were lucky

enough to collect more than one species. The

first question is, whether there is a greater re-

semblance in courtship displays between species

of one species group than between species from

different groups. Resemblances and differences

were found indeed, and we will return to this

point in the discussion. Below we give descrip-

tions of the displays of different species groups

of Tetrastichus. A few additional species which

belong to related genera (and which happened

to turn up in our collection) are also discussed.

We have tried to follow a standard pattern in

our descriptions, and abbreviate as much as pos-

sible to avoid repetition and save space. The

groups are arranged in an alphabetical order.

Fig. 2. T. arundinis

group; a, high phase posi- a

tions; b, low phase posi-

tions of 7. spec. 9 from ZON

aside; c, position of male

antennae of 7. spec. 9

seen from front, and d,

seen from top; e, T. arun-

dinis, positions of male

head in low and high

phases; f, T. arundinis,

low phase positions seen

from side. d

a. Tetrastichus, group arundinis; species ob-

served: 7. arundinis, T. gratus, T. spec. 9 (fig.

In general there is a distinct alternation of

“high, and less frontal” and “low, and more to

the front” phases. The male front tarsi are

placed on the female’s head, usually on her face.

During the high phase the male holds his anten-

nae in a curved position above the female’s and

vibrates them at a high frequency. At the onset

of a low phase the male bows low, lowering his

antennae and waving them in front of the female

(e.g., in spec. 9; fig. 2b, c, d), or, instead, the

male actually grasps the female’s antennae brief-

ly, holding his antennae in such a way that his

flagellar parts point backwards and touch the

female’s mouth parts. At the onset of a high

phase the male raises his antennae slowly. The

female indicates receptivity at the end of a low

phase. Copulation lasts about 5 secs. No post-

copulatory courtship observed.

In 7. arundinis males the alternation of low

and high (in this species it is far more an alterna-

tion between less frontal and extremely frontal)

was most pronounced; in spec. 9 it was far less

pronounced. The alternation appeared com-

pletely stereotyped in 7. arundinis, contrary to

spec. 9 where the high-phase duration was vari-

able. Vibrating the antennae above the female’s

antennae during the high phase was most con-

spicuous by far in 7. gratus. Embracing the fe-

male’s antennae during the low phase was most

pronounced in 7. arundinis, the males grasping

the female’s antennae from the sides with their

own antennae, pushing them together, freezing

in this posture for a second or so, their claval |

parts in contact with the female’s mouth parts

ge Ge |

208

di on

Fig. 3. 7. brevinervis group; T. lasiocera; a, high

phase, b, low phase positions, in sideview.

(fig. 2f). Leg movements (series of drumming

the front tarsi on the female’s head) were ob-

served in T. gratus; they did not occur in the

other species.

b. Tetrastichus, group brevinervis; species ob-

served: T. lasiocera (fig. 3).

There is no clear difference between high and

low positions of the male. Alternation of phases

stereotyped, both of equal and short duration.

The male front tarsi are placed on female’s

head (but sometimes on the antero-lateral mar-

gin of her thorax, henceforward to be called her

“shoulders”, especially so during post-copula-

tory courtship?). Antennae point upwards in

both sexes, male antennae in quivering motion

all the time (fig. 3a).

Male lowers antennae sideways and up again

in a single rapid movement. Female receptive

with male antennae in low position. Genital

contact about 2 seconds. Very short post-copu-

latory courtship.

c. Tetrastichus, group caudatus; species ob-

served: T. dotus, T. epicharmus (fig. 4).

There is an alternation of high and low

phases, the duration of the high phase is vari-

able.

fae

a b

Fig. 4. T. caudatus group; T. epicharmus; a, high

phase positions, and b, low phase positions, in side-

view.

Male front tarsi are placed on the female’s

“ » > =

shoulders”. The female’s antennae point up-

wards throughout, the male’s antennae point

upwards during high phases only.

TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

A low phase starts with the male bending

slightly forward. His antennae are bent low and

are in a quivering motion in front of the female’s

head. The male’s body is rocking from side to

side and his wings are vibrating. The male an-

tennal movements are very simple, there is no

raising sideways nor rubbing against or hitting

on the female’s antennae.

Copulation follows at the end of a low phase.

Males of 7. epicharmus pressed their “face”

against the female flagellar parts at the onset of

the low phase, and their wings vibrated in very

distinct pulses. During the low phase they

swayed their middle legs far to the front. Such

leg movements seem to be exceptional in Te-

trastichus, this is the only species in which we

observed them. Similar movements occur in all

Melittobia. At the end of a low phase males of

T. epicharmus quiver the antennae in front of

the female’s antennae; males of T. dotus hold

the antennae alongside the female’s antennae for

a short moment and then resume the high-phase

position.

d. Tetrastichus, group daira; species observed:

T.daira, TN diaphantus, 1. specs > MER peli

(fig. 5).

There appears to be some discrepancy be-

tween the display of 7. daira and those of the

other species. However, there are also striking

similarities.

In 7. daira alternation of phases occurs not so

much between high and low but rather between

more-frontal and less-frontal. The less-frontal

phase is shorter than the more-frontal.

Males of daira place the front tarsi on the fe-

male’s “shoulders”. In the less-frontal phase the

female’s antennae point forward, as do those of

the male.

At the start of the more-frontal phase the

male performs series of antennal quivering

(moving his antennae in between her’s, while

knocking them on her head) and he rocks his

body from side to side (fig. 5a). At the end of

the quivering episode, the female has raised her

antennae (fig. 5b, c). The male raises his side-

ways, keeps them thus for short moment (in a

position resembling that of miser-group species)

and proceeds with series of (about three) lashing

movements along the outside of the female’s an-

tennae, hitting her on or near her mouth parts

(fig. 5d). These hits are accompanied by wing

flicks.

Females become receptive at the end of lash-

ing series; a receptive female’s head is only

VAN DEN AsseM, Gijswijr & NùseL: Courtship and mating behaviour 209

a b

c d e

f g

ETTI ae

Fig. 5. 7. daira group; a—e, T. daira; f—ı, T. di-

aphantus c.s.; a, antennal quivering posture, and b,

hitting the female’s head; c, antennae raised; d, anten-

nae hitting female’s head in sideview, the arrow indi-

cates the way of raising antennae in between hits; e,

antennae raised in between hits, from front; f, g, high

and low phase positions; h, antennal hits, seen from

front, and i, from top.

slightly forward. Genital contact about 5—10

seconds. 7. daira males always performed post-

copulatory courtship and females showed re-

newed receptivity, which resulted in all of them

in a succession of four copulations.

In the other daira-group species differences

between high and low were more distinct (fig.

5f, g) than in 7. daira. In these species male

front tarsi are placed on the female’s head (most

extreme in 7. diaphantus with front feet placed

on her face). Female antennae never point up-

wards and male antennae are curved and held

forward above the female’s during high phase.

During the low phase male antennae follow a

daira-like pattern. There is a quivering episode

at the beginning and male antennae make rhyth-

mical contact with the female’s flagellar parts

near her pedicel. His mouth parts are near her

pedicel. The male’s body rocks slightly from

side to side. The male proceeds with raising the

antennae sideways and hits the female’s anten-

nae from the outside a few times in rapid suc-

cession (fig. 5h, i). Antennal hits are accompa-

nied by wing flicks. Males of spec. 4 drum the

hind legs on the female’s wing edges following

antennal hits.

Female receptivity occurs during the anten-

nal-lashing episode. Genital contact about 5 sec-

onds. There were sequences of post-copulatory

courtship.

e. Tetrastichus, group evonymellae; species ob-

served: 7. evonymellae, T. galactopus (fig. 6).

The alternation of high and low phases is very

distinct, the high phase is of a variable duration.

Male front tarsi are placed on the female’s

“shoulders”. Female antennae point upwards

throughout. During the high phase the male’s

antennae are in a similar position, forming a

sharp figure V, his wings standing up (fig. 6a).

At the beginning of a low phase the male low-

ers his head until it almost touches the female’s

vertex, while spreading his antennae sideways.

This movement is followed by rapidly rubbing

his antennae along, or knocking them against

the outside of hers (fig. 6b). The female’s head

did not appear to be hit (movements are ex-

a Cc

Fig. 6. 7. evonymellae group; a, head positions and

positions of the antennae during the high phase, in

sideview and from front; b, and c, idem, during low

phase.

210 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

Fig. 7. T. fulvipes group; a, high phase, and b, low

phase positions of 7. calamarius; c, copulation postu-

re of female.

tremely rapid) (fig. 6c). Knocks come in series

of one to three, at the end of a series the male

resumes the high-phase position.

The female becomes receptive during anten-

nal knocking, she lowers the frontal part of her

body, stretching her head to the front,

sometimes almost resting it on the substrate, ab-

domen maximally raised, a posture characteris-

tic of all Tetrastichinae (see p. 206). Genital

contact during less than one second. Short se-

quences of post-copulatory courtship were ob-

served in 7. evonymellae.

f. Tetrastichus, group fulvipes; species observed:

T. calamarius (fig. 7).

High and low phases alternate distinctly. The

low-phase position of the male is very extreme;

his high-phase position is mainly due to stretch-

ing of the middle legs, the front legs are fre-

quently taken off the female. Duration of low

phase about 2 seconds.

Male front tarsi are placed on the female’s

head. During the high phase the female’s anten-

nae point to front, male antennae are raised

sideways and to front.

At the onset of the low phase the male bends

far over the female’s head, drumming his front

tarsi on her eyes, his antennae touching or

drumming on her mouth parts, his mouth parts

very near her mouth parts and his wings vibrat-

ing. Male antennal movements are very simple.

The female’s antennae are folded in front of

head during the low phase. Males may perform

a kind of stepping movements with their hind

legs on the female’s wing edges during the low

phase.

Female receptivity occurs at end of antennal

drumming. Genital contact lasts about 30 sec-

onds. There was always some post-copulatory

display. A female may signal renewed receptivi-

ty, and, if so, another copulation will follow be-

fore the male walks off.

g. Tetrastichus, group lyadas; species observed:

T. abydenus, T. pallipes, T. strobilanae, T. spec.

5, T. spec. 6 (fig. 8).

In general the alternation of high and low

phases is rather distinct. Low phase usually of

short duration, high phase longer and more

variable.

Male front tarsi are placed on the female’s

IT goee 6,

Fig. 8. 7. lycidas group;

a, and c, positions during

high phase; b, and d, po-

sitions during low phase; DÌ 17

e, low phase positions of

Cc d

VAN DEN Assem, Gijswijr & NÜBEL: Courtship and mating behaviour Dil

“shoulders”. Female antennae point upwards brates his wings and brings his mouth parts near

throughout. The position of the male’s antennae the female’s pedicel, sometimes pressing his face

seems to vary between species. (Upwards in against her flagella. His antennae are lowered

pallipes, curved to front in abydenus, spread and spread sideways to the front. At the same

sideways in strobilanae, fig. 8a.) time he starts series of head-shaking movements

At the beginning of a low phase the male vi- which bring his antennae in alternate contact

eo 8.

alal

Fig. 9. T. miser group; a—c, T. asparagi c.s.; dI, T. atriclavus; a, and d, positions during high phase; b, c, e, f,

male heads and antennae during drumming; j—l, male antennal postures in sequential order; j is the initial postu-

re, adopted during the high phase, i, is similar to k, but seen from aside, | is the drumming posture, seen trom

front.

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

with the female’s antennae from the outside, at

the same time rubbing his face against her fla-

gellar parts. Females were not receptive.

The display of 7. spec. 6 differed in some

points and seems to be of considerable interest

(see p. 217). High- and low-phase positions of

the male were more distinctly different than in

the other species. Male front tarsi were placed

on the female’s head. Male antennae were

curved to front during high phase. Instead of

head shakes, the male performed rocking

movements with the entire body during the low

phase: the tip of his abdomen remained station-

ary, his frontal parts swayed from side to side,

wings vibrating. Male antennae seemed to

“grasp” the female’s antennae at the ipsolateral

side during these rocking episodes.

h. Tetrastichus, group miser; species observed:

T. asparagi, T. atriclavus, T. hylotomarum, T.

pass 12 (ng, 9).

In general there is a distinct alternation of

high and low phases. The duration of the high

phase is variable, but in atriclavus durations are

fixed and both high and low phase are of an

equally short duration.

Male front tarsi are placed on the female’s

“shoulders” (asparagi, spec. 12, hylotomarum),

or on her head (atriclavus, presumably also in

incertus, fide Miller, 1966). Female antennae

point upwards throughout, forming a sharp fig-

ure V. Male antennae in a similar posture during

the high phase only (fig. 9a, d).

The low phase starts with the male bending

forward to some extent, vibrating his wings. A

sequence of antennal movements follows: an-

tennae are spread sideways (fig. 9b, e) and kept

in this position for a moment, then bent and

drummed on the female’s head in bouts of very

rapid movements (fig. 9c, f). The male abdomen

is raised very conspicuously, especially during

the low phase.

Female receptivity occurs during antennal

drumming. Genital contact one second or less.

No post-copulatory courtship.

Females of 7. atriclavus were observed to

head-shake during the low phase as long as they

remained unreceptive.

1. Tetrastichus clavicornis, species sola.

There is a distinct alternation of high and low

phases.

Male front tarsi are placed on the female’s

“shoulders”. Female antennae point to the

front, male antennae are raised, pointing high

and to the front.

At the beginning of a low phase the male low-

ers his body, vibrating his wings. His mouth

parts come near the female’s pedicels, his anten-

nae are bent, vibrating outside the female’s an-

tennae; there are no additional antennal

movements.

Only incomplete sequences of courtship were

observed, females were very unreceptive. The

observed position of the female’s antennae may

not be the normal position for females engaged

in courtship.

DISPLAYS OF OTHER TETRASTICHINAE

a. Courtship of Crataepus marbis (fig. 10).

Crataepus marbis reminds one of mole-crick-

ets, its external morphology being probably an

adaptation to moving through the dense pilosity

of the natural habitat (composite flower heads).

In Crataepus there is an extreme difference be-

tween positions in low and high phase. High

and low phases alternate very fast but they do

not seem to be completely rigid.

Male front tarsi are placed on the female’s

head, front legs are stretched, antennae are

curved above female’s.

At the onset of a low phase the male ands his

front legs, brings his mouth parts in contact

with the female’s pedicels and folds his antennae

against the female’s antennae from the outside.

The tips of his flagellae are pressed against the

female’s mouth parts. Male antennae are neither

spread nor raised. There are no wing vibrations

nor other movements with legs. Females recep-

tive at the end of a low phase; the female’s head

is only slightly forward in the copulation pos-

ture, her antennae point upwards. Genital con-

tact about 15 seconds.

Wes

Fig. 10. Crataepus marbis; a, high phase, and b, low

phase positions; c, and d, female head from front with

a male’s front leg on top (stretched during high, bent

during low phase).

VAN DEN AssEM, Gijswijr & NÜBEL: Courtship and mating behaviour 213

a b

Fig. 11. Nesolynx albiclavus; a, low phase position; b,

posture of male’s middle legs during the drumming

episodes.

b. Courtship of Nesolynx albiclavus (fig. 11).

The male assumes a low-phase position fol-

lowing mounting and the relative position of

male and female heads does not change during

the display.

Male front tarsi are placed on female’s face

near her scapi. The male raises his antennae,

spreading them slightly, and flutters his wings at

the same time. Then his antennae are spread fur-

ther and folded over the female’s antennae in

such a way that her antennae are pinched in be-

tween his flagellar parts. Males clearly search

for a hold of female antennae before proceeding

with the embrace. At the end of a pinching epi-

sode the male starts drumming his middle legs

on the female’s thorax with high frequency.

Synchronously he drums on her dorsum with

the tip of his abdomen which is bent down.

During drumming male antennae rub up and

down against female antennae.

Onset of female receptivity is at the end of an

antennal embrace; females apparently need rela-

tively long bouts of courtship to become recep-

tive. Genital contact 1—2 seconds. No post-co-

pulatory display.

c. Courtship of Melittobia acasta (fig. 12).

Melittobia species exhibit strong sexual di-

morphism. Males bear peculiarly shaped anten-

nae with inflated scapes which play an impor-

tant role in courtship. Melittobia acasta is the

only European species. Comparative work on

this genus will be published in a separate paper.

a b

Fig. 12. Melittobia acasta; a, high phase, and b, low

phase positions in sideview.

Male front tarsi are placed on the female’s

neck, the male grasps the flagellar parts of the

female and holds them in the concave distal area

of his scapes. A high phase follows immedi-

ately: male loosens his grip, lets female antennae

go, spreads and raises his antennae, stretching

front legs to some extent, and performs swaying

movements with his hind legs, more or less syn-

chronously, up and down the female’s thorax.

Then, the male bends forward again, grasps the

female’s antennae which were kept more or less

folded in front of her head, and starts quivering

his flagellae. At the end of a quivering series the

male claval parts touch the female’s mouth

parts. Quivering is concluded by a distinct

pinch by the pedicel and scape as the male flag-

ella and scape are drawn together. Contact be-

tween antennae is broken off again. The next

high phase, with hind leg movements etc., fol-

lows.

In the course of the display antennal contact

becomes permanent and the male no longer

takes up the high-phase position although the

rhythmic sequence of antennal and hind leg

movements goes on. Alternations occur more

and more rapidly until the movements almost

overlap. At this point the male’s finale sets in:

coordination of hind leg movements changes

conspicuously, they are rubbed against the fe-

male thorax, for a few seconds, like riding a bi-

cycle. The hind legs are placed on the female

again and now middle legs are taken off the fe-

male and moved far to the front (as far as the fe-

male’s head or even beyond) and back again. It

is only here that a female Melittobia acasta may

indicate sexual receptivity. There are no excep-

tions to this rule. The display of Melittobia dif-

fers from all the other Tetrastichinae in this im-

portant aspect: a sequence is not simply a repe-

tition of similar elements but is built up in time,

with new elements being added during the dis-

play.

A number of Melittobia species (e.g., mem-

bers of the hawaitensis-complex) do not alter-

nate between high and low-phase episodes but

relative position of heads remains about con-

stant from the beginning and antennal contact is

permanent throughout (Dahms, 1973). Melitto-

bia sequences last much longer than those of

other species (in M. acasta they are relatively

short, in the order of 1—5 minutes, in M. aus-

tralica may be up to half an hour!). Genital con-

tact during about 7 seconds. There is no post-

copulatory courtship.

214 TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

COMPARISONS AND DISCUSSION

Similarities between species groups

The species that were available for study be-

long to four genera: Tetrastichus Haliday, 1844;

Crataepus Förster, 1878; Nesolynx Ashmead,

1905 and Melittobia Westwood, 1847. Tetrasti-

chus species represent eight species groups and

one species sola. All species are enumerated in

the Appendix.

If we consider entire displays, our species fall

into two different categories. The first category

accommodates those species whose males per-

form a more or less elaborate antennal display

including relatively long contacts with the fe-

male’s antennae. Often it involves embraces: ei-

ther the male grasps the female’s antennae with

his own and holds them, or he pushes them to-

gether in between his, while he bends forward

over her head, sometimes touching her mouth

parts with his clava or even with his mandibular

palpi. In this category are placed Crataepus

marbis, Nesolynx albiclavus, Melittobia species,

the Tetrastichus arundinis group and Tetrasti-

chus calamarius.!)

Our second category accommodates the re-

maining species groups of Tetrastichus, whose

males neither hold the female’s antennae for

some time in some sort of embrace nor bend

forward over the female’s head. This second

category is not uniform, two or three clusters of

species groups can be distinguished. The general

nature of male display in this category is sugges-

tive of some kind of dichotomy. In one branch,

with the species groups lycidas and caudatus,

males exhibit distinct sideways tendencies in the

low phase of courtship, rocking the entire body

from side to side as in caudatus species or in ly-

cidas species 6, or shaking the head, as in the

other species of the lycidas group. In the other

branch, with the species groups evonymellae,

miser and brevinervis, the general direction ot

movements is parallel to the length axis of the

22, The position of the daira group is not

clear (see below).

The courtship display of the two observed

evonymellae group species is rather simple: al-

1) Although 7. calamarius is currently placed with

the fulvipes group, it seems to be an aberrant spe-

cies which may deserve the status of a separate ge-

nus; if it actually belongs to the fulvipes group, it

is a highly specialized species (Graham, pers.

comm.). For this reason we do not take it to repre-

sent the entire fulvipes group.

ternation of distinct high and low phases, and in

the low phase a simple antennal display along-

side the female’s antennae. To some extent it re-

sembles the displays of those males of the muser

group which court from a position on the fe-

male’s thorax (“shoulders”) but in those males

the antennae perform series of knocking

movements, hitting the female’s head, while the

male’s abdomen is raised conspicuously during

the display. Courtship in the brevinervis group,

with only one species observed, however, is like

a simplified version of evonymellae or miser.

There are no up and down movements with the

antennae, but only one upward stroke following

low posturing. The posture of the male’s abdo-

men suggests an affinity with miser. Males of

the daira group exhibit sideways tendencies

which seem to point to the lycidas-caudatus

branch and caudatus males quiver the antennae

in front of the female in a dazra-like fashion but

there is some ambiguity with the display of 7.

daira itself. It was the only daira group species

observed with the male courting on the female’s

thorax, and it exhibits elements in its display

which are suggestive of the miser group, e.g.,

the female antennae point upward and the male

performs series of knocking movements with

his antennae, hitting the female’s head. The

more frontally courting daira group males ex-

hibit a more or less similar feature, but instead

of hitting the female’s head hit, her antennae

from the outside.

Our two categories do not cover entirely the

currently accepted division, i.e., the genus 7e-

trastichus versus the other genera of Tetrastichi-

nae. Judged by behavioural criteria some species

of Tetrastichus belong to our first category (i.e.

the species of the arundinis group, and 7. ca-

lamarius). There are some morphological fea-

tures which also point this way. The fulvipes

group may have affinities to Melittobia Westw.

For example, the inflated scapes of males of

some species suggest such relation. 7. calamari-

us has a peculiar, flattened habitus which is very

suggestive of Melittobia species, as does the

male’s position during courtship. Most species

of the arundinis group can be distinguished

from the other groups of Tetrastichus but mor-

phological criteria were insufficient to separate

them all. With the addition of behavioural crite-

ria as we now know them, we would suggest to

move this species group, and 7. calamarius (and

perhaps the entire fulwipes group) out of Tetra-

stichus s.s.

Some kind of dichotomy in our second cat-

VAN DEN AssEM, Gijswijr & NUBEL: Courtship and mating behaviour 215

egory is also indicated on morphological crite-

ria. The groups miser and brevinervis are char-

acterized by having only one single bristle on

the subcostal vein of the fore wings (and as such

constituted the former genus Tetrastichus). The

groups evonymellae and daira (and some

others) have the pygostylar setae of nearly equal

length; the groups lycidas and caudatus (and

some others) have one bristle far longer than the

others. These two clusters have been recognized

as separate species groups by Graham (1961)

and Domenichini (1965). Whether or not one or

more of these clusters are to be regarded as sep-

arate genera or subgenera is not a question to be

settled on our present evidence. More could be

said, no doubt, with more observations on a

greater variety of species.

Our conclusion on the available evidence is

that within species groups, species have many

features of their courtship display in common.

Some groups are much more similar to each

other than other groups. Further, there seems to

be a parallel to similarities and differences of

certain morphological features, features be-

lieved to be unrelated to courtship movements.

Evolution of patterns of courtship behaviour in

the Tetrastichinae

In the preceding section we have pointed out

differences and conformities: certain groups re-

semble each other much more than others.

However, mere resemblances do not guarantee

a close relationship. In this section we are going

to make a few suggestions on relationships be-

tween groups judged by shared characteristics

which are to be understood as modifications of

more original conditions. Groups not showing

such modifications would qualify as being more

primitive in that respect, i.e., having retained a

more original condition.

Of course, our observations cover only a

minute fraction of the Tetrastichinae known to

exist. Therefore, we present our data for what

they are worth. However, we believe that our

data indicate several transformations from more

original to more modified situations which have

occurred in the evolutionary history of

courtship of Tetrastichinae.

The performances of category I males are of a

considerable complexity and it seems very diffi-

cult to label them as original because of this

complexity. They rather appear to represent

modifications; most obviously this is true for

those species where holding the female’s anten-

nae for long periods goes together with the

presence of peculiar morphological structures

serving such a function, e.g., Melittobia, where

the male antennae turn out to be effective clasp-

ers. Males of all species which practise clasping

or embracing or contacting the female’s mouth-

parts, court from a frontal position, i.e., on the

female’s head or neck. In these species the fe-

male holds her antennae in a forward position

or she folds them in front of her head.

Neither frontal position of the male nor a for-

ward position of the female’s antennae are tied

to grasping or embracing (e.g., several species of

the group dazra, or T. lasiocera group breviner-

vis). Placement of the male’s fore tarsi and posi-

tion of the female’s antennae seem to be inde-

pendent to some degree but two combinations

occurred most frequently: tarsi on head/anten-

nae forward or folded, and tarsi on

“shoulders”/antennae upright. The first combi-

nation is most likely to represent a derived con-

dition, as we pointed out above. It follows, on

the assumption that both categories are genu-

inely related, that the second combination qual-

ifies as a more original condition. The alterna-

tive possibility of being another specialisation is

unlikely: an upright position of female antennae

during courtship appears to be a widespread

phenomenon, occurring in diverse groups.

Moreover, we also observed it in Elachertinae, a

group of Eulophidae believed to be near the ori-

gin of Tetrastichinae. The postures of female

antennae are illustrated in fig. 13.

It should be noted that we found the female’s

antennal posture to vary within one group (dai-

J oo

Fig. 13. Female antennal posture during low phase in

different groups. Further explanations in the text.

216 TijpscHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

ra) and the placement of the male’s fore tarsi in

at least three (lycidas, daira, miser). There are

no good reasons for splitting these groups on

this basis as we feel transitions have occurred in

separate groups independently a number of

times.

A frontal placement of the male seems to be

derived from a more caudally located place-

ment. Earlier we have reached this conclusion

with Pteromalidae where the original position

for courting is the same as the position for mat-

ing. Based upon Van den Assem (1976), we

have arguments to assume that in this group the

switch of courtship position of the male from

the rear to the front, the development of an an-

tennal receptivity signal by the female, and the

reduction of male size constitute a complex of

derived characters. In Eulophidae a parallel de-

velopment seems to have taken place although

we did not find truly caudal courters. An inter-

esting point in this respect is that the antennal

posture indicative of sexual receptivity of fe-

male Tetrastichinae is the reverse of the anten-

nal signal of Pteromalinae, ı.e., antennae

stretched and upright in Tetrastichinae versus

antennae bent down and drawn to the head in

Pteromalinae. Reduction of male size is a com-

mon phenomenon in many Chalcidoidea in-

cluding Tetrastichinae. Within the Tetrastichi-

nae Melittobia are a notable exception. How-

ever, taking account of the reproductive

strategy of Melittobia, this exception is at once

clear enough (Van den Assem, Gijswijt &

Nübel, 1980).

A few additional evolutionary changes may

be pointed out (see table 1 for a summary of our

ideas). In many species or species groups there

is a regular alternation of low and high-phase

episodes, in others there is not. Species of our

category I may serve as example of this latter

condition. In some category I species a regular

alternation occurred (Crataepus marbis, Tetras-

tichus calamarius, species of the arundinis

group), in some alternations were only observed

in the first part of a sequence of courtship, the

high phase being dropped later (several species

of Melittobia) whilst in some there never was a

true alternation at all, males being continuously

in a low-phase position (Nesolynx albiclavus,

several species of Melittobia). In contrast, most —

of the category II males exhibited distinct alter-

nation of high and low-phase episodes over the

entire sequence of courtship. We believe that

absence loss of high-phase episodes can be un-

derstood as a derived condition.

Another development has to do with the tim-

ing of alternations. In some species the dura-

tions of high and low phases vary, most often

the high phases have no fixed duration (e.g., Te-

trastichus spec. 9), in others the timing of alter-

Table 1. Evolutionary modifications of courtship behaviour of Tetrastichinae.

More original condition

alternation of high and low

phases

alternation between more caudal

and more frontal positions of

3 only slight

timing of alternations variable

patterns of & antennation as

rubbing or lashing 9 antennae

or head

display without leg movements

3 display a repetition of

similar elements

events tied to female’s sexual

receptivity occur at intervals

derived condition

high phases dropped underway

high phases never occur

alternation between more caudal

and more frontal positions of d

far more extreme

timing of alternations stereotyped

d embraces ® antennae

d antennae (claval parts) make

contact with ® mouth parts

® antennae pointing to the front

® antennae folded in front of head

6 front tarsi on £ neck

6 front tarsi on £ head

display includes ritualized leg

movements

new elements come in underway

female receptivity occurs at only

one single point of the male’s display

VAN DEN AssEM, Gijswijr & NÜBEL: Courtship and mating behaviour 217

nations has become completely rigid (e.g., 7. at-

riclavus, T. arundinis). The latter condition is

most probably to be understood as a derived

: development towards a high degree of

en is found in many groups, in fact it

appears to be a very general phenomenon.

Yet another development applies to the onset

of female sexual receptivity. Usually only virgin

females will respond to courtship by a conspeci-

fic male. In those species where the male’s dis-

play is a repetition of similar elements, female

receptivity usually sets in at fixed points (e.g., at

the end of an antennal lashing series in 7. dazra;

at the end of an antennal embrace in Nesolynx).

In Melittobia (except for the most primitive

species, Van den Assem et al., in prep.) female

receptivity occurs at only one single point in

time which is immediately following the male’s

“finale” performance (Van den Assem, 1975).

Lastly we draw attention to leg movements as

part of the courtship ritual. Movements of many

types with any pair of legs may be performed,

e.g., there are drumming the fore legs on the fe-

male’s head or eyes, hammering middle legs on

the female’s thoracic pleura and swaying middle

or hind legs back and forth, sometimes reaching

the female’s head or beyond. Bristles of many

kinds are placed on all parts of legs and they

may provide an array of stimuli to different

parts. Leg movements of some kind are includ-

ed in displays of almost all category I males

whereas they appear to be very rare in the cat-

egory II group; 7. epicharmus is such a rarity.

Most probably leg movements constitute later

modifications of the courtship ritual in the Te-

trastichinae.

Above we have pointed out a number of tran-

sitions which we believe to have taken place in

the evolutionary history of the Tetrastichinae.

Our ideas on this point are summarized in table

1. Many of the characters which we understand

as modifications occur in the displays of catego-

ry I males. As these characters are shared by the

whole group, they render it a certain cohesion

which seems to point to common ancestry. Such

a cohesion does not seem to exist in category II.

Perhaps the groups included here are not closely

related, although they share several features.

These features which include the following, are

likely to be of a more primitive character than

those of category I: male courtship position

more to the rear (his front tarsi placed on the fe-

male’s thorax); distinct alternation of high and

low phases, and duration of high phase episodes

variable; female’s antennae upright throughout

the male’s display, male’s antennae upright in

the high phase and moving up and down along

the female’s antennae in the low phase; male

tarsi remain in a fixed position throughout.

‘According to our views the group evonymellae

yields a picture of a rather generalized, original

pattern of tetrastichine courtship behaviour.

Domenichini (1965) placed the group strobi-

lanae (= lycidas + caudatus) at the base of his

reconstruction of the phylogeny of the Tetrastı-

chus complex because of what he called its mor-

phologically non-specialized nature. Courtship

characteristics of this group do not seem to sup-

port such a conclusion without reservations, but

they do seem to support another. Domenichini

derived, among others, the groups viridimaca-

latus (= arundinis) and fulvipes from its

branches. It is worth mentioning that some sim-

ilar conclusion might be drawn from a compari-

son of courtship patterns (with many reserva-

tions!). 7. spec. 6 (group Lycidas) males perform

antennal movements which resemble grasping

movements to some extent; 7. dotus (group

caudatus) males align their antennae along the

female’s for a moment and the female’s claval

parts align the (somewhat inflated) scapi of the

male. Both these displays are suggestive of inci-

pient stages of behaviour patterns of category I

displays.

As has been stated above, our observations

cover a limited number of species. Moreover,

our sample appears to be heavily biased: the

majority of our species are parasitic on gall

midges. As a rule, such a state of affairs is highly

unfortunate but apparently it is not for a com-

parative study of courtship behaviour. Other

kinds of behaviour (e.g., parasitisation behav-

iour) are much more prone to host influences.

No doubt this is the reason why Domenichini

was unable to find such separating features

when he tried to find behavioural characteristics

for separating Tetrastichus species-groups and

based his observations on parasitisation behav-

iour and preferences for certain types of hosts.

He came across a number of very striking con-

vergencies in species from different groups par-

asitising similar host species. Our observations

confirm that patterns of courtship behaviour are

highly independent of host characteristics, a re-

sult which did not surprise us.

Male courtship displays have presumably

evolved as the (to the males) most effective way

to overcome female “coyness”. It has led to the

production of highly specific stimuli. Taxonom-

ists can be helped considerably when trying to

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

solve classificatory riddles by taking notice of

the insects’ own advertisements, especially in

cases of clusters of sympatric, sibling species,

which is commonplace in Chalcidoidea. There

can be no doubt about the role which behaviou-

ral characters (courtship characteristics in par-

ticular) are going to play in future taxonomic

work. We have results which indicate that etho-

logical, isolating mechanisms (e.g., sex phero-

mones) may be already fully effective before en-

tities can be separated on morphological fea-

tures. In a forthcoming paper on Melittobia we

will return to this point and provide a few ex-

amples. If such a situation would turn out to be

a general phenomenon in parasıtic Hymeno-

ptera — and we believe it will — behaviour

studies will even become indispensable. For the

time being there can be no doubt about the val-

ue of behavioural characters as supplementary

to morphological data.

ACKNOWLEDGEMENTS

A number of colleagues sent us living speci-

mens for our observations. Their help is greatly

appreciated. All material received has been list-

ed in the Appendix. We are greatly indebted to

Dr M. W. R. de V. Graham (Oxford) for identi-

fications and for discussions, both at Leiden and

Oxford. Drs Graham and E. C. Dahms (Bris-

bane) commented on a manuscript draft of the

paper and helped to improve the final version.

The investigations were supported by the Foun-

dation for Fundamental Biological Research

(BION), which is subsidized by the Nether-

lands Organisation for the Advancement of

Pure Research (ZWO).

REFERENCES

Assem, J. van den, 1974. Male courtship patterns and

female receptivity signal of Pteromalinae, with

consideration of some evolutionary trends and a

comment on the taxonomic position of Pachycre-

poideus vindemiae. — Neth. J. Zool. 24: 253—

278.

, 1975. Temporal patterning of courtship behav-

iour in some parasitic Hymenoptera, with special

reference to Melittobia acasta. — J. Entom. (A)

50: 137—146.

——, 1976. Male courtship behaviour, female recepti-

vity signal, and size differences between the sexes

in Pteromalinae, and comparative notes on other

chalcidoids. — Neth. J. Zool. 26: 535—548.

, M. J. Gijswijt & B. K. Nübel, 1980. Observa-

tions on courtship- and mating strategies in a few

species of parasitic wasps (Chalcidoidea). — Neth.

J. Zool. 30: 208—227.

, & G. D. E. Povel, 1973. Courtship behaviour of

some Muscidifurax species (Hym., Pteromalidae):

a possible example of a recently evolved ethologi-

cal isolating mechanism. — Neth. J. Zool. 23:

465—487.

——, & F. A. Putters, 1980. Patterns of sound pro-

duced by courting chalcidoid males and its biolog-

ical significance. — Ent. exp. & appl. 27: 293—

302.

Boutek, Z., 1977. Descriptions of Tachinobia gen. n.

and three new species of Tetrastichinae (Hym.,

Eulophidae), with a tentative key to genera. —

Bull. ent. Res. 67: 17—30.

Dahms, E. C., 1973. The courtship behaviour of Me-

littobia australica Girault. — Mem. Qd Mus. 11:

411414.

Domenichini, G., 1965. I Tetrastichini (Hym., Eu-

lophidae) paleartici ed i loro ospiti. — Boll. Zool.

agr. Bachicolt. (II) 6: 61—205.

Graham, M. W. R. de V., 1961. The genus Aprostoce-

tus Hal. s.l., notes on the synonymy of European

species. — Ent. Month. Mag. 97: 34—64.

Miller, M. C., 1966. Emergence and mating of 7. in-

certus, a parasite of the alfalfa weevil. — J. econ.

Entom. 59: 1532.

VAN DEN Assem, Gijswijr & NÜBEL: Courtship and mating behaviour 219

APPENDIX

LIST OF SPECIES MENTIONED IN THE TEXT AND ORIGIN OF OUR MATERIAL

Species marked (*) have been observed in courtship.

Material has been deposited in the collections of M. J.

Gijswijt (G) and Institute for Taxonomic Zoology,

Univ. of Amsterdam, The Netherlands (ITZA).

mine of Phyllonorycter sp. (Lep.) on Fagus sylvati-

cus, Hortus Botanicus Leiden, The Netherlands,

19769,

* Tetrastichus epicharmus (Walker), group caudatus;

* Tetrastichus abydenus (Walker), group Iycıdas;

from galls of Rhabdophaga terminalis (Dipt.) on

Salix sp., Voorschoten, The Netherlands, viii.1978,

7 NOS (CAMSNE

* Tetrastichus arundinis Giraud, group arundinis;

from puparia of Diptera on Phragmitis australis,

Kroonpolders, Vlieland, The Netherlands, 111.1977,

coll. J. H. Mook, fair numbers (ITZA).

* Tetrastichus asparagi Crawford, group miser; from

eggs and larvae of Crioceris asparagi (Col.) on As-

paragus officinalis, Meijendel, Wassenaar, The

Netherlands, vi.1975, ıx.1978, coll. J. van Alphen,

10d 169 (G, ITZA). Geysteren, The Netherlands,

vii.1976, 16 49 (G).

* Tetrastichus atriclavus Waterston, group miser;

from pupae of Acigona ignefusalis (Lep.), via

IRAT, Montpellier, France, from Sénégal, West Af-

rica, x11.1978, leg. J. Brenière, several hundreds, sex

mmol 32 (VANS

Tetrastichus brevicornis (Panzer), group brevicor-

nis; from seed pods of Sarothamnus scoparius,

Bentheim, Westph., Germany, iv.1977; Beilen, The

Netherlands, ix.1977, 22 (G).

* Tetrastichus calamarius (Graham), group fulvipes;

from galls of Giraudiella inclusa (Dipt.) on Phrag-

mitis australis, Leiden, The Netherlands, 11.1974,

Kroonpolders, Vlieland, The Netherlands, iv.1977,

coll. J. H. Mook, large numbers, many more fe-

males than males (G, ITZA).

* Tetrastichus clavicornis (Zetterstedt), species sola;

from catkins with Semudobia spp. (Dipt.) on Betula

sp., Meijendel, Wassenaar, The Netherlands,

11.1978, 223 369; Oberstdorf, Allg., Germany,

x11.1977, 86 62 (G).

* Tetrastichus daira (Walker), group daira; from

flower heads of Cynara cardunculus, Ronda, Anda-

lusia, Spain, ix.1978, 110 specimens, 10—20% d

(ITZA).

* Tetrastichus diaphantus (Walker), group daira;

from galls of Biorhiza pallida (Hym.) on Quercus

pubescens, M. Ventoux, Dep. Vaucluse, France,

vu.1978, 166 609 (G, ITZA).

* Tetrastichus dotus (Walker), group caudatus; from

galls of Dasineura ulmariae (Dipt.) on Filipendula

ulmaria, Voorschoten, The Netherlands, vii.1976,

v.1980, 38 159 (G, ITZA).

Tetrastichus ecus (Walker), group ecus; from leaf

from seeds of Papaver dubium, St. Auban s.

Ouvèze, Dep. Drôme, France, viii.1978, 146 119

(GS

Tetrastichus eriophyes Taylor, group lycidas; from

galls of Phytoptus avellanae (Acar.) on Corylus

avellana, Brochterbeck, Westph., Germany,

v.1976, thelytokous (G).

Tetrastichus escherichi (Szelényi), group lycidas;

from catkins with Semudobia spp. (Dipt.) on Betula

sp., Meijendel, Wassenaar, The Netherlands,

v.1976, 12 (G).

* Tetrastichus evonymellae (Bouché), group evony-

mellae; from pupae of Yponomeuta padellus (Lep.)

on Crataegus monogyna, Vogelenzang, The Neth-

erlands, vu.1974; on Prunus padus, Leiden, The

Netherlands, vi.1976, coll. J. de Groot, fair num-

bers.

* Tetrastichus galactopus (Ratzeburg), group evony-

mellae; hyperparasite of a braconid in a caterpillar

of Pieris brassicae, Wageningen, The Netherlands,

ix.1973, leg. K. W. R. Zwart, fair numbers (G).

* Tetrastichus gratus Giraud, group arundinis; from

puparia of Diptera on Phragmitis australis, Kroon-

polders, Vlieland, The Netherlands, 111.1977, coll. J.

H. Mook, fair numbers (G).

* Tetrastichus hylotomarum (Bouché), group miser,

from pupae of Arche rosae (Hym.) on Rosa sp. (cul-

uvar), Schoonebeek, The Netherlands, ix.1976,

coll. A. van Frankenhuyzen, 148 199 (G).

* Tetrastichus lasiocera Graham, group brevinervis;

from galls of Perrisia persicariae (Dipt.) on Polygo-

num amphibium, Oegstgeest, The Netherlands,

x.1978, 28 99 (G).

Tetrastichus legionarius Giraud, group miser; from

puparia of Lipara lucens (Dipt.) on Phragmitis aus-

tralis, Oostelijk Flevoland, The Netherlands,

vi.1978, coll. J. H. Mook; 34 specimens.

Tetrastichus lycidas (Walker), group lycidas; from:

galls of Hartigiola annulipes (Dipt.) on Fagus sylva-

tica, ’s-Graveland, The Netherlands, ix.1977; 2d

29 (G).

Tetrastichus lysippe (Walker), species sola; from

galls of Dasineura crataegi (Dipt.) on Crataegus

monogyna, ‘’s-Graveland, The Netherlands,

vii.1976, 36 209 (G).

* Tetrastichus pallipes (Dalman), group lycidas; from

220

galls of Semudobia sp. (Dipt.) on Betula sp., Mei-

jendel, Wassenaar, The Netherlands, v.1976, few

specimens; Oberstdorf, Allg., Germany, ix.1977,

1d 39 (G).

Tetrastichus spartu (Ratzeburg), group lycidas;

from seed pods of Sarothamnus scoparıns, Bent-

heim, Westph., Germany, iv.1977, 19 (G).

Tetrastichus strobilanae (Ratzeburg), group lycidas;

from cones of Picea abies, Oberstdorf, Allg., Ger-

Many 141977,x11.19785368, 22:38 92(G)!

The following species of Tetrastichus are new spe-

cies which will be described by Dr. M. W. R. de Vere

Graham (Oxford).

Tetrastichus spec. 1, group brevinervis; from seed

pods of Papaver dubium, Entrechaud, Dep. Vau-

cluse, France, vii.1978, 109.

Tetrastichus spec. 2, group caudatus; on Verbascum

sp., Serrania de Ronda, Andalusia, Spain, ix.1978,

29,

* Tetrastichus spec. 3, group daira; from seed pods

Papaver dubium, Dep. Vaucluse, France, summer

1975.

* Tetrastichus spec. 4, group daira; from flowerheads

of Centaurea sp., Oberstdorf, Allg., Germany,

77590 BSL.

* Tetrastichus spec. 5, group lycidas; from Foenicu-

lum vulgare, Carratraca, Andalusia, Spain, ix.1978,

26 49.

* Tetrastichus spec. 6, group lycidas; from galls of

Plagiotrochus sp. on Quercus ilex, Alhaurin de la

Torre, Andalusia, Spain, 11.1979, 18 19.

Tetrastichus spec. 7, group lycidas; from gall on top

of twigs on Salix sp., St. Auban s. Ouveze, Dep.

Drôme, France, vi1.1978, 19.

Tetrastichus spec. 81), group lycidas; from galls of

Dasyneura epilobu (Dipt.) on Chamaemerion an-

1) 7.8 and 7. 11 may be the same species.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 125, AFL. 7 (1982)

gustifolium, _ Voorschoten, The Netherlands,

viii.1978, few specimens.

* Tetrastichus spec. 9, group arundinis; from Phrag-

mitis australis, Kroonpolders, Vlieland, The Neth-

erlands, 111.1977, coll. J. H. Mook, few specimens.

Tetrastichus spec. 10, group ?; from galls of Rho-

dites spinosissimae on Rosa pimpinellifolia, Bloe-

mendaal, The Netherlands, ix.1977, 16 112.

* Tetrastichus spec. 11!), group ?; from galls of Dasi-

neura epilobii on Chamaenerion angustifolium, ’s-

Graveland, The Netherlands, vi11.1976, 18 269.

* Tetrastichus spec. 12, group miser; from eggs of

Crioceris duodecimpunctata on Asparagus officina-

lis, Meijendel, Wassenaar, The Netherlands, coll. J.

van Alphen, few specimens.

* Crataepus marbis (Walker); from De of

Carduus sp., Malaucene, Mt. Serain, Dep. Vau-

cluse, France, viii.1978; idem of Cynara carduncu-

lus, Ronda, Andalusia, Spain, ix.1978, numerous

specimens, sex ratio ca. 1 : 1.

Hyperteles elongatus (Foerster); from galls of Mi-

kiola fagi on Fagus sp., Bentheim, Westph., Ger-

many, iv.1977, 18 19; Oberstdorf, Allg., Ger-

many, 1x.1977, 26 49.

Hyperteles luteus (Ratzeburg); from galls of Mikio-

la fagi on Fagus sp., Bentheim, Westph., Germany,

WM WW 29,

* Melittobia acasta (Walker), parasite of Odynerus

spinipes (Hym.), Losser, The Netherlands, vi.1974,

coll. G. A. Bekke; parasite of Omalis aureus

(Hym.) and Ancistrocerus sp. (Hym.), Ede, The

Netherlands, vi.1976, coll. R. Leys; ca. 5% d in

lab. cultures.

* Nesolynx albiclavus (Kerrich), from puparia of

Glossina sp. (Dipt.), obtained via Lab. Exp. Ento-

mology Amsterdam, from Salisbury, Rhodesia, leg.

G. A. Vale, iv. 1977, very skewed sex ratio.

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TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

* Een sterretje duidt aan een naam nıeuw voor de wetenschap

* An asterisk denotes a name new to science

ACARINA

avellanae, Phytoptus 219

COLEOPTERA

asparagi, Crioceris 219

duodecimpunctata, Crioceris

[220

DIPTERA

albimanus, Platycheirus 31, 34

angustatus, Platycheirus 31

annulipes, Hartigiola 219

annulipes, Melanostoma 34

crataegi, Dasineura 219

clypeatus, Platycheirus 30, 33

epilobii, Dasyneura 220

fagi, Mikiola 220

fasciatum, Melanostoma 25, 30

fulviventris, Platycheirus 32, 33

Glossina 220

granditarsa, Pyrophaena 25, 32

immaculatus, Platycheirus 25

(recte: immarginatus!)

immarginatus, Platycheirus 32

inclusa, Giraudiella 219

lucens, Lipara 219

marginatus, Mesograpta 25

mellinum, Melanostoma 28, 33

peltatus, Platycheirus 32

perpallidus, Platycheirus 33

persicariae, Perrisia 219

scalare, Melanostoma 25, 29

scambus, Platycheirus 32

scutatus, Platycheirus 27, 31, 34

Semudobia 219, 220

terminalis, Rhabdophaga 219

ulmariae, Dasineura 219

HETEROPTERA

alluaudi, Hebrus 7, 10

“bimaculatus, Hebrus 1, 7, 10

*bongaensis, Hebrometra 1, 15,

[16

caeruleis (coeruleis), Hebrus 10

campestris, Hebrus 10

gerardi, Hebrus 5, 7

*gidshaensis, Hebrus 1, 7, 12

*Hebrometra 1, 15 sqq

Hebrus 1 sqq

Hebrusella 3

Heterocleptis 3

houti, Hebrus 3

Hyrcanus 1, 3

jeanneli, Hebrus 7

jihafana, Saldula 15

katompei, Hebrus 7, 9, 12

Madeovelia 3

*malawiensis, Hebrometra 1, 18

mancinii, Hebrus 7, 16

Merragata 1, 3

Microvelia 15

mizae, Hebrus 12

*niemeri, Hebrometra 1, 17, 18

Paratimasiellus 3

Paratimasius 3

pelengei, Hebrometra 1, 7, 16

*pseudopusillus, Hebrus 1, 10

pusillus, Hebrus 3, 5, 8, 12

ruficeps, Hebrus 3, 5, 7

Saldula 15

somaliensis, Hebrus violaceus

[ssp. 7, 10

soudani, Hebrus 10

spanganii, Hebrus mancinii f. 17

*spinitibialis, Hebrus 1, 7, 9, 12,

[15

Timasielloides 3

Timasiellus 1

Timasius 3

violaceus, Hebrus 10

wygodzinskyi, Hebrus 7

HYMENOPTERA

abydenus, Tetrastichus 210, 219

acasta, Melittobia 213, 220

albiclavus, Nesolynx 213, 214,

[216, 220

Ancistrocerus 220

arundinis, Tetrastichus 207, 219

asparagi, Tetrastichus 206, 212,

[219

atriclavus, Tetrastichus 212, 219

aureus, Omalis 220

baueri, Ophion 83

brevicornis, Ophion 70, 86, 87

brevicornis, Tetrastichus 219

brevinervis, Tetrastichus 207

calcaratus, Ophion 77

calamarius, Tetrastichus 210,

[214, 216, 219

clavicornis, Tetrastichus 212,

[219

“crassicornis, Ophion 70, 84

costatus, Ophion 74, 86, 93

daira, Tetrastichus 208, 216,

[217,219

diaphantus, Tetrastichus 208,

[219

distans, Ophion 77

dotus, Tetrastichus 208, 217,

[219

ecus, Tetrastichus 219

elongatus, Hyperteles 220

epicharmus, Tetrastichus 208,

[217,219

eriophyes, Tetrastichus 219

escherichi, Tetrastichus 219

evonymellae, Tetrastichus 209,

[219

forticornis, Ophion 67, 83

fulvipes, Tetrastichus 210

galactopus, Tetrastichus 209,

[219

gratus, Tetrastichus 207, 219

hylotomarum, Tetrastichus

[212,219

impressus, Ophion 75

lasiocera, Tetrastichus 207, 215,

[219

legionarius, Tetrastichus 219

longicornis, Ophion 76

longigena, Ophion 70, 86

luteus, Hyperteles 220

luteus, Ophion 67, 77

222

lycidas, Tetrastichus 216, 219

lysippe, Tetrastichus 219

marbis, Crataepus 212, 214,

[216, 220

minutus, Ophion 63, 74

miser, Tetrastichus 216

mocsaryi, Ophion 74, 92

obscuratus, Ophion 70, 79

obscurus, Ophion 79

pallida, Biorhiza 219

pallipes, Tetrastichus 210, 219

parvulus, Ophion 71, 74, 90

“perkinsi, Ophion 70, 71, 87

Plagiotrochus 220

pteridis, Ophion 71, 88

rosae, Arche 219

scutellaris, Ophion 63, 76, 88

slaviceki, Ophion 77

spartii, Tetrastichus 220

spinipes, Odynerus 220

spinosissimae, Rhodites 220

stigmaticus, Ophion 76

strobilanae, Tetrastichus 210,

[220

ventricosus, Ophion 63, 75

LEPIDOPTERA

aceris, Acronicta 96

Acronicta 91

Agrochola 83

agathina, Amathes 83, 96

albida, Ipthima 100, 101, 104,

[143

albida occidentalis, Ipthima

[102

albida uniformis, Ipthima 102,

[144

ambusta, Atethmia 93, 96

antennata, Ipthima 100, 101,

[102, 103, 118

*antennata cornesi, Ipthima 120

Apamea 82

asterope, Ipthima 100, 101,

[102, 103, 104

asterope asterope, Ipthima 101,

[103

asterope hereroica, Ipthima 101,

[103

augur, (Graphiphora) 83, 96

aurantiaria, Agriopis 75, 95

baja, (Xestia) 83, 96

baldus, Iphtima 145

bicruris, (Hadena) 90, 96

brassicae, Mamestra 90

brassicae, Pieris 219

capsincola, (Hadena) 89

castanea, Xestia 96

chamomillae, Cucullia 86, 96

comes, Triphaena 82, 96

condamini, Ipthima 100, 102,

[104, 115

*condamini nigeriae, Iphtima 116

congoana, Ipthima 102, 103,109

coryli, Colocasia 96

cruda, Orthosia 96

cuspis, Acronicta 96

defoliaria, Erannis 75

diplommata, Ipthima 102, 104,

[142

doleta, Ipthima 101, 102, 104,

[141

Erannis 75

fasciata, Ipthima 145

ferrago, Mythimna 96

fimbriata, Noctua 77, 83, 92, 96

gothica, Orthosia 91, 96

granulosa, Ipthima 103, 126

icteritia, Xanthia 93

ignefusalis, Acigona 219

impura, Ipthima 101, 103, 104,

[130, 135

impura paupera, Ipthima 103,

TIJDSCHRIFT voor ENTOMOLOGIE, DEEL 125, 1982

unanimis, Apamea 96

upemba, Mashuna 145

varia, Lycophotia 82

verbasci, Cucullia 94, 96

*vuattouxi, Ipthima 102, 104, 121

yatta, Ipthıma 103, 108

Xanthia 83

xanthographa, (Xestia) 77, 96

ODONATA

Agriogomphus 38

Archaeogomphus 37 sqq

densus, Archaeogomphus 37,

[42, 45, 48

Epigomphus 38

furcatus, Archaeogomphus 41,

[42, 44,49

hamatus, Archaeogomphus 41,

[42, 44, 48

infans, Archaeogomphus 37, 41,

[42, 48

nanus, Archaeogomphus 41, 42,

[45, 49

[137 ORTHOPTERA

impura, Mythimna 82, 83, 96

interjecta, Noctua 83, 96

jacksoni, Ipthima 104

*lamto, Ypthima 103, 104, 123

leporina, Acronicta 96

lota, Agrochola 93, 96

lucipara, Euplexia 91, 96

marginaria, Agriopis 95

mashuna, Mashuna 145

myrtilli, Anarta 82

nigra, Aporophila 83, 85, 96

oleracea, Diataraxia 90, 96

padellus, Yponomeuta 219

Phyllonorycter 219

pilosaria, Apocheima 75, 76, 96

pisi, (Ceramica) 89, 90, 96

porphyria, Lycophotia 96

praecox, Ochropleura 79, 84, 96

praestans, Ipthima 102, 104, 139

progemmaria, Agriopis 75

pulchra, Ipthima 102, 104, 138

pupillaris, Ipthima 101, 103,

[104, 130, 133

pupillaris obscurata, Iphtima

[103, 134

recta, Ipthima 103, 104, 124

retusa, Ipimorpha 96

rhodesiana, Ipthima 102, 103,

rivularis, Hadena 96

scrophulariae, Cucullia 96

simplicia, Ipthima 102, 103, 113

trapezina, Cosmia 96

aegaeus, Poecilimon 188

beieri, Poecilimon 162

berlandi, Poecilimon 169

bosphoricus, Poecilimon 191

brevicauda, Poecilimon 188

brunneri, Poecilimon 169

bulgaricus, Poecilimon 177

chopardi, Poecilimon 178

cretensis, Poecilimon 171

deplanatus, Poecilimon 188

distinguendis, Poecilimon 171

ebneri, Poecilimon 174

Eupoecilimon 155

fieberi, Barbitistes 156

geoktschaicus, Poecilimon 191

gracilis, Poecilimon 161

hadjisarandou, Poecilimon 191

hamatus, Poecilimon 171, 190

hoezeli, Poecilimon 158

*ikariensis, Poecilimon 171

jonicus, Poecilimon 165

*jonicus lobulatus, Poecilimon

[167

*klisuriensis, Poecilimon 174

laevissimus, Poecilimon 164

lemnoticus, Poecilimon 169

macedonicus, Poecilimon 168

mavrovi, Poecilimon 161

miramae, Poecilimon 177

mytilensis, Poecilimon 188

nobilis, Poecilimon 162

obesus, Poecilimon 161

orbelicus, Poecilimon 177

TijpscHRIFT voor ENTOMOLOGIE, DEEL 125, 1982

ornatus, Poecilimon 156

pancici, Poecilimon 158

pergamicus, Poecilimon 170

*pindos, Poecilimon 160

Poecilimon 155

propinquus, Poecilimon 181,

[185, 188

sanctipauli, Poecilimon 188

syriacus, Poecilimon 172

tessellatus, Poecilimon 168

thessalicus, Poecilimon 181, 184

thoracicus, Poecilimon 163

veluchianus, Poecilimon 178

walteri, Poecilimon 168

werneri, Poecilimon 167

zimmeri, Poecilimon 181

zwicki, Poecilimon 175

PLANTAE

Asparagus officinalis 219, 220

Betula sp. 219, 220

Carduus 220

Carex 34

Centaurea 220

Chamaenerion angustifolium

[220

Corylus avellana 219

Crataegus monogyna 219

Cynara cardunculus 219, 220

Cyperus 34

Fagus 220

Fagus sylvatica 219

Filupendula ulmaria 219

Foeniculum vulgare 220

Melampyrum 31

Molinia coerulea 33

Papaver dubium 219, 220

Phragmitis australis 219, 220

Picea 29

Picea abies 220

Pinus 29

Plantago coronopus 27

Plantago lanceolata 25 sqq

Plantago major 27

Plantago maritima 27, 34

Plantago media 27, 34

Polygonum amphibium 219

Prunus padus 219

Quercus ilex 220

Quercus pubescens 219

Rhynchospora 33, 34

Rosa 219

Rosa pimpinellifolia 220

Salix 219, 220

Salix repens 93

223

Sarothamnus scoparius 219, 220

Typha angustifolia 25

Typha latifolia 31

Verbascum 220

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