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DEEL 124 1981

TIJDSCHRIFT

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

Tijdschrift voor Entomologie, deel 124, 1981

NEDERLANDSE ENTOMOLOGISCHE VERENIGING

BESTUUR (BOARD)

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The journal serves the publication of papers on Insecta, Myriapoda and Arachnoidea.

Subscription rate: D.Fl. 225,— per year.

Afleveringen 1—5 verschenen 20.VII.1981

Aflevering 6 verscheen 31.XII.1981

ISSN 0040-7496

INHOUD VAN DEEL 124

Aukema, B. — A survey of the Dutch species of the subgenus Hy/opsallus of Psallus (Hemipte-

Ka-Leteropteray Mini dae) aware N eee

| Boucek, Z., A. Watsham & J. T. Wiebes. — The fig wasp fauna of the receptacles of Ficus

thonningi (rymenopteray Chalcidoidea) rr

Chaudhuri, P. K., D. K. Guha & S. K. Das Gupta. — Taxonomic studies of Chironominae

(Diptera, Chironomidae) from India. The genus Polypedilum Kieffer ............

Das Gupta, S. K., zie Chaudhuri, P. K.

Guha, D. K., zie Chaudhuri, P. K.

| Newton, P. J., zie Wilkinson, C.

i Watsham, A., zie Bouéek, Z.

| Wiebes, J. T., zie Boutek, Z.

Wilkinson, C. — A supplement to the genus Ectoedemia Busck (Nepticulidae: Lepidoptera) in

North America, dealing with some difficult species and also some new ones ......

Wilkinson, C., & P. J. Newton. — The microlepidopteran genus Ectoedemia Busck (Nepticuli-

dao)linıN oLth-American SA RR Te

ERRATUM

On p. 168 the title of the new chapter should read Sycoecinae instead of Sycoryctini.

WA En

DEEL 124 AFLEVERING | 1981

TIESCHRITTL a

HARVArıy

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

B. AUKEMA. — A survey of the Dutch species of the subgenus Hy/opsallus of

Psallus (Hemiptera-Heteroptera, Miridae), p. 1—25, figs. 1—36.

Tijdschrift voor Entomologie, deel 124, afl. 1 Gepubliceerd 20-VII-1981

A SURVEY OF THE DUTCH SPECIES OF THE SUBGENUS

HYLOPSALLUS OF PSALLUS (HEMIPTERA-HETEROPTERA,

MIRIDAE)

(Mededeling EIS-Nederland, no. 6)

BEREND AUKEMA

Instituut voor Taxonomische Zoölogie, Amsterdam

With 36 text-figures

ABSTRACT

The Dutch material of the subgenus Psallus (Hylopsallus) and of the related Asthenarius quercus

(Kirschbaum, 1856) is extensively revised. The occurrence of Psallus (Hylopsallus) perrisi (Mulsant,

1852), P. (H.) variabilis (Fallen, 1829) and A. quercus in the Netherlands is confirmed, whereas P. (H.)

wagneri Ossiannilsson, 1953 is recorded now for the first time from this country. It is shown that

measurements of external morphological characters do not allow identification of the species, as is

suggested in most keys of Hylopsallus. Of the Dutch species and P. (H.) assimilis Stichel, 1956, which

may be expected, a key to the males, descriptions, and drawings of the male genitalia are given, and the

distribution of the Dutch species in the Netherlands is figured. Finally the status of P. (H.) wagneri is

discussed.

INTRODUCTION

The genus Psallus Fieber, 1858, belongs to the subfamily Phylinae Handlirsch,

1925, tribe Phylini Douglas & Scott, 1865. The subgenus Hylopsallus was esta-

blished by Wagner in 1952, and in the same publication Psallus variabilis (Fallen,

11829) was designated as the type of the subgenus.

| The Dutch species of Hylopsallus are characterized by the pale antennae (base

lof segment 1 more or less dark), the black, dark-brown, red-brown or red body,

the dark-brown or red femora, which are pale only at their extreme apices and

either unspotted or provided with a few obscure dark spots beneath, the pale tibiae

having dark spines arising from dark spots (at least at hind tibiae), the tarsal

‘segment 3 being shorter than segment 2, the male genital segment, which is

ventrally distinctly keeled, and the shape and processes of the aedeagus.

Wagner (1975) included 10 species in Hylopsallus, of which quercus (Kirsch-

baum, 1856) was transferred to the genus Asthenarius Kerzhner, 1962, in a later

‚publication of the same author (Wagner, 1975a). Four of the nine species consti-

‘tuting the subgenus now are found in the Netherlands or adjacent countries.

Probably because the identification of the species of Hylopsallus is difficult

‘Without examination of the male genitalia, Dutch entomologists have paid little

2 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

attention to the representatives of Hylopsallus. Reclaire (1932, 1948) included

quercus and variabilis in his checklist of Dutch species and Meurer (1956) added

perrisi (Mulsant, 1852).

My unsuccesful attempts to identify some Dutch material of Hylopsallus led to

this critical evaluation of all Dutch material of the subgenus available (cf.

collections mentioned under Acknowledgements). Because Asthenarius quercus

closely resembles Hylopsallus and since this species was often confused with

variabilis by Dutch entomologists I have incorporated this species in the present

study.

This study confirms the occurrence of P. (H.) perrisi, variabilis and quercus in the

Netherlands, whereas wagneri is recorded from this country for the first time.

Another species which could be expected in the Netherlands is P. (H.) assimilis

Stichel, 1956. This species was described after material from England and was later

recorded from West Germany (Rieger, 1975, 1976). Since assimilis was not found

in museum material from the Netherlands, I have undertaken special efforts to

find assimilis on its host plant, common maple (Acer campestre L.). However, all

attempts were unsuccessful.

Furthermore my study of the Dutch Hylopsallus material led to a critical

evaluation of the characters used in the literature for separating the species. The

genitalia of all available Dutch males of Hylopsallus were dissected and examined.

For reasons of comparison some Swedish material of wagneri and some British

material of perrisi, wagneri, variabilis and assimilis was examined. A key to the

males, drawings of the male genitalia and descriptions of the species are included.

TAXONOMIC CHARACTERS

The species of Hylopsallus have a very similar general appearance, although

perrisi and wagneri are somewhat different in size and colour from assimilis and

variabilis.

Absolute as well as relative measurements of external morphological structures

are most frequently used in separating species of Hylopsallus. Especially the ratio

of width of vertex to width of eye, the ratio of length of antennal segment 2 to

length of segment 1, the ratio of length of antennal segment 2 to width of pronotal

base, and the absolute length of the body including the wings are used in keys for

the identification of species in Hylopsallus (Kerzhner & Yachevski, 1964;

Southwood & Leston, 1959; Stichel, 1956, 1958; Wagner, 1952, 1961, 1967 and

1975; Wagner & Weber, 1964; Woodroffe, 1957 and Zaytseva, 1969).

However, precise measurements of great numbers of specimens show that these

measurements do not provide an reliable basis for identification because of the

great intraspecific variation and the great interspecific overlap (tables 1—3). All

the keys mentionned are more or less misleading in suggesting the possibility of

identification on the ground of simple measurements. The ratio of width of vertex

to width of eye for instance is used by Wagner (1952, 1961, 1967 and 1975) and

Woodroffe (1957) to separate perrisi/wagneri, assimilis, variabilis and quercus from

each other. Comparison of their ratios with the ratios from my mearurements

(table 4) shows that their ratios do fall within the range of my measurements. On

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Table 4. Comparison between the ratio of width of vertex to width of eye according to Woodroffe

(1975), Wagner (1975) and the measurements presented in this paper.

Species sex Woodroffe Wagner measured

1957 1975 range average

perrisi 3 ~ 2.0 1.9—2.0 1.54—2.12 1.76

perrisi/wagneri Q ~ 2.0 2.0—2.2 1.70—2.38 1.97

wagneri 3 ~ 2.0 1.9—2.0 1.60—2.00 1.76

assimilis 3 >2.0 — 1.90—2.11 1.98

variabilis d <2.0 1.70—1.75 1.39—2.16 1.78

Q — 1.80—1.85 1.80—2.22 2.04

quercus d ~ 1.3 1.5 1.30— 2.00 1.47

Q AMI 1.75 1.72—2.26 2.00

the other hand identification on species level on account of the ratio of width of

vertex to width of eye proved to be impossible.

Measurements were made according to the method followed by Wagner (1966:

10, 1970: 20—21) with a Zeiss stereo microscope at magnifications of 20 times

(length and width of the body including the wings), 40 times (length of antennae

and antennal segments, length of rostrum and width of pronotal base) and 50 times

(width of head and width of vertex). The ratio of width of vertex to width of eye

was calculated as: width of vertex: 0.5 times (width of head — width of vertex),

whereas width of head and width of vertex were measured as shown in fig. 1.

Fig. 1. Measurements of the head: V = width of vertex; E = width of eye; H = width of head.

The length of the rostrum in relation to the coxae, used by Stichel (1956) as a

distinguishing character, does not provide a reliable character since the extension

of the rostrum depends on the slant of the head, on the position of the coxae and

on the curvation of the abdomen. Neither does the actual length of the rostrum in

relation to the length of the body provide a reliable distinction since a few

measurements already show an overlap (tables 1—3).

Small differences between Hylopsallus species as regards the shape of the claws

have been described (e.g. Wagner, 1952), but these differences are too subtle.

The differences shown by the aedeagi (figs. 23—26) present the only reliable

characters for separating all species of Hylopsallus concerned. The right parameres

(figs. 8—11) are very similar throughout the subgenus, whereas the left parameres

(figs. 13—16, 18—21) and the thecas (figs. 28—31) only enable separation of the

AUKEMA: Dutch species of Hylopsallus 7

imm

Figs. 2—7. Habitus. 2—3. Psallus perrisi, male, female (Amsterdam, the Netherlands); 4—5. P. variabi-

lis, male, female (Amsterdam, the Netherlands); 6—7. Asthenarius quercus, male, female (Heerde, the

Netherlands).

8 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

Figs. 8—12. Right parameres. 8. Psallus perrisi (Amsterdam, the Netherlands); 9. P. wagneri (Heerde,

the Netherlands); 10. P. variabilis (Amsterdam, the Netherlands); 11. P. assimilis (Wychwood, N.B.,

Oxon, England); 12. Asthenarius quercus (Heerde, the Netherlands).

AUKEMA: Dutch species of Hylopsallus 9

Figs. 13—17. Left parameres, SP = sensorial process. 13. Psallus perrisi (Amsterdam, the Netherlands);

14. P. wagneri (Heerde, the Netherlands); 15. P. variabilis (Amsterdam, the Netherlands); 16. P. assimi-

lis (Wychwood, N.B., Oxon, England); 17. Asthenarius quercus (Heerde, the Netherlands).

10 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

Figs. 18—22. Left parameres from opposite view, SP = sensorial process. 18. Psallus perrisi (Amster-

dam, the Netherlands); 19. P. wagneri (Heerde, the Netherlands); 20. P. variabilis (Amsterdam, the

Netherlands): 21. P. assimilis (Wychwood, N.B., Oxon, England); 22. Asthenarius quercus (Heerde, the

Netherlands).

AUKEMA: Dutch species of Hylopsallus

‘(puej3ug ‘uox0 “d°N ‘POOMYIÁM) SYIMISSD ‘4 “CZ ‘(SPUE]I9Y19N 9} ‘apioaH) wausom ‘d ‘pz ‘(SPUE[ISUJON 9} ‘Wepio1swy) 191442d

Snjjosd ‘ET ‘9j919Qm ayepnanusp = LQ ‘ssa501d jesıde = AV ‘ssa901d jeseq = qg ‘210ydouo3 Krepuosos = DS ‘1Beopov ‘S7—€7 SIA

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

ww 1°O

‘(spuejloyJaN OU} ‘apiaoH) sn04anb snıavuayısy

‘LT ‘(SPUEJISYIAN oy) ‘wepiorsww) sıyıgpıava SDS ‘97 ‘SS9901d [eseq = dg ‘a10ydouo3 Arepuooss

DS Beopov LT ‘SUA

AUKEMA: Dutch species of Hylopsallus 13

Figs. 28—32. Thecas. 28. Psallus perrisi (Amsterdam, the Netherlands); 29. P. Wagneri (Heerde, the

Netherlands); 30. P. variabilis (Amsterdam, the Netherlands); 31. P. assimilis (Wychwood, N.B., Oxon,

England); 32. Asthenarius quercus (Heerde, the Netherlands).

14 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

EIER - rn

33 ZA

@ PRE 1950

@ 1960 onwarDs

@ PRE 1960 AND ONWARDS

Seed LI

Figs. 33—34. Maps of Dutch distributions. 33. Psallus perrisi; 34. P. wagneri.

AUKEMA: Dutch species of Hylopsallus 15

@ PRE 1%0

@ 1960 onwarps

@ PRE 1960 an onwarDs

Figs. 35—36. Maps of Dutch distributions. 35. Psallus variabilis; 36. Asthenarius quercus.

16 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

males of perrisi and wagneri from those of assimilis and variabilis.

Females can only be assigned to either perrisi/wagneri or assimilis/variabilis by

the colour of the hemelytra, the colour of the base of antennal segment | and the

colour of the femora. Whether the sclerotized rings of the bursa copulatrix might

present a good character for separating the species is not known since these

structures have not been studied for Hy/opsallus.

A key to the males of Hylopsallus and Asthenarius quercus is presented hereafter.

A key to the females was not constructed because I am unable to separate the

females of wagneri from those of perrisi and the females of assimilis from those of

variabilis. The differential characters for separating the females of perrisi and

wagneri from those of assimilis and variabilis can be read from the descriptions of

the different species. The presented key resembles mostly the key of Woodroffe

(1957) minus the use of ratios.

KEY TO THE MALES

1. Scale pubescence white. Coxae and trochanters greyish white. Aedeagus short

and C-shaped, secondary gonophore close to the apex; apex with a long twisted

denticulate band ending in a long and narrow apical process (fig. 27) quercus’

— Scale pubescence golden. Coxae and trochanters red-brown, dark-brown or

black. Aedeagus differently shaped, with lateral processes; secondary

gonophore distant from the apex (figs, 2326) rr MRS 2

2. Body black-brown to black, only immature specimens lighter. Basal one fourth

of first antennal segment dark-brown to black. Femora red-brown to dark-

brown or black, its extreme apices often yellowish. Sensorial process of left

paramere apically acutely angled (figs. 13—14, 18—19). Aedeagus long and

slender, provided with three lateral processes and a denticulate tubercle near

its apex (figs.232=24), RE ee. NE 3

— Body yellow-brown, red-brown or bright orange-red, sometimes dark-brown.

First antennal segment basally only very small dark. Femora dark-brown, their

apices broadly red to orange-red. Sensorial process of left paramere apically

broadly rounded (figs. 15—16, 20—21). Aedeagus robust, with two lateral

processes (figs: 25-26)" PR EN a LE RE 4

3. Aedeagus with apical process short and hook-shaped, strongly expanded

before apex, its apex just reaching the denticulate tubercle (fig. 23) .. perrisi

— Aedeagus with apical process long and narrow, its apex always reaching far

beyond the denticulate tubercle (fig: 24) Ren wagneri

4. Aedeagus with basal process larger, apically not hooked (fig. 25) .. assimilis

— Aedeagus with basal proces smaller, hooked apically (fig. 26) .... variabilis

SPECIES ACCOUNTS

In the next section the descriptions of the species are preceded by references to

the most important literature and synonyms, and by a list of material examined;

they are followed by remarks on biology and distribution.

AUKEMA: Dutch species of Hylopsallus 17

Psallus (Hylopsallus) perrisi (Mulsant, 1852)

(figs. 2—3, 8, 13, 18, 23, 28, 33)

Mulsant & Rey, 1852: 139—140; Wagner, 1952: 90—94, figs. 1—2; Wagner, 1952a: 176—177, figs.

108, 109a; Stichel, 1956: 287—288, figs. 741—742; Woodroffe, 1957: 258, 260, 262, 267, figs. 1, 23, 43b,

46b; Southwood & Leston, 1959: 221, 224, figs. 76, 85; Wagner, 1967: 145—146, figs. 98F, 99D; Wagner,

1975: 181, 186, figs. 751d, 752b, 753d. — Apocremnus anticus Reuter, 1876: 22; Wagner, 1959: 348— 349.

Material examined. — The Netherlands: 246 specimens from 58 localities (159

3/49 loc., 87 9/29 loc.). Great Britain: 17 specimens from 13 localities (8 3/6 loc.,

9 9/8 loc.), coll. British Museum (Natural History), London.

Description

A rather small-sized species, oval in shape (figs. 2—3), body of males and

females 2.2—2.6 as long as wide. Black-brown to black, only immature specimens

lighter. Pubescence consisting of both shiny golden scales, which are easily rubbed

off, and semi-erect black hairs.

Head. — Dark-brown to black, hind margin of vertex often yellowish-brown.

Width vertex/width eye g: 1.5—2.1, 9: 1.7—2.4. Antennae light yellow-brown,

basal one fourth of segment 1 dark-brown to black; length segment 2/length

segment | g: 3.8—4.7, ©: 4.0—5.0; length segment 2/length segments 3 + 4 &:

0.9—1.3, 9: 0.9—1.2; length segment 2/width pronotal base g: 0.6—0.9, 9:

0.7—0.9. Rostrum dark-brown to black, segments 2 and 3 often yellow-brown and

base of segment 1 ventrally reddish; length 2: 1.2—1.4 mm, 9: 1.3—1.5 mm.

Thorax. — Pronotum dark-brown to black. Mesoscutum dark-brown, outer

margins often with a reddish tinge. Scutellum dark-brown, sometimes yellow-

brown. Hemelytra dark-brown to black. Anterior parts of clavus, corium and

embolium more or less brown and tinged with red. Corium near fracture tinged

with red and along fracture sometimes whitish. Cuneus red-brown to dark-brown,

its central part often darker, basally broadly and apically narrowly whitish. Wing

membrane dark-grey with a clear spot just behind outer cell; base and central part

sometimes colourless too; veins colourless, sometimes tinged with light-yellow-

brown. In females basal part of hemelytra usually totally yellow-brown, sharply

contrasting with the darker pronotum and head. Femora red-brown to dark-

brown, or black, their extreme apices often yellowish. Hind femora sometimes

with a few obscure dark spots beneath. Tibiae yellow, with black spines which

arise especially on hind tibiae from dark-brown to black spots. Tarsi yellow,

segment 3 dark. Claws yellow-brown, slender; pseudarolia very small.

Ventral surface. — Dark-brown to black, but sometimes more reddish-brown,

especially in females. Margins of propleura, sternum and episternum, basalar

plate, thoracic spiracle and hind margin of ostiolar peritreme yellowish-white,

sometimes tinged with red.

Male genitalia. — Genital segment of male with distinct ventral keel. Right

paramere small, oblong (fig. 8). Left paramere small, with sensorial process

acutely angled apically (figs. 13, 18). Aedeagus long and slender, its apical process

18 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

short, hook-shaped and strongly expanded before the apex, just reaching the

denticulate tubercle (fig. 23). Theca small, triangular and more or less equally

tapering apically, with a small elevation just before the apex (fig. 28).

Length of body &: 3.1—4.0 mm, 9: 3.0—4.1 mm.

Biology. — In the Netherlands very common on Quercus, especially on Quercus

robur L. Most records from other plants are probably due to stray specimens.

Probably phytophagous as well as zoophagous. Imagines are found from mid-May

until early August. There is one generation a year and the eggs overwinter in

cracks in the bark of year-old oak wood (Southwood & Leston, 1959).

Distribution. — Psallus perrisi is a widespread West Palearctic species, thusfar

recorded from Finland, Sweden, Norway, Denmark, West Germany, England, the

Netherlands (fig. 33), France, Austria, Bulgaria, Greece, South Russia, Asia

Minor, Syria and Palestina (Coulianos & Ossiannilsson, 1976; Josifov, 1970;

Stichel, 1958; Wagner, 1959, and Wagner & Weber, 1964).

Psallus (Hylopsallus) wagneri Ossiannilsson, 1953

(figs. 9, 14, 19, 24, 29, 34)

Ossiannilsson, 1953: 2—3, figs. 1—5; Stichel, 1956: 288; Woodroffe, 1957: 258, 260, 262, 267, figs. 2,

24; Wagner, 1967: 146, figs. 98E, 99E; Wagner, 1975: 182, 186; Zaytseva, 1969: 526, 529— 530, figs.

22— 23,

Material examined. — Holotype: Uppland, Djursholm, Ösbysjön, 20.vi.1943, F.

Ossiannilsson, g, coll. Zoological Institute, University of Lund (Mus. Lund).

Paratypes: Uppland, Solna, 23.vii.1942, F. Ossiannilsson, g, coll. Mus. Lund;

Östergötland, Tjänste, Haglund, &, coll. Swedish Museum (Natural History),

Stockholm; Bohuslän, 15.vi, A. Tullgren, g, and Uppland, Experimentalfältet,

22.vi.1917, A. Tullgren, g, both in the coll. of the Swedish Plant Protection

Institute, Solna; Bohuslän, Ljung, Lyckorna, B. Tjeder, Z, and same data but

24.vi.1946, &, both in the coll. of B. Tjeder. The Netherlands: 39 males from 19

localities. Great Britain: 9 males from 9 localities, coll. British Museum (Natural

History), London. Sweden: 3 males from 2 localities, coll. F. Ossiannilsson.

Description

This species is very similar to perrisi. Females are thusfar indistinguishable from

those of perrisi. Males can only be separated from those of perrisi on account of

the different shape and size of the apical process of the aedeagus. A description of

these distinguishing features of the present species is given below. The

measurements of wagneri are provided, since the range of variation in wagneri

appeared to be somewhat different from those of perrisi.

Males.

Body 2.3—2.8 as long as wide, 3.1—3.9 mm long.

Head. — Width vertex/width eye: 1.5—2.0. Antennae: length segment 2/length

AUKEMA: Dutch species of Hylopsallus 19

segment 1: 3.8—4.8; length segment 2/length segments 3 + 4: 0.9—1.1; length

segment 2/width pronotal base: 0.7—0.9.

Male genitalia. — Right paramere (fig. 9) small and oblong, as in perrisi. Left

paramere (figs. 14, 19) with sensorial process acutely angled apically as in perrisi.

Aedeagus (fig. 24) as in perrisi (fig. 23), but with apical process long and narrow, its

apex always reaching well beyond the denticulate tubercle. Theca (fig. 29) as in

perrisi.

Biology. — In the Netherlands fairly common on Quercus, especially Quercus

robur L. Often together with perrisi imagines are found from mid-May until the end

of July. There is one generation a year and the eggs overwinter.

Distribution. — Psallus wagneri is a West Palearctic species, thusfar recorded

from Finland, Sweden, Denmark, West Germany, the Netherlands (fig. 34), the

Baltic States, Bulgaria and Central and South Russia (Coulianos & Ossiannilsson,

1976; Josifov, 1970; Stichel, 1958; Wagner, 1959; Wagner & Weber, 1964 and

Zaytseva, 1969). Probably a widespread species, which is however often confused

with perrisi and variabilis.

Psallus (Hylopsallus) variabilis (Fallén, 1829)

(figs. 4—5, 10, 15, 20, 26, 30, 35)

Fallen, 1829: 98; Wagner, 1952: 90—94, figs. 1—2; Wagner, 1952a: 177—178, figs. 108, 109b; Stichel,

1956: 289; Stichel, 1958: 794, figs. 744, 973; Woodroffe, 1957: 258, 260, 262, 267, figs. 6, 26; Carvalho,

1958: 33; Southwood & Leston, 1959: 221, 225, figs. 77, 82; Wagner, 1967: 145, figs. 97G, 98C, 99C;

Wagner, 1975: 181, 185, figs. 751c, 752a, 753c.

Material examined. — The Netherlands: 229 specimens from 57 localities (81 &/

34 loc., 148 9/50 loc.). Great Britain: 7 specimens from 5 localities (3 8/2 loc.

4 9/3 loc.), coll. British Museum (Natural History), London.

Description

A small species, oval in shape (figs. 4—5), body of males 2.3—2.8 and females

2.1—2.6 as long as wide. Yellow-brown, red-brown, bright-orange-red, sometimes

dark-brown. Pubescence as in perrisi.

Head. — Dirty yellow-brown to dark-brown, sometimes with a reddish tinge.

Width vertex/width eye g: 1.42.2, 9: 1.8—2.2. Antennae yellow-brown,

segment | basally only very small dark; length segment 2/length segment 1 &:

3.6—4.8, ©: 3.8—4.6; length segment 2/length segments 3 + 4 &, 9: 0.9—1.1;

length segment 2/width pronotal base g, 9: 0.7—0.9. Rostrum yellow-brown,

segments | and 4 more or less dark-brown, and base of segment 1 ventrally whitish

with a red tinge; length g: 1.4—1.5 mm, 9: 1.4—1.6 mm.

Thorax. — Pronotum light-yellow-brown to dark-brown, tinged with black and

sometimes anteriorly and/or posteriorly with dark markings. Pronotum especially

in females sometimes tinged with red or orange-red. Mesoscutum and scutellum

coloured as pronotum; lateral margins of both mesoscutum and scutellum and

20 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

apex of scutellum usually somewhat lighter. Hemelytra light-yellow-brown to

dark-brown, especially near apex more or less tinged with red or orange-red.

Central parts sometimes dark-brown to blackish, especially in males. Cuneus

yellow-brown, red-brown to deep-red, basally more or less white along fracture.

Wing membrane dark-grey to black, with a clear spot just behind the outer cell.

Veins colourless to greyish, usually tinged with red. Femora dark-brown, apically

broadly red to orange-red, sometimes with a few indistinct dark spots beneath.

Tibiae light-yellow with black spines arising from distinct dark spots, especially on

hind tibiae. Tarsi yellow, segment 3 dark apically. Claws yellow-brown, slender;

pseudarolia very small.

Ventral surface. — Light-red-brown to dark-brown, with the same yellowish-

white parts as in perrisi, sometimes with a red tinge.

Male genitalia. — Genital segment of maie with a distinct ventral keel. Right

paramere large and oblong (fig. 10). Left paramere large, with sensorial process

broadly rounded apically (figs. 15, 20). Aedeagus oblong and robust, with two

lateral processes; basal process short and apically hooked (fig. 26). Theca

somewhat larger than in perrisi and wagneri, with a more distinct elevation just

before the apex (fig. 30).

Length of body &: 3.5—4.4 mm, 9: 3.3—4.3 mm.

Biology. — In the Netherlands common on oak, especially on Quercus robur L.

Imagines are found from mid-May until early August. There is one generation a

year and the eggs overwinter.

Distribution. — A widespread West Palearctic species, known from Finland,

Sweden, Norway, the Baltic States, West Russia, Poland, Germany, Denmark,

Scotland, Ireland, England, the Netherlands (fig. 35), Belgium, France, Spain,

Algeria, Italy, Switzerland, Austria, Czechoslovakia, Hungary, Yugoslavia,

Albania, Greece, Bulgaria, Roumania, South Russia and Turkestan (Stichel, 1958;

Zaytseva, 1969).

Psallus (Hylopsallus) assimilis Stichel, 1956

Gigs Whe 1672162553 1)

Stichel, 1956: 289; Woodroffe, 1957: 258, 260, 262, 267, figs. 3, 27; Stichel, 1958: 795, fig. 973; South-

wood & Leston, 1959: 221, 224, fig. 87; Wagner, 1967: 145, figs. 98b, 99F.

Remarks. — Woodroffe (1957) pointed out that Apocremnus simillimus Douglas

& Scott, 1865 (nec Kirschbaum, 1856) represents a good species. Since the

publication of Reuter (1878) simillimus Douglas & Scott, 1865, was considered a

variety of Psallus (Hylopsallus) variabilis (Fallen, 1829), which was renamed f.

assimilis by Stichel in 1956 since Douglas & Scott’s name was a homonym of the

earlier Kirschbaum species. Following Stichel (1956) Woodroffe (1957) used the

name Psallus (Hylopsallus) assimilis Stichel, 1956, for the species originally

described by Douglas & Scott. When I tried to establish the identity of the

specimens after which Douglas & Scott described their Apocremnus simillimus it

AUKEMA: Dutch species of Hylopsallus 21

appeared that the three specimens in the Hope Department of Entomology,

Oxford University Museum collected by Douglas from Maple at Box Hill, Surrey,

are females. Consequently the identity of simillimus remains uncertain. The fact

however that the specimens of Douglas were collected in England and on Maple

strongly supports Woodroffe’s interpretation of simillimus. Whether or not

Stichel’s interpretation of simillimus was correct, I regard the name assimilis

Stichel, 1956 as a valid replacement name for simillimus.

Material examined. Great Britain: 48 specimens from 19 localities (15 3/9 loc.,

33 9/17 loc.). Coll. Hope Department of Entomology, Oxford University Mu-

seum: Box Hill, Surrey, J. W. Douglas, Maple, 3 9; 1870, J. W. Douglas, 3 9;

Reigate, vi, 1882, 1 &, 2 9; Renhold, Beds., 6.vii.1962, D. Leston, Maple, 3 3, 3

Q; the same, 30.vi.1963, 1 & (slide Xk4). Coll. British Museum (Natural History),

London: Chingford, 10.vi.1911, E. A. Butler, 1 9; East Malling, Kent, 27.vi.1945,

A. M. Massee, 1 9; Stondon Massey, Essex, 12.vi.1953, A. M. Massee, 1 9;

Epping Forest, Essex, 15.vi.1953, A. M. Massee, 1 9; Ditton, 24.vi.1954, Poplar, 1

Q; the same, 1.vii.1954, 1 © ; Eaglefieldern, 10.vii.1954, G. E. Woodroffe, Maple, 1

d, 1 9; Hedgerley, vi.1955, Maple, 1 9; Runnymede, 10.vi.1955, G. E.

Woodroffe, 3 &, 3 9; Windsor Forest, Berks., 16.vi.1955, G. E. Woodroffe,

Maple, 1 &, 3 ©; Ham Street, Kent, 12.vi.1960, A. M. Massee, 2 9; Old

Burghclere, Beacon Hill, Hants., 23.vii.1962, G. E. Woodroffe, Maple, 1 3,3 9;

Aston Ravent, N. R., 1.vii.1965, G. E. Woodroffe, 1 3; 88—11, 7.vi.1966, Scott, |

gd; Wychwood, N. B., Oxon, 6.vii.1966, G. E. Woodroffe, 1 g; the same,

8.vii.1969, Maple, 1 &,2 9; Power, Birch Wood, 69—31, Birch, 1 &.

Description

This species is very similar to variabilis. Males only can be separated on account

of differences in the aedeagi, but I am not able to separate the females from those

of variabilis. Woodroffe (1957) also separated the males of assimilis from those of

variabilis by the thickening of antennal segment 2: distinctly thickened in variabilis

and only slightly so in assimilis. Basing myself upon the material available for this

study I believe that such a difference does not exist. In Britain however, where

variabilis is seemingly rare, the host plant may be a valuable guide for

identification.

In the following lines I give a differential diagnosis for assimilis, including the

ranges of variation of the measurements, which are somewhat different from those

of variabilis.

Length body/width body &: 2.2—2.5, 9: 2.1—2.4; length of body g: 3.4—3.8

mm, 9: 3.5—4.0 mm.

Head. — Width vertex/width eye g: 1.9—2.1, 9: 1.9—2.4; length antennal

segment 2/length segment 1 &: 4.0—4.7, ©: 3.8—4.7; length segment 2/length

segments 3 + 4 g, 9: 0.9—1.1; length segment 2/width pronotal base ¢: 0.8, 9:

0.7—0.9.

Male genitalia. — Right paramere (fig. 11) and left paramere (figs. 16, 21) as in

22 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

variabilis. Aedeagus (fig. 25) with basal process longer than in variabilis, not

hooked apically. Theca (fig. 31) as in variabilis.

Biology. — Host plant: Acer campestre L. In England imagines are found from

early June until the end of July. There is one generation a year and eggs

overwinter.

Distribution. — Psallus assimilis is thusfar recorded only from England and West

Germany (Stichel, 1958; Rieger, 1975; 1976). The occurence of assimilis in Finland

as mentioned by Stichel (1958) has not been confirmed (Coulianos &

Ossiannilsson, 1976).

Asthenarius quercus (Kirschbaum, 1856)

(figs. 6, 7,12, 17,22,27,32,36)

Kirschbaum, 1856: 253; Wagner, 1952: 176, fig. 108; Stichel, 1956: 290, fig. 745; Woodroffe, 1957:

258, 260, 262, 265, figs. 4, 25; Carvalho, 1958: 128—129; Southwood & Leston, 1959: 220, 225, figs. 75,

83; Kerzhner, 1962: 232; Wagner, 1967: 145, figs. 98D, 99B; Zaytseva, 1969: 526, 528—529, figs. 10—14;

Wagner, 1975: 183—184, figs. 751b, 753b; Wagner, 1975a: 242—243, fig. 6. — Capsus simillimus Kirsch-

baum, 1856: 233; Wagner, 1943: 34— 36.

Material examined. — The Netherlands: 170 specimens from 46 localities (74 G/

30 loc., 96 9/38 loc).

Description

A relatively large species; males elongate, rather parallel-sided (fig. 6), females

smaller, elongate-oval (fig. 7). Length body/width body 3: 2.3—2.9, 9: 2.1—2.7.

Dark-brown to red-brown, females often lighter than males. Pubescence

consisting of both shiny white scales, which are easily rubbed off, and semi-erect

black hairs.

Head. — Dark-brown to red-brown, sometimes blackish. Width vertex/width

eye &: 1.3—2.0, 2: 1.7—2.3. Antennae yellow-brown, especially in segments | and

2 tinged with red. Segment 1 basally very small dark. Length segment 2/length

segment | g: 4.2—5.4, ©: 4.2—5.3; length segment 2/length segments 3 + 4 &:

1.0—1.3, 9: 0.9—1.3; length segment 2/width pronotal base &: 0.9—1.l, 9:

0.9—1.0. Rostrum dark-brown to red-brown, base of segment | ventrally white

with reddish tinge; length ¢: 1.6 mm (only 2 specimens measured), 9: 1.6—1.9

mm.

Thorax. — Pronotum dark-brown to red-brown, sometimes blackish, but

posterior angles or entire hind margin often lighter. Mesoscutum and scutellum

coloured as pronotum, laterally often lighter. Hemelytra dirty grey-brown to dark-

brown, basally often lighter, sometimes greyish-yellow. Apex and lateral margin

more or less tinged with red. Cuneus red-brown, with base along fracture narrowly

whitish. Wing membrane greyish to black-grey with a clear spot just behind outer

cell. Veins colourless, sometimes tinged with red-brown or red. Femora basally

AUKEMA: Dutch species of Hylopsallus 23

dark-brown to red-brown, apically coloured lighter, bright-red or yellow-brown

with a red tinge. Hind femora sometimes with a few obscure dark spots beneath.

Coxae and trochanters greyish-white in males and white in females. Tibiae yellow

with red tinge, especially in hind tibiae; apices small blackish. Tibiae with strong

black spines arising from large black to dark-brown spots, which are however

smaller in mid and fore tibiae. Tarsi yellow-brown, segment 3 mainly dark. Claws

yellow-brown; pseudarolia small.

Ventral surface. — Red-brown to dark-brown; thoracic spiracle, epimeron,

ostiolar peritreme, basalar plate and margins of propleura whitish with a red tinge.

Male genitalia. — Genital segment with a distinct ventral keel. Right paramere

small and oblong (fig. 12); left paramere small, with sensorial process acutely

angled apically (figs. 17, 22). Aedeagus short and C-shaped, without lateral

processes, its apex with a long twisted denticulate band ending in a long and

narrow apical process (fig. 27). Secondary gonophore situated near apex. Theca

small and strongly tapering apically without elevation before apex (fig. 32).

Length of body ¢: 3.8—4.7 mm, 9: 3.5—4.3 mm.

Biology. — In the Netherlands common on its host plant oak, especially on

Quercus robur L. Imagines are found from mid-May until the end of July. There is

one generation a year and eggs overwinter.

Distribution. — A very widespread West Palearctic species, known from

Finland, Norway, Poland, Germany, Denmark, Scotland, England, the

Netherlands (fig. 36), Belgium, France, Spain, Portugal, Italy, Switzerland,

Austria, Czechoslovakia, Hungary, Yugoslavia, Bulgaria, Roumania and South

Russia (Stichel, 1958).

CONCLUDING REMARKS

Very little is known of the biology of the Dutch species of Hylopsallus, whereas

knowledge of that kind is essential for understanding processes of speciation

within such a group of closely related species. Especially in the case of perrisi and

wagneri, which differ only in size and shape of the aedeagus, and occur on the same

host plant in the same period of the year, it is difficult to understand their

relationships. The same applies for Psallus (Hylopsallus) kiritshenkoi, which is

described by Zaytseva (1969) from South Russia. This species differs from perrisi

and wagneri only in size and shape of the apical process of the aedeagus, which is

intermediate between those of perrisi and wagneri!

It is clear that further study of the biology of these species and rearing and cross-

breeding experiments are necessary to unravel the relationships of these

morphologically ill-defined species.

ACKNOWLEDGEMENTS

For the loan of specimens I am greatly indebted to the following institutions and

persons: British Museum (Natural History), London (Mr. W. R. Dolling); Hope

24 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 1, 1981

Department of Entomology, Oxford University Museum (Dr. I. Lansbury); Dr. B.

Tjeder (Lund); Dr. F. Ossiannilsson (Uppsala); National Institute for Plant

Protection, Solna (Dr. G. Nordlander); Swedish Museum of Natural History,

Stockholm (Dr. P. I. Persson); Zoologiska Institutionen, Lund; Mr. J. S. Beernink

(Zaandam); Mr. W. H. Gravestein (Amsterdam); Mr. P. Poot (Maastricht); Mr. J.

H. Woudstra (Zaandam); Landbouwhogeschool, Wageningen (Dr. Ir. R. H.

Cobben); Natuurhistorisch Museum, Maastricht (Mrs. F. N. Dingemans-Bakels);

Rijksmuseum van Natuurlijke Historie, Leiden (Dr. P. H. van Doesburg).

Furthermore I am greatly indebted to Dr. J. P. Duffels (Instituut voor

Taxonomische Zoölogie, Zoölogisch Museum, Amsterdam) for giving me the

opportunity to do this investigation, for his valuable help and advise and for

correcting the English text.

REFERENCES

Carvalho, J. C. M., 1958. Catälogo dos Mirideos do mundo. — Archos Mus. nac. Rio de J. 45 (2):

1—216.

Coulianos, C. C. & F. Ossiannilsson, 1976. Hemiptera-Heteroptera. In: Catalogus Insectorum Sueciae.

2nd Ed. VII. — Ent. Tidskr. 97: 135—173.

Douglas, J. W. & J. Scott, 1865. Hemiptera-Heteroptera. In: The British Hemiptera. I: xii + 627 pp. —

Ray Soc., London.

Fallen, C. F., 1829. Hemiptera Sueciae: viii + 186 pp. — Lund.

Josifov, M., 1970. Ergebnisse der Albanien-Expedition 1961 des Deutschen Entomologischen Institu-

tes. — Beitr. Ent. 20: 825—956.

Kerzhner, I. M., 1962. Materials on the taxonomy of Capsid bugs (Heteroptera, Miridae) in the fauna

of the USSR. — Ent. Obozr. 41: 372— 387.

Kerzhner, I. M. & T. L. Yachevski, 1964. Heteroptera. In: Bei-Bienko, Key to insects of the European

part of the USSR: 1: 1—936.

Kirschbaum, C. L., 1856. Die Capsinen der Umgebung Wiesbadens. — Jb. Ver. Naturk. Wiesbaden 10:

163— 348.

Meurer, J. J., 1956. Waarnemingen van Wantsen (Hem.-Het.) met behulp van een vanglamp. — Ent.

Ber., Amst. 16: 54—63.

Mulsant, E. & C. Rey, 1852. Description de quelques Hémiptéres Hétéropteres nouveaux ou peu con-

nus. — Annls Soc. Linn. Lyon 1: 76—141.

Ossiannilsson, F., 1953. Psallus (Hylopsallus) wagneri n. sp. A new Mirid (Hemipt. Heteropt.) from Swe-

den. — Opusc. ent. 18: 122—123.

Reclaire, A., 1932. Naamlijst der in Nederland en het omliggend gebied waargenomen wantsen (He-

miptera-Heteroptera). — Tijdschr. Ent. 75: 59—258.

—, 1948. Se Vervolg op de Naamlijst der in Nederland en omliggend gebied waargenomen wantsen

(Hemiptera-Heteroptera). — Tijdschr. Ent. 89: 39—64.

Rieger, Chr., 1975. Nachweiss des Psallus masseei Woodroffe in Süddeutschland (Heteroptera, Miri-

dae). — NachrBI. bayer. Ent. 24: 57—58.

——., 1976. Die Wanzenfauna des mittleren Neckartales und der angrenzenden Albhochflache

(Landkreis Nürtingen, Reutlingen, Tübingen). 3. Nachtrag. — Veröff. Naturschutz Landesp-

flege Bad.-Württ. 43: 162— 169.

Reuter, O. M., 1876. Diagnoses praecursoriae. — Petites Nouv. Ent. 2 (144): 21—22.

, 1878. Hemiptera Gymnocerata Europae. I. — Acta Soc. Sci. Fenn. 13: 1—187, 8 pls.

Southwood, T. R. E. & D. Leston, 1959. Land and water bugs of the British Isles: ix + 436 pp., 63 pls.

— London.

Stichel, W., 1956— 1958. Illustrierte Bestimmungstabellen der Wanzen. II. Europa. 2: 170—907. Berlin.

Wagner, E., 1943. Zwei Kirschbaumsche Arten, die sind nicht aufrecht erhalten lassen. — Mitt. ent.

Ges. Halle 20: 33—36.

AUKEMA: Dutch species of Hylopsallus 25

——, 1952. Zur Systematik von Psallus variabilis (Fall.) (Hem. Het. Miridae). — Opusc. ent. 17:

92—94.

—, 1952a. Blindwanzen oder Miriden. In: Tierwelt Deutschlands 41: iv. + 218 pp. — Jena.

—, 1959. Ueber einige Mediterrane Miridae (Hem. Het.). — Acta ent. Mus. natn. Pragae 33 (553):

339— 358.

—, 1961. Heteroptera-Hemiptera. In: Tierwelt Mitteleuropas IV (Xa): 1— 173. — Leipzig.

——, 1966. Wanzen oder Heteropteren. I. Pentatomorpha. In: Tierwelt Deutschlands 54: vi + 235

pp. — Jena.

———, 1966a. Phylidea Reuter, 1899 und Psallus Fieber, 1858 (Hemiptera, Heteroptera, Miridae). —

Reichenbachia 8: 11—20.

——, 1967. Wanzen oder Heteropteren. II. Cimicomorpha. In: Tierwelt Deutschlands 55: iv + 179

pp. — Jena.

—, 1970. Die Miridae Hahn, 1831, des Mittelmeerraumes und der Makaronesischen Inseln (He-

miptera, Heteroptera). I. — Ent. Abh. Mus. Tierk. Dresden 37, Suppl.: iii + 484 pp.

—, 1975. Idem. III. — Ent. Abh. Mus. Tierk. Dresden 40, Suppl. 1: 1—483.

———, 1975a. Ueber Asthenarius Kerzhner, 1962 (Hemiptera, Heteroptera, Miridae). — Reichenba-

chia 15 (28): 233— 244.

Wagner, E. & H. H. Weber, 1964. Heteropteres Miridae. In: Fauna de France 67: 592 pp. — Paris.

Woodroffe, G. E., 1957. A preliminary revision of the British Psatlus Fieber (Hem., Miridae), with a

description of a new species. — Entomologist’s mon. Mag. 93: 258—271.

Zaytseva, I. F., 1969. A survey of species of the genus Psallus Fieber (Heteroptera, Miridae) from the

Caucasus. — Ent. Rev. Washington 47: 526—533.

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TIJDSCHRIFT

HARVARD

VOOR ENTOMOLOGIE ~

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

C. Wırkınson and P. J. Newron. — The microlepidopteran genus Ectoedemia

Busck (Nepticulidae) in North America, p. 27—92, figs. 1—71.

C. Wırkmson. — A supplement to the genus Ectoedemia Busck (Nepticulidae:

Lepidoptera) in North America, dealing with some difficult species and also

some new ones, p. 93—110, figs. 1-9.

Tijdschrift voor Entomologie, deel 124, afl. 2 & 3 Gepubliceerd 20-VII-1981

THE MICRO-LEPIDOPTERAN GENUS ECTOEDEMIA

BUSCK (NEPTICULIDAE) IN NORTH AMERICA

CHRISTOPHER WILKINSON

Department of Animal Systematics and Zoogeography, Vrije Universiteit, Amsterdam

and

P.J. NEWTON

20 Mellor Brow, Mellor, Blackburn, England

With 71 text-figures

ABSTRACT

This paper, together with the monograph by Wilkinson & Scoble (1979) forms a comprehensive taxo-

nomic account of the genus Ectoedemia Busck in North America. Species are here fully diagnosed and

illustrated with drawings of the external features and genitalia. Notes on their biology are also included.

CONTENTS

FAND SCT AC ent EA eer e EVO RS o Re a O p. 27

Introduction Re RO ih ARNO NERI EE it ETA PLONE p.27

Methods paves a Warn cer Re se DIRO RE Blest AE IRE p. 28

Ab breviati Onsite LA RIT Een AREN EN AA p. 28

Acknowledgements ende ne CD CR p. 29

MAXONOMICINISEORY NI a RE DE RE ER N ARE. p. 29

Tiaxonomieteonsiderationsn VENIER RA RI EEE RER p. 31

GenericidescriptionW eeen RE RORE EREN des pase

Generic differentialidiagnosissamr sari ac Ee p. 34

eheckelisttorpenussspeciestandisubspeeie Se ay erie TI. p. 36

ICV ALOISPECIESLAT ASI SER 0 TIA AD SER VAS NES I IAN TI SIAE ants ee p. 37

Table for the identification of Ectoedemia species from host plants and damage .......... p. 40

he populella TOUR SS Den: AIA NATI p.4l

DIEIPIAANC RIO PRA SARO p. 51

he bo ie a TOURMENTE Ee NRN p.61

TWO CaS tan CAE TOUS). Lion Bon |e, MEN RER EEE STE p. 72

References ME PRE CAT ee er a ee I OO CE Re RE p. 86

An index to the names of taxa dealt with in the present paper can be found at the end of the fol-

lowing paper.

INTRODUCTION

In this revision twenty-two species are considered. Five are new combinations,

two are new species and nine are diagnosed and figured which have previously

28 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

been described by Wilkinson & Scoble (1979). Those species attacking bark are

described at the end and form a clear group separable on the basis of genitalia

from the leaf and petiole miners. In the same way other groups amongst the latter

are recognisable. However, those attacking petioles, e.g. E. populella and canutus,

are included in one of the groups of leaf-miners. This action is supported by the

presence of an intermediate habit. In argyropeza the larva first bores into the pet-

iole and later continues to mine the leaf.

Thus in Ectoedemia it is now definitely established that three sites of attack exist:

leaf, petiole and bark. There is also a reference to attack in the cortex. Both galls

and mines are found in these locations.

There has been much discussion in the past regarding the boundaries of the

genus. This revision, with the advantage of additional material, sheds light on this

debate.

METHODS

The methods used are similar to those given in Wilkinson & Scoble (1979) as are

also the terms used, although in the five years that have elapsed since the former

work was completed we have modified some of our terminology. We have come to

realise that a thorough comparative morphological study of the various nepticulid

genera is long overdue, not only to ascertain homologies within the group but also

with other families. At present one is not always certain that parts of genitalia

given the same names as those in the Ditrysia are indeed homologous. For ex-

ample the interpretation of the term “saccus” by Beirne (1945) is unclear and has

not been adopted by other workers to mean a part of the integument as he seems

to interpret it. Here “‘saccus” refers to the anterior extension of the vinculum in

front of the ventral plate and is the area often extended into a bilobed protrusion.

Whether this is homologous with the saccus of Ditrysia is uncertain and therefore

the continued use of the term is open to question.

“Pseuduncus” is a posterior extension of the tegumen and a true uncus is not

present in Ectoedemia. However, some species have a thickened area in the region

where an uncus might be expected to arise. Scoble (verbal communication) sug-

gests that at least in the African species he has examined, the thickening rep-

resents a forward fold of the pseuduncus and therefore the uncus is still absent.

Expendable material of trinotata would be particularly useful for the investigation

of this point.

“Anellar lobes” refers to the large spines located at the end of the aedeagus. It

is not always certain if they are homologous. It is assumed they provide the normal

supporting role for the aedeagus. Sometimes, however, they might be more re-

ferable to the juxta or aedeagus proper.

Scale. — On the genitalia figures the scale line represents one tenth millimeter

(1/10 mm) unless otherwise indicated. The magnification of the external figures

can be ascertained from the al.ex. measurements (taken from centre of meso-

thorax to wing tip and doubled) given for each species.

ABBREVIATIONS

Institutes from which material was borrowed:

WILKINSON & NEWTON: Ectoedemia in North America 29

USNM — United States National Museum of Natural History, Smithsonian In-

stitution, U.S.A.

CNC — Canadian National Collection, Biosystematics Research Institute,

Ottawa, Canada

FIS — Forest Insect Survey, British Columbia, Canada

SOO — Forestry Service, Environment Canada, Sault Ste. Marie, Canada

ANS — Academy of Natural Sciences, Philadelphia, U.S.A.

BM (NH) — British Museum (Natural History), London, U.K.

MCZ — Museum of Comparative Zoology, Cambridge, Massachusetts,

U.S.A.

DFF — Department of Fisheries and Forestry, Forestry Service, (now Envi-

ronment Canada), Ste. Foy, Quebec, Canada.

ACKNOWLEDGEMENTS

Our thanks are due to Mej. R. Kloos and B. J. van Cronenburg who assisted

with the final production of the drawings and to Dr. Georgina Bryan who read the

final manuscript and made helpful suggestions. P. J. N. wishes to acknowledge the

support of Portsmouth Polytechnic during his tenure as research assistant to C. W.

TAXONOMIC HISTORY

The question of the significance of larval feeding habit and the pattern of

forewing venation has led to some confusion in the past and is now the focus of

discussions on the composition of this genus and its phylogeny with respect to the

other members of the family. The genus was originally described by Busck (1907)

as monotypic, because the species populella which, despite affinities with Stigmella

Schrank could not be included within its confines. The genus and author were

cited incorrectly by Busck as Nepticula Zeller. His action was right but his reason

was wrong. It was that the gall-forming habit of populella excluded it from the leaf-

mining species of Stigmella. Busck also gives major morphological differences be-

tween the types of Stigmella and Ectoedemia, including the closed cell of the

forewing venation formed by the medial cross-vein, which is actually also present

in other genera. For some time Braun had been aware of the two basic forms of

forewing venation, those with a medial cross-vein and those without. This feature,

however, was not given taxonomic status by her for a number of reasons. Firstly,

she regarded otherwise closely related species and sometimes male and female in-

dividuals of the same species as showing both conditions of the pattern of venation

(Braun, 1917: 157, and figs. 1 and 2). This conclusion was the result of misidentifi-

cation of a single species (nyssaefoliella), which she thought had both forms of ve-

nation and, more generally, is due to the absence of any study of the genital

morphology at that time. Again Braun and other of her contemporary workers re-

garded the mining habit of the larvae as the most significant indication of phy-

logeny. It was not until later examination of the genital morphology that evidence

in favour of uniting leaf-mining species and gall-forming species within a single

genus, became available.

30 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Beirne (1945) divided the family into two basic groups on the form of the male

genitalia, those with a ring-shaped vinculum and those with an incomplete vin-

culum. This led him to erect the genus Dechtiria for those leaf-mining species with

distinct differences in genitalia from the leaf-mining Stigmella (and Nepticula sensu

Beirne) species. A study of the group by Svensson (1966: 200) with a comparison of

genital morphology and venation of North American types, led him to synonymise

the European genus Dechtiria with the North American Ectoedemia Busck. This

decision has been accepted by Wilkinson & Scoble (1979), who included seven

leaf-mining species, which were earlier assigned to Stigmella, in Ectoedemia. In the

present study a further five species are combined with Ectoedemia, all of which are

leaf-miners. Beirne (1945: 204) noted that in the genus Dechtiria, with the ex-

ception of two species only, all species formed leaf-mines which terminated in

blotches. In the present revision only one included species, virgulae, forms a linear

leaf-mine. The remainder form either true blotches or linear mines terminating in

blotches.

Immediately after the publication by Zimmermann (1940) of work on the bark-

miner, Ectoedemia liebwerdella Zimmermann, Hering (1940) erected the genus

Zimmermannia, with this species as the type. Hering proposed this division on the

basis of a difference in larval feeding habit, comparing the North American type

populella Busck, which forms petiole galls in poplar, and the European species lieb-

werdella, which burrows in the bark of beechwood. Hering noted that the two

species were indistinguishable in the pattern of wing venation but that they dif-

fered in that liebwerdella possesses a row of sharp bristles on the inner side of the

hind-tibiae. Borkowski’s study (1972: 693) points out that this character is present

in all the European Ectoedemia species he examined, whether gall forming or bark

burrowing. This view is also held here for the American species but there is some

variation in the prominence of the bristles.

Schönherr (1957: 127; 1958: 6) subsequently synonymises Zimmermannia with

Ectoedemia and gives it subgeneric rank. This is confirmed by Borkowski (1972),

who points out that the morphological differences given by Hering are not suffi-

cient to warrant generic status being attached to Zimmermannia. He also presents

a summary of the treatment of the three genera at the time and supports the di-

vision of Ectoedemia sensu lato into three subgenera corresponding to larval

feeding habit: the petiole-miners, Ectoedemia Busck; the leaf-miners, Dechtiria

Beirne; and the bark-miners, Zimmermannia Hering. The group is treated similarly

by Emmet (1976) in his revision of the British fauna.

Although Borkowski is in some doubt that sufficient work was done by Svensson

in his synonymy of Dechtiria with Ectoedemia, it is felt that the work by Wilkinson

& Scoble (1979) on the Canadian fauna and the present study confirms the syn-

onymy. On the basis of genitalia and wing venation the authors also support the

synonymy of Zimmermannia with Ectoedemia. However, from this study it is pos-

sible to identify morphological divisions, mainly based on genitalia structures,

within the genus, which correspond with the sites of larval attack.

Johansson (1971: 241) is of the opinion that the family Nepticulidae may be split

into two genera, Nepticula and Trifurcula. He argues that Trifurcula has been subdi-

vided into several genera including Ectoedemia and Dechtiria, but that these should

WILKINSON & NEWTON: Ectoedemia in North America 31

be included as subgenera of Trifurcula on the basis of male genitalia. This scheme

is followed more recently by Karsholt & Schmidt Nielsen (1976) in the catalogue

of Danish Lepidoptera. Although there is merit in the proposal, this classification

is not adopted in this revision for reasons given by Wilkinson & Scoble (1979). It

was pointed out earlier that there are no significant structural differences between

the bark-miners and bark gall-formers. For example, heinrichi and castaneae

cannot be reliably separated without knowledge of the host plant and damage.

However, examination of the genitalia figures shows that species which attack

bark (whether galls or mines) have characteristic genitalia and leads one to guess

at the habits of those whose life history is unknown. The three species recorded

here as attacking petioles can be placed in the same morphological group on the

basis of genital structures, but the group also contains leaf-miners.

TAXONOMIC CONSIDERATIONS

The similarity in morphological structures, even genitalia, between castaneae

and heinrichi, highlights problems over the species concept in this group of Lepi-

doptera. Knowledge of the life history leads us to separate two species which

otherwise might have been regarded as one. Speciation is much more extensive

than has hitherto been recognised and many species are only known through the

excellent rearing work of Braun and others. In this genus alone, many species are

shown to have more than one form in the externals, particularly wing markings,

which is not matched by differences in the genitalia. At first sight a species appears

to be di- or polymorphic, but without knowledge of the life cycles, we cannot be

sure that these variants are not distinct species. For example, three wing pattern

forms occur in what has been called E. similella. Although they have identical gen-

italia, one form occurs in Ohio, New Jersey and Virginia, another form in Florida.

A single specimen with different markings has been collected from Arkansas. Host

plants are only known for the Ohio form (Quercus palustris and Q. rubra).

A specimen (nr. 7) probably represents a new species close to rubifoliella. It

differs in ground colour and certain aspects of the genitalia. It was reared by

Braun from a collection of similar mines on Quercus platanoides but the rest turned

out to be Stigmella flavipedella (Braun) comb. n. This is one of a number of ex-

amples which shows that each larva should be reared separately, as far as practi-

cable, and also have a separate rearing number. The fact that the host plants of ru-

bifoliella and specimen 7 are so different, supports the view that specimen 7 should

be treated as a new species but this has not been done in case there is anything

spurious about its breeding record. Otherwise its similarity to rubifoliella is quite

striking. The male genitalia figures are drawn to accentuate the differences which

may not be so easy to diagnose as they at first seem.

Of course, the problem must also be approached from the other direction. It is

quite possible that a species could attack more than one part of a plant. It is

known that E. argyropeza downesi mines both the petiole and the lamina. It is also

possible that the same species may create mines or galls depending upon the site of

oviposition. Although many species are monophagous not all are, and we should

therefore be prepared for dimorphism or different reactions from different host

32 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

plants to the same species. Some authorities think this unlikely but there is consid-

erable danger in the philosophy — two species of host plant, therefore ipso facto

two species of miner! However, hopefully these problems will encourage experi-

mental work designed to discover the flexibility in miner and host plant

relationships within the Nepticulidae. Meanwhile these cases underline the point

that no one character, or series of characters, can be taken as necessarily species-

diagnostic throughout the Nepticulidae or even for Ectoedemia.

There has been debate regarding the division of the genus into sub-genera and

also whether Ectoedemia comprises, in reality, more than one genus. Much more

information on life cycles is needed to answer this, but this paper shows that the

criteria previously used for separations are inadequate. The genitalia structures

allow us to divide Ectoedemia into broad groups and it seems that those species at-

tacking bark (both mining and gall-forming) are particularly diagnostic. However,

we do not yet know whether all those species assigned to the group do, in fact,

attack bark. In other groups it can be argued that one or two species (trinotata,

nyssaefoliella) have males belonging to a different group from the females, thus

suggesting that the groups are not fundamental. Our present level of knowledge

recommends us to regard these species as respresenting one genus and our studies

of Ectoedemia on other continents will shed further light on these conclusions. It is

interesting to note how some genera of Nepticulidae are worldwide in their distri-

bution whilst others are comparatively localised.

GENERIC DESCRIPTION

External features: ¢ 9. Head: palps extending well beyond labrum, pale grey or

white; antennae approximately half length of forewings, pale or fuscous; tuft on

front of head usually ochreous, sometimes white or brown, vertex concolourous;

eye-caps always whitish; collar white or ochreous, sometimes fuscous. Thorax

dark brown to black, sometimes white or buff and irrorate, usually iridescent. Ab-

domen grey or brown, iridescing metallic grey beneath. Venation as in figs. | or 2.

Forewings: Media coalescing with cubitus at base and passing obliquely to radius

at or beyond R,,, and anastomosing beyond middle of wing; cubitus usually

reaching margin; R, and R, separate; anal vein sometimes meeting cubitus dis-

tally. Hindwings: Media single-branched. Forewings: elongate and ovate in shape,

ground colour of dorsal surface generally fuscous with each scale darker at the tip,

rarely pale and irrorate with fuscous; fringe pale grey or brown and iridescent,

usually marked apically with a band of wing-scales or variously irrorate; markings

usually in the form of up to four streaks or patches, never with more than one

complete fascia. Hindwings: narrow and lanceolate, sometimes with costa

emarginated; pale grey or brown and sometimes with white or ochreous patches of

specialised scales. Legs: grey or brown, sometimes with scattered paler areas;

proximal pair of spurs on hind-tibiae at or below the middle (fig. 3); hind-tibiae

with many sharp bristles inwardly directed and variable in prominence.

Male genitalia: Vinculum always ring-shaped; tegumen fused with vinculum dor-

sally and produced into broad tapering, pointed or bluntly rounded pseuduncus.

Uncus often not readily visible, only present as membranous lobe at base of pseu-

WILKINSON & NEWTON: Ectoedemia in North America 33

1 Ectoedemia obrutella d

2 Ectoedemia similella Q

Fig. 1. Ectoedemia obrutella (Zeller) Busck, &, wing venation. Fig. 2. E. similella (Braun), 9, wing ve-

nation.

duncus. Gnathos with medial arms fused to form central boss or tapering process,

lateral arms rarely extended or bifurcate. Saccus bilobed to various degrees.

Valves usually tapering and inwardly curved distally, sometimes weakly bifurcate

and in some cases adorned with digitate setae. Transtillae usually U-shaped; trans-

verse bars continuous, fused medially. Juxta absent. Aedeagus: generally regular in

shape and longer than the capsule; vesica usually adorned with small denticles and

cornuti; also with a plate of minute papillae. Anellus with heavily sclerotised

spines, sometimes elaborate, rarely absent.

Female genitalia: Apophyses usually shorter than ductus but posteriores very

long in the castaneae group. Ductus at colliculum with double sclerotised ring and

associated denticulate sac, or with single plate, or without sclerotisation. Bursa

34 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Fig. 3. Ectoedemia species. Legs showing position of tibial spurs.

copulatrix: large and variously pectinate; signum double, comprising ovate

patches of reticulate cells.

Larvae: may be divided into three groups: leaf-miners usually forming linear

tracts terminating in blotches; petiole miners and bark miners both sometimes

producing galls.

GENERIC DIFFERENTIAL DIAGNOSIS

Characters which differentiate North American genera of the family Nepticu-

lidae are given.

Ectoedemia Busck, 1907.

Venation: Media of forewing coalescing with Cubitus at base, passing obliquely

to Radius at or beyond R,,; and anastomosing to a point beyond middle of wing;

R, and R, separate; Cubitus usually approaching margin; Media of hindwing

single. Proximal pair of spurs on hind tibiae sometimes in the middle. Male geni-

talia with gnathos W- or V-shaped, may vary according to method of mounting;

vinculum ring-shaped and without associated lateral bars; tegumen extended into

tapering or lobed pseuduncus; uncus absent or weakly membranous; valves in-

wardly curved distally sometimes with digitate setae; juxta absent; aedeagus

regular in shape with elaborate cornuti and usually anellar spines. Female genitalia

with or without complex sclerotisation of the ductus and spiculate accessory lobe;

apophyses shorter than ductus; signa comprising patches of reticulate cells. Larvae

may mine in, or form galls on leaves, petioles, bark or cortex.

WILKINSON & NEWTON: Ectoedemia in North America 35

Stigmella Schrank, 1802.

Venation: Media of forewing coalescing with Radius at base and anastomosing

to a point beyond the middle of the wing; R, coincident with R,; Cubitus arising

separately, approaching middle of the wing; Media of hindwing single. Forewings

usually uniform and dark in colour, with one or two complete fasciae or patches;

fringe with diffuse margin. Proximal pair of spurs on hind tibiae above the middle.

Male genitalia usually with U-shaped vinculum; tegumen strap-like, articulating

with vinculum dorsally; uncus bilobed; juxta, if present, membranous; aedeagus

usually flask-shaped, vesica usually with many denticulate cornuti orientated in a

ridge and rarely with plate-like cornuti at the anellus. Female genitalia with simple

ductus and accessory sac; bursa copulatrix usually without signum, but if present

often single and weakly sclerotised. Larvae mining leaves of trees and shrubs and

sometimes herbs.

Microcalyptris Braun, 1925.

Venation: Reduced; Media of forewing coalescing with Radius from base and

anastomosing to a point beyond the middle of the wing; R, coincident with R,;

Cubitus vestigial; Media of hindwing single, unbranched. Ground colour of dorsal

surface of forewing usually pale and variously irrorate. Proximal pair of spurs on

hind tibiae below middle. Male genitalia, with membranous pseuduncus and

strongly sclerotised bridge-like uncus; sclerotised gnathos with complex anterior

and posterior projections; lateral arms of vinculum usually with associated sclero-

tisations. Female genitalia with complex sclerotisations of the ductus; posterior

apophyses very long, longer than the ductus; signa usually comprising linear row

of spinose cells or plates. Larvae mining leaves.

Fomoria Beirne, 1945.

Venation: Media coalescing with Cubitus from base, both passing obliquely to

Radius at R,,; and anastomosing to beyond middle of wing; Cubitus becoming ob-

solete; R, and R, separate; Media of hindwing single, unbranched. Male genitalia

with membranous pseuduncus and uncus as a spatulate sclerotisation; Y- or V-

shaped gnathos; saccus weakly bilobed; valves sometimes with dorsal spine; ae-

deagus regular in shape and usually with complex anellar spines and cornuti.

Female genitalia, colliculum with sclerotised funicular antrum or complex plates;

simple ductus; signa comprising linear patches of reticulate cells. Larvae often re-

corded pupating within the leaf-mine.

Obrussa Braun, 1915.

Venation: Media of forewing coalescing with Cubitus at base, both passing

obliquely to Radius at R,,, and anastomosing to beyond middle of wing; Media

and Cubitus separate terminally; R, and R, separate; Media of hindwing single.

Ventral surface of forewing and dorsal surface of hindwing in males with patch of

brightly coloured specialised scales. Proximal pair of tibial spurs below middle of

hind tibiae. Male genitalia with ring-shaped vinculum; tegumen extended into ta-

pering pseuduncus; convex saccus; valves blunted distally and each with large

dorsal arm projecting transversely to reach opposite side of capsule; vesica with

36 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

transverse plate expanded laterally. Female genitalia with plate-like sclerotisation

at colliculum; signa comprising ovate reticulate patches. Larvae only known to

mine fruits of Acer sp.

Glaucolepis Braun, 1917.

Venation: Media of forewing coalescing with Cubitus at base, both passing

obliquely to Radius at R,,, and anastomosing to beyond middle of wing; Media

and Cubitus separate terminally; R, and R, separate; Media of hindwing bifurcate.

Hindwing of male with patch of brightly coloured specialised scales. Proximal pair

of spurs on hind tibiae in the middle. Male genitalia with tegumen extending into

tapering pseuduncus, gnathos with large transverse arms and medial dorso-lateral

arms fusing terminally; valves markedly bifurcate distally; aedeagus with lateral

cornuti extending full length of vesica and digitate distally. Female genitalia with

simple ductus; signa comprising linear patches or rows of pectinations. Larvae

mining leaves.

Manoneura Davis, 1979 (Replacement for homonymic name Oligoneura Davis,

1978).

Venation: greatly reduced; only two branches of Radius present; Media un-

branched and arising from stem of R,,,; Cubitus absent; hindwing extremely

slender and Media unbranched. Forewing dark fuscous with a single, narrow, pale

golden yellow fascia at distal third. Proximal pair of spurs on hind tibiae near apex.

Male genitalia with uncus vestigial; gnathos well developed, complex, consisting of

two, largely separate, transverse sclerites of a highly irregular, but symmetrical

outline; vinculum broad, quadrate; aedeagus moderately short and stout, with a

relatively complex apex and no cornuti. Female genitalia unknown. Presumably a

leaf mining genus. The type-species mines Coccoloba uvifera (L.).

Artaversala Davis, 1978.

Venation: greatly reduced; Radius unbranched, terminating well short of apex;

Media unbranched, extending almost to apex; Cubitus shortened, indistinctly

present; hindwing extremely slender and Media unbranched. Forewings with a

pale yellow to whitish apex and a single, broad, median fascia. Proximal pair of

spurs on hind tibiae near apex. Male genitalia with tegumen reduced to an ex-

tremely slender dorsal ring; uncus absent; vinculum well developed and V-shaped

valves deeply divided and aedeagus relatively simple, without cornuti. Female gen-

italia with slender and elongate ductus; bursa copulatrix membranous; signa

absent. Larvae mining leaves.

CHECK-LIST TO GENUS, SPECIES AND SUBSPECIES

Ectoedemia Busck, 1907. Type-species by original designation and monotypy: Ectoedemia populella

Busck.

Zimmermannia Hering, 1940. Type-species by original designation: Ectoedemia liebwerdella Zim-

mermann, 1940: 264.

Dechtiria Beirne, 1945. Type-species by original designation: Tinea subbimaculella Haworth, 1828: 583.

WILKINSON & NEWTON: Ectoedemia in North America

populella Busck, 1907

argyropeza downesi Wilkinson & Scoble, 19791)

canutus Wilkinson & Scoble, 1979

trinotata (Braun, 1914)

marmaropa (Braun, 1925)

platanella (Clemens, 1861)

maximella (Chambers, 1873)

clemensella (Chambers, 1873)

similella (Braun, 1917)

virgulae (Braun, 1927)

lindquisti (Freeman, 1962)

rubifoliella (Clemens, 1860)

ulmella (Braun, 1912)

nyssaefoliella (Chambers, 1880)

quadrinotata (Braun, 1917)

obrutella (Zeller, 1873)

bosquella (Chambers in Hayden, 1878a)

acanthella sp.n.

piperella sp.n.

heinrichi Busck, 1914a

castaneae Busck, 1913

phleophaga Busck, 1914b

chlorantis Meyrick, 1928a

mesoloba Davis, 1978

KEY TO THE SPECIES OF NORTH AMERICAN Ectoedemia

1. Male genitalia with aedeagus totally lacking anellar projections. Female geni-

talia with excessively broad anterior apophyses together with unequal signa

andmolcCollicul Um VERE ROR SANS II MAATEN E RISSA. 2

Male genitalia with anellar spines on aedeagus. Female genitalia without

aboverceombinationolicharacters SP PM O IR TI 3

. Male genitalia with valves relatively short; as in fig. 28. Female genitalia as in

fig. 29. Externals as in figs. 58, g, 59, 9. Larva forms ophistigmatonome on

INDSSOMIEAVEST IH UN RA ARNO RER A nyssaefoliella (p. 67)

Male genitalia with valves relatively long; as in fig. 30. Externals as in fig. 60.

Remalemotiknowni ast ame Seren. ante specimen 8 (p. 70)

. Male genitalia with long tapering pseuduncus extending well beyond valves.

Female genitalia with strongly sclerotised colliculum in the form of loops.

Leaf and petiole miners and petiole gallformers ................. 4

Male genitalia with large rounded pseuduncus not usually projecting beyond

valves and never far beyond. Female genitalia without colliculum or if weakly

sclerotised, never in form of loops. Mostly species attacking bark ..... 13

. Male genitalia with palmate setae on the dorsal surface of valves, (not well de-

veloped in one species). Female genitalia with enlarged spinose sac in asso-

ciationswithisclerotisedicolliculum Zr een EEE TERRE 5

Male genitalia without palmate setae on valves. Female genitalia without

spinoselobetatzcollieulumer a 6oonoge655bbo5bGn00000700000 8

1) The nominate subspecies of E. argyropeza (Zeller, 1839) is European and therefore not included

10.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

. Male genitalia with small three-pronged setae on dorsal surface of valves as in

fig. 15. Hindwing of male without lance-shaped chitinous plate on costal

margin. Female genitalia with equal signa, thin colliculum and minute spines

on ductus as in fig. 16. Larva makes ophistigmatonome on leaves of Platanus

occidentalis with long linear tract and large blotch ... clemensella(p. 55)

Male genitalia with strongly palmate setae on dorsal surface of valves.

Hindwing of male with lance-shaped chitinous plate on costal margin. Female

genitalia not with above combination of characters ............... 6

. Male genitalia with strongly bilobed saccus; sharply pointed valves bearing

palmate setae with shaft approximately equal to or shorter than digitate pro-

cesses as in fig. 17c. Female genitalia with small signa, unequal in length and

very small spicules on lobe associated with colliculum as in fig. 18. Externals

occur in two forms as in figs. 54 and 55. Larva makes ophistigmatonome in

leaves oh Ouereusäspint sak in Geel dt ae AE APE similella (p. 56)

Male genitalia without strongly bilobed saccus; valves not sharply pointed and

bearing palmate setae with shaft much longer than digitate processes. Female

genitalia with signa approximately equal in length and large spicules on lobe

associated with colliculum . none NN ee 1

. Male genitalia variable, but as in fig. 19; some palmate setae on valves with

shaft up to three times length of digitate processes. Female genitalia with signa

short and narrow about half length of bursa copulatrix; short rows of spines on

lobe associated with colliculum, as in fig. 20c. Externals in two forms as in figs.

56 and 57. Larva makes ophionome in Corylus sp. leaves virgulae (p. 59)

Male genitalia as in fig. 13; palmate setae on valves with shaft approximately

twice length of digitate processes. Female genitalia with long, broad signa ex-

tending almost full length and covering most of bursa copulatrix; very large

single spines on the lobe associated with colliculum as in fig. 14c. Externals as

in fig. 53. Larva makes ophistigmatonome in leaves of Platanus ..........

IRENE ROER Le) MEA BR | ere) A eee e RA platanella (p. 51)

. Male genitalia with long triangular pseuduncus tapering to point; aedeagus as

in fig. 4 or 21. Female genitalia never with spiculate lobe associated with colli-

Culumetars ei vrank Mala ee 9

Male genitalia with pseuduncus not triangular and rounded at least terminally.

Female genitalia with spiculate lobe associated with colliculum ...... 10

. Male genitalia with dorsal arms of transtillae extending beyond saccus and

comparatively small anellar projections; as in fig. 21. Female genitalia with

thickened ring of colliculum and relatively long signa; as in fig. 22. Externals

as in fig. 43. Larva forms stigmatonome on Betula sp. .. lindquisti(p. 61)

Male genitalia with dorsal arms of transtillae broad and short, not extending

beyond saccus; large anellar projections; as in fig. 4. Female genitalia with

thin ring of colliculum and short signa; as in fig. 5. A petiole gall former on

Populus sp; ER EDDIE AR popullella (p. 41)

Male genitalia with small valves only about half length of capsule; anellus with

bifurcate spines totalling more than six, as in fig. 11. Female genitalia with

large single or double spines on spiculate lobe; signa broad and short; as in fig.

12. Externals as in fig. 52. Larva makes ophistigmatonome on Rosa sp. .....

14.

16.

WILKINSON & NEWTON: Ectoedemia in North America 39

SE inten Beten Ned ste Tee marmaropa (p. 49)

Male genitalia usually with valves larger than half capsule length; anellus with

four spines. Female genitalia with spiculate lobe not as above; signalong 11

. Male genitalia with large broad pseuduncus and anellus with four large spines

as in fig. 7b. Female genitalia with broad signa tapering distally as in fig. 8. Ex-

ternals as in fig. 44. Larva is probably a petiole gall-former on Populus

SPERI RE Ue REESE EP SAI, gine Lote. cei tera FA) canutus (p. 45)

Genitaliamotiasiabove! CH Oetker I ae e 12

. Male genitalia with highly characteristic pseuduncus and shape of anellar

spines and valves; as in fig. 9. Female genitalia with long narrow signa and

large spines on spiculate lobe of colliculum; as in fig. 10. Externals as in fig. 51.

Larva makes ophistigmatonome on Carya sp. ........ trinotata (p. 46)

Males non-existent. Parthenogenetic females with genitalia as in fig. 6. Larva

mines first in petiole and then makes stigmatonome in lamina of Populus

SPAREN RI MA N en ln a argyropeza downesi (p. 45)

Male genitalia with not more than four large, simple spines near phallotreme;

valves markedly incised along inner margin. Female genitalia without any

thickening or sclerotisation at colliculum; apophyses usually not markedly dif-

ferentindengthenssn, Certes ZEN Bell SEN NE RASTA 14

Male genitalia with more than four spines near phallotreme; valves not

markedly incised. Female genitalia with colliculum thickened; posterior

apophyses usually markedly longer than anteriores .............. 16

Male genitalia with saccus strongly concave; anterior arms of valves long; ae-

deagus with thickened circular band of cornuti; as in fig. 35. Female not

known ae soi tene er acanthella (p. 75)

Male genitalia not with above combination of characters. Female genitalia as

IMB DAO NE AI N STERN ee. o 15

. Forewings mottled, indistinct fascia, dull. Male genitalia with valves extending

as far as, or further than, extremity of pseuduncus; dorsal arms of transtillae

short as in fig. 26. Female genitalia as in fig. 27. Larva makes ophistigma-

tonomesinwleaves#oHaUlmusEsp re ulmella (p. 65)

Forewing with distinct fasciae, shining silver as in fig. 49. Male genitalia with

valves not extending beyond pseuduncus; dorsal arms of transtillae reaching

extremity of capsule as in fig. 23. Female genitalia as in fig. 24. Larva makes

ophistigmatonome in Rubus sp. leaves ............ rubifoliella (p. 63)

Forewings marked with four small silver patches as in fig. 48. Male genitalia

with valves extending well beyond pseuduncus; crown of thorn-like spines

present near phallotreme as well as large anellar spines; as in fig. 31. Female

genitalia with simple circular thickening at colliculum. Anterior apophyses

broad and unusually long reaching as far as posteriores as in fig. 32. Larva

makes ophionome with blotch later, in leaves of Ostrya, Carpinus, Corylus and

BEAR SPIRA EE I RE A ANETTA e: quadrinotata (p. 70)

Male genitalia with valves not extending far beyond pseuduncus; anellar

spines and cornuti characteristic, similar to figs. 33 and 37. Female genitalia

with complex thickened folds at colliculum. Anterior apophyses always much

shortenthan posteriorest CEL ii. GROSSO SO 17

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

. Male genitalia with valves just reaching end of pseuduncus; as in fig. 37.

Female genitaliaasinfigg38 Pam ea Nr ER eee A 18

Genitalia not as above

Larva mines the bark of Quercus sp. Externals as in fig. 61 ............

EN ROR AMAR PT Se ORR a ta SEIT heinrichi (p. 78)

Larva makes galls on twigs of Castanea. Externals as in fig. 62

castaneae (p. 80)

ee leje 'e (a ce 0). 0; ‘0; ce, er ce) ie) se; MORTO oat zer ORIO: ot an 0 OD eee em enen Oo O. LO

. Male genitalia without swellings or characteristic knobs on inner surface of

valves; as in fig. 36. Externals as in fig. 55. Female not known

E MORTAIO BIO (SE INA TOR TORO e RI CASE IT OE piperella (p. 77)

Male genitalia with characteristic knobs on inner surface of valves. Female

genitalia with spicules on colliculum 20

ee © © © © © © © © © © © © ee ee ee ee @

TABLE FOR THE IDENTIFICATION OF ECTOEDEMIA SPECIES FROM HOST PLANTS

AND DAMAGE

Ectoedemia host host site of larval form of larval probable Type

species genus species damage damage No. of locality

generations

populella Populus tremuloides petioles galls 1 —

grandidentata

argyropeza Populus _tremuloides Ist petioles ophionome 1, ? only Ontario

downesi ?grandidentata 2nd lamina stigmatonome

canutus Populus balsamifera petioles galls 117 Ontario

trinotata Carya cordiformis lamina ophistigmatonome 2 Ohio

marmaropa Rosa woodsii lamina ophistigmatonome - Utah

platanella Platanus occidentalis lamina ophistigmatonome 3 —

clemensella Platanus occidentalis lamina ophistigmatonome 3 Kentucky

similella Quercus palustris lamina ophistigmatonome up to 3 Ohio

rubra

virgulae Corylus americana lamina ophionome 2 Ohio

lindquisti Betula papyrifera lamina stigmatonome 1 Ontario

lutea

rubifoliella Rubus sp. lamina ophistigmatonome 2 Ohio

ulmella Ulmus rubra lamina ophistigmatonome 2 Ohio

americana

racemosa

nyssaefoliella Nyssa sylvatica lamina ophistigmatonome 2or3 Ohio

quadrinotata Carpinus caroliniana lamina ophistigmatonome 2 Ohio

Corylus americana

Ostrya virginiana

Betula lutea

obrutella Quercus? sp. bark? = = Texas

acanthella = — bark? — = New Jersey

piperella ~ = bark? — = Arkansas

heinrichi Quercus palustris bark, twigs spiral ophionome 1 Virginia

castaneae Castanea sp. bark, twigs galls 2 or 3 Virginia

phleophaga Castanea dentata bark, cambium ophionome 1 Virginia

chlorantis - — bark? — = Ontario

mesoloba ~ si bark? — Florida

For Amelanchier see nyssaefoliella description.

WILKINSON & NEWTON: Ectoedemia in North America 41

20. Male genitalia with simple rounded knob on inner side of valves; valves not

reaching beyond pseuduncus; as in fig. 42. Externals as in fig. 50. Female not

KR OW esha AA ee NE mesoloba (p. 85)

— Male genitalia different from above and valves extending beyond pseuduncus.

Onlyztemalesyknownsinone species ee 21

21. Genitalia as in figs. 33 and 34. Male genitalia with three small knobs on inner

side of valves. Anterior arms of valves long extending beyond capsule. Ex-

ternalsgasintigsAlee MR EME en. obrutella (p. 72)

== Gemtalaandexternalsnobas ADO EP EE RE CUT 22

22. Genitalia as in figs. 39 and 40. Male genitalia with characteristic knobs on

inner side of valves. Dorsal arms of transtillae short. Female genitalia with

comparatively simple ductus bursae. Externals in fig. 63. Larva makes a bark

ODHIONOMEMNICAS ATEN RE EEEN phleophaga (p. 81)

— Female genitalia as in fig. 41. Complex thickening in ductus bursae. Externals

ASINI PALO Male NOT KNOWN sn... chlorantis (p. 84)

THE POPULELLA GROUP

The North American Ectoedemia can be divided into four species groups. The

populella group contains the first five species. All males have a pointed pseuduncus

except argyropeza downesi where males are not known to occur. They lack the

multi-branched setae on the inner sides of the valves found in the platanella group.

All females have a sclerotised ring-shaped colliculum and associated denticulate

patch. E. lindquisti could belong in this group but it does not have the same charac-

teristic anellar processes and has perhaps more affinity with the rubifoliella group.

This group includes all the species which attack petioles: populella, argyropeza

and probably canutus, as well as the leaf miners trinotata and marmaropa.

Ectoedemia populella Busck

(figs. 4, 5)

Ectoedemia populella Busck, 1907: 98.

Ectoedemia populella Busck; Braun, 1917: 197.

Ectoedemia populella Busck; Braun in Forbes, 1923: 83.

Ectoedemia populella Busck; McDunnough, 1939: 107 (no. 9783).

Ectoedemia populella Busck; Borkowski, 1972: 697.

Ectoedemia populella Busck; Wilkinson & Scoble, 1979: 74.

Description. — Fully described from Canada by Wilkinson & Scoble (1979).

Diagnosis. — Usually larger than the other members of the genus studied here

and characterised by the cupreous reflections of forewings. Differs from canutus in

the valves, which are shorter and broader, and the gnathos, which is not bifid lat-

erally, in populella. The gall forming habit differentiates the larvae from those of

argyropeza which produce more irregular callus tissue.

Distribution. — USA: — New York, New Hampshire, Massachusets, Ohio

(Braun). Canada: — Ontario, Manitoba.

Material examined. — In USNM — USA: New York, Monroe Co.; on Populus

42 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

4a 2 4b

Fig. 4. Ectoedemia populella Busck, & (a) and 9 (b) genitalia.

sp.; 8 g, 2 9, 2 ex., 1—16.v.1948 (Kimball). “Awene, Man.”; 1 ©, 24.v.1921

(Criddle). In CNC — Canada: Ontario, Bells Corner; 2 9, 30.v.1965 (Sattler);

Lake Erie, Cultus; on Populus tremuloides; 1 &, 1.iii.1962; La Passe; on Populus sp;

1 g, 10.xii.1970; 1 3, 1.iii.1971 (Lewis); Minden, | g, 27.v.1931 (McDunnough).

In SOO — Ontario, Bradford; on Populus tremuloides, 1 3,2 9, 1 ex., 28.v.1970; on

Populus grandidentata, 2 3,5 9,3 ex., 28.v.1970; Nestor Falls; on Populus tremu-

loides; 1 &, 11.1.1963, 1 &, 7.iii.1963.

Galls examined: In FIS — Canada: Ontario, Bradford; several galls on Populus

grandidentata preserved in alcohol; 16.vii.1969, 695—2125—01; Uxbridge; several

galls on Populus tremuloides preserved in alcohol; 11.ix.1969, 675—4171—01.

Syntypes. | &, 1 ©, have also been examined from the collection in the

Transvaal museum. However a lectotype is not here designated. This will be done

WILKINSON & NEWTON: Ectoedemia in North America 43

CPAS

ZI? LEE

Fig. 5. Ectoedemia populella Busck, head, anterior (a) and posterior (b) view (x 70).

in “The Moths of North America north of Mexico” from the type-series in the

USNM.

Biology. — The larvae form galls on the leaf petioles of Populus tremuloides and

P. grandidentata. These are swellings situated close to the lamina. The larvae

mature in October and the imagines emerge in May of the following year (Busck,

1907; Braun, 1917). The specimens in the CNC that emerged earlier than this were

probably “forced” in the laboratory.

Voltinism. Univoltine in both Canada and the U.S.A.

Ectoedemia argyropeza (Zeller)

(fig. 6)

Lyonetia argyropeza Zeller, 1839: 215.

Nepticula argyropeza (Zeller); Zeller, 1848: 320.

Nepticula argyropeza (Zeller); Petersen, 1930: 78.

Nepticula argyropeza (Zeller); Hering, 1951: 232.

Nepticula turbidella Zeller sensu Herrich-Schäffer, 1853: 357; Downes, 1968: 1078.

Dechtiria argyropeza (Zeller); Beirne, 1945: 205.

Dechtiria argyropeza (Zeller); Emmet, 1971: 242.

Dechtiria argyropeza (Zeller); Borkowski, 1972: 698.

Stigmella argyropeza (Zeller); Borkowski, 1969: 107.

Stigmalla (sic) turbidella (Zeller) sensu Herrich-Schäffer, 1853: 357; Cochaux, 1969: 12.

Trifurcula (Ectoedemia) argyropeza (Zeller) sensu Johansson, 1971: 245.

Ectoedemia argyropeza (Zeller); Emmet, 1976: 189.

Ectoedemia argyropeza (Zeller); Wilkinson & Scoble, 1979: 77.

Description. — Fully described from Canada by Wilkinson & Scoble (1979) and

is regarded as being represented by two subspecies. The female genitalia of the

subordinate subspecies are shown in fig. 6.

44 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

Fig. 6. Ectoedemia argyropeza (Zeller), 9 genitalia.

Diagnosis. — Differs from canutus in the forewing ground colour, which is gen-

erally darker and less irrorate in argyropeza. The wider and less pointed ovipositor

distinguishes the genitalia from those of canutus.

Discussion. — Closely related to the following species, canutus, in both externals

and genitalia.

Ectoedemia argyropeza argyropeza (Zeller)

Diagnosis. — Differs from the subordinate subspecies in the paler forewing

ground colour. Only known in Europe whilst downesi is North American.

Material examined. — 9 Lectotype: “Gross Glogau, Silesia, Zeller 183: 101291,

WILKINSON & NEWTON: Ectoedemia in North America 45

Zeller Collection Walsingham Collection 1910—427, 101291, Argyropeza Zeller

1839: 215. Lyonetia A. Bucculatrix Isis 1839: 215”; in BM (NH).

Other specimens: In BM (NH) —Locality as lectotype; 1 9, 22.v.1853 (Zeller);

Walsingham Coll. ex Zeller Coll., 1910—427; 101289.

Ectoedemia argyropeza downesi Wilkinson & Scoble

Ectoedemia argyropeza downesi Wilkinson & Scoble, 1979: 80.

Diagnosis. — Differs from the nominate subspecies in the ground colour of the

forewings which are grey and heavily irrorate with purplish brown scales distally.

Occurs only as parthenogenetic females.

Distribution. — Canada: — Ontario, Quebec.

Material examined. — 9 Holotype: Canada: “Ontario, Ottawa”; on Populus

tremuloides; 29.v.1969 (Downes); in CNC. Paratypes: In CNC — data as holotype;

12 9, 29.v.1969, 5 ©, 28.v.1968, 2 9, 31.v.1968, 4 9, 4.vi.1968, 2 9, 7.vi.1968

(Downes). Ontario, Ottawa; on Populus sp.; 20 9, 6.v.1941 (Freeman); Ste. Anne

de Bellevue, P.Q., Morgan Arboretum; 4 9, 1.vi.1969 (Sheppard).

Biology. — Egg. Laid on the petioles of Populus tremuloides and P. grandidentata.

The number of eggs per leaf depends on leaf size and the position of the leaf on

the tree: eggs are more abundant on leaves at lower levels.

Mine. Begins as a petiole mine and later becomes of stigmatonome in the leaf

blade. Whilst a certain amount of callus tissue is formed in the petiole of Populus

tremuloides, in P. grandidentata this may, apparently, block the mine, preventing

the larvae from developing: the larvae seldom reach maturity on this host. A green

zone extends along the mined portions of the leaf-blade, even after the leaf has

turned yellow and fallen to the ground (‘‘green-islands” of Hering).

Larva. Remains in the mine until after leaf fall in late October and early No-

vember. Over-winters in ground litter or soil in cocoon.

Pupation. Occurs in early spring.

Voltinism. One generation per year.

Other notes. Cochaux (1969: 12) suggests that the very high populations of this

species may be due to the parthenogenetic reproduction.

Ectoedemia canutus Wilkinson & Scoble

(figs. 7, 8, 44)

Ectoedemia canutus Wilkinson & Scoble, 1979: 81.

Description. — Fully described by Wilkinson & Scoble (1979).

Diagnosis. — Generally darker than argyropeza argyropeza and more uniformly

irrorate than argyropeza downesi. The valves of the male canutus are larger and nar-

rower than in populella. It also differs in the bifid nature of the gnathos.

Distribution. — USA: — California. Canada: — Ontario.

Material examined. — & Holotype: Canada: Ontario, Angus; on Populus balsa-

mifera; 4.iii.1969, 68.5.3397.01; SOO no. 9; in SOO. Paratypes: In SOO — Canada:

data as in Holotype; 6 &, 3 9, 3—11.iii.1969; 5 g, 1 9, 2 ex., 28.v.1970,

705.0074.01.

46 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Figs. 7, 8. Ectoedemia canutus Wilkinson & Scoble, g (7) and © (8) genitalia.

Other material. — USA: 4 &, 2 ©, California, Los Angeles Co., Big Rock

Wash., near Lovejoy Buttes, TEN R9W Sec 30, elev 2770’, 31.iii.1972 (Donahue).

Biology. — Probably a gall former on the petioles of Populus balsamifera. Galls

examined: There are several galls on petioles, preserved in alcohol, which are in-

distinguishable from those of populella. The host and locality suggest that they may

belong to canutus. In SOO — Canada: Ontario, Angus; on Populus balsamifera;

3.1x.1968 (Bowser).

Ectoedemia trinotata (Braun) comb. nov.

(figs. 9, 10, 51, 66)

Nepticula trinotata Braun, 1914: 18.

WILKINSON & NEWTON: Ectoedemia in North America 47

Figs. 9, 10. Ectoedemia trinotata (Braun), 3 (9) and 9 (10) genitalia.

Nepticula trinotata Braun; Braun, 1917: 169.

Nepticula trinotata Braun; Braun in Forbes, 1923: 87.

Nepticula trinotata Braun; McDunnough, 1939: 107 (no. 9723).

Description. — External features: g. Head: palps whitish; antennae dark

brown; tuft on front of head ochreous, vertex paler; eye-caps shining white; collar

off-white. Thorax very dark grey-brown with purple and blue reflections. Ab-

domen grey, shining metallic grey beneath. Forewings: ground colour of dorsal

surface very dark brown, basal third with purple reflections, apical two thirds ir-

rorate, with base of each scale greyish; fringe whitish, shining silver, with apical

band of dark brown wing scales; small diffuse antemedial spot on costal margin

which is occasionally eclipsed by greyish reflections when viewed from certain

angles, followed by two distinct postmedial spots, shining silver, one at each

margin. Hindwings: ground colour and fringe brownish grey, shining metallic grey.

Legs dark brown, shining metallic grey behind. Fig. 51.

Female. As ¢ except for a pair of external convex pockets ventro-medially

placed on the fourth abdominal segment.

48 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

Wing expanse: &: 3.8—5.3 mm (6 specimens); 9: 4.4—5.4 mm (7). Holotype: 5.0

mm.

Genitalia: & (fig. 9). Pseuduncus with a single lobe as in fig. 9 (a). Gnathos: W-

shaped as in fig. 9 (a) with long medial process; dorso-lateral arms short. Vin-

culum: lateral arms broad; ventral plate very narrow. Saccus approximately five

times the width of the ventral plate, weakly bilobed. Valves not reaching beyond

the pseuduncus, quadrate and with a triangular inner process, arising distally.

Transtillae: an inverted U-shape as in fig. 9 (a); lateral arms broad and short;

ventral arms narrow, reaching just beyond ventral plate; transverse bars fused to

form a narrow arcuate strap. Aedeagus: regular in shape, approximately equal to

length of the capsule; vesica with cornuti as many small denticles evenly dis-

tributed and with plate of minute papillae in the shape of figure eight; anellus com-

prising a pair of lateral spines and a pair of large canine tooth-like spines.

Female (fig. 10). Ductus bursae short with colliculum as a sclerotised double

ring associated with a large lobe bearing scattered spicules as in fig. 10 (c). Ac-

cessory duct arising from area of dilation of the ductus, distally spiralled. Bursa

copulatrix: very large and covered with small scallop-shaped chains of pectinations

on striations of the bursa; signum double, comprising a pair of long cellular

patches, equal in area as in figs. 10 (a) and 10 (d). Anterior apophyses long and

broad. Posterior apophyses straight and broad, equal in length to the anteriores.

Host plant: Carya cordiformis (Hickory).

Mine: An upper surface ophistigmatonome.

Diagnosis. — Easily differentiated from other members of the genus in this

study by the forewing markings, the lobe of the pseuduncus and the form of the

transtillae. The mine differs from the serpentine tract produced by Stigmella jug-

landifoliella (Clemens) also found on Carya spp.

Discussion. — Although the externals resemble those of quadrinotata there is no

similarity in the genitalia. The form of the colliculum in the female relates this

species to several members of the genus.

Distribution. — USA: — Ohio, Arkansas, Illinois.

Material examined. — 9 Holotype: USA: “B.499; Cincinnati, Ohio, Annette F.

Braun, i, 3.viii.1913; Type Collection of Annette F. Braun; Nepticula trinotata

Braun Type”; on Carya sp.; slide no. 144-PJN; in ANS. Paratypes: In ANS data as

holotype; 3 ©, 1,2,3.viii.1913, 1 &, 1 ex., 31.vii.1913, 2 &, 8,11.viii.1913, B.499

(Braun). In USNM — same data as Holotype: 2 9, 6,15.viii.1913, B.499 (Braun);

slide nos: 145-PJN, 146-PJN, CNC 3480.

Other specimens: In ANS — USA: Ohio, Cincinnati; on Carya sp.; 1 9,

15.viii.1914, 1 g, 17.vi.1917, 1 &, 1 9, 10.vii.1917, B.499 (Braun). In USNM — Ar-

kansas, Washington County; I &, 10.vii.1966 (Hodges). Illinois, Putnam County; 2

3,19, 1ex., 21.vii.1962, 1.vi.1963, 6.viii.1964, 12.vii.1965.

Mines examined: In ANS — USA: Ohio, West Fork Woods; 4 mines on Carya

sp.; 15.vii.1909, B.499 (Braun).

Biology. — Egg. Laid on the upper or lower surface, usually near a vein.

Mine. At first a very narrow, much contorted linear mine which rapidly enlarges

into an extensive blotch. The frass is scattered throughout the mine (fig. 66).

Larva. Emerges from the mine on the upper surface of the leaf.

WILKINSON & NEWTON: Ectoedemia in North America 49

Pupa. Cocoon pale brown in colour.

Voltinism. Bivoltine in Ohio, adults emerging from the overwintering generation

in June and from the other generation in late July and early August.

Ectoedemia marmaropa (Braun) comb.n.

(figs. 11, 12, 52, 67)

Nepticula marmaropa Braun, 1925: 225.

Nepticula marmaropa Braun; McDunnough, 1939: 107 (no. 9778).

Description. — External features: &. Head: palps buff-grey; antennae grey; tuft

on front of head ochreous, vertex darker; eye-caps shining white; collar pale och-

reous. Thorax dark brown with gold and bronze reflections. Abdomen brown with

Figs. 11, 12. Ectoedemia marmaropa (Braun), & (11) and 9 (12) genitalia.

50 TIIDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

gold reflections above, shining metallic grey beneath. Forewings: ground colour of

dorsal surface dark brown with bronze and gold reflections, becoming irrorate dis-

tally, each scale being darker at the tip; fringe grey, shining silver at apex, with an

apical band of dark brown wing scales; basal patch extending to one-fifth, dusted

with grey, followed by a single postmedial fascia, convex and shining silver.

Hindwings: ground colour and fringe brownish grey, shining metallic grey. Legs

dark brownish with silver reflections on both surfaces. Fig. 52.

Female. As 4 except for a pair of external convex pockets ventro-medially

placed on the fourth abdominal segment.

Wing expanse: 9: 4.2—4.4 mm (3 specimens). Holotype ¢: 4.0 mm.

Genitalia: ¢ (fig. 11). Pseuduncus with a single tapering lobe, rounded termi-

nally. Gnathos: W-shaped as in fig. 11 (a); transverse ventral plate with a short

central boss; dorso-lateral arms long and thin. Vinculum: lateral arms broad;

ventral plate narrow. Saccus approximately twice width of ventral plate, bilobed.

Valves reach not much further than base of pseuduncus, quadrate with a short,

broad style and very broad cuiller. Transtillae: broadly W-shaped as in fig. 11 (a);

lateral arms narrow; ventral arms straight, reaching beyond the ventral plate;

transverse bars fused to form a narrow strap. Aedeagus: flask-shaped, just greater

than length of the capsule; vesica with cornuti as many small denticles evenly dis-

tributed and with a pair of large digitate cornuti at the phallotreme, also with a

cup-shaped plate of minute papillae; anellus comprising a pair of large plates with

bifurcate spines.

Female (fig. 12). Ductus bursae short with colliculum as a sclerotised double

ring associated with a small lobe bearing scattered spicules as in fig. 12 (c). Ac-

cessory duct arising from dilation of the ductus, spiral medially. Bursa copulatrix:

very large, covered with short chains of pectinations proximally; signum double,

comprising a pair of equal, ovate cellular patches as in figs 12 (a) and 12 (d). An-

terior apophyses long and arcuate. Posterior apophyses straight and narrow

reaching beyond the anteriores.

Host plant: Rosa woodsii.

Mine: An upper surface ophistigmatonome.

Diagnosis. — The lustrous basal patch of the forewings separates this species

from any other in the genus with a single fascia. Resembles lindguisti in the overall

form of the male genitalia but differs in the nature of the valves and the more

complex anellar processes in marmaropa. The aedeagus is larger than that of lind-

quisti.

Discussion. — Related to several members of the genus in the form of the

female genitalia.

Distribution. — USA: — Utah, Wyoming. Ohio?

Material examined. — & Holotype: USA: “B.1191; Cache Co., Utah,

i.23.iv.1925. Annette F. Braun; Type Collection of Annette F. Braun; Nepticula

marmaropa Braun Type’; on Rosa woodsii; slide no. 131-PJN; in ANS. Paratypes:

In ANS data as Holotype; 2 9, 3,17.iv.1925 (Braun); slide nos: 132-PJN, 133-PJN.

Other Specimens: In ANS — USA: Wyoming, Phelps Lake, Grand Teton Na-

tional Park; 1 9, 17.v.1935, B.1462 (Braun).

WILKINSON & NEWTON: Ectoedemia in North America 51

Mines Examined: In ANS — USA: Ohio; 2 mines on Rosa woodsii; no date,

B.1191 (Braun).

Biology. — Egg. Laid on the lower surface of the leaf and, in both cases ob-

served, next to the midrib.

Host. The specimen from Wyoming bears the A. F. Braun breeding number

B.1462; the record and the material cannot be located for identification of the host

plant.

Mine. Begins as a very narrow linear mine but abruptly enlarges into a blotch

which may consume half the area of the leaf. The frass is scattered throughout the

blotch (fig. 67).

Larva. Emerges on the upper surface of the leaf.

Pupa. Cocoon at first bluish green, later turning dark brown in colour.

THE PLATANELLA GROUP

The following four species — platanella, clemensella, virgulae and similella — can

be separated to form a discrete group, when this is seen to be an advantage.

The males have the same tapering pseuduncus and W-shaped gnathos of the last

group; the aedeagus has very pronounced anellar lobes. The most notable char-

acter is the presence of multi-branched setae on the inner surface of the valves —

more weakly developed in clemensella.

The colliculum is present as a single or double sclerotised ring in females.

Externally there is a lance-shaped chitinous plate along the fore-edge of the hind

wing in the males, except in clemensella. Another minor feature is the forewing

marking. All have a biconcave fascia reduced in some to a pair of marginal

streaks. All are leaf-miners.

Ectoedemia platanella (Clemens)

(figs. 13, 14, 53, 68)

Nepticula platanella Clemens, 1861: 83.

Nepticula platanella Clemens; Clemens, 1862a: 133.

Nepticula plantanella (sic) Clemens; Clemens, 1862b: 149, 150.

Nepticula platanella Clemens; Clemens, 1865: 146.

Nepticula platanella Clemens; Clemens in Stainton, 1872: 173, 183, 192.

Nepticula platanella Clemens; Chambers, 1873: 125.

Nepticula platanella Clemens; Chambers in Hayden, 1878b: 158.

Nepticula platanella Clemens; Dyar, 1903: 546.

Nepticula platanella Clemens; Busck, 1903: 209.

Nepticula platanella CLemens; Braun, 1917: 187.

Nepticula platanella Clemens; Braun in Forbes, 1923: 92.

Nepticula platanella Clemens; McDunnough, 1939: 107 (no. 9760).

Ectoedemia platanella (Clemens); Wilkinson & Scoble, 1979: 89.

Nepticula maximella Chambers, 1873: 126 (syn. by Braun, 1917: 187).

Nepticula maximella Chambers; Chambers in Hayden, 1878b: 158.

Nepticula maximella Chambers; Dyar, 1903: 546.

Description. — External features: g. Head: palps buff; antennae dark brown;

tuft on front of head pale ochreous, vertex darker; eye-caps shining white; collar

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

pale ochreous. Thorax and abdomen dark greyish brown. Forewings: ground

colour of dorsal surface dark brown with bronze lustre; fringe whitish, with an

apical band of dark brown wing-scales; a pair of white, medial streaks marginally,

which are occassionally linked in the middle by a few white scales to form a com-

plete fascia. Hindwings: ground colour and fringe pale grey; a lance-shaped

chitinous plate extends along the fore edge to the middle of the costa. Legs buff,

with areas of dark brown. Fig. 53.

Female. As ¢ except that chitinous plate of hindwings is absent. With a pair of

convex, external pockets ventro-medially placed on the fourth abdominal

segment.

Figs. 13, 14. Ectoedemia platanella (Clemens), & (13) and 9 (14) genitalia.

WILKINSON & NEWTON: Ectoedemia in North America 53

Wing expanse: G': 4.0—6.8 mm (25 specimens); 9: 3.6—6.8 mm (23). Neotype:

6.0 mm. |

Genitalia: & (fig. 13). Pseuduncus with a single tapering lobe. Gnathos: W-

shaped as in fig. 13 (a); transverse ventral plate with broad central boss; dorso-

lateral arms long. Vinculum: lateral arms narrow; ventral plate narrow. Saccus

more than twice the width of the ventral plate and weakly bilobed. Valves: not

reaching beyond the pseyduncus, tapering slightly; with terminally digitate setae

on the dorsal surface distally as in fig. 13 (c). Transtillae: with short, narrow lateral

arms; ventral arms reaching beyond the ventral plate; transverse bars fused to

form a continuous narrow strap. Aedeagus: approximately equal to length of the

capsule, regular in width; vesica with cornuti as many small denticles evenly dis-

tributed and with a comma-shaped plate of minute papillae; anellus comprising

two pairs of simple spines as in fig. 13 (b).

Female (fig. 14). Ductus bursae long, colliculum as a sclerotised double ring ring

with serrate inner margin, as in fig. 14 (b), and with associated lobe bearing scat-

tered spicules as in fig. 14 (c). Accessory duct arising from area of dilation of the

ductus, spiral distally. Bursa copulatrix very large and covered with small scallop-

shaped chains of pectinations on striations of the bursa; signum double, com-

prising a pair of long reticulate patches, equal in area as in figs 14 (a) and (d). An-

terior apophyses long and broad. Posterior apophyses straight and narrow, equal

in length to the anteriores.

Host plant: Platanus occidentalis.

Mine: An upper surface ophistigmatonome.

Diagnosis. — Usually larger and browner than clemensella and the males have

the hindwing scale which is absent in clemensella. The linear portion of the mine is

longer in clemensella and the enlargement into the blotch more gradual than in pla-

tanella. The male genitalia differ in the nature of the saccus, which is markedly

bilobed, and the scales of the valves, which are shorter and less furcate, in clemen-

sella. The female genitalia may be separated from those of similella and virgulae by

the spines of the accessory sac, which are large and single, in platanella.

Discussion. — Clemens (1861: 83) originally described this species from mines

found on the “Button-Wood Tree or Sycamore”. In 1862 he described an adult

caught at light which he gave the same name and labelled as type. The original

mine(s) described actually constitute the type(s) but they have never been men-

tioned in print and are no longer extant. To resolve this problem a neotype is here

designated. It is the specimen described by Clemens and also Busck (1903: 209).

Chambers (1873: 126) described adults which he named maximella and associated

them with a mine earlier described by Clemens (1862b: 149) as Sycamore miner

no. 3 and thought to be the product of a different species. In spite of this, Braun

(1917: 187) later synonymised maximella with platanella although she gave no

reasons. The mine of maximella is characteristic in the quadrate nature of the ter-

minal blotch as opposed to the round blotch of platanella material. Examination of

the genitalia shows that the male syntype of maximella lacks the bifurcate anellar

process and the saccus is less markedly bilobed than in examples of platanella.

However, this preparation is distorted so we do not regard the differences ob-

served to be sufficient to withdraw maximella from synonymy at this stage.

54 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Distribution. — USA: — Ohio, New Jersey, Washington D.C., Pennsylvania,

Kentucky, Massachusetts, Florida, Alabama. Canada: — Ontario.

Material examined. — Sesignated as neotype the 9 examined by Busck (1903):

USA: “Type no. 7494, Nepticula platanella B. Clemens, Specimen no. 118, A.B.

1903”; slide no. CNC 3464; in ANS. Syntypes of maximella: 1 &, “2) Nepticula

maximella Cham.”; Type no. 525 USNM; slide no. CNC 3477; 1 9, “2) Cham.

Type”; slide no. CNC 3502; in USNM.

Other Specimens: In ANS — USA: Ohio, Cincinnati; on Platanus sp.; 1 ©,

12.vii.1909, 2 &, 13.vii.1909, 1 &, 29.vii.1909, 1 9, 4.vii.1911, 1 g, 1 9,5.vii.1911,

1 9, 29.vii.1913, B.348 (Braun); 1 ©, 30.vii.1903 (Braun). Ohio, Cincinnati; 5 g, 2

©, 27.vi.1916 (Braun); 1 g, “Lot 72”, (Heimbach). No locality data; 3 Q1, “2,

Nepticula maximella Cham = N. platanella Clem”. In USNM — Ohio, Cincinnati;

on Platanus sp.; 1 ©, 10.vii.1909, B.348 (Braun). Ohio, Cincinnati; 1 &, 31.vii.1903,

1 &, 4.viii.1903, 1 ©, 22.v.1906, 5 &, 1 9, 6 ex., 27.vi.1916 (Braun). No locality

data; on Platanus sp.; 1 &, 17.vi.1885, 1 ©, 5.v.1885 (Murfeldt); 1 3, “1) N. plata-

nella Clem. Ch.”; 1 ©, 5.vi.1885 (Fernald Coll.). Washington, D.C.; on Platanus

sp.; Aug. 1902 (Busck); 12 ex., 15.vii.1908. Massachusetts, Barnstaple; 1 ex.,

28.vii.1952 (Kimball). Alabama, Open Pond Cpgd., S. Andalusia, Covington Co.; 1

d, 18.iv.1976 (Heppner). Florida, Oneco, Manatee Co.; 1 g, 3 ex., 5—14.v.1953,

7.vi.1953 (Dillman). Siesta Key, Sarasota Co.; 2 ex., 30.iv. + 1.vi.1957 (Kimball).

Pensacola; 1 9, 7.ix.1961 (Hills). Near Gainsville, Alachua Co., Archer Road

Lab.; 1 g, 3 ex., 13.vi.1975, 4—5.v.1976 (Heppner); Gainsville; 1 ex., 11.x1.1974

(Greenbaum). Archbold Biol. St., Lake Placid, Highlands Co.; 2 g,1 9,6 ex., I

8.v.1975 (Heppner). In CNC — Canada: Ontario, Normandale; on Quercus rubra; |

3, 16.vi.1962 (Freeman and Lewis).

Mines examined: In ANS — USA: Ohio, Madisonville Swamp; 1 mine on Pla-

tanus occidentalis; 7.ix.1909, B.561!) (Braun).

Biology. — Egg. Laid on the lower surface of the leaf, sometimes near a vein.

Host. Braun reports that the mines of this species are abundant on the leaves of

Platanus spp. and that adults may be found in large numbers on the trunks of the

host. The single male bred from Quercus rubra is the only record of the species

from that host.

Mine. Begins as a much contorted linear mine which is usually filled with frass.

Several days before pupation, the mine abruptly enlarges into an almost circular

blotch, which often eclipses the earlier portion of the mine. The frass is scattered

evenly throughout the blotch (fig. 68).

Larva. Pale green in colour, emerges on the upper surface of the leaf.

Pupa. The cocoon is brown in colour.

Voltinism. Braun (1917) reports that there are three generations per year; the

material examined here shows specimens to have emerged from early May to late

July and reaching a peak in late June and early July. One specimen from Florida is

dated November.

!) Some specimens of Stigmella apicialbella comb. n. were incorrectly labelled by Braun with this same

breeding number.

WILKINSON & NEWTON: Ectoedemia in North America 55

Ectoedemia clemensella (Chambers)

(figs. 15, 16)

Nepticula clemensella Chambers, 1873: 125.

Nepticula clemensella Chambers; Chambers in Hayden, 1878b: 157.

Nepticula clemensella Chambers; Dyar, 1903: 545.

Nepticula clemensella Chambers; Braun, 1917: 188.

Nepticula clemensella Chambers; Braun in Forbes, 1923: 93.

Nepticula clemensella Chambers; McDunnough, 1939: 107 (no. 9761).

Ectoedemia clemensella (Chambers); Wilkinson & Scoble, 1979: 86.

Description. — Fully described from Canada by Wilkinson and Scoble (1979).

Diagnosis. — Generally smaller and paler in colour than platanella; the males

lack the chitinous plate of the hindwing which is found in platanella, similella and

15 16

Figs. 15, 16. Ectoedemia clemensella (Chambers), g (15) and 9 (16) genitalia.

56 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

virgulae. The setae of the valves are smaller and less furcate than those of any

other species in the group and the saccus more markedly bilobed than in similella.

The female genitalia are very similar to those of platanella.

Discussion. — Material has been examined from New York State; a locality not

previously cited.

Distribution. — USA: — Ohio, Maryland, New York. Pennsylvania, Kentucky,

Florida, Maine. Canada: — Ontario.

Material examined. — 9 Lectotype: USA: Kentucky, (Chambers); slide no.

CNC 3490; in MCZ. Paralectotype: In MCZ — 1 ex., “Peak Acad”, “Type no.

14955”.

Other specimens: In ANS — USA: Ohio, Cincinnati; on Platanus sp.; 2 &,

1.viii.1902, 1 g, 25.vii.1908, 3 ©, 8,25,26.vii.1912, 1 &, 26.vi.1916, 1 2, 27.vi.1916,

1 g, 13.vii.1916 (Braun). In USNM — USA: Maryland, Plummers Island; 1 ex.,

v.1906 (Busch). Florida, Gainsville; on Platanus occidentalis; 3 3,7 2,8 ex., 11 +

14.v.1964 (Denmark); 1 ex., 5.viii.1963 (Denmark); 1 ex., 26.viii.1966 (O’Berry);

Archbold Biol. Sta., Lake Placid, Highlands Co; 1 ex., 1.v.1975 (Heppner). Maine,

Bar Harbour; 1 9, 27.v.1936 (Brower). In CNC — New York, Cornell University;

1 ©, 1 ex., 6.iv.1885 (Murfeldt). Canada: Ontario, St. Williams; continue over on

Platanus occidentalis; 3 3,29, 16—26.ii.1960 (Freeman and Lewis). In SOO — On-

tario, St. Williams; on Platanus sp.; 1 3°, 20.iii.1962.

Mines examined: In ANS — USA: Ohio, Cincinnati; 2 mines on Platanus sp.;

26.vi.1909, B.452 (Braun); 7.ix.1909, B.562 (Braun). In CNC — Canada: Ontario,

St. Williams; 19 mines on Platanus occidentalis; 3.ix.1959, 59—152, 59—153

(Lewis). In FIS — Ontario, St. Williams; several larvae, pupae and mines pre-

served in alcohol, 25.viii.1961, 561—7153—01.

Biology. — Mine. An upper surface ophistigmatonome in Platanus occidentalis

comprising long linear tract terminating in small blotch. The frass line is central in

the ophionome but absent in most of the blotch.

Voltinism. Reported as trivoltine (Braun, 1917).

Ectoedemia similella (Braun) comb.n.

(figs. 2, 17, 18, 54, 55, 69)

Nepticula similella Braun, 1917: 188.

Nepticula similella Braun; Braun in Forbes, 1923: 93.

Nepticula similella Braun; McDunnough, 1939: 107 (no. 9762).

Description. — External features: G 9. As platanella except that: tuft on front of

head orange ochreous, vertex paler. Ohio form: Forewings: ground colour of

dorsal surface brown with scattered blue and purple reflections, distal half with

each scale darker at the tip; single medial fascia, oblique, shining silver and occa-

sionally reduced to marginal streaks. Fig. 54. Florida form: Forewings: ground

colour very dark brown to black and uniformly dusted with grey reflections;

markings barely visible as marginal patches of not more than five whitish scales;

extreme apex of fringe creamy white, giving impression of an apical spot. Fig. 55.')

Wing expanse: &: 4.6—5.6 mm (22 specimens); © 4.2—6.0 mm (16). Holotype:

52 mm}:

1) See footnote page 58.

WILKINSON & NEWTON: Ectoedemia in North America 57

pA oy €

an cots 5

ian

4 JA È;

DICH

Hayy”

ose.

; eve”.

eared!

esas ts

(a)

Figs. 17, 18. Ectoedemia similella (Braun), & (17) and 9 (18) genitalia.

Genitalia: & (fig. 17). As platanella except that saccus is markedly bilobed, each

lobe being as long as broad at base. Valves: rounded distally with small apical

style; setae of dorsal surface broad and fan-like as in fig. 17 (c). Aedeagus:

markedly shorter than the capsule; anellus comprising a pair of blunted, digitate

processes.

Female (fig. 18). As platanella except that sclerotised double ring with inner

margin weakly serrate, as in fig. 18 (b) and lobe of ductus with small triple spicules

as in fig. 18 (c). Bursa copulatrix: signa small in relation to the bursa and unequal

in size.

Host plant: Quercus palustris, Quercus rubra.

Mine: An ophistigmatonome.

Diagnosis. — More often smaller and more iridescent than platanella and clem-

ensella and differs in the irrorate nature of the forewing ground colour. This

species has the lance-shaped chitinous plate on the male hindwing which is absent

in clemensella. The saccus is more markedly bilobed than in platanella and virgulae

but less than in clemensella. The broad, fan-like scales of the valves differ from

those of platanella, clemensella and virgulae. The female genitalia resemble those of

58 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

the other members of the group in most respects: the unequal size of the signa and

the very small spicules of the lobe are the only consistent differences. The mine is

easily separated from the lower surface ophionome produced by Stigmella altella

(Braun) comb.n., which is also found on Quercus palustris.

Discussion. — The pattern of wing markings, although resembling that of other

members of the species group, is most variable in similella. There are two forms:

those with a complete fascia which is occasionally reduced to marginal streaks as

in platanellaa and clemensella, which is found in Ohio, New Jersey and Virginia and

referred to here as the Ohio form; and those with greatly reduced marginal

patches and a diffuse apical patch of off-white ciliae, which is known only from

Florida.!) The Ohio form is that originally described by Braun (/oc.cit) from ten

bred specimens on Quercus palustris in Ohio. There is a single female (Specimen 6)

from Arkansas, with externals intermediate between the two forms: the forewings

are dark chocolate brown with an apical patch of cream, as in the Florida form,

and there is a single medial fascia, as in the Ohio form.

Distribution. — USA: — Ohio, New Jersey, Illinois, Virginia, Florida, Arkansas,

Kentucky, Maine?

Material examined. — ¢ Holotype: USA: “B.649; Cincinnati, Ohio. Annette F.

Braun, i. 21.vi.1914; Type Collection of Annette F. Braun; Nepticula similella

Braun Holotype”; on Quercus palustris; slide no. 109-PJN; in ANS. Paratypes: In

ANS — USA: Ohio, Cincinnati; on Quercus palustris; 1 g, 18.vi.1914, 1 g,

22.vi.1914, 1 &, 2 9, 24.vi.1914, 1 9, 27.vi.1914, 1 9, 30.vi.1914, 1 9, 7.vii.1914,

B.649 (Braun). In DFF — Ohio, Cincinnati; on Quercus palustris; 1 9, 22.vi.1914,

B.649 (Braun); slide nos. 107-PJN, 108-PJN, CNC 3479.

Other specimens: In ANS — USA: Ohio, Cincinnati; on Quercus palustris; 1 3,

2.vi.1922, 2 9, 17.vi.1922, 1 g, 3 9, 21.vi.1922, 1 9, 28.vi.1922, B.649 (Braun); on

Quercus rubra; 1 9, 10.vi.1918, B.965 (Braun). In USNM — New Jersey, Anglesea,

1 ©, “V.30” (Kearfott). Virginia, Mountain Lake; 1 g, 24.vii.1940 (Milne and

Milne). Illinois, Putnam Co.; 2 &, 14.vii.1969, 23.vii.1970 (Glenn). Maine, Milli-

nocket; 1 ex., 11.vii.? Florida, Archbold Bio. Sta., Lake Placid; 1 &, 1—7.v.1964, 1

3, 2 9, 8—15.v.1964, 3 g, 16—22.v.1964 (Hodges). Oneco, Manatee Co.; 8 g,

5—14.v.1953, 3.ix.1953, 2.xi.1953 and 17.xi.1954 (Dillman). Gulf Coast Exp. Sta.,

Bradenton; 1 &',24.ii.1955 (Keisheimer).

Specimen 6: In USNM — USA: Arkansas, Devil’s Den St. Pk., Washington

County; 1 9, 24.vi.1966 (Hodges); al. ex. 5.6 mm; slide no USNM 17256.

Mines examined: In ANS — USA: Ohio, Cincinnati; 2 mines on Quercus pal-

ustris; 30.viii.1911, B.649 (Braun); 3 mines on Quercus rubra; 21.x.1917, 8.965

(Braun).

Biology. — Egg. Laid on the upper surface.

Host. Braun (1917 and breeding records) reports that similar mines have been

!) Footnote Post Scriptum: Since the description of this species was written, more material has become

available from Florida. Some of these specimens have complete fasciae, whilst retaining other char-

acters of the dark form. It still remains that Florida is the only state recording the form with incomplete

fasciae, but it now seems unlikely that the variation is geographically based as was first thought

probable (CW).

WILKINSON & NEWTON: Ectoedemia in North America 59

found on chestnut in Kentucky although the material in question cannot be lo-

cated.

Mine. The early part of the mine is very narrow and much contorted in close S-

shaped curves, with the frass scattered throughout its breadth. The larva then

forms a lower surface blotch in which the frass is deposited as a congealed mass at

the beginning (fig. 69).

Larva. Pale green in colour, emerging on the lower surface in most cases.

Pupa. Cocoon ochreous in colour.

Voltinism. Adults have been taken in May, throughout June and early July,

while late instar larvae are found from July to October. It is possible that there are

up to three broods per year.

Ectoedemia virgulae (Braun) comb.n.

(ps M9 2065770)

Nepticula virgulae Braun, 1927: 198.

Nepticula virgulae Braun; McDunnough, 1939: 107 (no: 9751).

Description. — External features: ¢ ©. As platanella except that tuft on front of

head orange-ochre, vertex paler. Thorax and abdomen dark brown with bronze re-

flections. Forewings: ground colour of dorsal surface brown to very dark choc-

olate brown with bronze and purple reflections, distally each scale darker at the

tip; single medial fascia, variable in breadth, oblique, shining silver and markedly

concave inwardly. Figs. 56 (@ Ohio) and 57 (4 Florida).

Wing expanse: 4: 3.8—5.0 mm (12 specimens); 9: 4.0—5.8 mm (19). Holotype:

4.0 mm.

Genitalia: & (fig. 19). As platanella except that valves markedly concave along

inner margin distally; setae of dorsal surface terminally digitate, with long basal

shaft as in fig. 19 (c). Aedeagus: markedly shorter than the capsule; anellus com-

prising a pair of bifurcate plates as in fig. 19 (b).

Female (fig. 20). As platanella except that sclerotised double ring with inner

margin markedly serrate, as in fig. 20 (b), and lobe of ductus with large triple

spines as in fig. 20 (c). Bursa copulatrix: signa small in relation to the bursa.

Variation. In externals, the ground colour of the forewings varies in intensity

from the pale form in Ohio to dark chocolate brown in Florida and Texas. The

breadth of the fascia varies and is broadest in those specimens examined from

Florida. Similarly, there is a greater degree of variation in the male genitalia than

is found in other members of the species group. The pseuduncus ranges in length

and shape from the short, blunted form in Florida to a longer more tapering form

in Ohio and Texas (fig. 19). The setae of the valves are more heavily sclerotised

and longer in Florida males than in those from Ohio and Texas illustrated in fig. 19

(©)

Host plant: Corylus americana.

Mine: An upper surface ophionome.

Diagnosis. — More often smaller than platanella or clemensella and differs in the

irrorations and reflections of the forewings; as far as observed the fascia is always

complete in virgulae. The saccus is more weakly bilobed and the valves more

60 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

Figs. 19, 20. Ectoedemia virgulae (Braun), & (19) and 9 (20) genitalia.

concave along the inner margin than in similella or clemensella. The setae of the

valves differ from those of platanella, clemensella and similella in the longer shaft

and the longer terminal ciliae. The female genitalia resemble those of the other

members of the group; the large, triple spines of the accessory lobe and the

smaller signa separate them from platanella. The mine is broader than that of Stig-

mella corylifoliella (Clemens), but narrower than Stigmella ostryaefoliella

(Clemens), both of which are also found on Corylus sp.

Discussion. — Originally described from four bred specimens on Hazel (Corylus

americana). The variation in externals of the material examined here shows a

similar geographical distribution to that in similella: a darker form being recorded

in the southern states (Florida and Texas). In virgulae, however, this also corres-

ponds with a difference in the male genitalia of the Florida specimens. In the two

males from Texas the transtillae are much stouter than in the type; this may be due

WILKINSON & NEWTON: Ectoedemia in North America 61

to distortion but is possibly a geographical or seasonal difference (these adults

were on the wing in September).

Distribution. — USA:— Ohio, Texas, Florida.

Material examined. — ¢ Holotype: USA: “B.1107, Clinton County, Ohio,

24.iv.1925; Type; Nepticula virgulae Braun type”; on Corylus americana; slide no.

100-PJN; in ANS. 9 Allotype: same data as Holotype; i.15.v.1925; slide no. 102-

PJN; in ANS. Paratypes: In ANS—USA: same data as Holotype; Ohio, Clermont

County; 1 &, 1 9, vii.1924; slide nos. 101-PJN, 103-PJN.

Other specimens: In USNM—USA: Texas, Bexar County; 2 g, 3 9, 11.ix.1959

(McGregor). Florida, Lake Placid, Archbold Bio. Sta.; 6 ¢, 5 9, 8—15.v.1964, 3

d, 7 9, 16—22.v.1964 (Hodges). Florida, Fisheating Creek, Palmdale; 2 9, 7—

10.v.1964 (Hodges).

Biology. — Mine. A gradually broadening linear mine with the frass scattered in

a broad tract (fig. 70).

Larva. Green in colour.

Pupa. Cocoon reddish brown in colour.

Voltinism. Possibly bivoltine in Ohio, with adults emerging in April, May and

July; Braun’s breeding records show that late instar larvae are present in August

and September. Adults have also been taken in May from Florida and in Sep-

tember from Texas.

THE RUBIFOLIELLA GROUP

This group has three species, rubifoliella, ulmella and quadrinotata with the pseu-

duncus prolonged backwards but bluntly rounded; the aedeagi have moderately

large, similar, anellar projections in males, and females are without thickened or

sclerotised collicula. The species nyssaefoliella is assigned here on the grounds that

the female is typical, having no colliculum. It has unusually broad and enlarged

anterior apophyses. The male however is more typical of the populella group with

the pointed pseuduncus. However the aedeagus (like specimen 8) is devoid of any

large spines or anellar projections. Another species which does not conveniently

fit into a species-group is lindquisti which in the male looks like quadrinotata but

with smaller anellar spines and more pointed pseuduncus. It also has similarities

with the preceding groups as indicated in the species diagnosis. The female has the

sclerotised ring-shaped colliculum reminiscent of populella.

Ectoedemia lindquisti (Freeman)

(figs. 21, 22, 43)

Nepticula lindquisti Freeman, 1962: 522.

Nepticula lindquisti Freeman; Lindquist, 1962; 524.

Ectoedemia lindquisti (Freeman); Wilkinson & Scoble 1979: 83.

Description. — Fully described from Canada by Wilkinson & Scoble (1979).

Diagnosis. — The darker tufts on the head and the generally darker ground

colour separate lindquisti from the members of the platanella group. Differs from

marmaropa in the absence of the basal patch of the forewings. The genitalia re-

62 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Figs. 21, 22. Ectoedemia lindquisti (Freeman), & (21) and 9 (22) genitalia.

semble those of the platanella group but differ in the absence of the setae of the

valves, in males, and the lobe of the colliculum, in females. The digitate processes

of the aedeagus are less complex than those in similella and marmaropa (compare

fig. 21 with figs. 17 and 11).

Discussion. — Additional specimens to those cited by Wilkinson & Scoble

(1979), from Maine indicate that the species has a wider distribution than has been

previously cited. One male has darker head tufts than is usual for the species, but

is otherwise indistinguishable. See also discussion for canadensis, p.

Distribution. — USA:— Maine. Canada:— Ontario.

Material examined. — & Holotype: Canada. Ontario, Lake Huron, Wiarton; on

Betula papyrifera; 13.vi.1959, 559.0049.01; Type no.: 7752; in CNC. Paratypes: In

CNC—Canada: data as Holotype; 26 g, 12 9, 8.vi.—1.viii.1959.

Other specimens: In ANS—USA: Maine, Bethel; 2 3, 1 ©, 29.vi.1946 (Braun).

WILKINSON & NEWTON: Ectoedemia in North America 63

Canada: data as Holotype; 1 ©, 8.vi.1959. In SOO—Ontario, Trout Creek; on

Betula alleghaniensis; 1 ©, 24.111.1958.

Biology. — A thorough study has been made by Lindquist (1962) for whom the

species was named by Freeman.

Egg. Laid on underside of Betula leaves (B. papyrifera, B. alleghaniensis = lutea),

sometimes in large numbers on one leaf.

Mine. Is an upper surface stigmatonome with a more or less rectangular blotch

between two leaf veins. Frass is deposited in circular patches.

Larva. Whitish in colour with translucent integument and pale yellow head.

Cocoon. Yellow-brown darkening with time and deposited in soil and litter.

Larva overwinters in cocoon.

Voltinism. Univoltine.

Ectoedemia rubifoliella (Clemens)

(figs. 23, 24, 49)

Nepticula rubifoliella Clemens, 1860: 214.

Nepticula rubifoliella Clemens; Clemens, 1865: 146.

Nepticula rubifoliella Clemens; Clemens in Stainton, 1872: 32, 42, 45, 152.

Nepticula rubifoliella Clemens; Chambers in Hayden, 1878b: 158.

Nepticula rubifoliella Clemens; Dyar, 1903: 547.

Nepticula rubifoliella Clemens; Busck, 1903: 208.

Nepticula rubifoliella Clemens; Braun, 1917: 183.

Nepticula rubifoliella Clemens; Braun in Forbes, 1923: 91.

Nepticula rubifoliella Clemens; McDunnough, 1939: 107 (no: 9750).

Ectoedemia rubifoliella (Clemens); Wilkinson & Scoble, 1979: 90.

Description. — Fully described from Canada by Wilkinson & Scoble (1979).

Diagnosis. — Resembles virgulae in externals, although rubifoliella is generally

darker and the fascia broader. The males lack the chitinous plate of the hindwings

found in virgulae. The nature of the pseuduncus and the relatively short aedeagus

separates the male genitalia from those of ulmella and quadrinotata, whilst the fe-

males differ only in the relative sizes of the signa.

Distribution. — USA:— Ohio, Kentucky, Pennsylvania (Clemens). Canada:—

Ontario, Quebec.

Material examined. — & Neotype: USA: Ohio, Cincinnati; on Rubus sp.;

23.v.1916 (Braun); slide no: USNM 17339; in ANS.

Other specimens: In USNM — USA: data as neotype; 2 9, 18.viii.1912,

1.vi.1916 (Braun). Kentucky, Red Bird River; 1 g, 19.viii.1933 (Braun). In CNC —

Canada: Ontario, Simcoe; on Rubus sp.; 1 3, 14.11.1966 (Freeman). Quebec, Hull;

on Rubus sp.; 2 9, 21.iii.1959 (Freeman and Lewis); | ex., 31.iii.1957 (Freeman).

Mines examined: In ANS — USA: Ohio, West Fork Woods; I mine on Rubus sp.;

6.ix.1909, B.558 (Braun); 1 mine on Rubus sp.; 4.vii.1909, B.558 (Braun). In CNC

— Canada: Ontario, Simcoe; 5 mines on Rubus sp.; 1965, 65—74 (Freeman).

Quebec, Hull; 3 mines on Rubus sp.; 1965, 56—257 (Freeman and Lewis).

There is a single male with externals and genitalia similar to rubifoliella except

that the ground colour is paler and the pseuduncus, valves and gnathos differ as in

fig. 25. According to the label data this specimen was reared from Quercus plata-

64 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

noides (the same series of mines from which Braun reared examples of Stigmella

flavipedella (Braun)). If this is correct then the mine differs from that of rubifoliella

in that it is a much contorted linear tract.

Specimen 7: In ANS — USA: Ohio, Cincinnati; on Quercus platanoides; | 3,

30.v.1922, B.653 (Braun); al.ex. 4.4 mm.; slide no: 130-PJN.

Figs. 23, 24. Ectoedemia rubifoliella (Clemens), & (23) and 9 (24) genitalia.

WILKINSON & NEWTON: Ectoedemia in North America 65

25 Sm —

Fig. 25. Ectoedemia, specimen 7, ¢ genitalia.

Biology. — Mine. A linear tract is made in Rubus (Blackberry) leaves and the

frass-line is broken and central. It then widens into a rather elongate blotch

forming an upper surface ophistigmatonome with the frass irregularly dispersed.

Pupa. Cocoon is dark brown.

Voltinism. Braun (1917) reports 2 generations.

Ectoemedia ulmella (Braun)

(figs. 26, 27)

Nepticula ulmella Braun, 1912: 87.

Nepticula ulmella Braun; Braun, 1917: 186.

Nepticula ulmella Braun; Braun in Forbes, 1923: 92.

Nepticula ulmella Braun; McDunnough, 1939: 107 (no: 9758).

Ectoedemia ulmella (Braun); Wilkinson & Scoble, 1979: 91.

Description. — Fully described from Canada by Wilkinson & Scoble (1979).

Diagnosis. — Larger and less iridescent than rubifoliella. The tapering valves

and the broad gnathos separate the males from those of rubifoliella, while the only

consistent difference in the females is the smaller and more ovate signa retinacula

of ulmella.

Discussion. — We have examined an additional male from New York which in-

dicates that the species has a wider distribution than has been previously cited.

Distribution. — USA:— Ohio, Pennsylvania, New York, Kentucky. Canada: —

Ontario, Quebec.

Material examined. — &. Lectotype: USA: Ohio, Cincinnati; on Ulmus sp.;

66 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

ALES

Figs. 26, 27. Ectoedemia ulmella (Braun), & (26) and 9 (27) genitalia.

18.viii.1912, B. 578 (Braun); slide no: USNM 16251; in ANS. Paralectotype: In

ANS — USA: Ohio, Clermont Co., on Ulmus sp.; 1 9, 10.viii.1912 (Braun).

Other specimens: In ANS - USA: Ohio, Cincinnati; Overbook; on Ulmus amer-

icana; | ex., 27.vii.1955 (Lewis); on Ulmus sp.; 1 ©, no date (Braun), 1 g,

24.vi.1905, 1 9, 8.vi.1912, 1 g, 1 ©, 3.vii.1907, 1 ex., 6.ix.1954 (Braun). In USNM -

Pennsylvania, Pittsburgh; 1 &, 23.vii.1906 (Engel). New York, Cornell University,

Ithaca; on Ulmus sp.; 1 &, no date (Murfeldt). In CNC — Canada: Quebec,

Kingsmere; on Ulmus fulva'); 2 3,3 92,2 ex., 12—30.vii.1956 (Lewis). Ontario, La

Passe; on Ulmus americana; 1 9, 25.11.1971, 1 g, 22.ii.1971, 1 g, 26.iii.1971

(Lewis).

WILKINSON & NEWTON: Ectoedemia in North America 67

Mines examined: In ANS — USA: Ohio, Anderson’s Ferry; 1 mine on Ulmus

fulva'); 20.ix.1909, B.578 (Braun); 1 mine on Ulmus racemosa; 21.vii.1913, B.578

(Braun). In CNC — Canada: Ontario, Kingsmere; 2 mines on Ulmus rubra;

11.ix.1955, 55—281 (Lewis). Ontario, La Passe; 1 mine on Ulmus americana;

16.ix.1970, 70—111A (Rockburne and Lewis). Ontario, Ottawa; 5 mines on Ulmus

americana; 7.viii.1955, 55—137A (Lewis). Quebec, Kingsmere; 3 mines on Ulmus

rubra; 10.ix.1955, 55—279 (Lewis); 2 mines on Ulmus fulva'); 10.1x.1955, 55—279

(Lewis).

Biology. — Mine. Is a much contorted frass filled tract in leaves of several

species of Ulmus including U. rubra (Slippery or Red Elm), U. americana (White

Elm) and U. thomasi') (Rock or Cork Elm). The tract expands into a blotch so pro-

ducing an upper surface ophistigmatonome. The frass is deposited in both patches

and lines.

Cocoon. Reddish brown and usually spun within the mine-blotch.

Voltinism. Bivoltine with mature larvae present in July and September (Braun,

1917).

Ectoedemia nyssaefoliella (Chambers) comb.n.

(ies28,29558. 59")

Nepticula nyssaefoliella Chambers, 1880: 66.

Nepticula nyssaefoliella Chambers; Dyar, 1903: 546.

Nepticula nyssaefoliella Chambers; Braun, 1909: 429.

Nepticula nyssaefoliella Chambers; Braun, 1917: 183.

Nepticula nyssaefoliella Chambers; Braun in Forbes, 1923: 91.

Nepticula nyssaefoliella Chambers; McDunnough, 1939: 107 (no: 9752).

Description. — External features: g. Head: Palps greyish; antennae dark

brown; tuft on front of head ochreous, vertex orange-ochreous; eye-caps shining

white; collar pale ochre. Thorax and abdomen dark brown, thorax with purple re-

flections, abdomen with silver reflections above, shining metallic grey beneath.

Forewings: ground colour of dorsal surface dark brown with bronze reflections

which become stronger towards the apex, distal scales darker at the tips; fringe

greyish brown, shining silver, with an apical band of dark brown wing-scales;

single medial fascia, shining silver, biconcave. Hindwings: ground colour and

fringe brownish grey; an oval patch of creamy white scales extending from base to

middle of dorsal surface, sometimes absent. Legs dark brown, shining silver

behind. Fig. 58.

Female. As ¢ except for hindwings without specialised scales. A pair of convex

external pockets on ventral side of the fourth abdominal segment, medial in po-

sition (fig. 59).

Wing expanse: &: 4.2—6.4 mm (14 specimens); 9: 4.6—6.4 mm (14). Neotype:

5.0 mm. -

Genitalia: & (fig. 28). Pseuduncus with a single tapering lobe. Gnathos: W-

shaped as in fig. 28 (a); transverse ventral plate with broad central boss. Vinculum:

1) N.B. Ulmus rubra Muhl. = U. fulva Michx. Ulmus racemosa Thomas = U. thomasi Sarg.

68 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

(a)

28 RT Tg 29

002mm

Figs. 28, 29. Ectoedemia nyssaefoliella (Chambers), g (28) and 9 (29) genitalia.

lateral arms broad, ventral plate narrow. Saccus more than two-times the width of

the ventral plate, bilobed. Valves not reaching beyond the pseuduncus, quadrate

with a triangular style arising distally. Transtillae: lateral arms narrow; ventral

arms long and narrow, reaching well beyond the ventral plate; transverse arms

fused. Aedeagus: regular in width, approximately equal to length of the capsule;

vesica with cornuti as many small denticles and with a cup-shaped plate of minute

papillae.

Female (fig. 29). Ductus bursae short and narrow with spiral duct arising me-

dially. Bursa copulatrix: large and covered with scallop-shaped chains of pectina-

tions on striations of the bursa; signum double, comprising a pair of unequal cel-

lular patches, one ovate and one constricted proximally, as in figs. 29 (a) and (b).

Anterior apophyses very broad basally, tapering markedly. Posterior apophyses

straight and narrow, reaching beyond the anteriores.

Host plant: Nyssa sylvatica (Sour Gum).

WILKINSON & NEWTON: Ectoedemia in North America 69

Mine: An upper surface ophistigmatonome.

Diagnosis. — Generally darker and the fascia narrower than in platanella or

members of the platanella group. The male genitalia are easily differentiated from

those of any other member of the genus, listed on page 37, by the absence of the

anellar processes. The females resemble those of rubifoliella, ulmella and quadri-

notata in the absence of a strongly sclerotised colliculum but may be separated by

the unequal signa, one of which is constricted proximally, in nyssaefoliella.

Discussion. — This species was originally described by Chambers (1880) from

the mine; adults reared from Nyssa sp. were described by Braun (1909) and given

the same name. Although the mine(s) constitute the original type material, they

were never preserved or designated. Braun (1909) does not mention the spe-

cialised scale patch present in some specimens in her first description of the

imagine, but does so in a later publication (1917). All the male specimens dated

1909 or earlier and presumably described by Braun (1909), possess the specialised

scale patch, those collected at later dates and presumably described by Braun

(1917) do not show evidence of specialised scale patches. We designate as Neotype

one of the males examined by Braun in 1909, which possesses specialised scale

patches.

It is not unlikely that these two forms of male represent separate species; but in

the absence of any further evidence we treat them as one. Several examples of the

loss of such specialized scales later in life are known within the Lepidoptera, al-

though this is the first possible example in the Nepticulidae.

There are two females reared from Amelanchier sp. by C. Heinrich with indis-

tinguishable genitalia from those of nyssaefoliella. The externals of wo females

differ from the type of nyssaefoliella (which is similar to the others) in the more ir-

rorate nature of the forewings and the broader fascia. The differences may result

from the different host plant but may indicate a separate species. “Nepticula ame-

lanchierella’’ Clemens, 1861, was described and is still only known from the mine.

Distribution. — USA:— Ohio, Virginia, New Jersey. Kentucky.

Material examined. — Designated as neotype: 1 4 from the series examined by

Braun 1909: USA: “B. 454, Cincinnati, Ohio. Annette F. Braun, 31.vii.1909: Nep-

ticula nyssaefoliella Cham.” — on Nyssa sylvatica; slide no: 138-PJN; in ANS.

Other specimens: In ANS — USA: Ohio, Cincinnati; 3 g, 4 9, 20.vi.1911

(Braun); on Nyssa sylvatica; 1 ©, 25.vii.1909, 3 g, 3, 16, 17.viii.1909, 2 g, 1 9,

27.vi.1911, 1 ©, 22.vi.1914, B.454 (Braun). In USNM — Ohio, Cincinnati; 1 9,

26.vi.1907 (Braun); on Nyssa sylvatica; 1 9, 27.vi.1911, B.454 (Braun); | g,

15.vi.1911;4 g, 2 9, 20.vi.1911, 1 9, 26.vi.1916 (Braun). New Jersey, Anglesea; |

©, v, 30 (Kearfott). In FIS — Virginia, Mountain Kale; “WH 3”; 2 3, 21.vii.1940

(Milne and Milne).

Tentative identifications: In USNM — USA: Virginia, Falls Church; “11153

Hopk. US”; on Amelanchier sp.; 1 9, 3.vii.1913 (Reared C. Heinrich). In DFF — |

Q, same data.

Mines examined: In ANS — USA: Ohio, Ferris Woods; 1 mine on Nyssa syl-

vatica; 28.vi.1909, B .454 (Braun).

Biology. — Egg. Laid on the lower surface and, in the single case observed, next

to the midrib of the Nyssa leaf.

70 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Fig. 30. Ectoedemia, specimen 8, ¢ genitalia.

Mine. Begins as a very narrow linear tract which abruptly broadens into an

elongate blotch on the upper surface. The frass is deposited as a continuous black

line in the centre of the linear portion and in a much contorted tract throughout

the blotch (fig. 71).

Larva. Emerges on the upper surface of the leaf.

Pupa. Pale green in colour.

Voltinism. Two or possibly three generations per year.

There is a single male with externals as nyssaefoliella except that the ground

colour of the forewings is darker and the al. ex. greater by almost 1 mm. The geni-

talia of this specimen resemble those of nyssaefoliella in overall form but differ in

the gnathos, which is more typical of Stigmella rather than Ectoedemia, and the rel-

atively larger aedeagus, see figs. 30, 60.

Specimen 8: In ANS — USA: Ohio, Cincinnati; 1 &, 20.vi.1911 (Braun); al. ex.

7.0 mm.; slide no: 135-PJN.

Ectoedemia quadrinotata (Braun)

(figs. 31, 32, 48)

Nepticula quadrinotata Braun, 1917: 168.

Nepticula quadrinotata Braun; Braun in Forbes, 1923: 86.

Nepticula quadrinotata Braun; McDunnough, 1939: 107 (no: 9722).

Ectoedemia quadrinotata (Braun); Wilkinson & Scoble, 1979: 95.

Description. — Fully described from Canada by Wilkinson and Scoble (1979).

WILKINSON & NEWTON: Ectoedemia in North America 71

Figs. 31, 32. Ectoedemia quadrinotata (Braun), & (31) and 9 (32) genitalia.

Diagnosis. — The externals resemble those of trinotata except that there are two

antemedial spots in quadrinotata. The tuberculate pseuduncus and the elaborate

anellar processess differentiate the male from other members of Ectoedemia

studied here. The female genitalia resemble those of rubifoliella, ulmella and nys-

saefoliella but differ in the presence of a weakly sclerotised colliculum in quadri-

notata.

Discussion. — A specimen found in USNM collection taken in Arkansas, indi-

cates that the species has a wider distribution than has been recorded previously.

Distribution. — USA:— Ohio, Arkansas, Kentucky. Canada:— Ontario.

Material examined. — 9. Holotype: USA: Ohio, Cincinnati; on Carpinus sp.;

28.v.1914, B.538 (Braun); slide no: USNM 17326; in ANS. Paratypes: In ANS —

USA: data as Holotype; 2 &, 1 2, 27—30.v.1914 (Braun); Sugar Grove; on Corylus

sp.;1g,1 ©, 1.vi.1915 (Braun).

Other specimens: In USNM — USA: Ohio, Cincinnati; on Carpinus sp.; 1 &,

72 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

30.v.1914, 1 ©, 31.v.1917, 2 g, 1 2, 2, 9.vi.1917, 1 ex., 26.v.1919, (Braun). Ar-

kansas, Washington Co.; 1 ©, 11.v.1966 (Hodges). Ontario, Sparrow lake; 8 &, 1

Q, 12.vii.1926 (Braun). Ontario, Severn; on Ironwood; 2 &, 1 9, 16—20.vi.1925

(MeDunnough). In SOO — Ontario, Trout Creek; on Betula alleghaniensis 1 9,

19.iii.1958, 19, 3.iv.1958.

Mines examined: In ANS — USA: Ohio, Still House Hollow; 2 mines on Car-

pinus sp. 24.viii.1909, B.538 (Braun).

Biology. — Mine. The mine begins as an ophionome — a narrow linear tract ex-

tending along the mid-rib or between two leaf veins and broadens, so filling the

space between them. Later the mine widens further and becomes virtually a

blotch. The frass is dark initially forming a broken line which becomes less

compact and later diffuse. The mine is principally the same regardless which of the

four host plants is attacked: Carpinus caroliniana (Hornbeam), Corylus americana

(Hazel), Ostrya virginiana (Ironwood), Betula alleghaniensis (now: B. lutea) (Yellow

Birch).

Voltinism. Bivoltine.

THE CASTANEAE GROUP

The following eight taxa resemble each other particularly in the genitalia. Of

special note are the following: very round pseuduncus, form of valves, complex

cornuti and anellar processes in males; the thickened colliculum which lacks the

characteristic sclerotised ring of other groups, very long posterior apophyses, es-

pecially in comparison with anterior ones, and the long narrow genitalia in fe-

males. Together with these features the similar venation with expanded costal

margin of hind wing (fig. 1) and similar wing markings show this to be a discrete

group. To this add the fact that all attack bark, as far as is known, but some form

galls and others mine.

In many ways this group presumably corresponds with the sub-genus Zimmer-

mannia Hering, 1940, adopted by several European workers. However the name is

not used here because the diagnosis which separates the gall makers from the bark

miners is invalid.

The group is called the castaneae group rather than by the older name obrutella

because of the uncertainty about the latter life history and site of larval attack.

Ectoedemia obrutella (Zeller)

(figs. 1, 33, 34, 47)

Trifurcula obrutella Zeller, 1873: 316.

Trifurcula obrutella Zeller; Dyar, 1903: 547.

Ectoedemia obrutella (Zeller); Busck, 1913: 103.

Ectoedemia obrutella (Zeller); Braun, 1917: 200.

Ectoedemia obrutella (Zeller); Braun in Forbes, 1923: 83.

Ectoedemia obrutella (Zeller); McDunnough, 1939: 107 (no: 9782).

Nepticula bosquella Chambers in Hayden, 1878a: 106 (syn. by Busck, 1903: 208).

Nepticula bosqueella (sic) Chambers; Chambers in Hayden, 1878b: 157.

Opostega bosqueella (sic) Chambers; Dyar, 1903: 547 (mistaken genus).

WILKINSON & NEWTON: Ectoedemia in North America 73

Nepticula bosquella Chambers; Busck, 1903: 208 (cited under Opostega albogaleriella).

Ectoedemia bosqueella (sic) (Chambers); Braun, 1917: 200.

Ectoedemia bosquella (Chambers); Meyrick, 1928b: 426.

Description. — External features: ¢,9. Head: palps off-white; antennae sandy

buff; tufts on front of head and vertex chocolate-brown; eye-caps creamy white;

collar chocolate-brown. Thorax creamy white with scattered brown scales above.

Abdomen greyish, shining metallic grey beneath. Forewings: ground colour of

dorsal surface creamy white, weakly lustrous, irrorate with brown scales partic-

Figs. 33, 34. Ectoedemia obrutella (Zeller) Busck, g (33) and 9 (34) genitalia.

74 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

ularly dense basally and terminally giving the impression of a medial, pale fascia;

fringe whitish, weakly lustrous. Hindwings: ground colour and fringe shining

white; with a pale yellow brush of ciliae at the base of the costa; costa emargin-

ated. Legs: buff with scattered yellow-gold reflections, darker behind; hind tibiae

densely covered inwardly with sharp bristles. Fig. 47.

Wing expanse: &: 7.4—9.5 mm (44 specimens); 9: 7.6—10.0 mm (27).

Genitalia: & (fig. 33). Pseuduncus with a single lobe, broad and flattened.

Gnathos: an inverted V as in fig. 33 (a); transverse ventral plate with a large medial

process; dorso-lateral arms very broad and weakly arcuate. Vinculum: lateral arms

broad, ventral plate broad. Saccus approximately half width of ventral plate,

weakly bilobed. Valves reaching beyond the pseuduncus and tapering markedly.

Transtillae: lateral arms broad; ventral arms very long, reaching beyond the saccus

laterally; transverse arms fused to form a continuous strap constricted medially.

Aedeagus: flask-shaped, just greater than length of the capsule; vesica with cornuti

as many small denticles evenly distributed and with a plate of minute papillae as in

fig. 33 (b); anellus comprising a pair of tooth-like spines with several denticles and

pair of palmate plates laterally.

Female (fig. 34). Ductus bursae long, weakly sclerotised proximally with

complex spiculate lobe as in fig. 34. Accessory sac small, arising from ductus me-

dially and with terminally spiral accessory duct. Bursa copulatrix small, covered

with irregular chains of pectinations on striations of the bursa; signum double

comprising two equal sized ovate patches of hexagonal cells. Anterior apophyses

short and broad. Posterior apophyses straight and narrow, reaching well beyond

the anteriores.

Host plant: Several adults have been taken from the trunks of Quercus sp.

Mine: Not known.

Diagnosis. — The very large al.ex. and the pale ground colour separates this

species from all other members of the genus except the castaneae group taxa. The

tuft is darker than in piperella and the forewings less irrorate with brown, in obru-

tella. It lacks the iridescent scales of the hindwings found in acanthella. The saccus

is more weakly bilobed than in piperella or acanthella. Compare figs. 33, 35 and 36

for other possible differences in the genitalia. See also diagnosis for heinrichi and

phleophaga.

Discussion. — It is unfortunate we do not know more of the life history of this

species. I (C.W.) suspect it is a bark miner. The genitalia are so very similar to

those of the other oak bark miner heinrichi and the differences in externals, only

slight, that how far they are taxonomically separate must remain in some doubt.

Originally described by Zeller (1873) as a member of Trifurcula Zeller, from a

male and a female specimen collected by Boll in Texas. This species was also de-

scribed by Chambers (1878a) from Texas material and given the name Nepticula

bosquella. Chambers (1878b) then listed the name incorrectly as bosqueella (sic).

Subsequently, Dyar (1903) listed bosqueella (sic) as a member of Opostega Zeller;

since he gives no reason for the combination and no previous reference to the

name can be traced it would seem to be an error by Dyar. Busck (1903) realised

this error, also comparing material with Zeller’s type of Trifurcula obrutella and

synonymising bosquella with obrutella. Busck (1913) later reports, in his description

WILKINSON & NEWTON: Ectoedemia in North America 75

of Ectoedemia castaneae, that obrutella has important differences in wing venation

from the type species of Trifurcula and he included obrutella in the genus Ectoe-

demia.

Distribution. — USA: — Texas, Pennsylvania, Massachusetts, Florida, Georgia,

Alabama, Mississippi.

Material examined. — d Type of bosquella: USA: “7/5; Collection of C.V.

Riley; Nepticula bosquella Cham. Texas Ch.; Wlsm 1106 1882; Type no: 528

USNM”; slide no: CNC 3478; in USNM.

Other specimens: In MCZ — USA: Pennsylvania, Oak Station, Allegheny

County; 1g, 8.v.1913, 4g, 29, 25.v.1913, 1g, 5.vi.1910, 6g, 29, 9.vi.1912

(Marloff). Pennsylvania, Jeannette; 14, 2.vi. (C.M. Acc. 2722, Klages). Pennsyl-

vania, Pittsburgh; 14, 3.vi., 1g, 7.vi.1904, 39, 20.vi. (C.M. Acc. 2723); 24, 39,

17.vi.1907 (C.M. Acc. 3495); On trunk of Quercus sp.; 23,79, 17.vi.1907 (Kahl);

19, 17.vi.1906 (C.M. Acc. 4067, Engel). Pennsylvania, New Brighton; 19,

12.vi.1907, 1g, 19, 14.vi.1907, 19, 15.vi.1907, 1g, 27.vi.1907 (C.M. Acc. 4067).

Massachusetts, Barnstable; 14, 26.vi.1958, 1 3, 4.vii.1958 (Kimball). In USNM—

Florida, Oneco, Manatee Co.; 34, 5.v.1953 (Dillman). Gulf Coast Exp. Sta. Bra-

denton; 38, 19, 11 ex., 13—24.iii.1955 (Keisheimer). Siesta Key, Sarasota Co.;

104, 19, 9 ex., 15.iii.1953—3.iv.1960 (Kimball). Gainsville, Alachua Co.; 2g,

14.iii.1955 (Morse), 1 ex., 1.v.1976. Pensacola; 13, 1 ex., 21 + 25.11.1962, 1g,

24.v.1964 (Hills). Mississippi, Choctaw Co.; 14, 22.iv.1976(Heppner). Three other

specimens probably belong here: Georgia, Crooked River St. Pk., Campden Co.;

14, 27.iv.1976 (Heppner). Alabama, Black Warrior River, Greenboro, Hale Co.;

19, 1 ex., 23.iv.1976 (Heppner).

Biology. — Immature stages unknown.

Ectoedemia acanthella sp.n.

(fig. 35)

Description. — External features: &. Head: palps off-white; antennae pale buff;

tufts on front of head and vertex dark brown; eye-caps creamy white; collar dark

brown. Thorax greyish white with scattered brown scales. Abdomen greyish white,

shining metallic beneath. Forewings ground colour of dorsal surface greyish white

with scattered brown scales, reflecting purple; fringe creamy white, weakly lus-

trous. Hindwings: ground colour and fringe buff with each scale shining pale bluish

purple; costa emarginated. Legs: pale buff with some gold reflections; sharp

bristles of hind tibiae pronounced.

Wing expanse: Holotype 7.2 mm.

Genitalia. & (fig. 35). Pseuduncus with single flattened lobe, broad. Gnathos: V-

shaped; transverse ventral plate broad with medial process; dorso-lateral arms

very broad and arcuate. Vinculum: lateral arms broad, ventral plate broad. Saccus

strongly bilobed with centre almost reaching edge of vinculum. Valves extending

just beyond pseuduncus but arising high up and tapering markedly. Transtillae:

lateral arms broad; ventral arms long, but not beyond anterior edge of saccus;

transverse arms fused to form continuous strap, constricted medially. Aedaegus:

flask-shaped, greater than length of capsule; vesica with cornuti as many small

76 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Fig. 35. Ectoedemia acanthella sp.n., 3 genitalia.

denticles evenly distributed and with plate of minute papillae as in fig. 35(b);

anellus comprising a number of tooth-like spines.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — Exteral features as obrutella except that abdomen is creamy white.

Forewings: ground colour of dorsal surface greyish white with scattered brown

scales, reflecting purple; fringe creamy white, weakly lustrous. Hindwings: ground

colour and fringe buff with each scale shining pale bluish purple. Genitalia as obru-

tella except that saccus is very narrow and markedly bilobed, each lobe as long as

broad at base. Aedeagus: broad and regular, greater than length of capsule; vesica

with cornuti as many small denticles evenly distributed and with a plate of minute

papillae as in fig. 35; anellus comprising a pair of tooth-like spines but no palmate

plates. Valves not so long and lack the inner corrugations and knobs of other

species.

Discussion. — The differences given in the diagnosis, in particular the markedly

bilobed saccus and the absence of the palmate anellar plates, exclude this

specimen from obrutella, piperella sp.n., heinrichi and castaneae. The absence of

inner knobs on the valves separates acanthella sp.n. from phleophaga and mesoloba.

Distribution. — USA: — New Jersey.

Material examined. — ¢ Holotype: USA; New Jersey, Essex County Pk.; “July

26 Trap W.D. Kearfott’’; slide no: USNM 17287; in USNM.

Biology. — Immature stages unknown.

WILKINSON & NEWTON: Ectoedemia in North America 77

Ectoedemia piperella sp.n.

(figs. 36, 45)

Description. — External features: ¢. Head: palps buff; antennae pale brown;

tufts on front of head and vertex orange-ochreous; eye-caps shining white; collar

pale brown. Thorax yellow-cream, irrorate with purplish brown. Abdomen grey,

shining metallic grey beneath. Forewings: ground colour of dorsal surface yel-

lowish, irrorate with purplish brown scales particularly at the base and along the

costal margin, reflecting purple; fringe creamy white, shining metallic grey.

Hindwings: ground colour and fringe buff, shining metallic grey; pale yellow brush

of ciliae at the base of the costal margin; costa emarginated. Legs: tibiae of fore

and midlegs dark brown, otherwise buff and shining metallic grey behind; hind

tibiae densely covered inwardly with sharp bristles. Fig. 45.

Wing expanse: ¢: 6.4—8.2 mm (3 specimens). Holotype: 7.2 mm.

Genitalia: & (fig. 36). Pseuduncus with a single, broad and flattened lobe.

Gnathos: an inverted V as in fig. 36(a); transverse ventral plate with large medial

process; dorso-lateral arms broad and weakly arcuate. Vinculum: lateral arms

broad; ventral plate broad. Saccus as wide as ventral plate, bilobed. Valves

reaching beyond the pseuduncus, broad basally but tapering markedly, inner

margin concave distally. Transtillae: lateral arms broad; ventral arms long and

narrow, reaching beyond the ventral plate; transverse bars fused to form a con-

tinuous strap, constricted medially. Aedeagus: broad and regular in width, approx-

(b)

Fig. 36. Ectoedemia piperella sp.n., & genitalia.

78 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

imately equal to length of capsule; vesica with cornuti as many small denticles

evenly distributed and with a plate of minute papillae as in fig. 36(b); anellus com-

prising a pair of tooth-like spines with several large single spines and a pair of

palmate plates laterally.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — Resembles both obrutella and acanthella but differs in the more

yellowish ground colour of the dorsal aspect and the more dense irroration of the

forewings, in piperella. It lacks the iridescent scales on the hindwings found in

acanthella. The saccus is more markedly bilobed than in obrutella but less than in

acanthella. The stouter valves and the relatively short aedeagus separate this from

obrutella.

The colours of the vertex and tufts on front of the head differ in this species

from heinrichi and castaneae. It also has purple and grey reflections on the wings

which are missing in these latter species. The male genitalia are similar but

compare figs. 36 and 37 for differences. E. piperella can be diagnosed from phleo-

phaga and mesoloba by the absence of knobs on the inner surface of the valves.

Distribution. — USA: — Arkansas.

Material examined. — & Holotype: USA: “Devil's Den St. Pk., Washington

Co. Ark., 9.vi.1966 R. W. Hodges”; slide no: USNM 17285; in USNM. Paratypes:

In USNM — USA: same data as Holotype; 34, 28.v.1966, 9.vi.1966, 16.vi.1966

(Hodges); slide no: USNM 17286.

Biology. — Immature stages unknown.

Ectoedemia heinrichi Busck

(figs. 37, 38, 61)

Ectoedemia heinrichi Busck, 1914a: 149.

Ectoedemia heinrichi Busck; Braun, 1917: 199.

Ectoedemia heinrichi Busck: Braun in Forbes, 1923: 83.

Ectoedemia heinrichi Busck; McDunnough, 1939: 108 (no: 9786).

Description. — External features: 3 ©. Head: palps buff; antennae light brown;

tuft on front of head and vertex dark brown; eye-caps and collar white. Thorax

white with some brown-tipped scales. Abdomen pale brown. Forewings: irrorate

with white and brown scales, variable on each wing as well as between specimens.

Hindwings: greyish buff; undersurface brownish buff. Legs buff. Fig. 61.

Wing expanse: g: 8.0—10.0 mm (15 specimens); 9: 8.0—10.5 (11). Holotype:

8.5mm.

Genitalia: & (fig. 37). Pseuduncus with a single broad lobe as in fig. 37 (a).

Gnathos: W-shaped, but lateral arms shorter than inner arms. Vinculum: lateral

arms broad; ventral projections narrow. Saccus broad and weakly bilobed. Valves

extending as far posteriorly as pseuduncus, triangular and slightly arcuate. Trans-

tillae: a narrow inverted U-shape; lateral arms broad and short; ventral arms

narrow, projecting anteriorly well beyond margin of ventral plate; transverse bars

fused to form a narrow arcuate strap. Aedeagus: approximately equal to length of

WILKINSON & NEWTON: Ectoedemia in North America 79

Figs. 37, 38. Ectoedemia heinrichi Busck, 3 (37) and 9 (38) genitalia.

genitalia; vesica with cornuti as small denticles and plate of minute papillae as in

fig. 37(b); anellus with pair of tooth-like spines and a pair of palmate plates.

Patches of scent scales present, one pair overlapping genitalia.

Female (fig. 38). Ductus bursae short, without sclerotised colliculum, but with

enlarged spiculate lobe. Accessory duct arising from dilation of ductus and distally

spiralled. Bursa copulatrix large, covered with pectinations; signum double, com-

prising two relatively small, but equal-sized patches of cells. Anterior apophyses

comparatively short and broad. Posterior apophyses straight and narrow, very

long, extending below swelling of ductus.

Host plant: Quercus palustris (Pin Oak).

Mine: An oval spiral in the bark.

Diagnosis. — This species is very similar to castaneae, even the larva and espe-

80 TUDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

cially the genitalia, although heinrichi appears to be a slightly larger species than

castaneae. They are very difficult to diagnose with respect to morphology, but with

regard to biology, heinrichi mines the bark of Quercus, whereas castaneae makes

galls on chestnut. Another similar species, obrutella, is distinguished from heinrichi

by a darker, brown, collar and paler hindwings and there are also small differences

inthe male and female genitalia (see figs. 33 and 34).

Discussion. — The wing markings are variable and the genitalia similar to those

of other species in the group. The life history of heinrichi is well known and was

first worked out by Carl Heinrich and detailed by Busck (1914a). This is fortunate

since its separate identity is most strongly supported by what is known of its bi-

ology.

Distribution. — USA: — Virginia, Ohio, Kentucky, Illinois.

Material examined. — ¢ Lectotype: USA: Virginia, “Falls Church, Va; Reared

May 22-14, C. Heinrich; Hopk. US 12107; Ectoedemia heinrichi cotype Busck;

USNM No. 19039; slide no. 16935; Quercus palustris”; in USNM. Paralectotypes:

In USNM-data as lectotype; 93,49, 1 ex, 8.v.1914 — 5.vi.1914.

Other specimens: In ANS—USA: Ohio, Cincinnati; 4g, 79, 4.vi.1903,

21.vi.1904, 3.vi.1905, 15.v.1906, 30.v.1906, 7.vi.1906, 20.v.1917, 5.vi.1917,

14.vi.1917 (Braun). In USNM—USA: Illinois, Putnam Co. 18, 25.v.1975 (Glenn).

ld, in ANS, data as lectotype, but not labelled cotype, was presumably given to

Braun in exchange for 1 4 from Cincinnati, Ohio, 4.vi.1913, in USNM.

Mines examined: In USNM—USA: Numerous mines (and pupae) from Busck

collection.

Biology. — Egg. Laid on the bark of young branches.

Mine. I have only seen this mine on the twigs of young saplings (C.W), although

Busck (1914a: 149) reported finding it also on the outer branches of larger trees. It

isa narrow linear track coiled into an oval spiral with the empty egg at the centre.

Larvae. Matures in the autumn, October and November, when it leaves the

mine and falls to the ground.

Pupa. Cocoon flattened, oval, reddish brown, 2—2.5mm x 3—4 mm.

Voltinism. Univoltine. Adults normally emerge in May—June. In the laboratory

specimens often emerge two or three months earlier.

Ectoedemia castaneae Busck

(fig. 62)

Ectoedemia castaneae Busck, 1913: 103.

Ectoedemia castaneae Busck; Braun, 1917: 198.

Ectoedemia castaneae Busck; Braun in Forbes, 1923: 83.

Ectoedemia castaneae Busck; McDunnough, 1939: 108 (no. 9784).

Description. — External features: SQ. Head: palps buff; antennae brown; tuft

on front of head chocolate brown, vertex brown; eye-caps white; collar white.

Thorax irrorate with brown and white scales. Abdomen brown. Forewings: ground

colour of dorsal surface white, but many scales are tipped with dark brown,

making the surface variously irrorate. Hindwings: brownish buff; undersurface

brownish grey. Legs buff or brownish buff. Fig. 62.

WILKINSON & NEWTON: Ectoedemia in North America 8

Wing expanse: &: 6.5—8.0 mm (3 specimens); 9: 7.0—8.0 mm (4); Lectotype:

6.5 mm.

Genitalia: 3,9. Similar to those of heinrichi which are illustrated in figs. 37 and

38.

Host plant: Castanea sp. (Chestnut).

Mine: In form of small galls around young twigs.

Diagnosis. — As discussed in the diagnosis of heinrichi, the genitalia of both

males and females of these two species are very similar; separate figures are not

helpful and diagnosis on these features is too difficult. However, castaneae and

heinrichi do differ in their life history and host plant and possibly also in size. Since

castaneae is so similar to heinrichi in externals and genitalia the diagnosis for the

latter species with respect to obrutella also holds for castaneae, which has a lighter,

white, collar and darker hindwings than obrutella and similar small differences in

the genitalia.

Discussion. — The life history of this species as a gall-former enables us to rec-

ognize its separate identity from heinrichi.

Distribution. — USA: — Virginia, Pennsylvania, Kentucky.

Material examined. — ¢ Lectotype: USA: Virginia, ‘“Vietch, Va., 23 April 1913,

Snyder; USNM No. 16333; slide no. 16713; Ectoedemia castaneae Busck cotype;

11236 Hopkins US”; in USNM. Paralectotypes: In USNM—data as lectotype;

1g, 29, 23—24.iv.1913. Other specimens: In USNM—USA: Pennsylvania, New

Brighton; 24,29, 14.vi.1907 (Meyrick Museum).

Biology. — Egg. Laid on the bark of Castanea twigs.

Larva. Typically flattened, with limb buds on segments 3 to 10 and 13, but very

rudimentary on segment 5.

Mine. No mine is made as such, but more a spherical gall resembling in form

and size “the egg masses of the forest tent-caterpillar”’ (Busck, 1913: 103).

Voltinism. Bivoltine or, more probably, trivoltine.

Ectoedemia phleophaga Busck

(figs. 39, 40, 63)

Ectoedemia phleophaga Busck, 1914b: 3.

Ectoedemia phleophaga Busck; Braun, 1917: 198.

Ectoedemia phleophaga Busck; Braun in Forbes, 1923: 83.

Ectoedemia phleophaga Busck; McDunnough, 1939: 108 (no. 9785).

Description. — External features: ¢ ©. Head: Palps buff; antennae light brown;

tuft on front of head and vertex white or buff; eye-caps white; collar brown.

Thorax brown. Abdomen pale brownish buff. Forewings: brown with some white

scales, sometimes forming a postmedial and antemedial spot and occasionally a

postmedial fascia. Thorax and basal half of forewing sometimes with dark bluish

fuscous reflections. Hindwings and ventral surface brownish buff. Legs buff. Fig.

63.

Wing expanse: &: 8.3—10.5 mm (32 specimens); 9: 8.7—10.6 mm (27); lecto-

type::9.0 mm.

ee}

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

39 40

Figs. 39, 40. Ectoedemia phleophaga Busck, & (39) and © (40) genitalia.

Genitalia: & (fig. 39). Pseuduncus with single rounded lobe. Gnathos: W-shaped

with lateral arms as long as inner arms. Vinculum: lateral arms and ventral plate

narrow. Saccus broad, weakly bilobed, concave anteriorly. Valves: fractionally

longer than genitalia; narrowly arcuate, with median boss. Transtillae: lateral arms

short and narrow; ventral arms short; transverse bars fused centrally. Aedeagus:

equal to length of capsule; vesica with many denticles and with large plate of small

papillae as in fig. 39(b); anellus comprising a pair of long, broad spines and pair of

simple plates laterally.

Female (fig. 40). Ductus bursae long, colliculum not heavily sclerotised. Acces-

sory duct arising from ductus bursae and spiral distally. Bursa copulatrix long, with

WILKINSON & NEWTON: Ectoedemia in North America 83

fine pectinations. Signum double, comprising a pair of long, reticulate patches, ap-

proximately equal in area. Anterior apophyses long, arcuate and narrow. Posterior

apophyses very long and straight, reaching well beyond the anteriores.

Host plant: Castanea dentata.

Mine: A bark ophionome.

Diagnosis. — This species is similar to castaneae but phleophaga is easily distin-

guished by its larger size, darker wings and lighter tufts and abdomen. The white or

buff tufts also distinguish it from obrutella and heinrichi in which they are dark

brown. Considering the male genitalia, phleophaga is the only bark-miner with the

inner arms of the W-shaped gnathos equal in length to the lateral arms. It also has

a central boss on the middle of the valves as does mesoloba, in which it is more sim-

ple in form. The female genitalia are typical of the Ectoedemia species attacking

bark described here: they lack the strongly sclerotised colliculum and have very

long posterior apophyses. It is possible that phleophaga can be diagnosed by the an-

terior apophyses, which are narrower and straighter than in other similar species,

especially obrutella, heinrichi and castaneae.

Discussion. — Again knowledge of the life history allows us to separate phleo-

phaga with certainty from other species associated with bark, especially castaneae,

which is also found on chestnut. The life history was worked out by Busck (1914b),

together with Snyder and Heinrich and independently at about the same time, by

Ruggles (1913). Because of its pest status (see Biology), phleophaga has been given

the common name of “Chestnut bastminer”.

Distribution. — USA: — Virginia, Pennsylvania, Massachusetts?

Material examined. 9 Lectotype: USA: Virginia, “Falls Church Va., 22 Sept.

1913; USNM No. 16900; slide no. 16633; Castaneae dentata; 11245 Hopk US”; in

USNM. Paralectotypes: In USNM—data as lectotype, with cotype labels; 11g,

89, 12—22.ix.1913.

Other specimens: In USNM—data as lectotype; 21 g, 19 9, 2 ex. 22.vi-

i—15.1x.1914.

Biology. — Egg laid on the bark.

Mine. In the lower layers of the bark and in the cambium. A slender, contorted,

serpentine mine; a number of centimetres long and a few millimetres wide, but the

width varying throughout its length.

Larva. On hatching from the egg, the larva burrows in the bark until the second

or third instar. By this time winter causes it to hibernate in its tunnel and activity

starts again in the spring. When fully grown it is white with dark brown sclerotisa-

tions of the head capsule, thoracic, sternal and anal plates. In April to early June

the larva tunnels out and drops to the ground. Only then does the tunnel become

apparent by means of the exit hole.

Pupa. The cocoon is spun amongst the debris on the ground or in a burrow in

loose soil. It is a reddish brown, closely woven cocoon, rather seed-like.

Voltinism. The species seems to be univoltine, the adults appearing in August

and September.

Pest status. According to Ruggles (1913: 852), phleophaga is associated with the

spread of the chestnut bark disease, Endothia parasitica. An enormous number of

larval exit holes appear at a time of year when blight spores of the fungus are parti-

84 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

culary prevalent (see also the Report of the State Forester of Massachusetts on the

Chestnut Bark Disease, 1912).

Ectoedemia chlorantis Meyrick

(figs. 41, 46)

Ectoedemia chlorantis Meyrick, 1928a: 462.

Ectoedemia chlorantis Meyrick; McDunnough, 1939: 108 (no: 9787).

Description. — External features: 9. Head: palps pale buff; antennae buff with

Fig. 41. Ectoedemia chlorantis Meyrick, 9 genitalia.

WILKINSON & NEWTON: Ectoedemia in North America 85

darker banding; tuft on front of head and vertex yellowish brown; eye-caps

whitish; collar brownish buff. Thorax buff, irrorate with light grey and abdomen

presumably similar. Forewings yellowish buff, speckled lightly with light grey or

fuscous and more heavily speckled in distal half. Hindwings and ventral surface

buff, tinged with grey. Legs buff, irrorate with fuscous on outer surface. Fig. 46.

Wing expanse: Holotype: 9 mm.

Genitalia: © (fig. 41). Ductus bursae short and broad, with sclerotised colli-

culum (not ring-shaped) and with spicules. Accessory duct spiral medially. Bursa

copulatrix long, with fine pectinations; signum double, comprising two approxi-

mately equal-sized ovate patches of cells. Anterior apophyses broad and arcuate.

Posterior apophyses particularly long and narrow.

Host plant: Not known.

Mine: Not known.

Diagnosis. — The forewing of ch/orantis has light background colouring, pep-

pered with darker irrorations, similar to that of mesoloba. There is not enough

material to know whether there are reliable diagnostic differences in the externals

of these two, but ch/orantis is possibly a much larger species. The female genitalia

of chlorantis lack the characteristic colliculum of most leaf- and petiole-mining

species, although there is a weak sclerotisation. The long posterior apophyses are

also of note as a possible indication of a bark-miner.

Discussion. — This species is only known from the female holotype, which ap-

pears to be distinct. It should be no great problem for additional material to be

collected in the type-locality to provide us with better information.

Distribution. — Canada: — Ontario.

Material examined. — 9 Holotype: Canada: Ontario, “Toronto, September

(Parish); in BM(NH).

Biology. — Unknown.

Ectoedemia mesoloba Davis

(figs. 42, 50)

Ectoedemia mesoloba Davis, 1978: 209.

Description. — External features. g. Head: palps buff or white; antennae

whitish proximally, brown distally, but paler above; tuft on front of head, vertex

and eye-caps buff to white. Thorax white with some pale brown above. Abdomen

buff with much pale brown marking above. Forewings: irrorate with buff and pale

brown. Hindwings and ventral surface pale buff. Legs buff with some brown scales

dorsally, particularly on pro- and mesothoracic legs. Fig. 50.

Wing expanse: Holotype: 5.5 mm.

Genitalia. & (fig. 42). Pseuduncus with a single, unspecialised, rounded lobe as

in fig. 42(a). Gnathos: W-shaped with central boss shorter than lateral arms. Vin-

culum: broad and short, anteriorly slightly concave. Saccus broad and hardly

concave. Valves not extending further than pseuduncus, narrow and lobed halfway

along costal margin. Transtillae: strongly arcuate lateral arms broad and short;

ventral arms extend well forward, beyond margin of ventral plate; transverse bars

fused centrally. Aedeagus: equal in length to or slightly longer than genitalia;

86 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

$ \

Fe re

ran

EF 100

‘

Fig. 42. Ectoedemia mesoloba Davis, & genitalia.

vesica with large cornuti and plate of minute papillae; anellus with pair of tooth-

like spines and spinose plates laterally, as in fig. 42(b).

Female: Unknown.

Host plant: Not known.

Mine: Not known.

Diagnosis. — Externally, mesoloba is mainly light coloured, like chlorantis, but

mesoloba is very much smaller. The male genitalia have the central part of the

gnathos markedly shorter than the lateral arms. This feature together with the

presence of a central boss on the valves separates mesoloba from the other Ectoe-

demia species reported here, especially the bark-miners whose genitalia are so

similar to those of mesoloba. There is also a boss on the valves in phleophaga male

genitalia, but it is more complex in form.

Discussion. — Although mesoloba is similar to chlorantis the type-localities are

widely separated and it seems unlikely that they occur throughout the intervening

area. Davis, who recently described mesoloba, feels that the diagnostic features

and type-locality are so characteristic that the specimen should be given specific

status. It is unfortunate that more material is not available. I suspect it is a bark-

miner, although it is somewhat smaller than the other bark-miner species dis-

cussed here (C.W.).

Distribution. — USA: — Florida.

Material examined. — ¢ Holotype: USA: “Pensacola, Escambia Co., Florida,

Nov. 12.1961, Shirley Hills”; slide no. 16835; in USNM.

Biology. — Not known.

REFERENCES

Beirne, B. P., 1945. The male genitalia of the British Stigmellidae (Nepticulidae) (Lepidoptera): —

Proc. R. Ir. Acad. (B) 50: 191—218, 81 figs.

WILKINSON & NEWTON: Ectoedemia in North America 87

Borkowski, A., 1969. Studien an Stigmelliden (Lepidoptera) Teil I. Zur Verbreitung, Biologie und Öko-

logie der Stigmelliden in den polnischen Sudeten. — Polskie Pismo ent. 39: 95—122, 20 figs.

—, 1972. Studien an Nepticuliden (Lepidoptera) Teil IV. Bemerkungen zur Nomenklatur und Sys-

tematik der Familie Nepticulidae. — Polskie Pismo ent. 42: 689— 709, 4 pls., 16 figs.

Braun, A. F., 1909. Notes on Chambers species on Tineina. — Ent. News. 20: 428—434.

, 1912. Notes on North American species of Nepticula, with descriptions of new species (Lepi-

doptera). — J. Cincinn. Soc. nat. Hist. 21: 84—101, 13 figs.

——, 1914. Notes on North American species of Nepticula with descriptions of new species (Lepi-

doptera). — Can. Ent. 46: 17—24, 8 figs.

———, 1915. New genera and species of Tineina. — Can. Ent. 47: 188—197, 6 figs.

—., 1917. Nepticulidae of North America. — Trans. Am. ent. Soc. 43: 155—209, 4 pls.

——, 1923. Superfamily Nepticuloidea. Family 5 Nepticulidae In: Lepidoptera of New York and

Neighbouring States pp. 79—98, 10 figs. Forbes W.T.W. (ed.). — New York.

—, 1925. Microlepidoptera of Northern Utah. — Trans. Am. ent. Soc. 51: 183—226.

, 1927. Descriptions of New Microlepidoptera. — Trans. Am. ent. Soc. 53: 191—199.

Busck, A., 1903. Notes on Brackenridge Clemens’ types of Tineina. — Proc. ent. Soc. Wash. 5: 181—

220.

—, 1907. New American Tineina. — Proc. ent. Soc. Wash. 8: 86—99, 11 figs.

—, 1913. Two microlepidoptera injurious to Chestnut. — Proc. ent. Soc. Wash. 15: 102—104, | fig.

—, 1914a. Descriptions of New Microlepidoptera of Forest Trees. — Proc. ent. Soc. Wash. 16:

143—150.

, 1914b. The Chestnut Bastminer (Lepidoptera, Tineidae). — Insecuter Inscit. Menstr. 2: 3—4.

Chambers, V. T., 1873. Nepticula. In: ‘*Micro-Lepidoptera”. — Can. Ent. 5: 124—128.

, 1878a. New Tineina from Texas. In: Hayden, F. V. (ed.). — Bull. U.S. geol. geogr. Surv. Territ.

4: 79—106.

—, 1878b. Index to the described Tineina of the United States and Canada. In: Hayden, F. V. (ed.).

— Bull. U.S. geol. geogr. Surv. Territ. 4: 125—167.

, 1880. Notes upon some Tineid larvae. — Psyche, Camb., 3: 63—68.

Clemens, B., 1860. Contributions to American Lepidopterology — No. 5. — Proc. Acad. nat. Sci.

Philad. 12: 203—221.

——, 1861!). Micro-Lepidopterous Larvae. Notes on a few species, the imagos of which are probably

undescribed. — Proc. ent. Soc. Philad. 1 (4): 75—87.

—, 1862a. New American Microlepidoptera. — Proc. ent. Soc. Philad. 1 (5): 131—137.

—, 1862b. North American Microlepidoptera. Nepticula miners of the Sycamore leaf. — Proc. ent.

Soc. Philad. 1: 149—151.

—, 1865. North American Microlepidoptera. — Proc. ent. Soc. Philad. 5: 133—147.

—, 1872. In: Stainton, H.T. (ed.). The Tineina of North America (being a collected edition of his

writings on that group of insects). With notes by the editor. XVI + 282 pp. — London.

Cochaux, P., 1969. A. European Leaf Miner Stigmella (Nepticula) turbidella [H.S.] in Quebec. — Bi-m.

Res. Notes. Can. Dep. Fish. For. 25 (2): 12.

Davis, D. R., 1978. New leaf-mining Moths of the Family Nepticulidae from Florida. — Florida Ento-

mologist. 61: 209—224.

—, 1979. Manoneura, a new name to replace the generic homonym Oligoneura Davis (Lepidoptera:

Nepticulidae). — Florida Entomologist. 62: 276.

Downes, J. A., 1968. A Nepticulid Moth feeding on the leaf nectaries of Poplar. — Can. Ent. 100:

1078—1079.

Dyar, H. G., 1903. List of North American Lepidoptera, and key to the literature of this order of in-

sects. — Bull. U.S. natn. Mus. 52: xix + 723 pp. — Washington.

!) Although the publication is dated Nov. 1861, it seems likely that this was the date that the Entomo-

logical Society of Philadelphia agreed on the contents for the issue. The printed copy was first received

by the society’s librarian on 13 Jan. 1862, so publication falls somewhere between and almost certainly

close to Jan. 13. Further explanation may be found in Brown, F. M., 1964, “Dates of publication of the

various parts of the Proceedings of the Entomological Society of Philadelphia”, Trans Amer. ent. Soc.

89: 305-308.

88 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.2, 1981

Emmet, Col. A. M., 1971. Notes on some of the British Nepticulidae (Lepidoptera). — Entomologist’s

Rec. J. Var. 83: 75—83, 136— 142, 163— 171, 240—8, 278— 282, 282— 287, 300— 304.

—, 1976. In: Heath, J. (ed.). Nepticulidae. The moths and butterflies of Great Britain and Ireland.

I, Micropterigidae-Heliozelidae: 171—267 pp., 56—73 figs., 19—113 maps., 1—9, 11-12 pls.

Freeman, T. N., 1962. A new species of Nepticula v. Heyd. on Birch (Lepidoptera: Nepticulidae). —

Can. Ent. 94: 522—523, 2 figs.

Haworth, A. H., 1828. Lepidoptera Britannica: sistens digestionem novam Insectorum Lepidopterorum

quae in magna Britannia reperiuntur 4: 513—569. — London.

Hering, E. M., 1940. Zimmermannia liebwerdella (Zimmermann). Ein Nachwort. — Mitt. zool. Mus.

Berlin. 24: 266.

, 1951. Biology of the leafminers: iv + 420 pp., 2 pls., 180 figs. — ’s-Gravenhage.

Herrich-Schäffer, G. A. W., 1853—55. Systematische Bearbeitung der Schmetterlinge von Europa, 5,

Die Schnaben und Federmotten: 394 pp. — Regensburg.

Johansson, R., 1971. Notes on Nepticulidae (Lepidoptera) I. A revision of the Nepticula ruficapitella

group. — Ent. Scand. 2: 241—262, 52 figs.

Karsholt, O. & E. Schmidt Nielson, 1976. Systematisk fortegnelse over Danmarks sommerfugle. Cata-

logue of the Lepidoptera of Denmark: 128 pp. — Klampenberg, Denmark.

Lindquist, O. H., 1962. A Biological study of a New Leaf Miner on Birch, Nepticula lindquisti Freeman

(Lepidoptera: Nepticulidae), in Ontario. — Can. Ent. 94: 524—530, 9 figs.

McDunnough, J., 1939. Checklist of the Lepidoptera of Canada and the United States of America, pt

II. Microlepidoptera. — Mem. sth. Calif. Acad. Sci. 2: 1—171.

Meyrick, E., 1928a. Exotic Microlepidoptera 3: 640 pp. (Vols. 1—5, 1912—1937. Republished in 4 vols.

by Classey, England, 1969).

, 1928b. A revised Handbook of British Lepidoptera: pp. vi + 914. — London.

Petersen, W., 1930. Die Blattminierer-Gattungen Lithocolletis und Nepticula (Lepidoptera). Teil II: Nep-

ticula Zeller. — Stettin. ent. Ztg. 91: 1—82, 3 pls., 16 figs.

Ruggles, A. G., 1913. Notes on a chestnut-tree insect. — Science, Dec. 12, 1913: 852.

Schönherr, J., 1957. Rindenminen an Waldbäumen. — Wanderversamml. dt. Ent., 8: 127—133.

, 1958. Biologie und Morphologie van Ectoedemia liebwerdella Zimmerm. unter Berücksich-

tigung der überigen rinderminierenden Nepticuliden (Lep.). — Dt. ent. Z. (N.F.). 5: 1—71.

Schrank, F. von P., 1802. Fauna boica. 2 (2): 412 pp. — Nurnberg.

Svensson, I., 1966. New and confused species of Microlepidoptera. — Opusc. ent. 31: 183—202, 4 pls.

Wilkinson, C., 1979. A taxonomic study of the microlepidoptera genera Microcalyptris Braun and Fo-

moria Beirne occurring in the United States of America. — Tijdsch. Ent. 122: 59— 90.

Wilkinson, C. & M. J. Scoble, 1979. The Nepticulidae (Lepidoptera) of Canada. — Mem. Can. Ent.

Soc. 107: 1—118, 61 figs., 6 tables, 10 pls.

Zeller, P.C., 1839. Versuch einer naturgemässen Einteilung der Schaben. — Isis, Jena 23: 167—220.

, 1848. Die Gattungen der mit Augendeckeln versehenen blattminierenden Schaben. — Linn.

ent. 3: 248— 344.

——., 1873. Beiträge zur Kenntnis der nordamericanischen Nachtfalter besonders der Microlepidop-

teren. — Verh. zool. bot. Ges. Wien 23: 201—334, 2 pls.

Zimmermann, F., 1940. Eine neue Nepticulide aus Deutschland (Lep.). — Mitt. zool. Mus. Berlin. 24:

264—265, 1 fig.

WILKINSON & NEWTON: Ectoedemia in North America 89

Be) \\ \ N

Figs. 43—50. Ectoedemia species, external features. 43, E. lindquisti (Freeman); 44, E. canutus Wilkinson

& Scoble; 45, 7. piperella sp.n., &; 46, E. chlorantis Meyrick, © ; 47, E. obrutella (Zeller) Busck; 48. E.

quadrinotata (Braun); 49, E. rubifoliella (Clemens); 50, E. mesoloba Davis, 3.

90 TIJDSCHRIFT VOOR ENTOMOLOGIE

. DEEL 124. AFL. 2, 1981

ey ef

N N

Figs. 51—60. Ectoedemia species, external features. 51, E. trinotata (Braun), ¢; 52. E. marmaropa

(Braun), 9; 53, E. platanella (Clemens), &'; 54, 55, E. similella (Braun), g, Ohio form (54) and Florida

form (55); 56, 57, E. virgulae (Braun), 9, Ohio form (56), g, Florida form (57); 58, 59, E. nyssaefoliella

(Chambers), & (58), 2 (59); 60, specimen 8, ¢.

WILKINSON & NEWTON: Ectoedemia in North America

‘yosng vâvydooryd 4

‘EQ ‘yosng aD9uDISDI ‘7 ‘79 ‘Nosng 14214U19Y ‘7 ‘19 ‘sainyeoy [euI9]X9

‘

soloads DIU2P9019] ‘¢9— 19 ‘SF

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 2, 1981

Figs. 64—71. Leaf-mines. 64, 65, fossil leaf-mines from the lower Eocene (loaned from P. Opler); 66—

71, Ectoedemia mines; 66, E. trinotata (Braun) on Carya cordiformis; 67, E. marmaropa (Braun) on Rosa

woodsii; 68, E. platanella (Clemens) on Platanus occidentalis; 69, E. similella (Braun) on Quercus palustris;

70, E. virgulae (Braun) on Corylus americana; 71, E. nyssaefoliella (Chambers) on Nyssa sylvatica. All

natural size.

A SUPPLEMENT TO THE GENUS ECTOEDEMIA BUSCK

(NEPTICULIDAE: LEPIDOPTERA) IN NORTH AMERICA,

DEALING WITH SOME DIFFICULT SPECIES AND ALSO

SOME NEW ONES

CHRISTOPHER WILKINSON

Department of Animal Systematics and Zoogeography, Vrije Universiteit, Amsterdam

With nine text-figures

ABSTRACT

This article attempts to shed light on some nomenclatorial problems in Ectoedemia. Since the mono-

graph “The microlepidopteran genus Ectoedemia Busck (Nepticulidae) in North America” was written,

approximately five hundred additional specimens have been made available which have provided new

species and elucidated other information given here. Four problematic species are discussed, four new

species are described and five significant specimens are detailed.

INTRODUCTION

In every major revision a few species names cause nomenclatorial problems.

This is often because there is inadequate type material to check the names against,

or no known specimens at all. The question of what to do with doubtful species

usually arises during taxonomic revisionary work. The inclination is simply to cite

them and leave them as doubtful in the forelorn hope that someone in the future

will acquire more material. However, for the Nepticulidae it seems most unde-

sirable to have doubtful species remaining in the literature when their identity has

been suspect for more than one hundred years. Here the problems are explained,

together with suggestions of how one might solve the doubtful identity of certain

species. This is done with a view to seeking opinion, not only from other workers

in the group but also of those who have dealt with similar situations. Four prob-

lematic species in Ectoedemia are dealt with and another four in the revision of the

genus Stigmella in North A merica (in press).

One proposal, especially for those species described from a mine (no longer

extant), with perhaps some vague reference to a larva, is to designate a neotype

from an already known species — assuming that the descriptions of early stages

agree. The two names then become synonymised and remain so, because of the

neotype designation, and the former vague status disappears. The disadvantage is

that in these cases the species may become known by the hitherto doubtful species

name, if that is the older. A further suggestion which has been made to me and is

an extension of the same idea, is that all already synonymised species lacking type

94 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

specimens should have neotypes designated to ensure and confirm the synonymy.

I would be pleased to hear taxonomists views on the points raised in this article.

If there is wide opinion in favour of neotype designation for the doubtful species

discussed, especially those of Clemens and Chambers, they can perhaps be as-

signed in the Nepticulidae volume of the ‘*Moths of North America, North of

Mexico” (in preparation).

It is now very unlikely that the holotype of Ectoedemia canadensis (Braun) — the

only known specimen — is still in existence, but curators are kindly referred to

that species description (p. 94) in the hope that they will check their collections.

SUPPLEMENTARY CHECK LIST TO GENUS Ectoedemia

Locality

canadensis Braun, 1917 (with discussion on E. lindquisti) B.C., Canada.

grandisella Chambers, 1880 Texas, USA

anguinella Clemens, 1861 Kentucky, USA

platea Clemens, 1861 Kentucky, USA

coruscella sp.n. Illinois, USA

specimen 90 Illinois, USA

andrella sp.n. Maine, USA

specimens 245 and 251 Illinois, USA

reneella sp.n. Florida, USA

helenella sp.n. Florida, USA

specimen 180 Alabama, USA

acanthella Wilkinson & Newton Florida, USA

Methods, Abbreviations and References are the same as in the preceding mono-

graph.

All genitalia slides have been given USNM numbers, except where otherwise

stated.

Acknowledgements: My thanks are due to Mej. R. Kloos and B. J. van Cronen-

burg for technical assistance and artwork.

Ectoedemia canadensis (Braun) comb.n.

Nepticula canadensis Braun, 1917: 185.

Nepticula canadensis Braun; McDunnough, 1939: 107 (no. 9757).

Description. — External features: Head: palps dull white; antennae greyish; tuft

on front of head and vertex black or very dark brown; eye-caps and collar white.

Thorax black with brownish purple tinge. Abdomen dull black. Forewings: ground

colour of dorsal surface very dark brown or black; single fascia, irregular in

outline, white and medial in position. Hindwings: ground colour and probably

fringe, grey. Legs dull black.

Wing expanse: Holotype: 5.5 mm.

Host plant: Alnus tenuifolia (Mountain Alder).

Mine: An upper surface ophistigmatonome.

WILKINSON: Supplement to Ectoedemia 95

Diagnosis. — Similar to E. lindquisti but the head and vertex are ochreous in this

species and the fascia is more postmedial in position. The mine of E. lindquisti is a

blotch (ophionome) and occurs on Betula sp. See also discussion.

Discussion. — It is most unfortunate that the only known specimen of this

species — the holotype — is no longer to be found in Braun’s collection, and at-

tempts to locate it in other collections have failed. Braun was unable to say where

it had gone when asked about it a few years ago. I have been to the type locality,

Rogers Pass, B.C., during August 1971 and 1974 in the hope of finding new

material. The locality has changed considerably since 1915 when Annette Braun

made her collection, and although A/nus is widespread she only found specimens

in one restricted place. It is not surprising, therefore, that later searches were un-

successful. Thus there remains no known specimen to examine.

From my experience of Ectoedemia I have developed the strong feeling that ca-

nadensis almost certainly belongs to this genus and not to Stigmella. Further, it has

striking similarities with lindquisti and since no one likes doubtful species re-

maining in the literature for many years, I at first considered the possibility of des-

ignating a neotype and then synonymising the species with lindguisti. Freeman

(1962: 523) also recognised the similarity but the differences he gave regarding the

lustreless forewings of lindquisti as compared with the purple reflections of ca-

nadensis is not correct since in certain lights strong purple and bronze iridescences

can be seen on lindguisti, including the types. Braun in her 1917 description does

not say that canadensis has reflections, but I believe Freeman once told me he had

compared the holotype of canadensis with his species. The head colour of lindquisti

is mostly pale ochreous or slightly darker, but that of canadensis is black, however

this is not necessarily significant. In Europe the head colour varies with region and

when there is debate relating to colour one should further remember the less so-

phisticated equipment in use at the time of the early workers.

Whilst there may well be differences in externals it is necessary to turn to the

life cycle to find the most telling information. I have compared the relevant mines

and rearing records from Braun’s and Freeman’s collections, and find them to be

most significant. The fact that lindquisti larvae mine Betula whilst canadensis mine

Alnus sp. cannot be overlooked, especially as Freeman (1962) says that his species

has not been observed to feed on A/nus, even though alder occurs amongst birch in

areas where lindquisti is very abundant. However, most importantly, if we examine

the European fauna we discover that these two species have their direct homo-

logues and therefore are most unlikely to be synonyms. E. mediofasciella (Ha-

worth) in Europe corresponds with canadensis, whilst E. argentipedella (Zeller) cor-

responds with lindquisti. The North American species will key out using Emmets

(1976) European key and the similarities in mine structure and host plant prefer-

ences are most striking.

E. argentipedella and lindquisti feed solely on Betula and make a similar but un-

usual blotch mine. Compare the figures given by Van Frankenhuijzen and De

Vries (1979)!) with those of Lindquist (1962). The accounts of life cycle and bi-

ology also show a large degree of correspondence.

') Waarnemingen aan Ectoedemia argentipedella (Zeller), een mineermot op berk (Lep. Nepticulidae),

Ent. Ber. 39: 133.

96 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

E. mediofasciella is well known to feed on Betula and Corylus but has also been

discovered in the Alps feeding on Alnus viridis (E. v. Nieukerken, verbal commu-

nication). The mine corresponds with that of canadensis which starts as a short

contorted serpentine mine which later abruptly enlarges into a blotch. Compare

the figure in Emmet (1976, pl. 6, fig. 12) with that of Braun (1917, pl. 6, fig. 26) as

well as the descriptions of the adults. Furthermore both species may be described

as very local, sometimes common in a restricted area; locally it may also be host

specific.

We have found similar correspondence between other species of North

American and European Nepticulidae, including genitalia structures. This work is

continuing and may bring about a number of synonymies, some of which may

solve taxonomic difficulties such as we have with canadensis.

Thus three possibilities exist for clarification of the systematic position of ca-

nadensis. Obviously finding new material from the type locality would be very sat-

isfactory; synonymy with a European or North American species after designation

of a neotype; or the last possibility is the remote chance that the original holotype

still exists somewhere. So this constitutes a plea to those in charge of North

American micro-lepidopteran collections to have a careful look to see if this

specimen has finished up amongst those specimens in their care.

Distribution. — Canada: — British Columbia.

Type citation. — Holotype: Canada: B.874; Bear Creek, above Rogers Pass,

British Columbia, 6000 ft., August 19, 1915, A. Braun.

Biology. — Mine. A short, narrow serpentine mine which abruptly enlarges into

an irregular blotch, often filling the space between two lateral veins. Often as

many as a dozen mines on one leaf. The parenchyma is entirely consumed. Distri-

bution is very local.

Pupa. Cocoon dark brown in colour.

Voltinism. Probably univoltine.

Ectoedemia grandisella (Chambers) comb.n.

(fig. 1)

Nepticula grandisella Chambers, 1880: 193.

Nepticula grandisella Chambers; Dyar, 1903: 546 (listed).

Nepticula grandisella Chambers; Braun, 1917: 192.

Nepticula grandisella Chambers; McDunnough, 1939: 107 (no. 9770).

Description. — External features: g. Head: palps off-white; antennae brownish

extending about or a little more than two-thirds wing length; tuft on front of head

and vertex ochreous to dirty brown; eye-caps white. Thorax and abdomen brown,

greyer beneath. Forewings: ground colour of dorsal surface brown speckled with

darker brown; fringe brown. Hindwings: ground colour and fringe brownish grey.

Legs brown.

Wing expanse: Holotype: 10 mm.

Genitalia: ¢ (fig. 1). Pseuduncus with single broad flattened lobe. Gnathos: V-

shaped; transverse ventral plate with medial process; dorso-lateral arms broad and

weakly arcuate. Vinculum: lateral arms and ventral plate broad. Saccus as wide as

WILKINSON: Supplement to Ectoedemia 97

reconstructed

Fig. 1. Ectoedemia grandisella (Chambers). Reconstructed ¢ genitalia.

ventral plate, weakly bilobed. Valves reaching beyond the pseuduncus, width not

varying much throughout length, inner margin somewhat concave. Transtillae:

lateral arms narrow, ventral arms not long but narrow; transverse bars fused to

form a continuous strap. Aedeagus: broad and regular in width, length approxi-

mately equal to that of genitalia capsule; vesica with cornuti as small denticles

evenly distributed and with a plate of minute papillae; anellus comprising a pair of

canine tooth-like spines as shown in fig. 1b.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — The wing markings are typical of the bark mining species. The

background colour may be somewhat darker than other species or this may be due

to the age of the specimen. In the genitalia, the valves seem to have a character-

istic shape, as do the spines on the aedeagus.

Discussion. — This species, because of its large size and genitalia structure is

likely to be a bark miner. The colour and other external features are in keeping

with this suggestion. The species is transferred to Ectoedemia from Stigmella

(‘’Nepticula’’) for the first time.

Distribution. — USA: — Texas.

Material examined. — ¢ Holotype: U.S.A.: “Texas Cham. “Nepticula gran-

disella” “Chambers” Type 1302”; slide no. CNC 3496; in MCZ.

Biology. — Immature stages unknown.

98 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

Ectoedemia anguinella (Clemens) comb.n.

Nepticula anguinella Clemens, 1861: 85.

Nepticula anguinella Clemens; Clemens in Stainton, 1872: 175.

Nepticula anguinella Clemens; Braun, 1917: 204.

Nepticula anguinella Clemens; MeDunnough, 1939: 107 (no: 9780).

Discussion. — For 120 years this species has only been known from the original

description of a few lines about the mine and larva. No type material was pre-

served and no one has been aware of collecting the species since.

Because the larva is described as having “ten square dark brown or blackish

spots” this suggests to me that the species belongs to Ectoedemia, this character

being typical of immature instars of the genus. The black spots are lost by the time

the final stage larva is reached. Therefore, we might assume that the mines de-

scribed by Clemens were also incomplete. The “narrow serpentine tract which is

filled or discoloured throughout its length by black excrement” could, in a mature

mine, have continued by opening into a blotch so producing a typical ophistigma-

tonome. When a mine remains narrow and/or becomes discoloured by a mess of

blackish excrement, it often indicates that the larva has been parasitised and

therefore these may not be true larval characters.

It seems strange that with a type locality like Kentucky and a host-plant like oak

that no one has collected it since, if indeed, it has a separate identity. Having rea-

lised that this species is described from an immature Ectoedemia larva on oak, it

enables a more accurate comparison to be made. In fact, Clemens’ original de-

scription corresponds in all points with the juvenile larva of similella and therefore

it seems desirable to synonymise the two names. To make this permanent would

necessitate the designation of a neotype for anguinella which would be an adult ex-

ample of similella. Unfortunately anguinella would be the senior name and

therefore the one to be adopted.

The advantage of the scheme is obvious — it would fix a name which has been

in doubt for 120 years, and if it seems a desirable procedure it will be adopted in

the forth’coming Nepticulidae volume of the ““Moths of North America, North of

Mexico”.

Ectoedemia platea (Clemens) comb.n.

Nepticula platea Clemens, 1861: 85.

Nepticula platea Clemens; Clemens in Stainton, 1872: 175.

Nepticula platea Clemens; Braun, 1917: 204.

Nepticula platea Clemens; McDunnough, 1939: 107 (no: 9781).

Discussion. — There is less evidence for the identity of this species than for an-

guinella and not so much justification for regarding it as Ectoedemia. On the other

hand, it may be said that that provides all the more reason to do something pos-

itive to remove it from its dubious status. There is nothing in the original de-

scription to suggest that this “species” represents anything other than a healthy

mature larva of the preceding species anguinella. The larva is described as purplish

with a pale green vascular line and a row of reddish brown dorsal dashes. Ectoe-

WILKINSON: Supplement to Ectoedemia 99

demia quinquella Beddell in Europe is yellow with a row of large spots on the

venter, as in anguinella. These spots are shed early in the final instar and the larva

is then pale greenish white with a bright green gut line and a chain of small linear

dashes on the venter — the larva mines ventral side up. This is now similar to

platea, except for the background colour — purple. As Braun (1917: 204) points

out, the purple colour of the larva is not conclusive, as the colour is often pro-

duced in larvae which feed on leaves with autumnal colouration.

I have examined the larvae and/or descriptions of the twelve or so north

American Nepticulidae which feed on oaks, or have been recorded on them. Most

can be eliminated for one reason or another as possibly conspecific with platea; of

the rest the most feasible and likely proposition, on present evidence, is that angui-

nella and platea are synonyms — the descriptions, therefore, representing different

stages in the growth of the larvae. If this seems a reasonable proposition a formal

synonymy can be made in the “Moths of North America, North of Mexico”.

Ectoedemia coruscella sp.n.

(fig.2)

Description. — External features: g. Head: palps off-white; antennae buff; tuft

on front of head and vertex brown; eye-caps white; collar brown. Thorax and ab-

domen light brown, shiny. Forewings: ground colour of dorsal surface buff or light

brown, heavily speckled with dark brown, iridescent scales; these are less dense

sub-basally, and sub-terminally a pair of light buff triangular spots exist without

Fig. 2. Ectoedemia coruscella sp.n., & genitalia.

100 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

the darker scales; one spot is costal, the other anal; fringe brownish-grey, lustrous.

Hindwings: ground colour and fringe shiny brownish grey. Legs buff.

Wing expanse: Holotype: 7 mm.

Genitalia: & (fig. 2). Pseuduncus with single, rounded but flattened lobe.

Gnathos: W-shaped but central projection unusually long. Vinculum: broad lat-

erally and ventrally. Saccus bilobed but not markedly so. Valves arising high on

capsule and extending just beyond pseuduncus; inner margin highly irregular with

medial papillate protuberance. Transtillae: with lateral arms well developed;

ventral arms extending to edge of saccus; transverse arms medially constricted.

Aedeagus: flask-shaped; vesica with extremely spinose cornutus and papillate

plate as in fig. 2b; anellus comprising two pairs of bifurcate spines.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — This species is externally typical of other members of its group,

but can be diagnosed from others treated here and in the foregoing revision by the

pair of triangular postmedial wing patches. The male genitalia shows the valves,

aedeagus and perhaps the gnathos to be characteristic. -

Discussion. — This species is similar to a number of bark miners, but the ab-

sence of palmate anellar plates excludes this species from obrutella. On the other

hand, the aedeagus is far more spinose than that of acanthella. The vinculum and

saccus also have a different shape and the valves have a unique inner margin.

Perhaps mesoloba has the most similar male genitalia to those of coruscella but me-

soloba has a smaller central boss on the gnathos and somewhat smaller valves. The

wings appear to lack the triangular patches of coruscella.

Distribution. — USA: — Illinois.

Material examined. & Holotype: USA: Illinois, “Putnam Co., Aug. 14.1960;

M.O. Glenn”; slide no. 21817; specimen no. 250; in USNM.

Biology. — Unknown. Possibly bark miner which may explain why a northern

states species is only now discovered, and by so little material. It is not surprising

that the mines of bark miners have remained largely unnoticed and therefore the

life cycles are mostly unknown.

Ectoedemia specimen 90

(fig. 3)

Description. — External features: &. Head: palps buff; antennae light brown;

tuft on front of head and vertex darker brown; eye-caps white; collar brown.

Thorax and abdomen brown with some white scales. Forewings: ground colour of

dorsal surface brown, irrorate and speckled with white scales, a patch of white

medial in position and smaller flecks elsewhere. Hindwings: ground colour and

fringe brownish-grey. Legs buff.

Wing expanse: ¢: 8 mm (1 specimen).

Genitalia: 3 (fig. 3). Pseuduncus with single broad and flattened lobe. Gnathos:

W-shaped but central boss approximately half length of outer arms. Vinculum:

broad laterally and ventrally. Saccus only weakly bilobed. Valves arising one third

WILKINSON: Supplement to Ectoedemia 101

ft,

le,

Fig. 3. Ectoedemia specimen 90, ¢ genitalia.

up genital capsule and extending barely as far as pseuduncus; inner margin pa-

pillate. Transtillae: lateral arms short and robust; ventral arms long, extending

beyond capsule; transverse arms united to form a strap. Aedeagus: as long or

longer than capsule; vesica with spinose cornutus and papillate plate; anellus com-

prising a number of very long spines as in fig. 3b.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — This specimen appears to have highly diagnostic characters in the

genitalia, even if not in the externals. Whilst the gnathos and valves are similar to

the preceding species coruscella, they are not identical and the aedeagus is very

different. E. coruscella has characteristic spots on the wings.

Discussion. — The male genitalia of this species are quite different from those

of all other species dealt with here or in the preceding monograph. The nearest are

those of the preceding species which also comes from the same locality. More

material is obviously urgently needed to clarify the species these specimens rep-

resent. There is no information as to how this material was obtained.

Distribution. — USA: — Illinois.

Material examined. — USA: Illinois, Putnam Co., 1g, 14.vii.1968? (or 19607).

M. O. Glenn; slide no. 20752; specimen no. 90 in USNM.

Biology. — Immature stages unknown.

102 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

Ectoedemia andrella sp.n.

(fig. 4a,c)

Description. — External features: 3. Head: palps whitish buff; antennae buff;

tuft on front of head white darkening to deep buff on vertex; eye-caps white;

collar dark brown. Thorax and abdomen brown. Forewings: ground colour of

dorsal surface dark brown, lighter proximally, as is each individual scale; weakly

oblique medial fascia white; fringe white terminally, buff elsewhere; all weakly lus-

trous. Hindwings: ground colour and fringe greyish buff; a patch of buff hair scales

probably androconia can be seen baso-ventrally. Legs buff.

Wing expanse: Holotype: 6 mm.

Genitalia: & (fig. 4a,c). Pseuduncus rounded but prolonged posteriorly.

Gnathos: W-shaped with long central projection. Vinculum: broader laterally than

ventrally. Saccus small and bilobed. Valves arise low down on capsule and extend

a little beyond it; tapering but regular inner margin. Transtillae: small lateral arms,

longer ventral arms and well developed transverse arms forming a lateral strap.

Aedeagus as in fig. 4c; flask-shaped; vesica with small denticles and papillate

plate; anellus comprising two large spines, tapering markedly to a point.

Female: Unknown, but see specimen 251 following.

Host plant: Not known.

Mine: Not known.

Diagnosis. — Similar to ulmella but differs externally in the more dowdy ap-

pearance and weaker lustre of andrella. In the male genitalia the pseuduncus is

Fig. 4. Ectoedemia andrella sp.n., & genitalia, a and c; Ectoedemia specimen 245, ¢ genitalia, a and b.

WILKINSON: Supplement to Ectoedemia 103

broader and larger; the gnathos is more W-shaped and the valves more evenly ta-

pering than in u/mella.

The following two specimens (245 and 251) probably also belong here. The male

genitalia only differ in the spines of the aedeagus, and externally the iridescence

and tufts of the head differ as does the locality. These points may not be signif-

icant.

Discussion. — Specimens 245 and 251 are in poor condition and do not warrant

being made paratypes, although most of the characters can easily be seen.

Distribution. — USA: — Maine.

Material examined. — ¢ Holotype: USA: “Maine, Augusta, July 9 1940; A. E.

Brower, at light” Slide no. 21819; specimen no. 98 in USNM.

Biology. — Unknown.

Ectoedemia specimens 245 and 251

(figs. 4a, b, 5)

Description. — External features: 4, ©. Head: palps and antennae buff; tuft on

front of head and vertex dark brown; eye-caps shining white; collar brown. Thorax

and abdomen shining dark brown. Forewings: ground colour of dorsal surface

uniform dark brown with a single shining white medial fascia; fringe dark brown

but terminally white. Hindwings: ground colour and fringe greyish brown. Legs

dark brown.

Wing expanse: 2: 5.5 mm (1 specimen).

Genitalia: & (fig. 4a,b). Pseuduncus rounded but prolonged posteriorly.

Gnathos: W-shaped with long central projection. Vinculum: broad laterally but not

so marked ventrally. Saccus small, bilobed. Valves arise low down on capsule and

extend beyond it, tapering, inner margin regular. Transtillae: small lateral arms,

longer ventral arms and well developed transverse arms forming a lateral strap.

Aedeagus: flask-shaped; vesica with minute denticles and papillate plate; anellus

comprising two large spines slightly bifurcate at tip, as shown in fig. 4b.

Female (fig. 5). Ductus bursae broad, not long, colliculum appears to be present

as an indistinct loop amongst folds and associated lobes. Bursa copulatrix: very

large with a pair of reticulate signa approximately equal in size. Anterior apo-

physes broad and unusually long. Posterior apophyses straight and narrow, short

extending no further anteriorly than anteriores.

Host plant: Not known.

Mine: Not known.

Discussion. — Judging from the male genitalia these specimens almost certainly

belong to the new species andrella. They only seem to differ in small details in the

aedeagus, and these differences may only be a consequence of viewing from dif-

ferent angles, since the capsules appear identical. The tufts on the front of the

head and vertex are much darker than in andrella.

Distribution. — USA: — Illinois.

Material examined. — USA: Illinois, Putnam Co. 13,19, 14.&23.iv.1967, M.O.

Glenn; slide nos. 21821 and 21822; specimen nos. & 245, 9 251 in USNM.

Biology. — Unknown.

104 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

Fig. 5. Ectoedemia specimen 251, © genitalia.

Ectoedemia reneella sp.n.

(fig. 6)

Description. — External features: &. Head: palps, antennae off-white with little

brown; tuft on front of head and vertex buff; eye-caps white; collar off-white.

Thorax and abdomen lustrous white tending to buff. Forewings: ground colour of

dorsal surface and fringe shining white tending to buff with very few scattered

darker buff or brown scales. Hindwings: ground colour and fringe as forewings.

Ventral surface both wings as upper, but more buff scales distally. Legs off-white.

Wing expanse: Holotype: 6 mm.

Genitalia: 3 (fig. 6). Pseuduncus smoothly rounded and not prolonged poste-

riorly. Gnathos: W-shaped. Vinculum: comparitively narrow laterally, broad ven-

WILKINSON: Supplement to Ectoedemia 105

Fig. 6. Ectoedemia reneella sp.n., & genitalia.

trally. Saccus large, bilobed. Valves arise high up capsule and do not extend

beyond pseuduncus; evenly curved papillate inner margin. Transtillae: small

lateral arms and long ventral arms; well developed transverse arms tapering to-

wards centre. Aedeagus as in fig. 6b, c: flask-shaped, vesica with many denticles,

papillate plates and strong spinose lobe. Anellus comprising two pairs of large

spines posteriorly and two pairs of digitate processes with spines.

Female: Not known.

Host plant: Not known.

Mine: Not known.

Diagnosis. — This species is unusually light coloured, having no dark coloured

parts. As such this could be diagnostic. The aedeagus of the male genitalia is also

characteristic having two pairs of anellar spines and two pairs of spinose digitate

plates. No other species known so far appears to have so much spiny ornamen-

tation and is without internal lobes to the valves.

Distribution. — USA: — Florida.

Material examined. — & Holotype: USA “Florida, Siesta Key, Sarasota Co.,

1.v.1960 C. P. Kimball”; slide no. 21823; specimen no. 144 in USNM (fig. 6c). Par-

atype: In USNM — USA: same data as Holotype; 1g, 18.iv.1960 (Kimball); slide

no; 18243; specimen no. 159 (fig. 6b).

Biology. — Unknown.

Ectoedemia helenella sp.n.

(fig. 7)

Description. — External features: &. Head: palps pale buff; antennae brown;

106 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

Fig. 7. Ectoedemia helenella sp.n., & genitalia.

tuft on front of head buff, and vertex, dark brown; eye-caps, collar, thorax and ab-

domen brown, shiny. Forewings: ground colour of dorsal surface buff with

scattered dark brown scales, iridescent bronze; lighter patches occur subbasally,

shining silver especially on wing tip; fringe brown, lustrous. Hindwings: ground

colour and fringe shine brownish grey. Legs buff.

Wing expanse: &: 6 mm (1 specimen). Holotype: 6 mm.

Holotype: 4 mm.

Genitalia: & (fig. 7). Pseuduncus with single, small, rounded, flattened lobe.

Gnathos: W-shaped. Vinculum: not so broad laterally but proportionately more so

ventrally. Saccus very broad and not bi-lobed but slightly depressed medially.

Valves long, tapering and extending beyond pseuduncus; inner margin relatively

smooth and regular. Transtillae: lateral arm well developed and leading to very

long ventral arms; transverse arms easily separated. Aedeagus: broad, regular;

vesica with spinose cornutus and papillate plate; anellus comprising bifurcate

spines and a pair of spinose palmate plates as in fig. 7b.

Female: Not known.

Host plant: Not known.

Mine: Not known.

WILKINSON: Supplement to Ectoedemia 107

Diagnosis. — This species is extremely small and in that resembles mesoloba.

However, in as far as one can judge by externals this species is much darker than

mesoloba and the valves of the genitalia differ. The genitalia are more similar to

those of heinrichi than perhaps the illustrations would suggest, however, the size,

wing-span and externals are markedly different.

Discussion. — Perhaps most closely related to heinrichi but very much smaller.

Distribution. — USA: — Florida.

Material examined. — ¢ Holotype: USA: “Florida, Siesta Key, Sarasota Co.,

23.11.1954, C. R. Kimball”; slide no. USNM 21818; specimen no. 155 in USNM.

Paratype: In USNM — USA: Florida, Siesta Key, Sarasota Co., 1 g, 17.iv.1953

(Kimball); slide no. USNM 21825; specimen no. 166.

Biology. — Unknown.

Ectoedemia specimen 180

(fig. 8)

Description. — External features: 9. Head: palps buff with dark brown scales;

antennae dark brown with lighter annulations; tuft on front of head and vertex

dark brown to black; eye-caps deep gold paling at edges; collar dark brown and

black. Thorax and abdomen dark brown to black. Forewings: ground colour of

dorsal surface uniform brown, lighter than thorax; individual scales basally grey

which would give lighter appearance to worn specimens; fringe and wing edge

dark buff; all weakly lustrous. Hindwings: ground colour uniform grey, markedly

different from fringe and wing edge which are buff. Legs buff with dark brown

scales. 3 unknown.

Wing expanse: 9: 12.5 mm (1 specimen).

Genitalia: 9 (fig. 8). Ductus bursae broad with lobes and spicules; colliculum

not ring-shaped. Accessory duct arising from ductus bursae and spiral medially.

Bursa copulatrix long, with fine pectinations. Signum double, comprising equal-

sized reticulate patches. Anterior apophyses long, arcuate and narrow. Posterior

apophyses much longer, narrower and straight.

Host Plant: not known.

Mine: not known.

Diagnosis. — An extremely large species with some, so far, unique characters

including the golden eye-caps, the intensely dark tufts and the striking grey

hindwings edged in buff. Perhaps most like obrutella, heinrichi and phleophaga but

the foregoing characters are clearly diagnostic. The genitalia are also similar to

obrutella.

Distribution. — USA:- Alabama.

Material examined: — USA: “Alabama, Open Pond Cpgd. 20 mi. S. Andalusia,

Covington Co., 19, 18.iv.1976 J. B. Heppner; slide no. 3029; specimen no. 180 in

USNM.

Biology. — Unknown. Barkminer?

Ectoedemia specimen no. 170 g. Might conceivably belong to the same species

at this 9. It has white eye-caps and is smaller (11 mm). Unfortunately it is broken

and the abdomen is missing.

108 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

Fig. 8. Ectoedemia specimen 180, 9 genitalia.

Material examined. — USA: “Alabama, Black Warrior River, 13 mi. SW

Greensboro, Hale Co., 1g, 23.iv.1976 J. B. Heppner”; specimen no. 170 in

USNM.

Ectoedemia acanthella Wilkinson and Newton

(fig. 9)

Description. — This species has been described earlier (Wilkinson & New-

ton, 1981, p. 75). Two additional male specimens probably belong here, which

WILKINSON: Supplement to Ectoedemia 109

Fig. 9. Ectoedemia acanthella, Wilkinson & Newton, & genitalia.

only became available after the original description was made and the holotype

had been returned to the USNM. Therefore they have not yet been directly com-

pared. The external features agree with the acanthella description. Fig. 9a, b, c,

shows the male genitalia of these specimens and their similarity with fig. 35 (acan-

thella holotype) in Wilkinson & Newton (1981) can be seen. Certain features in fig.

9 not shown in fig. 35 probably clarify details of the male genitalia rather than rep-

resenting characters which separate two different species. If this inference is

correct then it extends the previously cited distribution to include Maine and Ar-

kansas.

Distribution. — USA :— New Jersey, Maine, Arkansas.

Material examined. — USA: — Maine, Round Pond, 14, 26.vii.?, TI3R12; slide

no. 21826; specimen no. 176. Arkansas, Mt. Magazine, Brown Spring, 1g,

9.vi.1964 (Clarke); slide no. 21827; specimen no. 151 in USNM.

110

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 3, 1981

INDEX TO GENERA, SPECIES, SUBSPECIES AND OTHER IMPORTANT NAMES

(The index serves the two papers by Wilkinson & Scoble and Wilkinson)

acanthella 75, 108

altella 58

amelanchierella 69

andrella 102

anguinella 98

argentipedella 95

argyropeza 43

argyropeza argyropeza 44

argyropeza downesi 45

Artaversala 36

bosquella 72

bucculatrix 45

canadensis 94

canutus 45

castaneae 80

chlorantis 84

clemensella 55

corusella 99

corylifoliella 60

Dechtiria 30

downesi 45

Ectoedemia 34

flavipedella 31

Fomoria 35

Glaucolepis 36

grandisella 96

heinrichi 78

helenella 105

Juglandifoliella 48

liebwerdella 30

lindquisti 61

Lyonetia 43

Manoneura 36

marmaropa 49

maximella 51

mediofasciella 95

mesoloba 85

Microcalyptris 35

Nepticula 29

nyssaefoliella 67

Obrussa 35

obrutella 72

Oligoneura 36

Opostega 72

ostryaefoliella 60

phleophaga 81

piperella 77

platanella 51

platea 98

populella 41

quadrinotata 70

reneella 104

rubifoliella 63

similella 56

Stigmella 29, 35

subbimaculella 36

Tinea 36

Trifurcula 30

trinotata 46

turbidella 43

ulmella 65

virgulae 59

Zimmermannia 30

DEEL 124 AFLEVERING 4 1981

MUS, COMP, ZOOL

LIBRARY ~

AUGS 1991

TIJDSCHRIFT u»

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

P. K. ChaupHurI, D. K. Guna and S. K. Das Gupra. — Taxonomic studies of Chi-

ronominae (Diptera, Chironomidae) from India. The genus Polypedilum

Kieffer, p. 111—147, figs. 1—14.

Tijdschrift voor Entomologie, deel 124, afl. 4 Gepubliceerd 20-VII-1981

TAXONOMIC STUDIES OF CHIRONOMINAE

(DIPTERA, CHIRONOMIDAE) FROM INDIA.

THE GENUS POLYPEDILUM KIEFFER !)

P. K. CHAUDHURI, D. K. GUHA and S. K. DAS GUPTA 2)

Entomology Laboratory, Department of Zoology, University of Burdwan, Burdwan 713 104, India

With 14 text-figures

ABSTRACT

Fourteen species of non-biting midges belonging to the genus Polypedilum Kieffer are described

or revised from six geophysical zones of West Bengal, India. Polypedilum albipalpus, P. ascium, P.

chaudhurii, P. flagellatum, P. insolitum, P. lineatum, P. lucidum, P. nudiceps, P. numerus, P.

pseudoflagellatum, P. rufomarginalis and P. tripunctum are described as new species. Polypedilum

annulatipes Kieffer and P. griseoguttatum Kieffer, previously known from India and Africa, are

recorded and revised in the light of modern taxonomic usages. A key to the species occurring in the

state of West Bengal is presented. A check-list of Indian species of Polypedilum is also provided.

INTRODUCTION

Non-biting midges of the genus Polypedilum Kieffer are one of the most

common and widespread group of insects having a fairly wide latitude of

ecological tolerance, most of them occurring in lakes and lowland situations.

The immatures are known to inhabit running water and places usually avoided

by mosquitoes.

The material was collected mainly during the period of May, 1976 and June,

1979 with light-traps and sweeping net, while a few, as denoted, were reared in

our laboratory.

Polypedilum Kieffer is a well-defined genus of the family Chironomidae. It

was denoted as a genus for the first time by Kieffer (1913) on the basis of a

Palaearctic species, Polypedilum emarginatum Kieffer, which was fixed as its

type-species. Edwards (1929) treated it as a subgenus of genus Chironomus

Meigen. Goetghebuer (1936, 1937), while accepting the idea of Kieffer, fixed

Chironomus nubeculosus Meigen as its genotype in place of that made by

Kieffer. Later, the species was found to be similar to that of Kieffer. Townes

(1945) considered Polypedilum as a broad genus with three subgenera including

Pentapedilum Kieffer and Polypedilum Kieffer and this classification was

followed by most workers. Freeman (1958, 1961) accepted the view of Townes

1) Contribution no. 18 to the study of Chironomidae from the Univ. of Burdwan.

2) Professor & Head of the Department of Zoology, Presidency College, Calcutta 700073, India.

111

112 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

and synonymised a large number of genera with this genus. Sublette & Sublette

(1973) re-established the generic status of Polypedilum and placed about forty

species from the Oriental Region in this genus and also kept many more species

unplaced for obvious reasons. Prior to this study, only seven species had been

reported from India (Kieffer, 1910, 1911d and 1913).

For explanation of morphological terminologies and abbreviations, the works

of Frommer (1967), Guha & Chaudhuri (1979) and Saether (1971, 1974 and

1977) have been considered here. The figures in the first brackets in the text of

the paper indicate those found in the majority instead of the average number.

Types and other specimens of this study are at present kept in the collections of

insects at the Entomology Laboratory, Department of Zoology, University of

Burdwan, Burdwan and will be deposited in the British Museum (Nat. Hist.),

London, U.S.N.M., Washington D.C., Zoological Survey of India, Calcutta

and Zoologische Staatssammlung, München (West Germany) in due time.

Again, help without which this paper could not have been completed was

generously accorded by our co-workers, among whom special mention should

be made of Dr. D. C. Sinharay, Mr. R. K. Debnath (Raniganj), Mr. P.

Chaudhuri, Head of the Department of Zoology, University of Burdwan, is

ical Survey of India, Calcutta, is sincerely thanked for kindly permitting us to

examine the specimens at the Zoological Survey of India, Calcutta. Prof. D. K.

Chaudhuri, Head of the Department of Zoology, University of Burdwan, is

greatly acknowledged for giving laboratory facilities and permission to name a

species after him.

TAXONOMIC ACCOUNT

Polypedilum albipalpus n.sp.

(figs. 1 a—h)

Material: Holotype d (Type no. 78, B.U. Ent.), India, West Bengal, Rani-

ganj, 10.v.1977, D. K. Guha. Allotype ©, Burdwan, 11.11.1978, A. Chatterjee.

Paratypes: 3 4, 2 2, Burdwan, July, 1978, A. Chatterjee; 1 d, 1 2, Memari,

July, 1978, P. K. Chaudhuri; 278, 3 2, Ranigany, August, 00973888

Chaudhuri.

Remarks: This species is named Polypedilum albipalpus on the basis of the

white colour of its palpomere V. It shows similarities in wing and male hypopygi-

um with Polypedilum numerus n.sp. described in this paper and P. aegyptium

Kieffer (1925) described by Freeman (1958) from Africa, but it can easily be

distinguished as a distinct and valid species by, (i) colour and chaetotaxy of

maxillary palp, (ii) spots of wing, (iii) fore tibial scale, (iv) swollen inturned anal

point, and (v) appendage 2 with long setae.

Male: Body length 2.04 (1.98—2.24, n = 6) mm.

Head: Pale brown in colour. Vertex with 9 setae (postocular 1, outer verticals

7, inner verticals 1). Corona without setae. Clypeus with 14—16 setae, clypeal

ratio 0.75. Maxillary palp (fig. 1a) pale brown, palpomere V whitish, palpomere

III with a preapical pit bearing sensilla, length ratio of palpomeres I—V

CHAUDHURI, GUHA & DAs GUPTA: Polypedilum from India 113

Figs. la—h. Polypedilum albipalpus n.sp. g: a, maxillary palp; b, thorax; c, wing; d, tibial ends of

fore, mid and hind legs; e, hypopygium 2; f, antenna; g—h, genitalia.

5:6:14:15:27, L/W ratio 3.5. Eyes almost contiguous, bare, reniform, extended

dorsally, extension being 0.15 mm long. Frontal tubercle absent. Antenna

brown, densely setaceous, setae longer and dense on proximal flagellomeres and

gradually become shorter and sparse towards distal flagellomeres, apex of

flagellomere XII without seta, length ratio of flagellomeres I—XII

10:5:5:6:7:7:7:7:7:7:7:65; AR 0.92. Pedicel ratio 0.8. CA 0.65, CP 1.43.

Thorax (fig. 1b): Yellow in colour. Antepronotum thin without V-shaped

emargination. Acrostichals 10 arranged in pairs, dorsocentrals 11 in one row,

114 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

humeral 1, prealars 4. Scutellum with 10 setae, postscutellum light brown and

bare.

Wing (fig. 1c): Cloudy with veins light yellow. Brachiolum with 1 seta. R with

14, R, with 9, R,,; with 12 setae at the apical part; R,,, meets C very close to R,

being 0.02 mm from R,; ending of R,,, and C subacute; r—m proximal to f—Cu;

An ends below f—Cu. Wing with pale irregular spots distributed as follows: (i)

one at the base of cell R,,;, near r—m, and one linear at the apex of it, often

extending to cell R,,3, (ii) one very small spot near the base of M,,, and one

longitudinal streak present below R,,;, (ii) one covering the base of cell M,,,,

and (iv) one at the middle and apex of cell An. Sensory organ 1 at the base of R,,..

Squama with 5 setae. Haltere light brown at stem and knob and bare. Wing

length 1.0 (0.98—1.14, n = 6) mm, breadth 0.36 (0.34—0.38, n = 6) mm. CR

0.91, VR 1.25. Legs (fig. 1d): Yellow to light brown. Fore tibial scale in the shape

of isobilateral triangle with a spine-like apex. Spur of mid and hind tibia equal in

length, being 0.043 mm long, slightly curved, ratio of length of spur to the apical

diameter of mid and hind tibia 10 : 9. Tarsomere V of all legs dark. Pulvilli small

and split; empodium small.

Proportions and ratios of leg segments

Fe Ti ta, ta, ta; tay tas LR TR

Fore 24 16 28 14 10 7 5 1975 —

Mid 29 22 14 11 8 4 4 0.63 —

Hind 30 25 18 12 9 5 5 0.72 ES

Abdomen: Yellow with uniformly distributed setae. Segment VIII slightly

constricted at the base.

Hypopygium (fig. le): Anal point narrow at the base and middle part wide,

apex inturned, swollen and pointed, its base with 5—6 setae on each side.

Gonocoxite with 8—10 (10) setae; gonostylus long, slightly bowed with 6—7

setae at its inner apical margin, gonostylus with 7—9 (9) setae over it. Append-

age 1 elongated with S—6 gently curved setae at its apex; appendage 2 long,

elongated, little bent, with 7—8 (7) incurved setae at its apex. Transverse

sternapodeme 0.12 mm long, lateral sternapodeme 0.09 mm long, coxapodeme

0.038 mm long and phallapodeme 0.064 mm long. HR 0.71, HV 2.55.

Female: Body length 1.64 (1.56—1.74, n= 6) mm.

Similar to male with usual sex differences. Antenna (fig. 1f) light brown,

flagellomere V narrowed apically and with few sensilla, length ratio of flagello-

meres IV 19:13:15:9:26, AR 0.47. Wing and its chaetotaxy as in male. Wing

length 1.16 (1.12—1.18, n = 6) mm, breadth 0.42 (0.40—0.44, n = 6) mm.

Genitalia (figs. 1g—h): Notum 0.11 mm long. Coxasternapodeme bowed.

Gonapophysis VIII divided into a broad dorsomesal and finger-like ventrolate-

ral lobe. Apodeme lobe filamentous. Gonocoxite IX with 2 setae. Postgenital

plate V-shaped. Cerci with 10—12 long and numerous small setae. Labia

without microtrichia. Seminal capsules (fig. 1g) equal, oval with a neck, measur-

ing 0.047 mm by 0.03 mm; ducts of seminal capsules with one loop and open

separately.

CHAUDHURI, GUHA & DAs GUPTA: Polypedilum from India 115

Figs. 2a—f. Polypedilum annulatipes Kieffer 8; a, maxillary palp; b, wing; c, tibial ends of fore, mid

and hind legs; d, hypopygium ®; e, antenna; f, genitalia.

Polypedilum annulatipes Kieffer

(figs. 2 a—h)

Chironomus annulatipes Kieffer, 1910: 234.

Polypedilum annulatipes; Kieffer, 1921: 100; 1922: 30.

Polypedilum octostictum Kieffer, 1921: 101; 1922: 39.

116 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Polypedilum octomaculatum Goetghebuer, 1934: 195; 1936: 488.

Polypedilum annulatipes; Freeman, 1958: 277—278.

Material: 12 d, 14 ©, India, Orissa, Barkuda lake, 1.x.1922, N. Annandale

(C/O Z.S.I., Calcutta); 5 d, 2 ®, West Bengal, Burdwan, 11.111.1978, A.

Chatterjee; 3 4, Bankura, 27.ix.1977, S. Basak; 10 d, Coochbehar, 3.xi.1974,

S. K. Das Gupta; 3 d, Digha, 15.1v.1976, P. K. Chaudhuri; 3 d, Canning,

2.1v.1976, A. De. i

Remarks: This species appears to be distributed in Africa and India. It was

described by Kieffer as Chironomus annulatipes in 1910 and as Polypedilum

annulatipes in 1921 and 1922. The redescription of the species as presented here

is based on the identified collections received from the Zoological Survey of

India, Calcutta, and on those collected during the present investigation. Corre-

lating the findings of the present authors with that of Kieffer, it may be inferred

that P. annulatipes is a distinct species of the genus Polypedilum due to (i)

presence of single spur in mid and hind tibia, (ii) absence of frontal tubercles, (iii)

antepronotum not collar-like and (iv) abdominal segment VIII constricted at the

base.

Male: Body length 2.66 (2.52—2.69, n = 10) mm.

Head: Yellowish brown in colour. Vertex light yellow with 9 (postocular 1,

outer verticals 5, inner verticals 3) setae. Corona without setae. Clypeus with

14-16 (15) setae, clypeal ratio 1.5. Maxillary palp (fig. 2a) pale to yellow,

palpomere III with a preapical rounded pit bearing 3 cilia-like sensilla, palpo-

mere V longest with a straight apical seta, length ratio of palpomeres I—V

8:9:26:31:46, L/W ratio 4.33. Eyes bare, crescentic, extended dorsally, exten-

sion being 0.07 mm long. Frontal tubercle absent. Antenna brown to dark

brown, flagellomeres dumb-bell-shaped, flagellomere XII longest, pear-like,

setae on proximal flagellomeres very long and gradually becoming shorter

distad; length ratio of flagellomeres I-XII 2:2:2:2:2:2:2:2:2:2:2:40; AR 1.81.

Pediceliratio ls; EAU 53, CP 13:

Thorax: Yellow with two longitudinal dark brown bands on mesonotum.

Antepronotum very thin without V-shaped emargination. Acrostichals 16 in

pairs, dorsocentrals 14 in single row, humerals 2—3 (2), prealars 6. Scutellum

with 8 setae in oblique row, postscutellum very dark and bare.

Wing (fig. 2b): Whitish with veins yellow and conspicuous. Brachiolum with 1

seta. R with 22, R, with 22 and R,,, with 33—35 setae; R,,, meets C very close to

R, being 0.05 mm away from it; ending of R,,; and C subacute; r—m distinct and

proximal to f—Cu; An ends below f—Cu. Wing with several dark and light grey

irregular spots and bands distributed as follows: (i) cell R,,; with one broad

linear band covering the proximal half up to r—m, one very small near the apical

margin and a third one at the middle, (ii) cell M,,, with one linear band or broad

streaks along the mid part of M,,, and one small spot sometimes found to be

present at the apical margin of the cell, (iii) cell M ,,, with two spots, one covering

the area of f—cu extending along Cu, up to the margin and another near the

apical end below M,,,, (iv) cell An with three spots, one broad at the middle of

the cell, one slightly above and away from it and the third covering the area

between the end of Cu, and its margin. Sensory organ 1 at the base of R,. Squama

CHAUDHURI, GUHA & DAs GUPTA: Polypedilum from India 117

with 10 setae. Haltere bare with yellow stem and knob. Wing length 1.51 (1.51—

1.53, n = 10) mm, breadth 0.48 (0.44—0.48, n = 10) mm. CR 0.97, VR 1.27.

Legs (fig. 2c): Yellow in colour. Femur and tibia of all legs with a proximal and

subapical broad bands. Fore tibial scale elongated and blunt, bearing 0—1 (1)

seta. Spur of mid tibia single, curved apically, 0.051 mm long, ratio of length of

spur to the apical diameter of mid tibia 14:13; spur of hind tibia single, curved

apically, 0.059 mm long, ratio of length of spur to the apical diameter of hind tibia

16:14. Pulvilli split; empodium as long as claw.

Proportions and ratios of leg segments

Fe Ti ta, ta, ta; ta, tas LR TR

Fore 46 30 50 17 12 9 7 1.67 =

Mid 51 43 23 10 6 4 5 0.54 _

Hind 53 50 22 10 7 4 5 0.44 _

Abdomen: Intensively dark brown in colour, terga uniformly setaceous.

Segment VIII narrowed at base.

Hypopygium (fig. 2d): Anal point 0.03 mm long with sharply pointed apex,

anal point with S—6 setae at each basal margin and a few setae at the base.

Gonocoxite small with 6—8 long setae; gonostylus long, narrowed distally,

bearing 15—20 setae over it and 10—12 (12) setae along its inner margin.

Appendage 1 well developed, pad-like and minutely setaceous; appendage 2

long, bent with uniform diameter, apex slightly bent inward, bearing 15 incurved

setae. Laterotergite with 10—12 (10) long setae; transverse sternapodeme 0.17

mm long, lateral sternapodeme 0.14 mm long, coxapodeme 0.07 mm long and

phallapodeme 0.69 mm long. HR 0.6, HV 1.75.

Female: Body length 1.90 (1.90—1.92, n=8) mm.

Similar to male with usual sex differences. Antenna (fig. 2e) dark, flagello-

mere V longest, length ratio of flagellomeres I—V 20:14:15:10:34; AR 0.40.

Pedicel ratio 0.12. Wing with spots and bands similar to those of male but darker.

Wing length 1.44 (1.41—1.44, n=10) mm, breadth 0.40 (n=10) mm. Genitalia

(fig. 2f): Notum 0.012 mm long. Gonocoxapodeme wide and rounded, not

joined mesally. Coxasternapodeme transverse with a single broad bend. Gona-

pophysis VIII divided into a broad, posteriorly directed, dorsomesal lobe and a

tongue-like ventrolateral lobe bearing brush-like setae. Apodeme lobe moder-

ately developed. Tergum IX with 5—6 long setae on each side. Gonocoxite with

4—S (4) setae.

Postgenital plate prominently protruding and conical. Cerci toe-like and

minutely setaceous. Labia elongated with numerous microtrichia. Seminal

capsules (fig. 2f) unequal, oval or spherical with or without little neck, measuring

0.037 mm by 0.037 mm and 0.041 mm by 0.03 mm; ducts of seminal capsules long,

coiled with one or two loops or bends and open in common to the vagina.

Polypedilum ascium n.sp.

Material: Holotype d (Type no. 79, B.U. Ent.), India, West Bengal, Bolpur,

118 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 3a—e. Polypedilum ascium n.sp. $: a, maxillary palp; b, thorax; c, wing; d, tibial ends of fore,

mid and hind legs; e, hypopygium.

18.vi.1977, D. Chatterjee. Paratypes; 18 d, data same as holotype, 18—

20.vi.1977; 1 6, Burdwan, 3.iii.1979, S. K. Nandi; 4d, Calcutta, 23.1.1978, D.

K. Guha; 3 4, Rampurhat, 29.iv.1978, D. Roy.

Remarks: This species is named as Polypedilum ascium on the basis of the

axeshaped appendage 2 of its male hypopygium. It comes near to Polypedilum

acutus (var. acutus) Kieffer (1915) from the British Isles in certain features of the

male hypopygium but on the basis of the detailed morphology, such as colour of

the body, chaetotaxy of leg, wing and appendage 2 of male hypopygium, it is

considered here a distinct species.

Male: Body length 2.4 (2.2—2.62, n=10) mm.

Head: Yellow to brown in colour. Vertex light brown with 8 (postocular 1,

outer verticals 5, inner verticals 2) setae. Corona without setae. Clypeus with

16—18 setae, clypeal ratio 1. Maxillary palp (fig. 3a) brown, palpomere III with a

preapical pit bearing 3 small sensilla, palpomere V with 3 apical setae, length

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 119

ratio of palpomeres I—V 6:7:14:18:28,L/W ratio 2.8. Eyes bare, reniform,

extended dorsally, extension being 0.09 mm long. Antenna brown, densely

setaceous, setae longer at proximal flagellomeres and gradually becoming

shorter distad; length ratio of flagellomeres I—XII 3:3:3:3:3:3:3:3:3:3:3:62; AR

1.947 Pedieelratio33. CA 105) CP i'd.

Thorax (fig. 3b): Brown with dark brown patches on mesonotum. Anteprono-

tum very thin without any emargination. Acrostichals 12 arranged in pairs,

dorsocentrals 9 in single row, humerals 2, prealars 3. Scutellum with 6 setae,

postscutellum dark brown with a middle suture and bare.

Wing (fig. 3c): Smoky with veins yellow. Brachiolum with 1 seta. R with 11, R,

with 5 and R,,; with 16—18 setae; R,,; meets C a little away from R,, distance

being 0.08 mm; ending of R,,; and C subacute; r—m oblique and proximal to f—

Cu; An ends before f—Cu. Sensory organ 1 each on r—m and at the base of R,,..

Squama with 5 setae. Haltere with light brown stem and dark brown knob and

bare. Wing length 1.26 (1.14—1.35, n=10) mm, breadth 0.36 (0.32—0.40,

n=10) mm. CR 0.94, VR 1.33.

Legs (fig. 2d): Yellow to light brown in colour. Fore tibial scale elongated and

blunt, often with a short abrupt point. Spur of mid tibia single, bent at the apex,

0.05 mm long, ratio of length of spur to the apical diameter of mid tibia 13:11;

hind tibia also with a spur, 0.06 mm long, ratio of length of spur to the apical

diameter of hind tibia 14:12. Pulvilli very short, bilobed with a median, setaceous

empodium.

Proportions and ratios of leg segments

Fe Tht ta, ta, ta; ta, tas LR TR

Fore 38) 21 29 12 10 8 4 1.4 —

Mid 32 29 15 9 i 3 4 0.52 =

Hind 36 15 8 6 4 4 0.47 1.88

Abdomen: Dark brown in colour, terga uniformly setaceous. Segment VIII

constricted widely at the base.

Hypopygium (fig. 3e): Anal point 0.04 mm long, apex subacute slightly

inflated with 5—6 setae at each basal margin and 10—12 setae at the base.

Gonocoxite short with 5 long setae; gonostylus tapered with 5 long setae at its

inner apical margin and with 15—20(18) long setae over it. Appendage 1 long,

straight with a bent tip or incurved, bearing 3 setae at its base; appendage 2 long,

axe-shaped, moderately bent at the middle, with 1 long and 6 incurved setae at its

apex. Laterotergite with 16—18(18) long setae, transverse sternapodeme 0.14

mm long with a small concavity at the middle, lateral sternapodeme 0.077 mm

long, coxapodeme 0.043 mm long and phallapodeme 0.055 mm long. HR 0.36,

HV 1.83.

Female: Unknown.

Polypedilum chaudhurii n.sp.

(figs. 4a—g)

Material: Holotype & (Type no. 80, B.U. Ent.), India, West Bengal, Burd-

120 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

wan, 11.iii.1978, A. Chatterjee. Allotype 2, data same as holotype. Paratypes:

76, 29, data same as holotype; 2 d, Calcutta, 23.1.1978, D. K. Guha; 1 d,

Chinsura, 7.iii.1972, P. K. Chaudhuri; 26, 22, Durgapur, 7—8.i.1978, S.

Chaudhuri.

Remarks: This species is named after Prof. D. K. Chaudhuri, Head of the

Department of Zoology, University of Burdwan, India, in appreciation of his

sustained interest, enthusiasm and efforts towards the development of Biologi-

cal Sciences in this University. It can be distinguished from all other species

described in this paper by, (i) mottled brown thorax and its chaetotaxy, (ii)

colour pattern of the wing, (iii) colour band of the legs, (iv) fore tibial scale, (v)

two rows of scutellar setae, (vi) anal point with two projections, and (vii)

appendages 1 and 2 of male hypopygium. This species comes near Polypedilum

griseoguttatum Kieffer (1921) from Africa and Australia in regard to the wing

and male hypopygium and to P. prolixitarsis Lundstròm (1916) as described by

Edwards (1929) from Finland, but the above characters are sufficient for

distinguishing from them and for determining it as a distinct and valid species.

Male: Body length 2.51 (2.40—2.62, n=8) mm.

Head: Brown in colour. Vertex brown with 13 (postoculars 2, outer verticals 7,

inner verticals 4) setae. Corona without setae. Clypeus with 14—16(16) setae,

clypeal ratio 1. Maxillary palp (fig. 4a) light brown to pale, palpomere III with a

shallow preapical pit bearing 2 long sensilla, palpomere V with an apical seta;

length ratio of palpomeres I—V 10:10:28:32:52; L/W ratio 4.66. Eyes bare,

crescentic, well extended dorsally, extension being 0.13 mm long. Frontal

tubercle absent. Antenna pale brown, setae long in proximal flagellomeres and

shorter distad, apical part of flagellomeres with only a few setae; length ratio of

flagellomeres I-XII 2:2:2:2:2:2:2:2:2:2:2:40; AR 1.54. Pedicel ratio 1.37. CA

OS HER:

Thorax (fig. 4b): Mottled brown in colour. Antepronotum very thin and

without V-shaped dorsal emargination. Acrostichals 20 arranged in pairs,

dorsocentrals 16 arranged in single row, humerals 2, prealars 6. Scutellum with

20 setae in two rows, postscutellum black-brown and bare.

Wing (fig. 4c): Spotted with veins yellow. Brachiolum with 1 seta. R with 18,

R, with 15 and R,,; with 20 setae; R,,, meets C close to R,, distance being 0.04

mm from R,; ending of R,,; and C subacute; r—m distinct and proximal to f—

Cu; An ends before f—Cu. Wing with irregular grey spots distributed as follows:

(i) cell R,,; with two large and one small spots, smaller spot near the apical

margin, larger spots lying at the base of cell near r—m and at the middle of cell,

(ii) cell M with one at the middle and cell M,,, with one linear spot parallel to

proximal half of M,,,, (ili) cell M,,, with one broad spot covering the basal

‘margin, (iv) cell An with one broad spot at the middle, another broad spot at

apex of cell and the third at the tip on An. Squama with 9 setae. Haltere with

brown stem and knob and bare. Wing length 1.48 mm, breadth 0.44 cm. CR 0.96,

VR 1.2.

Legs (fig. 4d): Yellow brown to dark brown. Femur dark brown with apical

pale band, fore and hind tibia yellow, mid tibia dark brown with an apical and

basal yellow band. Fore tibial scale triangular and pointed at the apex, bearing

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 121

Figs. 4a—g. Polypedilum chaudhurii n.sp. 8: a, maxillary palp; b, thorax; c, wing; d, fore, mid and

hind legs; e, hypopygium. 2; f, antenna; g, genitalia.

3—4 long setae at the base. Spur of mid tibia bent, 0.055 mm long, ratio of length

of spur to the apical diameter of mid tibia 15:12; hind tibia also with bent spur,

0.059 mm long; ratio of length of spur to the apical diameter of hind tibia 16:15.

Pulvilli small and split, empodium normal.

122 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Proportions and ratios of leg segments

Fe ali ta, ta, ta; tay tas OR TR

Fore 19 11 18 9 7 4 5 1.63

Mid 20 18 10 9 Y 4 5 0.55 _

Hind 22 20 11 10 6 5 5 0.55 oil

Abdomen: Dark brown in colour and uniformly setaceous. Segment VIII

widely constricted at the base.

Hypopygium (fig. 4e): Anal point trifid, moderately stout, apex little nar-

rowed and curved with 6—7(6) setae at each side and numerous small setae at

base. Gonocoxite well developed with 12—14(13) long setae; gonostylus long,

straight, distal part narrowed with 12 long setae over it and 5 setae at its inner

apical margin. Appendage 1 well developed, with uniform width, gently bent at

the middle, bearing 4 setae and numerous small weak setae; appendage 2 long,

elongated, little bowed, apex with 12 incurved setae. Laterotergite with 4 long

setae, transverse sternapodeme 0.16 mm long, lateral sternapodeme 0.09 mm

long, coxapodeme 0.04 mm long and phallapodeme 0.074 mm long. HR 0.87,

HV 1.62.

Female: body length 1.65 (1.62—1.68, n=4) mm.

Similar to male with usual sex differences. Maxillary palp brown. Antenna

(fig. 4f) yellowish, flagellomere V dark; length ratio of flagellomeres I—V

26:21:22:14:38, AR 0.40. Wing with spots and veins similar to those of female.

Wing length 1.35 (n=4) mm, breadth 0.52 (0.52—0.54, n=4) mm. Genitalia (fig.

4g): Notum 0.14 mm long. Gonocoxapodeme broadened and joined distinctly at

the middle. Gonapophysis VIII divided into broad dorsomesal and a small

ventrolateral lobe. Apodeme lobe finger-like. Gonocoxite IX with 3 long setae.

Segment X with 8 setae on each side. Postgenital plate as small protuberance.

Cerci elongated. Labia with microtrichia. Seminal capsules oval or spherical,

subequal, measuring 0.092 mm by 0.059 mm and 0.085 mm by 0.051 mm; ducts of

seminal capsules straight with one or two bends, opening separately.

Polypedilum flagellatum n.sp.

(figs. Sa—h)

Material: Holotype d (Type no. 81, B.U. Ent.), India, West Bengal, Rani-

ganj, 25.iv.1977, D. C. Sinharay. Allotype 2, data same as holotype, date being

4.viii.1977. Paratypes: 4 d, Asansol, Sept., 1976, R. K. Debnath; 4 d, 2 2,

Bolpur, 18.viii.1977, D. Chaterjee; 8 6, 3 ®, Burdwan, June, 1977, S. K.

Nandi; 1 4, Rampurhat, 15.viii.1977, D. Chatterjee; 3 d, 3 2, Raniganj, 25—

28.1v.1977, D. C. Sinharay.

Remarks: The species is named Polypedilum flagellatum in view of the

presence of an unusually long flagellum-like seta at the apex of appendage 2 of

the male hypopygium. In certain morphological features, especially those

concerning the male hypopygium, it seems to be related to a number of species

such as Polypedilum bifalcatum Kieffer (1921, 1922) from Sudan (Africa), P.

brunneicornis Kieffer (1911) from Italy and P. flavescens Johannsen (1932) from

CHAUDHURI, GUHA & DAS GUPTA: Polypedilum from India 123

Figs. Sa—h. Polypedilum flagellatum n.sp. 6: a, maxillary palp; b, thorax; c, wing; d, tibial ends cf

fore, mid and hind legs; e, hypopygium. 2: f, antenna, g—h, genitalia.

Bogor, Java, Bali, Sumatra etc., but it can be considered a distinct species by

virtue of, (i) presence of seta in corona, (ii) comma-shaped eye, (iii) brachiolum

with 1 seta, (iv) light grey spots or bands on the wing and opaque haltere, (v) dark

brown abdomen, (vi) small and bare appendage 1 and long dorsoventrally

flattened appendage 2 bearing one unusually long and a number of usual

incurved setae at the apex, and (vii) longer anal point with its characteristic

chaetotaxy.

124 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Male: Body length 3.61 (3.19—3.91, n=10) mm.

Head: Dark brown to black in colour. Vertex dark brown with 16 (postocular

4, outer verticals 10, inner verticals 2) setae. Corona well developed with 4 setae.

Clypeus with 21 long setae, clypeal ratio 0.73. Maxillary palp (fig. Sa) dark

brown, palpomere III with a crescentic pit bearing 2 cilia-like sensilla; length

ratio of palpomeres IV 3:3:5:7:10, L/W ratio 3.33. Eyes bare, comma-shaped,

extended dorsally, extension being 0.11 mm long. Frontal tubercle absent.

Antenna brown to dark brown, densely setaceous, setae longer at the proximal

flagellomeres and gradually becoming shorter distad; length ratio of flagello-

meres I-XI 4:2:2:2:2:2:2:2:2:2:48; AR 2.0. Pedicel ratio 1.2. CA 0.55, CP 1.21.

Thorax (fig. 5b): Yellow with one longitudinal band in mesonotum and two

lateral bands. Antepronotum thin, collar-like without V-shaped emargination.

Acrostichals 10 in pairs, dorsocentrals 11, humerals 2—3 (3), prealars 2.

Scutellum with 10 setae arranged mostly in oblique row, postscutellum dark

brown and bare.

Wing (fig. 5d): Yellow in colour. Femur pale. Fore tibial scale blunt and

elongated bearing 4 long setae. Spur of mid tibia 0.034 mm long, ratio of length of

spur to the apical diameter of mid tibia 8:12; spur of hind tibia bent apically, 0.034

mm long, ratio of length of spur to the apical diameter of hind tibia 8:14.

Tarsomeres IV—V without long setae. Pulvilli split longitudinally into two

narrow lobes, empodium short, with setae.

Proportions and ratios of leg segments

Fe ii ta, ta, ta; tay tas LR TR

Fore 34 29 44 27 21 17 9 ILDIl =

Mid 40 35 22 11 7 6 4 0.62 —

Hind 40 41 33 17 15 9 5 0.80 1293

Abdomen: Dark brown in colour and heavily setaceous. Segment VIII deeply

constricted at the base.

Hypopygium (fig. 5e): Anal point 0.048 mm long and little narrowed with

slightly pointed end, bearing 4—5 setae at its basal margin. Gonocoxite stout,

narrowed and slightly attenuated at the apex, gonocoxite with 15—18 (18) long

setae; gonostylus broad at base and apex narrowed with 6 small setae at its inner

apical margin, gonostylus with 10—12 long setae over it. Appendage 1 bare,

gently bent at the middle, its base broad and gradually narrowed; appendage 2

stout, base narrowed, dorsoventrally flattened and apex wider with 15—16 (16)

long incurved setae, longest seta being 0.09 mm long. Laterotergite with 3 setae,

transverse sternapodeme 0.05 mm long, lateral sternapodeme 1.2 mm long,

coxapodeme 0.06 mm long, phallapodeme 0.08 mm long. HR 1.14, HV 2.20.

Female: Body length 2.61 (2.36—2.96, n=10) mm.

Similar to male with usual sex differences. Antenna (fig. 5f) light brown,

flagellomere V dark brown, AR 0.31. Wing colour, spots as in male but darker.

Wing length 1.4 (1.33—1.48, n=5) mm, breadth 0.51 (0.48—0.55, n=5) mm.

Genitalia (fig. 5 g—h): Tergum 0.12 mm long. Coxosternapodeme well devel-

oped with a notch and two bends. Gonocoxite VIII joined at the middle.

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 125

Gonapophysis VIII divided into a long dorsomesal and a well developed

ventrolateral lobe. Apodeme lobe weak. Tergum IX with 20—25 (24) setae,

tergum X with 22—24 setae. Gonocoxite IX with 3 setae. Postgenital plate a

small protuberance. Cerci long and broad with 20—25 setae. Labia without

microtrichia. Seminal capsules (fig. 5g) oval, 0.044 mm and 0.048 mm; ducts of

seminal capsules straight without any loops and opening separately.

Polypedilum griseoguttatum Kieffer

(figs. 6a—d)

Polypedilum griseoguttatum Kieffer, 1921: 100; 1922: 36.

Polypedilum hieroglyphicum Kieffer, 1921: 101; 1922: 38.

Polypedilum decem-maculatum Goetghebuer, 1934: 195.

Polypedilum vanbemmeli Kruseman, 1949: 254.

Polypedilum griseoguttatum; Freeman, 1958: 284; 1961: 709.

Material 2 ore bundwanee2 e193 DARK GUAM Calcutta 6222

13.11.1979, P. K. Chaudhuri.

Remarks: This species was described by Freeman (1958, 1961) in both sexes

from Africa and Australia. The present material, though based on only five

microslide mounts of males, resembles the above species as described by

Freeman. Despite the differences which exist in regard to the male hypopygium

it seems unwise to consider this a separate species unless exotic material of

the species is available for comparison. Pending the availability of the female

specimens this species better be treated as Polypedilum griseoguttatum Kieffer.

Male: Body length 3.84 mm.

Head: Yellowish brown in colour. Vertex with 9 (postocular 1, outer verticals

6, inner verticals 2). Corona without setae. Clypeus with 18—19 long setae,

clypeal ratio 1.2. Maxillary palp brown in colour, length ratio of palpomeres I—

V 8:9:28:32:40, L/W ratio 4.6. Eyes bare, reniform, extended dorsally, exten-

sion being 0.11 mm long. Frontal tubercle absent. Antenna dark brown, densely

setaceous, setae longer at proximal flagellomeres and gradually becoming

shorter distad, apex of flagellomere XII without setae; length ratio of flagello-

meres I—XII 2:2:2:2:2:2:2:2:2:2:2:58, AR 2.64. Pedicel ratio 1.12. CA 0.54, CP

12%

Thorax (fig. 6a): Brown with dark brown bands laterad. Antepronotum very

thin without V-shaped dorsal emargination. Acrostichals 28 in pairs, dorsocen-

trals 18, humeral 1, prealars 7, prescutellars 4. Scutellum with 36 setae in two

rows, postscutellum dark brown and bare.

Wing (fig. 6b): Opaque with veins yellow in colour. Brachiolum with 1 seta. R

with 32, R, with 22, R,,, with 35—37 setae; R,,, meets C very close to R,,

distance being 0.05 mm from R,; ending of R,,; and C acute; r—m distinct and

proximal to f—Cu; An ends below f—Cu. Sensory organ 1 on r—m. Wing with

complicated pattern of dark grey spots distributed as follows: (i) one small spot

distad of cell R,,, touching the margin, (ii) three in cell R,,;, (iii) one at the apex

of cell M and M,,,, (iv) two spots in cell M,,, which appear continuous with

dark, bent streak, (v) one at the middle and apex and another smaller in between

126 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 6a—d. Polypedilum griseoguttatum Kieffer g: a, thorax; b, wing; c, tibial ends of fore, mid and

hind legs; d, hypopygium.

the two. Besides the above, a linear dark streak runs below and parallel to M...

Squama with 15 setae. Haltere bare with stem yellow and knob dark brown.

Wing length 2.07 mm, breadth 0.59 mm. CR 0.96, VR 1.12.

Legs (fig. 6c): Yellow to brown in colour. Fore legs yellow except tarsomeres

II—V, each of which bear a dark brown basal band. Fore tibial scale triangular

bearing a sharply pointed seta at the apex. Mid and hind tibia dark brown

distally. Spur of mid tibia dagger-like, slightly bent inward, 0.051 mm long, ratio

of length of spur to the apical diameter of mid tibia 14:16; spur of hind tibia also

like that of mid tibia, 0.062 mm long, ratio of length of spur to the apical diameter

of hind tibia 17:18. Tarsomeres II—V with dark brown band, tarsomere IV with

two apical spurs. Pulvilli of moderate length and split; empodium short.

Proportions and ratios of leg segments

Be Ti ta, ta, ta; ta, tas LR TR

Fore 24 19 30 21 17 12 5 1.58 =

Mid 28 23 14 8 6 3 2 0.65 ==

Hind 29 27 20 12 9 5 3 0.74 1.67

CHAUDHURI, GUHA & DAS GUPTA: Polypedilum from India 127

Abdomen: Dark brown. Terga uniformly covered with setae which are

arranged irregularly. Segment VIII deeply constricted at the base.

Hypopygium (fig. 6d): Anal point well developed, tongue-like, with a con-

striction at the middle. Gonocoxite well developed with 12—15 (14) long setae;

gonostylus long, narrowed towards apex, with 6 setae at its inner apical margin,

gonostylus with 20—22 (22) long setae over it. Appendage 1 stout, broad,

roughly clubbed, covered with numerous small setae over it; appendage 2 long,

bowed with narrow base and apex, bearing 2 long slightly incurved setae.

Transverse sternapodeme 0.23 mm long, lateral sternapodeme 0.13 mm long,

coxapodeme 0.059 mm long, phallapodeme 0.14 mm long. HR 0.68, HV 1.73.

Female: Unknown.

Polypedilum insolitum n.sp.

(figs. 7a—d)

Material: Holotype d (type no. 82, B.U. Ent.), India, West Bengal, Digha,

15.1v.1975, P. K. Chaudhuri. Paratypes: 5 d, same data as holotype.

Remarks: The species is named Polypedilum insolitum in view of some

unusual characters like the presence of a frontal tubercle and absence of setae on

squama. It shows close resemblances with Lauterborniella fuscoguttata Kieffer

(1922) from France in its wing and fore tibia, and strong similarities with

Polypedilum longinervis Kieffer (1922) from (the then French) Cameroons in

respect of the abdomen and male hypopygium but by virtue of having the

characters like, (i) presence of frontal tubercle, (ii) absence of stripes on the

thorax, (ili) spots of wing, (iv) femoral bands, (v) appendage 1 bare and bilobed,

(vi) appendage 2 flattened without any incurved setae, (vii) segment VIII of

abdomen deeply constricted at the base and (viii) gonostylus with only 5 small

setae at its inner apical margin, the inclusion of the species in the genus

Polypedilum and distinctiveness as a separate species seems to be assured.

Male: Body length 3.62 mm.

Head: Dark brown in colour. Vertex with 8 (postocular 3, outer verticals 4,

inner verticals 1) setae. Corona without setae. Clypeus with 20 long setae,

clypeal ratio 1.12. Maxillary palp brown to dark brown, palpomere II with a

preapical pit bearing 2 flagella-like sensilla; length ratio of palpomeres I—V

4:4:9:6:11, L/W ratio 3.0. Frontal tubercles present. Eyes bare, reniform but

slightly extended dorsally, extension being 0.09 mm long. Antenna dark brown

to brown, length ratio of flagellomeres I—IV (rest of the flagellomeres broken in

all specimens) 5:3:3:3. Pedicel ratio 0.97.

Thorax (fig. 7a): Brown with dark brown patches. Antepronotum thin with a

small dorsal emargination. Acrostichals 12 arranged in pairs from tip of mesono-

tum, dorsocentrals 10 in a row, humerals 0, prealars 2—3 (3). Scutellum with 10

setae arranged irregularly, postscutellum dark brown and bare.

Wing (fig. 7b): Pale with veins yellow and conspicuous. Brachiolum with 2

setae. R with 14, R, with 18 and R,,; with 30—32 setae; R,,; meets C almost near

the middle, the distance being 0.11 mm from R,; ending of R,,, and C acute; r—

128 TIIDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 7a—d. Polypedilum insolitum n.sp. 9: a, thorax; b, wing; c, tibial ends of fore, mid and hind legs

with fore and mid femora and tibia; d, hypopygium.

m distinct and proximal to f—Cu; An ends before f—Cu. Wing with several

irregular grey spots distributed as follows: (i) 1 pair broad, unequal spots or

bands over the basal and middle of cells R,,; and M,,, with its vicinities, (ii) 1

each near the apical margin of cell R,,; and cell M,,,, (iii) 3 unequal spots in cell

M,,,, (iv) 1 large and 1 small linear spot placed one above the other at the distal

end of cell An. In addition to the above, linear dark streaks are observed along

the distal part of M,,, and in cell An. Sensory organ 1 on r—m. Squama bare.

Haltere bare and yellow in both stem and knob. Wing length 1.48 (1.48—1.50,

n=2) mm, breadth 0.48 (n=2) mm. CR 0.95, VR 1.20.

Legs (fig. 7c): Brown to dark brown in colour. Fore femur and tibia brown with

a distal broad dark brown band, femur and tibia of mid and hind legs with a broad

proximal and small preapical dark brown band. Fore tibial scale with a sharp

pointed spur 0.045 mm long. Spur of mid tibia bent at the apex, ratio of length of

spur to the apical diameter of mid tibia 4:4; spur of hind tibia 0.09 mm long, ratio

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 129

of length of spur to the apical diameter of hind tibia 6:4. Tarsomeres of all legs

brown to dark brown. Pulvilli split longitudinally. LR 1.48 in fore leg, LR 0.77 in

mid and LR 0.90 in hind legs. Abdomen: Brown with alternate dark brown bands

on terga I to VII and with uniformly distributed long setae. Segment VIII deeply

constricted at the base.

Hypopygium (fig. 7d): Anal point small, 0.025 mm long and rod-like and

narrowed distad. Gonocoxite with 16—18 (18) long setae; gonostylus small,

little expanded and abruptly attenuated, with 5 small setae along its inner apical

margin. Appendage 1 short, base broad, distal part slightly bent and bare;

appendage 2 elongated, dorsoventrally compressed, reaching almost the middle

of gonostylus, appendage 2 also without incurved setae but 1—2 setae near its

blunt apex. Laterotergite with 6—7 setae, transverse sternapodeme 0.22 mm

long, lateral sternapodeme 0.09 mm long, coxapodeme 0.06 mm long, phallapo-

deme 0.07 mm. HR 1.2, HV 3.27.

Female: Unknown.

Polypedilum lineatum n.sp.

(figs. 8a—e)

Material: Holotype d (Type no. 83, B.U. Ent.), India, West Bengal, Rani-

ganj, 28.x.1977, D. K. Guha. Paratypes: 8 9, Burdwan, 28—30.x.1977, A.

Chatterjee; 4 d, Siliguri, 21—22.iv.1979, S. K. Das Gupta; 3 6, Triveni,

4.1v.1979, S. Sarkar.

Remarks: The above species is named Polypedilum lineatum in view of its

conspicuous linear dark spots running parallel to M,,,. It seems close to

Polypedilum tonnoiri Freeman (1961) from Australia in regard to the wing and

the male hypopygium. The characters (i) chaetotaxy of thorax, nature of spots

on the wing, (ii) infuscation on the apical margin of wing, (ili) dark streak below

M,,,, (iv) bands on the mid and hind femur with its chaetotaxy, (v) size and

structure of anal point and (vi) appendages 1 + 2, and certain other characters

clearly justify its treatment as a distinct species. It should be noted also that the

flagellomeres of male antenna number thirteen instead of twelve as in other

species.

Male: Body length 1.81 mm.

Head: Dark brown in colour. Vertex dark brown with 11 (postocular 2, outer

verticals 8, inner vertical 1) setae. Corona with 1 seta. Clypeus with 16 setae,

clypeal ratio 1. Maxillary palp (fig. 8a) brown, palpomere III with a preapical pit

bearing sensilla, palpomere V with 1 straight apical seta, length ratio of palpo-

meres I—V 7:6:20:24:32, L/W ratio 4. Eyes bare, semicircular, little extended

dorsally, extension being 0.09 mm long. Frontal tubercles absent. Antenna dark

brown, setaceous, setae longer at proximal flagellomeres and shorter distad,

length ratio of flagellomeres I-XIII 5:6:8:8:8:8:8:9:9:9:9:9:75; AR 0.71. Ped-

icel ratio OZEAN EIERN 2:

Thorax (fig. 8b): Yellow to light brown with a median and two lateral brownish

bands on mesonotum. Antepronotum dark, very thin without dorsal emargina-

130 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 8a—e. Polypedilum lineatum n.sp. g: a, maxillary palp; b, thorax; c, wing; d, tibial ends of fore,

mid and hind legs; e, hypopygium.

tion. Acrostichals 16 in two rows, dorsocentrals 14 in one row, humeral 0,

prealars 3. Scutellum with 10 setae arranged in oblique row, postscutellum dark

brown and bare.

Wing (fig. 8c): Cloudy. Brachiolum with 1 seta. R with 13, R, with 6 setae at

the basal half and R,,; with 21 setae; R,,, meets C close to R,, distance being 0.05

mm from R,; ending of R,,, and C acute; r—m slightly proximal to f—Cu; An

ends below f—Cu. Wing with faint grey irregular spots distributed as follows: (i)

one at the base of cell R,,; near r—m, (ii) one near f—Cu in cell M,,,, (iii) one

linear dark streak running parallel to M,,,, (iv) one broad interrupted spot at the

middle and another at the apex of cell An, in some cases the apical margin of cell

R,,; and M,,, appear to be infuscated. Sensory organ 1 at the base of R, and

R,,s. Squama with 3 setae. Haltere brown, and bare. Wing length 0.97 (0.94—

0.99, n=6) mm, breadth 0.34 (0.33—0.35, n=6) mm. CR 0.97, VR 1.48.

Legs (fig. 8d): Yellow to light brown with numerous long weak setae. Femur of

mid and hind legs with a middle apical/subapical light greyish band. Fore tibial

scale triangular and pointed bearing 1 long seta. Spur of mid tibia slightly curved,

0.04 mm long, ratio of length of spur to the apical diameter of mid tibia 11:10;

spur of hind tibia equal to that of mid tibia, ratio of length of spur to the apical

diameter of hind tibia 11:12. Pulvilli longitudinally split with a long setaceous

empodium.

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 131

Proportions and ratios of leg segments

Fe Ti ta, ta; taz tay tas LR TR

Fore 31 17 Dj 12 8 5) 4 1.59 —

Mid 35 27 15 10 7 5 4 0.56 —

Hind 35 31 19 13 9 6 5) 0.61 1.46

Abdomen: Light brown with a longitudinal band at the middle of terga.

Segment VIII constricted at the base.

Hypopygium (fig. 8e): Anal point quite long (0.03 mm) long, little bent and

dilated at the apex, anal point with 10—12 setae at each basal margin. Gonocox-

ite small with 4—6 setae; gonostylus very long, little bowed with 15—17 (17)

small setae at its inner apical margin, gonostylus with 8—10 long setae. Append-

age 1 well developed, wider at the apex, with numerous small setae; appendage 2

long, slightly bent near base, with 2 long and 8 incurved setae. Transverse

sternapodeme 0.11 mm long, lateral sternapodeme 0.05 mm long, coxapodeme

0.03 mm long, phallapodeme 0.04 mm long. HR 0.68, HV 1.63.

Female: Unknown.

Polypedilum lucidum n.sp.

(figs. 9a—e)

Material: Holotype d (Type no. 84, B.U. Ent.), India, West Bengal, Burd-

wan, 12.11.1978, S.K. Nandi. Paratypes: 10 d, data same as holotype; 4 d,

Calcutta (reared in the laboratory), D. K. Guha; 2 ®, Panagar, 7.iv.1977, K.

Sinha.

Remarks: The present species is named as Polypedilum lucidum because of

the presence of pale infuscations or spots on the wing. It shows similarities in

relation to wing with Polypedilum longicrus Kieffer (1911) from Africa. Similar-

ly, it shows resemblances with P. numerus n.sp. described in this paper and

P. swlovatum Johannsen (1932) in the male hypopygium. However, P. lucidum

can be identified as a separate species by the colour pattern of wing and haltere

bands on legs, structure of fore tibial scale and the male hypopygium.

Male: Body length 2.29 (2.22— 2.40, n=8) mm.

Head: Brown to dark brown in colour. Vertex brown with 9 (postocular 1,

outer verticals 7, inner vertical 1) setae. Corona without setae. Clypeus with

22—24 (24) setae, clypeal ratio 1.15. Maxillary palp (fig. 9a) pale yellow to

brown; palpomere III with a preapical pit bearing 2 sensilla; palpomere V pale

with 1 long apical seta; length ratio of palpomeres I—V 7:9:21:27:42, L/W ratio

3.5. Eyes bare, ovoid, well separated, extended dorsally, extension being 0.14

mm long. Frontal tubercle absent. Antenna dark brown, densely setaceous,

setae longer at base and gradually become shorter distad, apex of flagellomere

XII without long seta; length rato of flagellomeres IE XII

2 DD D DDD EP DBs INNA, De dicelatiorl 2 SNCANOESD ACE NED

Thorax (fig. 9b): Brown in colour. Antepronotum very thin, collar-like

without V-shaped emargination. Acrostichals 16 in pairs, dorsocentrals 23,

132 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 9a—e. Polypedilum lucidum n.sp. 8: a, maxillary palp; b, thorax; c, wing, d, fore, mid and hind

legs with their tibial ends; e, hypopygium.

humerals 2, prealars 5. Scutellum with 10 setae in a simple transverse row,

postscutellum dark brown and bare.

Wing (fig. 9c): Cloudy with veins pale. Brachiolum with 1 seta. R with 24, R,

with 15 and R,,; with 28 setae; R,,, meets C very close to R, being 0.05 mm from

R,; ending of R,,; and C subacute; r—m distinct and proximal to f—Cu; An ends

a little proximal to f—Cu. Wing with infuscations often in the form of spots or

bands distributed as follows: (i) one spot at the base of cell R,,;, (ii) one linear

dark streak parallel to M,,, in cell M,,,, (iii) one spot at the base of cell M,, , near

f—Cu and (iv) two linear spots at the proximal half of cell An, and one spot at the

apex of the cell. Squama with 7 setae. Haltere with stem light brown and knob

dark brown and bare. Wing length 1.36 (1.29—1.48, n=8) mm, breadth 0.44

(0.39—0.48, n=8) mm. CR 0.97, VR 1.3.

Legs (fig. 9d): Brown to dark brown in colour. Fore femur with an apical dark

brown band, femur of mid and hind legs each with a broad median and a small

apical dark brown band; tibia of both mid and hind legs with a basal and apical

pale band. Fore tibial scale triangular with a short pointed tip. Spur of mid tibia

0.04 mm long, ratio of length of spur to the apical diameter of mid tibia 13:12;

CHAUDHURI, GUHA & DAS GUPTA: Polypedilum from India 133

spur of hind tibia slightly bent at the apex, 0.05 mm long, ratio of length of spur to

the apical diameter of hind tibia 16:14. Tarsomeres slightly pale. Pulvilli lobe-

like, split at the middle; with a long empodium equal to the claws.

Proportions and ratios of leg segments

Fe Ti ta, ta, taz tay tas LR TR

Fore 17 11 21 12 9 7 4 LO —

Mid 19 16 10 4 3 2 0.62 —

Hind 19 18 18 8 6 4 3 0.72 1.62

Abdomen: Light brown in colour, terga uniformly setaceous, setae on lateral

side arranged in rows. Segment VIII broadly constricted at its base.

Hypopygium (fig. 9e): Anal point with 3 setae at each basal margin and a fair

number of setae at its base. Gonocoxite well developed with 12—15 long setae;

gonostylus long, narrowed down with 5 setae at its inner apical margin, gonosty-

lus with 17—19 setae over it. Appendage 1 clublike bearing numerous small

setae; appendage 2 long, bowed, apex wide, bearing 8 long incurved setae.

Transverse sternapodeme 0.19 mm long, lateral sternapodeme 0.09 mm long,

coxapodeme 0.03 mm long, phallapodeme 0.07 mm long. HR 0.8, HV 1.55.

Female: Unknown.

Polypedilum nudiceps n.sp.

(figs. 10a—h)

Material: Holotype d (Type no. 85, B.U. Ent.), India, West Bengal, Darjeel-

ing, 5.ix.1977, S. K. Das Gupta. Allotype 9, data same as holotype. Paratypes:

50 d, 36 ?, Darjeeling, July-Oct., 1977, S. K. Das Gupta; 7 d, 3 ?, Ghum,

June, 1973, D. C. Sinharay; 5 d., Jorbangla, Sept., 1978, P. Chaudhuri; 11 3, 10

MP OVS, A ORL SG, | 2, Wons, Sent, IWS, S. Walle 8,3 2;

Lebong, Sept., 1978, P. K. Chaudhuri.

Remarks: The above species is named Polypedilum nudiceps in view of bare

appendage 1 of the male hypopygium. It comes close to Polypedilum bipustula-

tum Freeman (1958) from Africa in some morphological features specially in

relation to the male hypopygium but the characters like, (i) colour of the

specimens including haltere, (ii) absence of stripes in the mesonotum, (iii)

haltere without any setae, (iv) shape of appendage 1, (v) appendage 1 without

seta, (vi) appendage 2 with 2 long apical setae, (vii) long anal point, amply justify

its identity as a distinct species.

Male: Body length 2.77 (2.59—2.96, n=15) mm.

Head: Brown to dark brown in colour. Vertex brown with 12 (postocular 1,

outer verticals 7, inner verticals 4) setae. Corona without setae. Clypeus with

18—20(20) long setae, clypeal ratio 1.0. Maxillary palp (fig. 10a) yellow,

palpomere II dark brown, length ratio of palpomeres I—V 9:10:30:35:53; L/W

ratio 5.0. Eyes bare, hemispherical, slightly extended dorsally, extension 0.11

mm long. Antenna yellow to light brown in colour, flagellomere XII longest,

pear-shaped and densely setaceous, setae longer at the base and gradually

134 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 10a—h. Polypedilum nudiceps n.sp. 8: a, maxillary palp; b, thorax; c, wing; d, tibial ends of

fore, mid and hind legs; e, hypopygium. ©: f, antenna, g—h, genitalia.

narrowed distad, length ratio of flagellomers I—XII 15:8:10:10:10:11:11:11:

11:11:11:81; AR 0.62. Pedicel ratio 1.25. CA 0.6, CP 0.85

Thorax (fig. 10b): Deep yellow to brown in colour. Antepronotum very thin

without any V-shaped emargination. Acrostichals 14—16 in pairs, dorsocentrals

18, prealars 5, humeral 0—1. Scutellum with 12 long and 6 small setae arranged

in two oblique rows, postscutellum light brown and bare.

Wing (fig. 10c); Microtrichia dense and the veins brown in colour. Brachiolum

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 135

with 1 seta. R with 26, R, with 18, R,,; with 52 setae; R,,, meets C very close to R,

being 0.02 mm away from R,; ending of R,,; and C blunt; r—m oblique and

proximal to f—Cu; An ends a little proximal to f—Cu. Squama with 15 setae.

Haltere with yellow stem and knob and bare. Wing length 1.85 (1.66—1.96,

n=15) mm, breadth 0.55 (0.48—0.62, n=15) mm. CR 0.52, VR 1.3.

Legs (fig. 10d): Yellow in colour. Fore tibial scale abruptly pointed with 3—4

setae. Spur of mid tibia bent at the apex, 0.064 mm long, ratio of length of spur to

the apical diameter of mid tibia 15:12; spur of hind tibia also bent at the apex,

0.068 mm long, ratio of length of spur to the apical diameter of hind tibia 16:13.

Pulvilli split longitudinally; empodium as long as claws.

Proportions and ratios of leg segments

Be amt ta, ta, taz ta, tas LR TR

Fore 22 13 28 18 11 9 4 2.16 —

Mid 26 21 11 7 6 4 3 0.52 —

Hind 28 23 17 11 ©) 6 3 0.74 1.54

Abdomen: Yellow to light brown in colour. Terga uniformly setaceous and

with greyish broad bands. Segment VIII constricted basally.

Hypopygium (fig. 10e): Anal point long (0.062 mm), pointed with 7—8(8)

small setae at each basal margin. Gonocoxite short with 6 setae; gonostylus long,

little bent inward, apex narrowed down and tip in most cases abruptly pointed

with a long seta; in other cases, gonostylus straight, widened with little attenua-

tion at tip, gonostylus with 15—16 setae over it and 6—7 setae along its inner

margin. Appendage 1 bare, broad at base, narrowed down to apex and gently

bent inward, often bearing 2 long straight and 6—7 incurved setae. Laterotergite

with 6—7 long setae, transverse sternapodeme 0.086 mm long, coxapodeme

0.034 mm long, phallapodeme 0.077 mm long. HR 0.4, HV 1.83.

Female: Body length 2.40 (2.12—2.70, n=15) mm.

Similar to male with usual sex differences. Maxillary palp light brown, length

ratio of palpomeres I—V 11:12:36:43:68, L/W ratio 4.5. Antenna (fig. 10f) light

brown, length ratio of flagellomeres I—V 30:20:23:12:30. Genitalia (figs. 10g—

h): Notum 0.11 mm long. Coxosternapodeme well developed, with a notch and

two bends. Gonocoxapodeme VIII joined indistinctly at the middle. Gonapo-

physis VIII with a large dorsomesal lobe and brush-like ventrolateral lobe.

Apodeme lobe very weak. Tergum IX as usual. Tergum X with 6—7 setae on

each side. Gonocoxite IX with 6—8 setae. Postgenital plate protruded and

pointed at the apex. Cerci normal. Labia with dense microtrichia. Seminal

capsules oval, 0.055 mm by 0.051 mm long with a short neck; ducts of seminal

capsules mostly with two bends.

Polypedilum numerus n.sp.

(figs. 11a—d)

Material: Holotype d (Type no. 86, B.U. Ent.), India, West Bengal, Darjeel-

ing, 27.x.1977, P. Chaudhuri. Paratypes: 3 ©, data same as holotype; 3 d,

136 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 11a—d. Polypedilum numerus n.sp. @: a, thorax; b, wing; c, tibial ends of fore, mid and hind

legs; d, hypopygium.

Jorbangla, 3.x1.1978, P. Chaudhuri; 1 d, Kurseong, 22.x.1977, S. K. Das

Gupta.

Remarks: The species is named as Polypedilum numerus in view of the finger-

like appendage 2 of the male hypopygium. It comes close to Polypedilum

flagellatum n.sp. described in this paper and P. watsoni Freeman (1961) from

Australia in some aspects of wing and the male hypopygium. The characters like,

(i) setae in corona, (ii) presence of prescutellar setae, (iii) number and nature of

spots or bands on the wing, (iv) bands on the abdominal terga, (v) structure and

size of anal point and (vi) male hypopygium having short gonocoxite, long

gonostylus, club-shaped appendage 1 with numerous setae and finger-like long

appendage 2 with incurved setae, differentiate it as a distinct and valid species in

the genus Polypedilum. It will be worth mentioning in this connection that, since

the antenna was not available in any of the slide-mounts of this species examined,

it has not been possible in this study to present any information on it.

Male: Body length 2.73 mm.

Head: Yellow to light brown. Vertex with 12 (outer verticals 7, inner verticals

5) setae. Corona with 14 setae, clypeal ratio 1.09. Eyes bare, wider, extended

dorsally, extension being 0.11 mm long. Frontal tubercles absent. Maxillary palp

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 137

pale, palpomere III with a preapical pit (sensilla not seen), length ratio of

palpomeres I—V 6:5:15:20:28, L/W ratio 4.5. Antenna broken in all specimens.

Pedicel ratio 1.25.

Thorax (fig. 11a): Yellow to light brown in colour. Antepronotum thin,

without dorsal emargination. Acrostichals 12 in pairs, dorsocentrals 16, hume-

rals 3, prealars 6. Scutellum with 10 large and 8 small setae arranged in two rows,

postscutellum light brown.

Wing (fig. 11b): Pale brownish in colour. Brachiolum with 1 seta. R with 24, R,

with 17 and R,,; with 31 setae; R,,, meets C close to R,, distance being 0.05 mm

from R,; ending of R,,; and C acute; r—m proximal to f—Cu; An ends a little

proximal to f—Cu. Wing with light irregular grey spots or bands distributed as

follows: (1) two broad bands at the proximal and distal part of cell R,, 5, (ii) two in

cell M,,,, one being near f—Cu and another small one below the apex of M,,,,

(iii) two in cell An, one broad at the middle and another below apex of Cu,, (iv)

one linear grey streak parallel to M,,, in cell M,,,. Squama with 8 setae. Haltere

yellow and bare. Wing length 1.29, breadth 0.44 mm. CR 0.98, VR 1.23.

Legs (fig. 11c): Yellow in colour. Femur slightly dark proximally. Fore tibial

scale blunt with 1 long seta. Spur of mid tibia straight, 0.055 mm long, ratio of

length of spur to the apical diameter of mid tibia 15:13; spur of hind tibia slightly

bent apically, 0.059 mm long, ratio of length of spur to the apical diameter of hind

tibia 16:14. Pulvilli split in the middle; empodium slightly longer than pulvilli.

Proportions and ratios of leg segments

Fe Mi ta, ta, taz ta, tas LR TR

Fore 43 29 39 14 11 8 4 1.38 =

Mid 48 40 21 8 7 5 2 0.52 =

Hind 50 44 24 10 8 5 3 0.60 2.4

Abdomen: Terga I—III with brown bands and uniformly setaceous. Segment

VIII moderately constricted at the base.

Hypopygium (fig. 11d): Anal point short, 0.033 mm in length, with a few setae

at its base. Gonocoxite short, bearing 2—4(4) long setae and numerous small

setae at its inner margin; gonostylus long, little bowed, with 4 setae at its inner

apical margin, gonostylus with 8—10(10) setae over it. Appendage 1 broad,

roughly club-shaped with numerous small setae; appendage 2 elongated, finger-

like with 15—16(15) long incurved setae. Laterotergite with 2 long setae,

transverse sternapodeme 0.09 mm long, coxapodeme 0.03 mm long, phallapo-

deme 0.07 mm long. HR 0.46, HV 2.02.

Female: Unknown.

Polypedilum pseudoflagellatum n.sp.

(figs. 12a—e)

Material: Holotype d (Type no. 87, B. U. Ent.), India, West Bengal,

Durgapur, 5.xi.1976, R. K. Debnath. Paratypes: 2 d, Batanagar, 12.v.1975, S.

K. Das Gupta; 2 d, Burdwan, 2—3.vii.1977, A. Chatterjee; 1 d, Kharagpur,

138 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Figs. 12 a—e. Polypedilum pseudoflagellatum n.sp. 9: a, maxillary palp; b, thorax; c, wing; d, tibial

ends of fore, mid and hind legs; e, hypopygium.

14.x.1978, P. K. Chaudhuri; 2 &, Raniganj, 28.vii.1977, D. K. Guha.

Remarks: This species is named Polypedilum pseudoflagellatum because of its

close similarities with Polypedilum flagellatum n.sp. described in this paper.

Although this species resembles P. flagellatum in some features, it is held as a

distinct and valid species on the basis of the following:

P. flagellatum n.sp. P. pseudoflagellatum n.sp.

1. Corona with 4 setae 1. Corona without setae

2. Clypeus with 21 setae 2. Clypeus with 13 setae

3. Wing with spots 3. Wing without spots

4. Fore tibial scale elongated with blunt apex 4. Fore tibial scale triangular with pointed apex

5. HR 1.14 5. HR 0.62

Male: Body length 2.7 mm.

CHAUDHURI, GUHA & DAs GUPTA: Polypedilum from India 139

Head: Brown in colour. Vertex light brown with 9 setae (postocular 0, outer

verticals 6, inner verticals 3). Corona without setae. Clypeus with 13 setae,

clypeal ratio 1.22. Maxillary palp (fig. 12a) brown, palpomere III with an

rounded apical pit bearing bearing 1—2 sensilla, length ratio of palpomeres I—V

5:8:27:24:46, L/W ratio 4.5. Eyes bare, comma-shaped, extended dorsally,

extension pointed and being 0.37 mm long. Frontal tubercles absent. Antenna

dark brown, densely pubescent, setae longer at proximal flagellomeres and

gradually narrowed distad, length ratio of flagellomeres I—XII

DD DD DD 2A ASO Ak leg Re dicel ratio 2847 CAO 587 CPAS

Thorax (fig. 12b): Yellowish brown in colour. Antepronotum very thin

without V-shaped emargination. Acrostichals 10—12(10) in pairs, dorsocen-

trals 12 in a row, humerals 2, prealars 5. Scutellum with 10—12 setae arranged

irregularly, postscutellum dark brown to black and bare.

Wing (fig. 12c): Whitish without any clouds in most cases. Veins yellow.

Brachiolum with 1 seta. R with 19, R, with 16 and R,,; with 22—24(24) setae;

R,,, meets C very close to R,, distance being 0.05 mm from R,; ending of R,,;

and C acute; r—m distinct and proximal to f—Cu; An ending little below f—Cu.

Squama with 7 setae. Haltere bare with light brown stem and knob. Wing length

1.36 (1.34—1.38, n=5) mm, breadth 0.44 (0.42—0.45, n=5) mm. CR 0.94, VR

1,216

Legs (fig. 12d): Yellow in colour. Fore tibial scale triangular with a very short

and abrupt point bearing 3 long setae at the base. Spur of mid tibia bent, 0.03 mm

long, ratio of length of spur to the apical diameter of mid tibia 10:13; spur of hind

tibia also bent, 0.04 mm long, ratio of length of spur to the apical diameter of hind

tibia 13:13. Pulvilli small and bilobed; with a small setaceous empodium.

Proportions and ratios of leg segments

Ee Ti ta, ta, taz ta, tas LR TR

Fore 17 11 20 12 9 7 4 1.82 =

Mid 19 15 9 5 4 3 2 0.6 =

Hind 19 17 11 6 5 4 2 0.65 1.83

Abdomen: Dark brown in colour, terga with scattered setae, lateral setae

arranged in longitudinal row. Segment VIII constricted at base but less than in

Polypedilum flagellatum n.sp.

Hypopygium (fig. 12e): Anal point 0.51 mm long, apex subacute, bearing a

number of small setae at each basal margin. Gonocoxite short with 6—8(8)

setae; gonostylus moderately developed, slightly bowed and with 6 small setae at

its inner apical margin, gonostylus with 12 long setae over it. Appendage 1 well

developed, distal part bent inward and without seta; appendage 2 well devel-

oped, narrow at the base, pointed at the apex, bearing 12—14(12) long curved

setae of which outermost longer than that Polyepedilum flagellatum n.sp.

described in this paper. Laterotergite with 6—7(7) setae, transverse sternapo-

deme 0.2 mm long, lateral sternapodeme 0.09 mm long, coxapodeme 0.03 mm

long, phallapodeme 0.06 mm long. HR 0.62, HV 2.43.

Female: Unknown.

140 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Polypedilum rufomarginalis n.sp.

(figs. 13a—f)

Material: Holotype & (Type no. 88, B.U. Ent.), India, West Bengal, Burd-

wan, 4.ix.1878, A. Chatterjee. Allotype 2, data same as holotype. Paratypes: 8

d, 4 2, Burdwan, 3—4.ix.1977, 2.iii.1978, A. Chatterjee; 3 6, Berhampur,

22.ix.1976, P. K. Chaudhuri; 4 d, 1 2, Malda, 15.viii.1974, S. Chatterjee; 3 d,

1 2, Raniganj, 11—12.iii.1978, D. C. Sinharay; 2 d, Tribeni, 7.111.1975, S. Das.

Remarks: In view of the beautifully red coloured margin of the mesonotum,

this species is named Polypedilum rufomarginalis. It appears to bear some

similarities in colour pattern of the wing to Polypedilum nubifer (Skuse) from

Australia and in the male hypopygium to P. alticola Kieffer (1913) from Africa

and Chironomus nitidus Kieffer (1913) from Calcutta, India, but the colour

pattern of the mesonotum, wing, leg, tarsomere V, basal constriction of the

abdominal segment VIII and the shape of the male hypopygium altogether

support its distinctiveness from the above-mentioned species.

Male: Body length 4.25 (4.14—4.4, n=10) mm.

Head: Reddish brown in colour. Vertex red with 15 (postocular 3, outer

verticals 8, inner verticals 4) setae. Corona without setae. Clypeal with 22 long

setae, clypeal ratio 1.66. Maxillary palp dark brown, palpomere III quite long

with a preapical pit bearing 3 cilia-like sensilla; length ratio of palpomeres I—V

9:12:33:35:45, L/W ratio 3. Eyes bare, crescentic, extended dorsally, extension

0.14 mm long. Frontal tubercles absent. Antenna dark brown, densely seta-

ceous, setae longer at proximal flagellomeres and gradually becomes shorter

distad, flagellomere XII devoid of seta; length ratio of flagellomeres ITXII

LIZA 3 ARS 47 Rediceliratiol EA 0°63" CRM

Thorax (fig. 13a): Brown to dark brown with red patches, margin chocolate

red. Antepronotum dark red, thin, without V-shaped emargination. Acrosti-

chals 18—20(20) in pair from the tip of mesonotum, dorsocentrals 28, humeral 1,

prealars 6. Scutellum with 18—20 setae, postscutellum dark brown with a

median suture and bare.

Wing (fig. 13b): Pale yellowish in colour and the veins conspicuous. Brachio-

lum with 1 seta. R with 25, R, with 21 and R,,; with 24 setae; R,, meets C very

close to R,, distance being 0.09 mm from R,; ending of R,,; and C subacute; r—

m distinct and proximal to f—Cu; An ending below f—Cu. Wing with nine grey

irregular spots distributed as follows: (i) cell R,,; with three spots placed at the

base, middle and near the apex, (ii) cell M,,, with two spots placed near the base

of M,,, and behind the apex, (iii) cell M,,, with one spot near f—Cu and

continuous along Cu, as a streak, (iv) cell An with one broad at the middle, one

small behind f—Cu and another at the end of An. Sensory organ l on r—m.

Squama with 15 setae. Haltere light brown at stem and dark brown or black at the

knob. Wing length 2.07 (1.99—2.14, n=10) mm, breadth 0.55 (0.52—0.59,

n=10) mm. CR 0.96, VR 1.08.

Legs (fig. 13c): Yellow to brown in colour. Femur slightly dark, fore tibia pale

near the apex. Fore tibial scale blunt with 4—5 long setae. Spur of mid tibia bent

being 0.04 mm long, ratio of length of spur to the apical diameter of mid tibia

CHAUDHURI. GUHA & Das GUPTA: Polypedilum from India 141

Figs. 13a—f. Polypedilum rufomarginalis n.sp. 9: a, thorax; b, wing; c, tibial ends of fore, mid and

hind legs; d, hypopygium, ®; e, antenna, f, genitalia.

12:19; spur of hind tibia also bent, 0.05 mm long, ratio of length of spur to the

apical diameter of hind tibia 15:20. Tarsomere IV brown and tarsomere V dark

in all legs. Pulvilli narrow and split; a short setaceous empodium, shorter than

claws.

Proportions and ratios of leg segments

Fe Ti ta; ta2 ta; ta, tas LR TR

Fore 29 18 26 14 12 I 4 1.44 =

Mid 29 23 14 8 6 4 2 0.60 a=

Hind 28 27 19 10 ©) 5 3 0.70 1.9)

142 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

Abdomen: Dark brown in colour. Tergum 1 with a broad median depression;

setae on terga long and arranged in irregular rows. Segment VIII constricted like

a triangle having a very narrow base.

Hypopygium (fig. 13d): Anal point 0.07 mm long, tip blunt, base broader with

8—10(10) small setae. Gonocoxite well developed, long, with 18—20(18) long

setae; gonostylus short, stout, apex a little narrowed, bearing 4—6(4) small setae

at its inner apical margin. Appendage 1 hockey-stick-shaped; appendage 2

narrow at base and gradually becoming wider towards apex, with 18—20 long

incurved setae. Laterotergite with 3—4(3) setae, transverse sternapodeme 0.28

mm long, lateral sternapodeme 0.17 mm long, coxapodeme 0.08 mm long,

phallapodeme 0.13 mm long. HR 1.17, HV 3.80.

Female: Body length 3.6 mm.

Similar to male with usual sex differences. Maxillary palp pale to dark brown.

Palpomere 1 pale, others dark brown, length ratio of palpomeres I—V

10:9:28:25:45, L/W ratio 3.1. Antenna (fig. 13e) yellow to dark brown, flagello-

meres I—V dark brown and narrowed with 2 subapical setae, length ratio of

flagellomeres I—V 7:6:5:5:9, AR 0.39. Colour and spots of wings similar to that

of male. Wing length 1.96 (1.92—1.96, n=5) mm, breadth 0.66 (0.65—0.68,

n=5) mm. Genitalia (fig. 13f): Notum 0.16 mm long. Gonocoxapodeme VIII

caudally rounded and widely separated at the middle. Coxosternapodeme with a

small and broad bend. Gonapophysis VIII with a tongue-like dorsomesal lobe

and a small blunt brush-like ventrolateral lobe. Apodeme lobe moderate.

Gonocoxite IX with 3—4 long setae. Tergum IX with 8—10(8) setae on each

side. Tergum X normal. Postgenital plate as small protuberance. Cerci elong-

ated. Labia narrow, with microtrichia. Seminal capsules (fig. 13f) equal, oval,

with small neck, measuring 0.05 mm by 0.03 mm; ducts of seminal capsules

straight or with one bent and an ampulla before opening in common.

Polypedilum tripunctum n.sp.

(figs. 14a—e)

Material: Holotype d (Type no. 89, B.U. Ent.), India, West Bengal, Darjeel-

ing, 25.x.1970, S. K. Das Gupta. Paratypes: 4 d, Darjeeling, 2—3.1x.1973, P.

K. Chaudhuri; 1 &, Tindharia, 17.ix.1974, T. Sharma.

Remarks: The present species is named Polypedilum tripunctum in view of its

triradiate structure of anal point of the male hypopygium. It resembles Polypedi-

lum aegyptum Kieffer (1925), P. pruina Freeman (1958) and P. tridens Freeman

(1958) in certain features of the male hypopygium and P. tropicum Kieffer

(1913) in respect of wing and male hypopygium, but its separate entity as a valid

species can be justified by means of the following features: (i) spots on the wing,

(ii) spur of fore tibia, (iii) dark bands on legs, (iv) structure of the appendages 1

and 2 with its chaetotaxy and (v) shape of anal point.

Male: Body length 2.75 (2.53—3.0, n=5) mm.

Head: Brown to dark brown. Vertex brown with 12 (postocular 1, outer

verticals 8, inner verticals 3) setae. Corona without setae. Clypeus with 18—20

CHAUDHURI, GUHA & DAS GUPTA: Polypedilum from India 143

Figs. 14a—e. Polypedilum tripunctum n.sp. 6: a, maxillary palp; b, thorax; c, wing; d, femora and

tibia of fore, mid and hind legs with their tibial ends; e, hypopygium.

(18) long setae, clypeal ratio 0.83. Maxillary palp (fig. 14a) light brown,

palpomere V brown, length ratio of palpomeres I—V 7:7:20:23:42, L/W ratio

3.3. Eyes bare, hemispherical, extended dorsally, extension being 0.15 mm

long. Frontal tubercles absent. Antenna brown to pale brown, flagellomeres

XIII dark distally, length ratio of flagellomeres I—XII 11:13:10:12:13:13:13:

13:13:13:13:13:75, AR 0.5. Pedicel ratio 1. CA 0.54, CP 1.06.

Thorax (fig. 14b): Brown in colour. Antepronotum very thin and without any

V-shaped emargination. Acrostichals 12—14 in pairs, dorsocentrals 16—18

(18), humerals 2, prealars 5. Scutellum with 18—20 setae, postscutellum dark

brown and bare.

Wing (fig. 14c): Cloudy especially along the margin of wing. Veins dark

144 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

brown, anterior veins more prominent than the posterior ones. Brachiolum with

1 seta. R with 21, R, with 15 and R,,; with 35—38 setae; R,,, meets C slightly

away from R,, distance being 0.1 mm from R,; ending of R,,; and C blunt; r—m

oblique and proximal to f—Cu; An ending below f—Cu. Wing with four

irregular grey spots distributed as follows: (i) two in cell R,,; near r—m and at the

middle of the cell, (ii) one in cell M;,, covering the base nearer to f—Cu and (iii)

one at the middle of cell An. In addition to the above a grey streak runs below the

vein M and the space between the spots in cell R,,; along the apex. Wing length

1.64 (1.5—1.78, n=8) mm, breadth 0.6 (0.56—0.62, n=8) mm.

Legs (fig. 14d): Brown in colour. Femur with broad dark brown band at the

proximal two-third and a small apical dark brown band, tibia with an apical dark

brown band on all legs, rest of the legs pale to brown. Fore tibial scale narrowed

and slightly pointed at the apex. Spur of mid tibia slightly bent and sharply

pointed at the apex, 0.05 mm long, ratio of length of spur to the apical diameter of

mid tibia 12:10; spur of hind tibia also bent and pointed, 0.055 mm long, ratio of

length of spur to the apical diameter of hind tibia 13:12. Pulvilli split longitudinal-

ly.

Proportions and ratios of leg segments

Fe Ti ta, ta, ta; tay tas LR TR

Fore 31 22 41 28 15 7 5 1.87 —

Mid 38 31 16 10 7 5 3 0.51 —

Hind 41 33 25 13 11 7 4 0.76 LED

Abdomen: Brown to dark brown. Tergum I densely setaceous, rest of the

segments with scattered setae.

Hypopygium (fig. 14e): Anal point long, apex pointed, base with two hook-

like structures. Gonocoxite short and with 10—12 long setae; gonostylus very

long, bowed and with 15—18 setae. Appendage I well developed, broad apex

with numerous setae; appendage 2 long, apex slightly broad and with 8—10 long

curved setae. Lateral sternapodeme 0.094 mm long, coxapodeme 0.051 mm

long, phallapodeme 0.07 mm long. HR 0.66, HV 1.76.

Female: Unknown.

CHECK-LIST OF THE SPECIES OF THE GENUS PL YPEDILUM KIEFFER FROM INDIA

albipalpus n.sp., 6,9, Raniganj (West Bengal)

angustiforceps Kieffer, 6, Katihar (Bihar)

annulatipes (Kieffer), d, 9, Puri (Orissa)

ascium n.sp., &, Bolpur (West Bengal)

brumale Kieffer, ®, Giridih (Bihar)

chaudhurii n.sp., 4, 9, Burdwan (West Bengal)

fasciatipennis (Kieffer), 9, Calcutta (West Bengal)

flagellatum n.sp., 3, 2, Raniganj (West Bengal)

griseoguttatum Kieffer, 4, 2, Burdwan and Calcutta (West Bengal)

insolitum n.sp., 4 , Digha (West Bengal)

CHAUDHURI, GUHA & DAS GUPTA: Polypedilum from India 145

lineatum n.sp., 6, Raniganj (West Bengal)

lucidum n.sp., 4, Burdwan (West Bengal)

milnei Kieffer, 6, Berhampur (West Bengal)

nudiceps n.sp., 3, 9, Darjeeling (West Bengal)

numerus n.sp., 6, Darjeeling (West Bengal)

pseudoflagellatum n.sp., 6, Durgapur (West Bengal)

purimanus Kieffer, & , Calcutta (West Bengal)

rufomarginalis n.sp., 8, 2, Burdwan (West Bengal)

sealaenulus (Edwards), 3, 9, Barkuda Is. (Orissa)

tripartitum (Kieffer) 6, 2 (Orissa and West Bengal)

tripunctum n.sp., d, Darjeeling (West Bengal).

KEY TO THE SPECIES OF GENUS POL YPEDILUM KIEFFER IN WEST BENGAL

IWinsiwithidarkmarkinestand’elouds Zr EE 2

med Winesunmarked nee ee ar 14

Plhoraxowith bands Ormarkines, IE 3

=e NOLAXGwithOutL bands On markings neee GEER 8

SEeAppendagesishortandibroadwithisetaeger nn re 4

— Appendage 1 long and narrow without setae ........................................ 6

4. ARlessthan 1. Abdomen withmarkingsorbands .................. lineatum n.sp.

— AR more than 1. Abdomen without markings or bands ........................... 5

5. Fore tibial scale pointed. Anal point tongue-like, constricted at the middle … …

eN EEE griseoguttatum Kief.

Fore tibial scale blunt. Anal point narrow with pointed apex … …

SVN UNE, PRIS OR RE ENGINE AE DE annulatipes Kief.

6. Abdominal terga I—VII with alternate dark brown and pale bands................

RIT pote LU ee Fo OS Jane Ae denn insolitum n.sp.

— »Abdomen:withoutsuchbands urn EE atd deed CT 7

7. Thorax yellow in colour. Coronal setae 4. Appendage 2 with a long setae........

RO ATI RR SEA soa ended OSE at flagellatum n.sp.

— Thorax red in colour. Coronal seta absent. Appendage 2 with normal setae.....

Ree a ARR MR I EA Ra rufomarginalis n.sp.

STATA PONI AH iste Eee ME ARE ALOE EEE 9

— Analpointisimple Me Bee RL Re 11

9. Fore femur with dark brown apical band. Appendage 1 flattened ............ 10

— Fore femur without dark brown apical band. Appendage 1 narrow................

Be ee EG nr LG ah Let Ab chaudhurii n.sp.

10. Anal point lanceolate, lateral branches of anal point short and blunt..............

E ER RI eR A deb O Va o A MZ DI lucidum n.sp.

— Analpointlongandpointed -....:................................... tripunctum n.sp.

11. Legswithbandsonfemurandtibia .................................. brumale Kieffer

— Legs without bands on femur and tibia ............................................. 12

12. Appendage 1 club-shapedwithnumeroussetae .................... numerus n.sp.

= Appendagenisimplekwvatlmaktemise aCMEN PER EEE EEE 15

13. Analpointhood-likeandinturned .................................. albipalpus n.sp.

—_ wAnalpeintsimple TITTI ani milnei Kieffer

146 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 4, 1981

14. Thorax with bands or markings Mers 15

— ‘Thorax without bandstormarkings nn... RR 16

15. Mid and hind tibia with brown apical band. Abdominal segments II, III, V

and Vilwithdarkibrownibands) Zee mer fasciatipennis n.sp.

— Mid and hind tibia without bands. Abdominal segments without such bands ..

ee LE SSSR N ag pe ascium n.sp.

16. Abdominal segments with bands. Appendage 2 finger-like and uniform in

diameterbeanngineurved SCIARE RR nudiceps n.sp.

— Abdominal segments without band. Appendage 2 with narrow base and

wide apex bearing incurved setae, outermost seta unusually long ..................

ha ER LOR RIA NI ARR AIA RARE, RO pseudoflagellatum n.sp.

REFERENCES

(When marked with an asterisk, the original publication was not seen.)

Edwards, F. W., 1929. British non-biting midges (Diptera, Chironomidae).— Trans. R. ent. Soc.

Lond. 77: 279—430.

Freeman, P.,1958. A study of the Chironomidae (Diptera) of Africa south of Sahara. IV. — Bull. Br.

Mus. nat. Hist. Ent. 4: 263—363.

, 1961. The Chironomidae (Diptera) of Australia. — Aust. J. Zool. 9: 611—737.

Frommer, S., 1967. Review of the anatomy of adult Chironomidae. — Calif. Mosquito Control Ass.

Tech. Ser. Bull. 1: 1—39.

Goetghebuer, M., 1934. Ceratopogonidae et Chironomidae du Congo Belge. — Revue Zool. Bot.

afr. 25: 191—205.

—, 1936. Chironomidae du Congo Belge. — Ibid. 28: 453—492.

——, 1937. Ceratopogonidae et Chironomidae nouveaux ou peu connus d'Europe. — Bull.

Annls Soc. r. ent. Belg. 77: 273—280.

Guha, D. K., & P. K. Chaudhuri, 1979. A new species of genus Cryptochironomus Kieffer

(Chironomidae, Diptera) from West Bengal. — Bull. Zool. Surv. Ind. 2 (1): 95—98.

Johannsen, O. A., 1932. Chironomidae of the Malayan subregion of the Dutch East Indies. — Arch.

Hydrobiol. Suppl. 11: 503—552.

Kieffer, J. J., 1910. Etude sur les Chironomides des Indes Orientales, avec description de quelques

nouvelles espéces d’Egypte. — Mem. Indian Mus. 2: 181—242.

, 1911a. Contributions to the fauna of Yunnan. IV. Les Chironomides. — Rec. Indian Mus.

6: 27—30.

, 1911b. Descriptions de nouveaux Chironomides de l’Indian Museum de Calcutta — Ibid. 6:

113-117.

—, 1911c. Les Chironomides (Tendipedidae) de l'Himalaya et d’Assam. — Ibid. 6: 319—349.

———, 1911d. Diptera. Chironomidae der Seychellen-Inseln, aus der Sammlung von Mr. H. Scott.

— Trans. Linn. Soc. London, Zool., 14: 331—366.

—, 1913. Nouvelle étude sur les Chironomides de I’Indian Museum de Calcutta — Ibid. 9:

119—197.

, 1915. Fauna faeroensis: 2. Tendipedidae (Chironomidae). — Zool. Jb. 39: 103—120.

, 1921. Chironomides nouveaux ou peu connus de la région palaearctique. — Bull. Soc. Hist.

nat. Metz. 29: 51—109.

, 1922. Chironomides de l’Afrique Equatoriale. Ile partie. — Annls Soc. ent. Fr., 91: 1—72.

1925. Chironomides d’Egypte (Dipt.). — Bull. Soc. ent. Egypte 8: 244—313.

Kruseman, G., 1949. Note on Tendipedidae of the Suez Canal. — Bijdr. Dierk. 28: 249 —254.

* Lundström, C., 1916. Beitrage zur Kenntnis der Dipteren Finlands. X. Suppl. 4: Bibionidae,

Chironomidae, Tipulidae. — Acta Soc. Fauna Flora fenn. 44: 5—25.

Saether, O. A., 1971. Notes on general morphology and terminology of the Chironomidae

(Diptera). — Can. Ent. 103: 1237—1260.

CHAUDHURI, GUHA & Das GUPTA: Polypedilum from India 147

——, 1974. Morphology and terminology of female genitalia in Chironomidae (Diptera). — Ent.

Tidskr. Suppl. 95: 216—224.

—, 1977. Female genitalia in Chironomidae and other Nematocera; morphology, phylogenies,

keys. — Bull. Fish. Res. Bd. Can. 197: 1—210.

* Skuse, F. A. A., 1889. Diptera of Australia. Part VI. The Chironomidae. — Proc. Lin. Soc.

N.S.W. 2: 215—311.

Sublette, J. E., & M. S. Sublette, 1973. Family Chironomidae. — In: Delfinado, M., & E. D. Hardy

(Eds.): Catalogue of the Diptera of the Oriental region, Suborder Nematocera, 1: 289—

422. — University Press of Hawaii.

Townes, H. K., 1945. The Nearctic species of Tendipedini (Diptera: Tendipedidae — Chironomi-

dae). — Amer. Midl. Nat. 34: 1—206.

Tr se ei

TI) 799°

DEEL 124 AFLEVERING 5 1981

MUS, COMP. ZOOL

LIBRARY 7

AUG S «ac

AUG 5 19 81

THDSCURIFT ei

UNIVERSITY

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

Z. Boucex, A. WarsHam and J. T. Wieses. — The fig wasp fauna of the recep-

tacles of Ficus thonningii (Hymenoptera, Chalcidoidea), p. 149—233, figs.

1—158, four colour-plates.

Tijdschrift voor Entomologie, deel 124, afl. 5 Gepubliceerd 20-VII-1981

THE FIG WASP FAUNA OF THE RECEPTACLES OF FICUS

THONNINGII (HYMENOPTERA, CHALCIDOIDEA)

Z. BOUCEK!), A. WATSHAM?) and J. T. WIEBES?)

With 158 text-figures and four colour-plates

ABSTRACT

Records and descriptions of the chalcids reared from the receptacles of Ficus thonningii BI. (burkei

(Miq.) Miq., petersii Warb.) mainly in Zimbabwe-Rhodesia, among which the following new

species: Camarothorax brevimucro Boutek, spec. nov.; equicollis Boutek, spec. nov.; longimu-

cro Boutek, spec. nov. — Crossogaster odorans Wiebes, spec. nov. — Eurytoma ficusgallae

Boucek, spec. nov. — Ficomila Boutek, gen. nov., type-species Ficomila curtivena Boutek,

spec. nov.; gambiensis (Risbec) Boutek, comb. nov. from Decatoma. — Otitesella tsamvi Wiebes,

spec. nov. — Ormyrus flavipes Boutek, spec. nov.; subconicus Boutek, spec. nov.; watshami

Boucek, spec. nov. — Philotrypesis parca Wiebes, spec. nov. — Syceurytoma Boutek, gen. nov,

type-species Syceurytoma ficus Boutek, spec. nov. — Sycophila flaviclava Boutek, spec. nov.;

kestraneura (Masi) Boutek, comb. nov. from Decatoma; modesta Boutek, spec. nov.; naso

Boucek, spec. nov.; punctum Boutek, spec. nov.; sessilis Boutek, spec. nov. Sycoryctes hirtus

Wiebes, spec. nov.; remus Wiebes, spec. nov. — Sycoscapter cornutus Wiebes, spec. nov. —

Sycotetra Boutek, gen. nov., type-species Sycotetra serricornis Boutek, spec. nov. — Watshamiel-

la Wiebes, gen. nov., type-species Watshamiella alata Wiebes, spec. nov. — The following known

species are recorded from the receptacles of Ficus thonningii: Alfonsiella brongersmai Wiebes;

longiscapa Joseph. — Elisabethiella stuckenbergi (Grandi). — Phagoblastus barbarus Grandi. —

The following new species and combinations originate from other localities: Camarothorax africanus

(Wiebes) Boutek, comb. nov. from Sycobia (ex Ficus cf. eriobotryoides K. & B., Sierra Leone);

imerinensis (Risbec) Boutek, comb. nov. from Callimomus (Malagasy); orientalis (Wiebes)

Boutek, comb. nov. from Sycobia (ex Ficus reflexa Thunb., Aldabra). — Ormyrus chevalieri

(Risbec) Bouéek, comb. nov. from Monobaeus; decaryi (Risbec) Boutek, comb. nov. from

Tribaeus. — Sycophila, new combinations, all from Decatoma: aethiopica (Silvestri) Boucek;

cassinopsisi (Risbec) Boutek); nigrofasciata (Risbec) Boutek; ruandensis (Risbec) Boutek;

rubra (Risbec) Boutek; xiphigaster (Risbec) Boutek. — Watshamiella infida Wiebes, spec. nov.

(ex. Ficus religiosa L., Sri Lanka); stilifera (Wiebes) Wiebes, comb. nov. from Sycoscapteridea (ex

Ficus stupenda Miq., North Borneo).

A key is given to the chalcids associated with the receptacles of F. thonningii, irrespectively

whether coming from evidently galled ones or not. The species having a closer association with the

fig, especially the pollinators, are also mentioned in the introductory description of the fig.cycle and

the chalcid wasp fauna of the best studied species of Ficus are reviewed.

1) Commonwealth Institute of Entomology, c/o British Museum (Natural History), London,

England.

2) St. Ignatius College, Salisbury, Zimbabwe-Rhodesia.

3) c/o Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden, The Netherlands (requests for

reprints to be sent to this address).

149

150 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

INTRODUCTION

For some years, one of us has been observing fig insects from what is called

Ficus burkei on Chishawasha Mission, near Salisbury, Zimbabwe-Rhodesia. In

the present paper, the general chapter on the fig cycle and the colour-plates are

by his hand (A. W.). The insects collected were sent for identification to Europe,

where they were divided into two lots, viz., the Agaonidae and the groups known

as Sycoecini, Philotrypesini, Sycoryctini and Otitesellini (treated by J. T. W.),

and the Epichrysomallinae, Eurytomidae and Ormyridae (treated by Z. B.). In

a short chapter on the classification of chalcidoid fig wasps, some alterations are

suggested as to their placement in one of the families Torymidae and Pteromali-

dae (Z. B.)

Some additional material was collected by Dr. M. G. Bingham (Lusaka) in

Zambia, or taken from the BMNH collection. Also some colleagues from South

Africa, viz., Dr. C. Eardley (Pretoria), Dr. S. Neser (Stellenbosch) and Dr. M.

L. Penrith (Windhoek) kindly sent fig material when requested. In several

instances, our treatment led to a revision of the genus or group concerned, and

typical material of extralimital species or forms from other host Ficus, were

requested on loan. Thanks are due to the curators of the following collections,

for complying with our requests (the abbreviations of the institutes are those

used throughout the text): British Museum (Natural History), London

(BMNH); Cornell University, Entomology Department, Ithaca, N.Y.; Labo-

ratorio di Entomologia, Bologna (LEB); Muséum National d'Histoire Natu-

relle, Paris (MNHN); Rijksmuseum van Natuurlijke Historie, Leiden

(RMNH).

Acknowledgements are also due to Dr. C. C. Berg (University of Utrecht,

The Netherlands), for botanical help and reading portions of the manuscript.

Dr. R. B. Drummond of the Salisbury Herbarium arranged for the preserving of

the Ficus-specimen and was helpful with botanical advise.

Publication of the four colour-plates was made possible by a grant from the

Uyttenboogaart-Eliasen Stichting at Amsterdam.

THE FIG CYCLE

Ficus thonningii Blume is an extremely variable species in which can be

included Ficus burkei (Miquel) Miquel and F. petersii Warburg, hitherto sepa-

rated on minor differences in leaves and figs. Although their conspecificity is

recognized, the names F. burkei and petersii are still used here, as samples have

been sent under those names. In Ficus thonningii s.l. the trees are up to 15 m high.

The leaves are obovate to elliptic or even lanceolate, glabrous or beneath

pubescent, the apex is short-acuminate to obtuse, the base acute to rounded:

The figs are borne in the leave axils or just below the leaves, they are sessile or

pedunculate, sparsely puberulous to densely pubescent. Material of Ficus

thonningü is often confused with Ficus natalensis Hochstetter and F. craterosto-

ma Mildbread & Burret. The latter two species are sometimes treated as distinct,

by others considered as inseparable (e.g., Palmer, 1977: 85, 89).

When sectioned a fig can be seen to have a space in the centre, surrounded by

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 151

inhibited

ripening

until insects

leave

‘parasites!

oviposit from

emergence

of males 6

mating

emergence

È of females

oviposition

pollination anthers develop

'lodgers'

arrival of enter and pollinator

nematodes oviposit collects pollen

Figs. 1—6. Ficus thonningii BI. and Elisabethiella stuckenbergi (Grandi). 1, section through a young

fig (in the receptive stage B); 2, female florets, with styles of different lengths; 3, male floret (ripe in

stage D); 4, general appearance of a twig with leaves and figs; 5, female fig wasp, showing pollen

pockets (p); 6, the fig cycle, running from a young receptable A, over B (in which the pollinators and

the lodgers enter), C (in which most parasites oviposit) and D (from which the wasps of the new

generation emerge), to the ripe fig E, the fruits of which are dispersed, a.o. by birds. (Scale to figs and

insects of fig. 6).

florets, the outer layer being an enlarged receptacle (fig. 1). Bracts form an

ostiole by which insects may enter; entrance is limited to those insects that are

adapted to crawling through narrow places. Such adaptations of the female of

Elisabethiella stuckenbergi (fig. 5) are seen in the legs, the wings and the head and

antennae. The legs, hind and front, are powerfully developed, whereas the

middle pair are not. The tibiae are short and strong, with numerous spines. The

head has a sunken central membraneous region, which allows it to be deformed

when entering, and protects part of the antennae. The mandibular appendage

may assist her in forcing her way in. The trailing segments of the antennal club

and funicle break off and the wings detach when she enters the fig.

At the early stage (B, fig. 6) it is comparatively easy for the female to crawl in

through the ostiole. Inside there is a space for her, surrounded by an even layer of

152 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

stigmas (synstigma, Galil & Eisikowitch, 1968: 261 '). In the section of the fig it

can be seen that the styles are of different lengths (fig. 2). The insect has a short

ovipositor, which limits her ability to lay eggs in the florets with the shorter styles:

in these the new generation of wasps will develop. The florets with the longer

styles are more likely to become seeds. The receptacle is monoecious, the male

florets are scattered amongst the female and there are relatively few (about

10%).

Ovipositing and pollinating will commence when the wasp has entered and

proceed for some hours. If there are other insects also there, she will not be

disturbed, but her ovipositing will be restricted. Other species may enter: of

these Crossogaster odorans is the most common. Less common is Phagoblastus

barbarus. Recently, it was published that a species of Phagoblastus pollinates the

flowers of Ficus vogelii (Miq.) Miq. in Ghana, although it is not as efficient as is

the Agaonid (Allotriozoon heterandromorphum Grandi). According to Newton

& Lomo (1979: 29) this relative ineffectiveness of the Phagoblastus probably

results from two factors. One is that the only pollen carried into the fig by the

female wasp is that adhering to the outside of her body. The other factor is that

transfer of pollen from the insect’s body to the synstigma is probably a matter of

chance, brought about as the insect crawls around. In Ficus thonningii, pollina-

tion is a deliberate act by the Elisabethiella and she carries the pollen in special

pockets (fig. 5). After pollination, she cannot leave the fig, due to the arrange-

ment of the ostiole, however, she has only a short time to live. Nematodes inside

her will feed on her remains, growing rapidly to about 0.7 mm. They emerge

from the gastral end as a number of large nematodes, twisted together, and inert.

The species was recently described as Parasitodiplogaster sycophilon Poinar

(1979). A general account on figs and nematodes was published by Martin et al.

(1973).

Next to Elisabethiella stuckenbergi, there are two other Agaonids entering the

receptacle in stage B, viz., Alfonsiella brongersmai and longiscapa. They may be

lodgers in the fig, just as are the Crossogaster and Phagoblastus, and pollinate

only occasionally. They have functional pollen pockets and pollen can some-

times be seen in them, but nothing is known of their behaviour. It is known for the

Elisabethiella: while continuing oviposition, she periodically will stretch for-

ward both head and thorax, and at the same time fold the legs back so that the feet

touch the edges of the pollen pockets. It can be seen from the side that the thorax,

which is normally arched, flattens out, drawing the coxae of the front legs

forward and exposing the pockets. It is presumed that the pollen removed from

the pockets with the feet is then applied to the stigmal surfaces as she rubs the feet

about. The action may be done once, or repeated a number of times before

pausing. Thus, five species are found to oviposit inside the fig, viz.,

Elisabethiella stuckenbergi (Grandi) Crossogaster odorans Wiebes

Alfonsiella brongersmai Wiebes Phagoblastus barbarus Grandi

Alfonsiella longiscapa Joseph

1) General reference is here made to a series of papers by Galil and collaborators on the pollination

of figs (review in Galil, 1977). Some of the terms used in the present paper were taken from Galil et

al., as are the indications of the developmental stages of the receptacle.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 153

The fig develops inwards. — After pollination, the florets and galls enlarge,

completely filling the space, and crushing the remains of the insects. In this state,

the nematods may be overlooked, but if the fig is cut up and placed in water, they

can be found.

A number of species of chalcids oviposit through the skin of the fig. They have

various methods of forcing the ovipositor through the hard receptacle, as de-

scribed by several authors (review in Wiebes, 1977: 224—227, fig. 8). Some are

evidently adapted to this task by the structure of their ovipositing organs, which

they can raise high over the substratum, or they can guide the ovipositor by the

long valves. Other species are quite normal and have their ovipositor concealed

in the gaster. Some are known to behave as the cuckoo and have been termed

“cleptoparasites’’. They lay their eggs later than the wasps that enter the fig, and

they tend to emerge before, so their rate of development is faster. They cannot

themselves cause the florets to develop, so have to rely for their larval food on

one already started. Some take only plant tissue prepared by the Agaonid, e.g.

this is known for Philotrypesis caricae (L.) from the Common Fig. Other species

are supposed, by their being noticeably smaller than the other wasps, to be para-

sitoids of any of the other larvae, killing it and eating its body contents. The fol-

lowing is a list of those to be found ovipositing from the outside:

Philotrypesis parca Wiebes Ficomila curtivena Boutek

Watshamiella alata Wiebes Ficomila gambiensis (Risbec)

Sycoscapter cornutus Wiebes Sycophila kestraneura (Masi)

Sycoryctes remus Wiebes Sycophila punctum Boutek

Sycoryctes hirtus Wiebes Sycophila sessilis Boucek

Otitesella tsamvi Wiebes Sycophila modesta Boutek

Sycotetra serricornis Boutek

Camarothorax equicollis Boutek

Camarothorax longimucro Boutek

Camarothorax brevimucro Boutek

Eurytoma ficusgallae Boucek

Sycophila naso Boutek

Sycophila flaviclava Boutek

Ormyrus watshami Boutek

Ormyrus flavipes Boutek

Ormyrus subconicus Boutek

Syceurytoma ficus Boutek

It should be stated that we found it difficult to draw a line between the obligato-

ry fig wasps and those which apparently have a looser or even very loose associa-

tion with the figs. We have included in the study the chalcids which frequently oc-

cur in the galled receptacles, whether these finally develop into normal or sub-

normal syconia. However, there are a number of other insects that use figs as a

suitable niche, but are not closely associated with them. Even if they are some-

times present in considerable numbers, as e.g., parasites of dipterous eggs de-

posited into the tissues of the fig, or of other insects using the fig in a similar way,

they are regarded as being out of the scope of this paper. The chalcids associated

with such facultative inhabitants of the figs belong mostly to the genera Eupel-

mus Dalman, Anastatus Motschulsky, Pediobius Walker, Tetrastichus Haliday,

etc.

The fig grows outwards. — As the fig grows, the central space will be formed

again. This allows more freedom of movement. Nematode eggs can be found

between the bracts of the florets. Inside the eggs, the juvenile nematode can be

154 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

seen sliding about. As the space is formed in the centre of the fig, the male insects

start to emerge from their galls. This can be encouraged by opening the nearly

developed fig.

The male Elisabethiella makes an opening in his gall and sits on the edge of the

gall. He looks with the aid of his antennae for galls containing females of his

species, which are usually in one region. There is probably something in them

that attracts him, as he will continue to open a female gall even when she has been

removed. He bites a hole in the female gall and inserts the tubular segments of his

gaster and mates. This process takes about half a minute from finding to

insertion. He continues looking for females. :

Other wingless males emerge at the same time. These eae their own galls and

go in search of the females: in the species which oviposit from the outside,

perhaps they are more scattered than the Elisabethiella, and so need to be looked

for. The males bite an opening into the female gall and slip in to mate with her

inside. This behaviour may be different for the different species, but this can be

more easily studied now they have been distinguished and described.

As the males are very similar to each other, and differ completely from the

females, in most cases it was necessary to collect mating pairs. As each pair

mated, they were collected. The winged females can be collected on forceps and

presented to the males. The other species will reject them, while the correct one

will stroke her face and then mount and mate. Later in this stage (fig. 6, D), the

Elisabethiella males make several openings in the receptacle and the females

start to leave.

When the Elisabethiella female crosses an anther, she is stimulated to collect

pollen. She rubs her fore legs as though wiping the feet on a mat, and raises them

to place the pollen into the pockets. After having crawled out of the receptacle,

she flies off. The complete cycle may take about 50 to 60 days, though it is

variable. Ficus capensis Thunb. takes about this time, and it is quite easy to find

fruit at different stages, most of the year. The Ficus burkei seems to have a similar

lot of fruit available, but it is not always so easy to find. At certain times there is

much ripe fruit and many thousands of insects are emerging, but it is difficult to

find any trees with young fruit that would receive these insects. This may be due

to the fact that trees are cut for firewood. A careful search will reveal fruit at

different stages of development.

Nematodes can be seen in the newly emerged insects. At first they are crawling

over the insects in the moist conditions of the fig. More than twenty can be seen

easily. If the emerged insects are kept for a day in a plastic bag, fewer nematodes

are seen on the outside, and some are found inside the thorax (sometimes in the

pollen pockets) and gaster. They can be seen moving about, displacing the

contents of the gaster. At this time they are about 0.4 mm long.

The final ripening of the fig fruit. — According to Galil et al. (1973) opening of

the figs allows an increase of ethylene, which encourages the final ripening of the

fruit. So the insects are protected while they remain in the fig and no emergence

holes have yet been made. Once the insects emerge, the fig begins to ripen and

birds such as Glossy Starlings and other fruit-eating birds, also the Fruit Bats,

feed on the ripe figs and aid in the dispersal of the seeds.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 155

Other insects are to be found in the fig, but they are not concerned with the

cycle of the fig. These include Lepidoptera, weevils (Coleoptera) and several

Hemiptera. Diptera that are interested in ripening fruit may also be found (see

an interesting paper by Lachaise, 1977).

THE FIG WASP FAUNA FROM FICUS THONNINGII COMPARED WITH THAT FROM OTHER FIGS

The main purpose of the present paper is to give a survey of all chalcids reared

from one species of fig, in order to make their symbioses open to further

research. Compared to what we know of the entomofauna associated with other

species of Ficus, that of Ficus thonningti is now the best known, while that of the

Indian Ficus benghalensis is the next best. In general, however, our knowledge

of the fig-fauna is scanty: hopefully, our survey will persuade other entomolog-

ists to study the wasps of figs within their reach.

In table 1 a survey is presented of selected figs and their chalcid fauna. Most

sections of Old World Ficus (see Corner, 1965) are represented, but five had to

be excluded for lack of relevant data (viz., Stilpnophyllum Endl., Rhizocladus

Endl., Kalosyce (Miq.) Corner, Sinosycidium Corner and Adenosperma Cor-

ner). In the accompanying list, the references are given that can not be found in

the host catalogue (Wiebes, 1966 b). The data lead to the following conclusions.

Generally, the Agaonid genera, or in some instances species-groups, are specific

to their host-section(s): members of the subfamily Agaoninae are restricted to

two sections of Ficus, the African representatives of which also harbour Sycoeci-

nae (but see Wiebes, 1974 b for the Indo-Malayan relatives). The Sycophagini

(one species of Parakoebelea excepted, see Joseph, 1957: 100) and the Apocryp-

tini are restricted to figs pollinated by Ceratosolen. The Otitesellinae (with very

few exceptions, not listed; see Wiebes, 1974 d) are restricted to the sections of

subgenus Urostigma and to section Oreosycea.

Special attention is drawn to cases where more than one Agaonid are present

(nos. 18, 19, 35, 36, 78), where several species of parasitic wasps are present (nos.

8, 17, 41, 45, 48, 49, 50), or where one species was recorded from different

species of Ficus (nos. 71, 77; 47, 84; 48-2, 85; 49-5, 86). The list is not in all

instances definitive because some of the groups mentioned are still in need of

revision.

I. Section Urostigma (Gasp.) Miq.

Ficus religiosa L. (see Wiebes, 1967: 419—434): 1, Platyscapa quadraticeps (Mayr) (Wiebes, 1977

b: 214); 2, Philotrypesis anguliceps (Westwood) (Abdurahiman & Joseph, 1975 b: 78); 3, Sycoryctes

religiosae Wiebes; 4, Sycoscapteridea monilifera (Westwood); 5, Watshamiella infida Wiebes*; 6,

Eurytoma spec.

II. Section Conosycea (Miq.) Corner

Ficus benghalensis L. (see Wiebes, 1967: 400—419): 7, Eupristina masoni Saunders; 8, Philotrype-

sis transiens (Walker) & affinis (Westwood); 9, Sycoscapter stabilis (Walker) (S. insignis Westwood,

1883: 35, d; Indothymus crenulatus Joseph, 1953: 77, ®); 10, Sycoryctes spec.; 11, Sycoscapteridea

guruti Joseph & Abdurahiman (1969: 42); 12, Micranisa pteromaloides (Walker); 13, Walkerella

temeraria Westwood (Joseph & Abdurahiman, 1969: 44); 14, Sycobia bethyloides Walker; 15,

Sycophilodes moniliformis Joseph; 16, Sycophilomorpha saptapurensis Joseph & Abdurahiman

156

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Table 1. Groups and selected genera of fig insects, with the Ficus species from which they were

Fig wasps:

Blastophaginae

Agaoninae

Sycoecinae

Sycophagini

Apocryptini

Philotrypesini

Sycoryctini

Otitesellinae

Epichrysomallinae

Eurytominae

Ormyrinae

Platyscapa

Eupristina

Waterstoniella

Blastophaga

Dolichoris

Kradibia

Ceratosolen

Pleistodontes

Allotriozoon

Elisabethiella

Alfonsiella

Crossogaster

Phagoblastus

Sycophaga

Eukoebelea

Parakoebelea

Idarnes

Apocrypta

Philotrypesis

Sycoscapter

Arachonia

Sycoryctes

Sycoryeteridea

Sycoscapteridea

Watshamiella

Otitesella

Micranisa

Walkerella

Grandiana

Grassetana

Sycobia

Pembertonia

Sycophilodes

Sycophilomorpha

Sycotetra

Camarothorax

Eurytoma

Syceurytoma

Ficomila

Sycophila

Ormyrus

— religiosa

benghalensis

etupenda

reared.

macrophylla

eriobotryoides

thonningtt

callosa

eartea

exasperata

laterifolia

nota

variegata

racemosa

75,

eycomorus

(1969: 37); 17, Sycophila decatomoides Walker and other species (? Decatoma, Joseph & Abdurahi-

man, 1968).

Ficus stupenda Miq. (see Wiebes, 1966 b: 163—192): 18, Waterstoniella masii (Grandi); 19,

“Blastophaga” errata Wiebes; 20, Sycoscapter reticulatus Wiebes; 21, Arachonia borneensis Wiebes;

22, Sycoryctes hilli Wiebes; 23, Watshamiella stilifera (Wiebes)*; 24, Grandiana corneliae Wiebes.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 157

III. Section Malvanthera Corner

Ficus macrophylla Desf. ex Pers.: 25, Pleistodontes froggatti Mayr; 26, Philotrypesis aterrima

Saunders; 27, Sycoryctes australis (Froggatt); 28, Pembertonia ficicola Gahan.

IV. Section Galoglychia (Gasp.) Endl.

Ficus cf. eriobotryoides K. & B. (see Wiebes, 1971: 367—383): 29, Allotriozoon prodigiosum

Grandi; 30, Philotrypesis finitimorum Wiebes; 31, Sycoscapter montis Wiebes; 32, Sycoryctes

lomaensis Wiebes; 33, Otitesella royi Wiebes; 34, Camarothorax africanus (Wiebes)*.

Ficus thonningü Bl. (present paper): 35, Elisabethiella stuckenbergi (Grandi); 36, Alfonsiella

brongersmai Wiebes & longiscapa Joseph; 37, Crossogaster odorans Wiebes*; 38, Phagoblastus

barbarus Grandi; 39, Philotrypesis parca Wiebes*; 40, Sycoscapter cornutus Wiebes*; 41, Sycoryctes

remus Wiebes* & hirtus Wiebes*; 42, Watshamiella alata Wiebes*; 43, Otitesella tsamvi Wiebes*; 44,

Sycotetra serricornis Boutek*; 45, Camarothorax equicollis Boutek*, longimucro Boutek* &

brevimucro Boutek*; 46, Eurytoma ficusgallae Boutek*; 47, Syceurytoma ficus Boutek*; 48,

Ficomila curtivena Boutek* & gambiensis (Risbec)*; 49, Sycophila kestraneura (Masi)*, punctum

Boutek*, sessilis Boutek*, modesta Boutek*, naso Boutek* & flaviclava Boutek*; 50,

Ormyrus watshami Boutek*, subconicus Boutek* & flavipes Boutek*.

V. Section Oreosycea (Miq.) Corner

Ficus callosa Willd.: 51, Dolichoris malabarensis (Abdurahiman & Joseph) (Wiebes, 1979 a: 188);

52, Sycoryctes callosa Abdurahiman & Joseph (1975 a: 103); 53, Sycorycteridea keralensis Abdurahi-

man & Joseph (1975 a: 99); 54, Grasseiana callosa Abdurahiman & Joseph (1967: 14).

VI. Section Ficus L.

Ficus carica L.: 55, Blastophaga psenes (L.); 56, Philotrypesis caricae (L.); 57, Sycoscapteridea

raoi Joseph.

VII. Section Sycidium Mid.

Ficus exasperata Vahl: 58, Kradibia gestroi (Grandi) (Wiebes, 1978 a: 176); 59, Philotrypesis

quadrisetosa (Westwood); 60, Sycoscapteridea longipalpus (Joseph).

Ficus laterifolia Vahl: 61, Kradibia cowani Saunders (Wiebes, 1978 a: 174); 62, Sycoscapter gibbus

Saunders (Wiebes, 1978 b: 187).

VIII. Section Sycocarpus Miq.

Ficus nota (Blanco) Merr.: 63, Ceratosolen notus (Baker); 64, Eukoebelea nota (Baker); 65,

Apocrypta larvalis (Baker); 66, Philotrypesis similis Baker; 67, Sycoryctes bakeri Wiebes.

IX. Section Neomorphe King

Ficus variegata Bl.: 68, Ceratosolen appendiculatus (Mayr); 69, Eukoebelea spinitarsus (Mayr);

70, Philotrypesis bimaculata Mayr; 71, Sycoryctes patellaris Mayr.

X. Subgenus Sycomorus (Gasp.) Miq.

Ficus racemosa L.: 72, Ceratosolen fusciceps (Mayr); 73, Eukoebelea brevitarsis (Grandi); 74,

Parakoebelea stratheni Joseph; 75, Idarnes testacea (Mayr); 76, Apocrypta westwoodi Grandi; 77,

Sycoryctes patellaris Mayr.

Ficus sycomorus L. (see Wiebes, 1968: 307—320): 78, Ceratosolen arabicus Mayr & galili Wiebes;

79, Sycophaga sycomori (L.); 80, Eukoebelea sycomori Wiebes; 81, Parakoebelea gigas (Mayr); 82,

Idarnes gracilis Wiebes; 83, Apocrypta longitarsus Mayr; 84, Syceurytoma ficus Boutek*; 85,

Ficomila gambiensis (Risbec)*; 86, Sycophila naso Boutek*.

REFLECTIONS ON THE CLASSIFICATION OF CHALCIDOID FIG WASPS

It is still rather premature to give proper taxonomic assignment of most

* New species or new combination published in the present paper.

158 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

chalcidoid groups associated with the figs, to the families as they are recognised

today. It can be said that the closer is this association, the more difficult it is to

trace the links with the most closely related non-fig-associated groups. Despite

certain controversies most authors agree that the genus Ficus was already

present and abundant in the Cretaceous period. However, there is but one fossil

record of an Agaonid (Brues, 1910) and also records of the other Chalcidoidea

are very scanty, especially of the groups of the “pteromaloid complex”, i.e.,

families most closely related to Pteromalidae, including Torymidae, Eurytomi-

dae and Agaonidae. So far the main source has been an analysis of the morpho-

logical characters, supplemented by some biological evidence. This analysis

seems to suggest that at least some of the relevant groups have nothing to do with

the Torymidae under which most of the fig wasps (apart from Agaonidae) have

been recently classified.

Agaonidae are best regarded as a separate family. However, it seems possible

that the “‘unplaced”’ Sycoecinae are related to them, probably as a plesiomorphic

(less specialised) group, pointing perhaps to a common origin of both groups in

the pteromaloid complex. All the same it must be stressed that a placement of

Sycoecinae in Agaonidae is still very problematic. |

Apart from the neotropical genus Physothorax Mayr, which undoubtedly

belongs to Torymidae, subfamily Toryminae, two other groups seem to have

closer links with Torymidae, viz., Apocryptini and Sycophagini. They might be

regarded as two tribes of a special subfamily Sycophaginae. Another, but more

questionably torymid subfamily may constitute the Sycoryctinae. They may

include two tribes, Sycoryctini (here also the neotropical Critogaster Mayr) and

Philotrypesini. Both are undoubtedly closely related, but there are almost as

many reasons to classify them under Pteromalidae as are in favour of the

Torymidae.

A few further groups seem to be placed best in the family Pteromalidae. Here

belong Otitesellinae which include, apart from several palaeotropical genera

(listed e.g., in Hill, 1967) also the neotropical Aepocerus Mayr and Heterandri-

um Mayr. Yet another pteromalid subfamily, although completely different

from Otitesellinae, may be Epichrysomallinae. They may retain the subfamily

status, being sufficiently different from, although related to, the Brachyscelidi-

phaginae (Epichrysomalla Girault was originally described along with several

genera of the latter group), possibly as their apomorphic sister group, as

suggested by Leeweniella Ferrière. Both groups are of phytophagous habits, but

Epichrysomallinae are associated exclusively with figs (palaeotropical), galling

the female florets as do the Agaonidae, but apparently ovipositing through the

fig wall. Otherwise the coiled ovipositor reminds us much of certain Eurytomi-

dae. However, Epichrysomallinae do not seem to be closely related to the

eurytomids, as found independently, on other evidence, also by M. Copland (in

litt.).

The remaining groups Eurytomidae and Ormyridae do not pose many prob-

lems, but it should be noted that the latter group shows close relationship with

Torymidae and should perhaps be better regarded as a subfamily of Torymidae,

as they were in the past.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 159

Thus, in the present paper the following classification is used:

Family Subfamily Tribe

Agaonidae Agaoninae

Blastophaginae

(not placed) Sycoecinae

Torymidae Toryminae

Sycophaginae Sycophagini

Apocryptini

Ormyrinae

Torymidae Sycoryctinae Philotrypesini

or Pteromalidae Sycoryctini

Pteromalidae Otitesellinae i

Epichrysomallinae

Eurytomidae Eurytominae

In the following descriptive part, the groups are treated in the sequence of the

above classification, which is that given by their association with the figs; for the

reason of their rather loose association with Ficus, however, the Ormyrinae are

treated last. The Blastophaginae and Sycophaginae are not represented in Ga-

loglychia-figs, as is evident from table 1.

pay

KEY TO THE FIG WASPS FROM FICUS THONNINGII

MEullwWineedsfemalesandimales eee ere EEE RENE COORTE 2

APLeTOUSONbDEACRYpDIE TOUS AMAlES ONLY PE CEE EEE EC RE ERE 36

. Females: gaster with distinct tail formed either by the ovipositor and its

sheaths, or also by extremely narrowed last one or two tergites ............ 3

Females without such a slender tail: ovipositor sheaths short, hardly ex-

serted; or males, i.e., with last sternite virtually reaching apex of gaster .... 12

WINGED FEMALES WITH TAILS

Narrow gastral tail formed for a great part by one or two ultimate tergites ... 4

Last tergite short, tail formed by the ovipositor and its sheaths ............ 8

Tail formed in anterior part by two ultimate tergites (the proximal of which is

recognizable by the very small spiracular peritremes) and, in hinder portion,

by long protruding ovipositor and sheaths (Philotrypesini) ................

NEE EBEN BEN ED ieee HORDE ESS E DE EE Philotrypesis parca

Tail formed by the last tergite, i.e., the apparent gaster ending with the

eighth urotergite, which bears the spiracular peritremes (Sycoryctini) ...... 5

. Stigmal vein produced into a boot-like stigma (fig. 64), the fore wing with

long setae in the marginal-stigmal angle; the epistomal margin produced

intojantacutesprojection(figs65) rr er Sycoscapter cornutus

Stigmal vein more normal, the wings at most with some long setae along the

veins (but may be pilose all over); the epistomal margin straight or with wide

TODESPRPARMA NEE EN DAN LE MER AE LO Or 6

. Epistomal margin widely lobed (fig. 68); the wings hyaline but for some

160

10.

JUL

12}

13;

14.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

Setacalongtheveins(Syconyctes) EEE E EE EE TEE 7

Epistomal margin straight (fig. 62); the wings evenly pilose ................

cre Anka an he sie RR Watshamiella alata

The legs up to and including the tibiae dark .............. Sycoryctes remus

The legs from the femora onwards yellowish ............ Sycoryctes hirtus

. Antennal scape strongly broadened; prognathous head with mandibles

provided with a denticulate appendage; marginal vein normal and slender,

postmarginal short but not rudimental (Agaoninae) ..................... 9

Antenna of more normal form; mandibles without such appendage, al-

though they bear teeth directed backwards; marginal vein thickened and

longer than rudimental postmarginal (Sycoecinae) ..................... 11

Funicular segments of antenna with sensilla linearia. General aspect, fig. 5 . . . .

A Seed eet rey ite Re IREH RER. For Elisabethiella stuckenbergi

siella) - fatten de i ate Re: Ia ei MR SEHEN DAR Ohr O 10

Head scarcely longer than wide (fig. 14), mandibular appendage with

twenty ventral rows of teeth; antennal scape not produced..................

iran iena ati Babe 2 Alfonsiella brongersmai

Head distinctly longer than wide (fig. 15), mandibular appendage with

thirty-five ventral rows of teeth; antennal scape with an apical process. . ......

RAMUS VARO ODE SPR ATE OSM ER CR Alfonsiella longiscapa

Antenna with one anellus, funicle with long sensilla chaetica; fore tibia with

adorso-apicaly bidentate hook Ere wer SRO Crossogaster odorans

Antenna with two anelli, funicle with sensilla linearia; fore tibia with a comb

oficonicaliteettà i SIC RO ... Phagoblastus barbarus

WINGED FEMALES WITHOUT TAILS AND WINGED MALES

Gaster with at least some coarse sculpture; marginal vein very long, at least

seven times as long as short stigmal vein; antenna thirteen-segmented, with

twolanell'(Ormyrinae) am me Ze ee ie ee EEEE 13

Gaster without any coarse sculpture, either smooth or virtually so; marginal

vein never so much longer than the stigmal vein ....................... 15

Mid lobe of mesoscutum in both sexes mainly bare, pilosity reduced to few

dark bristles on posterior half; in 2 gaster mostly not longer than head plus

thorax combined, dorsally keeled; femora and tibiae dark; in d legs and

sides of thorax testaceous, gaster flat and shiny, two middle tergites usually

each with two exposed rows of deep foveolae .......... Ormyrus watshami

Mesoscutum (and scutellum) all over with regular subdecumbent pilosity;

othercharacters also pattly.different.... nen. er ne EEEN: 14

Femora and tibiae in both sexes testaceous; in © gaster anteriorly rather

high, compressed, but its apical third (from middle of sixth tergite) abruptly

narrowed and slightly upturned; middle tergites each with a row of raised

short ridges but deep foveolae at base of tergites often concealed; in d

gaster slightly dull (also between foveolae), fairly densely punctulate ........-.

Set t o, Gail CORO Pet Ormyrus flavipes

Femora, sometimes also tibiae, in both sexes more or less infuscate and

15}

16.

18.

20.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 161

pedicellus slightly shorter than in alternate; in © gaster not high, almost

gradually tapering, subconical, usually tergites 3 to 5 with exposed deep

foveolae at base, just in front of serrate rows of short ridges; in d gaster with

distinct smooth interspaces between deep foveolae .. Ormyrus subconicus

Body with metallic gloss, very finely reticulate; in 9 apex of gaster slightly

eunyineäyentrad.(Otiteselinae)er nee een Otitesella tsamvi

Body non-metallic, frequently either almost smooth or with distinct sculp-

ture on thorax and mostly also on head; apex of gaster in © often curving

dorsadsinever VentEad wen. Aula lie eee ARI AIRES 16

Thorax (and head) with distinct sculpture and pubescence; stigmal vein

usually at acute angle or short marginal vein distinctly expanded (Eurytomi-

naer he entre re bonsai eta RENE 17

Body without distinct sculpture, although often microscopically alutaceous;

thorax always with pubescence reduced to pairs of bristles; marginal vein not

so short or, if so, then stigmal vein arising at nearly right angle ........... 26

Gena posteriorly with strong and distinctly raised carina extending up the

temple; thorax densely umbilicately punctured ........................ 18

Gena posteriorly with an edge which is blunt or at most indistinctly carinate

in middle or at mandible; umbilicate punctures, if present on thorax, usually

notiveryidenseir tr A AE ETA EME 19

Marginal vein distinctly swollen; gaster on distinct petiole which has ante-

riorly a transverse crest; fore coxa antero-ventrally flat; mesosternal shelf

very short, in middle with small rounded projection which is hollowed from

both sides; genal carina smoothly joining mouth margin; body usually

extensively rufous, in darkest specimens at least ventrally . . Syceurytoma ficus

Marginal vein normal, slender; gaster in 2 virtually sessile; fore coxa on

antero-ventral surface with strong oblique carina; mesosternal shelf sepa-

rated by sinuate cross-carina; genal carina with a tooth at base of mandible,

body black, legs beyond coxae usually orange ........ Eurytoma ficusgallae

Mesosternal part of thorax with distinct though not high median projection

excavated from the sides and, sublaterally, with slight carina marking off

mesosternal shelf; marginal vein in © not longer than stigmal vein which is

usually distinctly curved; wing sometimes without fuscous spot (macula)

(Economia. BR, meh RETTET NIE 20

Mesosternum regularly sloping, posteriorly with median hairy depression

(at mid coxae), but without any indication of delimited mesosternal shelf;

marginal vein always with fuscous macula and frequently longer than the

stema Velni(SY RUE ZI

In © gaster dorsally broad, with fourth tergite about twice as long as the

third; thorax dorsally almost always black, with dense umbilicate punctura-

tion and conspicuous white pubescence; fore wing whitish, without infus-

cate macula, its pilosity, including marginal fringe, very short, inconspicu-

ousadiunknown ME. FETI, SI REIT SI AREN Ficomila curtivena

In ® gaster compressed from sides, dorsally subcarinate, its fourth tergite

hardly longer than the third; thorax usually yellowish, its pilosity and

puncturation often not conspicuous; fore wing with small fuscous macula on

Maroinalavcin wee aes n Ri ho biro Ficomila gambiensis

162

JAY,

22%

23)

24.

25}

26.

DAR

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

Pedicel in both sexes distinctly shorter than first funicular segment; marginal

vein swollen, i.e. with its lower margin convex, often strongly so (in dark

larger 6); in 2 wing below submarginal vein bare or with up to 2 hairs and

petiole distinctly elongate (about 1.5 x 1), gaster compressed, dorsally

keeleden tedere eo ee corte Sycophila kestraneura

Pedicel nearly as long as or even longer than first funicular segment;

marginal vein widening distad but its lower margin (sometimes obscured by

macula) either straight or even concave; partly different also in other

characters fi sur. whee). Lenci ERE 22

Exposed part of prepectus with some hairs (fig. 144); in 2 wing below

submarginal vein bare or with up to 2 hairs only; marginal vein only

moderately widened and rather short, macula small, usually round and not

reaching base of marginal vein; in d head subglobose, stout, 1.5—1.65

times as broad as long in dorsal view .................. Sycophila punctum

Prepectus completely bare (occasionally with 1—2 hairs in sessilis); in 9

wing below submarginal vein at least with a sparse hairline on basal fold; in

6 head more transverse thaninalternate oe AREE 23

Gaster in © virtually sessile (fig. 148) as dorsal part of petiole is strongly

transverseso unknown an SESSI SS Sycophila sessilis

Gaster distinctly petiolate in both sexes, petiole in 9 mostly elongate in

dorsal view: gites une: sarai alice lose EEE 24

Malar space slightly shorter than, or at most virtually as long as half of

longest diameter of eye, lower extremity of eye subangular (fig. 150); in ©

antenna darkened but with yellow apex; interantennal crest rather steep

into:serobakeavit ya ee Sycophila flaviclava

Malar space at least slightly longer than half of longest eye diameter, eye

broadly rounded in lower part (figs. 149, 151, 152); antenna in © uniformly

darkened; not yellowiatiapex «u... Inner. moeren GEE 25

Fore wing on dorsal surface of costal cell in 9 with distinct row of erect hairs;

interantennal crest gradually descending into scrobal cavity; © gaster

usually with darker bands on anterior tergites; pedicel at most about 2.2

timesiasdongias broad ayy a ne Er Sycophila modesta

Dorsal surface of costal cell in 2 bare or at most with odd hair; interantennal

crest mostly perpendicularly descending into scrobes, forming a subrectan-

gular tooth; 2 gaster usually unicoloured, yellowish or blackish; pedicel in

both sexes with flat dorsal outline, usually more than 2% times as long as

broadis sh ade er Sycopnila naso

Only males: fore wing wholly pilose, narrow, marginal vein very long and

postmarginal fully twice as long as the stigmal; antennal flagellum not or

hardlyllongerthaniscapusia Ario Era MR ae Watshamiella alata

Both sexes; in male venation different, wing at base extensively bare,

flagellum longer: voos uier EIER IT EEE Di

Only males: marginal vein thickened and at least 1.9 times as long as the

stigmal, wing in area below marginal vein and parastigma pubescent; head

flat;:with long mandibles (Sycoecinae) tre: ERRATE 28

Both sexes: in male marginal vein not so long and thickened, area below this

vein at least partly bare; head different (Epichrysomallinae)

ee ee ee RO ee ee

28.

29}

36.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 163

Head virtually as long as wide, with antennal toruli partly below lower ocular

line; marginal vein fully twice as long as the stigmal; outer tooth of mandible

shorter than the inner tooth (fig. 36) ................ Crossogaster odorans

Head slightly but distinctly longer than wide, with antennal toruli distinctly

above ocular line; marginal vein not quite twice as long as the stigmal; outer

tooth of mandible longer than inner tooth (fig. 37) .. Phagoblastus barbarus

Tarsi 4-segmented; notauli very superficial, in form of dark lines not nearly

reaching transscutal line; scutellum anteriorly separated from axillae only

by lines, not by grooves; funicular segments in both sexes separated by

distinct incisions, in © funicle serrate and 6-segmented, in d 5-segmented ....

PRA LR Sycotetra serricornis

Tarsi 5-segmented; notauli groove-like, complete or nearly so (some d);

axillar grooves also complete; flagellar segments in ? never serrate, flagel-

lum fairly compact and sometimes with different number of segments; in d

funicle either with 5 or 7 segments (Camarothoraxd) ae Se RER ERRT 30

. Females (gaster compressed, high, dorsally keeled) .................... 31

Males (gaster flat, much shorter than thorax) ......................... 33

. Antenna with 7 funicular segments plus small anellus; pedicel subglobose .......

BETEN TIT VITO: EOS] EIERN TOGA ENT Camarothorax equicollis

Only 6 funicular segments, a small anellus and an elongate pedicel (at least

PPA RA een EN URES PRONE tisi nice 32

. Gaster strongly compressed, about as long as head plus thorax, its hypopygi-

um not quite reaching two-thirds along ventral edge but bearing at apex a

long thin whitish projection; clypeus dorsally separated from supraclypeal

akeaibyidistinct lino EN Camarothorax longimucro

Gaster relatively broader and only about as long as thorax, hypopygium

reaching near to apex of gaster and bearing a short projection; clypeus

dorsally not separated from supraclypeal area.. Camarothorax brevimucro

. Antenna with 5 funicular segments; clypeus not separated dorsally from

supraclypeal area; body yellowish ............. Camarothorax brevimucro

With 7 funicular segments; clypeus separated dorsally by a dark line from

supraclypeal area; body often brownish .............................. 34

. Head and pronotum unusually long, each longer than broad; notaular

grooves almost meeting at scutellum ............ Camarothorax equicollis

Head and pronotum shorter, each at least slightly broader than medially

lone-noraulimonezaparspostenionlyrz te SARNO RO OR AE OE 35

. Head only slightly broader than long, also pronotum relatively long; body

brown, legs partly yellowish ........ (? a form of Camarothorax equicollis)

Head obviously shorter than broad, its genae converging; pronotum trans-

versesbodysyellowar Sem an ee Camarothorax longimucro

APTEROUS MALES

Solenogastrous i.e., the last four or five gastral segments tubularly length-

ened; antenna reduced to five or six segments (Agaoninae) ............. 57

Last gastral segments not tubularly lengthened; antenna consisting of at

leastitenisepments! or NAA NN 39

164

41.

42.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

. Antennae situated in a common groove (fig. 7), the club slender, not much

wider than the funicle (fig. 10) ................ Elisabethiella stuckenbergi

Antennae situated in separate sockets, the club much wider than the funicle

(iesl3)Alfensiellay nen hr.) MER ee 38

. Head rather wide in front, the width just behind the eyes little smaller than

the maximum width. General aspect, fig. 13 ...... Alfonsiella brongersmai

Head distinctly narrowing frontad, the width just behind the eyes four-fifths

ofthemaximum width er ee SE Alfonsiellalongiscapa

. Antennae inserted high on face, at the upper ocular line, their toruli wide

apart; tarsi 4-segmented, antennae 10-segmented (Otitesellinae)............

Atesina bay. Be of il a oe Ro Otitesalla tsamvi

Antennae inserted near to mouth, below lower ocular line; tarsi 5-seg-

mented, antennae consisting of 11 or 12 segments (Sycoryctinae) ......... 40

. Segments of antennal flagellum normal, gradually changing in shape; two

basal segments of hind tarsus strongly expanded dorsally; hypostomal

margin of characteristic shape: deeply incised between sublateral lobes (fig.

49) (Philotrypesini) Ar er DE Philotrypesis parca

Flagellum with partly alternating larger and smaller segments, especially

second funicular segments always smaller than the first; hind tarsus at most

with first segment expanded; hypostomal margin at most moderately emar-

ginate (or straight) (Sycoryctiml) 22... 22.0... 4-4). 20 ee 41

Tarsi with enlarged last segment but basal segments small, especially on fore

tarsus (fig. 58); antenna 11-segmented .............. Sycoscapter cornutus

Tarsi different, last segment not greatly enlarged, on hind tarsus first

segment distinctly longer than the second; antenna 12-segmented (Sy-

Conyetes) en. anal. uve ia herten 42

Head posteriorly with conspicuous cervical shield; head and thorax with

abundant pilosity; antennal club usually much darker than the slender

funiculus; hind basitarsus at most only moderately broadened ..............

atleta Sooke Le soe N RS ERO Sycoryctus hirtus

Head posteriorly only with the usual raised cross-carina; head and prono-

tum, apart from usual bristles, nearly glabrous; funicular segments of

antenna rather wide, club less distinctly set off, pale; hind basitarsus often

ereatly;enlarseda e nat doren ted ee er Sycoryctes remus

AGAONINAE

In the present material, three species of Agaoninae are represented, viz., two

of Alfonsiella Waterston and one of Elisabethiella Grandi. Evidently, either can

act as a pollinator, but the true relationships are not understood.

A key to the African genera of the Agaonidae was published by Wiebes

(1974a: 34—36), one to the species of Elisabethiella in the same paper (p. 30—

31). The species here treated is a member of the species group of E. enriquesi

(Grandi), with which it will be compared in the description below.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 165

Elisabethiella stuckenbergi (Grandi)

(figs. 5, 7—12; pl. 1 fig. 1, 3)

Blastophaga (Elisabethiella) stückenbergi Grandi, 1955: 85—87. Holotype ?, South Africa: Gra-

hamstown (LEB) [examined].

Female. — Length of the head, thorax and gaster ca. 1.4 mm. Colour dark

brown to blackish, the antennal scape and the tarsi of the legs lighter.

Head as long as wide across the compound eyes; the eye as long as the cheek.

Mandible with five ventral ridges, two apical teeth and two glands; the append-

Figs. 7—12. Elisabethiella stuckenbergi (Grandi), female (12) and male (711). 7, head and thorax;

8, fore tibia and tarsus; 9, hind tibia and tarsus; 10, left antenna, ventral aspect; 11, left mandible,

ventral aspect; 12, hypopygium. Figs. 13—15. Alfonsiella, sketches of: 13, male of A. brongersmai

Wiebes; 14, female of A. brongersmai Wiebes; 15, female head and part of thorax of A. longiscapa

Joseph.

166 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

age four times as long as wide, with about twenty rows of ten to twelve

denticulations (except for the proximal and distal rows, which have less).

Labium with two apical setae, the maxilla with two subapicals. Antenna: the

scape three times as long as the rotundate pedicel; the segments, from the fifth

onwards, with rather long sensilla covering the whole length of the segment.

Thorax with large mesosternal pollen pockets; shallow coxal corbiculae and a

distinct comb. Fore tibia with an apical comb of three teeth and with one ventral

tooth and a spur; the tarsal segments approximately in ratio 7 : 3 : 3 : 3 : 6. Mid

leg: tarsal ratio 4:3:3:3:4. Hind tibia with two bidentate antiaxial and a

slender, simple axial teeth; the tarsal segments approximately in ratio

8:4:3:2:5. Fore wing (ca. 2 : 1) 1.3 mm long, the submarginal, marginal,

stigmal and postmarginal veins approximately in ratio 10 : 2: 3 : 4; the hind

wing (4 : 1) 0.7 mm long.

Gaster: the hypopygium (fig. 12) has a rather short spine with a blunt apex

(unknown for E. enriquesi). The ovipositor is as long as the body, over two times

as long as the gaster (27 : 13).

Male. — Length of the head and thorax ca. 0.9 mm. Colour yellow-brown.

Head (fig. 7) as long as wide; the eye approximately one-fifth of the length of

the head. The dorsal surface, especially in the anterior third, has rather stout

spines directed backwards. Mandible (fig. 11) with one apical tooth and with one

gland; other mouthparts atrophied. Antennae (fig. 10) situated in a common

groove; there are five free segments (formula 1121), the third and fourth of

which are annuliform; the club is more than twice as long as wide (20 : 9).

Thorax, fig. 7. The pronotum has a distinct collar, the sclerite itself is shorter

than the combined lengths of the posterior sclerites (10 : 11) and longer than

wide posteriorly (20 : 19). As seen from the lateral indentations, the meso-,

metanotum and propodeum are subequal in length; the mesonotum is wider

than the metanotum (19 : 14), the propodeum is narrower (12). The spiracles

occupy one-third of the lengh of the propodeum. Fore tibia (fig. 8) with three

teeth in a dorso-apical crest and two short conical spines on the dorsal margin, a

bidentate ventral apex; the tarsal segments all distinctly separate, approxi-

mately in ratio 8 : 3 : 2: 1 : 7, the first and second with two and one axial cones,

respectively. Mid leg: tarsal ratio 6:5:6:5:7. Hind tibia (fig. 9) with a

bidentate apical crest at the antiaxial disk and a faintly bicuspidate tooth axially

(Grandi mentioned a tridentate crest for E. enriquesi, which must have been the

bidentate crest and the axial tooth seen as one); the tarsal segments approxi-

mately in ratio 8:5:4:5: 10, the basitarsus has four conical spines in two

rows (in one line in E. enriquesi) and the second has one ventral cone.

Gaster: the genitalia simple.

Material. — Series 236, Zimbabwe-Rhodesia: Makumbi Mission, Chinamo-

ra Reserve, ex Ficus burkei, xii.1976 (A. Watsham); Salisbury, ex Ficus burkei,

series £ 8, ii-v.1977 (A. Watsham). Zambia: Kafue National Park, Ngoma, ex

Ficus burkei, 6 8, 8.xii.1977 (M. G. Bingham, no. 2465); Lusaka, Chelston

School, ex Ficus burkei, 7.1.1979 (M. G. Bingham, no. 2472); Chipata Golf

Course, ex Ficus burkei, 14 2, 11.1.1979 (M. G. Bingham, no. 2474); Lusaka,

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 167

Cathedral of the Holy Cross, ex Ficus burkei, 50 2 50 6, 16.11.1979 (M. G.

Bingham, no. 2490). The material is deposited in the RMNH, Leiden, some

duplicates are in the BMNH, London and in the Watsham Collection.

Comments. — The number of crenulations in the transverse rows on the

mandibular appendage was not clear from Grandi’s description of the female;

now, the data used in my key (Wiebes, 1974a: 31) prove to be incorrect. The only

differential character that remains with the female of E. enriquesi is the longer

head. The male, too, should be compared with E. enriquesi and also with E.

allotriozoonoides (Grandi), to which Grandi referred when describing E.

enriquesi. The following couplets have to be inserted into my key to accomodate

for the species mentioned.

1. Female: the antennal pedicel angular in outline, longer than wide. Male: the

head distinctly longer than wide (7:6), the pronotum as long as the

composite posteriorsclerite. Eritrea, ex Ficusspec.......... allotriozoonoides

— Female: the antennal pedicel rotundate in outline. Male: the head, if at all,

only little longer than wide, the pronotum shorter than the posterior sclerite.. 2

2. Female: the head distinctly shorter than wide across the compound eyes

(6 : 7). Male: the fore tibia without conical spines on the dorsal margin; the

secondtofifthtarsalsegmentsfused. Angola, ex Ficusspec.......... enriquesi

— Female: the head as long as wide. Male: the fore tibia with two short conical

spines on the dorsal margin; all tarsal segments free. South Africa, Rhodesia

ANdZAMDIA EX ALCUS DUREE ECC ER EEE CO O stuckenbergi

Alfonsiella Waterston

There are four described species of Alfonsiella, mainly known from light

catches. In the present material (which is retained in the RMNH, Leiden for

further revision) two species are represented, viz., A. brongersmai Wiebes and

A. longiscapa Joseph, but in different samples. The two species are recognizable

by the following characters (Wiebes, 1972).

Female. — Length of the head, thorax and gaster I—2 mm. Colour yellowish.

Head scarcely longer than wide in A. brongersmai (fig. 14), distinctly longer

than wide in A. longiscapa (fig. 15); the compound eye 1/2—2 times as long as

the cheek. Two ocelli. Mandibular appendage with about twenty (A.

brongersmai) or thirty-five ventral rows of teeth (A. longiscapa). Antenna: the

scape with an apical process in A. longiscapa, without such a process in A.

brongersmai; the third segments rather simple, the funicular sensoria very long

and flexible.

Thorax with mesosternal pollen pockets, and also the fore coxa with a

corbiculum. Fore tibia with two dorso-apical teeth and one ventral. Hind tibia

with two ventral spurs. Postmarginal vein of fore wing as long or half as long as

the stigmal (A. longiscapa and brongersmai, respectively).

Gaster: the ovipositor about as long as the gaster.

Male. — Length of head and thorax 1 mm. Colour yellowish, but the head may

be darker, especially the large mandibles; the mid femur and tibia dark and

setose.

168 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Head little longer than wide in A. brongersmai (fig. 13), distinctly longer than

wide (5 : 4) in A. longiscapa; with rather large compound eyes, one-quarter of

the length of the head. Mandible heavy, triangular in lateral aspect. Antenna six-

segmented (formula 1131).

Thorax consisting of two sclerites. Fore tibia with two dorsal hook-like teeth

and one simple ventral, the dorsal edge with conical spines; the tarsus oligomer-

ous. Mid leg strikingly darker than the other legs, the femur and tibia setose.

Hind tibia: two ventral spurs, the dorsal margin with conical spines.

Gaster: genitalia simple.

Alfonsiella brongersmai Wiebes

(figs. 13, 14)

Alfonsiella brongersmai Wiebes, 1972: 326. Holotype ©, Kenia: Nairobi (RMNH, Leiden) [exam-

ined].

Material. — Zimbabwe-Rhodesia: Salisbury, ex Ficus burkei, 2 d, 11.1977

(A. Watsham). Zambia: Chipata, ex Ficus petersii, 14 2 64, 11.1.1979 (M. G.

Bingham, no. 2475); Lusaka, Cathedral of the Holy Cross, ex Ficus burkei, 1 9,

16.11.1979 (M. G. Bingham, no. 2490).

Alfonsiella longiscapa Joseph

(fig. 15)

Alfonsiella longiscapa Joseph, 1959: 30. Holotype ®, Guinea: Mts. Nimba (MNHN, Paris)

[examined]; Wiebes, 1972: 328—330.

Material. — Zambia: Luapula Prov., Mununshi Banana Estate, ex Ficus cf.

burkei, 23 2 5 3, 9.xii.1978 (M. G. Bingham, no. 2471).

SYCORYCTINI

The Sycoecinae have alate males, and females modified for penetrating the fig

ostiole. They are easily distinguished by the wing veins in that the marginal is just

longer than the stigmal, and thickened; the postmarginal is subequal to the

stigmal but very sharply tapering (Hill, 1967: 98).

Grandi (1955: 101, nota) concluded when making his key to the genera of the

Sycoecinae (Sycophagini in his sense, pro parte), that the classification of this

group is still unstable. Even now Lipothymus Grandi and Eujacobsonia Grandi

are allocated to the Otitesellinae and Sycophaga Westwood to the Sycophagini s.

str. (Wiebes, 1966a), there still is some ambiguity in the definitions of Crossogas-

ter Mayr and Phagoblastus Grandi. Six species are included: Crossogaster

triformis Mayr (1885: 189—192, figs. 20-22; redescribed by Grandi, 1928:

203-206, figs. xlii-xliii, the apterous male excluded), Crossogaster silvestrii

Grandi (1916: 253—264, figs. xli-xliv), Crossogaster atrata Masi (1917: 125—

126, fig. 3), Crossogaster odorans Wiebes, spec. nov., described below, Phagob-

lastus barbarus Grandi (1955: 102—106, figs. viii-ix), the male described below,

and Phagoblastus liodontus Wiebes (1979b: 397—400, figs. 20—37).

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 169

It appears that there are three species groups, as follows: Crossogaster s. str.,

species C. triformis and odorans. — Epistomal margin entire; antenna with one

anellus; labial palpus consisting of one segment, the maxillary one- or indistinct-

ly two-segmented!); female fore tibia with a dorso-apical, bidentate hook;

stigma of the eighth urotergite especially in the female large, ovoid.

Phagoblastus, species P. barbarus and liodontus. — Epistomal margin entire;

antenna with two anelli; labial palpus consisting of two segments, the maxillary

of three; female fore tibia with a row of teeth.

Crossogaster, group of C. silvestrii and atrata. — Epistomal margin cleft in the

middle; antenna with two anelli; labial palpus of silvestrii consisting of one

segment, maxillary of two (atrata: 2, 3); female fore tibia with a row of conical

spines.

Table 2. Host relations of Crossogaster and Phagoblastus.

Ficus Crossogaster Phagoblastus Elisabethiella Allotriozoon references

vasta triformis!) socotrensis Wiebes, 1977b: 210

burkei odorans barbarus stuckenbergi present paper

petersii barbarus stuckenbergi Grandi, 1955: 106

leprieuri liodontus Wiebes, 1979b: 397

reflexa spec.?) reflexa Wiebes, 1975: 234

vogelii silvestrii3) spec.) heterandromorphum Grandi, 1916: 264

baroni spec.>) heterandromorphum Wiebes, 1974c: 140

nautarum cf. atratus heterandromorphum Wiebes, 1975: 233

1) 8°9, Ethiopia, leg. Friis no. 2263 (RMNH, Leiden, no. 2905).

2) Crossogaster spec, Wiebes, 1975.

3) Host record inferred from the locality data.

4) Newton & Lomo, 1979; the identity has to be checked.

5) Series 2 &, Madagascar, leg. L. Blommers (RMNH, Leiden, no. 2363).

Crossogaster s. str. and Phagoblastus are associated with Agaonidae of the

genus Elisabethiella, as is apparent from the records listed in table 2. It should be

noted that also Philocaenus Grandi may be found in Elisabethiella-figs. The two

aberrant species of Crossogaster seem to be associated with Agaonidae of the

genus Allotriozoon Grandi. The species may be distinguished by the following

key.

Key to the species of Crossogaster and Phagoblastus

(RE pistomalimargmiele finite middle SB REESE 2

ME PISONIANO ERE RS OTO ERE O I 3

2. Labial palpus consisting of two segments, the maxillary of three. Male

unknown. Seychelles and Aldabra, ex Ficusnautarum Bak ............. atrata

— Labial palpus consisting of one segment, the maxillary of two. Senegal, ex

Jes VOGAMIMUG) o's 0269000050000 q000cecccccsaasasanopsooaseas silvestrü

1) Grandi (1928: 204, fig. xlii, 3) corrected Mayr (1885: 189) and stated that the maxillary palpus has

two segments. In my material, from Ficus vasta Forsk. as well as from F. burkei, the basal segment of

the maxillary palpus is short, in some instances merely consisting of a sclerotization of the connective

membrane.

170 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

3. Antenna with one anellus. Female fore tibia with a dorso-apical, bidentate

hook vu. atome niem rain ae 4

— Antenna with two anelli. Female fore tibia with a comb of three to seven

conicaliteethi nach «hth se eee alta III 5

4. Female: antenna with sensilla linearia. Male: the hind tibia without very

Stoutspines Socotra ex FicuswastaForsk 22.2.2 TE triformis

— Female: antenna with long sensilla chaetica. Male: hind tibia rather heavily

armed with dorsal and ventral rows of spines. Rhodesia and Zambia, ex

Ficusburkeby Sees WE OE Sarre EP E odorans

5. Female: mandible smooth, without teeth except at the apex. Male: the

compound eye more than twice as long as the cheek (12 : 5). Gabon, ex

FicusleprieuriMig. ERE ANNEER aoc SERRE liodontus

— Female: mandible with a row of ventral teeth. Male: the compound eye not

quite twice as long as the cheek. South Africa, Rhodesia and Zambia, ex

Ficusburkeiandpetersü.. … a See ans enne see TIE barbarus

Crossogaster odorans Wiebes, spec. nov.

(figs. 16—24, 29—36; pl. 1 figs. 2, 4)

Female. — Length of the head, thorax and gaster 1.3—1.6 mm, the ovipositor

valves ca. 0.15 mm long. Colour dark, especially the head and thorax, the gaster

and the greater part of the legs lighter.

Head (fig. 22) about as long as wide across the compound eyes; the eye almost

half as long as the head (5: 11), slightly longer than the cheek (5 : 4). Three

ocelli. The face has a triangular median impression, with the antennal toruli

about half way between the epistomal margin and the median ocellus. Epistomal

margin weakly convex. Antennal toruli close together. Mandible (fig. 19) with

two apical teeth, two glands, and with a longitudinal row of approximately

fourteen small teeth at the ventral surface. Labium and maxillae (fig. 24): the

palpi unisegmented. Antenna (fig. 29) eleven-segmented, formula 1115(3); the

scape five times as long as wide, 2! times as long as the pedicel; the funicular

segments and the club have long rod-like sensilla in an irregular row of about ten

per segment (in some instances the sensilla are somewhat longer than drawn in

the figure).

Thorax glabrous except for some long setae on the pronotum (which is wider

than long, 5:4), the scutum (with complete parapsidal furrows) and the

scutellum; and with a row of short setae on the mesosternum running from

slightly behind the wing-basis almost to the coxal cavity of the mid leg!). Fore leg

(fig. 17): the tibia has long setae, the armature consists of a dorso-apical

bidentate hook, one ventral cone, and a bifid spur, next to two peg-like spines at

the axial apex; the tarsal segments approximately in ratio 7:2:2:2:6, with

peg-like spines at the axial surface. Mid leg: tarsal ratio 12: 7 : 6 : 5 : 6. Hind

leg (fig. 16): the tibia heavily armed with setae and about eight conical spines

1) I mentioned such a patch of setae for Phagoblastus liodontus Wiebes (1979b: 397, 399); it may be

adaptive to (topocentric) pollination.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 171

along the dorsal margin, one robust cone and a long spur-like spine in the ventral

angle; the tarsal segments approximately in ratio 12 :8:7:6:7, with many

ventral and axial spines. Fore wing (7:3) 1.1 mm long, the submarginal,

marginal, stigmal and postmarginal veins approximately in ratio 16 :7:4:2

ihe

\| LY 27 28 AA

4 I

Figs. 16-24. Crossogaster odorans Wiebes, spec. nov., female holotype. 16, hind tibia and

metatarsus; 17, fore tibia and tarsus; 18, venation of fore wing, detail; 19, mandible, ventral aspect;

20, stigma of eighth urotergite; 21, pygostyle; 22, head; 23, hypopygium; 24, mouthparts. Figs. 25—

28. Phagoblastus barbarus Grandi, female. 25, hypopygium; 26, head; 27, mouthparts; 28, fore tibia

and tarsus.

172 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

(fig. 18), the membrane glabrous in the proximal third, with small microtrichia _ |

distally; hind wing (4 : 1) 0.9 mm long.

Gaster. The posterior edges of the segments crenulate. Hypopygium (fig. 23)

with short setae and small warts along the arms of the V, the spine short. Stigma

(fig. 20) of the eighth urotergite large, ovoid in outline; the pygostyle (fig. 21)

with five long setae.

Male. — Length of the head, thorax and gaster, ca. 1.5 mm. Colour yellowish,

the long setae on the head, thorax and gaster blackish.

Head (fig. 31) about as long as wide across the compound eyes; the eye almost

half as long as the head (5 : 11), distinctly longer than the cheek (5 : 3). Three

ocelli. The antennal toruli situated just below an imaginary line connecting the

lower rims of the eyes. Clypeus slightly impressed, the epistomal margin weakly

convex. Mandible (fig. 36) with two teeth, the subapical of which situated on the

outer side; two glands. Maxillary palpi two-segmented (1:5), labial palpi

consisting of one segment, which is as long as the apical one of the maxillary

palpi. Antenna (fig. 30) eleven-segmented, formula 1115(3); the scape five

times as long as wide, more than twice as long as the pedicel (21 : 9); the club

(segments in ratio 2 : 3 : 4) much wider than the funicle.

Thorax with a rather long pronotum (length: width, 3 : 5) and short mesoscu-

tum (5 : 12), with short parapsidal furrows; the scutellum distinctly transverse

(8 : 5), the propodeum wide (1 : 3). Fore leg (fig. 32): the tibia with two short

dorso-apical teeth, one ventral conical spine and one bifid spur; axially, there are

some slender spines next to the spur; the tarsal segments approximately in ratio

4:1:1:1:2, with peg-like spines on the axial surface. Mid leg: tarsal ratio

12:7:6:5:8. Hind leg (fig. 33): the tibia rather heavily armed with dorsal

and ventral rows of spines and with two long ventral spurs; the tarsal segments

approximately in ratio 15:9:7:5:9, with many setae and spines. Wings

much as in the female, the submarginal, marginal, stigmal and postmarginal

veins of the fore wing (12 : 5, 1.2 mm long) approximately in ratio 16 : 6 : 5 : 2;

the hind wing (5 : 1) 0.9 mm long.

Gaster. The pygostyle (fig. 35) with five setae; the claspers of the genitalia (fig.

34) with three claws.

Type material. — Holotype © (slide-mounted), Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2970). Paratypes: series 2 d and two mating pairs, same data as holotype (coll.

dates ii-iv. 1977); Chishawasha nr. Salisbury, 15 © 106, vii, x1.1974, i, iv,

vi.1975, vii.1976, 1.1977; Makumbi Mission, Chinamora Reserve, ex Ficus

burkei, xii.1976 (A. Watsham). Zambia: Kafue National Park, Ngoma, ex Ficus

burkei, 69, 8.xii.1977 (M. G. Bingham no. 2465); Chipata Golf Course, ex

Ficus burkei, 1 6, 11.1.1979 (M. G. Bingham, no 2474). The material is

deposited in the RMNH, Leiden, some duplicates are in the BMNH, London

and in the Watsham Collection.

Comments. — Compared with Crossogaster triformis Mayr, the new species

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 173

can at once be distinguished by the sensilla of the female antenna. The male of C.

triformis was not fully described and for that reason cannot be compared with the

present sample.

‘PIE

ELS

Figs. 29—36. Crossogaster odorans Wiebes, spec. nov., female holotype (29) and male (30—36). 29,

female antenna, axial aspect; 30, male antenna, axial aspect; 31, head; 32, fore tibia and tarsus; 33,

apex of hind tibia, and metatarsus; 34, armature of genitalia; 35, pygostyle; 36, apex of mandible.

Figs. 37—40. Phagoblastus barbarus Grandi, female (38) and male (37, 39, 40). 37, apex of mandible;

38, female antenna, antiaxial aspect; 39, male antenna, axial aspect; 40, head.

174 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Phagoblastus barbarus Grandi

(figs. 25—28, 37—40)

Phagoblastus barbarus Grandi, 1955: 102—106. Holotype 2, South Africa: Grahamstown (LEB)

[examined].

Female. — Length of the head, thorax and gaster ca. 1.8 mm, the ovipositor

valves ca. 0.2 mm long. Colour much as in Crossogaster odorans.

Head (fig. 26) variable, its length equal to its width in large specimens, up to

distinctly larger than the width (11 : 10) in more slender examples; the eye more

than half as long as the head (13 : 25), almost twice as long as the cheek (13 : 7).

Mandible much as in fig. 19; the two segments of the labial palpus approximately

in ratio 3 : 4, the three of the maxillary palpus 5 : 2 : 4 (fig. 27). Antenna (fig.

38), formula 1125(3): the scape six times as long as wide, 212 times as long as the

pedicel; the funicular segments with few linear sensilla.

Thorax, differential characters with Crossogaster odorans: fore tibia (fig. 28)

with four or five teeth in the dorsal comb (seven in Grandi’s specimens), the

tarsal segments approximately in ratio 6:3:2:2:6. Mid leg: tarsal ratio

4:3:3:2:3. Hind tibia with a row of five or six conical spines along the dorsal

margin and one conical spine next to the long spur at the ventral apex, much as in

Crossogaster odorans (fig. 16); the tarsal ratio approximately 8 : 7 : 5 : 5 : 6.

Gaster. Hypopygium (fig. 25) with a short, rather wide spine. Stigma of the

eighth urotergite approximately one-third of that of Crossogaster odorans; the

pygostyle with four long setae.

Male. — The male was not known to Grandi; it is here compared to that of

Crossogaster odorans, described above. Length of the head, thorax and gaster

ca. 1.7 mm; the colour darker than in Crossogaster odorans. Head (fig. 40) as

long as wide across the compound eyes, which are not quite half as long as the

head (11 : 24) and much longer than the cheek (5 : 3). The antennal toruli are

situated above an imaginary line connecting the lower rims of the eyes; the

epistomal margin is almost straight. Mandible (fig. 37): the subapical tooth

truncate at the tip or even bicuspidate, situated on the inner side of the large

apical tooth; two glands. Maxillary palpus three-segmented (7 : 3 : 6), labial

palpus two-segmented (9 : 11). Antenna (fig. 39), formula 1125(3); the scape

slightly expanded distad, the anelli unequal (2:5), the funicular segments

transverse, with lateral sensilla, the club wider, with more sensilla.

Thorax. The armature of the legs similar to that in Crossogaster odorans, the

tarsal ratios as follows: fore leg, 3:1:1:1:3; mid leg, 10:7:6:4: 8; hind

leo 132.9 FAT ESE 9:

Gaster. Claspers of the genitalia with four or five claws.

Material. — Series 26, Zimbabwe-Rhodesia: Makumbi Mission, Chinamo-

ra Reserve, ex Ficus burkei, xii.1976 (A. Watsham); Salisbury, ex Ficus burkei,

series 23, ii.1977 (A. Watsham). Zambia: Chipata, ex Ficus petersü, S226,

11.1.1979 (M. G. Bingham, no. 2475). The material is deposited in the RMNH,

Leiden, some duplicates are in the BMNH, London and in the Watsham

Collection.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 175

Comments. — Although the two specimens from Grandi’s original series

differ from the present specimens in the number of teeth on the fore tibia (seven

versus four or five), I could not find any other characters warranting a specific

distinction from Phagoblastus barbarus.

PHILOTRYPESINI

The Philotrypesini form a group of fig wasps immediately recognized by the

tubularly lengthened eighth and ninth urotergites of the female and by the

deeply emarginate hypostomal edge of the male. There are some twenty known

species of Philotrypesis Förster (revised by Grandi, 1930), and one of Philotry-

pomorpha Abdurahiman & Joseph (1976) from India. In general, the differ-

ences between the species are rather slight, especially in the female sex, while in

the male sex the polymorphism may make differentiation and identification

difficult. Philotrypesis appears to be associated with species of all sections of

Ficus. There are five species of Philotrypesis known from Africa. The present

species cannot be confused with two of those because they have the sensilla of the

female antenna situated in several rows per segment (P. africana Grandi,

longicornis Grandi). From the other three viz., P. erythraea Grandi, finitimo-

rum Wiebes and selenetica Grandi, it can be distinguished by the characters used

in the following key.

Key to some African species of Philotrypesis

1. Female: setae on the antennal funicle dispersed over the segment. Male:

hind basitarsus 3—4 times as long as the second segment ................. 2

— Female: setae on the antennal funicle situated at the base of the segment.

Male: hind basitarsus not quite twice as long as the second segment ........ 3

2. Female: funicular segments of the antenna relatively short, e.g., the first

only slightly longer than wide. Hind tibia with four conical spines in the

dorso-apicalcorner. Male unknown. Guinea, ex Ficus vogeliiMiq.... selenitica

— Female: funicular segments of the antenna longer, e.g., the first 12 times as

long as wide. Hind tibia with two conical spines in the dorso-apical corner.

Sierra Leone, ex Ficus cf. eriobotryoidesK.&B. ............... finitimorum

3. Female: the stigmal vein half as long as the postmarginal. Male: fore tibia

with two stout spines proximad of the ventral spur; mid tibia with conical

spines along the dorsal margin; hind femur without apical spines, the tibia

without stout spines at one-third ofitslength. Eritrea, ex Ficusspec. .. erythraea

— Female: the stigmal vein one-third of the length of the postmarginal. Male:

fore tibia with three stout spines proximad of the ventral spur; mid tibia

without a row of dorsal conical spines; hind femur with two spines at the

dorsal apex, the tibia with two stout spines at one-third of its length (fig. 47).

Rhodesia and Zambia, ex Ficus burkei parca

176 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

Philotrypesis parca Wiebes, spec. nov.

(figs. 41—57; pl. 2 figs. 1, 2, pl. 3fig. 1)

Female. — Length of head, thorax and apparent gaster 1.8 mm, the tubular

segments and the ovipositor valves ca. 3 mm. Colour yellowish, with black marks

on the scutellum and propodeum, and on the dorsum of the gastral tergites.

Head (fig. 42) distinctly shorter than wide across the compound eyes (3 : 4);

the compound eye almost two-thirds of the length of the head, 112 times as long

as the cheek. Mandible (fig. 44) with two apical teeth, one of which is truncate;

| two glands; the labial palpus consisting of two segments (2 : 3), the maxillary

(fig. 48) of four (4: 6: 3 : 11). Antenna (fig. 41) consisting of thirteen seg-

ments, formula 1135(3); the scape is three times as long as the pedicel; the

funicular segments about as long as wide, each has one rather regular row of long

sensilla and long, basal setae. |

Thorax. The tibial armature of the fore leg (figs. 45, 46) consists of three

conical antiaxial spines, viz., one ventral and two dorsals, and two slender axial

spines next to the ventral spur; the tarsal segments are approximately in ratio

6:3:3:2:6. Mid leg: tarsal ratio 8: 4 : 3 : 2 : 3. Hind leg: the tibial arma- _

ture (fig. 47) consists of two unequal ventral spurs accompanied by two slender

antiaxial spines and a row of axials, and four to six spines at the antiaxial dorsal

apex; the tarsal segments are approximately in ratio 11 : 7 : 4 : 3 : 4. Fore wing

(5 : 2) 1.3 mm long, the submarginal, marginal, stigmal and postmarginal veins

approximately in ratio 6 : 4 : 1 : 3; the hind wing (4 : 1) 0.8 mm long.

Gaster. The hypopygium (fig. 43) has a short, wide spine. The eighth uroter-

gite is only slightly shorter than the combined lengths of the proximal segments

(20 : 23), the ninth is not quite half as long as the eighth (9 : 20). The ovipositor

and the valves are distinctly more than twice as long as the eighth and ninth

urotergites combined (22 : 9).

Male. — Length of head and thorax 0.8—0.9 mm. Colour yellowish. All

specimens studied are eumegetic, acanthocephalous, macrognathous (triodon-

tous) and apterous.

Head (fig. 50) about as long as wide across the compound eyes; the compound

eye over one-fifth of the length of the head (2 : 9), longer than the cheek (5 : 3).

The head bears long dorsal setae, but no ventrals. The antennal toruli are

situated in a shallow depression above the epistomal edge. Mandible falcate,

with two glands (fig. 49, also showing the hypostomal edge, so characteristic for

all species of Philotrypesis), the labial palpus consisting of two segments

(11 : 10), the maxillary of four (2:2:1:3). Antenna (fig. 51) eleven-seg-

mented, formula 1115(3); the scape is less than five times as long as the pedicel

(23 : 5); the five funicular segments are gradually widening distad, the second to

fourth have apical sensilla.

Thorax with the pronotum much longer than the other segments combined

(5 : 3), with long setae in the anterior and posterior corners; the mesonotum is

(in the middle) more than twice as long as the metanotum and the propodeum

taken together. Fore leg: the tibial armature (figs. 53, 56) consists of three stout

axial spines, proximad of the ventral spur and one antiaxial, one robust spine at

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 177

the axial and antiaxial apices, and one in the dorso-apical angle accompanied by

four conical spines; the tarsal segments are approximately in ratio

Figs. 41—52. Philotrypesis parca Wiebes, spec. nov., female (41—48) and male holotype (49—52).

41, female antenna, axial aspect; 42, head; 43, apex of hypopygium; 44, clypeus and left mandible,

dorsal aspect; 45, apex of fore tibia, and tarsus, axial aspect; 46, do., detail in antiaxial view; 47, apex

of hind tibia; 48, mouthparts, palpi; 49, hypostomal margin and right mandible, ventral aspect; 50,

head and thorax; 51, male antenna, axial aspect; 52, genitalia.

178 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 53—57. Philotrypesis parca Wiebes, spec. nov., male holotype. 53, apex of fore tibia, and

tarsus, axial aspect; 54, apex of mid tibia, and tarsus; 55, hind tibia, axial aspect; 56, apex of fore tibia,

and part of tarsus, antiaxial aspect; 57, apex of hind tibia, and tarsus (long setae not fully drawn).

Figs. 58—60. Sycoscapter cornutus Wiebes, spec. nov, male holotype. 58, fore tibia and tarsus; 59,

mid tibia, and part of tarsus; 60, apex of hind tibia, and tarsus.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 179

3:1:1:1:15. Mid leg: the tibia (fig. 54) has a small dorsal cone and two more

slender ventral spines at the apex (one axial and one antiaxial), the ventral

margin has six large spines, the dorsal margin is without the row of conical spines

described and figured for all forms of P. erythraea; the tarsal segments are

approximately in ratio 11 : 4 : 5 : 4 : 24. Hind leg (figs. 55, 57): the femur have

two spines, one behind the other, at the dorsal apex; the tibial armature consists

of two spines (situated just beyond one-third of the length of the tibia) that are

not apparent in P. erythraea; some more spines, most of which small and conical,

occur along the dorsal margin and at the antiaxial apex; two longer, slender

spines accompany the unequal ventral spurs, and three larger spines are situated

along the distal third of the ventral margin; the tarsal segments (measured along

the dorsal margin) approximately in ratio 8:5:1:1:7, the proximal two

expanded dorsad, with long setae.

Gaster. The claspers of the genitalia (fig. 52) with four claws.

Type material. — Holotype ® (slide-mounted) Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2964). Paratypes: series 9 5 d, same data as holotype; Chishawasha nr.

Salisbury, 5 2 5 &, 1976, vi.1977. Zambia: Chipata Golf Course, ex Ficus

burkei, 32 4 3, 11.1.1979 (M. G. Bingham, no. 2474). The material is deposited

in the RMNH, Leiden, some duplicates are in the BMNH, London and in the

Watsham Collection.

Comments. — The present material differs only slightly from Grandi’s

Philotrypesis erythraea, just enough — although originally with some doubt — to

consider them separate taxa. A similar situation exists with Elisabethiella

stuckenbergi and socotrensis (discussed above) and Otitesella tsamvi and epica-

rioides (compared below). For the alate male (?), see below, under Watshamiel-

la alata.

SYCORYCTINI

A key to the Indo-Australian genera of this group was published by Wiebes

(1966c: 173). Revisionary notes on some were given by Wiebes (1967: 428,

Sycoscapteridea Ashmead; 1978b: 185—186, Sycoscapter Saunders in West-

wood). Abdurahiman & Joseph (1975a: 99) added a new genus Sycorycteridea,

which they compared with Sycoryctes Mayr. As a group, the Sycoryctini are

recognizable in the female sex by the tubularly lengthened ninth urotergite,

which makes the impression that the apparent gaster ends with the eighth

segment; the males are mostly apterous, subapterous in some species and, as will

be apparent from one of the descriptions below, alate in others. The apterous

males can be distinguished at a glance from those of Philotrypesis by the straight

or almost straight hypostomal edge.

As many samples of fig insects bred from one fig contain several species and

genera of Sycoryctini (up to four were recorded by Wiebes, 1966), it is not always

easy to correlate the sexes. For the present study mating pairs were available of

all species, so as to allow of a certain correlation of males with their females. In

180 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

general, the generic classification of the Sycoryctini is still unsatisfactory and this

is why I here confine myself to the differentiation of the genera treated in the

general key (p. 159).

Sycoscapter Saunders in Westwood

A survey of the species now recognized in the genus Sycoscapter was given

with a recent redescription of its type-species (Wiebes, 1978): the African

species are S. gibbus Saunders and S. montis Wiebes. The new species now

added may be differentiated from its African congeners by the following key.

The male sexes are recognizable as a group by the ventral spur of the fore tibia

being bifid, instead of simple as in the Indo-Malayan species.

Key to the African species of Sycoscapter

1. Female: the fore wing with about fifteen large setae in the marginal-stigmal

angle, some ten along the stigmal vein and a few in the wing-apex; the ninth

urotergite about six times as long as the apparent gaster. Male: all tarsi

consisting of four segments; wing remnant present. Sierra Leone, ex Ficus

chseriobotryoidesK-& Bi Lu ut ans LO EER montis

— Female: the fore wing with thirty or more large setae in the marginal-stigmal

angle, and 15—20 along the stigmal vein; the ninth urotergite two to three

times as long as the apparent gaster. Male: all tarsi consisting of five

segments; No:wingremnant Vi Cor Er o ee - noe eee 2

2. Female: the fore wing with many setae in the apical part, so as to make the

surface distinctly hirsute. Male: the epistomal margin almost straight or with

two slightly protruding bulges. Madagascar, ex Ficus soroceoides Baker and

Réunion, ex Ficuslaterifolia Vahl... e E .. gibbus

— Female: the fore wing with only a few large setae in the apical part, otherwise

glabrous. Male: the epistomal margin distinctly produced into two rather

acute lobes. Rhodesia and Zambia, ex Ficus burkeiandpetersii ...... cornutus

Sycoscapter cornutus Wiebes, spec. nov.

(figs. 58—60, 64—66, 75—76, 83; pl. 2 fig. 3)

Female. — Length of head, thorax and apparent gaster ca. 1.4 mm, the ninth

tergite (the “tail”) ca. 2 mm, i.e., 2/2—3 times as long as the apparent gaster.

Colour blackish bronze, the basal segments of the legs up to the tibiae brown,

sometimes also the dorsal part of the tibiae dark, mostly the tibiae and tarsi

yellow.

Head (fig. 65) shorter than wide across the compound eyes (9 : 11), the eye

half as long as the head and as long as the cheek. Antennal toruli slightly below

the middle of the face; deep scrobes run from the toruli to the median ocellus.

Epistomal margin produced into an acute median projection. Mandible with two

rather robust apical teeth; two glands; the labial palpus consisting of two

segments (4:3), the maxillary of four (6: 7:5: 13). Antenna (fig. 66),

formula 1125(3); the scape 212 times as long as the pedicel; the funicular

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 181

segments with one row of long sensilla, and with a basal row of long setae.

Thorax. Fore leg with two dorsal cones and one antiaxial spine on the tibial

apex, and with a bifid ventral spur accompanied by a slender spine; the tarsal

ratio approximately 5 : 3 : 2: 2: 6. Mid leg: tarsal ratio 9:5:4:3:5. Hind

leg: the tibia with about six to nine small conical spines along the distal two-thirds

of the dorsal margin, and one or two short spines in the ventral angle, next to the

two spurs; the tarsal segments approximately in ratio 13 : 10:5:4:7. Fore

wing (2 : 1) 1.2 mm long, the submarginal, marginal, stigmal and postmarginal

veins approximately in ratio 5 : 4 : 3 : 6, the stigma (fig. 64) boot-shaped; ca. 35

long setae occur in the marginal-stigmal angle, some fifteen on or along the

stigmal vein, and a few in the part distad of this vein; hind wing (4 : 1) 0.8 mm

long.

Gaster. Hypopygium (fig. 83) with a short and wide spine, and one pair of long

setae.

IG DI

— =

e SG — y, 4

« —

ug e 2

El A

Fig. 61—69. Details of some female Sycoryctini: 61—63, Watshamiella alata Wiebes, gen. & spec.

nov.; 64—66, Sycoscapter cornutus Wiebes, spec. nov.; 67—69, Sycoryctes remus Wiebes, spec. nov.

61, 64, 67, veins of fore wing; 62, 65, 68, head; 63, 66, 69, scape, anelli and first funicular segment of

antenna.

182 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Male. — Length of head and thorax ca. 1.0 mm. Colour and general habitus as

in pl. 2 fig. 4, but the epistomal edge with more distinct horn-like projections (fig.

75) and the tarsal segments much shorter, especially those of the fore leg (fig.

58).

Head (fig. 75) longer than wide across the compound eyes (10 :9); the

compound eye one-quarter of the lengt of the head, about as long as the cheek.

The toruli are close together at the epistomal margin, which bears two horn-like

projections with a seta on top. Mandible robust, with a serrate molar edge and

with two glands; the labial palpus consisting of two segments (2 :1), the

maxillary of four (5 : 6 : 2 : 4). Antenna (fig. 76) eleven-segmented, formula

1115(3); the first and third funicular segments larger than the others, with an

apical sensillum (axial in the one, antiaxial in the other); the club segments

unequal (3 : 1 : 1), with sensilla on the apical and subapical.

Thorax (fig. 75): the pronotum rather short, 1% times as wide as long in

flattened position, also much wider than the mesonotum (10 :7) and the

posterior tergite (10 : 9) representing the metanotum (the lateral lobes) and the

propodeum. All legs bear relatively few tibial spines. The fore coxa has an

antiaxial, hyaline edge; except for one long axial, all spines of the fore tibia are

fully visible in antiaxial aspect (fig. 58); the ventral spur is rather long and bifid;

the five tarsal segments are approximately in length ratio 5:1:1:1:17. The

mid tibia (fig. 59) is more robust than those of the other Sycoryctini treated here;

it bears pairs of ventral spines in the distal half, one simple spur that is as long as

the first three tarsi combined, one apical on either side, and up to ten dorsals in

the distal half; the tarsal ratio is 3 : 2 : 2 : 2 : 10. The hind tibia (fig. 60) has

small spines along two-thirds of the dorsal margin and four longer spines near the

ventral apex next to the spur, which is as long as the basitarsus; the tarsal ratio is

3 les 21 elen

Gaster: the claspers of the genitalia with three or four claws.

Type material. — Holotype d (slide-mounted), Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, 11.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2968). Paratypes: series 2d and 13 mating pairs, same data as holotype (coll.

dates iii-v.1977); Chishawasha nr. Salisbury, 22 ® 28 6, vii.1974, v.1975, 1976,

i, vi.1977; Makumbi Mission, Chinamora Reserve, ex Ficus burkei, series 9 9

d, xii.1976 (A. Watsham). Zambia: Kafue National Park, Ngoma, ex Ficus

burkei, 4 3, 8.xii.1977 (M. G. Bingham, no. 2465); Chipata Golf Course, ex

Ficus burkei, 1 3, 11.1.1979 (M. G. Bingham, no. 2474); Chipata, ex Ficus

petersii, 2 2, 11.1.1979 (M. G. Bingham, no. 2475); Lusaka, Cathedral of the

Holy Cross, ex Ficus burkei, 19 © series d, 16.11.1979 (M. G. Bingham, no.

2490). The material is deposited in the RMNH, Leiden, some paratypes in

BMNH, London and the Watsham Collection.

Sycoryctes Mayr

The type-species of Sycoryctes, S. patellaris Mayr, originates from Java. It

seems that most Indo-Malayan species of the genus are distinct in having long

setae on the male hind basitarsus (Mayr, 1885, pl. 13 fig. 30) instead of shorter

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 183

setae as in the African species (e.g., fig. 78). Many of the African species are

insufficiently known; the new species described below are compared with the

following: S. caelebs Wiebes, coccothraustes Mayr, lomaensis Wiebes, sjoestedti

Mayr, truncatus Mayr. Two species are known in the female sex only, viz., S.

dagatiguyi (Risbec), which has a longer postmarginal vein than the specimens

from Ficus burkei (where the postmarginal is scarcely longer than the stigmal);

and S. sebertianus Masi, which seems to be rather larger (6.6 mm versus 5 mm,

including the ‘tail’’). In general, the females are difficult to differentiate, as in

most cases the descriptions are very short and mostly refer to colour characters.

In the present paper, too, the females of the two new species are distinct mainly

in colour. The males can be distinguished by the characters used in the following

key. Much as the present novelties appear well characterized, the differentiation

of some of the others is still unsatisfactory.

Key to the males of African Sycoryctes

ieee llead:and thorax hirsute dorsallys. 34.52 Reel SECTOR ee 2

— Head and thorax with some long setae, but not hirsute ................... 3

2. Spurofthehindtibiaaslongasthebasitarsus. Cameroons, ex Ficusspec.. .......

DEE N AE EN N RUE ATE EB. eh Er mA sjoestedti

— Spur of the hind tibia reaching to the third tarsal segment. Rhodesia, ex

CUS DUR CL EL IIS LIE TREE SUSA NE RE IA ATAR hirtus

3. Spur of the hind tibia long, reaching up to the fourth or fifth segment ...... 4

— Spur ofthe hind tibia much shorter, reaching to the third segment ......... 5

4. Eye small, one-third of the cheek. Basitarsus of the hind leg not expanded.

SocotrasexhicusvastaN allen SSR a: coccothraustes

— Eye almost as long as the cheek. Basitarsus of the hind leg in most instances

distinctly expanded. Rhodesia and Zambia, ex Ficus burkei and petersii. . . remus

5. Head distinctly longer than wide (3 : 2), the lateral margins straight. Sierra

BeoneneskicuschsenobotnyvoiderksCiB E EROE lomaensis

Head transversenthe lateral Martins CUVE NP ORE EEE 6

6. Antennawithone anellus. Socotra, ex Ficus vasta Vahl ............ truncatus

— Antennawithtwoanelli. Aldabra, ex Ficusavi-aviBl................. caelebs

Sycoryctes remus Wiebes, spec. nov.

(figs. 67—69, 72—74, 77—79, 84; pl. 2 fig. 4)

Female. — Length of head, thorax and apparent gaster 1.3—1.6 mm, the

ninth tergite ca. 2 mm, i.e., 2/2 times as long as the apparent gaster. Blackish,

also the legs up to and including the tibiae dark; the wings hyaline.

Head (fig. 68) distinctly shorter than wide across the compound eyes (23 : 28),

the eye more than half as long as the head (12 : 23) and about twice as long as the

cheek. Antennal toruli situated in the lower third of the face, at a distance from

the epistomal margin equal to their longitudinal diameter; shallow scrobes run to

the median ocellus. Epistomal margin with three wide lobes. Mandible biden-

tate; two glands; the labial palpus consisting of two segments (4:5), the

184 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 70—71. Sycoryctes hirtus Wiebes, spec. nov., male holotype. 70, head and thorax; 71, antenna,

axial aspect. Figs. 72—74. Sycoryctes remus Wiebes, spec. nov., male holotype. 72, head; 73, lateral

lobe of mesothorax, and wing remnant; 74, antenna, axial aspect. Figs. 75—76. Sycoscapter cornutus

Wiebes, spec. nov., male holotype. 75, head and thorax; 76, antenna, axial aspect.

maxillary of four (2: 4: 1:5). Antenna (fig. 69), formula 1125(3); the scape

three times as long as the pedicel; the funicular segments with one row of long

sensilla, and with long basal and shorter apical setae.

Thorax. Fore leg: the tibial armature consisting of two dorso-apical spines and

one subapical; ventrally, there is the bifid spur and some accompanying spines,

viz., one distal of the spur and two, more slender, proximad; the tarsal segments

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 185

approximately in ratio 16 : 8 : 8 : 5 : 12. Mid leg: tarsal ratio 19:9:6:4:7.

The tibial armature of the hind leg consisting of two unequal spurs and some

stout spines along the distal half of the ventral margin: typical is a number of

three dark spines and some lighter more proximally; the distal third of the dorsal

margin has four or five conical spines and two apicals; the tarsal segments

approximately in ratio 22: 10 : 7 : 4 : 8. Fore wing (5 : 2) 1.25 mm long, the

submarginal, marginal, stigmal and postmarginal veins approximately in ratio

35: 18:9: 11 (fig. 67), the wing hyaline but for some setae along the veins;

hind wing (4 : 1) 0.9 mm long.

Gaster. Hypopygium (fig. 84) broadly rounded, with two setae of medium size

at the apex; a patch of two or three setae occur on either side, where the arms of

the V appear broken.

Male. — Length of head and thorax 0.7—0.9 mm. General habitus pl. 2 fig. 4;

in most specimens, the basitarsus of the hind leg is more distinctly enlarged.

Head (fig. 72) quadrate or slightly transverse, more elongate in small speci-

mens; almost similar in shape to that of Sycoscapter cornutus, but the epistomal

margin less distinctly cornute or even almost straight in the middle; the posterior

edge of the head rather sharp, laterally produced into acute angles. Also

mandible similar; the segments of the labial palpus approximately in ratio 3 : 2,

those of the maxillary palpus 7 : 14 : 4 : 9. Antenna (fig. 74), formula 1125(3).

Thorax much as in Sycoscapter cornutus, but with a distinct wing remnant (fig.

73). The fore leg (fig. 79) resembles that of Sycoryctes hirtus, to be described

hereafter, but the dorsal and ventral spines are less in number and there are two

apical spines instead of one; the tarsus is more compact, its ratio 5: 3: 2:2:6.

The mid leg is long and slender; the tibia (fig. 77) has many spines along the

dorsal and ventral edges and a long ventral spur reaching to the third tarsal

segment; the tarsal ratio approximately 8 : 4 : 4 : 3 : 9. Hind leg (fig. 78) much

like that of S. hirtus, but two ventral spines are sometimes situated on a small

tibial protuberance, so as to make them very conspicuous, the tibial spur reaches

to the fourth tarsal segment; the basitarsus distinctly expanded, the ratio

13:24:27 PRES PES

Gaster: the claspers of the genitalia wide, with three to five claws.

Type material. — Holotype d (slide-mounted). Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in the RMNH, Leiden

(no. 3713). Paratypes: series ? d and eight mating pairs, same data as holotype

(coll. dates ii-v.1977); Chishawasha nr. Salisbury, 12 2 10 d, 1976, vi.1977;

Makumbi Mission, Chinamora Reserve, ex Ficus burkei, series 2 15 d, xii.1976

(A. Watsham). Zambia: Kafue National Park, Ngoma, ex Ficus burkei, 1 2%

3, 8.xii.1977 (M. G. Bingham, no. 2964); Chipata Golf Course, ex Ficus burkei,

5 © 3 6, 11.1.1979 (M. G. Bingham, no. 2474); Chipata, ex Ficus petersü, 2 6,

11.1.1979 (M. G. Bingham, no. 2475); Luapula Prov., Mumunshi Banana

Estate, ex Ficus cf. burkei, 1 2, 9.xii.1978 (M. G. Bingham, no. 2471); Lusaka,

Cathedral of the Holy Cross, ex Ficus burkei, 30 2 24 6, 16.11.1979 (M. G.

Bingham, no. 2490). The material is deposited in the RMNH, Leiden, some

paratypes are in the BMNH, London and in the Watsham Collection.

186 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

Sycoryctes hirtus Wiebes, spec. nov.

(figs. 70, 71, 80, 81)

Female. — Length of head, thorax and apparent gaster ca. 1.5 mm, the ninth

tergite ca. 2.5 mm long, i.e., three times as long as the apparent gaster. Blackish

as in S. remus, but more dull, the legs from the femora onwards yellowish. In all

morphological characters this species very much resembles Sycoryctes remus,

but the number of ventral spines on the hind tibia tends to be larger (up to six) and

there usually also are more dorsal spines (six to eight). In some of the samples I

could not identify all female specimens with any certainty.

Male. — Length of head and thorax 0.9—1.0 mm. General habitus as in pl. 2

fig. 4, but the head and thorax hirsute dorsally, the antenna slender with a

brownish apical knob, and the head with a cervical sclerite (fig. 70); the

basitarsus of the hind leg is not expanded.

Head (fig. 70): shape and proportions much as in Sycoscapter cornutus, but the

epistomal margin straight; the pubescence of the dorsal surface especially

distinct in lateral aspect, as a short fur in between the longer setae — the same

applies to the thorax. The posterior part of the head is raised in the middle, so as

to form a small cervical shield; the lateral angles of the head are slightly

produced. The molar teeth of the mandible stout; the segments of the labial

palpus approximately in ratio 1 : 1, those of the maxillary palpus 7:10:3:5.

Antenna (fig. 71), formula 1125(3).

Thorax, fig. 70. Compared to Sycoscapter cornutus, the fore tibia (fig. 80)

more slender, with a series of five ventral spines proximad of the bifid spur, one

apical spine visible in antiaxial aspect, and three dorsals in the apical third,

inserted more axially; the tarsal ratio approximately 6 : 6 : 3 : 2 : 8. Hind tibia

(fig. 81): long dorsal spines among a row of cones in the distal two-thirds;

ventrally, there are three or four conspicuous spines in the distal third and one

long spur, reaching to the third tarsal segment; the tarsal ratio approximately

Sr ZE

Gaster: the claspers of the genitalia with four claws.

Type material. — Holotype dg (slide-mounted), Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2967). Paratypes: 9 2 76 and seven mating pairs, same data as holotype (coll.

dates iii-iv.1977); Chishawasha nr. Salisbury, 10 2 4 d, vii.1974, 1976, vi.1977;

Makumbi Mission, Chinamora Reserve, ex Ficus burkei, 2 2 3 6, xii.1976 (A.

Watsham). Zambia: Lusaka, Catheral of the Holy Cross, ex Ficus burkei, 9 9

13 &, 16.11.1979 (M. G. Bingham, no. 2490). The material is deposited in the

RMNH, Leiden, some paratypes in the BMNH, London and in the Watsham

Collection.

Watshamiella Wiebes, gen. nov.

Type-species: Watshamiella alata Wiebes, spec. nov.

Sycoryctini with alate male. Toruli of the antennae widely spaced, situated

: Fig wasp fauna 187

Z. BOUCEK, A. WATSHAM & J.T. WIEBES

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188 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

rather close to the epistomal margin, which is straight. Mandible bidentate (most

males) or tridentate (females). Antenna with three anelli. Pronotum long;

parapsidal furrows complete or almost so. Front femur inflated. Hind basitarsus

at least as long as the second and third segments combined. Wings hirsute; the

postmarginal vein 2—3 times as long as the stigmal. Gaster shortly petiolate; the

ninth urotergite of the female with lateral rows of setae, and with small pygo-

styles; also the male ninth urotergites with pygostyles.

Comments. — The correlation of the female of W. alata with an alate male

made me reconsider the generic identification of such males, up to now classified

with Philotrypesis Förster, viz., P. anguliceps (Westwood), finitimorum Wiebes

and longicornis Grandi. The first mentioned appears to be congeneric with the

one here treated and thus should be allocated to Watshamiella. P. finitimorum

has short wing remnants and, for the time being, there is no reason to alter its

generic status; the wings in P. longicornis are longer, but this male, which I did

not study myself, is so similar to its female that I prefer to leave it in Philotrypesis.

Other species to be compared were up to now classified with Sycoscapteridea

Ashmead, viz., the female identified with S. monilifera by Wiebes (1967) and S.

stilifera Wiebes (1966). Both are now allocated to Watshamiella.

It should be clearly stated that in classifying Watshamiella with the Sycorycti-

ni, much weight is given to the structure of the ovipositing organs (Wiebes,

1966a), while the resemblance to the Philotrypesini is neglected e.g., in the three

antennal anelli, the situation and number of spines on the legs, the long

postmarginal vein of the fore wing. The Watshamiella-males, known from their

association with the females with which they were found in copula, are very

similar to a type of male that I know from the Indo-Malayan region and of which

we now also have at least one from Ficus thonningii. AW pictured it (pl. III fig.

1), but JTW could not yet study it in any detail: it has a bare strip along the

marginal vein, just as Philotrypesis-females usually have. Probably, this is an

alate male of Philotrypesis!

The reclassification of the species mentioned makes necessary some nomen-

clatorial alterations, as follows:

Watshamiella alata Wiebes, spec. nov., described below.

Watshamiella infida Wiebes, spec. nov., description in Wiebes, 1967: 426, figs. 77—84 (Sycoscapteri-

dea monilifera (Westwood), female), 433—434, figs. 102—108 (Philotrypesis anguliceps (West-

wood), alate male).

Watshamiella stilifera (Wiebes, 1966c), nov. comb.

These species may be recognized by using the following key.

Key to the species of Watshamiella

1. Funicular segments of the antenna distinctly longer than wide, with about

two (sometimes irregular) rows of sensilla. Male unknown. North Borneo,

ex Ficus stupenda MUG sn SAI stilifera

— Funicular segments of the antenna not longer than wide, with one row of

sensilla. Female: ninth urotergite three times as long as the apparent gaster.

Male: hind basitarsus longer than the three following segments combined... 2

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 189

2. Female: the head distinctly transverse (16 : 19); the eye not quite 1/2 times

as long as the cheek (11 : 8). Male: the compound eye distinctly longer than

thecheek (= 2)A@eylonsexkieusreligiosaE nr re infida

— Female: the head almost as long as wide (23 : 25); the eye twice as long as the

cheek. Male: the compound eye as long as the cheek. Rhodesia, ex Ficus

Dürkeise ES BEER. EI EED OE alata

Watshamiella alata Wiebes, spec. nov.

(figs. 61—63, 82, 85—92; pl. 2 figs. 5, 6)

Female. — Length of head, thorax and apparent gaster ca. 2 mm, the ninth

urotergite ca. 3 mm i.e., three times as long as the apparent gaster. Colour

yellow-brown, with black markings on the dorsum of the gaster.

Head (fig. 62) almost as long as wide across the compound eyes (23 : 25), the

eye more than half as long as the head (14 : 23) and twice as long as the check.

Antennal toruli widely spaced, situated rather low in the face, close to the

epistomal margin; shallow scrobes run to about the middle of the face. Epistomal

margin straight. Mandible tridentate, two glands; the labial palpus consisting of

two segments (3 : 7), the maxillary of four (3:2:2:5). Antenna (fig. 63),

formula 1135(3); the scape three times as long as the pedicel; the funicular

segments with one row of sensilla, and with scattered setae.

Thorax. Pronotum long, about of the same length as the scutellum (which

bears two long, posterior setae); the parapsidal furrows complete. Fore leg: the

femur inflated, the tibial armature consisting of two dorso-apical spines and a

bifid ventral spur, accompanied by some spines; the tarsal segments approx-

imately in ratio 12:4:4:3: 14. Mid leg: tarsal ratio 17:9:6:5:7. Hind

leg: the tibia (fig. 89) with a rather straight apical edge, the armature consisting of

stout spines next to the ventral spurs and six conical spines along the distal fifth of

the dorsal margin; the tarsal ratio 8 : 5 : 3 : 2 : 3, i.e., the basitarsus as long as

the combined lengths of the second and third segments. Fore wing (5 : 2) 1.6 mm

long, the submarginal, marginal, stigmal and postmarginal veins approximately

in ratio 15: 10 : 3 : 9 (fig. 61), the surface hirsute; hind wing (4 : 1) 1.1 mm

long.

Gaster shortly petiolate. Hypopygium (fig. 82) with an acute spine and three

pairs of long setae. The ninth urotergite with small pygostyles.

Male. — Length of head, thorax and gaster ca. 1.7 mm. Colour yellow-brown.

Head (fig. 87) almost as long as wide across the compound eyes (13 : 14), the

eye as long as the cheek. Three ocelli. Antennal toruli widely spaced, situated

close to the epistomal margin, which is straight. Mandible bidentate, with two

glands, but the inner tooth truncate and divided at the apex in some specimens;

the labial palpus consisting of two segments (1:2), the maxillary of four

(8 : 8 : 7 : 14). Antenna (fig. 85), formula 1135(3); the scape 212 times as long

as the pedicel; the funicular segments with some large sensilla and mainly apical

setae.

Thorax. Pronotum almost as long as the mesonotum; the parapsidal furrows

almost complete. Fore leg (figs. 86, 91—92): the femur inflated, the tibial

190 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 85—92. Watshamiella alata Wiebes, gen. & spec. nov., male (85—88, 90—92) and female (89).

85, right antenna, antiaxial aspect; 86, right fore leg; 87, head; 88, apex of hind tibia, and basitarsus;

89, apex of female hind tibia, axial aspect; 90, apex of gaster, with genitalia; 91, apex of fore tibia, and

two tarsal segments; 92, do., detail in axial view.

armature much as in the female, the tarsal segments approximately in ratio

9:3:3:2:7. Mid leg: tarsal ratio 6:3:3:2:6. Hind leg (fig. 88) with

seven conical spines along the distal third of the dorsal tibial margin and some

ventral spines next to the spurs; the tarsal segments approximately in ratio

23 :9:6:3: 11. Fore wing (5 : 2) 1.3 mm long, the submarginal, marginal,

stigmal and postmarginal veins approximately in ratio 26 :23:9:18, the

membrane hirsute; hind wing (5 : 1) 0.9 mm long.

Gaster shortly petiolate. Genitalia (fig. 90) with four claws on the claspers;

pygostyles present.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 191

Type material. — Holotype © (slide-mounted), Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2966). Paratypes: series 2 and four mating pairs, same data as holotype (coll.

dates ii-v.1977); Chishawasha nr. Salisbury, 5 d, 1976 & vi.1977; Makumbi

Mission, Chinamora Reserve, ex Ficus burkei, 5 2 3 &, xii. 1976 (A. Watsham).

The material is deposited in the RMNH, Leiden, some paratypes in the BMNH,

London and in the Watsham Collection.

OTITESELLINAE

The Otitesellinae held my attention for some time (Wiebes, 1967: 435—441,

1969: 443-444, 1974: 145-161). They were formerly classified with the Tory-

midae, but now are transferred to the Pteromalidae. The females have a short

ovipositor and for that reason were supposed to oviposit from within the

receptacle; some have special features (rasps, stout spines) evidently related to

the way of entering through the ostiole, but most have not. Otitesella Westwood,

to which the species from Ficus burkei belongs, does not enter the receptacle for

oviposition. The males are peculiar by their oversized head and mandibles. The

main character by which the species of Otitesella and its close relatives can be

distinguished, is found in the male thorax (fig. 94), the terga of which are all free;

the female wings are almost hyaline. Immediate African relatives of the species

here treated are Otitesella africana Grandi, epicarioides Grandi and royi

Wiebes, with which the species from Ficus burkei is being compared in the

following key.

Key to some African species.of Otitesella

1. Female: epistomal margin with a median lobe in between the lateral lobes.

Male: head with a large anterior concavity reaching the antennal toruli.

ERICA ENIRICUSISDEC MR IT III OI epicarioides

— Female: epistomal margin with a median indentation in between the lateral

lobes. Male: head convex dorsally, only faintly impressed in front ........ 2

2. Female: the compound eye four times as long as the cheek. Male: toruli of

the antennae distinctly situated between the eyes. Guinea, ex Ficus vogelii

MINI AN DER EAN RE SEE DORE RR EN AN: africana

— Female: the compound eye about twice as long as the cheek. Male: toruli of

the antennae almost totally behind the posterior edge of the eyes .......... 3

3. Female: the antennal anelli long, the distal one half as long as wide. Male:

antenna with two distinct anelli. Guinea, ex Ficus cf. eriobotryoides K. & B. …

EI Ath RA ra A ANALI Eh RIO I LA O i PRET royi

— Female: the antennal anelli (fig. 101) shorter, disk-like, up to five times as

wide as long. Male: antenna with only one anellus. Rhodesia and Zambia, ex

KICUSIDUIKCIOR PASO tsamvi

Otitesella tsamvi Wiebes, spec. nov.

(figs. 93—108; pl. 1 figs. 5, 6)

Female. — Length 1.4—1.7 mm. Colour blackish, the tibiae and tarsi of the

legs lighter.

192 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 93—108. Otitesella tsamvi Wiebes, spec. nov., male holotype (93—98) and female (99—108).

93, apex of antennal scape, pedicel and three flagellar segments; 94, head and thorax; 95, apex of fore

tibia, and tarsus, axial aspect; 96-97, antennal scapes of a large- and small-sized specimen,

respectively; 98, apex of male fore tibia; 99, apex of female fore tibia, and basitarsus; 100, right

mandible, ventral aspect; 101, basal segments of antenna, axial aspect; 102, outline of head; 103,

epistomal margin; 104, apex of hind tibia, and basitarsus; 105, maxillary palpus; 106, labial palpus;

107, part of wing-venation; 108, hypopygium.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 193

Head (fig. 102) shorter than wide across the compound eyes, twice as long as

the cheek. The epistomal margin (fig. 103) broadly bilobate. Mandible

(fig. 100) with three teeth; the labial palpus (fig. 106) consisting of two segments

(11 : 16), the maxillary (fig. 105) of four (4:6:3:10). Antenna (fig. 101),

formula 1135(3); the scape more than three times as long as the pedicel (13 : 4);

the flagellum with five or six sensilla and long mainly basal setae visible in axial

view of each segment, more and shorter setae over the whole of the disk and less

sensilla in antiaxial view.

Thorax. The legs (hind leg, fig. 104) are quite similar to those of O. epica-

rioides. Fore wing (2 : 1) 1.3 mm long, the submarginal, marginal, stigmal and

postmarginal veins approximately in ratio 30 : 9 : 8 : 4 (fig. 107, but in some

specimens the stigmal and postmarginal are of almost equal length), hyaline, no

fringe; the hind wing (4 : 1) 1.0 mmlong, with a normal fringe.

Gaster. The hypopygium (fig. 108) has a short spine.

Male. — Length of the head (without mandibles) and thorax 0.9—1.0 mm.

Colour yellowish.

Head (fig. 94) slightly longer than wide (greatest length: greatest width,

1.0 : 1.1), only faintly depressed above the epistomal edge and without a sulcus

parallel to the posterior margin. The eyes large, more than thrice as long as the

cheek (7 : 2) and more than half as long as the remaining posterior part of the

head. The toruli of the antennae are situated in the posterior half of the head,

well behind an imaginary line connecting the posterior margins of the eyes.

Mandible two-thirds of the length of the head; three glands; the labial palpus

consists of one segment, the maxillary of two (4 : 5). Antenna (figs. 93, 96—97),

formula 1115(2); the scape distinctly expanded, only slightly so in smaller

specimens; the pedicel about a quarter of the length of the scape; one anellus (it is

difficult to distinguish between the stalk of the third segment and a possible extra

anellus; even if the stalk is counted as a second anellus, there is a difference with

O. epicarioides and royi, which then have three anelli: see Grandi, 1922, fig. vi

2); five subequal funicular segments, each with one antiaxial sensillum, and a

divided club (2 : 1).

Thorax (fig. 94): the pronotum 1/2 times as wide as long, three times as long as

the mesonotum is in the middle, and thrice as long as the propodeum is laterally.

Wing remnant present. Legs (fore tibia, figs. 95, 98) much as in O. epicarioides.

Gaster. The claspers of the genitalia bear three-claws.

Type material. — Holotype d (slide-mounted), Zimbabwe-Rhodesia: Salis-

bury, ex Ficus burkei, ii.1977 (A. Watsham); deposited in RMNH, Leiden (no.

2971). Paratypes: series 2d and 11 mating pairs, same data as holotype (coll.

dates iii-iv.1977); Chishawasha nr. Salisbury, 3 ® 22 d, i, v.1975, ix.1976, i,

vi.1977; Makumbi Mission, Chinamora Reserve, ex Ficus burkei, 35 2 10 6,

xii.1976 (A. Watsham). Zambia: Kafue National Park, Ngoma, ex Ficus burkei,

2 d, 8.xii.1977 (M. G. Bingham, no. 2465); Chipata Golf Course, ex Ficus

burkei, 1 2, 11.1.1979 (M. G. Bingham, no. 2474); Chipata, ex Ficus petersii, 3

2 7 6, 11.1.1979 (M. G. Bingham, no. 2475); Lusaka, Cathedral of the Holy

Cross, ex Ficus burkei, 20 2 20 6, 16.11.1979 (M. G. Bingham, no. 2490). The

194 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

material is deposited in the RMNH, Leiden, some paratypes in the BMNH,

London and in the Watsham Collection.

Derivation of the name. — Tsamvi is the vernacular name (Shonatribe) for

Ficus burkei.

EPICHRYSOMALLINAE

Most of the described genera of this group were listed by Hill (1967), but so far

no key to them has been published and most of them were not critically studied.

Most of the descriptions were based on slide-mounted material in which it is

difficult to see certain important characters and usually too much significance is

_ ascribed to tiny differences in bristles on legs, wings and some other body parts.

Unlike many other groups, in Epichrysomallinae the number of antennal

segments and their shape is often different even in closely related species, often

also in different sexes of the same species, and, if not corroborated by other

features, seems to be of problematic value as a generic character. Unfortunately

most of the existing generic descriptions were based on such characters, some-

times even on the still less reliable difference in the palpal segments. The validity

of such genera was first questioned by Wiebes (1971: 382) when describing a new

species from Sierra Leone. He decided then to place the species under the oldest

name in the group, Sycobia Walker, 1871, and called it S. africana. Later (1975)

he placed similarly another new species from the Aldabra Islands in the Indian

Ocean, Sycobia orientalis. To my knowledge there is only one further species

coming into our consideration, described by Risbec (1955a) from the Malagasy

Republic (Madagascar) as Callimomus imerinensis (although some Indian

species have been compared as well).

Epichrysomallinae seem to be confined to the tropics of the Old World and

probably all are associated with figs. I have studied taxonomically at least two

dozen species, some of which could be named, but am not yet quite sure about

the range of some genera. The oldest genus, Sycobia Walker, is known to me

only in the males of its type-species, S. bethyloides Walker from India. These

males are quite distinct from the males of all African species (described or

undescribed) and because it seems to me doubtful that the rather ordinary

looking female assumed by some authors to belong to S. bethyloides really

belongs there, I am reluctant to accept the name Sycobia for these species. The

second oldest name is Camarothorax Mayr, 1906, based on C. obscurus Mayr

from Java. Although I have not seen any material certainly belonging to this

species (and the type material could not be located in the Mayr collection in

Vienna), the description suggests a genus acceptable for most of the African

species known to me. Therefore I transfer the three previously described

African species to this genus and they should be known as Camarothorax

africanus (Wiebes) comb. nov., C. orientalis (Wiebes) comb. nov. (both from

Sycobia) and Camarothorax imerinensis (Risbec) comb. nov. (from Callimo-

mus).

According to the emerging evidence Epichrysomallinae seem to oviposit into

the female florets of figs. They gall them in a similar way as do the Agaonidae, but

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 195

they oviposit from outside. They are no pollinators and attack only figs of smaller

size, with florets within reach of their coiled ovipositors (see Galil & Copland,

1981). If this picture of their biology proves accurate it remains to explain why

their bodies are invariably so smooth (unsculptured).

Altogether four species have been found to occur in the figs of F. burkei, one

belonging to a new genus Sycotetra, the remaining three to Camarothorax Mayr,

as already mentioned also in the key above.

Sycotetra Boucek, gen. nov.

Type-species: Sycotetra serricornis Boucek, spec. nov.

Head in female with convex vertex, parascrobal areas and temples, and

relatively deep scrobes; no interantennal ridge; occipital carina distinct only

above level of foramen; in facial view head not as high as broad, with rounded

converging genae; antennae inserted slightly above centre of face, distinctly

above lower ocular line. Clypeus with bilobed lower margin, its median incision

accompanied by round depression; dorsally not separated from supraclypeal

area; its lateral margins strongly converging and just above tentorial pits less

apart than one ocellus diameter, each margin then curving towards inner third of

the closely approached antennal toruli. Mouth and mandibles of normal size.

Antennal formula 1106(3), i.e., without distinct anellus; funicular segments

very clearly separated by incisions, asymmetric, appearing distinctly serrate in

the type-species (fig. 111); claval segments almost completely fused.

Thoracic dorsum virtually smooth, with few pairs of short bristles, only the

pair at apex of scutellum long, upturned forwards. Pronotum medially shorter

than mesoscutum; latter with strongly converging incomplete notaular lines

(these not groove-like) which turn caudad at posterior end, at the medio-

posterior pair of setae, still about a quarter of their length before transscutal

suture. Scutellum rather broadly bordering on mesoscutum, this border about as

broad as each axilla, but axillar lines anteriorly not marked as grooves on

surface. Propodeum without significant sculpture; spiracle partly covered by

transparent outer flange. Metapleuron hairy, its hairs much shorter than hairs on

the flat propodeal callus. Mesopleuron with deep grooves separating subalar

sclerite and lower epimeron, but groove indicating pleural suture (running from

mid coxa to fore wing) completely effaced in ventral half. All tarsi four-seg-

mented, basitarsus of mid and hind legs very long. Fore wing venation normal for

the group, postmarginal vein however longer than half of the stigmal, which is

curved and shorter than marginal vein.

Gaster sessile, slightly compressed from sides; epipygium with cerci situated

dorso-laterally; hypopygium slightly exceeding three-quarters of gastral length.

In male head stouter than in female, especially temples more tumid; antennal

formula 1105(3), antennal insertion below centre of face, lower margins of toruli

slightly below lower ocular line; flagellum not serrate, its first segment usually

smaller than any of subequal segments 2 to 5. Pronotum distinctly longer than in

female and pubescence of fore wing more conspicuous. Gaster flat, subsessile.

Comments. — Sycotetra differs from most of the known genera of Epichryso-

196 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

mallinae in having only tetramerous tarsi. These are also found in Odontofrog-

gatia Ishii (1934), from which Sycotetra differs in having incomplete notauli,

normal male mandibles (falcate in Odontofroggatia) and a subsessile male gaster

(petiolate in Odontofroggatia, armed with lateral or ventral spines).

Sycotetra serricornis Boutek, spec. nov.

(figs. 109—113; pl. 3 fig. 2)

Female. — Stretched body 2.0—2.7 mm. Testaceous, gaster usually darker

brown; wings hyaline including most of marginal vein, venation otherwise

brownish.

Head; relative measurements: breadth 85, length (dorsally) 47, height includ-

ing clypeal lobes 72, breadth of frons 56, POL : OOL as 21: 16, eye 40x32.5,

malar space 19, mouth breath 39, distance between toruli and median ocellus 26,

scapus 20X7.5, flagellum plus pedicellus 93. Upper face with shallowly engraved

striae merging at antennae with lengthened engraved reticulation. Malar sulcus

obliterated, indicated merely by a narrow streak of denser reticulation. Anten-

nal toruli about one-third of torulus apart. Flagellar segments (fig. 111) with

Figs. 109—113. Sycotetra serricornis Boutek, gen. & spec. nov., female (109—111) and male (112,

113). 109, head and thorax; 110, gaster and hind leg; 111, antennae, left one in lateral aspect, right

antenna (part) in dorsal aspect; 112, male pronotum, dorsal aspect; 113, male antenna. Figs. 114,

115. Camarothorax equicollis Boutek, spec. nov., female. 114, venation of fore wing; 115, clypeus.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 197

longitudinal plus trichoid sensilla, trichoid ones sparse on funicular segments,

first segment ventrally flattened and there devoid of sensilla.

Pronotum about 0.8 of breadth of mesoscutum, its sides subparallel, anterior-

ly broadly rounded (fig. 109); its dorsum smooth. Flat; both hind and anterior

margins of pronotum emarginate, length in median line only about 0.4 of

maximum breadth and (in presumably normal position) about half as long as

mesoscutum. Relative length of scutellum (less axillae) 60, breadth 50. Fore

femur (less trochantellus) about 2.8 times as long as broad, about as broadened

as hind femur; fore basitarsus dorsally as long as the second, mid basitarsus

dorsally as long as rest of tarsus less claws, hind basitarsus relatively very slightly

shorter. Fore wing venation see pl. 3 fig. 2.

Gaster hardly longer than thorax; in normal position tip of hypopygium

reaching very nearly as far as apex of the spiracle-bearing tergite (fig. 110), most

of hypopygium bare, except for few hairs at tip which is shortly produced

(mucro).

Male. — Usually 1.5—1.8 mm (non-stretched specimens shorter, then length

of body about equal to length of forewing). Antennae paler than in ?. Head only

1.6 times as broad as long in dorsal view, with temples longer, almost parallel just

behind eyes. For antenna see fig. 113. Pronotum (fig. 112) dorsally flat, its

median length about 0.8 of the breadth, hind margin fairly deeply emarginate.

Mesoscutum with distinct engraved striation generally parallel to notaular lines.

Fore wing pubescence beginning at base of marginal vein (in 9 bare area

stretches to base of stigmal vein), similar to that shown in fig. 122.

Type material. — Holotype ® (on card), Zimbabwe-Rhodesia: Makumbi

Mission nr. Salisbury, ex Ficus burkei, xii.1976 (A. Watsham); deposited in

BMNH, London. Paratypes: 44 ® 33 d (1 d observed mating with holotype

2), same data as holotype; Chishawasha nr. Salisbury, 3 gd, xii.1974, 5 ®,

ix.1974, iii. and v.1975 (A. Watsham). Some paratypes in RMNH, Leiden and

the Watsham Collection.

Camarothorax Mayr

Type-species: Camarothorax obscurus Mayr.

For discussion on Camarothorax see above, under the heading Epichrysomal-

linae.

Camarothorax equicollis Boutek, spec. nov.

(figs. 114—121; pl. 3 fig. 3)

Female. — 2.2—2.7 mm. Reddish testaceous but often with gaster dark brown

(except for paler base), as well as mesopleura, metathorax plus propodeum,

antennal flagellum and legs except tarsi. Wings hyaline, including parts of

venation which alternate with darker colour at base, middle and apex of

submarginal vein and at base and apex of stigmal vein.

Head orthognathous, scarcely broader than mesoscutum (36 : 34), in dorsal

198 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

view fully 1.9 times as broad as long (stout), in facial view (fig. 117) barely 1.2

times as broad as high. Relative measurements: head width 89, frons 53,

POL : OOL as 23 : 13, eye 40x34, malar space 27, mouth 44; toruli 30 from

lower margin of clypeal lobes, 20 from eye and 32 from median ocellus; slender

scapus 31, flagellum plus pedicellus 86. Vertex very weakly convex, with bristles

as in 6 but without short ones mesad and laterad of ocelli; also sculpture and

pilosity similar to that of male. Occipital carina conspicuous and complete,

reaching mouth margin on hind side of head; temples rounded, short. Lower

face with some very short hairs, longer hairs present only at mouth margin and on

mandibles; clypeus subtriangular, its lower margin with two deeply separated

lobes (fig. 115), dorsal margin against supraclypeal area indicated by dark line,

supraclypeal area itself narrower than one torulus, almost parallel-sided, about

three times as high as broad. Malar groove distinct. Antenna (fig. 118): pedicel-

lus short; flagellum stout except for thin anellus, with seven funicular segments,

all of them transverse, distal ones usually about twice as broad as long; formula

1117(3).

Figs. 116—121. Camarothorax equicollis Boutek, spec. nov., female (116—118) and male (119—

121). 116, apex of gaster in lateral view; 117, head; 118, antenna; 119, male antenna; 120, male head;

121, male thorax, dorsal aspect.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 199

Thorax shiny, with similar fine engraved striae as head; dorsally with two

bristles on each side of pronotum, two posteriorly at each outer corner of

mesoscutum, the meso-posterior pair more than twice from each other than

from scutellum; on scutellum one pair of lateral bristles in middle of length and

another pair at hind margin. Pronotum distinctly narrower than mesoscutum, its

median length to breadth in ratio as 26 : 62; anterior corners broadly rounded.

Notaular grooves shallow, posteriorly sinuate and meeting scutellum slightly

inside of axillar grooves; latter grooves separated from each other by little less

than breadth of one axilla (21 : 26). Relative length of scutellum 59, breadth 53,

median length of propodeum 20. Propodeum with fine median carina accompa-

nied by a groove connected anteriorly with a broader groove running along

anterior margin, narrowing laterad and subdivided in several areolae; short

carinae arising also from raised petiolar margin of propodeum; spiracle round,

less than its diameter from metanotum, anteriorly overroofed by transparent

flange; callus with long hairs, these reduced posteriorly to hardly more than one

row. Metapleuron convex, reticulate, bare. Mesopleuron ventrally very flat,

dorsally with longitudinal striation; a curved groove separating subalar sclerite,

a distinct bent groove delimiting lower epimeron (katepimeron). Fore femur

and hind femur each about three times as long as broad. Fore wing pilosity

strongly reduced, hairs extremely short, sparse, marginal fringe absent; bristles

of veins reduced to a few on darker parts of venation (fig. 114); relative

measurements: costal cell length about 115, marginal vein 26, postmarginal 9,

stigmal 18.

Gaster sessile, only moderately compressed from sides, hardly as long as

thorax. Cerci close to each other, on ventral side. Hypopygium ending slightly

beyond middle of gaster but with thin white median projection (mucro, fig. 116)

about of length of hind basitarsus.

Male (pl. 3 fig. 3). — Stretched body 1.9—2.7 mm (holotype 2.6 mm). Dark

brown, but mesothorax dorsally sometimes more or less testaceous; antennae,

fore tibia and all tarsi testaceous. Wings slightly infumate but hyaline in basal

one-third sending out a curved hyaline streak, also quadrangular area below

marginal vein hyaline; venation brown, but marginal vein pale. |

Head subprognathous, elongate-subquadrangular (fig. 120). Relative mea-

surements: breadth 65, thickness 44, height (here = length) 78, breadth of frons

39, POL : OOL as 19 : 12, eye 38x26, malar space 23, width of mouth 44,

distance between toruli and median ocellus 42, scapus length 37, pedicellus plus

flagellum 69. Surface of body shiny, with very fine shallow engraved striation;

short hairs scattered on lower face, spreading partly along inner orbits and across

genae to temples; on vertex regular bristles: one in front of each lateral ocellus,

one antero-laterad from that ocellus at eye margin, four bristles in cross-arch on

vertex and a short bristle mesad of each lateral ocellus. Occipital carina even

dorsally indistinct. Malar sulcus fine but distinct. No supraclypeal area, and

clypeus ill-defined dorsally, very transverse, lobes of lower margin as if converg-

ing, separated by a subtriangular incision. Antennal insertion about half-way

between ocular line and mouth margin, toruli close to each other. For antenna

see fig. 119, formula 11071; scapus straight; first flagellar segment undivided,

200 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

distinctly longer than following one but scarcely as long as pedicellus; clava not

distinctly subdivided.

Thorax with fine striation similar to that on head, dorsally slightly flattened, in

shape slightly widening caudad (fig. 121), in stretched position about 2.5 times

as long as broad. Pronotum about 1.2 times as long as broad, its hind margin only

shallowly emarginate; sides subparallel. Mesoscutum (if pronotum in horizontal

position) fully twice as broad as long, only half as long as pronotum; notauli

extremely shallow though groove-like, very nearly complete, almost meeting on

scutellar border. Scutellum about 1.2 times as long as broad, most of border with

axilla indistinct. Propodeum subhorizontal, without significant sculpture, only

medio-anteriorly with faint converging grooves separated by short fine carina.

Metapleuron and mesopleuron as in female, only with odd hairs in front of mid

coxae. Legs strong but tibiae and tarsi relatively long and slender; hind coxa

pyriform, dorsally and laterally bare; fore femur stout, 2.1 times as long as

broad, hind femur about 2.9 times as long as broad; mid basitarsus dorsally

slightly shorter than combined dorsal length of segments 2 to 4. Proximal third of

fore wing bare except basal cell and apex of costal cell, bare area stretching to

base of stigmal vein; otherwise distinctly pubescent, hairs moderately dense;

marginal fringe conspicuous; bristles on submarginal vein absent only on a short

stretch just before parastigma.

Gaster flat, mostly short-oval.

Type material. — Holotype d (on card), Zimbabwe-Rhodesia: Makumbi

Mission nr. Salisbury, ex F. burkei, xii.1976 (A. Watsham); deposited in

BMNH, London. Paratypes: 12 © 5 d, same data as holotype; Chishawasha nr.

Salisbury, 11 © 8 6, ix. and xii.1974, vii.1976, iii. and iv.1977 (A. Watsham).

South Africa: Pondoland, Port St. Johns, 1 ©, vii.1923 (R. E. Turner). Uganda:

Kawanda, 1 ©, ii.1939 (T. H. C. Taylor). Some paratypes in RMNH, Leiden

and the Watsham Collection.

Comments. — This species, especially by its long “horse” head and pronotum

(hence its name equicollis) reminds one much of Camarothorax orientalis

(Wiebes) described in Sycobia from the Aldabra Islands (1975). C. equicollis

differs from C. orientalis (of which I could examine the type material) in the

female mainly by the lateral ocellus being nearer to the median ocellus than to

the eye (the reverse in orientalis), a different disposition of hairs on the vertex, a

slightly more transverse head (in frontal view), a much shorter pronotum

(median length to breadth in orientalis as 13 : 26), and by the distinctly com-

pressed gaster (not compressed in orientalis); in the male by the relatively less

elongate head with larger eyes which are much longer than part of head anterior

to eyes in equicollis, but only about as long in orientalis.

The above material of C. equicollis comes mainly from the Rhodesian Ficus

burkei, but a very similar form is known to us from the figs of F. petersü from

South West Africa (Namibia). The latter form cannot be separated morphologi-

cally but is much darker, having only the pronotum and lower parts of the head

pale testaceous. I regard it as a form of C. equicollis, although the material is not

included as paratypes (it may be a climatic form, or does the colour difference

suggest a beginning of speciation?)

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 201

Camarothorax longimucro Boucek, spec. nov.

(figs. 122-125; pl. 3 fig. 4)

Because of great similarity of this species to C. equicollis the stress is made in

the following description on the diagnostic characters.

Female. — 2.05—2.35 mm. Testaceous, but propodeum and gaster some-

times slightly darker than the rest; wings hyaline, venation with a pattern similar

to equicollis but generally paler.

Head in dorsal view transverse-quadrangular, about 1.85 times as broad as

long, with swollen temples which are subparallel for short distance at eyes; in

facial view (fig. 125) head nearly 1.2 times as broad as high. Sculpture on head,

as well as on thorax, similar to that of equicollis, but deeper, therefore surface

duller. Relative measurements: head breadth 83, frons 51, POL: OOL as

22 : 14, eye 34x29, malar space 24,-mouth 42, toruli 25 from lower margin of

clypeal lobes and 33 from median ocellus (lower torular margin on lower ocular

line), scapus 31, flagellum plus pedicellus 82. Ocellar angle about 135°, ocellar

area about three times as broad as high. Compared with equicollis eyes are

smaller, supraclypeal area (separated by distinct line from clypeus, fig. 125) is

only about twice as high as broad, as broad as one torulus; antennae inserted

slightly lower on face, pedicellus fully 1.6 times as long as broad, flagellum

slightly more slender, with only six funicular segments.

Pronotum parallel-sided, only 0.75 times as broad as mesoscutum, its median

length about 0.47 the breadth; anterior corners (shoulders) subrectangular.

Notaular grooves meeting scutellum slightly outside of axillar grooves; latter

grooves separated anteriorly by distance subequal to half breadth of axilla.

Scutellum about 1.15 times as long as broad, posteriorly more regularly semicir-

cular than in equicollis, but propodeum and sculpture of pleura as in that species,

except that metapleuron is narrower, hind lateral corner of propodeum slightly

more produced, the groove delimiting subalar sclerite ventrally is straight. Legs

relatively slender; fore femur about 2.5 times, hind femur nearly 3 times as long

as broad; hind basitarsus dorsally about as long as rest of tarsus (less pulvillus).

Fore wing as in equicollis, relative length of costal cell 120, marginal vein 26,

postmarginal 10, stigmal vein 21.

Gaster sessile, fairly compressed from sides, about as long as head plus thorax

combined. Hypopygial mucro slender (fig. 124), about as long as two basal

segments of hind tarsus combined. Cerci situated ventrally, close to each other.

Male. — About 1.7—1.9 mm (if stretched). Pale testaceous (only eyes, ocelli

and pulvilli dark); fore wing venation (fig. 122) pale, only parastigma and

stigmal vein more pigmented.

Head almost orthognathous (mouth downwards), in facial view fully 1.1 times

as broad as high. Relative measurements: breadth of head 71, length (dorsally)

about 40, height 62, frons 42, eye 34x26, malar space 16, scapus 25X9, flagellum

plus pedicellus 55. Occipital carina fine and developed only dorsally. Head

narrower in temples than across eyes but temples parallel for short distance just

behind eyes; a few additional hairs posteriorly on vertex; two pairs of dorsal

adorbital bristles (one pair in equicollis). Antennal toruli situated below lower

202 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

ocular line; supraclypeal area short, with sides diverging upwards, ventrally

separated by distinct line from clypeus. Antenna short and stout, clavate

(fig. 123); scapus slightly narrowed basally; flagellum more distinctly tapering

towards base; seven distinct funicular segments, no anellus.

Thorax long, with large pronotum which is about 0.85 times as broad as

mesoscutum, but normally (in subhorizontal position) distinctly longer; maxi-

mum length of pronotum equal to breadth, but because of emargination of hind

margin, in median line shorter than broad as 45 : 54, broadly rounded anterior-

ly; pronotum dorsally nearly smooth, with 4 to 6 bristles in an anterior cross-line,

one pair of sublateral bristles behind this row and 4 bristles at hind margin

(submedian ones more apart). On propodeum the T-shaped groove obliterated;

Figs. 122-125. Camarothorax longimucro Boutek, spec. nov., male (122, 123) and female (124,

125). 122, venation of fore wing; 123, antenna; 124, female gaster and hind leg; 125, female head and

antenna. Figs. 126-128. Camarothorax brevimucro Boutek, spec. nov., female (126, 127) and

male (128). 126, head; 127, apex of gaster; 128, male antenna.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 203

also grooves on mesopleuron almost obliterated. Legs strong and stout; fore

femur hardly 2.1 times, hind femur about 2.4 times, as long as broad; fore tarsal

segments 1 to 4 very short; hind tibia dorsally bristly; hind basitarsus dorsally

about as long as segments 2 plus 3 combined (dorsally). Fore wing (fig. 122) with

pilosity similar to that of d equicollis; on lower surface pubescence extending

partly to marginal vein.

Gaster flat, much shorter than thorax.

Type material. — Holotype © (on card), Zimbabwe-Rhodesia: Chishawasha

nr. Salisbury, ex F. burkei, vi.1977 (A. Watsham); deposited in BMNH,

London. Paratypes: same origin as holotype, 2 d, 3.xii.1974 and 5 ©, 1976.

Some paratypes deposited in RMNH, Leiden, PPRI (Plant Protection Research

Institute), Pretoria and the Watsham Collection.

Comments. — The name of C. longimucro refers to the unusually long spine-

like projection (mucro) of the female hypopygium. In this respect the species is

similar to C. equicollis. The female looks otherwise very much as that of

C. brevimucro, especially in the antennae, but the males of the two species have

very different antennae and the female of C. brevimucro, as the name suggests,

has only a short projection on its hypopygium. Camarothorax longimucro is also

similar to C. africanus (Wiebes) from West Africa. The latter species differs

from C. longimucro by its larger size, and in both sexes by a more robust body

with a stout head which looks subquadrangular dorsally and is, in female, only

about 1.65 times as broad as long, with the occipital carina nearly reaching the

level with ocelli (this carina much lower in longimucro), the pronotum with

almost parallel sides and longer, in female medially about 0.6 times as long as

broad, in both sexes of C. africanus the axillar grooves are rather strongly

converging forwards, thus considerably narrowing the anterior margin of the

scutellum, and the clypeus is distinctly separated from the supraclypeal area (as

in C. longimucro, fig. 125, but unlike in the otherwise also similar C. brevimu-

cro, fig. 126).

Camarothorax brevimucro Boutek, spec. nov.

(figs. 126—128; pl. 3 fig. 5)

Female. — 1.3—2.5 mm (holotype 1.8 mm). Testaceous to dark brown,

sometimes either pale or dark, usually pale colour spreading first on sides of

pronotum, then on lower parts of head and in a streak on sides of thoracic dorsum

including fore coxa, prepectus and sides of scutellum (so in holotype). Wings

hyaline, venation uniformly pale brown. Otherwise very similar to C. longimu-

cro, hence stress is made on diagnostic characters.

Head with occipital carina reaching higher up than in longimucro, therefore

vertex less sloping posteriorly. Ocellar area about twice as broad as long, angle of

ocelli about 120°. In facial view (fig. 128) head rather transverse, about 1.37

times as broad as high; clypeal lobes rather short, clypeus dorsally confluent with

supraclypeal area. Relative measurements: breadth of head 78, frons 48,

POL : OOL as 18: 11, eye 34x29 (rather large), malar space 20 (relatively

204 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

short), scapus 22, flagellum plus pedicellus about 60. Flagellar segments mostly

more transverse than in /ongimucro.

Thorax slightly stouter than in /ongimucro; median carina on propodeum

reduced to mere rudiment anteriorly; several bristles on dorsal part of metapleu-

ron. Fore and hind femur each about 2.5—2.6 times as long as broad. Relative

measurements in fore wing: costal cell length 95, marginal vein 18, postmarginal

about 6, stigmal vein 18 (the marginal relatively short).

Gaster sessile, only moderately compressed from sides, about as long as

thorax. Hypopygium reaching near to apex of gaster, its median projection

(mucro) short (fig. 127; hence the name).

Male. — 1.4—1.9 mm. Mainly yellowish but often with darkened gaster and,

usually less distinctly darker on vertex and on median parts of thoracic dorsum.

Wings mostly hyaline but sometimes slightly infumate, mainly with a faint

median streak and more infumate in basal half where a strongly curved line

following the cubital vein and another longitudinal streak along hind margin are

left hyaline.

Very similar to & of longimucro but body less slender and antennae only with

five segments between pedicellus and clava (fig. 128); clypeus dorsally fused

with supraclypeal area; vertex subhorizontal behind ocelli, relative distance

between lateral ocelli (POL) 18, both only 14 from distinct occipital carina;

length of pronotum medially about 0.65 the breadth, in two-thirds of length with

a pair of submedian bristles. Fore femur nearly 2.3 times as long as broad (hind

femur barely more slender); fore tarsus very short; hind basitarsus about as long

as three following segments combined, measured dorsally. Fore wing pilosity

about as in C. equicollis: relative length of marginal vein 22, postmarginal about

3 (indistinct), stigmal vein 19. Gaster subpetiolate, flat, short oval; median

length of petiole barely a third of maximum breadth.

Type material. — Holotype © (on card), Zimbabwe-Rhodesia: Makumbi

Mission nr. Salisbury, ex F. burkei, xii.1976 (A. Watsham); deposited in

BMNH, London. Paratypes: same data as holotype, 100 9 109 3; Chishawasha

Reserve, 1 ©, iii.1975 (all A. Watsham). South Africa: Pondoland, Port St.

Johns, 1 ®, viii.1923 (R. E. Turner). Some paratypes deposited in RMNH,

Leiden, PPRI, Pretoria and the Watsham collection.

Comments. — The main characters separating this species are summarized in

the key and in the comments on C. longimucro. The male of C. brevimucro is

similar in many features to that of Sycotetra serricornis, but it has complete

notaular grooves, five-segmented tarsi and differs otherwise in various details

mentioned in the description.

EURYTOMINAE

Figs, even if inhabited only by the pollinating Agaonidae, represent an

ecological niche virtually identical to many plant galls and as such attract many

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 205

species of gall-inhabiting Eurytomidae. All the African species associated with

figs belong to subfamily Eurytominae, as defined by me earlier (e.g., in Peck,

Boutek & Hoffer, 1964 : 23) and quite recently by Subba Rao (1978). There

seems to be no good reason to split the subfamily, as some authors traditionally

did. Eurytominae include a variety of genera, with numerous species both of

phytophagous and entomophagous habits.

There are still considerable difficulties with the range of many genera and it

proved the best way to place two of the species associated with Ficus burkei into

two new genera, whilst the majority belongs to Sycophila Walker and one

species can be undoubtedly classified as Eurytoma Illiger. Their separation is

summarised in the key, based mainly on practical characters (not necessarily of

generic value, as some, e.g., in Eurytoma ficusgallae).

Eurytoma Illiger

One species, apparently undescribed, belongs to this well known genus.

Eurytoma ficusgallae Boucek, spec. nov.

(figs. 129—130; pl. 3 fig. 6)

Female. — 2.5—3.4 mm (holotype 3.3 mm). Black; head and thorax with

short silvery pubescence; antennae, tegulae, venation and legs beyond coxae

rufous, tarsi paler, sometimes pedicels infuscate, rarely also femora; wings

hyaline.

Head dorsally twice as broad as long, anteriorly 1.4 times as broad as high;

densely umbilicately punctured, including malar space, but on lower face

punctures confluent into grooves separated by raised striae radiating from

depressed clypeal margin. Frons convex but not bulging; deep scrobes barely

narrower than parascrobal area, its edges carinate, especially in upper third,

united at ocellus, in a slight depression. Inner orbits with slight carina. Hairs on

face very dense and flattened, arranged as if radiating from antennal base; lower

edge of toruli at half distance between median ocellus and the emarginate mouth

margin; on lower face median stria raised slightly more than others. In frontal

view, mouth margin medially distinctly emarginate and depressed (including

clypeus) but raised laterad of clypeus, narrowly notched above base of each

mandible so that genal lamina projects there as a tooth; malar sulcus absent.

Relative measurements: head width 55, frons 34, POL 12.5, OOL 7, eye 22x20,

malar space 18.5, scapus 17X5, flagellum plus pedicellus 52. Antenna 11153;

scapus reaching top of median ocellus, distinctly tapering in apical third

(fig. 129); pedicellus subglobose; funicular segments oblong, the first distinctly,

the fifth only slightly longer than broad, all with sparse longitudinal sensilla.

Thorax: densely punctate; dorsum about 1.43 times as long as mesoscutum

broad; pronotal sides rounded, anteriorly slightly carinate (carina vertical).

Hind margin of mid lobe of mesoscutum in middle slightly produced. Tegula

distinctly striate-reticulate at hind margin. Scutellum rather flat, at apex broadly

rounded (almost truncate) and with hairs directed towards median line; axillula

impunctate, well delimited even dorsally, its posterior subvertical border out-

206 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 129, 130. Eurytoma ficusgallae Boutek, spec. nov. 129, female antenna; 130, male antenna.

Figs. 131—134. Ficomila gambiensis (Risbec), female (131—133) and male (134). 131, venation of

fore wing; 132, antenna (less scapus); 133, body in lateral view; 134, male antenna. Figs. 135—137.

Ficomila curtivena Bouéek, gen. & spec. nov., female. 135, venation of fore wing; 136, gaster; 137,

lateral aspect of part of thorax, showing prepectus (prp), mesopleuron with characteristic feature of

the genus, and mid coxa (mc).

standing. Propodeum steep; the irregularly round large median area only

slightly depressed, with low weak cross-carinae arranged as a tree: lower ones

nearly horizontal, upper ones increasingly turning upwards; bottom between

them reticulate; margins of area partly carinate against the foveolate-areolate

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 207

convex sublateral parts. Mesopleuron in posterior part horizontally rugulose-

striate, anteriorly with epicnemial area delimited by subparallel low keels and

containing a partly doubled row of piliferous puncta. Mesosternal shelf distinctly

delimited by wavy carina, its minimum length (in front of each mid coxa) equal to

breadth of fore tibia; ventral surface hairy, with a few hairs also in front of the

shelf carina. Fore coxa anteriorly hairy, with strong oblique lamina beyond basal

third; mid coxa without auricular lamella. Hind tibia on dorsal edge with a row of

about 9 slightly alternating bristles, longest one as long as inner spur. Fore wing

with marginal, postmarginal and stigmal veins subequal in length; basal cell

hairy in distal third; costal cell pubescent on ventral surface but dorsally bare.

Gaster sessile, about as high and long as thorax but narrower, compressed,

though not keeled dorsally; in median line tergites 2, 3 and 4 subequal in length.

Surface smooth, but beginning with fourth tergite finely punctulate; fourth

tergite sublaterally at base with a row of hairs which is more complete on the fifth,

sixth tergite wholly hairy; epipygium very short, barely up-turned.

Male. — 1.7—3.0 mm. Pubescence on head and thorax, especially on face,

golden. Flagellum brownish, with 4 bristly funicular and 3 claval segments

(fig. 130). Scapus with distinct granulate boss beyond middle; first funicular

segment distinctly elongate, segments 2 to 4 subequal, very slightly longer than

broad. Antennal insertion distinctly above middle of face. Median area of

propodeum more coarsely sculptured than in 2. Gastral petiole about 1.5 times

as long as broad, laterally and anteriorly carinate, dorsally rather flat, granulate,

anterior crestlike margin arcuate.

Type material. — Holotype 9: Zimbabwe-Rhodesia: Chishawasha nr. Salis-

bury, xii.1974 (A. Watsham); deposited in BMNH, London. Paratypes: 13 ©

and 9 gd, same origin as holotype, mostly reared from galled figs of F. burkei,

11.1974, iii.-x1.1975, 11.1976; 15 © and 4 d, Makumbi Mission nr. Salisbury,

same host, xii.1976 (all A. Watsham). Some paratypes deposited in RMNH,

Leiden and in the Watsham collection.

Comments. — Eurytoma ficusgallae seems to be easily separable from all the

described African species of the genus. By the combination of the well delimited

(carinate) mesosternal shelf and the presence of an oblique laminate carina on

fore coxa, E. ficusgallae approaches the nodularis-group of Eurytoma. Howev-

er, unlike the latter group, it has the coarse radiating striation on the lower face,

with a medially emarginate and depressed mouth margin. This character,

combined with the mesosternal shelf, is present in the braconidis-group (para-

sites in cocoons of Braconidae and Ichneumonidae, etc.), but there the shelf is

differently formed (not well delimited laterally), the massive genal carina

merges smoothly with the sublateral mouth border and there is a pair of

outstanding carinae below the antennae, instead of a median carina as in

ficusgallae. The facial radiation reminds one also of the robusta-group but in

those species the fore coxa has on its anterior face just a shallow longitudinal

channel, not a laminate carina. This carina is further present in the latrodecti-

group (called Desantisca by Burks; parasites in spider egg-cocoons) the species

208 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

of which much resemble E. ficusgallae also in many other characters and are also

similar and closely related to the braconidis-group. The latrodecti-group differs

in having a relatively deep reticulation on the tegulae and in female sex loose

funicular segments.

Syceurytoma Boutek, gen. nov.

Type-species: Syceurytoma ficus Boucek, spec. nov.

Body generally in shape and sculpture as in most Eurytoma species, i.e., with

head and thorax densely umbilicately punctured, with very short pubescence.

Head transverse; scrobes reaching ocellus, not broader than parascrobal area.

Interantennal crest small but high, doubled by broad median groove. Lower face

coarsely striate, median stria slightly raised; striae radiating from lower clypeal

margin which is slightly bilobed: lobes separated by depression in emargination;

tentorial pits indistinct. Malar sulcus slightly indicated, about as long as shorter

diameter of the relatively small eye. Gena posteriorly with strong laminate

carina joining ventrally the reflexed-carinate mouth border. Occipital carina

low, ending near middle of posterior genal carina by a small tooth and this

connected by horizontal carina with hypostomal carina (which delimits the

cavity containing mouth organs). Antennae inserted slightly above middle of

face, formula in female 11153, with scape narrowed apically and with small

anellus; in male 11143, with funicular segments more distinctly constricted at

both ends, longer and more hairy: hairs semidistant and about as long as breadth

of segments and mostly not distinctly whorled (fig. 140).

Pronotum large and virtually as broad as, and slightly longer than, the

mesoscutum; antero-laterally with vertical carina. Scutellum as in Eurytoma;

axillulae not well delimited. Propodeum steep, medially depressed, irregularly

alveolate, anteriorly with two carinae diverging from middle as in Sycophila.

Mesopleuron anteriorly delimited by epicnemial strip which is reduced in

ventral half to single carina; this carina becomes high laminate where turning

mesad, there marking off short mesosternal shelf; more mesad carina lowered

and joining in median line a double rounded projection (fig. 138); this projec-

tion is strongly excavated from either side and anteriorly connected with two

weaker subparallel carinae, each with a side branch. Fore coxa laterally carinate

but its anterior face rather flat, with slight meso-distal depression. Fore wing

much as in Sycophila: marginal vein strongly enlarged (more so in male than in

female), accompanied by a slight infuscation; postmarginal and stigmal veins

short (fig. 139).

Gastral petiole subquadrate in female, in male distinctly elongate, dorsally

flat, anteriorly expanded on sides (fig. 141), with sharp anterior crest. Gaster

globose, in female broad and dorsally only weakly convex, with third tergite the

longest.

Comments. — The new genus is proposed, rather reluctantly, for a distinctive

species which shows affinities both to Eurytoma Illiger and Sycophila Walker. It

is described after an attempt at a critical analysis of the characters used previous-

ly as generic characters, but also including a number of new ones. The only

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 209

alternative to this generic separation would be a union of many genera of the

former Eurytominae, Eudecatominae and Harmolitinae, which, indeed, I

regard all as belonging to Eurytominae (in Peck, Boutek & Hoffer, 1964: 23).

These subfamilies are still maintained by some authors, in my opinion without

good reason.

Another genus which to some extent intergrades between Eurytoma and

Sycophila is Paradecatoma Masi. It was originally based on a single female

named Paradecatoma bannensis Masi from Somalia but is known to me now in

both sexes from Ethiopia (Harrar), Yemen, Tanzania and South Africa (Port St.

Johns). Some specimens were apparently reared but no host data are attached.

Paradecatoma has the posterior genal carina (I think that “‘postgenal carina” of

Claridge is rather occipital carina) strong and complete as in Syceurytoma and

most Eurytoma, but has, apart from other characters, an impunctate boss on

malar space, the lower margin of clypeus with a small median tooth, a cross-

striate and hairy anterior face of fore coxa, a coarsely punctured and hairy long

mesosternal shelf which is anteriorly not carinate but abruptly turning smooth,

and the male antennal flagellum with six two-whorled funicular segments plus an

indistinctly 3-segmented clava. The bristly male flagellum puts Paradecatoma

nearer to Eurytoma, but again the marginal vein is enlarged and stigmated by

infuscation, although short and with certain similarity to the genus described

here below, Ficomila. In the Eurytominae the posteriorly rounded gena seems

to indicate, at least to some extent, a degree towards phytophagy, as suggested

by the rounded gena in Tetramesa Walker, Systole Walker, Bruchophagus

Ashmead (and the extremely close Risbecoma Subba Rao, 1978, based on the

supposedly parasitic Eurytoma bruchocida Risbec which is in fact phytophagous

in the Acacia seeds!). If this hypothesis works, both Syceurytoma and Paradeca-

toma should be parasitic in habit.

Syceurytoma ficus Bouéek, spec. nov.

(figs. 139—141; pl. 4 figs. 1, 2)

Female. — 2.1—3.4 mm. Ochreous to rufous, sometimes (especially in small

specimens) with black spreading from median line on thorax and vertex.

Head; relative measurements: breadth 68, length 36, height 52, frontovertex

45, POL 13, OOL 13, eye 24.520, malar space 22, scape 21.5, flagellum plus

pedicellus 59. Fine granulate reticulation on scapus confluent near base into

cross-striation. Narrow interstices of umbilicate punctures on head and thorax

reticulate. Punctured thoracic dorsum only about 1.44 times as long as broad.

Anterior margin of pronotal collar broadly emarginate in middle, its sides longer

than half its breadth (33 : 62). Scutellum barely longer than broad. Propodeum

rather steep, convex, except for shallow median depression; mainly irregularly

alveolate behind two carinae diverging from middle anteriorly and arcuately

turning to anterior end of spiracles; sometimes median depression partly

delimited by subparallel irregular carinae (more often in smaller specimens).

Exposed part of prepectus concave, about as large as tegula, separated by a

carina from the normally concealed part which is broad channel-like. Fore wing:

costal cell dorsally nearly bare but densely pubescent underneath; basal third of

210 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Figs. 138—141. Syceurytoma ficus Boutek, gen. & spec. nov., female (138) and male (139—141).

138, characteristic sculpture on mesosternal part; 139, venation of male fore wing; 140, male

antenna; 141, gastral petiole of male. Figs. 142, 143. Sycophila kestraneura (Masi), female. 142,

antenna; 143, venation of fore wing. Figs. 144, 145. Sycophila punctum Boutek, spec. nov., 144,

male head with antenna and part of thorax; 145, venation of female fore wing. Fig. 146. Sycophila

sessilis Bouëek, spec. nov., venation of female fore wing.

wing below submarginal vein mainly bare, but basal fold with hairs in 1 or 2 rows;

parastigma distinctly enlarged; marginal vein broader than length of stigmal

vein, its lower margin slightly convex.

Gastral body broadly convex; exposed part of third tergite nearly to about

twice as long as the second, both smooth; epipygium very short, transverse,

barely up-turned.

Male. — 1.45—3.4 mm. Thorax usually with black widespread, more than in

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 211

darkest female, sometimes whole dorsal side of body black, rarely wholly

ochreous as in most females. Antenna in most specimens not appearing very

bristly (fig. 140) but bristles often more outstanding in smaller specimens and

then fourth funicular segment with two whorls. For gastral petiole see fig. 141,

for fore wing venation fig. 139, made from a specimen of size over 3 mm; smaller

specimens have hairs relatively sparser, in large specimens also wing blade is

partly infuscate.

Type material. — Holotype 9: Zimbabwe-Rhodesia: Chishawasha nr. Salis-

bury, from F. burkei (= “natalensis”), 10.xii.1974 (A. Watsham); in BMNH,

London. Paratypes: 104 © and 61 d, same origin as holotype, xi.1974—vi.1975,

ii.1976; 38 © and 5 d, Makumbi Mission nr. Salisbury, iv.1975 and xii.1976 (all

A. Watsham). Uganda: Kampala, “ex stem galls on Ficus sycomorus”, 3 ®, 3

d, 8.x.1935 (H. C. Taylor). South Africa: 1 &, Zululand, Eshowe, iv.1926 (R.

E. Turner); 1 2, 1 d, Natal, Weenen, xi.1923 (H. P. Thomasset); 3 ©,

Pondoland, vii. and xi.1923 (R. E. Turner); 1 ©, East London, v.1923 (N. K.

Munroe). Paratypes are deposited in RMNH, Leiden and in the Watsham

collection.

Ficomila Boucek, gen. nov.

Type-species: Ficomila curtivena Boutek, spec. nov.

Puncturation on head obliterate but on thorax often distinctly umbilicate,

becoming relatively less dense and obliterate in smaller and pale-coloured

specimens, usually absent on median part of scutellum in males. Scrobes deep

but not carinate on margins, not reaching ocellus; interantennal ridge ending in

abrupt angular tooth; antennal toruli slightly above middle of face. Clypeus

delimited laterally by grooves reaching tentorial pits; lower margin bilobed.

Malar groove distinct; gena posteriorly edged as in Sycophila; occiput with weak

and low occipital fold (normally developed as carina running from upper margin

of foramen obliquely towards mouth corners) only half way down, ending at

change of plane. Female antenna 11153; scapus tapering apically; funicular

segments short; male antenna 11143; first funicular segment the longest, con-

stricted at base, longer than pedicel.

Pronotal collar on shoulders only bluntly edged, not carinate. Scutellum with

axillulae indistinctly separated. Propodeum: median part flat or weakly con-

cave, moderately finely reticulate in larger specimens, in smaller ones sculpture

partly obliterate; median area anteriorly delimited by arcuate carina which

broadly touches metanotal margin (as in some Sycophila, slightly less broadly

than in Systole and Bruchophagus). Mesepimeron horizontally striate, sepa-

rated from mesepisternum by fine carina; mesosternal shelf partly delimited:

sublaterally by short cross-carina as continuation of epicnemial edge (weak in

some males) and in middle by a rounded or slightly depressed (but still very

distinct) projection which is hollowed from both sides into a shape of a little

elongate window (fig. 137), similar to that of Syceurytoma (fig. 138). Fore wing

with short marginal vein widening distally, stigmal vein curved on its upper

margin and longer than short postmarginal vein.

212 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Gaster of female very nearly sessile (petiole barely visible dorsally), about as

long as thorax, convex or compressed; fourth tergite much longer (F. curtivena)

or hardly longer (F. gambiensis) than the third; epipygium short. In male petiole

longer than hind coxa.

Comments. — This is another peculiar group, in some ways again intermed- |

late between Eurytoma and Sycophila. Although at present Eurytoma includes a

whole range of sometimes very different species-groups, they all differ from

Ficomila at least in the venation, especially in the postmarginal vein, which is

always relatively longer, and also in the occipital characters. By the genae,

scrobes and the venation Ficomila approaches more Sycophila, but in that

genus, although its limits were slightly extended recently (Boucek, 1974: 267—

268), the mesosternal region of the thorax is remarkably uniformly shaped,

without any separation of the “mesosternal shelf”, in contrast to Ficomila, as

stressed in the key above.

The genus is represented in the material by two species: of these the type-

species F. curtivena is known only in the females, the other, which I identify as

F. gambiensis (Risbec), in both sexes. Because of certain doubts as to the

identity of the latter species I designate the first as type-species.

Ficomila curtivena Boutek, spec. nov.

(figs. 135—137)

Female. —1.7—2.5 mm. Black, but often with mouth margin, pronotum

lateroanteriorly and subalar sclerite testaceous, sometimes pale brown spreads

on lower face, eye orbits and most of pronotum, with vague spots even on sides of

mesoscutum and base of gaster; legs and antennae mainly pale coloured or with

parts infuscate, beginning with coxae, then more or less femora and flagellum.

Wings hyaline to whitish, veins yellowish.

Head with inner eye orbits converging upwards; a few hairs present between

scrobes and ocellus. Relative measurements: breadth of head 92, length 51,

height 72, frontovertex 51, POL 23, OOL 8, eye 43x38, malar space 27, scapus

34, flagellum plus pedicellus 80. Funicular segments 2 to 5 moderately trans-

verse, each slightly shorter than pedicellus; first funicular segment swollen

dorsally, about 1.4 times as long as broad.

Thoracic dorsum sculptured and about 1.57 (less than 1.6) times as long as

broad; pronotum barely narrower than mesoscutum, relative length laterally 33

(compare head). Propodeal, upper diverging cross-carina ending by angular

tooth sublaterally (there turning downwards); area anterior to carina subdivided

by several longitudinal carinulae; in profile propodeum sloping at about 70° as to

plane mesoscutum-scutellum. Fore and hind femora moderately thickened;

hind tibia with a dorsal row of about 9 outstanding bristles, longest bristle about

as long as 0.8 of tibial breadth. For fore wing venation see fig. 135.

Gaster (fig. 136) smooth; slightly compressed from sides, dorsum only weakly

convex; fourth tergite about twice as long as the third, bearing one cross-line of

hairs which is broadly interrupted dorsally. Apex slightly up-turned.

Male. — Not known (one male which might belong to this species, cannot be at

present separated reliably from those of F. gambiensis).

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 213

Type material. — Holotype ?, Zimbabwe-Rhodesia: Chishawasha nr. Salis-

bury, ex F. burkei (natalensis), 26.xi.1974 (A. Watsham); in BMNH, London.

Paratypes: same origin as holotype, 22 ©, xi.-xi1.1974, ili.and ix.1976, ii1.1977;

Makumbi Mission nr. Salisbury, 4 ?, iv.1975 and xii.1976 (all A. Watsham).

South West Africa: Gross Otavi, ex F. petersü, 4 ©, xi.1973 (A. Watsham).

Paratypes deposited in RMNH, Leiden and in the Watsham collection.

Ficomila gambiensis (Risbec) comb. nov.

(figs. 131—134)

Decatoma gambiense Risbec, 1955b: 561—564. Holotype 9, Gambia. Mocquerys (MNHN, Paris)

[examined].

Female. — 2.1—3.2 mm, holotype 4 mm. Usually honey yellow, flagellum

and gaster slightly darker, tiny dark spots at ocelli; in holotype thorax except

pronotum and almost whole gaster dark, blackish, in dark-coloured southern

specimens whole body with all coxae almost black, only sides of pronotum and

base of gaster ventrally paler, femora partly infuscate. Fore wing always with

distinct macula, normally small, but somewhat expanded below level of stigmal

vein in holotype.

Head dorsally about 1.8 times (1.75—1.85X) as broad as long, with only

slightly receding temples, hence appearing rather quadrangular. Scrobes ending

one diameter from ocellus. Antenna: flagellum plus pedicellus less than 1.1

times as long as breadth of head; pedicellus nearly as broad as the broad-filiform

funicle and distinctly shorter than its first segment (fig. 132) which is as if swollen

on dorsal side and slightly concave ventrally where short hairs replace longitudi-

nal sensilla; this segment about twice as long as broad in large holotype but

usually relatively shorter in smaller specimens.

Thorax about 1.75 times as long as broad. Pronotum shallowly emarginate

posteriorly. Outline of propodeum in profile straight; neck distinct. Marginal

vein of fore wing slightly more widening (fig. 131) than in F. curtivena (fig. 135).

Gastral body slightly longer than thorax (fig. 133), fairly strongly compressed

from sides, but dorsal keel rather blunt; hypopygium reaching about one-

quarter along convex ventral edge.

Male. — Size and colour range about as in female (except holotype). Scapus

distinctly broadened in basal two-thirds (fig. 134); pedicellus much shorter than

first funicular segment which is dorsally swollen and bears short hairs in

subconcave basal part ventrally, similar to female. Fore wing strongly hairy in

basal part, unlike in female, marginal vein moderately to very strongly thick-

ened. Petiole dorsally slightly shorter than scutellum, narrowed anteriorly,

about 3.5 times as long as broad.

Material (apart from holotype). — Gambia: Fajara, 2 ©, i.1978 (L. Huggert).

Uganda: Kawanda, ex Ficus sycomorus, 1 ®, ii.1939 (T. H. C. Taylor).

Zimbabwe-Rhodesia: Chishawasha and Makumbi Mission nr. Salisbury,

mostly ex or on F. burkei, 50 2 and 23 6, 1970-1977 (A. Watsham). South

West Africa: Gross Otavi, ex F. petersü, 3 2, 6 à, xi.1973 (A. Watsham). South

Africa: Hartebeeste Poort Dam, 1 2, viii.1974 (A. Watsham).

214 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Comments. — This is apparently a widespread African species which I hope I |

have correctly interpreted, although in the wide variation of the specimens |

actually none is extremely similar to the holotype. However, the difference ,

seems to be due mainly to the unusual size of the holotype. |

Certainly of interest is the fact that this species was found associated with a

least three different fig species, which again must reflect in some variation, apart ,

from the geographical range from Gambia to Uganda and South Africa.

Sycophila Walker |

Type-species: Sycophila decatomoides Walker. |

The genus Sycophila Walker was earlier known under the name Decatoma,

but this name given by Spinola was later recognised, according to its type- |

species, as a junior synonym of Eurytoma Illiger. Decatoma was then replaced by |

Eudecatoma Ashmead, until this name itself was synonymised, together with |

Tineomyza Rondani, under Sycophila (by Boutek, 1974). The synonymy

slightly widened the earlier concept of the genus (earlier based only on North

American and European species), mainly as to the length of the gastral petiole

and the form of the expanded marginal vein. These characters vary considerably

within the genus, especially in many tropical species partly or mostly associated

with galled figs. Even now, however, Sycophila is one of the better delimited

genera of the whole subfamily Eurytominae. It includes species of relatively

usual eurytomid shape of body, with the gena posteriorly more or less edged but

never sharply carinate; the scrobes deep, the parascrobal areas flat or slightly

convex; antennae in female with five free funicular segments (11153), in male

with four funicular segments (11143), without whorls of bristles. The marginal

vein of fore wing is more or less widened (varying as to species and sometimes

also as to sexes: more widened in males) and very nearly always bearing an

infuscate spot which sometimes extends below the vein. The propodeum has

always a median depression which, especially if it is shallow, is anteriorly

delimited by slightly oblique cross-carinae diverging from the middle of the base

(anterior margin). Otherwise the gaster in the males is always petiolate, in the

female the petiole is sometimes shortened. Legs usually normal, but in some

species, especially in the males, mainly fore and hind femora are thickened,

often strongly so.

The intraspecific variation, especially of size and colour, is frequently wide.

Some species vary from completely yellow to completely black, but in the partly

pale specimens a certain pattern can be recognised. The size affects also

sculpture and density of pubescence. Some of the variation is certainly con-

nected with temperature and humidity, some is more host-dependent or climatic

and may offer a geographic pattern. Apparently at least some Sycophila are not

host-specific and it seems that the host range of some species may be fairly wide.

List of described African and Malagasy species of Sycophila:

Sycophila aethiopica (Silvestri, 1915) comb. nov. (from Decatoma), Eritrea

Sycophila cassinopsisi (Risbec, 1952) comb. nov. (from Decatoma), Madagascar

Sycophila hilla Watsham (1977), Zimbabwe-Rhodesia

Sycophilla kestraneura (Masi, 1917) comb. nov. (from Decatoma), Seychelles

Sycophila nigrofasciata (Risbec, 1952) comb. nov. (from Decatoma), Madagascar

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 215

Sycophila ruandensis (Risbec, 1957) comb. nov. (from Decatoma), Rwanda

Sycophila rubra (Risbec, 1952) comb. nov. (from Decatoma), Madagascar

Sycophila xiphigaster (Risbec, 1955c) comb. nov. (from Decatoma), Kenya

Dr. J. R. Steffan of the Paris Museum (MNHN) kindly enabled me to examine

most of Risbec’s species. One of them, S. ruandensis, is extremely close to S.

aethiopica, of which I could examine a syntypic couple coming from Nefasit,

Eritrea, thanks to Prof. Dr. G. Viggiani of Portici: the female is here designated

(and was accordingly labelled by me) as lectotype. The two may be just forms of

one species, the main difference being more compressed female gaster in

aethiopica, apart from colour and extension of the sub-parastigmal patch of

hairs, which is certainly prone to intraspecific variation. Also, S. aethiopica

seems to be distributed over most parts of Africa. Most of these species are

probably never associated with figs, only S. kestraneura apparently is, and

probably also S. xiphigaster. The newly described species are compared with the

old ones wherever it is relevant.

Decatoma plectroniae Risbec (1952) from Madagascar does not belong to

Sycophila. It has a percurrent carina bordering the gena and temple posteriorly.

Sycophila kestraneura (Masi)

(figs. 142, 143; pl. 4 fig. 5)

Decatoma kestraneura Masi, 1917: 138—139. Holotype ®, Seychelles (BMNH, London) [exam-

ined].

This seems to be a very variable species and it took me a long time to recognise

it from the small holotype specimen (in BMNH), a dwarf as I now believe, from

the Seychelles Islands. A redescription seems necessary.

Female. — 2.1—3.0 mm. Usually yellow with poor blackish markings in

median line of body, including ocellar area, occiput, median streak on thorax

expanding on base of propodeum to inside of wing bases, petiole and dorsal keel

of gaster. This pattern may be either reduced or slightly extended. Fore wing

with elongate fuscous spot on swollen marginal vein, spot expanding slightly

below the vein in large darker specimens.

Head dorsally about 1.85 times as broad as long; relative measurements:

breadth of frontovertex 27.5, short oval eye 21 x 18, malar space 13.5. Scrobes

ending about half diameter from ocellus; interantennal keel narrow and not

protruding, rather gradually sloping into scrobal cavity. Flagellum plus pedicel-

lus barely longer than breadth of head, subclavate (fig. 142); pedicellus about 1.4

times as long as broad, narrower than basal funicular segments first of which is

usually about 1.8 times as long as broad and distinctly longer than pedicellus (and

bearing two rows of longitudinal sensilla), the fifth quadrate; clava about twice

as long as broad.

Pronotum nearly as broad as mesoscutum, its sides in dorsal view moderately

rounded. Umbilicate punctures on thorax usually fairly dense, but less so in

smaller specimens. Axillar grooves anteriorly wide apart, only slightly inside of

notauli: base of scutellum slightly broader than each axilla. Propodeum with

conspicuous anterior converging keels, anterior to them (submedially) with

216 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

broad irregular areolae; median part shallowly convex, broad, anteriorly often

with indication of median carina; posterior quarter, to one-third, narrowed,

neck-like. Legs relatively slender, more slender than in S. naso. Fore wing

pubescence rather reduced, dorsal surface basally extensively bare, occasionally |

with 1 or 2 hairs on basal fold; costal cell dorsally bare, on ventral surface |

generally with one row of hairs in proximal third, expanding to about 3 irregular |

rows distally; marginal vein distinctly longer than the subhyaline stigmal vein,

swollen, its lower margin convex (fig. 143).

Petiole 1.5—1.8 times as long as broad (dorsally); gastral body fully as long as

thorax, smooth, strongly compressed from sides, with distinct dorsal keel.

Hypopygium ending shortly before middle of gastral body, its apex with usually |

two pairs of hairs.

Male. — 1.5—3.4 mm. Small specimens often similar to female in colour, but

all differ mainly in antennae, fore wing pilosity and veins, and in gaster. Dark |

colour often spreads with larger size, including fore wing macula which may then

be combined with brownish infumation of the blade proximad and distad of

speculum; also femora get much stouter with larger size.

Scapus and pedicellus hardly different from female, but flagellum is filiform, |

itself (less pedicellus) distinctly longer than breadth of head; all funicular |

segments elongate, with rather dense subdecumbent hairs, first segment cylin-

drical, always much longer than pedicellus, in larger specimens fully twice so;

following three segments subequal, each fully twice as long as broad; clava

nearly four times as long as broad. Pronotum expanding forwards in larger

specimens. Fore wing pilosity dense, long, extensive, especially long in basal

part; in small specimens marginal swelling not much larger than in female but in

larger specimens becoming more conspicuous, broader and further accentuated

by expanding infuscation which may become higher than broad, as in pl. 4 fig. 5, .

which also shows the form of gaster, including the rather slender curved petiole |

which is slightly longer than the scutellum.

Material. — S. kestraneura was reared from figs of Ficus burkei in Zimbabwe- |

Rhodesia, from F. petersii in South West Africa (Gross Otavi, xi.1973, A.

Watsham), from unidentified figs in Uganda (Kawanda, ii.1939, T. H. C.

Taylor) and collected also in South Africa: Pondoland, Port St. Johns, v.1923

(R. E. Turner). In Zimbabwe-Rhodesia the collected or reared material comes

from Chishawasha and Makumbi Mission nr. Salisbury and the Chinamora

Reserve (all A. Watsham).

Comments. — In Africa there seems to be another species very close to S.

kestraneura (Masi), with similarly shaped marginal vein but shorter funicular

segments and the hypopygium (in female) fairly long. Otherwise S. kestraneura,

among the African species, seems to be the one most closely related to the type-

species of the genus S. decatomoides Walker from India.

Sycophila punctum Bouéek, spec. nov.

(figs. 144, 145)

Female. — 1.8—3.1 mm (holotype 2.5 mm). Usually predominantly black,

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 217

with pronotum on shoulders broadly pale yellow, also antennae pale yellow;

hind coxa, femur and tibia mainly infuscate, on mid and especially fore legs pale

colour more extended; sometimes head yellowish, mainly ventrally, rarely also

most of pronotum, legs and even parts of ventral thorax and gaster anteriorly

pale. Wings hyaline, with fuscous spot small and usually round, not reaching pale

base of marginal vein.

Head finely reticulate, without umbilicate punctures; white pubescence

unusually short, subdecumbent, longer and more erect only at mouth margin. In

dorsal view head about 1.6—1.68 times as broad as long, with lateral ocellus

‚ nearer to eye than to median ocellus; the latter fully by its diameter from scrobes;

i interantennal crest not high but slightly tooth-like. Relative measurements:

| head width 47, length 28.5, height 37, frontovertex 25.5, eye 23 x 19, malar

space 12, scapus 19, flagellum plus pedicellus 46. Centres of antennal toruli on

lower ocular line. Pedicellus 1.7—1.8 times as long as broad, at least slightly

| longer (but not broader) than first funicular segment which is usually 1.5 times as

long as broad. Flagellar segments stout-filiform, preclaval one often quadrate;

flagellar pubescence subdecumbent.

Sculpture on thorax fine dense reticulation, surface not shiny; puncturation

obliterated, pubescence white, short and almost decumbent, absent from

central area of scutellum. Anterior edge of lateral panel of pronotum usually

finely carinate. Propodeal surface in profile convex in upper one-third. Hind

tibia with a row usually of 7—10 dorsal bristles, middle ones longer than half

breadth of tibia. Fore wing with basal third bare (as in kestraneura); marginal

vein gradually expanding, relatively short, about as long as the stigmal (fig. 145).

Gastral petiole dorsally (measured as usual from anterior cross-ridge) sub-

quadrate. Body of gaster about as long as thorax less propodeum, distinctly

compressed, tergites 3 to 5 distinctly keeled, fourth slightly longer than third and

with one short row of sparse hairs on either side; tergites with extremely fine and

shallow cross-striae.

Male. — 1.9—2.5 mm. Head extensively to completely yellowish; pronotum

dorsally yellow, except median triangle, but lower part of lateral panel abruptly

black, often other parts of thorax pale. Head very stout (fig. 144), but variable,

1.35—1.55 times as broad as long in dorsal view. Antennal toruli entirely below

lower ocular line. Pronotum distinctly longer than in ®, its collar fairly flat,

about 1.5 times as broad as laterally long. Fore and hind femora distinctly

thickened. Fore wing macula in one specimen extended down as curved streak.

Gastral petiole stout in lateral view, dorsally finely reticulate and about 2.1 times

as long as broad, slightly tapering forwards.

Type material. — Holotype ® (plus 7 ® and 1 d paratypes), Zimbabwe-

paratypes, xii.1976 (all A. Watsham); deposited in BMNH, London. Further

paratypes: Chishawasha nr. Salisbury, 73 9 2 dg, 1969—1977, mostly ex F.

burkei; St. Pauls Mission, 1 2, 1.1975 (A. Watsham). Some paratypes are

deposited in RMNH, Leiden and in the Watsham collection.

218 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Comments. — Sycophila punctum, as stressed in the key, is recognisable |

mainly on its pilose prepectus (fig. 144), further in the female on the usually,

predominantly black colour with large pale spots on pronotal shoulders, small |

wing macula and a stout head. The head apparently becomes stouter in larger

males, also the wing macula may be expanded. It is striking that so few males

were reared or collected, although in this species they seem to be quite safely

recognisable.

Sycophila sessilis Boucek, spec. nov.

(figs. 146, 148, 149)

Female. — 2.0—2.7 mm. Colour as in paler specimens of S. punctum, 1.e., |

with pale yellow shoulders but yellowish colour otherwise spread also over most

parts of body, but flagellum slightly brownish. |

Head dorsally slightly less than 1.8 times as broad as long. Interantennal ridge

gradually descending into scrobal cavity. Relative measurements: head width

48, frontovertex 25, eye 19.5 x 17 (smaller than in punctum), malar space 13.5,

scapus 20, flagellum plus pedicellus 51. All funicular segments elongate, the first

very slightly longer than pedicel (fig. 149).

Umbilicate punctures on thorax shallow and not very dense, but fairly distinct, |

pubescence longer than in S$. punctum. Exposed part of prepectus bare, occa-

sionally with 1 or 2 hairs. Fore wing with sparse bristles on basal fold, rarely with

only two or three; fuscous macula similar to that of S. punctum but marginal vein

longer (fig. 146).

Gastral petiole dorsally much broader than long, gaster almost sessile; its

body about as long as thorax, more strongly compressed than in S. punctum, |

surface smooth.

Male. — Not known.

Type material. — Holotype 9, Zimbabwe-Rhodesia: Chishawasha nr. Salis- |

bury, ix.1974 (A. Watsham); in BMNH, London. Paratypes: Chishawasha,

Turner).

Comments. — The species should be recognisable from the key and the

additional characters in the description. However, I could not find any males —

which I could safely associate with the females of S. sessilis.

Sycophila modesta Boutek, spec. nov.

(fig. 147)

Female. — 2.2—2.6 mm. Predominantly yellow, with tiny dark spots at ocelli —

and brown colour on flagellum, pedicellus, median part of propodeum, petiole, |

broad bands at hind margins of first three tergites, sides of fourth tergite and —

apical half of hind tibia. Dark markings may be slightly reduced or expanded.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 219

Fig. 147. Sycophila modesta Boutek, spec. nov., venation of female fore wing. Figs. 148, 149.

Sycophila sessilis Boutek, spec. nov., female. 148, propodeum, gaster and hind leg; 149, head,

tilted lateral view. Fig. 150. Sycophila flaviclava Boutek, spec. nov., female head in tilted lateral

view. Figs. 151—153. Sycophila naso Boutek, spec. nov., female. 151, head in a larger specimen

with a more rectangular interantennal lobe; 152, head in a smaller specimen with more rounded

interantennal lobe; 153, gaster.

Wing macula (fig. 147) as in S. naso.

In colour, size, sculpture, pubescence of fore wing and length of petiole very

similar to S. naso, but for following characters. Lateral ocellus slightly nearer to

220 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

eye than to median ocellus, interantennal ridge only gradually sloping into

scrobes; pedicellus up to 2.2 times as long as broad, subequal in length to first |

funicular segment; scutellum slightly broader at base (border with mesoscu- |

tum); propodeum rather flat in median part, ground nearly smooth, wide-

areolate, with stronger carinae delimiting a pentagonal area, with a cross-carina

separating the neck; costal cell dorsally in distal half usually with distinct line of

erect hairs (fig. 147; rarely reduced to several hairs only). Body of gaster broader

and shorter, only moderately compressed from sides, not distinctly keeled

dorsally.

Male. — Not known (I am not sure about their specific identity).

Makumbi Mission nr. Salisbury, ex F. burkei, xii.1976 (A. Watsham); in

BMNH, London. Further paratypes: Chishawasha nr. Salisbury, mostly ex F.

burkei, 12 2, ix. and xi.1974, iii. and vi.1975, xii.1976, vi.1977 (A. Watsham).

Comments. — This species seems to be safely recognisable in the females,

although similarly coloured females of other closely related species, e.g., of S.

naso, may occur. The wing pilosity seems to be a good character, but not for the

males in which larger and darker specimens of the same species often exhibit

much greater density of pubescence, along with broader veins, femora, etc.

Sycophila naso Boutek, spec. nov.

(figs. 151—153)

Female. — 2.3—3.5 mm (holotype 2.8 mm). Body usually yellow (as in

holotype) but colour variable: dark colour appearing first in a streak on dorsal

keel of gaster, then at ocelli, on lower occiput, pronotal collum, on deeper parts

inside wings, some specimens have also darker mesoscutum anteriorly and

scutellum posteriorly, in extreme case whole gaster and underside of thorax

including propodeum are dark, also partly mid and hind legs; flagellum brown- |

ish; fore wing with dark elongate spot on marginal vein, with about half of macula |

(or more) below the vein.

Puncturation, especially on thorax, rather dense and conspicuous; pubes- |

cence moderately short but distinct. Head dorsally about 1.75 times as broad as —

long, with temples rather strongly receding; POL nearly twice OOL; lateral

ocellus hardly nearer to median one than to eye; scrobes ending by blunt edge —

about half diameter before ocellus; interantennal keel high, fairly broad ventral- —

ly, dorsally forming a distinctly protruding subrectangular tooth (figs. 151, 152;

hence the name, naso = nose). Parascrobal area subequal in breadth to scrobes

at level where its straight edge turns towards ocellus. Pedicellus dorsally fully —

twice as long as broad (17 : 7), slightly to obviously longer than first funicular

segment (17 : 15 in holotype), its dorsal side nearly straight; fifth funicular

segment slightly elongate; clava about 2.5 times as long as broad.

Thorax about twice as long as broad (mesoscutum); pronotum slightly wid-

ening forwards, with subrectangular protruding corners, laterally longer than in

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 221

middle; lateral panel not distinctly carinate anteriorly. Mid lobe of mesoscutum

posteriorly narrow, scutellum still more narrowed anteriorly, axillae little apart.

Propodeum shorter than scutellum (about 3 : 4); its median depression shallow;

anterior diverging keel distinct, pentagonal median area subdivided into irregu-

lar areolae. Metasternal region medially usually with three distinct carinae

converging caudad into one. Legs fairly slender. Fore wing with distinct line of

bristles on basal and most of cubital fold, usually also some bristles on basal cell

(as in fig. 147); costal cell with numerous short hairs on ventral surface but dorsal

surface bare, occasionally with one hair; marginal vein broadening distad, its

lower margin almost straight.

Gastral petiole dorsally fully 1.5 times as long as medially broad (20 : 13 in

holotype), sides subparallel. Gastral body (fig. 153) slightly longer than thorax

(including propodeum), fairly strongly compressed from sides, dorsally with

blunt but distinct keel; epipygium very shortly protruding, usually also tips of

sheaths visible; tergites 1 to 5 virtually bare and smooth. Hypopygium (not

always visible) rarely exceeding basal quarter of gaster, at apex with two thin

long bristles, before them a pair of shorter hairs.

Male. — In size and colour and in most characters very similar to female.

Pedicellus dorsally about 3 times as long as broad, almost as long as, to slightly

longer than, first funicular segment (anellus excluded), latter slightly longer

than any of following three subequal segments; flagellum almost filiform, clava

as long as two preceding segments combined. Basal third of fore wing with many

more hairs than in female, hairs much denser in bigger and darker specimens;

ventral surface of costal cell with extremely dense short pubescence, distal half of

dorsal surface with a row of hairs which is usually partly doubled. Lower margin

of marginal vein almost straight (but looking slightly convex in certain lights in

darker specimens), relatively longer than in S. kestraneura, about twice as long

as the pale stigmal vein which is subparallel to front margin of wing; greater part

of fuscous macula often below the vein. Petiole dorsally slightly shorter than

scutellum.

bury, ex galls on F. burkei, v.1975 (A. Watsham); deposited in BMNH.

Paratypes: same origin as holotype, 65 2 and 52 d, ix.—xii.1974, ii., v.—

A. Watsham); Uganda: Kampala, ex galls on stem of Ficus sycomorus, 2 9,

8.x.1935 (T. H. C. Taylor); South West Africa: Gross Otavi, ex F. petersii, 1 9,

x1.1973 (A. Watsham). The females of the Chishawasha material emerged

between May and July are yellow-coloured, those emerged between September

and March are dark-coloured, whilst the dark males appear throughout the year,

although the yellow ones only together with yellow females. Some paratypes in

RMNH, Leiden and the Watsham collection.

Variation. — Apart from the range of variation already mentioned I find some

puzzling features in some further specimens (not included in the type material of

naso). In a few females the pedicellus is relatively shorter (fig. 152) and so it is in a

222 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

number of males. I wonder whether still another species may be involved, but as |

to the males, some of them may belong also to Sycophila sessilis or S. modesta,

whose males are not yet known for sure.

Comments. — S. naso is recognisable mainly on the relatively long and slender

pedicel (in both sexes), the strongly converging axillar grooves and the small

wing macula on the straight lower margin of the marginal vein. Furthermore the

female has a fairly long gastral petiole and hairy proximal part of the fore wing,

but virtually no hairs on the dorsal surface of the costal cell.

Another similar species is Sycophila cassinopsisi (Risbec), comb. nov. origi-

nally described in Decatoma (Risbec, 1952: 278—281) from Madagascar. Dr. J.

R. Steffan from the MNHN, Paris, kindly sent me the type material for

examination. I remounted most of the syntypes on cards and designated one

female (emerged 6.iv.1950) as lectotype and labelled accordingly. S. cassinopsi-

si is smaller but less slender than S. naso, has the puncturation on thorax much

more obliterated, the head is stouter, about 1.5 times as broad as long (dorsally

70 : 47), the sides of pronotum are parallel, the collar is regularly convex, in

middle almost as long as at sides, the pedicellus is distinctly broader than the

proximal half of flagellum and much longer than the first funicular segment. The

fore wing pubescence is short and rather reduced towards base, with only few

hairs on basal fold. The gastral body of female is relatively short (not the petiole)

and not keeled dorsally. By the stouter head it reminds one somewhat of

Sycophila punctum, but that species differs greatly in colour, the pilose prepec-

tus, the shorter marginal vein and petiole, etc.

Sycophila flaviclava Boutek, spec. nov.

(fig. 150; pl. 4 figs. 3, 4)

Female. — 2.0—3.2 mm (holotype 3.2 mm). Predominantly yellow, with

small black spots at occipital foramen, base of fore coxa, mid part of prepectus

(normally concealed), a T-shape pattern on propodeum, metasternum and

petiole; sometimes dark colour more spread, or less so (pl. 4 fig. 3); flagellum

slightly infuscate but most of clava (except base) yellow (hence the name).

Head shaped about as in S. naso but white pubescence distinctly denser,

shorter and more decumbent (this to lesser extent also on thorax); lateral ocelli

slightly nearer to eye than to median ocellus; eye distinctly larger than in S. naso

and its lower extremity distinctly subangular; parascrobal areas relatively

narrower, also temples and genae; interantennal crest descending slightly

obliquely into scrobes; antenna very slightly clavate, pedicellus dorsally only

1.75 times as long as broad and slightly shorter (7 : 8 in holotype) than first

funicular segment. Relative measurements: head breadth 54, length 30, height

(with clypeal lobes) 45, frontovertex breadth 29, eye 29 x 21.5, malar space 14,

scapus 24, flagellum plus pedicellus 52.

Thorax slightly more robust than in S. naso; median part of metasternum in

profile angular at level of anterior edges of hind coxae; metanotal hairline dense.

Gastral body distinctly less compressed than in S. naso, slightly shorter than

thorax less propodeum; dorsally not keeled.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 223

paler apically, pubescence slightly longer and more erect. Longest eye diameter

in ratio to malar space as 23 : 13; pedicel dorsally twice as long as broad, scarcely

longer than fourth funicular segment. Pronotum posteriorly hardly emarginate,

this margin subparallel to anterior edge of collar; black petiole dorsally fully

three times (28 : 9) as long as broad.

bury, 11.1977 (A. Watsham); in BMNH, London. Paratypes: same origin as

holotype, partly reared from F. burkei galled figs, 9 2, 1 6, ix.—xi.1974, iii. and

(A. Watsham).

Comments. — The description is based on difference between S. flaviclava

and S. naso. Both species belong to a group of closely related species which might

be called the naso-subgroup and includes also S. modesta and S. sessilis, whilst

the other two species treated here in detail, S. punctum and S. kestraneura,

belong to other subgroups.

ORMYRINAE

Ormyrus Westwood

Type-species: Ormyrus punctiger Westwood; by monotypy.

The species of this group possess a strongly sclerotised gaster, often bearing

characteristic rough sculpture (figs. 155—157). This sculpture (although not

present in all species) combined with the obliterated notaular grooves and the

virtually non-exserted ovipositor with which again a reduction of the epipygial

cerci (to low tubercles) is correlated, were regarded recently as important

enough to warrant the group a family status, i.e, independent of Torymidae.

Riek (1970: 921) even transferred them, on the basis of the reduced cerci, as a

subfamily to the Pteromalidae. However, an analysis of various critical charac-

ters seems to demonstrate strong links with Torymidae and thus they seem to be

best placed as their subfamily, as they were mostly in the past.

Several generic names have been proposed in this group, all based on the

number of the reduced proximal segments of the flagellum (anelli) and partly on

the gastral sculpture. As they do not denote any evidently natural groups, only

one genus is recognised, Ormyrus Westwood.

All Ormyrus species have associations with plant tissues, especially galls,

although a few species probably develop in grass stems. Their actual role within

the galls is not yet known, although they are supposed to be at least partly

entomophagous, parasitic, rather than feeding solely on plant tissues (see under

O. flavipes, below).

Three species were found in association with the galled figs of Ficus burkei:

Ormyrus watshami spec. nov., O. subconicus spec. nov. and O. flavipes spec.

nov. Otherwise the following species have been described from Africa south of

the Sahara and from Malagasy:

224 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

O. australis Risbec, 1957, from Madagascar

O. chevalieri (Risbec, 1955 b) comb. nov. (originally in Monobaeus), from Chad

O. decaryi (Risbec, 1955 a) comb. nov. (originally in Tribaeus), from Madagascar, but widely

distributed in southern Africa including Zimbabwe-Rhodesia

O. eugeniae Risbec, 1955 a, from Madagascar

O. harongae (Risbec, 1952) from Madagascar, originally described in Wania, but transferred to

Ormyrus by Risbec (1955a: 176) himself

O. ornatus (Risbec, 1951) from Senegal, originally described in Wania, but transferred to Ormyrus

by Risbec (1954: 535)

O. sculptilis Crosby, 1909, from Malawi (formely Nyasaland)

O. striatus Cameron, 1907, from South Africa (the single extant male type in BMNH, London, Type

Hym. 5—64, is here designated lectotype).

The new species are compared with the earlier described species following a

study of their types.

Ormyrus watshami Boucek, spec. nov.

(fig. 157)

Female. — 1.7—2.0 mm. Bright metallic blue (mainly thorax) to green

(vertex, occiput), dorsum of darker bluish gaster from apex of tergite 1 to base of

tergite 3 black; antennae dark brown but scapes, fore tibiae, all tarsi, partly also

knees, trochanters, mid tibiae and fore femora, testaceous. Wings hyaline.

Head fairly shiny, with sculpture weaker than in most known species of

Ormyrus, on vertex consisting of curved engraved cross-striae; hairs on vertex

reduced to one pair between posterior ocelli, two hairs at eye outside of posterior

ocellus and one hair in front of this ocellus; piliferous punctures very conspicu-

ous; parascrobal areas and lower face with very short and rather sparse white

hairs. Anterior margin of clypeus short, bilobed, with median incision. Occipital

carina situated low, near occipital foramen; a blunt edge slightly indicated not far

behind posterior ocelli. Relative measurements: head width 85, height 67,

length (thickness) 43, frons width 50, POL 24, OOL 8, eye 44 x 34, malar space

16, scapus 28 x 7, flagellum plus pedicellus 74. Pedicellus subglobose (beyond

basal constriction); first anellus very thin, second larger but still more than 3

times as broad as long; all six funicular segments transverse, the first slightly so,

the sixth twice as broad as long, each with one row of sensilla.

Thorax about 1.33 times as long as (mesoscutum) broad, strongly convex,

shiny. Mesoscutum finely but not very densely cross-striate, mid lobe mainly

bare, only posteriorly with 3—5 irregular pairs of bristles (fig. 157). Scutellum

fully 1.1 times as long as broad, dorsally with concentric striae, with only 4 pairs

of bristles leaving the disc broadly bare; apex of scutellar rim truncate-subemar-

ginate and jutting out over the steep smooth propodeum; latter with numerous

hairs on lateral callus. Mesopleuron almost smooth, bare; broad metapleuron

with a few hairs. Dorsal edge of hind tibia with a row of about 10 strong bristles

and more outside with another row of slightly shorter bristles; longer spur

curved, its basal half distinctly hairy. Fore wing unusually sparsely pubescent,

basal third including large speculum (which extends on dorsal surface to stigmal

vein) bare, but usually with some hairs in place where basal fold approaches

cubital fold.

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 225

Gaster not or hardly longer than thorax, fairly compressed from sides, with

distinctly up-turned epipygium; tergites 2 to 5 with simple keel in median line.

Hind margin of first tergite medially excised; second tergite short, punctured;

third tergite longer, in basal half sublaterally with full 3 rows of coarse puncta

(rows interrupted in median line), apex with simple puncturation (as most

tergites); fourth tergite sublaterally hardly showing any puncta (they are mostly

Figs. 154-156. Ormyrus flavipes Boutek, spec. nov., female (154, 155) and male (156). 154,

venation of fore wing; 155, gaster; 156, male gaster, dorsal aspect. Fig. 157. Ormyrus watshami

Boutek, spec. nov., female. Fig. 158. Ormyrus subconicus Boutek, spec. nov., female gaster with

hind leg.

226 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

concealed by third tergite). Hypopygium not prominent, not exceeding middle

of gaster.

Male. — 1.6—1.9 mm. Bright brassy green but sides of thorax including

tegulae testaceous, as well as all legs and antennal scapes; rest of antenna more

brownish; usually also a broad indefinite cross-band on gaster just before middle

testaceous.

Head and thorax very much as in female but anelli more conspicuous; fore

wing pilosity denser and more conspicuous, speculum slightly reduced, reaching

only two-thirds of marginal vein. Gaster flat, elongate, its sculpture generally

weaker than in @; third and fourth tergites basally with large puncta (pits)

arranged in very irregular rows, more than two of which are usually exposed on

the third, and less than two rows on the fourth tergite.

Type material. — Holotype 9, Zimbabwe-Rhodesia: Makumbi Mission nr.

Salisbury, ex Ficus burkei galls, xii.1976 (A. Watsham); deposited in BMNH,

Chishawasha nr. Salisbury, x. and xii.1974, iii. and v.1975, ili. and ix.1976, vi.

and x.1977.

Comments. — Ormyrus watshami can be immediately recognised in both

sexes by the reduced pilosity on the dorsum of the head and thorax, leaving the

mid lobe of mesoscutum bare, except for a few hairs posteriorly. This is unique

among all other known species of the genus. In addition, the female has a

characteristic short form of the gaster (fig. 157) and the male is distinct by the

combination of the gastral sculpture and the widespread pale colour of the body.

Otherwise it belongs to the punctiger-group.

Ormyrus flavipes Boutek, spec. nov.

(figs. 154—156; pl. 4 fig. 6)

Female. — 2.1—4.3 mm (holotype 4.1 mm). Bright metallic green, partly

more golden, especially on genae and sides of pronotum, but on hinder part of

thorax and its sides more bluish to violaceous; violaceous mainly on dorsal edge

of hind coxa, on dorsal part of first tergite and on the coarsely sculptured basal

parts of tergites 3 to 5; epipygium dark purpureous; testaceous are: antennal

scapes, all legs including fore coxae and distal parts of mid and hind coxae, then

tegula with subalar area, partly lateral panel of pronotum and underside of

gaster from where the pale colour usually spreads broadly dorsad, at least along

hind margins of tergites, especially the fifth and sixth; rest of antenna black.

Wings hyaline.

Head and thorax dorsally fairly dull, generally cross-striate, rather densely

clothed with subdecumbent pubescence. Piliferous punctures on vertex little

conspicuous in dense cross-striation; an edge behind ocelli, occipital carina

situated close to foramen. Face, except for the rather deep and smooth scrobes,

with dense rugose striation; on lower face a slightly raised semicircular carina in

half distance between toruli and mouth (separating also clypeus from supracly-

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 227

peal area), at sides ascending towards eye margins. Gena in a strip just behind

malar groove very finely but distinctly striate; posteriorly striation much coars-

er. Lobes of clypeal margin low, separated by shallow emargination. Relative

measurements: head width 111, height 83, length (thickness) 52, frons width 60,

POL 24, OOL 12, eye 55 x 44, malar space 24, distance between toruli and

mouth margin 31, scapus 33 x 10, flagellum plus pedicellus 101. Pedicellus

dorsally nearly twice as long as broad (13 : 7); first anellus thin, second nearly

half as long as broad; first funicular segment about 1.1 times, last (sixth) about

1.5 times as broad as long, each with dense row of sensilla; clava only 1.7 times as

long as broad.

Thorax dorsally about 1.45 times as long as broad; its sculpture on mesoscu-

tum consisting of cross-striation' formed by numerous fine strips with mostly

raised hind margins, with hairs arising on hind slope of raised margin. Scutellum

about 1.15 times as long as broad, its apex rounded; anteriorly it is cross-striated

with piliferous punctures, posteriorly striae are arranged concentrically. Propo-

deum subvertical, short, medially smooth, submedially alutaceous. Hind tibia

dorsally with many rather short bristles which are only partly arranged in about

three irregular rows. Fore wing with rather dense pilosity, cubital hairline

reaching base of wing, basal hairline complete, basal cell with a dorsal row of

bristles subparallel to submarginal vein; costal cell dorsally bare but ventrally

with a hairline which is trebbled in distal third; stigma close and subparallel to

postmarginal vein (fig. 154).

Gaster fully twice as long as thorax, fairly compressed from sides, with simple

median keel distinct from tergite 4 to saddle of tergite 6 (fig. 155); distal third of

gaster conical and moderately up-turned; tergites 3 to 5 each punctured, with

basal part raised and ending by a vertical row of short ridges (each ridge with a

bristle at caudal tip), virtually without any exposed foveae; on sixth tergite the

ridges shortened into tubercles and additional weaker tubercles are scattered on

basal part; ridges on fourth tergite mostly longer than distance between them.

Sculpture as wel as dimensions indicated in fig. 156.

Male. — 1.4—2.4 mm. Gaster wholly metallic, long oval (fig. 156), slightly

longer than thorax to as long as head plus thorax combined; first tergite dorsally

with broad-meshed reticulation; second very short; tergites 3 to 5 each with dark

purple basal band of deep foveolae arranged in at least 2 rows, with smooth

narrow interspaces.

Biology. — After finishing this paper one of us (A. W.) observed a white larva

attached to the pupa of Syceurytoma ficus in a galled fig. He separated the

parasitised pupa from the rest and after a few days the parasite pupated and

proved to be Ormyrus flavipes.

bury, ex F. burkei galls, vi.1975 (A. Watsham); deposited in BMNH, London.

Paratypes: 16 2, 12 d, same data as holotype; 12 9 and 20 d, same origin, ii.—

ix.1975, 1 d ix.1976; 1 d Makumbi Mission nr. Salisbury, xii.1976 (A.

Watsham).

228 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL.5, 1981

Comments. — Among the African species known to me (described or

undescribed) several species belong to a species-group of O. orientalis Walker

characterised by the double carina on at least some gastral tergites in the females,

with generally rough sculpture of gaster in both sexes. From the African species

listed above O. eugeniae Risbec, O. ornatus (Risbec) and O. striatus Cameron

belong to this orientalis-group and can be therefore easily separated from O.

flavipes. Another easily separable species is O. decaryi (Risbec) belonging to the

species with no dorsal keel on the gaster in females. Our O. flavipes, with its

simple keel on the female gaster, belongs to the group of O. punctiger Westwood

(a European species), along with the African O. chevalieri (Risbec) and O.

sculptilis Crosby, and the Malagasy O. harongae (Risbec). O. chevalieri is easily

recognisable by its short rounded female gaster. O. harongae, of which I

examined the two syntypes (mounted on a slide) thanks to my colleagues from

the Paris MNHN, differs from O. flavipes mainly by the predominantly yellow-

ish body with the apical part of the female gaster still more produced and with

shorter ridges in rows on the tergites. On the other hand O. sculptilis, of which I

was enabled to examine and remount the original material (thanks to my

colleagues in the Cornell University in Ithaca) and selected and labelled a female

in good condition as lectotype (here designated), has the conical apical part of

the female gaster much shorter than in O. flavipes, and not up-turned. The male

can be more easily confused and the assessment of its diagnostic characters has to

await a revision of the African species.

Ormyrus subconicus Boucek, spec. nov.

(fig. 158)

Female. — 1.7—3.3 mm (holotype 2.5 mm). Mainly bright green, sometimes

slightly bronze or brassy, but on head and thorax often in places bluish to

violaceous or even purpureous, on gaster foveolate bases of middle tergites

usually dark violet or purpureous; scapes (except apically), tegulae, trochant-

ers, knees and tarsi testaceous, but often pale colour spreading to whole tibiae

and most of femora, then with only hind femora mainly metallic. Wings hyaline.

Sculpture and form of head and thorax much as in smaller specimens of O.

flavipes, but lateral ocelli slightly nearer to eyes, POL nearly 3 times OOL;

scapus slender, about 4 times as long as broad.

Gaster (fig. 158) about 1.3 times as long as head plus thorax combined, only

slightly compressed from sides, with simple dorsal keel. Basal part of tergites 3, 4

and 5 with distinct exposed belts of foveolae, usually in two rows on each tergite;

tergite 6 usually as long as the fifth, its saddle very shallow; epipygium much

shorter than preceding sixth tergite, not or scarcely up-turned.

Male. — 1.4 mm. Very similar to male of O. flavipes but hind femur distinctly

infuscate, antennal pedicellus subglobose, i.e., beyond its basal constriction

only slightly longer than broad (more evidently so in flavipes) and tergites 4 and 5

each with full two rows of basal foveolae (irregular and not fully two rows in

flavipes, fig. 156).

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna 229

Type material. — Holotype 9, Zimbabwe-Rhodesia: Makumbi Mission nr.

Salisbury, F. burkei, xii.1976 (A. Watsham); in BMNH, London. Paratypes: 1

xi.—xii.1974 (A. Watsham).

Comments. — This species also belongs to the punctiger species-group and,

being close to O. flavipes, the above description stresses the diagnostic charac-

ters in relation to that species. This species-group is rich in species in Africa.

Among the previously described species the nearest is O. sculptilis Crosby which

differs from O. subconicus mainly in the still less saddled sixth tergite in the

female, with longer fifth tergite exposing usually three regular rows of foveolae

(in this it differs also from O. flavipes). I find these more extensive foveolate

bands also in two males reared from figs of F. petersii in South West Africa and

therefore regard them as probably specifically different from O. subconicus. It

seems that as to the host-specificity possibly O. watshami is a typical fig-asso-

ciated species, whilst O. flavipes and O. subconicus may develop also in some

other galls or gall-like plant parts. However, the questions of the host-specificity

have been hardly considered in the African chalcids so far.

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232 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5, 1981

INDEX OF GENERA AND SPECIES (MAINLY CHALCIDOIDEA)

Aepocerus 158

aethiopica 214

affinis 155

africana (Otitesella) 191

africana

(Philotrypesis) 175

africanus

(Camarothorax) 157,194

alata 153, 157, 160, 162,

[189

Alfonsiella 156

Allotriozoon 156, 167,

[169

allotriozoonoides 167

Anastatus 153

anguliceps 155, 188

Apocrypta 156

appendiculatus 157

arabicus 157

Arachonia 156

aterrima 157

atrata 168, 169

australis (Ormyrus) 224

australis (Sycoryctes) 157

bakeri 157

bannensis 209

barbarus 152, 157, 160,

[163, 170

bethyloides 155, 194

bimaculata 157

Blastophaga 156

borneensis 156

braconidis 207

brevimucro 153, 157, 163,

[203

brevitarsis 157

brongersmai 152, 157,

[160, 164, 168

bruchocida 209

Bruchophagus 209, 211

caelebs 183

Callimomus 194

callosa (Grasseiana) 157

callosa (Sycoryctes) 157

caricae 153,157

Camarothorax 156, 194,

[197

cassinopsisi 214, 222

Ceratosolen 155, 156

chevalieri 224, 228

coccothraustes 183

corneliae 156

cornutus 153, 157, 159,

[164, 180

cowani 157

crenulatus 155

Critogaster 158

Crossogaster 156, 169

curtivena 153, 157, 161,

[212

dagatiguyi 183

decaryi 224, 228

Decatoma 156, 214

decatomoides 156, 214,

[216

Desantisca 207

Dolichoris 156

Elisabethiella 156

enriquesi 164, 167

epicarioides 191

Epichrysomalla 158

equicollis 153, 157, 163,

[197

errata 156

erythraea 175

Eudecatoma 214

eugeniae 224, 228

Eujacobsonia 168

Eukoebelea 156

Eupelmus 153

Eupristina 156

Eurytoma 155, 156, 205,

[208, 214

ficicola 157

Ficomila 156, 209, 211

ficus 153, 157, 161, 208,

[209, 227

ficusgallae 153, 157, 161,

[205

finitimorum 157, 175, 188

flaviclava 153, 157, 162,

[222

flavipes 153, 157, 160,

[224, 227

froggatti 157

fusciceps 157

galili 157

gambiensis 153, 157, 161,

[213

gestroi 157

gibbus 157, 180

gigas 157

gracilis 157

Grandiana 156

Grasseiana 156

guruti 155

harongae 224, 228

Heterandrium 158

heterandromorphum

[152, 169

hilla 214

hilli 156

hirtus 153, 157, 160, 164,

[186

Idarnes 156

imerinensis 194

Indothymus 155

infida 155, 188, 189

insignis 155

keralensis 157

kestraneura 153,157,

[162, 215

Kradibia 156

larvalis 157

latrodecti 207

Leeweniella 158

liodontus 168, 170

Lipothymus 168

lomaensis 157,183

longicornis 175, 188

longimucro 153, 157, 163,

[201

longipalpus 157

longiscapa 152, 157, 160,

[164, 168

longitarsus 157

malabarensis 157

masii 156

masoni 155

Micranisa 156

modesta 153, 157, 162,

[218

monilifera 155,188

moniliformis 155

Monobaeus 224

montis 157,180

naso 153, 157, 162, 220

nigrofasciata 214

nodularis 207

nota 157

notus 157

obscurus 194

Odontofroggatia 196

odorans 152, 157, 160,

[163, 170

orientalis

(Camarothorax) 194

orientalis (Ormyrus) 228

Ormyrus 156, 223

ornatus 224, 228

Otitesella 156, 191

Paradecatoma 209

Parakoebelea 155, 156

Parasitodiplogaster 152-

[154 (nematod)

Z. BOUCEK, A. WATSHAM & J.T. WIEBES: Fig wasp fauna

parca 153, 157, 159, 164,

[176

patellaris 157,182

Pediobius 153

Pembertonia 156

Phagoblastus 152, 156,

[169

Philocaenus 169

Philotrypesis 156, 175

Philotrypomorpha 175

Physothorax 158

Platyscapa 156

plectroniae 215

Pleistodontes 156

‚ prodigiosum 157

psenes 157

pteromaloides 155

punctiger 223, 228

punctum 153, 157, 162,

[216

quadraticeps 155

quadrisetosa 157

raoi 157

reflexa 169

religiosae 155

remus 153, 157, 160, 164,

[183

reticulatus 156

Risbecoma 209

royi 157,191

ruandensis 215

Acacia 209

Adenosperma 155

avi-avi 183

baroni 169

benghalensis 155

burkei 149-229

callosa 157

capensis 154

carica 157

Conosycea 155

craterostoma 150

eriobotryoides 157,175,

[180, 183, 191

Ficus 155,157

exasperata 157

Galoglychia 157

rubra 215

saptapurensis 155

sculptilis 224, 228

sebertianus 183

selenitica 175

serricornis 153, 157, 163,

[196

sessilis 153, 157, 162, 218

silvestrii 168, 169

similis 157

sjoestedti 183

socotrensis 169

spinitarus 157

stabilis 155

stilifera 156, 188

stratheni 157

striatus 224, 228

stuckenbergi 151,157,

[160, 164, 165

subconicus 153,157,161,

[228

Syceurytoma 156,208,

[211

Sycobia 156,194

sycomori

(Eukoebelea) 157

sycomori (Sycophaga) 157

Sycophaga 156,168

Sycophila 156,205, 208,

[211, 214

Sycophilodes 156

INDEX OF PLANT NAMES

Kalosyce 155

laterifolia 157,180

leprieuri 169

Malvanthera 157

macrophylla 157

natalensis 150, 211, 213

nautarum 169

Neomorphe 157

nota 157

Oreosycea 155, 157

petersii 150, 169, 180,

[183, 185, 193, 200, 213,

[216, 221, 229

racemosa 157

reflexa 169

religiosa 155, 189

Sycophilomorpha 156

sycophilon 152-154

[(nematod)

Sycorycteridea 156, 179

Sycoryctes 155, 156, 179,

[182

Sycoscapter 156, 179, 180

Sycoscapteridea 156, 179,

[188

Sycotetra 156, 195

Systole 209, 211

temeraria 155

testacea 157

Tetramesa 209

Tetrastichus 153

Tineomyza 214

transiens 155

Tribaeus 224

triformis 168, 170

truncatus 183

tsamvi 153, 157, 161, 164,

[191

Walkerella 156

Wania 224

Waterstoniella 156

watshami 153, 157, 160,

[224, 229

Watshamiella 156, 186

westwoodi 157

xiphigaster 215

Rhizocladus 155

Sinosycidium 155

soroceoides 180

Stilpnophyllum 155

stupenda 156,188

Sycidium 157

Sycocarpus 157

Sycomorus 157

sycomorus 157,211,213,

4 [221

thonningii 149, 150, 155-

[157, 159

Urostigma 155

variegata 157

vasta 169, 183

vogelii 152, 169, 175, 191

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TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5 PL. 1

Fig. 1, Elisabethiella stuckenbergi (Grandi), female; 2, Crossogaster odorans Wiebes, spec. nov., fe-

male; 3, Elisabethiella stuckenbergi (Grandi), male; 4, Crossogaster odorans Wiebes, spec. nov., male;

5, Otitesella tsamvi Wiebes, spec. nov., male; 6, do., female.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5 PL. 2

„5 6

Fig. 1, Philotrypesis parca Wiebes, spec. nov.; 2, do., male; 3, Sycoscapter cornutus Wiebes, spec.

nov., female; 4, Sycoryctes remus Wiebes, spec. nov., male; 5. Watshamiella alata Wiebes, gen. et

spec. nov., female; 6, do., male.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5 PL.3

5 6

Fig. 1, Philotrypesis parca Wiebes, spec. nov., alate male; 2, Sycotetra serricornis Boutek, gen. et

spec. nov., female; 3, Camarothorax equicollis Boutek, spec. nov., male; 4, Camarothorax longimu-

cro Bouéek, spec. nov., female; 5, Camarothorax brevimucro Boutek, spec. nov., male; 6, Euryto-

ma ficusgallae Boutek, spec. nov., female.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 5 PL. 4

Fig. 1, Syceurytoma ficus Boutek, gen. et spec. nov., female; 2, do., male; 3, Sycophila flaviclava

Boutek, spec. nov., female; 4, do., male; 5, Sycophila kestraneura (Masi), male; 6, Ormyrus flavipes

Boutek, spec. nov., female.

USE

DEEL 124 AFLEVERING 6 VU

1981

COMP. ZOOL

LISRARY

WOE

TIJDSCHRIFT sur

CMNVYERSITY

VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

MUS

sl.

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

INHOUD

J. KRIKKEN, C. VAN ACHTERBERG, P. H. VAN DOESBURG, R. DE Jong and K. W. R.

ZWART. — Samuel Constant Snellen van Vollenhoven (1816-1880) and his

entomological work, p. 235—268, figs. 1—3, one colour-plate.

Tijdschrift voor Entomologie, deel 124, afl. 6 Gepubliceerd 31-XII-1981

SAMUEL CONSTANT SNELLEN VAN VOLLENHOVEN

(1816-1880) AND HIS ENTOMOLOGICAL WORK

J. KRIKKEN, C. VAN ACHTERBERG, P. H. VAN DOESBURG,

R. DE JONG!) AND K. W. R. ZWART?)

With one plate and three text-figures

Hat sich nun ein Entomolog um unsre Wissenschaft nicht bloss in seinem

engern Vaterlande und amtlichen Wirkungskreise, sondern auch nach vielen

und bedeutenden Richtungen im Grossen und Ganzen so hochverdient

gemacht wie Samuel Constant Snellen van Vollenhoven, so gebührt ihm ein

Zeugniss der dankbaren Anerkennung.

C. A. Dohrn, 1881,

Stett. ent. Ztg. 42: 371

ABSTRACT

The significance of the entomological work of S. C. Snellen van Vollenhoven (1816-1880) is descri-

bed by means of a concise biography, an annotated bibliography, and an annotated list of taxa he pro-

posed. The bibliography includes 184 items. The list of taxa includes the names of 9 genera and 471

species proposed by Vollenhoven in the insect orders Dictyoptera, Orthoptera, Hemiptera, Coleopte-

ra, Diptera, Hymenoptera and Lepidoptera.

CONTENTS

RTÉLACE RE TOA IR E TNA NO A CR IA A IONI 235

BIOLrAPhy: SR N Een 239

Bibliography i I LO SR eenen 240

PIstofgenus-prOUpiandspecies:SIOUPMAMESERT RT TO I 248

Indexofisenus-group,and'species-SToUp names Re RT O TT 263

PREFACE

Now that we are compiling this account of the entomological activities of S. C.

Snellen van Vollenhoven?) a century has passed since he died: the first truly scien-

tific entomologist in the Netherlands. Others had preceded him (e.g., Cramer,

Stoll, Voet, Merian, Sepp), but their contributions to Dutch entomology were li-

mited from a scientific point of view. Vollenhoven’s achievements still keep us

1) Rijksmuseum van Natuurlijke Historie, Leiden

2) Dept. of Entomology, Agricultural University, Wageningen

3) The full family name Snellen van Vollenhoven is in this paper abbreviated Vollenhoven. Vollenho-

ven should not be confounded with the Dutch lepidopterist P. C. T. Snellen (1832-1911).

235

236 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Fig. 1. Samuel Constant Snellen van Vollenhoven, around 1870.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 237

L00 Hat tH |

a. oh ark Le

—

Soa) n

Fig. 2. Handwriting of Snellen van Vollenhoven. Section of a letter written in 1860.

238 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Fig. 3. Stone plate in wooden carved frame, commemorating Snellen van Vollenhoven’s decease in

1880. It shows the symbols of two of his life-time’s ambitions: a butterfly (top of frame) and a sketch-

book and drawing-pen (bottom). Rijksmuseum van Natuurlijke Historie, Leiden.

busy, were it alone for the fact that, as museum curators, we are still dealing with

the numerous insects he assembled and described. Therefore, we thought it a

good idea to spend part of 1980, the centennial year of his death, on writing an

account of Vollenhoven’s entomological work. This account consists of a concise

biography, a bibliography and a list of the genera and species he described. Spe-

cialists interested in Vollenhoven’s work will find numerous verified details in the

following pages; they will, however, not find solutions to any taxonomic problems

they might have with respect to Vollenhoven’s species. The list of taxonomic na-

mes given below is only intended as a preliminary survey of Vollenhoven’s taxo-

nomic activities. We hope that the present account will show our fellow entomolo-

gists, besides technical details, a man at the roots of Dutch entomology: to Vol-

lenhoven life was nearly synonymous with the study of insects.

We are indebted to the Uyttenboogaart-Eliasen Foundation for financing the

publication of plate 1.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 239

BIOGRAPHY

Samuel Constant Snellen van Vollenhoven was born in Rotterdam on 18 Octo-

ber 1816. At the age of 18, after his secondary school education, he started to stu-

dy law at the University of Leiden and graduated on 25 September 1839. Soon

after his graduation he registered as a lawyer in The Hague, but there is no evi-

dence that he ever actually practised. From his childhood Vollenhoven appeared

to be more interested in the natural sciences, and, consequently, after his marria-

ge with Jeanne Everdine Meijer, he moved to Leiden again with the explicit pur-

pose of acquiring a more serious knowledge of animals and plants. He frequently

visited the Rijksmuseum van Natuurlijke Historie and the Hortus Botanicus, and

was delighted in conversing with the zoologists and botanists working in these in-

stitutions. After this second Leiden period Vollenhoven, disliking city life, took

up his residence at the Gliphoeve, a country seat south of Haarlem. Here he

spent, according to his own saying, the happiest years of his life. Vollenhoven

now devoted himself almost entirely to insect studies. An early result of these stu-

dies is his publication on insects harmful or beneficial to horticulture (1843a).

Before Vollenhoven’s time Dutch entomology had been largely the domain of a

few amateurs and non-specialist biologists, and the scanty knowledge available on

the Dutch insect fauna was totally unreliable. This changed drastically through

the efforts of Vollenhoven: soon after its foundation in 1845 he joined the Neder-

landsche Entomologische Vereeniging (Netherlands Entomological Society), and

immediately impressed his fellow members with his scientific approach to entomo-

logy. They elected him President of the Society, an office he held till 1872, and

again from 1878 till his decease in 1880. In 1858 he took the initiative to publish a

Society journal, the Tijdschrift voor Entomologie, which was profusely illustra-

ted, in the first place by Vollenhoven himself. Of the 277 plates published in the

journal up till 1880 (23 vols.) no less than 155 had been prepared by Vollenhoven,

who shared his artistic ambitions with contemporary entomologists, like J. O.

Westwood, of Oxford, with whom he had a very intense correspondence.

The government recognized Vollenhoven’s capacities by appointing him ento-

mological curator of the Leiden museum in September 1854. What he found there

seems to have been a mess, which was partly due to the protracted illness of his

predecessor Dr. W. de Haan. The notorious C. A. Dohrn, of Stettin, on visiting

Leiden shortly after Vollenhoven’s appointment, noticed the curator’s embarrass-

ment over the state the collections were in. Vollenhoven, however, set to work

energetically, and Dohrn clearly appreciated the improvement during a later visit,

as he mentioned in Vollenhoven’s obituary (1881; see quotation above). Vollen-

hoven’s private collection was purchased by the museum and incorporated. His

appointment necessitated Vollenhoven to move to Leiden once again. His cura-

tor’s salary was not particularly high, and the resources of the museum were also

very limited. Luckily, Vollenhoven was financially more or less independent, and

he regularly spent private money to pursue his entomological endeavours. While

continuing his work on the Dutch insect fauna, Vollenhoven’s attention was also

drawn to the exotic treasures kept in the Leiden museum. He described many no-

velties from the Dutch East Indies and other parts of the world, partly in well-il-

lustrated monographs. From 1871 on Vollenhoven had rheumatic complaints, and

240 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

considered that resignment from his curatorial duties would improve his health. In

1873 he was succeeded by his able assistant C. Ritsema. Bureaucratic decisions

barred Vollenhoven from further pursuing his survey of the East Indian insect

fauna, and his attention now shifted to the Dutch Ichneumonoidea and Hemipte-

ra, although his broad interest remained. He died on 22 March 1880, at the age of

63.

Indeed, Vollenhoven was the first truly scientific Dutch entomologist: reliable

literature on the Dutch insect fauna was virtually non-existent until he compiled

the first critical lists of Dutch Coleoptera (see bibliography, 1848), Diptera

(1852c, 1853a), Hemiptera (1852d), Hymenoptera (1858a), Orthoptera (1859g),

and their various sequels. He was keenly interested in economic entomology (cf.

1843a, 1852a). He published dependable popular surveys of Dutch arthropods

(1859f, 1861g) and of insects in general (1870d). Vollenhoven described the meta-

morphoses and habits of Dutch Symphyta with numerous fine illustrations (1858g,

etc.). His other important contributions to entomology include monographs of

East Indian insect groups (1863k, 18651, 1868f), a treatise on Dutch Hemiptera

(1878j), and the Pinacographia (1875, etc.), being an atlas of Hymenoptera. All

in all we counted 480 new names proposed by Vollenhoven, as follows:

genera species genera species

Dictyoptera 1 1 Coleoptera 2 97

Orthoptera 1 3 Hymenoptera 5 103

Heteroptera 1 155 Diptera 1 15

Homoptera 0 9 Lepidoptera 0 88

Vollenhoven’s scientific efforts were rewarded by memberships of several fo-

reign learned societies. In 1860 he became a member of the Koninklijke Acade-

mie van Wetenschappen (Royal Academy of Sciences). In April 1862 the Univer-

sity of Groningen bestowed on him the title of Philosophiae naturalis doctor hono-

ris causa.

An extensive obituary was given by F. M. van der Wulp, 1880, Tijdschr. Ent.

24: LXXXIX-CVII, in Dutch, with a portrait.

BIBLIOGRAPHY

All publications with separate title headings are included. Casual observations,

not published under a separate heading and without any substance, can be found

in meeting reports of the Netherlands Entomological Society as published ın the

Tijdschrift voor Entomologie; these have been omitted (we have a list of 35 addi-

tional items). Other minor contributions (e.g., to a dictionary) have also been

omitted. Annotations are added where necessary, especially translations of titles

in the Dutch language.

1843a. De schadelijke insecten in tuinen met de middelen tot derzelver verdelging, benevens eene

opgave der voor tuinen nuttige insecten, p. 1—128, pls. I—5. — Rotterdam. (Treatise of no-

xious and beneficial insects in gardens).

1843b. Over de larve van de Cimbex lucorum. — Tijdschr. Natgesch. Phys. 10: 97—102, figs. 1—6.

(Description of larva).

1844.

1848.

1852a.

1852b.

1852c.

1852d.

1853.

1854a.

1854b.

1854c.

1855.

1856a.

1856b.

1858a.

1858b.

1858c.

1858d.

1858e.

1858f.

1858g.

1858h.

1859a.

1859b.

1859c.

1859d.

1859e.

1859f.

1859g.

1860a.

1860b.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 241

Beschrijving eeniger larven van Tentredinidae. — Tijdschr. Natgesch. Phys. 11: 157—163.

(Descriptions of larvae).

Bijdrage tot de fauna van Nederland. Naamlijst van de schildvleugelige insecten, p v + (1) +

1—50. — Haarlem (His first checklist of the Dutch Coleoptera; cf. 1854c, 1870e).

De insecten welke den landbouwer schaden, p. 1—136, figs. 1—69. — Arnhem. (Noxious

insects in agriculture; also published in 1856 by Kruseman at Haarlem).

Nederlandsche blad- en houtwespen. — Bouwst. Faun. Ned. (Herklots) 1: 104-112. (Dutch

Symphyta).

Naamlijst van inlandsche Diptera. — Bouwst. Faun. Ned. (Herklots) 1: 138—151 (Checklist

of the Dutch Diptera, with F. M. v. d. Wulp).

Nederlandsche Hemiptera. — Bouwst. Faun. Ned. (Herklots) 1: 169—186; (with H. W. de

Graaf; Dutch Hemiptera).

Naamlijst van inlandsche Diptera. — Bouwst. Faun. Ned. (Herklots) 1: 188—206. (2nd part

of checklist, cf. 1852c; with F. M. v. d. Wulp).

Over het tweede deel van den “Schouburg der rupsen, wormen, maden en vliegende dier-

kens daaruit voortkomende” van Steph. Blankaart. — Alg. Konst- en Letterbode 66: 159—

160. (About a manuscript by S. Blankaart).

(Discussion about the genus Canopus (Hemiptera)). — Hand. Ned. Ent. Ver. 1: 63—64.

Naamlijst van de Nederlandsche schildvleugelige insekten. — Bouwst. Faun. Ned. (Her-

klots) 2: 1—70. (Checklist of the Dutch Coleoptera).

Mierengasten. — Jaarb. Natura Artis Magistra: 140—148. (Popular short paper on ant guests

in the province of Gelderland).

Nieuwe bijdragen voor de Fauna van Nederland. — Alg. Konst- en Letterbode 68: 178.

(Short faunistical list of Coleoptera and Hemiptera of the Netherlands).

Naamlijst van inlandsche Diptera. — Bouwst. Faun. Ned. (Herklots) 2: 89—117. (With F.

M. v. d. Wulp; checklist of Dutch Diptera).

Naamlijst van Nederlandsche vliesvleugelige insekten (Hymenoptera). — Bouwst. Faun.

Ned. (Herklots) 2: 221—275. (Checklist of Dutch Hymenoptera).

Beschrijving der nieuwe soorten. — Bouwst. Faun. Ned. (Herklots) 2: 276—283. (Descrip-

tions of new species of Hymenoptera).

Description de quelques espèces nouvelles de Coleoptères Lamellicornes. — Tijdschr. Ent.

1: 20—27, pl. 2.

Notes diptérologiques. — Tijdschr. Ent. 1: 88—93.

De rups van Hepialus sylvinus. — Tijdschr. Ent. 1: 98. (The caterpillar of H. sylvinus).

Phyllodes Verhuellii, nieuwe vlindersoort uit Java. — Tijdschr. Ent. 1: 159—168, pl. 8. (New

moth from Java).

De inlandsche bladwespen in hare gedaanteverwisselingen en levenswijze beschreven (1). —

Tijdschr. Ent. 1: 133—154, pls. 5—7. (First part of the series on Dutch Symphyta).

Id. (2). — Tijdschr. Ent. 1: 171—194, pls. 9—12. (Second part on Dutch Symphyta).

Id. (3). — Tijdschr. Ent. 2: 63—78, pls. 3—5. (Third part on Dutch Symphyta).

Phyllodes Eyndhovii, nieuwe vlindersoort uit Java. — Tijdschr. Ent. 2: 86—89, pl. 6. (New

moth from Java).

De inlandsche bladwespen in hare gedaanteverwisselingen en levenswijze beschreven (4). —

Tijdschr. Ent. 2: 134—151, pls. 8—10. (Fourth part on Dutch Symphyta).

Eenige aanteekeningen van gemengden inhoud. — Tijdschr. Ent. 2: 152—158. (Some mis-

cellaneous notes).

Over de verdedigingsmiddelen der bladwespen larven. — Jaarb. Natura Artis Magistra, p.

125—133. (Short paper on the defense-mechanisms of sawflies, mainly against parasites).

Natuurlijke historie van Nederland. De dieren van Nederland. Overzigt der gelede dieren 1:

i-vi+ 1—296, many text-figs., pls. 1—19. — Haarlem. (Popular survey of the Dutch Arthro-

poda).

Naamlijst van Nederlandse regtvleugelige insekten (Orthoptera). — Bouwst. Faun. Ned.

(Herklots) 3: 34—39. (Checklist of the Dutch Orthoptera).

Over eenige nieuwe soorten van dagvlinders uit Oost-Indië. — Tijdschr. Ent. 3: 35—45, pls.

1—4. (New butterflies from the Dutch East Indies).

Description d'une nouvelle espèce de lépidoptère du genre Papilio, suivie d’une énumération

242

1860c.

1860d.

1860e.

18608.

1860h.

18601.

1860j.

1861a.

1861b.

1861c.

1861d.

1861e.

1861f.

1861g.

1862a.

1862b.

1862c.

1862d.

1862e.

1862f.

1862g.

1862h.

18621.

1862j.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

des espèces de ce genre, qui se trouvent au Musée royal des Pays-bas à Leide. — Tijdschr.

Ent. 3: 67—89, pl.6.

De inlandsche bladwespen in hare gedaanteverwisselingen en levenswijze beschreven. (5).

— Tijdschr. Ent. 3: 99—119, 3 pls. (Fifth part on Dutch Symphyta).

Deux Lépidoptères nouveaux. — Tijdschr. Ent. 3: 125—127.

Beschrijving van eenige nieuwe soorten van bladwespen. — Tijdschr. Ent. 3: 128—130.

(Descriptions of some new species of sawflies).

Over de larve van Carabus auratus L. — Tijdschr. Ent. 3: 166—167, pl. 12: A, B. (Note

about larva of C. auratus).

Tweede naamlijst van inlandsche Hemiptera, eerste gedeelte. — Tijdschr. Ent. 3: 168—195.

(With H. W. de Graaf and G. A. Six). (Checklist of the Dutch Hemiptera, cf. 1862b; first

part).

Iets over de Columbatscher-mug (Simulium maculatum Meig.). — Jaarb. Natura Artis Ma-

gistra: 128—135. (Short note about the midge Simulium maculatum).

Etwas über die Columbatscher Mücke (Simulium maculatum Meig.). — Stett. ent. Ztg 21:

306—319. (Introduction and translation of 1860h. by C. A. Dohrn).

Beschrijvingen en afbeeldingen van Nederlandsche vlinders. In: Sepp’s Nederlandsche Insec-

ten, 2nd series, 1860-1900, pt. I-IV. — ’s-Gravenhage. (Descriptions and illustrations of

Dutch Lepidoptera).

Over drie bladwespen en Sesia formicaeformis. — Tijdschr. Ent. 4: 44—46. (Notes on three

sawflies, and Sesia formicaeformis).

Aantekening omtrent het leven en de wetenschappelijke werkzaamheden van Q. M.R. Ver

Huell, in leven Ridder van onderscheidene orden, gepensioneerd Schout-bij-nacht, lid van

verschillende geleerde Genootschappen, enz. — Tijdschr. Ent. 4: 55—63. (Obituary of Q.

M. R. Ver Huell).

De inlandsche bladwespen in hare gedaanteverwisseling en levenswijze beschreven (6). —

Tijdschr. Ent. 4: 65—87, pls. 1—4 (Sixth part on Dutch Symphyta).

Beschrijving van eenige nieuwe soorten van Lucanidae. — Tijdschr. Ent. 4: 101—115, pls.

5—7. (Description of new Lucanidae).

Description de quelques espèces nouvelles de Lepidoptères. — Tijdschr. Ent. 4: 157—163,

pls. 8—10.

Bij de afbeelding der larve en pop van Rhyssa persuasoria. — Tijdschr. Ent. 4: 176—177, pl.

12. (Note on figures of Rhyssa persuasoria).

Natuurlijke historie van Nederland. De dieren van Nederland. Overzigt der gelede dieren 2:

297—558, pls. 20—35. — Haarlem (cf. 1859f).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (7). — Tijd-

schr. Ent. 5: 49—71, 4 pls. (Seventh part on the Dutch Symphyta).

Tweede naamlijst van inlandsche Hemiptera. — Tijdschr. Ent. 5: 72—95. (With H. W. de

Graaf and G. A. Six), (Second part; cf. also 1860g.).

Beschrijving van Dryobius riparius v. Voll. — Tijdsch. Ent. 5: 95—96, pl. 5, figs. 1—3. (Des-

cription of Dryobius riparius).

Iets over het cocon van Sagra Boisduvalii Dej. — Tijdschr. Ent. 5: 97—99, pl. 5 figs. 69.

(Cocoon of Sagra Boisduvalii, Coleoptera).

Beschrijving eener nieuwe soort van Kakkerlak uit Sumatra, Archiblatta Hoevenii. — Tijd-

schr. Ent. 5: 106—110, pl. 6, figs. 1—2. (Description of Archiblatta Hoevenii).

Bijdrage tot de kennis van het vlindergeslacht Adolias. — Tijdschr. Ent. 5: 181—207, pls.

10—12. (Contribution to the knowledge of the genus Adolias, Lepidoptera).

Description d’un Lépidoptère nouveau de la tribu des Saturnides, appartenant à la faune son-

daique. — Rev. Mag. Zool. Pure Appl. (2nd series) 14: 335—339, pl. 14 fig. 1.

Diagnoses specierum novarum e genere Lepidopterum diurnorum Adolias, insulas Indiae

orientalis inhabitantium. — Versl. Meded. K. Akad. Wet. Natuurk., Amst. 13: 270-273. (14

new species of Adolias are described from Indonesia).

Over Stekaas en wat daaruit voortkomt. — Jaarb. Natura Artis Magistra: 143—153. (Popu-

lar paper about caddis-flies).

Levensschets van Dr. J. Wttewaall. — Ned. Spectator, p. 290—292 + 298—299. (Obituary

of Dr. J. Wttewaall).

1863a.

1863b.

1863c.

1863d.

1863e.

1863f.

1863g.

1863h.

18631.

1863j.

1863K.

1864a.

1864b.

1864c.

1864d.

1865a.

1865b.

1865c.

1865d.

1865e.

1865f.

18658.

1865h.

18651.

1865j.

1865k.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 243

Over de inlandsche soorten van het geslacht Delphax Fabr. — Tijdschr. Ent. 6: 10—12. (No-

tes on the Dutch spp. of Delphax, Homoptera).

Mededeeling. — Tijdschr. Ent. 6: 13—16. (Short descriptions of some papers published in

“Schriften der Königlichen physikalisch-ökonomischen Gesellschaft zu Königsberg”).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (8). — Tijd-

schr. Ent. 6: 65—86, pls. 4. (Eighth part on Dutch Symphyta).

Bijvoegselen tot de naamlijst van Nederlandsche schildvleugelige insecten. — Tijdschr. Ent.

6: 90—116. (New records of Dutch Coleoptera).

Beschrijving van drie nieuwe soorten, behoorende tot het geslacht van Hemiptera, Oplomus.

— Tijdschr. Ent. 6: 121—123. (Descriptions of three new species of Oplomus).

Description de quelques nouvelles espèces de lépidoptères des Indes Orientales. — Tijdschr.

Ent. 6: 129— 144, pls. 8—10.

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (9). — Tijd-

schr. Ent. 6: 179—187, 2 pls. (Ninth part on Dutch Symphyta).

Bijdrage tot de kennis van het vlindergeslacht Leptosoma Boisd. — Ned. Tijdschr. Dierk. 1:

35—51. (Contribution to the knowledge of the genus Leptosoma, Lepidoptera).

Over een merkwaardige soort van Tweevleugelig Insekt, Antidoxion fulvicorne. — Ned.

Tijdschr. Dierk. 1: 346—349. Same paper appeared in Versl. Med. K. Akad. Wet. Natuurk.,

Amst. 15: 1—4. (About the dipteron Antidoxion fulvicorne).

Beschrijving van eenige nieuwe soorten van Diptera. — Ned. Tijdschr. Dierk. 1: 349— 355.

Same paper appeared in Versl. Med. K. Akad. Wet. Natuurk., Amst. 15: 8—18. (Descrip-

tions of some new Diptera).

Essai d’une faune entomologique de l’Archipel Indo-Neérlandais. Première Monographie:

famille des Scutellérides. p. 1—64, 4 col. pls. — La Haye.

Verslag van de negentiende algemeene vergadering der Nederlandsche Entomologische Ver-

eeniging te Leiden, den 29sten Augustus 1863. — Tijdschr. Ent. 7: 1—31 (perhaps already

published in 1863) (Report of the 19th general meeting of the Netherlands Entomological So-

ciety, with description of a new genus and species of Scutelleridae.).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze (10). — Tijdschr. Ent.

7: 59—74, pls. 1—3. (Tenth part on the Dutch Symphyta).

Description de quelques espèces nouvelles de Coleoptères. — Tijdschr. Ent. 7: 145—170,

pls. 9—12.

Bijdrage ter aanvulling van de naamlijst der inlandsche Waternimfen. — Bouwst. Faun.

Ned. (Herklots) 3: 188. (Addition to checklist on Odonata).

Verslag van de twintigste algemeene vergadering der Nederlandsche Entomologische Veree-

niging gehouden te Amersfoort, den 30sten Julij 1864. — Tijdschr. Ent. 8: 1—29. (Report of

the 20th general meeting of the Netherlands Entomological Society, with notes on some Or-

thoptera).

Description d’une nouvelle espèce de Calodema (Calodema Johannae) de Waigeou. — Tijd-

schr. Ent. 8: 61—62, pl. 1. (Buprestidae).

Un genre nouveau d’Hémiptères Scutellérides. — Tijdschr. Ent. 8: 63—64, pl. 1 figs. 3—5.

Beschrijving van een nieuwe Tetrix-soort, Tetrix limosina. — Tijdschr. Ent. 8: 65—66, pl. 1

figs. 6—8. (New species of Tetrix).

Over een rups van Clostera curtula. — Tijdschr. Ent. 8: 69—70, pl.2 figs 34. (Description

of the caterpillar of Clostera curtula).

De inlandsche bladwespen in hare gedaantewisselingen en levenswijze beschreven (11). —

Tijdschr. Ent. 8: 73-93, pls. 3—6. (Eleventh part on Dutch Symphyta).

Macrolyristes, een nieuw geslacht van Orthoptera. — Tijdschr. Ent. 8: 106110, pl. 7, pl. 8

figs. A—C. (Description of a new genus).

Opmerkingen omtrent de vangst van kleine insecten. — Tijdschr. Ent. 8: 132—136. (Notes

on the capture of small insects).

Sur quelques Lucanides du Muséum Royal d’Histoire Naturelle à Leide. — Tijdschr. Ent. 8:

137—156, pls. 10-11.

Over eene galwespsoort, die nieuw is voor onze fauna. — Tijdschr. Ent. 8: 160—168. (Des-

cription of an unnamed species of gall-wasp).

Beschrijving van eenige nieuwe soorten van Curculioniden, uit het geslacht Apoderus. —

Ned. Tijdschr. Dierk. 2: 158—167, 3 figs. (Descriptions of new Attelabidae).

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Essai d’une faune entomologique de l’Archipel Indo-Néerlandais. Seconde Monographie:

Famille des Piérides, p. 1—70, pls. 1—7. — La Haye.

Description de deux nouvelles espèces de Piérides et de la femelle de P. Polisma. —Tijdschr.

Ent. 9: 57—60, pls. 1—2.

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (12). —

Tijdschr. Ent. 9: 189—208, pls. 7—9. (Twelfth part of his series on Dutch Symphyta).

Description de deux nouvelles espèces de Lépidoptères. — Tijdschr. Ent. 9: 209—210, pl.

10.

Eenige nieuwe soorten van het geslacht Dalcantha Am. & Serv. — Tijdschr. Ent. 9: 216—

221, pl. 11 figs. 5—9. (Some new species of the genus Dalcantha, Hemiptera).

Beschrijving van eenige nieuwe soorten van Coleoptera uit Oost-Indië. — Tijdschr. Ent. 9:

222—229, pls. 11—12. (Descriptions of new Indonesian Coleoptera).

Naamlijst der soorten van de familie der Pieriden behoorende tot de fauna van Nederlandsch

Oost-Indië. — Ned. Tijdschr. Dierk. 3: 53—61. (Checklist of Indonesian Pieridae).

Mededeelings omtrent de toezendingen in de laatste jaren aan ’s Rijks Museum van Natuur-

lijke Historie gedaan, in betrekking tot de entomologie. — Versl. Meded. K. Akad. Wet.

Natuurk., Amst. 1 (2): 210—223. (Notes on the Entomological collections sent to the Rijks-

museum van Natuurlijke Historie at Leiden).

Jan Swammerdam’s Catalogus. — Ned. Spectator, p. 125—127. (Notes on a catalogue by Jan

Swammerdam on objects owned by his father).

Twee waarnemingen van wijlen Dr. J. Wttewaall. — Tijdschr. Ent. 10: 20—25, pl. 1 figs. 1—

6. (Two observations by the late Dr. J. Wttewaall).

Bijschrift bij het opstel van Dr. Mayr. — Tijdschr. Ent. 10: 118—119. (Note on the paper by

Dr. Mayr).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (13). —

Tijdschr. Ent. 10: 165—182, pls. 6—10. (Thirteenth part on the Dutch Symphyta).

Drie nieuwe soorten van inlandsche Hymenoptera. — Tijdschr. Ent. 10: 222—226, pl. 10.

(According to the title three new species are described from the Netherlands, but actually

two new species are named).

Naamverwisseling. — Tijdschr. Ent. 11: 128. (Note on the synonymy of Euploea superba

Voll. with E. schlegelii Felder).

De inlandsche Hemiptera beschreven en meerendeels ook afgebeeld. Eerste stuk met twee

platen. —Tijdschr. Ent. 11: 129—172 pls. 5—6. (Monograph on the Dutch Hemiptera, first

part; cf. 1869a, 1870c, 1873d, 1876c, 1877e, 1878d.).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (14). —

Tijdschr. Ent. 11: 197—209, pls. 8—10. (Fourteenth part on the Dutch Symphyta).

Diagnosen van eenige nieuwe soorten van Hemiptera Heteroptera. — Versl. Meded. K.

Akad. Wet. Natuurk., Amst. 2 (2): 172—188. (Diagnoses of some new species of Heteropte-

ra).

Schetsen ten gebruike bij de studie der Hymenoptera. Ist part, 3 pls. — ’s-Gravenhage.

(Sketches for the study of Hymenoptera. First part contains line-drawings of the Ichneumoni-

dae, for 2nd part (Braconidae) cf. 1869d, for 3rd part (Pteromalidae) cf. 1871e, and for 4th

part (Proctotrupoidea) cf. 18731).

Essai d'une faune entomologique de l’Archipel Indo-Néerlandais. Troisième Monographie:

famille des Pentatomides, Ire partie. p. 1—54, 4 col. pls. — La Haye. (Monograph on the

Indonesian Pentatomidae).

Natuurlijke Historie van Nederland. De gelede dieren, 2 pts. — Amsterdam. (Reprint of the

popular treatise on the Dutch Arthropoda, cf. 1859f).

Iets over galnoten. — Jaarb. Natura Artis Magistra: 137—146. (Short popular paper about

galls and gallwasps).

De inlandsche Hemipteren beschreven en meerendeels ook afgebeeld. Tweede stuk met

twee platen. — Tijdschr. Ent. 12: 49—74, pl. 1—2. (Second part of the paper on Dutch He-

miptera).

Nieuwe naamlijst van Nederlandsche Vliesvleugelige insecten (Hymenoptera). — Tijdschr.

Ent. 12: 89—127, pl. 3. (Checklist of the Dutch Hymenoptera, followed by descriptions of

several new species of Symphyta and Cynipoidea).

Description de six espèces nouvelles de Hémiptères Hétéroptères. — Tijdschr. Ent. 12:

1869d.

1869e.

1870a.

1870b.

1870c.

1870d.

1870e.

1870f.

1870g.

1871a.

1871b.

1871c.

1871d.

1871e.

1871f.

1872a.

1872b.

1872c.

1872d.

1873a.

1873b.

1873c.

1873d.

1873e.

1873f.

1873g.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 245

255— 260, pl. 11, figs. a—e. (Description of six new species of Heteroptera).

Schetsen ten gebruike bij de studie der Hymenoptera. 2nd part. 3 pls. (Braconidae; cf.

1868e).

Recherches sur la Faune de Madagascar et de ses dépendances, d’après des découvertes de

Frangois P. L. Pollen et D. C. van Dam, Insectes, 5(1): 1—14, 2 pls. — Leiden. (with E. de

Sélys Longchamps; list and descriptions of some new species collected in Madagascar and

surrounding islands).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (15). —

Tijdschr. Ent. 13: 55—74, pls. 1—4. (Fifteenth part on Dutch Symphyta).

Bijschrift bij Plaat 6. — Tijdschr. Ent. 13: 158, pl. 6. (Text of plate 6).

De inlandsche Hemipteren beschreven en meerendeels ook afgebeeld. Derde stuk met drie

platen. —Tijdschr. Ent. 13: 263—302, pls. 10—12. (Third part of paper on Dutch Hemipte-

ra).

Gedaantewisseling en levenswijze der Insecten. p. 1—457, figs. 1—251, 4 pls. — Haarlem.

(The insects, their biology; a general survey; reprinted in 1876 (Amsterdam)).

Laatste lijst van Nederlandsche schildvleugelige insecten (Insecta Coleoptera), p. i—

iv+(1)+1—146. (Checklist of the Dutch Coleoptera).

Jean Théodore Lacordaire — Ned. Spectator, p. 321—322. (Obituary of J. T. Lacordaire).

Verslag van de vier-en-twintigste algemene vergadering der Nederlandsche Entomologische

Vereeniging, gehouden te Zwolle, den 31sten Julij 1869. — Tijdschr. Ent. 13: 17—20. (Re-

cord of Polysphincta carbonator Grav. as ectoparasite of Miranda cucurbitina and notes on

other parasites of spiders).

(Comment on Darwin’s theory of evolution). — Tijdschr. Ent. 14: 28—29.

Quelques espèces nouvelles de Curculionites et de Longicornes. — Tijdschr. Ent. 14: 101—

112, pls. 4—5.

Les Batocérides du Musée de Leide. — Tijdschr. Ent. 14: 211—220, pl. 9.

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (16). —

Tijdschr. Ent. 14: 237—254, pls. 10—12. (Sixteenth part on the Dutch Symphyta).

Schetsen ten gebruike bij de studie der Hymenoptera. 3rd part, 4 pls. Pteromalidae. (cf.

1868e).

[Description of Eupoecila balteata, in Mohnike, O., Uebersicht der Cetoniden der Sunda-In-

seln und Molukken]. — Arch. Naturg. 37: 277, pl. 6 fig. 3. (Letter of Vollenhoven quoted

verbatim, his figure published).

Bladvulling. — Tijdschr. Ent. 15: 111—112. (Notes on some Tenthredinidae in the Ritsema

Collection).

Opmerkingen omtrent een miskenden Sumatraanschen kever. — Tijdschr. Ent. 15: 125—

128. (Notes on Rhomborrhina, Cetoniidae).

Beschreibung einer neuen Paussus-Art aus Ost-Indien, Paussus Ludekingii. — Stett. ent.

Ztg. 33: 82.

Over de ziekte van de wijnstok in Frankrijk en de Phylloxera vastatrix. — Alb. Nat. Haar-

lem, p. 33—46. (Paper on the pest by Phylloxera vastatrix in France).

Aderen in de ondervleugels der Hemiptera en andere mededelingen. In: Verslag van de zes-

de wintervergadering der Nederlandsche Entomologische Vereeniging, gehouden te Leiden,

den 28 December 1872. — Tijdschr. Ent. 16: LIV—LVIII. (On the veins of the hind wings of

Hemiptera).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (17). —

Tijdschr. Ent. 16: 1—15, pls. 1—3. (Seventeenth part on the Dutch Symphyta).

Trois espèces nouvelles du genre Rhyssa. — Tijdschr. Ent. 16: 67—70, pl. 4.

De inlandsche Hemipteren beschreven en meerendeels ook afgebeeld. Vierde stuk met drie

platen. —Tijdschr. Ent. 16: 75—122, pls. 5—7. (Fourth part of the paper on Dutch Hemipte-

ra).

Nieuwe naamlijst van Nederlandsche vliesvleugelige insecten (Hymenoptera), tweede stuk.

Beschrijving der nieuwe soorten. Bijlage tot de lijst van inlandsche Hymenoptera. — Tijd-

schr. Ent. 16: 209—220, pl. 9. (Descriptions of 13 new species of Ichneumonoidea).

Description d’un Bombus nouveau de l’île de Sumatra. — Tijdschr. Ent. 16: 229—230, pl. 10.

246

1873h.

18731.

1873j.

1873k.

18731.

1874a.

1874b.

1874c.

1875a.

1875b.

1875c.

1875d.

1875e.

1875f.

1875g.

1875h.

1876a.

1876b.

1876c.

1876d.

1876e.

1876f.

1876g.

1876h.

1877a.

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Description de trois espèces nouvelles de Lépidoptères des Indes Orientales. — Tijdschr.

Ent. 16: 244—247, pls. 11—12.

Schetsen ten gebruike bij de studie der Hymenoptera. 4th part, 4 pls. — ’s-Gravenhage.

(Proctotrupoidea).

Vijf entomologische wandelingen, p. 1—162, ill. — Haarlem. (Five “entomological walks”

are described).

Een snuitkever en een sluipwespje. — Jaarb. Natura Artis Magistra: 155—168. (Popular pa-

per about some observations by Stollwerk concerning the hymenopterous parasite of Attela-

bus curculionoides).

Een indringster en hare gevechten. — Jaarb. Natura Artis Magistra: 173—180. (Popular pa-

per on the biology of the ant genus Solenopsis).

Verslag van de acht-en-twintigste zomervergadering der Nederlandsche Entomologische

Vereeniging, gehouden te Breda den 7 Junij 1873. — Tijdschr. Ent. 17: xii—xv. (Notes on

Dryinidae and Tenthredo agilis Klug).

Over de groep der Dryiniden in de familie der Proctotrupiden, met beschrijving eener nieu-

we soort — Versl. Meded. K. Akad. Wet., Amst. (2) 8: 150—162, figs. 1—6. (Notes on the

Dryinidae, with description of Dryinus spectrum Snellen van Vollenhoven).

Verslag van de zevende wintervergadering der Nederlandsche Entomologische Vereeniging,

gehouden te Leiden den 6 December 1873. — Tijdschr. Ent. 17: LXV—LXVI. (List of Hy-

menoptera new to the Dutch fauna, i.a. Dryinus spectrum n.sp.).

Verslag van de negen-en-twintigste zomervergadering der Nederlandsche Entomologische

Vereeniging, gehouden te Arnhem den 29 Augustus 1874. — Tijdschr. Ent. 18: xx—xxii+xx-

xi—xxxil. (Notes on Dutch Lepidoptera, Hemiptera and Hymenoptera).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (18). —

Tijdschr. Ent. 18: 33—49, pl. 3—5. (Eighteenth part on the Dutch Symphyta).

Systematische lijst der in dit Tijdschrift beschreven gedaantewisselingen van bladwespen. —

Tijdschr. Ent. 18: 50—52. (List of Dutch Symphyta of which metamorphosis was described in

this journal).

De inlandsche Hemipteren, beschreven en meerendeels ook afgebeeld. Vijfde stuk met drie

platen. —Tijdschr. Ent. 18: 150—185, pls. 8—10. (Fifth part on Dutch Hemiptera).

Pinacographia. — Illustrations of more than 1000 species of North-West-European Ichneu-

monidae sensu Linnaeano, p. 1—8, pls. 1—5. (Series of papers on about 400 species of Ich-

neumonidae, Braconidae, Chalcidoidea, and Proctotrupoidea, all illustrated and shortly des-

cribed. Incomplete when Vollenhoven died. N.B. Preface (p. I—IX), index and p. 65—68

are by G. A. Six (1880)).

Id. — p. 9—16, pls. 6—10.

De Colorado-kever. — Alb. Nat., Haarlem: 97—104. (Popular paper on Leptinotarsa decem-

lineata Say).

De wurm in ’t hout (Anobium pertinax). — Alb. Nat., Haarlem: 187—189. (Popular paper

on some Anobium spp. and their parasite).

Verslag van de dertigste zomervergadering der Nederlandsche Entomologische Vereeniging,

gehouden te Amsterdam op Zaturdag 24 Julij 1875. — Tijdschr. Ent. 19: XL—XLII. (Notes

on some Lepidoptera and Hymenoptera).

Verslag van de negende wintervergadering der Nederlandsche Entomologische Vereeniging,

gehouden te Leiden op Zaturdag 18 december 1875 des avonds ten 7 ure. — Tijdschr. Ent.

19: CIX—CXIII. (Comparison of the fauna of Heteroptera of the Netherlands and Great

Britain).

De inlandsche Hemipteren, beschreven en meerendeels ook afgebeeld. Zesde stuk met 3 pla-

ten. —Tijdschr. Ent. 19: 65—132, pls. 3—S. (Sixth part on Dutch Hemiptera).

Iets over Otiorhynchus sulcatus L. — Tijdschr. Ent. 19: 210. (Record from the Netherlands).

Bijvoegsel tot de nieuwe naamlijst van Nederlandsche vliesvleugelige insecten (Hymenopte-

ra). —Tijdschr. Ent. 19: 211—257. (Additions to the checklist of Dutch Hymenoptera).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (19). —

Tijdschr. Ent. 19: 258—277, pls. 10—12. (Nineteenth part of series on Dutch Symphyta).

Pinacographia, p. 17—24, pls. 11—15. (Third part of this series on Hymenoptera).

Id., p. 25—32, pls. 16—20 (Fourth part of this series).

Verslag van de een-en-dertigste zomervergadering der Nederlandsche Entomologische Ver-

1877b.

1877c.

1877d.

1877e.

1877f.

18778.

1878a.

1878b.

1878c.

1878d.

1878e.

1878.

18789.

1878h.

18781.

1879a.

1879b.

1879e.

1879d.

1879e.

1879f.

1880a.

1880b.

1880c.

1880d.

1880e.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 247

eeniging, gehouden te Middelburg. — Tijdschr. Ent. 20: xiii—xv. (Some notes on galls of

Andricus ramuli L. and on the caterpillar of Meterocampa margaritaria L.).

Verslag van de tiende wintervergadering der Nederlandsche Entomologische Vereeniging,

gehouden te Leiden op Zaturdag 23 December 1876. — Tijdschr. Ent. 20: Ixxiv—Ixxvii.

(Lists of galls, and of scarce Dutch Heteroptera are given; Coleophora onosmella Brahns and

a Coccus species).

Bijdrage tot de kennis der gedaantewisseling van Diptera. — Tijdschr. Ent. 20: 57—63, pl. 4

figs. 1—5. (Notes on the biology of Machaerium maritimum Hal.).

Tryphon praerogator Grav. — Tijdschr. Ent. 20: 64. (Short note on its identity).

De inlandsche Hemipteren, beschreven en meerendeels ook afgebeeld. Zevende stuk met

vier platen. — Tijdschr. Ent. 20: 90—167, pls. 7—10. (Seventh part of the series on Dutch

Hemiptera).

Pinacographia, p. 33—39, pls. 21—25. (Fifth part of the series).

Determinatie der platen in het werk van Joannes Goedaert. — Alb. Nat., Haarlem: 307—

318. (On the identification of insects figured by Joannes Goedaert).

Verslag van de twee-en-dertigste zomervergadering der Nederlandsche Entomologische Ver-

eeniging, gehouden te Utrecht op Zaturdag 30 Junij 1877. — Tijdschr. Ent. 21: xvii—xix.

(On an Ephemera species, Cheimatobia and some spp. from the Holmgren Collection).

Verslag van de elfde wintervergadering der Nederlandsche Entomologische Vereeniging, ge-

houden te Leiden op Zaturdag 22 December 1877. — Tijdschr. Ent. 21: LXXIV—LXXVII.

(On Indonesian Ichneumonidae and the checklist of Dutch Hymenoptera).

Un Ephialtes de Java, Ephialtes Melanomerus de Haan. — Tijdschr. Ent. 21: 48.

De inlandsche Hemipteren, beschreven en meerendeels ook afgebeeld. Achtste stuk met

twee platen. — Tijdschr. Ent. 21: 49—80, pls. 3—4. (Eighth part on the Dutch Hemiptera).

Oproeping. — Tijdschr. Ent. 21: 126—128. (Appeal concerning Orthoptera).

Especes nouvelles ou peu connues d’Hymenopteres térébrants. — Tijdschr. Ent. 21: 153—

177, pls. 9—11.

Pinacographia, p. 41—48, pls. 26—30. (Sixth part of the series).

Id. p. 49—56, pls. 31—35. (Seventh part of the series).

Hemiptera Heteroptera Neerlandica. De inlandsche ware Hemipteren. (Land- en water-

wantsen) beschreven en meerendeels ook afgebeeld, p. 1—368, pls. 1—22. ’s-Gravenhage.

(Treatise on the Dutch Heteroptera).

Verslag van de drie-en-dertigste zomervergadering der Nederlandsche Entomologische Ver-

eeniging, gehouden te Nijmegen op Zaturdag 29 Junij 1878. — Tijdschr. Ent. 22: ix—

xilit+xxvi. (Notes on Typhlocyba and other faunistic data; description of Proctotrupes sixia-

nus).

Verslag van de twaalfde wintervergadering der Nederlandsche Entomologische Vereeniging,

gehouden te Leiden op Zondag 22 December 1878. — Tijdschr. Ent. 22: xc. (On Ephippige-

ra vitium Serv. and Cratomus megacephalus F.).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (20). —

Tijdschr. Ent. 22: 1—20, pls. 1—4. (Twentieth part of the series on Dutch Symphyta).

Bijvoegsel tot de lijst der inlandsche Hemiptera Heteroptera. — Tijdschr. Ent. 22: 227—231,

pl. 12, figs. C—F. (Additions to the checklist of Dutch Heteroptera).

Einige neue Arten von Pimplarien aus Ost-Indien. — Stett. ent. Ztg. 40: 133—150.

Pinacographia, p. 57—64, pls. 36—40. (Eighth part of the series).

Verslag van de vier-en-dertigste zomervergadering der Nederlandsche Entomologische Ver-

eeniging. — Tijdschr. Ent. 23: XIV—XVI. (Short enumeration of some interesting species of

Hymenoptera new to the Dutch fauna and a note on Typhlocyba).

De inlandsche bladwespen in hare gedaantewisseling en levenswijze beschreven (21). —

Tijdschr. Ent. 23: 4—16, pls. 1—3. (21st part on the Dutch Symphyta).

Pinacographia, p. 65—68, pls. 41—45. (Ninth part of the series, text by G. A. Six).

Levensschets van Mr. Pierre Lyonet. — Alb. Nat., Haarlem: 1—14, 1 portrait. (Sketch of

the life of P. Lyonet).

Lyonet’s Microskoop. — Alb. Nat., Haarlem: 62. (Short note about the fate of Lyonet’s mi-

croscope).

248 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

GENUS-GROUP AND SPECIES-GROUP NAMES FIRST PUBLISHED BY SNELLEN VAN

VOLLENHOVEN

To our knowledge, Vollenhoven published nine new genus-group names and

471 new species-group names. The following list gives all the names we have

found, in their original combination, alphabetically arranged within the families in

which they are currently placed. Each species-group name is followed by a refe-

rence to the bibliography and many are followed by the name of the genus in

which the species is currently placed. Furthermore type-localities are cited, usual-

ly in the original spelling. Frequently a country name is added between brackets.

The status of type-material is given as follows. Holotype: Vollenhoven undoub-

tedly had only one specimen at hand; syntypes: Vollenhoven undoubtedly had

more specimens at hand or did not specify the number. If not indicated otherwise,

the specimens are present in the Leiden museum. The number of syntypes is the

number of specimens found in the collections, and does not indicate the number

of types originally at hand. So “1 syntype” means that one specimen has been

found; this might in fact be the only specimen Vollenhoven had.

In the Hymenopterous family Ichneumonidae most of the types of Dutch spe-

cies were found in the collection of the Dept. of Entomology in Wageningen. For

this family the presence of type-material in Leiden en Wageningen is mentioned

separately. |

Vollenhoven did not label his types as such. In many cases he even did not add

a name to the pin. This will certainly have caused the loss of various types and

makes it difficult to recognise them.

A few specimens bear lectotype labels, but we are not always sure whether the

designations were actually published. The reverse may also be true. Where the

designation of a lectotype is known to have been published, this is indicated. No-

ne of the statements in the present list should be interpreted as a nomenclatorial

action taken by the compilers, i.e. neither the statements concerning lectotypes,

nor those concerning recombinations, synonymies, etc. We have not included in-

formation on current species-level synonymies except those proposed by Vollen-

hoven himself. The reason is that these synonymies are only rarely based on a stu-

dy of the type-specimens.

The lists for the various orders have been compiled by the authors, viz., J. Krik-

ken (Coleoptera), P. H. van Doesburg (Dictyoptera, Orthoptera, Hemiptera), C.

van Achterberg (Hymenoptera), K. W. R. Zwart (Hymenoptera: Ichneumonidae

in the Wageningen collection) and R. de Jong (Lepidoptera). P. J. van Helsdin-

gen supplied information concerning Diptera. All are curating the collections of

these groups in the Leiden museum or the collection of the Dept. of Entomology

in Wageningen. Any enquiries should be addressed to the curator of the group

concerned. |

ORDER DICTYOPTERA

Family Archiblattidae

Archiblatta, 1862e: 106, type-sp. A. hoevenii Vollenhoven, 1862 (monotypy).

Archiblatta hoevenii, 1862e: 106, pl. 6 figs. 1—2, Padangse Bovenlanden, or Agam (Sumatra), holoty-

pe, 9.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 249

ORDER ORTHOPTERA

Family Tettigoniidae

Macrolyristes, 1865g: 106, type-sp. M. imperator Vollenhoven, 1865 (monotypy).

Macrolyristes imperator, 1865g: 108, pl. 7 figs. 1—2, pl. 8, figs. A, B, C, Java, Borneo, 1 d syntype,

Pengorang (Borneo).

Family Acrididae

Oedipoda virgula, 1869e: 3, 11, pl. 2 fig. 2, Madagascar, 2 d syntypes.

Family Tetrigidae

Tetrix limosina, 1865d: 65, pl. 1 figs. 6—8 (to Ophiotettix), Isl. Gebeh, 2 9 syntypes.

ORDER HEMIPTERA

Suborder HETEROPTERA

Family Berytidae

Berytus driebergensis, 1869a: 64, pl. 1 fig. 9 (to Berytinus), De Bildt, Sterkenburg nr. Driebergen (Ne-

therlands), 2 d syntypes.

Family Coreidae

Choerommatus niger, 1869e: 5, 14, pl. 1 fig. 8, Mayotte (Comores), holotype, 2.

Coreus difficilis, 1869a: 56, pl. 1 fig. 5 (to Ceraleptus), Den Haag, (Netherlands), holotype, à.

Family Cydnidae

Acatalectus flavomarginatus , 1868d: 177 (to Adrisa), Nova Caledonia, 4 syntypes.

Acatalectus luteomarginatus , 1868d: 177 (to Adrisa), Timor, Flores, 3 syntypes; 1868f: 19, pl. 2 fig. 9.

Aethus pallidicornis, 1868d: 177 (to Geotomus), Bezoeki (Java), Ceram, 8 syntypes; 1868f: 17, pl. 2

fig. 8.

Cyrtomenus insignis, 1868d: 177 (to Scoparipes), Java, Sumatra, Borneo, 6 syntypes; 1868f: 16, pl. 2

fig. 7.

Family Microphysidae

Microphysa sanguinea, De Graaf & Snellen van Vollenhoven, 1852d: 175 (to Loricula), Leiden, (Ne-

therlands), 1 9 syntype.

Family Miridae

Lopus subpatellatus, 1875d: 172, pl. 9, figs. 2, 2a (to Conostethus), Netherlands, Texel, Velp, (Nether-

lands), 2 syntypes.

Lygus aurantiacus, 1876c: 103, (to Phylus), Holland, holotype, d.

Family Pentatomidae

Alcimus collaris, 1868d: 182 (to Alcimocoris), Timor, 1 d syntype.

Alcimus venustus, 1868d: 182 (to Alcimocoris), Gorontalo, (Sulawesi), 1 2 syntype.

Asopus bernsteinii, 1868f: 14, pl. 2 fig. 6 (to Amyotea), Nova Guinea, 1 2 syntype.

Asopus distigma, 1868f: 13, pl. 2 fig. 4 (to Amyotea), Amboina, Ternate, 3 2 syntypes.

250 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Aspidestrophus lineola, 1863k: 44, pl. 3 figs. 10, 10a, Borneo, 2 d, 1 9 syntypes.

Canthecona acuta, 1868f: 9, pl. 2 fig. 1 (to Cantheconidea), Timor, Halmaheira, 1 d,3 9 syntypes.

Canthecona apicalis, 1868f: 6, pl. 1 figs. 3, 3a (to Platynopus), Ternate, Halmaheira, Batjan, Morotai,

Gebeh (Moluccas), 1 4,4 9, 1 ?, syntypes.

Canthecona biguttata, 1868f: 8, pl. 1 fig. 6 (to Platynopus), Aru, holotype, 2.

Canthecona mitis, 1868f: 7, pl. 1 fig. 5 (to Cantheconidea), Amboina, Timor, 3 2 syntypes.

Canthecona plebeja, 1868f: 7, pl. 1 fig. 4 (to Cantheconidea), Ternate, holotype, d.

Cazira coccinelloides, 1868f: 4 (to Blachia?), Hindostan, 1 d syntype.

Chlaenocoris pusillus, 1863k: 45, pl. 3 figs. 11, 11a (to Sepontia), Java, holotype, 2.

Chlorocoris roseus, 1868d: 178, patria ignota, 1 9 syntype.

Coenus punctatissimus, 1868d: 183, Wisconsin (U.S.A.), 1 d syntype.

Cuspicona basimaculata, 1868d: 188 (to Vitellus), Waigeou, 2 6, 2 2 syntypes.

Dalcantha sanctifargavii, 1866d: 218, pl. 11 fig. 6 (to Pycanum), Sylhet (India), 1 d syntype.

Dalcantha servillei, 1866d: 220, pl. 11 fig. 9 (to Pseudopycanum), Malacca, 1 &, 1 nymph, syntypes.

Dalcantha stalii, 1866d: 220, pl. 11 fig. 8, Sylhet (India), 4 d syntypes.

Dalpada aenea, 1868d: 179, Timor, 1 d syntype.

Dalpada triguttata, 1868d: 179, Java, Borneo, 2 &,2 9, syntypes.

Diceraeus euschistoides, 1868d: 180, Wisconsin (U.S.A.), 1 6 syntype.

Diceraeus sellula, 1868d: 179, Java, 19 syntype.

Dryptocephala divergens, 1868d: 178, Rio de Janeiro (Brasil), 1 9 syntype.

Eysarcoris geminatus, 1868d: 182, Java, 1 6 syntype.

Eysarcoris lineola, 1868d: 183, Timor, Borneo, syntypes. Not seen.

Eysarcoris obscurus, 1868d: 182, Java, Sumatra, 2 d syntypes.

Eysarcoris rufoscutellatus, 1868d: 183 (to Menida), Timor, 1 9, syntype.

Hoplistodera decora, 1868d: 181, Manilla, 3 9 syntypes.

Hoplistodera gibba, 1868d: 181 (to Paracritheus), Gorontalo, (Sulawesi), 1 6 syntype.

Hoplistodera schwaneri, 1868d: 182, Borneo, 1 d syntype.

Loxa minor, 1868d: 179 (to Fecelia), Portorico, 1 & syntype.

Lyramorpha perelegans, 1868f: 35, Nouvelle Hollande (Australia), 1 &, type ?, = L. rosea Hope, co-

type.

Megarhynchus fuscus, 1868f: 43, 44 (to Megarrhamphus), Ningpo (Japan), 2 4 syntypes.

Megymenum parallelum, 1868f: 48, pl. 4 figs. 10, 10a (to Pseudaradus), Java, Sumatra, 1 2, syntype

(Java), 2 syntypes lost.

Megymenum quadratum, 1868f: 46, pl. 4 fig. 7 (to Anoplocephala), Morotai, 2 syntypes lost.

types, 2 syntypes lost.

Mormidea hoplites, 1868d: 181 (to Cuspicona), Timor, 1 d syntype.

Mormidea vidua, 1868d: 180 (to Padaeus), Guatemala, 1 à , type.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 251

Mucanum patibulum, 1868f: 22, pl. 3 figs. 1, la, Sumatra, 2 d, 1 2, syntypes.

Oncomerus bernsteinii, 1868f: 30, pl. 4 fig. 1, Halmaheira, Morotai, 2 d,1 2, syntypes.

Oncomerus flavicornis Guér. var. chrysoptera, 1868f: 30, pl. 4, fig. 2, Waigeou, 2 syntypes.

Oplomus flavoguttatus, 1863e: 123, Suriname, 2 & syntypes.

Oplomus haematicus, 1863e: 122, Suriname, 3 9 syntypes.

Pentatoma arlechino, 1868d: 184, Sumatra, 1 d,2 2 syntypes.

Pentatoma hilaris, 1868d: 184 (to Antestia), Java, 1 9 syntype.

Pentatoma ignobilis, 1868d: 184 (to Antestia), Tondano (Sulawesi), 1 &, 1 9 syntypes.

Pentatoma nurus, 1868d: 185 (to Antestia), Java, 1 9 syntype.

pes.

Pentatoma plebeja, 1868d: 184 (to Antestia), Java ?, 1 & syntype.

Podops breviscutum, 1869e: 5, 13 (to Thoria), Nossi-Bé (Malagasy), holotype, à.

Podops serratus, 1863k: 42, pl. 3 figs. 9, 9a (to Scotinophara), Borneo, Celebes, 5 5, 4 9 syntypes.

pes.

Proxys rhododactylus, 1868d: 180, Caracas. Not seen.

Pygoplatys minax, 1868f: 23, pl. 3 fig. 3, Borneo, holotype, 2.

Pygoplatys roseus, 1868f: 24, Malacca, holotype, 9.

Pygoplatys subrugosus, 1868f: 23, pl. 3 fig. 2, Ambon, Buru, 2 d,4 ® syntypes.

Rhaphigaster ludekingii, 1868d: 187 (to Glaucias), Java, Sumatra, Borneo, 3 4,2 9 Syn Den

Rhaphigaster melanosticticus, 1868d: 187 (to Glaucias), Java, 1 9 syntype.

Rhaphigaster nitens, 1868d: 187 (to Menida), Sumatra, 1 9 syntype.

Spudaeus modestus, 1868d: 178 (to Ectenus), Waigeou, 2 9 , 1 ?, syntypes.

Strachia coelestis, 1868d: 186 (to Stenozygum), Nova Hollandia (Australia), 1 9 syntype.

Strachia instabilis, 1868d: 186 (to Antestia), Timor, Isl. Wetar, 3 4,5 9 , syntypes.

Strachia quincunx, 1868d: 186 (to Antestia), Waigeoe, 2 9 syntypes.

Strachia rubescens, 1868d: 186 (to Stenozygum), Celebes, 2 9 syntypes.

syntypes.

Tesseratoma javanica Thunberg var. stictica, 1868f: 26 (var. nov.), Java, holotype, 2.

Tesseratoma javanica Thunberg var. timorensis, 1868f: 26, pl. 3 fig. 5 (not 4!) (var. nov.), Timor, Ado-

nara, 2 8,3 2 syntypes.

Vulsirea tau, 1868d: 186, Rio de Janeiro, 1 d, type.

Family Plataspididae

Brachyplatys crux, 1863k: 54, pl. 4 fig. 9, Sumatra, holotype, 9.

Brachyplatys pallifrons, 1863k: 55, pl. 4 figs. 10, 10a, Timor, 2 4, 10 9 syntypes.

Brachyplatys pauper, 1863k: 54, Java, Ternate, Celebes, 3 4,3 9, 1? syntypes.

Brachyplatys radians, 1863k: 53, pl. 4 fig. 7, Celebes, 3 9 syntypes.

Coptosoma marmoratum, 1868k: 48, pl. 4 fig. 3, Celebes, 1 d, 1 2 syntypes.

Captosoma (sic!) modestum, 1863k: 47, pl. 4 fig. 1, Timor, holotype, d.

Coptosoma muelleri, 1863k: 49, pl. 4 fig. 5, Borneo, holotype 9.

Coptosoma tondanense, 1863k: 48, pl. 4 fig. 2, Celebes, 2 9 syntypes.

252) TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Poseidon, 1864a: 30 (to Cronion Bergroth, 1891, new name), type-sp. P. malayanus Vollenhoven,

1864.

Family Scutelleridae

Callidea bilunulata, 1863k: 33, pl. 3 fig. 1 (to Chrysocoris), Sumatra, holotype, d.

Callidea bosschei, 1863k: 34, pl. 3 fig. 4, (to Chrysocoris), Banca, holotype, 3.

Callidea croesus, 1868d: 176 (to Lampromicra), Aru, Salawatti, Gebeh, 7 4, 7 2, syntypes; 1869c:

2S9Rpla lino re:

Callidea daedalica, 1868d: 176 (to Chrysocoris), patria ignota, 1 9, syntype.

Callidea elongata, 1868d: 175 (to Calliphara), Salawatti, 1 6, 1 2 syntypes; 1869c: 257, pl. 11. fig. b.

Callidea eximia, 1863k: 20, pl. 1, figs. 8, 8a (to Calliphara), Sumatra, Ambon, Morotai, Ternate, 3 g,

6 © syntypes.

Callidea hypherythra, 1863k: 34, pl. 3 fig. 3 (to Chrysocoris), Borneo, holotype, d.

Callidea hypomelaena, 1863k: 33, pl. 3 fig. 2 (to Chrysocoris), Borneo, holotype, 2.

Callidea modesta, 1863k: 29 (to Chrysocoris), Sumatra, holotype, à,

Callidea puella, 1868d: 176 (to Chrysocoris), Cochinchina, 1 6 syntype.

Callidea schlegelii, 1863k: 24, pl. 2 figs. 3, 3a, 3b (to Chrysocoris), Halmaheira, Bajoa, Ternate, 3 9

syntypes.

Cantao rudis, 1863k: 60, Isls. Kajoa & Morotai, 2 4,6 2 syntypes.

Coleotichus pallidus, 1863k: 4, Adelaide (Australia), 1 2 syntype.

Eurygaster ligneus, 1863k: 39, Java, Celebes, 2 ? syntypes.

Hotea fusca, 1863k: 38 (to Tylonca), Java, holotype, sex ?

Libyssa westwoodii, 1868d: 175 (to Calidea), Zambesi, 1 &, 1 ?, syntypes; 1869c: 256.

Pachycoris tigrinus, 1868d: 176 (to Polytes), Colombia, 1 d syntype.

Poecilocoris aeneiventris, 1863k: 7, pl. 1 fig. 3 (to Pachycoris), “India Orientali” (New World spe-

cies!). Not seen.

Scutellera pilosa, 1863k: 13 (as junior synonym of S. amethystina Germar).

Tetrarthria marginepunctata, 1863k: 13, pl. 1, fig. 6, a, b, c, d, Celebes, Sylhet (Bangla Desh), Java ?,

2 d syntypes.

Suborder HOMOPTERA

Aphidoidea

Family Lachnidae

Dryobius riparius, 1862c: 91, 95, pl. 5 figs. 1—3 (to Tuberolachnus). Culemborg (Netherlands), 5

alate syntypes.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 253

Auchenorrhyncha

Family Cicadellidae

Jassus apicalis, 1862b: 83, Doorn (Netherlands), not seen.

Jassus longeciliatus, 1862b: 84, Noordwijk, Voorhout (Netherlands), 1 syntype (coll. Fokker).

Paropia vilis, 1862b: 81 (to Megophthalmus), Gliphoeve, Zutphen, (Netherlands), 2 syntypes.

Typhlocyba agathina, 1862b: 87 (to Linnavuoriana), Den Haag (Netherlands), holotype, à.

Typhlocyba aurantiaca, 1862b: 85 (to Eurhadina) Wassenaar (Netherlands), holotype, 2.

Typhlocyba fastuosa, 1862b: 86 (to Alebra), Driebergen (Netherlands), not seen.

Typhlocyba medenbachii, 1879a: XII (nom. nud. to Eurhadina), Arnhem (Netherlands).

Sternorrhyncha

Family Lecaniidae

Lecanium rosarum, 1862b: 94, Deventer (Netherlands), not seen.

ORDER COLEOPTERA

Vollenhoven’s names were all traced in the Coleopterorum Catalogus of Junk

& Schenkling for ascertaining the current combination of the species-group name.

Meanwhile (i.e. since the publication of the respective parts of this catalogue)

combinations may have been revised, but it is simply impossible to check this wi-

thin a short period of time. Equally, lectotype designations may have been publis-

hed of which we are unaware.

Family Apionidae

Chalcocybebus, 1866e: 225, incl. 2 species.

Chalcocybebus alboguttatus, 1866e: 226, Salawatti, 2 syntypes.

Chalcocybebus nitens, 1866e: 225, Waigeou, 1 syntype.

Family Attelabidae

Apoderus anser, 1865k: 162 (to Cycnotrachelus), Timor, 5 syntypes.

Apoderus erythrogaster, 1865k: 165 (to Compsapoderus), Japan, holotype.

Apoderus niger, 1865k: 166, Gorontalo (Sulawesi), 2 syntypes.

Apoderus olor, 1865k: 161, Java, 2 syntypes.

Apoderus palliatus, 1865k: 162 (to Paracycnotrachelus), Sumatra, Java, 5 syntypes.

Apoderus pardalis, 1865k: 164 (to Paroplapoderus), Japan, 2 syntypes.

Apoderus quadrillum, 1865k: 163 (to Leptapoderus), Java, 2 syntypes.

Family Bostrichidae

Bostrichus iracundus, 1869e: 10, pl. 1 fig. 7, 7a-b (to Xylothrips), Nossi-Bé (Malagasy), 2 syntypes.

Family Buprestidae

Calodema johannae, 1865b: 61, pl. 1 figs. 1, 2, Waigeou, 1 syntype.

Catocantha hemixantha, 1864c: 160, pl. 11 figs. 1, 2 (to Megaloxantha), Banca, holotype.

Chalcophora amabilis, 1864c: 163, pl. 11 fig. 5 (to Chalcophorella), Japan, syntypes. Not seen.

Chalcophora pyrostictica, 1864c: 162, pl. 11 fig. 4 (to Chrysodema), Sumatra, 2 syntypes.

Chalcophora pyrothorax, 1864c: 162 (to Chrysodema), Borneo, 3 syntypes.

Chrysochroa ludekingii, 1864c: 161, pl. 11 fig. 3, Sumatra, 1 syntype.

254 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Chrysobothris pulcherrima, 1864c: 164, Sumatra, Banca, 2 syntypes.

Psiloptera mayottensis, 1869e: 9, pl. 1 fig. 6, 6a (to Polybothris), Mayotte (Comores), syntypes. Not

seen.

Family Cerambycidae

Batocera irregularis, 1871c: 220, pl. 9 fig. 3 (to Abatocera), Menado (Sulawesi), Sangir, at least 2 syn-

types.

Batocera obligua, 1871c: 219, pl. 9 fig. 3, Buru, 3 syntypes.

Epipedocera perelegans, 1871b: 104, pl. 4 fig. 4, Timor, 2 syntypes.

Eurycephalus wieneckii, 1871b: 105, pl. 4 fig. 5 (to Purpuricenus), Timor, 1 syntype.

Leprodera felderii, 1871b: 109, pl. 5 fig. 9, Sumatra, 1 syntype.

Megacriodes guttatus, 1871b: 110, pl. 5 fig. 10 (to Batocera), Sumatra, 2 syntypes.

Phymasterna humeralis, 1869e: 10, pl. 1 fig. 1, la (to Megalofrea), Nossi-Bé (Malagasy), holotype.

Protemnemus rosenbergii, 1871b: 108, pl. 4 fig. 8, Doreh (West New Guinea), holotype. Not seen.

Tmesisternus isabellae, 1871b: 107, pl. 4 fig. 7, Salawatti, 2 syntypes.

Tragocerus heraldicus, 1871b: 106, pl. 4 fig. 6, New Guinea, holotype.

Family Cetoniidae

Chalcothea affinis, 1858c: 23, pl. 2 fig. 2, Borneo, holotype.

Clinteria dives, 1864c: 153, Borneo, Billiton, 2 syntypes.

Clinteria egens, 1864c: 152, Borneo, holotype.

Clinteria vidua, 1864c: 152, pl. 4 fig. 6, West coast of Sumatra, holotype.

Euryomia forsteni, 1864c: 156, pl. 10 fig. 4 (to Glycyphana), Tondano (Sulawesi), 1 syntype.

Euryomia quadriguttata, 1864c: 158 (to Glycyphana), Sumatra, Halmaheira, Batjan, Morotai, 5 synty-

pes.

Euryomia regalis, 1864c: 157 (to Glycyphana), Tondano (Sulawesi), 1 syntype.

Euryomia sieboldii, 1864c: 158 (to Glycyphana), Japan, holotype.

Euryomia sumatrensis, 1864c: 157 (to Glycyphana), Sumatra, holotype.

Heterorhina infuscata, 1864c: 151 (to Smaragdesthes), St. George del Mina (Ghana), 1 syntype.

Macroma flavoguttata, 1864c: 159, pl. 10 fig. 6, (to Campsiura), 2 syntypes. Lectotype, Krikken, 1977.

Macronota aciculata, 1864c: 153 (to Penthima), Java (“vrai habitat?”), holotype.

Macronota aurantiaca, 1858c: 26, pl. 2 fig. 5 (to Taeniodera), Sumatra, holotype not seen.

Macronota forsteni, 1858e: 24 (to Mecinonota), Tondano (Sulawesi), 1 syntype.

Macronota luctuosa, 1858e: 25, pl. 2 fig. 4 (to Mecinonota), Amboina (Moluccas), 2 syntypes.

Macronota ludekingii, 1864c: 154, pl. 10 fig. 1 (to Eumacronota), Fort de Kock (= Bukittinggi, Suma-

tra), 1 syntype.

Macronota suturalis, 1858e: 25, pl. 2 fig. 3 (to Meroloba), Sumatra, 2 syntypes.

Parachilia polleni, 1869e: 9, pl. 1 fig. 5, Sa-c (to Heterosoma), Nossi-Bé (Malagasy), holotype.

Schizorhina nigerrima, 1864c: 156 (to Lethosesthes), Morotai (Indonesia), 2 syntypes.

Schizorhina balteata, 1871f: 277, pl. 6 fig. 3 (to Dilochrosis), Waigeou, holotype.

Schizorhina sanguinolenta, 1864c: 155 (to Sternoplus), Tondano (Sulawesi), 1 syntype.

Trichius septemdecemguttatus, 1864c: 159, Japan, 1 syntype.

Family Cucujidae

Hectarthrum nodicorne, 1864c: 145, pl. 9 fig. 1, Borneo, holotype.

Family Curculionidae

Arachnopus frenatus, 1866e: 228, pl. 12 fig. 5, Salawatti, at least 1 syntype.

Arachnopus geometricus, 1866e: 227, pl. 12 fig. 4, Tondano (Sulawesi), 1 syntype.

Arachnopus persona, 1866e: 226, pl. 12 fig. 3, Waigeou, 2 syntypes.

Danae, 1864c: 16 (to Rhinoscapha), type-sp. Danae lunulata Vollenhoven, 1864 (monotypy).

Danae lunulata, 1864c: 166, pl. 12 fig. 2 (to Rhinoscapha), Morotai (Indonesia), holotype.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 255

Episomus stellio, 1864c: 169, pl. 12 fig. 5, Sumatra, holotype.

Eupholus aurifer, 1864c: 167, pl. 12 fig. 3, Wahaai (South Ceram), holotype.

Eupholus bandanus, 1871b: 102, pl. 4 fig. 2, Banda (Moluccas), at least 5 syntypes.

Eupholus vilis, 1864c: 168 (to Rhinoscapha), Sumatra, at least 2 syntypes.

Macropterus verlorenii, 1871b: 101, pl. 4 fig. 1 (to Esamus), Atapoepoe (Timor) 1 syntype.

Pachyrhynchus forsteni, 1864c: 168, pl. 12 fig. 4, Ternate, Halmaheira, Sumatra, at least 7 syntypes.

Pachyrhynchus morotaiensis, 1864c: 169, Morotai (Indonesia), 2 syntypes.

Protoceraius laetus, 1866e: 228, pl. 12 fig. 1, Sulawesi, syntypes. Not seen.

Rhinoscapha batjanensis, 1864c: 164, Batjan (Moluccas), Kajoa (Sulawesi?), at least 7 syntypes.

Rhinoscapha dohrnii, 1864c: 165, pl. 12 fig. 1, Morotai (Indonesia), 5 syntypes.

Family Dynastidae

Oryctes cristatus, 1858c: 21, pl. 2 fig. 1, St. George del Mina (Ghana), holotype.

Stypotrupes candezei, 1866e: 222, pl. 11 figs. 1—4 (to Ceratoryctoderus), Celebes, 1 syntype.

Trichogomphus alcides, 1864c: 151, pl. 9, Borneo, 2 syntypes. Lectotype.

Trichogomphus simson, 1864c: 150, Sumatra, 3 syntypes. Lectotype.

Family Gyrinidae

Dineutes bidens, 1869e: 7, pl. 1 figs. 1, 2, Mayotte (Comores), at least 2 syntypes.

Family Lucanidae

Dorcus forceps, 1861d: 109, pl. 6 fig. 2 (to Prosopocoilus), Sumatra, 2 syntypes. Lectotype.

Dorcus niponensis, 1861d: 113, pl. 7 fig. 3 (to Macrodorcas), Japan, 1 syntype.

Dorcus pilifer, 1861d: 112, pl. 6 fig. 4 (to Serrognathus), Japan, 2 syntypes.

Dorcus purpurascens, 1861d: 111, pl. 7 figs. 1, 2 (to Serrognathus), Sumatra, 3 syntypes.

Dorcus rectangulus, 1861d: 114, pl. 7 fig. 7 (to Aegus), Sumatra, 1 syntype.

Dorcus tragulus, 1861d: 113, pl. 7 figs. 4, 5, 6 (to Prosopocoilus), Sumatra, 2 syntypes. Lectotype,

Bomans.

Eurytrachelus alcides, 1865i: 150, pl. 10 figs. 2, 3, Sumatra, 1 syntype.

Eurytrachelus rubrofemoratus, 1865i: 152, pl. 11 figs. 1, 2, Japan, syntype.

Gnaphaloryx miles, 1865i: 155, pl. 11 fig. 5, South Halmaheira and Gebeh (Indonesia), 2 syntypes.

Lucanus brookeanus, 1861d: 107, pl. 6 fig. 1 (to Odontolabis), Borneo, 2 syntypes. Lectotype, Bom-

ans.

Lucanus lacordairei, 1861d: 105, pl. 5 fig. 1 (to Odontolabis), Sumatra, 1 syntype.

Lucanus ludekingii, 1861d: 105, pl. 5 fig. 2 (to Odontolabis), Sumatra, 2 syntypes.

Lucanus dejeani var. nigra, 1861d: 108 (to Odontolabis), no type-loc. given, 1 specimen (syntype)

from Sumatra.

Family Melolonthidae

Anomala bivittata, 1869e: 8, pl. 1 fig. 4 (to Prodoretus), Nossi-Bé (Malagasy), holotype.

Parastasia atra, 1864c: 149, Amboina, 1 syntype.

Parastasia degenerata, 1864c: 147, Borneo, Sulawesi, Batjan, Morotai (Indonesia), 5 syntypes.

Parastasia ephippium, 1864c: 148, pl. 9 fig. 4, Sumatra, 2 syntypes.

Parastasia pileus, 1864c: 147, pl. 9 fig. 3, Borneo, Sumatra, Batjan, Halmaheira, 5 syntypes.

Patastasia vittata, 1864c: 149, Amboina, Buru, 6 syntypes.

Pleophylla unicolor, 1869e: 8, pl. 1 fig. 3 (to Phylloserica), Nossi-Bé (Malagasy), holotype.

Family Mordellidae

Anaspis testacea, 1854c: 70, Netherlands, syntypes.

Anaspis assimilis, 1854c: 70, Netherlands, syntypes.

256 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Family Paussidae

Paussus ludekingii, 1872c: 82, Sumatra, 1 syntype (of 2).

Family Scaphidiidae

Apoderus spectrum, 1865k: 159, pl. 3 figs. (to Diatelium), Sumatra, 2 syntypes.

Family Scarabaeidae

Onthophagus schwaneri, 1864c: 146, pl. 9 figs. 2, 2a, Borneo, 2 syntypes.

Family Staphylinidae

Staphylinus pygmaeus, 1854c: 70 (to Philonthus), Netherlands, syntypes. Not seen.

Stenus roscidus, 1854c: 70, Netherlands, 1 syntype.

ORDER DIPTERA

Family Bombyliidae

Systropus blumei, 1863j: 8, pl. 1 fig. 4, Java, holotype, d.

Systropus tessellatus, 1863j: 9, Sumatra, holotype, d.

Family Calliphoridae

Calliphora nigribarbis, 1863j: 17, Japan, 4 2 syntypes. Lectotype, Van Helsdingen, 1980.

key, 1965.

Family Conopidae

Conops lugens, 1863 j: 10, pl. 1 fig. 5 (to Physocephala), Borneo, holotype, 2 (not d!).

Family Dryomyzidae

Family Rachiceridae

Antidoxion, 1863i: 6 (to Rhachicerus Walker), type-sp. Antidoxion fulvicorne Vollenhoven (monoty-

Py).

Family Stratiomyidae

Ptilocera amethystina, 1858d: 92, Java, Celebes, 2 ? syntypes.

Ptilocera smaragdina, 1858d: 92, Menado (Sulawesi), 2 2 d syntypes.

Family Syrphidae

Milesia apicalis, 1863j: 13, Java, holotype, d.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 257

Milesia diardi, 1863j: 14, pl. 1 fig. 6, Borneo, 3 d syntypes.

Milesia undulata, 1863j: 12, Japan, holotype, 9.

Family Tachinidae

Crosskey, 1973.

ORDER HYMENOPTERA

Many of the Hymenoptera were described from the Netherlands; localities lac-

king country names are situated in the Netherlands. The families are arranged ac-

cording to the usual grouping in Symphyta, Parasitica and Aculeata. In the non-

Ichneumonid families records of type material mean that the specimens are in the

Leiden museum. For the Ichneumonid types the presence in Leiden and Wage-

ningen is specified. In a forthcoming paper K. W. R. Zwart will give more details

concerning Ichneumonid types, including some lectotype designations.

SUBORDER SYMPHYTA

Family Argidae

Hylotoma flava, 1860e: 129 (to Arge), Japan, 2 9 syntypes. Not seen.

Hylotoma humeralis, 1860e: 129 (to Arge), Japan, holotype not seen.

Hylotoma microcephala, 1860e: 130 (to Arge), Java, 3 2 syntypes.

Hylotoma similis, 1860e: 128 (to Arge), Japan, 4 ? syntypes. Not seen.

Family Cephidae

Cephus elongatus, 1858b: 280 (to Calameuta), Sterkenburg near Driebergen; near Heemstede; 4 9, 1

3; 1 syntype from first locality seen.

Family Cimbicidae

Amasis sanguinea, 1878f: 154, pl. 9 (to Corynis), Morocco, holotype. Not seen.

Family Diprionidae

Lophyrus catocalus, 1858b: 276 (to Gilpinia), Groesbeek, 2 9, 2 d syntypes.

Family Pamphiliidae

Lyda maculifrons, 1858b: 179 (to Neurotoma), near Utrecht, holotype, d. Not seen.

Lyda nigricornis, 1858b: 279 (to Pamphilius), near Den Haag, holotype, d.

Family Tenthredinidae

Dolerus busaei, 1858b: 277, Haarlem, 4 syntypes.

Emphytus majalis, 1869b: 121, pl. 3, near Utrecht. Holotype not seen.

Macrophya histrionica, 1878f: 155, surroundings of Baireuth, holotype, 2. Not seen.

Macrophya klugii, 1869b: 124, Noord-Brabant; near Den Haag, 2 d syntypes. Not seen.

258 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Nematus aquilegiae, 1866b: 202, pl. 9 (to Pristiphora), Renkum, 4 syntypes.

Nematus catachloris, 1869b: 120, pl. 3, near Leyden. Type not seen.

Nematus consobrinus, 1871d: 237, pl. 10, Haarlem, 10 9, 1 & syntypes. Not seen.

Nematus immaculatus, 1869b: 119, Schoten; near Rotterdam, syntypes. Not seen.

Nematus lugdunensis, 1871d: 243, pl. 11, dunes near Noordwijk, 9 d syntypes. Not seen.

Nematus pectoralis, 1867d: 223, surroundings of Utrecht, holotype. Not seen.

Nematus solea, 1870a: 59, pl. 1, Netherlands, from Larix, holotype, 4. Not seen.

Nematus trimaculatus, 1862a: 69, pl. 4, Leiden, 2 syntypes (of 6).

Nematus vallator, 1858h: 191, pl. 12 (to Pristiphora), Leiden, 2 syntypes.

Nematus wttewaalli, 1862a: 65, pl. 4, Voorst; Utreht; Velp; near Leyden; Sterkenburg near Drieber-

gen; Ter Wadding between Leyden and Voorschoten; only one syntype present.

Phyllotoma pinguis, 1869b: 121, pl. 3 (to Heterarthrus), near Den Haag, holotype, 2.

Selandria albomarginata, 1858b: 278 (to Sciapteryx), near Den Haag; near Utrecht, 3 syntypes. Not

seen.

Selandria cereipes, 1873b: 13, pl. 3, Gelderland, “d. Roo, Velp”, 3 syntypes seen (out of 12 9).

Selandria humeralis, 1869b: 122 (to Blennocampa), Gliphoeve, holotype, 9.

Selandria phthisica, 1869b: 123, pl. 3 (to Stromboceros), Beekbergerwoud, holotype.

Selandria sixii, 1858b: 278, near Utrecht, holotype, 9.

Selandria soror, 1869b: 123, Leyden, near Utrecht, syntypes. Not seen.

SUBORDER APOCRITA

Division Parasitica

Family Braconidae

Alysia theodori, 1878f: 174, pl. 11 fig. 3 (to Phaenocarpa), dunes of Loosduinen, holotype, 2.

Laccophrys medenbachii, 1878f: 173, pl. 11 fig. 2 (to Opius?), Surroundings of Arnhem, holotype, 9.

Not seen.

Laccophrys villaenovae, 1876e: 243 (nom. nud.); 1878h: 172, pl. 11 fig. 1 (to Opius), Naaldwijk; Zee-

land; d 2 syntypes. Not seen.

Macrocentrus flavus 1878h: 54, pl. 34 fig. 3, Glogau (Poland), 2 d syntypes. Not seen.

Mesora analis, 1878f: 175, pl. 11 fig. 4 (to Dacnusa), Biggekerke in Zeeland, holotype, 4. Not seen.

Microdus compeditus, 1876e: 243 (nom. nud.), near Venlo; specimens not seen, synonymized by Vol-

lenhoven (1878f: 173) with Agathis rugulosus (Nees, 1834).

Phylax aestivalis, 1858b: 282 (to Aleiodes), near Den Haag, holotype, 9. Synonymized by Vollenho-

ven (1873e: 193) with Rogas dissectus Nees, 1834.

Polemon albimanus, 1873f: 219 (to Coelinius), near Vogelenzang. Type not seen.

Rogas pubescens, 1879a: XXVI (nom. nud.), St. Jansberg, Mook.

seen. Synonymized by Vollenhoven (1878g: 46) with Agathis syngenesiae Nees, 1814.

Family Cynipidae

Aulax albinervis, 1869b: 126. Driebergen, holotype, 2. Not seen.

Family Diapriidae

Loxotropa sabuleti, 1879d: 62, pl. 40 fig. 2 (to Spilomicrus), Netherlands. Type not seen.

Spilomicrus major, 1879d: 63, pl. 40 fig. 7, Burgdorff (Switzerland), Den Haag, 2 d syntypes. Not

seen.

Family Eucoilidae

Eucoila inflata, 1867d: 224, pl. 10, figs. 2—3, surroundings of Utrecht; Driebergen, 2 syntypes. Not

seen.

J. KRIKKEN ET AL.: Snellen van Vollenhoven 259

Family Ichneumonidae

Agrothereutes batavus, 1873f: 209, pl. 9 fig. 1, Zuid-Holland, holotype, 9 . (Wageningen).

Agrothereutes destitutus, 1879f: 58, pl. 37 fig. 4, no type locality mentioned, holotype, ?. Not seen.

Anomalon bucephalum, 1858b: 281 (to Heteropelma), Walcheren and Warmond, 4 g syntypes (Lei-

den and Wageningen). Synonymized by Vollenhoven (1876e: 237) with Anomalon amictum F.

Anomalon melanocneme, 1878f: 168, pl. 10 fig. 5, near Montpellier (France), holotype, 9 . Not seen.

Aptesis longicauda, 1873f: 210, Wort-Rhede in Gelderland, holotype, ? (Wageningen).

Atractodes spiniger, 1876e: 239 (nom. nud.); 1878f: 171, pl. 10 fig. 8, near Leyden, holotype, d. Not

seen.

Bassus monticola, 1875e: 4, pl. 1 fig. 9 (to Sussaba), Switzerland, holotype (sex not given). Not seen.

Bassus quadriguttatus, 1873f: 211, Leyden, holotype, 2 (Wageningen).

Coleocentrus abiae, 1873f: 215, pl. 9 fig. 3 (to Perispuda), no type-locality mentioned (in the Nether-

lands), holotype, 4 (Wageningen).

Coleocentrus sixii, 1873f: 216, pl. 9 fig. 4, near Utrecht, holotype, d. Not seen.

Colpotrochia affinis, 1875f: 12, pl. 8 fig. 2, Arnhem, holotype (sex not given). Not seen.

Cremastus balteatus, 1878f: 169, pl. 10 fig. 6, Breda, holotype, d. Not seen.

Cremastus cabalisticus, 1876e: 239 (nom. nud.). In 1878f: 169 Vollenhoven states that the specimen

does not belong to a new species, nor is it a Cremastus. No specimen seen.

Cremastus sabulosus, 1876e: 239 (nom. nud.); 1878f: 170, pl. 10 fig. 7, near Scheveningen, 2 d synty-

pes. Not seen.

Crypturus niger, 1878f: 158, pl. 9 fig. 3, (Southern) France, holotype, d (?). Not seen.

Eclytus semiluctuosus, 1878h: 51, pl. 32 fig. 5, near Driebergen and Roozendaal, syntypes (sex not

given) (1 syntype Leiden; heavily damaged).

Ephialtes melanomerus, 1878c: 48 (to Dolichomitus), Java, holotype, ? (Leiden).

Euceros unifasciatus, 1876e: 229 (nom. nud.); 1878f: 159, near The Hague, Leyden, [var. near Breda],

2 2 syntypes. Not seen. In 1878h: 53 Vollenhoven states that it might be a variety of Euceros egre-

gius.

Exetastes cimbicis, 1858b: 281, Zwammerdam ex Cimbex aenea (1858a: 244) (to Perispuda), d + ?

syntypes (1 d, 1 2 Wageningen, 2 specimens Leiden).

Exochus semilividus, 1875f: 13, pl. 8 fig. 11, Switzerland, holotype (sex not given). Not seen.

Exophanes [sic !] fulvescens, 1875e: 5, pl. 2 fig. 5, Rotterdam, holotype 9. Not seen.

Glypta elegans, 1873f: 214, pl. 9 fig. 2, Beverwijk, 2 d syntypes (Wageningen).

Lissonota cruenta, 1858b: 281, near Brummen (1858a: 241), 3 syntypes (2 Wageningen, 1 Leiden).

Synonymized with Lissonota lateralis Grav. bij Vollenhoven 1873e: 178.

Notopygus fulvipes, 1878h: 50, pl. 31 fig. 5, no type locality given, probably from Sweden, holotype

(sex ?). Not seen.

Notopygus minkii, 1878h: 49, pl. 31 fig. 4, no type locality given, probably from Germany, holotype,

9. Not seen.

Ophion monostigma, 1879f: 62, pl. 39 fig. 8 (to Enicospilus), no type locality given, holotype (sex ?).

Not seen.

Ophion tournieri, 1879f: 61 (to Enicospilus); in 1878g: pl. 28 fig. 7 figured as Ophion repentinus Hlg.,

type locality not given, probably near Geneva (Switzerland), holotype d . Not seen.

Ophion unicallosus, 1878f: 167, pl. 10 fig. 4 (to Enicospilus), Espagne (Spain), holotype ?. Not seen.

Paniscus ornatus, 1973f: 218 (to Netelia), Drever in Groningen, holotype, 2 (Wageningen).

Perilissus singularis, 1878h: 52, pl. 32 fig. 8, near Rotterdam, holotype, d (Leiden).

Phidias, 1878f: 164 (= Diacritus Foerster, 1868), type-sp. Phidias aciculatus Voll., 1878 (monotypy).

Preoccupied by Phidias Rafinesque, 1815.

Phidias aciculatus, 1878f: 165, pl. 10 fig. 2 (to Diacritus), Angleterre (England), holotype, 2. Not

seen.

Phytodietus exareolatus, 1876e: 236 (nom. nud.); 1878f: 163 (as Phytodictus [sic !], pl. 9 fig. 5, Voorst

in Gelderland, holotype, 2. Not seen.

Pimpla basalis, 1879e: 148 (to Lissopimpla), Sumatra, holotype, 4 (Leiden).

Pimpla caligata, 1877f: 34, pl. 21 fig. 8, near Scheveningen, holotype, d. Not seen.

Pimpla elegans, 1879e: 147 (to Xanthopimpla), Java, holotype, ? (Leiden).

Pimpla laevidorsum, 1873f: 213 (to Scambus), Voorst, holotype, 2 (Wageningen).

Pimpla sagrae, 1878b: LXXVI (nom. nud.); 1879e: 149 (to Sericopimpla), Buitenzorg, Java, lectotype

260 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Pimpla transversalis, 1879e: 146 (to Xanthopimpla), Sumatra, Borneo, Timor, Thibet, lectotype 2

Pimpla vincta, 1873f: 212, Holland, Zeeland, Utrecht, Gelderland (Provinces), 1 9 syntype (Wage-

ningen).

Pimpla zebra, 1879e: 147 (to Theronia), Ambarawa, Java, holotype, 9 (Leiden).

Platylabus pictus, 1878f: 157, pl. 9 fig. 2, surroundings of Montpellier (France), holotype, d. Not

seen.

Rhyssa bernsteinii, 1879e: 138 (to Myllenyxis), Salawatti or Waigeou (Indonesia) (label destroyed du-

Rhyssa doreica, 1873c: 69, pl. 4 fig. 3 (to Lytarmes), Dorey, New Guinea, holotype, 2 (Leiden).

syntypes (Leiden).

den).

Rhyssa fulva, 1879e: 136 (to Lytarmes), Obi (Indonesia), holotype, & (Leiden).

Rhyssa laniaria, 1873c: 68, pl. 4 fig. 2 (to Megarhyssa), Amboine (= Ambon) (Indonesia), holotype,

2 (Leiden).

Rhyssa mülleri, 1879e: 140 (to Myllenyxis), Java, holotype, 9 (Leiden).

Scolobates corrallinus, 1873f: 211, Wikkenberg near Schalkwijk, 4 9 syntypes (Wageningen) (of 5).

Scolobates marshalli, 1878f: 166, pl. 10 fig. 1, Pyrenées (France), holotype, 4. Not seen.

Syene, 1878b: LXXVI (= Echthromorpha Hlg., 1868), no species included, type-sp. Cryptus notulato-

rius F. (= E. agrestoria notulatoria), designated by Townes, 1960.

Trichomastix, 1878f: 160 (= Bioblapsis Foerster, 1868), type-sp. Trichomastix polita Voll., 1878 (mo-

notypy).

Trichomastix polita, 1878f: 161, pl. 9 fig. 4 (to Bioblapsis), Scheveningen, holotype, 2. Not seen.

Family Proctotrupidae

Proctotrupes sixianus, 1879a: XIII, Netherlands, Type not seen.

Family Pteromalidae

Hetroxys gribodii, 1878f: 176, pl. 11, North Italy, 2 9 syntypes. Not seen.

Lagynodes pallipes, 1873e: 205 (nom. nud.); 1876e: 253 (nom. nud.). Synonymized by Vollenhoven

(1876e: 249) with Theocolax formiciformis Westwood. No specimens seen from “Leyden”.

Division ACULEATA

Family Apidae

Bombus senex, 1873g: 229, pl. 10, Sumatra, holotype, 9.

Nomada batava, 1858b: 283, near Leiden, holotype not seen, may be a specimen under Nomada goo-

deniana Kirby, 1802.

Family Dryinidae

Dryinus spectrum, 1874c: (LXVI) (nom. nud.); 1874b: 159, figs. 34, from “Vogelzang” (= Vogelen-

zang), holotype, ?.

Goniozus tibialis, 1876e: 253 (nom. nud.); 1878f: 177, pl. 11, near Den Haag, one of the 2 syntypes

seen.

ORDER LEPIDOPTERA

For the family classification of the Papilionoidea Kristensen (1976) has been

followed. The family classification of the other Lepidoptera is in conformity with

J. KRIKKEN ET AL.: Snellen van Vollenhoven 261

the dictionary of Laithwaite et al. (1975). No attempt has been made to indicate

the genera the taxa described by Snellen van Vollenhoven are at present suppo-

sed to belong to, as there is no general agreement on the generic classification of

many families. Most species-group names of the list below can be found in the Le-

pidopterorum Catalogus, where further references are given.

Family Agaristidae

Ophthalmis bernsteinii, 1863f: 132, pl. 9 fig. 1, Morotai, holotype, 9.

Family Arctiidae

Crambomorpha (?) aurora, 1873h: 246, pl. 12 fig. 2, Salawatti. Type not seen.

Gnophria (?) ceramensis, 1873h: 246, pl. 12 fig. 1, Ceram, holotype, d (?).

Hypsa orbona, 1863f: 137, pl. 9 fig. 4, Halmaheira, Morotai, 4 syntypes.

Leptosoma assimile, 1863h: 39 (to Nyctemera, the following species as well), Java, 2 d syntypes.

Leptosoma clathratum, 1863h: 48, Amboina, holotype, 9.

Leptosoma herklotsii, 1863h: 39, Java. Type not seen.

Leptosoma inconstans, 1863h: 47, Java, Borneo, 4 4,5 2 syntypes.

Leptosoma luctuosum, 1863h: 42, Batjan, holotype, d.

Leptosoma ludekingii, 1863h: 49, Sumatra. Type not seen.

Leptosoma macklotti, 1863h: 50. Amboina, holotype, 9.

Leptosoma mülleri, 1863h: 41. Sumatra, 2 9 (?) syntypes.

Leptosoma noviespunctatum, 1863h: 42. Java, 3 2 syntypes.

Leptosoma nubecula, 1863h: 45. Java, 2 d syntypes.

Leptosoma pallens, 1863h: 45. Java, holotype, d.

Leptosoma quadriguttatum, 1863h: 40, Java, 2 d syntypes.

Numenes trigonalis, 1863f: 140, pl. 10 fig. 1, Sumatra, 2 2 syntypes.

Numenes virginalis, 1863f: 142, Sumatra, holotype, ®.

Spilosoma roseiventer, 1863f: 143, Japan. 2 d, 1 9 syntypes.

Family Elachistidae

Atteva apicalis, 1863f: 139, pl. 9 fig. 5, Java, 2 syntypes.

Atteva basalis, 1863f: 140, pl. 9 fig. 6, Morotai, 7 syntypes.

Family Geometridae

Milionia flammula, 1863f: 134, pl. 9 fig. 2, Nova Guinea, holotype, d (?).

Milionia fulgida, 1863f: 133, pl. 8 fig. 4, Java, 1 d, 1 9 syntypes.

Family Lymantriidae

Lymantria (Liparis) hilaris, 1863f: 143, pl. 10 figs. 2, 3, Sumatra (interior), 1 4,3 2 syntypes.

Family Noctuidae

Phyllodes eyndhovii, 1859b: 86, pl. 6, Java. Type not seen.

Phyllodes verhuellii, 1858f: 159, pl. 8. Java, holotype, 3.

Family Nymphalidae

Acraea dammi, 1869e: 12, pl. 2 fig. 4, Nossi-Bé (Malagasy), holotype, à.

262 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Adolias agnis, 1862f: 202, pl. 12 fig. 2; 1862h: 273, Java, 1 9 syntype.

Adolias apicalis, 1862f: 186, pl. 10 fig. 1; 1862h: 270, Borneo, holotype, à.

Adolias apsarasa, 1862f: 198, pl. 11 fig. 3; 1862h: 272, Borneo, holotype, d.

Adolias bipunctata, 1862f: 191, pl. 10 fig. 4; 1862h: 271, Borneo, holotype, d.

Adolias blumei, 1862f: 205, pl. 12 figs. 3, 4; 1862h: 273, Java, Sumatra (Padang), syntypes, probably

several specimens.

Adolias clathrata, 1862f: 205, pl. 12 fig. 5; 1862h: 273, Borneo, holotype, 4.

Adolias diardi, 1862f: 188, pl. 10 fig. 2; 1862h: 270, Borneo, holotype, à.

Adolias ludekingii, 1862f: 189, pl. 10 fig. 3; 1862h: 270, Padang (Sumatra), holotype, 9.

Adolias octogesima, 1862f: 193, pl. 10 fig. 5, pl. 11 fig. 1; 1862h: 271, Java, Borneo, 1 2 syntype.

Adolias pardalis, 1862f: 197, pl. 11 fig. 5; 1862h: 272, Java, 1 d syntype.

Adolias surjas, 1862f: 200, pl. 12 fig. 1; 1862h: 272, Java, holotype, 2.

Charaxes cognatus, 1861e: 159, pl. 9 figs. 1, 2, Moluques, holotype, 3.

Charaxes concha, 1861e: 162, pl. 10 figs. 1, 3, Java, Padang (Sumatra), 2 &, 1 9 syntypes.

Charaxes rayi, 1861e: 161, pl. 9 fig. 3, pl. 10 fig. 2, Ambon, holotype, 4.

Drusilla anableps, 1860a: 40, pl. 1 fig. 3, pl. 2 fig. 1, Otaheiti, holotype, à.

Drusilla artemis, 1860a: 37, pl. 1 figs. 1, 2, Nieuw-Guinea, holotype, 4.

Drusilla dioptrica, 1860a: 38, pl. 2 figs. 2, 3, Nieuw-Guinea, holotype, à.

Drusilla diops, 1863f: 131, pl. 8 fig. 3, Morotai, Halmaheira, 2 d syntypes.

Euploea mesocala, 1873h: 244, pl. 11 figs. 1,2, Waigeou, 1 d,1 2, syntypes.

Euploea superba, 1866c: 209, pl. 10 fig. 1, Gorontalo (Sulawesi), 6 d syntypes.

Idea tondana, 1860a: 41, pl. 4, Tondano (Sulawesi), 1 &, 1 9 syntypes.

Melanitis stellaris, 1861e: 159, pl. 8 fig. 3, Nouvelle Guinée, holotype, 9.

Minetra tigrina, 1866c: 210, pl. 10 fig. 2, Salawatti, 2 d syntypes.

Mynes mulderi, 1863f: 129, pl. 8 figs. 1, 2, Batjan, Halmaheira, 3 d syntypes.

Vanessa royeri, 1861e: 157, pl. 8 figs. 1, 2, Nouvelle Guinée. holotype, 9.

Family Papilionidae

Papilio trogon, 1860b: 69, pl. 6 figs. 1, 2, Sumatra, 2 d syntypes.

Family Pieridae

Pieris affinis, 18651: 40, pl. 5 fig. 2, Celebes sept., holotype, d.

Pieris amalia, 18651: 23, pl. 3 fig. 6, Sumatra, Banca, 1 d syntype.

Pieris candida, 18651: 11, pl. 3 fig. 2, Batjan. Type not seen.

Pieris chrysomelaena, 1866a: 57, pl. 1 figs. 1, 2, Kajoa, holotype, à.

Pieris chrysorrhoea, 18651: 6, pl. 2 fig. 4, Sumatra, holotype, à.

Pieris cornelia, 18651: 5, pl. 2 fig. 2, Borneo, holotype, d.

Pieris dice, 18651: 39, pl. 4 fig. 7, Nova Guinea, Waigeou, 2 d,1 2 syntypes.

Pieris elisa, 1869e: 12, pl. 2 fig. 3, Mayotte (Comores), 1 4, 1 9 syntypes.

Pieris emma, 18651: 24, pl. 4 fig. 2, Batjan, 2 9 syntypes.

Pieris haemorrhoea, 18651: 10, pl. 2 fig. 5, Banca, 1 6, 1 2 syntypes.

Pieris hagar, 18651: 38, pl. 4 fig. 6, Padang (Sumatra), holotype, d.

Pieris herodias, 18651: 14, pl. 3 fig. 4, Galela (Halmaheira), 1 6, 1 9 syntypes.

Pieris poecilea, 18651: 13, pl. 3 fig. 3, Halmaheira, Morotai, 1 4, 2 9 syntypes.

Pieris rosenbergii, 18651: 11, pl. 2 fig. 6, pl. 3 fig. 1, Celebes sept., 3 4,2 2 syntypes.

Pieris zoe, 18651: 37, pl. 4 fig. 5, Batjan, holotype, à.

J. KRIKKEN ET AL.: Snellen van Vollenhoven

263

Pontia lignea, 18651: 4, pl. 2 fig. la, b, Bone, Gorontalo (Sulawesi), 3 d syntypes.

Terias impura, 18651: 70, pl. 7 fig. 5, Timor, holotype, 1 d (?).

Terias tominia, 18651: 66, pl. 7 fig. 4, Celebes sept., 1 4,2 2 syntypes.

Thestias ludekingii, 1860d: 125, Sumatra (interior), holotype, 3.

Thestias reinwardtii, 1860d: 126, Moluques, holotype, 4.

Family Pyralidae

Deiopeia occultans, 1869e: 13, pl. 2 fig. 5, Nossi-Bé (Malagasy), holotype, d.

Saturnia insularis, 1862g: 335, pl. 14 fig. 1, Java, Sumatra, 4 d syntypes.

Family Saturniidae

Family Zygaenidae

Agalope westwoodii, 1863f: 136, pl. 9 fig. 3, Japan, holotype, d.

Chalcosia candida, 1863f: 135, Sumatra, 2 syntypes.

Leptosoma arcuatum, 1863h: 45, Java, 2 d syntypes.

Leptosoma marginale, 1863h: 43, Java, holotype, à.

Abatocera 254

abiae 259

Acatalectus 249

aciculata 254

aciculatus 259

Acraea 261

Acrosternum 251

acuta 250

Adolias 262

Adrisa 249

Aegus 255

aenea (Cimbex) 259

aenea (Dalpada) 250

aeneiventris 252

aestivalis 258

Aethus 249

affinis (Colpotrochia) 259

affinis (Chalcothea) 254

affinis (Pieris) 262

Agalope 263

agathina 253

Agathis 258

agnis 262

agrestoria 260

Agrothereutes 259

albimanus 258

albinervis 258

alboguttatus 253

albomarginata 258

INDEX

alcides (Eurytrachelus) 225

alcides (Trichogomphus)

[255

Alcimocoris 249

Alcimus 249

Alebra 253

Aleiodes 258

Alysia 258

amabilis 253

amalia 262

Amasis 257

amethystina (Ptilocera) 256

amethystina (Scutellera)

[252

amictum 259

Amyotea 249, 250

amyoti 250

anableps 262

anacanthum 250

analis 258

Anaspis 255

Anomala 255

Anomalon 259

Anoplocephala 250

anser 253

Antestia 251

antica 250

Antidoxion 256

apicalis (Adolias) 262

apicalis (Atteva) 261

apicalis (Canthecona) 250

apicalis (Jassus) 253

apicalis (Milesia) 256

Apoderus 253, 256

apsarasa 262

Aptesis 259

Arachnopus 254

aquilegiae 258

Archiblatta 248

arcuatum 263

Arge 257

arlechino 251

artemis 262

Asopus 249, 250

Aspidestrophus 250

Aspongopus 250

assimile 261

assimilis 255

atra 255

Atractodes 259

Atteva 261

Aulax 258

aurantiaca (Macronota)

[254

aurantiaca (Typhlocyba)

253

aurantiacus 249

aurifer 255

264 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

aurora 261

balteata 254

balteatus 259

bandanus 255

basalis (Atteva) 261

basalis (Pimpla) 259

basimaculata 250

Bassus 259

batava 260

batavus 259

batjanensis 255

Batocera 254

bernsteinii (Asopus) 249

bernsteinii (Oncomerus)

[251

bernsteinii (Ophthalmis)

[261

bernsteinii (Rhyssa) 260

Berytinus 249

Berytus 249

bidens 255

biguttata 250

bilunulata 252

Bioblapsis 260

bipunctata 262

bison 251

bivittata 255

Blachia 250

Blennocampa 258

blumei (Adolias) 262

blumei (Systropus) 256

Bolbocoris 252

Bombus 260

bosschei 252

Bostrichus 253

Brachycerocoris 251

Brachyplatys 251

breviscutum 251

brookeanus 255

bucephalum 259

busaei 257

cabalisticus 259

caesar 252

Calameuta 257

Calidea 252

caligata 259

Callidea 252

Calliphara 252

Calliphora 256

Calliscyta 252

Calodema 253

Campsiura 254

candezei 255

candida (Chalcosia) 263

candida (Pieris) 262

Cantao 252

Canthecona 250

Cantheconidea 250

Captosoma 251

Carbula 250

carnifex 249

Carpona 250

catachloris 258

catocalus 257

Catoxantha 253

Cazira 250

celebensis (Callidea) 252

celebensis (Rhaphigaster)

[251

Cephus 257

Ceraleptus 249

ceramensis 261

Ceratoryctoderus 255

cereipes 258

Certonotus 260

Chalcocybebus 253

Chalcophora 253

Chalcophorella 253

Chalcosia 263

Chalcothea 254

Charaxes 262

Chlaenocoris 250

chloris 251

Chlorocoris 250

Choerommatus 249

Chrysobothris 254

Chrysochroa 253

Chrysocoris 252

Chrysodema 253

chrysomelaena 262

chrysoptera 251

chrysorrhoea 262

Cimbex 259

cimbicis 259

cingulatus 256

clathrata 262

clathratum 261

Clinteria 254

coccinelloides 250

coecus 250

coelestis 251

Coelinius 258

Coenus 250

cognatus 262

Coleocentrus 259

Coleotichus 252

collaris 249

Colpotrochia 259

compeditus 258

Compsapoderus 253

concha 262

Conops 256

Conostethus 249

consobrinus 258

consul 252

Coptosoma 251

Coreus 249

Coridius 250

cornelia 262

corrallinus 260

Corynis 257

Crambomorpha 261

Cremastus 259

cristatus 255

croesus 252

Cronion 252

cruenta 259

crux (Brachyplatys) 251

crux (Dalpada) 250

Crypturus 259

Cryptus 260

Cuspicona 250

Cyclopelta 250

Cycnotrachelus 253

Cyrtomenus 249

Dacnusa 258

daedalica 252

Dalcantha 250

Dalpada 250

dalpadoides 250

dammi 261

Danae 254

decora 250

decorata 250

degenerata 255

Deiopeia 263

dejeani 255

destitutus 259

Diacritus 259

diardi (Adolias) 262

diardi (Idea) 262

diardi (Milesia) 257

Diatelium 256

dice 262

Diceraeus 250

difficilis 249

Dilochrosis 254

Dineutes 255

diops 262

dioptrica 262

dissectus 258

distigma 249

ditissima 252

divergens 250

dives 254

dohrnii 255

Dolerus 257

Dolichomitus 259

Dorcus 255

doreica 260

driebergensis 249

dromedarius 251

Drusilla 262

Dryinus 260

Dryobius 252

Dryomyza 256

Dryptocephala 250

Echthromorpha 260

Eclytus 259

Ectenus 251

effluens 258

egens 254

egregius 259

elegans (Glypta) 259

elegans (Pimpla) 259

Elemana 251

elisa 262

elongata 252

elongatus 257

eltio 250

emarginatus 252

emma 262

Emphytus 257

Enicospilus 259

Ephialtes 259

ephippium 255

Epipedocera 254

Episomus 255

erythrogaster 253

Esamus 255

Euamphibolia 257

Euceros 259

Eucoila 258

Eumacronota 254

Eupholus 255

Euploea 262

Eurhadina 253

Eurycephalus 254

Eurygaster 252

Eurytrachelus 255

Euryomia 254

euschistoides 250

Eusthenes 250

exareolatus 259

Exetastes 259

eximia 252

Exochus 259

Exophanes 259

eyndhovii 261

Eysarcoris 250

fastuosa (Callidea) 252

fastuosa (Typhlocyba) 253

fatime 262

J. KRIKKEN ET AL.: Snellen van Vollenhoven

Fecelia 250

felderi 262

felderii 254

flammula 261

flava 257

flavescens 261

flaviceps 260

flavicornis 251

flavoguttata 254

flavoguttatus 251

flavomarginatus 249

flavus 258

forceps 255

formiciformis 260

Formosia 257

forsteni (Coptosoma) 251

forsteni (Euryomia) 254

forsteni (Macronota) 254

forsteni (Pachyrhynchus)

[255

fractinervis 260

frenatus 254

fulgida 261

fulva 260

fulvescens 259

fulvicorne 256

fulvipes 259

fusca 252

fuscus (Coleotichus) 252

fuscus (Megarhynchus) 250

gandarva 262

geminatus 250

geometricus 254

Geotomus 249

gibba 250

gibbosa 252

gigas 256

Gilpinia 257

Glaucias 251

gloriosa 252

Glycyphana 254

Glypta 259

Gnophria 261

Goniozus 260

goodeniana 260

gribodii 260

guttatus 254

Gynenica 250

haematica 250

haematicus 251

haemorrhoea 262

hagar 262

Hectarthrum 254

hemichloris 251

hemixantha 253

265

heraldicus 254

herklotsii 261

herodias 262

hester 262

Heterarthus 258

Heteropelma 259

Heterorhina 254

Heterosoma 254

Hetroxys 260

hilaris (Lymantria

[(Liparis)) 261

hilaris (Pentatoma) 251

histrionica 257

hoevenii 248

Hoplistodera 250

hoplites 250

Hotea 252

humeralis (Hylotoma) 257

humeralis (Phymasterna)

[254

humeralis (Selandria) 258

Hylotoma 257

hypherythra 252

hypomelaena 252

Hypsa 261

Hyrmine 251

Idea 262

ignobilis 251

immaculatus 258

imperator 249

impura 263

inconstans 261

indras 262

inflata 258

infuscata 254

insignis 249

instabilis 251

insularis (Saturnia) 263

insularis (Vipio) 258

intermedius 250

Iphias 262

iracundus 253

irregularis 254

isabellae 254

Jassus 253

javanica 251

johannae 253

klugii 257

Laccophrys 258

lacordairei 255

laetus 255

laevidorsum 259

Lagynodes 260

266 TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

Lampromicra 252

laniaria 260

latefasciata 252

lateralis 259

Lecanium 253

Leprodera 254

Leptapoderus 253

Leptosoma 261, 263

Lethosesthes 254

Libyssa 252

lignea 263

ligneus 252

limosina 249

lineola (Aspidestrophus)

[250

lineola (Eysarcoris) 250

Linnavuoriana 253

Liparis 261

Lissonota 259

Lissopimpla 259

longeciliatus 253

longicauda 259

Lophyrus 257

Lopus 249

Loricula 249

Loxa 250

Loxotropa 258

Lucanus 255

luctuosa 254

luctuosum 261

ludekingii (Adolias) 262

ludekingii (Chrysochroa)

[253

ludekingii (Leptosoma) 261

ludekingii (Lucanus) 255

ludekingii (Macronota) 254

ludekingii (Paussus) 256

ludekingii (Rhaphigaster)

[251

ludekingii (Thestias) 263

lugdunensis 258

lugens 256

lunulata 254

luteomarginatus 249

Lyda 257

Lygus 249

Lymantria 261

Lyramorpha 250

Lytarmes 260

macklotti 261

Macrocentrus 258

Macrodorcas 255

Macrolyristes 249

Macroma 254

Macronota 254

Macrophya 257

Macropterus 255

maculifrons 257

majalis 257

major 258

malayanus 252

marginale 263

marginepunctata 252

marmoratum 251

marmoratus 250

marshalli 260

mayottensis 254

Mecinonota 254

medenbachii (Laccophrys)

[258

medenbachii (Typhlocuba)

[253

Megacriodes 254

Megalofrea 254

megalops 251

Megaloxantha 253

Megarhynchus 250

Megarhyssa 260

Megarrhamphus 250

Megaspis 256

Megophthalmus 253

Megymenum 250

Melanitis 262

melanocneme 259

melanomerus 259

melanosticticus 251

Menida 250, 251

Meroloba 254

mesocala 262

Mesora 258

microcephala 257

Microdus 258

Microphysa 249

miles 255

Milesia 256, 257

Milionia 261

minax 251

Minetra 262

minkii 259

minor (Eusthenes) 250

minor (Loxa) 250

mitis 250

modesta 252

modestum 251

modestus 251

monostigma 259

monticola 259

Montrouzierellus 250

Mormidea 250

morotaiensis 255

Mucanum 251

muelleri (Aspongopus) 250

muelleri (Coptosoma) 251

mulderi 262

mülleri (Leptosoma) 261

mülleri (Rhyssa) 260

Myllenyxis 260

Mynes 262

Nematus 258

Netelia 259

Neurotoma 257

niger (Apodorus) 253

niger (Choerommatus) 249

niger (Crypturus) 259

nigerrima 254

nigra 255

nigribarbis 256

nigricornis 257

niponensis (Conops) 256

niponensis (Dorcus) 255

nitens (Chalcocybebus) 253

nitens (Rhaphigaster) 251

nitidissima 256

nodicorne 254

Nomada 260

Notopygus 259

notulatorius 260

noviespunctatum 261

nubecula 261

Numenes 261

nurus 251

Nyctemera 261

obliqua 254

obscurus 250

occultans 263

octogesima 262

Odontolabis 255

Oedipoda 249

olor 253

Oncomerus 251

Onthophagus 256

Ophion 259

Ophiotettix 249

Ophthalmis 261

Opius 258

Oplomus 251

orbona 261

ornatus 259

Oryctes 255

Pachycoris 252

Pachyrhynchus 255

Padaeus 250

pallens 261

palliatus 253

pallidicornis 249

pallidiventris 251

pallidus 252

pallifrons 251

J. KRIKKEN ET AL.: Snellen van Vollenhoven 267

pallipes 260 Polemon 258 rosenbergii (Protemnemus)

Pamphilius 257 polita 260 [254

Paniscus 259 polleni 254 roseus (Chlorocoris) 250

Papilio 262 Polybothris 254 roseus (Pygoplatys) 251

Parachilia 254 Polytes 252 royeri 262

Paracoleotichus 252 Pontia 263 rubescens 251

Paracritheus 250 Poseidon 252 rubrofemoratus 255

Paracycnotrachelus 253 pretiosa 257 rubropictus 251

parallelum 250 Pristiphora 258 rudis 252

Parastasia 255 Proctotrupes 260 rufescens 250

pardalis (Adolias) 262 Prodoretus 255 rufoscutellatus 250

pardalis (Apoderus) 253 Prosopocoilus 255 rugulosus 258

Paropia 253 Protemnemus 254 Rutilia 257

Paroplapoderus 253 Protoceraius 255

patibulum 251 Proxys 251 sabuleti 258

pauper 251 Pseudaradus 250 sabulosus 259

Paussus 256 Pseudopycanum 250 sagrae 259

pectoralis 258 Psiloptera 254 sanctifargavii 250

Pentatoma 251 Ptilocera 256 sanguinea (Amasis) 257

Penthima 254 pubescens 258 sanguinea (Microphysa)

perelegans (Epipedocera) puella 252 [249

[254 pulcherrima 254 sanguineguttata 251

perelegans (Lyramorpha) punctatissimus 250 sanguinolenta 254

[250 Purpuricenus 254 Saturnia 263

Perilissus 259 pusillus 250 Scambus 259

Perispuda 259 Pycanum 250 Schizorhina 254

persona 254 pygmaeus 256 schlegelii 252

Phaenocarpa 258 Pygoplatys 251 schwaneri (Callidea) 252

Phidias 259 pyrosticta 253 schwaneri (Hoplistodera)

Philonthus 256 pyrothorax 253 [250

phthisica 258 schwaneri (Onthophagus)

Phylax 258 quadratum 250 [256

Phyllodes 261 quadriguttata 254 Sciapteryx 258

Phylloserica 255 quadriguttatum 261 Scolobates 260

Phyllotoma 258 quadriguttatus 259 Scoparipes 249

Phylus 249 quadrillum 253 Scotinophara 251

Phymasterna 254 quadrimaculata 252 scurra 251

Physocephala 256 quincunx 251 Scutellera 252

Phytodictus 259 Selandria 258

Phytodietus 259 radians 251 sellula 250

Phytomyia 256 rayi 262 semilividus 259

pictus 260 rectangulus 255 semiluctuosus 259

Pieris 262 regalis 254 semivestitum 250

pileus 255 reinwardtii 263 semiviolaceus 250

pilifer 255 repentinus 259 senex 260

pilosa 252 Rhachicerus 256 Sepontia 250

Pimpla 259 Rhaphigaster 251 septemdecemguttatus 254

pinguis 258 Rhinoscapha 254, 255 Sericopimpla 259

Placosternum 251 rhododactylus 251 serratus 251

Platylabus 260 Rhyssa 260 Serrognathus 255

Platynopus 250 riparius 252 servillei 250

plebeja (Canthecona) 250 Rogas 258 sieboldii 254

plebeja (Pentatoma) 251 rosarum 253 Silbomyia 256

Pleophylla 255 roscidus 256 similis 257

Podops 251 rosea 250 simson 255

poecilea 262 roseiventer 261 singularis 259

Poecilocoris 252

rosenbergii (Pieris) 262

sixianus 260

268

'. sixii (Coleocentrus) 259

sixii (Selandria) 258

smaragdesthes 254

smaragdina 256

Solea 258

Soror 258

spectrum (Apoderus) 256

spectrum (Dryinus) 260

Spilomicrus 258

Spilosoma 261

spiniger 259

Spudaeus 251

stalii (Callidea) 252

stalii (Dalcantha) 250

Staphylinus 256

stellaris 262

stellio 255

Stenozygum 251

Stenus 256

Sternoplus 254

stictica 251

Strachia 251

Stromboceros 258

Stypotrupes 255

subpatellatus 249

subrugosus 251

sulphurea 262

sumatrana 252

sumatranus 252

sumatrensis 254

superba 262

surjas 262

Sussaba 259

suturalis 254

Syene 260

syngenesiae 258

Systropus 256

Taeniodera 254

Tarisa 251

tarsalis 251

tau 251

tenebrosa 252

Terias 263

tessellatus 256

Tesseratoma 251

testacea 255

Testrica 252

Tetrarthria 252

Tetrix 249

Theocolax 260

theodori 258

Theronia 260

Thestias 263

Thoria 251

tibialis 260

tigrina 262

tigrinus 252

timorensis 251

Tmesisternus 254

Tolumnea 251

tominia 263

tondana 262

tondanense 251

tournieri 259

Tragocerus 254

tragulus 255

transversalis 260

Trichius 254

Trichogomphus 255

Trichomastix 260

trigonalis 261

triguttata 250

trimaculata 250

trimaculatus 258

trisignata 250

trogon 262

Tuberolachnus 252

Tylonca 252

Typhlocyba 253

TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

undulata 257

unicallosus 259

unicolor 255

unifasciatus 259

vallator 258

Vanessa 262

variabilis (Callidea) 252

variabilis (Canthecona) 250

varians 251

varuna 262

venustus 249

verhuellii 261

verlorenii 255

vermiculatus 251

vidua (Clinteria) 254

vidua (Mormidea) 250

vilis (Eupholus) 255

vilis (Parapia) 253

villaenovae 258

vincta 260

Vipio 258

virginalis 261

virgula 249

Vitellus 250

vittata 255

Vulsirea 251

westwoodii (Agalope) 263

westwoodii (Dalcantha)

[250

westwoodii (Libyssa) 252

wieneckii 254

wttewaalli 258

Xanthopimpla 259, 260

Xylothrips 253

zebra 260

zoe 262

Bra TIJDSCHRIFT VOOR ENTOMOLOGIE, DEEL 124, AFL. 6, 1981

‘© ON.

Figs. 1—S. Adolias species described by Snellen van Vollenhoven, after his original water colours.

.The names of Snellen van Vollenhoven are followed by the names of the taxa (in brackets) Snellen van

Vollenhoven’s species are currently considered to belong to.

1. Adolias diardi, holotype (subspecies of Euthalia cocytina Horsfield). 2. Adolias ludekingii, holotype

(Euthalia cocytina cocytina Horsfield). 3. Adolias bipunctata, holotype (subspecies of Euthalia monina

Fabricius). 4. Adolias octogesima, $ (Euthalia alpheda alpheda Godart; the female of A. octogesima is

considered to be the same as Euthalia monina salia Moore). 5. Adolias varuna (subspecies of Tanaecia

lutala Moore).

TIJDSCHRIFT VOOR ENTOMOLOGIE

UITGEGEVEN DOOR

DE NEDERLANDSE ENTOMOLOGISCHE VERENIGING

* Een sterretje duidt aan een naam nieuw voor de wetenschap

* An asterisk denotes a name new to science

The articles by Bouéek, Watsham and Wiebes (on p. 232) and Krikken, Van Achterberg, Van Does-

burg, De Jong & Zwart (on p. 263) have their own index.

DIPTERA

aegyptium 112, 142

*albipalpus 111, 112, 144

alticola 140

angustiforceps 144

annulatipes 111, 116, 144,

[145

*ascium 111, 117, 144, 146

bipustulatum 133

brumale 144, 145

*chaudhurii 111, 119, 144,

[145

Chironomus 111

emarginatum 111

*fasciatipennis 144, 146

*flagellatum 111, 122, 136,

[144, 145

fuscoguttata 127

griseoguttatum 111, 120,

[125, 144, 145

*insolutum 111, 127, 144,

[145

Lauterborniella 127

“lineatum 111, 129, 145

longicrus 131

longinervis 127

*lucidum 111, 131, 145

milnei 145

nitidus 140

nubeculosus 111

nubifer 140

*nudiceps 111, 133, 145,

[146

“numerus 111, 112, 131,

[135, 145

Pentapedilum 111

Polypedilum 111 sqq

proxilitarsis 120

pruina 142

*pseudoflagellatum 111,

[137, 145, 146

purimanus 145

*rufomarginalis 111, 140,

[145

sealaenulus 145

tonnoiri 129

tridens 142

tripartitum 145

*tripunctum 111, 142, 145

tropicum 142

watsoni 136

HETEROPTERA

*assimilis 2, 16, 20

Asthenarius 1, 22

Hylopsallus 1 sqq

kiritshenkoi 23

perrisi 1, 2, 16, 17

Psallus 1 sqq

quercus 1, 2, 16, 22

variabilis 1, 2, 16, 19

*wagneri 2, 16, 18

LEPIDOPTERA

acanthella 39, 75, 108

*andrella 102

anguinella 98

argentipedella 95

argyropeza 28, 31, 43, 44

Artaversala 36

bosquella 72

canadensis 94

canutus 28, 39, 45

castaneae 31, 40, 80

chlorantis 41, 84

clemensella 38, 55

*coruscella 99

Dechtiria 30

ssp. downesi (E. argyro-

[peza) 31, 39, 45

Ectoedemia 27 sqq

flavipedella 31

Fomoria 35

Glaucolepis 36

grandisella 96

heinrichi 31, 40, 78

*helenella 105

liebwerdella 30

lindquisti 38, 61, 95

Manoneura 36

marmaropa 39, 49

maximella 53

mesoloba 41, 85

mediofasciella 85

nyssaefoliella 29, 37, 67

Obrussa 35

obrutella 41, 72

phleophaga 41, 81

*piperella 40, 77

platanella 38, 51

platea 98

populella 28, 29, 30, 38,

[41

quadrinotata 39, 70

*reneella 104

rubifoliella 31, 39, 63

similella 38, 56

stigmella 29, 31, 35

Trifurcula 30

trinotata 39, 46

ulmella 39, 65

virgulae 30, 38, 59

Zimmermannia 30

PLANTAE

Voor de namen in Bou-

tek, Watsham & Wiebes

zie p. 233

*Acer campestre 2, 22

Betula 38

Betula lutea 40

Betula papyrifera 40, 62

Carpinus 39, 72

Carya 39

Carya cordiformis 40, 48

Castanea 40, 41, 81