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HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

Volume 134, no. I, 1991

Tijdschrift

voor

Entomologie

A journal of systematic and evolutionary

entomology since 1858

Published by the Nederlandse Entomologische Vereniging si

Tijdschrift voor Entomologie

A journal of systematic and evolutionary entomology since 1858

Scope

The ‘Tijdschrift voor Entomologie’ (Netherlands Journal of Entomology) has a

long tradition in the publication of original papers on insect taxonomy and

systematics. The editors particularly invite papers on the insect fauna of the

Palaearctic and Indo-Australian regions, especially those including evolutionary

aspects e.g. phylogeny and biogeography, or ethology and ecology as far as

meaningful for insect taxonomy. Authors wishing to submit papers on disciplines

related to taxonomy, e.g. descriptive aspects of morphology, ethology, ecology

and applied entomology, are requested to contact the editorial board before

submitting. Usually, such papers will only be published when space allows.

Editors

E. J. van Nieukerken (elected 1986) and J. van Tol (1985)

Co-editors

A. W. M. Mol (1990) and R. T. A. Schouten (1990)

Advisory board

M. Brancucci (Basel), Y. Hirashima (Fukuoka) and M. R. Wilson (London).

The ‘Tijdschrift voor Entomologie’ is published in two issues annually by the

‘Nederlandse Entomologische Vereniging’ (Netherlands Entomological Society),

Amsterdam.

Editorial address

c/o National Museum of Natural History,

Postbus 95i7, 2300 RA Leiden, The Netherlands.

Correspondence regarding membership of the society, subscriptions and

possibilities for exchange of this journal should be addressed to:

Nederlandse Entomologische Vereniging

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Subscription price per volume Hfl. 300.- (postage included).

Special rate for members of the society. Please enquire.

Instructions to authors

May be obtained from the editors.

Graphic design

Ontwerpers B.V., Aad Derwort, ’s-Gravenhage

C. GIELIS

National Museum of Natural History, Leiden

MCZ

LIBRARY

JUL 5 1 1991

PTEROPHORUS SCHAFFER IN THE ETHIOPIAN

REGION (LEPIDOPTERA: PTEROPHORIDAE: HARVARD

PTEROPHORINAE): NEW SPECIES AND

CHECKLIST

UNIVERSITY

Gielis, C., 1991. Pterophorus Schiffer in the Ethiopian region (Lepidoptera: Pteropho-

ridae: Pterophorinae): new species and checklist. - Tijdschrift voor Entomologie 134: 1-

8, figs. 1-21. [ISSN 0040-7496]. Published 1 July 1991.

Pterophorus dallastai sp. n., P. uzungwe sp. n. and P. massai sp. n. are described. A

provisional checklist of the Ethiopian Pterophorus species is given, with notes on all

species. A lectotype is designated for P. cleronoma (Meyrick). P. centrocrates (Meyrick)

is synonimised with P. rhyparias (Meyrick). The status of P. endophaea (Meyrick) and P.

bacteriopa (Meyrick) is discussed.

Correspondence: C. Gielis, Mr. Haafkensstraat 36, 4128 CJ Lexmond, The Netherlands.

Keywords. - Lepidoptera; Pterophoridae; Prerophorus; new species; Africa.

When the opportunity arose a number of Ethi-

opian Pterophoridae from the collections of the

Zoological Museum of the University of Copenha-

gen (ZMUC), the Museum National d’Histoire na-

turelle, Paris (MNHN) and the Royal Museum of

Central Africa in Tervuuren (MRAC) were exam-

ined. The results are here combined with data ob-

tained from the collection of the British Museum

of Natural History, London (BMNH), the Instituut

voor Taxonomische Zoologie, Amsterdam (ZMAN)

and the authors’ collection (CG). Some of the spe-

cimens resembled, on superficial examination, pa-

laearctic species. A closer examination of these

species, however, showed a considerable variabil-

ity, especially in the genitalia. This was particularly

so in the genus Pterophorus, the subject of this

publication. The species have usually poorly

marked wings, which show a colour, ranging from

purely white to pale grey-brown. Especially in the

group of purely white specimens examination of

the genitalia is essential. It is to be expected that

more species will be recognized if more material

from this region becomes available.

The genus Pterophorus has its main distribution

centre in the Palaearctic region. A high number of

species (approximately 60) is known from West

and Central Europe, the Mediterranean area, Tur-

key and adjacent areas. From there the genus has

a continuous distribution into the Southeast Asiatic

region. Apart from Africa, further a few species are

recorded from the Pacific area and the new world.

Literature on the african representatives of the

genus Pterophorus is scattered over a few smaller

publications and mainly comprises single descrip-

tions of species, without illustrations of externals

or genitalia. Bigot (1969), in his review of the

collection of MRAC, describes and illustrates the

specimens in this collection, in this way fixing the

new species described by him accurately. Since

then, Bigot has given his attention mainly to the

fauna of Madagascar. Only Arenberger (1986) and

Gielis (1990) have published since on the Ethio-

pian fauna, but on other subfamilies: Agdistinae

and Platyptiliinae.

Provisional checklist of Ethiopian Pterophorus

Pterophorus Schäffer, 1766

Aciptilia Hiibner, [1825]

Merrifieldia Tutt, 1905

Porrittia Tutt, 1905

Alucita auct., nec Linnaeus, 1758

1. albidus (Zeller, 1852) comb. n.

2. rhyparias (Meyrick, 1907) comb. n.

centrocrates (Meyrick, 1933) syn. n.

. ischnodactyla (Treitschhe, 1833)

endophaea (Meyrick, 1930) comb. n.

bacteriopa (Meyrick, 1922) comb. n.

ceraunia (Bigot, 1969) comb. n.

lampra (Bigot, 1969) comb. n.

N DW mw

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

8. spissa (Bigot, 1969) comb. n.

9. lindneri (Amsel, 1963) comb. n.

10. candidalis (Walker, 1864) comb. n.

11. massat sp. n.

12. uzungwe sp. n.

13. dallastai sp. n.

14. cleronoma (Meyrick, 1920) comb. n.

TAXONOMIC TREATMENT

Pterophorus Schaffer, 1766

Type-species: Phalaena Alucita pentadactyla Linnaeus,

1758

Diagnosis. - Forewing cleft from % or less. Top

of lobes acute, without a terminal margin. R1 ab-

sent, R4 and RS fused, M3 and CU? forked or fused.

Hindwing without a scale-tooth.

Male genitalia asymmetrical, with asymmetrical

costal processes.

Female genitalia with a blister-like bursa copula-

trix in which no, one or two signa, an often pro-

nounced vesica seminalis, and often absent apo-

physes anteriores.

1. Pterophorus albidus (Zeller, 1852) comb. n.

(figs. 1, 10)

Aciptila albida Zeller, 1852: 397.

Diagnosis. - The species is characterized by the

faint yellow colour. Male genitalia with a hooked

costal process in the left valve and a large club-like

process in the right valve. The female genitalia

have the junction between the ductus bursae and

the bursa copulatrix in the shape of a bulging pro-

trusion with numerous sclerotized ridges.

Remarks. - This species has originally been des-

cribed from South Africa. This species can be easily

recognized by its yellowish-white to greyish-white

colour and the minute punctuation. After studying

the male (fig. 1) and female (fig. 10) genitalia, the

species turned out to have a wide distribution into

the Indo-Australian region. Specimens have been

Figs. 1-9. Pterophorus spp., male genitalia. — 1, P. albidus

(Zeller) Rep. of S. Africa, slide cG 1694 (ZMAN); 2, P.

rhyparias (Meyrick), British East Africa, slide CG 1870

(MNHN); 3, P. ischnodactyla (Treitschke, 1833). Rep. of S.

Africa, slide CG 1695 (ZMAN); 4, P. ceraunia (Bigot),

paratype, Zaire, slide Bigot (MRAC); 5, P. lampra (Bigot),

paratype, Zaire, slide Bigot (MRAC); 6, P. spissa (Bigot),

Central African Republic, slide cG 1878 (MNHN); 7, P.

candidalis (Walker), Ethiopia, slide CG 1879 (MNHN); 8,

P. dallastai sp. n., holotype; 9, P. wzungwe sp. n., holotype.

GIELIS: Ethiopian Pterophorus

examined from: Republic of South Africa, Zim-

babwe, Kenya, Zaire, Tanzania, Gambia, Cöte

d'Ivoire, Vietnam, Indonesia, New Guinea.

2. Pterophorus rbyparias (Meyrick, 1907) comb. n.

(figs. 2, 11)

Alucita rhyparias Meyrick, 1907: 489. Lectotype 9 (here

designated): Rep. of S. Africa, Transvaal, Pretoria

(Janse), genitalia slide BM 17993 (BMNH) [examined |.

Alucita centrocrates Meyrick, 1933: 425. Holotype &:

Belgian Congo (Zaire), N.E. Kivu, La Mutura, iii (Sey-

del), genitalia slide Bigot (MRAC) [examined]. Syn. n.

Diagnosis. - The species is characterized by the

pale ochreous-white colour with some small black

dots along the costa and dorsum of both forewing

lobes. Male genitalia (fig. 2) with a long and short

costal process in the left valve and a large club-like

process in the right valve. The female genitalia

(fig. 11) shows a distinct crescent-shaped distal

margin of the 7th tergite and a many folded ap-

pearance of the ductus bursae.

Variation. - The colour of the species ranges

from grey-white to pale ochreous-white. The

number of dots on the forewing may be reduced.

Distribution. - Rep. of S. Africa, Zaire, Kenya,

Tanzania.

Remarks. - Since the type-specimens of P. rhy-

partas and centrocrates are respectively a female

and a male, the synonymy, suggested by Aren-

berger (personal communication) could only be

proven by additional material. The evidence was

found in a small series, present in ZMUC, where

both sexes are present.

This species is difficult to separate from the next,

P. ischnodactyla Treitschke, and can only be distin-

guished by the distinct genitalia.

3. Pterophorus ischnodactyla (Treitschke, 1833)

(fig. 3)

Alucita ischnodactyla Treitschke, 1833: 223.

Diagnosis. - The species is characterized by the

pale ochreous-white colour and the small black dots

along the costa and dorsum of the forewing. Male

genitalia (fig. 3) with a widely forked costal process

in the left valve and a hooked single process in the

right valve. The female genitalia show an exca-

vated shape of the distal margin of the 7th tergite,

the ductus bursae is bulged out, with hardly any

transverse ridges.

Remarks. - The Zoological Museum, Amster-

dam (ZMAN) holds a specimen from the Republic

of South Africa, Bloemfontein, 30.1.1951 (Ee), ge-

nitalia CG 1695. This specimen represents the sole

verified African record of the species known to me.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 10-15. Prerophorus spp., female genitalia. - 10, P. albidus (Zeller), Tanzania, slide cc 4018 (zMuc); 11, P.

rhyparias (Meyrick), Zaire, slide Bigot (MRAC); 12, P. ceraunia (Bigot), Benin, slide cG 1881 (MNHN), 13, P. lampra

(Bigot), Côte d'Ivoire, slide CG 2178 (CG); 14, P. spissa (Bigot), paratype, Zaire, slide Bigot (MRAC); 15, P. candidalis

(Walker), Côte d'Ivoire, slide CG 2182 (CG).

This means an extension of the distribution from

the Mediterranean area towards the south of

Africa.

4. Pterophorus endophaea (Meyrick, 1930)

comb. n.

Alucita endophaea Meyrick, 1930:576.

Distribution. - Mozambique.

Remarks. - See under P. bacteriopa

5. Pterophorus bacteriopa (Meyrick, 1922)

comb. n.

Alucita bacteriopa Meyrick, 1922: 41.

Distribution. - Tanzania.

Remarks. - P. endophaea and P. bacteriopa have

not been examined, since I was unable to locate the

type specimens. The description of both species

indicates a relationship with the difficult to separ-

ate species P. rhyparias and P. ischnodactyla. A

synonymy based on the descriptions is suggestive,

but not proven yet.

6. Pterophorus ceraunia (Bigot, 1969) comb. n.

(figs. 4, 12)

Aciptilia ceraunia Bigot, 1969: 204.

Diagnosis. - The species belongs to the group of

white species, with distinct black fringe markings.

Differs in male genitalia (fig. 4) in having an asym-

metrical pair of costal processes in the left valve

and an equal long pair of processes in the right

valve. In female genitalia (fig. 12) in having long

ductal, longitudinally ridged, segments in both the

junctions to the bursa copulatrix as to the vesica

seminalis. Antrum longitudinal.

Distribution. — Zaire, Benin.

7. Pterophorus lampra (Bigot, 1969) comb. n.

(figs. 5, 13)

Aciptilia lampra Bigot, 1969: 204.

Diagnosis. - The forewings are purely white

with some black scales near the base and inner

margin of the first lobe. Differs in male genitalia

(fig. 5) in having a single costal process in the left

valve; slightly longer than the valve and a single

long costal process in the right valve which is

slightly shorter than the valve, in female genitalia

(fig. 13) in having an arched 7th tergite, with the

lateral positioned antrum, laterally progressing via

GIELIS: Ethiopian Pterophorus

the short ductus bursae into the blister-like bursa

copulatrix.

Distribution. - Zaire, Gabon, Côte d'Ivoire.

8. Pterophorus spissa (Bigot, 1969) comb. n.

(figs. 6, 14)

Aciptilia spissa Bigot, 1969: 202.

Diagnosis. - Forewings pure white, with small

black spots at the inner margin of both lobes. The

fringes show a distinct black and white pattern.

Differs in male genitalia (fig. 6) in having two

costal processes in the left valve, of which the

shorter is “th of the length of the longer. In the

right valve two costal processes of equal length and

a small transverse, short hook; in female genitalia

(fig. 14) in having the ductus bursae slightly longer

than the ductus to the vesica seminalis. Antrum

large, almost square.

Distribution. - Zaire, Central African Republic.

9. Pterophorus lindneri (Amsel, 1963) comb. n.

Aciptilia lindneri Amsel, 1963: 6.

Diagnosis. - The species is shining white, with-

out any dark scales. Bursa copulatrix without a

signum.

Male genitalia. - Unknown.

Female genitalia. - See diagnosis.

Distribution. - Ethiopia.

Remarks. - I have not examined the type-spec-

imen, since the description by Amsel (1963) is

adequate. It indicates the absence of a signum,

which character differentiates the species from all

its allies, which have one or two rosette-like signa

in their bursa copulatrix.

10. Pterophorus candidalis (Walker, 1864)

comb. n.

(figs. 7, 15-17)

Aciptilus candidalis Walker, 1864: 948.

Redescription. - Wingspan 15-28 mm. Head

white. Palpae short, white and greyish at top. An-

tennae white, shortly ciliated. Thorax, mesothorax,

tegulae and abdomen white. Legs white. Spurs of

unequal length. Forewings cleft from 74, colour

white with a faint yellowish grey gloss. Sparse dark

scales at the base of the cleft and near the apex of

the first lobe. Fringes white, at dorsum of both

lobes with two faint greyish interruptions. Hind-

wings white. Fringes white, at dorsum of second

lobe a greyish interruption.

Male genitalia (fig. 7). - Valvae asymmetrical. In

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 16-21. Pterophorus spp., female genitalia. - 16, P. candidalis (Walker), Chad, slide CG 2186 (cG); 17, Idem,

Tanzania, slide cc 4020 (zMuc); 18, P. dallastai sp. n., paratype, slide CG 1851 (MRAC); 19, P. massat sp. n., holotype,

20; P. uzungwe sp. n., paratype, slide cc 4040 (ZMUC); 21, P.

cleronoma Meyrick, parailecitotype, slide Bigot (MNHN).

right valve a pair of saccular spines; the ventral

spine twice as long as the centrally placed spine.

The left valve with two saccular spines; the ventral

spine abruptly ending in a slender, acute top. The

dorsal spine slender, length “4th of the length of

the ventral spine. Tegumen simple. Uncus slender,

of moderate length. Vinculum arched. Aedeagus

straight.

Female genitalia (figs. 15-17). — Antrum in

shape of a slender rectangular plate, with a rounded

top around a small hole. Ductus bursae and ductus

seminalis well-developed with longitudinal ridges.

Bursa copulatrix vesicular, with a single rosette-

like signum.

Variation. - The female genitalia show variation

in the length of the ductus seminalis. In specimens

from Céte d'Ivoire (fig. 15) it is longer than the

ductus bursae; from Chad and Ethiopia (fig. 16) of

equal length, and in Tanzania (fig. 17) shorter than

the ductus bursae.

Distribution. — Ethiopia, Kenya, Tanzania,

Zaire, Chad, Côte d'Ivoire, Sierra Leone.

Remarks. — The type specimen from Sierra

Leone (BMNH) is missing its abdomen. After com-

paring with the specimens from areas relatively

close to the type locality, I agree with Bigot (1969)

about the identity of the species.

11. Pterophorus dallastai sp. n.

(figs. 8, 18)

Type material. - Holotype 4: Zaire [Congo Belge], P.

N. A., Secteur Tschiaberimu, Mont Kamatondi pres Ki-

zungu, 2850 m., 22.iv.1955 (Vanschuytbroeck & Fontein,

no. 12785), genitalia slide cc 1850 (MRAC). — Paratypes:

1 9, Zaire [Congo Belge], P. N. A., Secteur Tschiaberimu,

Mont Musimba près Musabaki, 2450 m., 25.11.1954 (Van-

schuytbroeck & Synave, no. 7830), genitalia slide cc 1849

(MRAC); 2 9, Zaire [Congo Belge], P. N. A, Secteur

Tschiaberimu, Riv. Musavaki, affl. Talya Nord, 2720 m,

26-29.111.1954 (Vanschuytbroeck & Synave, no. 7828-29),

genitalia slides CG 1851, 1852 (MRAC, CG).

Diagnosis. - The species is closely related to P.

candidalis (Walker). It is characterized by the

slightly ochreous tinge of the forewing, the saccular

spines in the male genitalia and the shape of the

antrum in the female genitalia.

Description. - Male, female. Wingspan 24-28

mm. External characters as in P. candidalis

(Walker). The colour of the forewings is slightly

ochreous tinged.

Male genitalia (fig. 8). — Valvae asymmetrical.

Right valve narrow, lanceolate, with a long curved

and a short stout saccular spine. Left valve widen-

ing towards %th of valve length; with a long curved

and a short, wavy saccular spine. Tegumen bilobate.

Uncus slender, moderate. Vinculum narrow,

GIELIS: Ethiopian Pterophorus

arched. Aedeagus tube-like, towards top gradually

narrowing.

Female genitalia (fig. 18). — Antrum large,

rounded, without a central hole. Ductus bursae

longer than ductus seminalis, both ducti with lon-

gitudinal ridges. Bursa copulatrix vesicular, with a

single rosette-like signum.

Variation. - The specimens show some variation

in the intensity of the black scaling on the fore-

wings.

Biology. - The moths were collected in March

and April. Immature stages unknown.

Distribution. - Zaire.

Etymology. - A noun in genitive case, named in

honour of Dr. U. Dall’asta, curator of Lepidoptera

in MRAC.

12. Pterophorus massai sp. n.

(fig. 19)

Type material. - Holotype 9: Kenya (Afr. Or. Angl.),

Nairobi, Plaine Massai, viii.1904 (Alluaud), genitalia

slide cG 1875 (MNHN).

Diagnosis. - The species is closely related to P.

candidalis (Walker) and P. dallastai Gielis, but char-

acterized by the female genitalia: the ratio of the

length of the ductus bursae and the ductus semina-

lis is three to one.

Description. - Female. Wingspan 25 mm. Exter-

nal characters as in P. candidalis (Walker).

Male genitalia. - Unknown.

Female genitalia (fig. 19). - Antrum large, end-

ing roof-like, with an oval hole. Ductus bursae

three times longer than ductus seminalis, both with

longitudinal ridges. At the junction of the ductus

bursae and ductus seminalis two rows of small

spines. Bursa copulatrix vesicular, with a rosette-

like signum.

Biology. - The moth was collected in August.

Immature stages unknown.

Distribution. - Kenya.

Etymology. - Massai, a noun in apposition,

named after the type locality.

13. Pterophorus uzungwe sp. n.

(figs. 9, 20)

Type material. - Holotype &: Tanzania, Uzungwe

Mts., Mwanihane Forest above Sanje, 1000 m, 1.viii.1981

(Stoltze & Schaff), genitalia slide cc 4039 (ZMUC). Par-

atype 9: same locality and data, genitalia slide cG 4040

(GS,

Diagnosis. - The species is characterized by the

shape of the saccular arms in the male and the

double signum in the female. The size is relatively

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

small for species of its group.

Description. - Male, female. Wingspan 18-19

mm. External characters as in P. candidalis

(Walker).

Male genitalia (fig. 9). - Valvae asymmetrical.

Right valve with a long, almost straight, saccular

spine. Left valve with a curved saccular spine, with

two small lateral thorns. Tegumen bilobate. Uncus

slender, moderate. Vinculum rather wide. Aedea-

gus tube-like.

Female genitalia (fig. 20). - Antrum three times

longer than wide, club-like, with a large central

distal hole. Ductus bursae and ductus seminalis of

equal length, both with longitudinal ridges. Bursa

copulatrix with two rosette-like signa.

Biology. - The moths were collected in August.

Immature stages unknown.

Distribution. - Tanzania.

Etymology. - Uzangwe, a noun in apposition,

named after the type locality.

14. Pterophorus cleronoma (Meyrick, 1920)

(fig. 21)

Alucita cleronoma Meyrick, 1920: 41. Lectotype 9 (here

designated): Brit. E. Africa, Kenya, Mt. Kenya, i-

ii.1912 (Alluaud & Jeannel), genitalia slide BM 17987

(BMNH) [examined].

Other material examined. — Paralectotype Q: locality

and data as lectotype, genitalia slide Bigot (MNHN)

[examined].

Diagnosis. - The species is characterized by the

shape of the female genitalia. Especially the grad-

ually narrowing antrum, in combination with the

absence of a signum is diagnostic.

Female genitalia (fig. 21). - Antrum gradually

narrowing. Ductus bursae slender. Bursa copulatrix

vesicular. No signum. Apophyses anteriores ab-

sent. Apophyses posteriores three times longer

than papillae anales.

Distribution. - Kenya.

Remarks. - In his description Meyrick did not

designate a holotype. Both BMNH and MNHN ap-

peared to hold a specimen labelled ‘type’, the BMNH

specimen is herewith designated as lectotype.

ACKNOWLEDGMENTS

I wish to thank Mr. B. Brugge (ZMAN), Dr. U.

Dall’asta (MRAC), Mr. O. Karsholt (ZMUC), Dr. G.

C. Luquet (MNHN) and Mr. M. Shaffer (BMNH) for

the opportunity to study the Pterophoridae in their

museum, Mr. H. W. van der Wolf for correcting the

English text and Mrs. van de Sant for typing the

manuscript.

REFERENCES

Amsel, H. G., 1963. Kleinschmetterlinge aus Aethiopien.

- Stuttgarter Beiträge zur Naturkunde 121: 1-12.

Arenberger, E., 1986. Die Agdistis-Arten der Aethiopi-

schen Region, 1. Beitrag. - Mitteilungen der Schwe-

izerischen entomologischen Gesellschaft 59: 187-196.

Bigot, L., 1969. Les Lepidopteres Pterophoridae du musee

royal de l’Afrique Centrale, à Tervuren. - Revue de

Zoologie et de Botanique Africaines 79: 165-206.

Gielis, C., 1990. Neue Pterophoridae der Aethiopischen

Fauna. - Entomologische Zeitschrift, Frankfurt 100:

113-125.

Meyrick, E., 1907. Notes and descriptions on Pteropho-

ridae and Orneonidae. - Transactions of the Entomo-

logical Society of London 1907: 471-511.

Meyrick, E., 1920. Microlepidoptera. - In: Voyage de Ch.

Alluaud et R. Jeannel en Afrique Orientale (1911-

1912). Résultats scientifiques. Lepidoptera, II: 33-120.

Paris.

Meyrick, E., 1922. New microlepidoptera of the German

entomological institute. - Entomologische Mitteilun-

gen 11: 44.

Meyrick, E., 1930 (reprint 1969). Exotic microlepidop-

tera 3 (18): 545-576. - Hampton, England.

Meyrick, E., 1933 (reprint 1969). Exotic microlepidop-

tera 4 (14): 417-448. - Hampton, England.

Treitschke, F., 1833. Die Schmetterlinge von Europa 9: 1-

294. - Leipzig.

Walker, F., 1864. List of the specimens of lepidopterous

insects in the British Museum 30. Lepidoptera Heter-

ocera: 926-953. - London.

Zeller, P. C., 1852. Revision der Pterophoriden. - Linnaea

Entomologia 1: 319-413.

Received: 5 June 1990

Revised version accepted: 2 January 1991

R. V. HENSEN

Utrecht, The Netherlands

REVIEW OF MALESIAN SPHECINA

(HYMENOPTERA, SPHECIDAE, SPHECINAE)

Hensen, R. V., 1991. Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae).

- Tijdschrift voor Entomologie 134: 9-30, figs. 1-65. [ISSN 0040-7496]. Published 1 July

1991.

The species of Sphex and Isodontia (subtribe Sphecina) in the Malesian region are

reviewed. Twenty species are recognized in Isodontia, which are segregated into five

species-groups, while 21 species are recognized in Sphex, divided among three species-

groups. Keys are provided to the genera, species-groups and species. Synonymy and

distribution of the species is discussed. Twelve new species are described, viz. I. pilipes

(Burma to Sumatra, Borneo), I. cestra, I. elsei (both Sulawesi), I. nidulans (Philippines),

I immaculata, 1. capillata, I. delicata, I. papua, S. wilsoni, S. rex (all New Guinea), S.

solomon (Solomon Islands), and S. walshae (NE India to Sumatra, Borneo). The following

new synonymies are established: Sphex morosus Smith, 1861, S. volatilis Smith, 1861, S.

triodon Kohl, 1890, S. abditus Kohl, 1895 and S. abditus nugenti Turner, 1910 are

synonymized with Isodontia aurifrons (Smith, 1859); I. diodon alemon Van der Vecht,

1957 with I. severini (Kohl, 1898); S. wstulatus Kohl, 1890, S. simplex Kohl, 1898 and

S. insularis Cameron, 1901 with I. praslinia (Guérin-Méneville, 1831); S. albohirtus

(Turner, 1908) with I. sepicola (Smith, 1859) and S. fulvohirtus Bingham, 1890 with S.

diabolicus Smith, 1858.

Raymond V. Hensen, W.A. Vultostraat 147, 3523 TW Utrecht, The Netherlands.

Key words. - Malesia, Oriental region, Australian region, Sphecidae, Sphex, Isodontia,

taxonomy.

In this paper the species of the subtribe Sphecina

comprising the genera Sphex Linnaeus and Isodon-

tia Patton are briefly treated as far as they occur in

the Malesian area. The Malesian area is used here

in a broad sense, i. e. comprising Southern Thai-

land, Malaysia, Indonesia, the Philippines, New

Guinea and the Solomon Islands.

Sphex and Isodontia are large, active insects, and

some of the species treated here are among the

most commonly encountered digger wasps in the

area. The last comprehensive publication dealing

with the Malesian species was that by Kohl (1890),

but the taxonomy of the group was updated in

many smaller papers (eg. Kohl 1895, 1898, Van der

Vecht 1957, 1973). Therefore, it is surprising that

so many new species remained undetected. Furth-

ermore, a considerable number of new synonyms

were discovered in the course of this study.

Comprehensive diagnoses of the morphology

and biology of the two genera were given by Bohart

& Menke (1976), and need not be repeated here.

One of the aims of this study is to define and

diagnose a number of species-groups. Most of these

groups are considered monophyletic, being based

on inferred synapomorphies. Others, in particular

the Sphex subtruncatus-group are possibly para-

phyletic. It seems premature to pay more attention

to the phylogeny of the subtribe here, as a thorough

treatment of this subject will require study of all

species in these genera.

This paper deals strictly with species: it does not

present a classification below the species-level. Al-

though past workers (eg. Van der Vecht & Krom-

bein 1955) have recognized subspecies in the Male-

sian Isodontia, especially for colour differences from

island to island, I have not recognized them in this

paper. Subspecific names recognized in the past are

simply treated here as synonyms of the nominate

form. On the other hand, several taxa regarded as

subspecies in the past are elevated to species rank.

This study was initiated by dr. J. van der Vecht,

and in many instances I have relied on his notes

concerning literature, type material and specimens.

As far as type specimens are concerned, I have

indicated examination by Van der Vecht with a

single exclamation mark, by myself with a double

exclamation mark after the type depository (e.g.

BMNH!!). A question mark behind an abbreviation

of a museum means that a type was looked for but

not found in that particular museum.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

A few thousand specimens were examined in the

course of this study. I have refrained from listing

these explicitly, except for the type material of new

species. The outlines of distribution, as given for

each species are entirely based on examination of

specimens by me or by Van der Vecht, except as

noted.

Abbreviations. - Names of museums in which

types are preserved are abbreviated as follows.

BMNH Natural History Museum, London, Eng-

land.

BRI Biosystematics Research Institute, Ot-

tawa, Canada.

BPBM Bernice P. Bishop Museum, Honolulu,

Hawaii, USA.

CAS California Academy of Sciences, San

Francisco, USA.

DEI Deutsche Entomologische Institut,

Eberswalde, Germany.

IRSN Institut Royal de Sciences Naturelles,

Brussels, Belgium.

MCG Museo Civico di Storia Naturale, Genova,

Italy.

MCZ Museum of Comparative Zoology, Cam-

bridge, Massachusetts, USA.

MNHN Muséum National d’Histoire Naturelle,

Paris, France.

NMB Naturhistorisches Museum, Basel, Swit-

zerland.

NMW Naturhistorisches Museum, Vienna,

Austria.

OUM Oxford University Museum, Oxford,

England.

RMNH Nationaal Natuurhistorisch Museum,

Leiden, The Netherlands.

TMB Termeszettudomanyi Muzeum, Budap-

est, Hungary.

USNM National Museum of Natural History,

Smithsonian Institution, Washington, D.

CAUSA

UZMC Universitetets Zoologisk Museum, Co-

penhagen, Denmark.

ZMB Zoologisches Museum der Humboldt-

universität, Berlin, Germany.

The following morphological terms are abbre-

viated.

IOC Shortest interocular distance across the

clypeus.

IOV Shortest interocular distance across the

vertex.

ACKNOWLEDGEMENTS

Dr. J. van der Vecht donated his personal notes

and provided many of the specimens that formed

the foundation of this study. The following persons

aided in various ways, including arranging loans of

material, assistance and hospitality at my visits to

various museums, etc. In alphabetical order: Dr.

ing. C. van Achterberg, Leyden; Dr. J. M. Carpen-

ter, Cambridge, Mass.; Dr. J. Casevitz-Weulersse,

Paris; Dr. P. Dessart, Brussels; Dr. M. Fischer,

Vienna; Dr. F. Koch, Berlin; Dr. ©. Lomholdt,

Copenhagen; Dr. A. S. Menke, Washington DC;

Mr. G. M. Nishida, Honolulu; Dr. J. Oehlcke,

Eberswalde; Dr. C. O’Toole, Oxford; Dr. J. Papp,

Budapest; Dr. W. J. Pulawski, San Francisco; Dr. M.

Sharkey, Ottawa; Mr. C. R. Vardy, London.

I wish to express my sincere gratitude to these

persons, and others who took a constructive inter-

est in my activities. Special thanks are due to Dr.

A. S. menke for reviewing the manuscript.

A visit to the Natural History Museum, London,

and a collecting trip to Indonesia were supported

by the Uyttenboogaart-Eliasen Stichting.

SYSTEMATIC PART

Key to genera and species-groups

1. Sides of propodeum with complete spiracular

groove; basal vein of second submarginal cell

as long as or longer than anterior vein; female

with well developed tarsal rake, composed of

long, flattened spines (fig. 62) (Sphex) ..... 2

- Sides of propodeum without spiracular groove;

basal vein of second submarginal cell usually

shorter than anterior vein; female without tar-

salfrakenlsodontsa) tae CRE EN 4

2. Metanotum medially with pair of tubercles or

with single tubercle, that is more or less im-

Pressed/imediallyar ae RR SERBIEN ORA

eh eo a group of S. argentatus (p. 20)

- Metanotum without tubercles, flat or slightly

CONVEXIMECIA VARE ee eae tea eee ee ee 3

3. Scutellum of female flat, shiny (sometimes

densely covered with tomentum); male flagel-

lum without placoids; claw teeth small, per-

pendicularstorclawA(ti 0859) EUR ER

ee group of S. resplendens (p. 22)

— Scutellum of female convex, often with median

furrow, usually dull; at least fifth flagellomere

of male with placoid (figs. 43-45); claw teeth

large, at sharp angle to claw (fig. 60)...........

VELEN SEE group of S. subtruncatus (p. 26)

AE AMES ATI 9197 IA NIIR WI DEE AO NOA 5

SHRemales Ent MNT LAME DIE ONE dana 9

5. Mandibles bidentate ............................ 6

— Mandibles tridentate ........................... 7

6. Mandibles very slender, nearly straight (fig. 8);

inner orbits strongly converging below; petiole

10.

11.

19%

longer than hind basitarsus; small, slender spe-

cies, body length 13-19 mm

ES eg A group of I. diodon (p. 14)

Mandibles stout and arcuate (fig. 6); inner or-

bits parallel or slightly diverging below; peti-

ole shorter than hind basitarsus; large, stout

species, body length 20-28 mm

group of I. chrysorrhoea (p. 13)

. Fourth flagellomere with well defined placoid

(figs. 35-38) group of I. praslinia (p. o

Flagellum without placoids

. Basal inner tooth of mandible strongly jee

loped, much stronger than second inner tooth

(fig. 10); basal half of clypeus with blunt lon-

Ritudinalicasina e een:

group of I. ochroptera (p. 19)

Basal inner tooth of mandibles smaller (fig. 2);

basal half of clypeus at most with trace of

longitudinal carina

group of I. aurifrons (p. 11)

. Widest part of frons slightly narrower than

clypeus in the middle, inner orbits slightly di-

verging below; outer side of hind tibia with

more than 20 short spines; body length 29-35

mm group of I. chrysorrhoea (p. 13)

Inner orbits parallel or converging below;

outer side of hind tibia with at most 10-12

short spines; body length 14-26 mm

Mandibles very slender, straight, apically bifid

(fig. 7); hind tibiae with 1-3 short spines on

outer side; fourth segment of hind tarsi wider

than long eee to group of I. diodon (p. 14)

Mandibles robust, curved, apically tridentate

(fig. 9); hind tibiae with 5-10 spines on outer

side; fourth segment of hind tarsi elongate

ao agri ident Retest rae Beiden! 11

Labrum with single median longitudinal car-

ina, that does not project apically; apical mar-

gin of clypeus with pair of well differentiated

teeth (fig. 9); pilosity of head and mesosoma

black group of I. ochroptera (p. 19)

Labrum apically with two parallel, raised, pro-

jecting carinae; apical clypeal margin medially

notched (fig. 11), or with pair of short teeth

(fig. 1); pilosity variable

First flagellomere distinctly longer than se-

cond; antennae not or hardly longitudinally

ridged (fig. 21); mandible stout (fig. 11)

group of I. praslinia (p. 19)

First flagellomere hardly longer than second

(fig. 19), or (I. pilipes) mandibles slender (fig.

3); antennae longitudinally ridged (fig. 19)

group of I. aurifrons (p. 11)

HENSEN: Malesian Sphecina

Genus Isodontia Paton

Group of Isodontia aurifrons

Diagnosis

Body length 4 25-23 mm, 9 17-26 mm. Labrum

of female with a pair of short parallel projecting

carinae apically; mandible tridentate in both sexes,

sometimes with weak third inner tooth near apex;

clypeus of female with median notch; first flagel-

lomere as long as second; flagellum of female us-

ually with longitudinal sulci on second to tenth

segment; third to seventh flagellomere in male

with convex, papillate sensory areas; outer side of

hind tibia with 5-9 short spines; petiole usually

shorter than hind basitarsus.

Species included

Species treated. - I. aurifrons (Sm.), formosicola

(Strand), pilipes sp. n.

Other included species. - I. mazdli (Yasumatsu)

from Japan.

Key to species

ile

Females dente ee Ce 2.

MAESTRI I TIE TR 4

First flagellomere much longer than second

(fig. 20); Burma, Sumatra, Borneo

pilipes sp. n.

First flagellomere not or scarcely longer than

second (fig. 19)

. Metasoma largely or entirely dull red; clypeus

with median carina over most of its length;

Taiwan, China, S. Thailand, Malaysia, Sumatra,

Borneo formosicola (Strand)

Metasoma entirely black; clypeus with short

carina near base or medially; Sikkim to China

and Malaysia, Malesian region, Queensland

aurifrons (Smith)

. Metasoma largely dull red; third flagellomere

as long as second (fig. 23); clypeus apically with

pair of short teeth; wings yellowish

formosicola (Strand)

Metasoma entirely black; third flagellomere

longer than second; clypeus apically without

teeth (fig. 2); wings brown or clear

. Petiole as long as hind basitarsus; first flagel-

lomere longer than second (fig. 24)

pilipes sp. n.

Petiole shorter than hind basitarsus; first fla-

gellomere as long as second (fig. 22)

aurifrons (Smith)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 1-12. Isodontia, clypeus and mandibles. - 1, aurifrons, 9; 2, aurifrons, 6; 3, pilipes, 9; 4, pilipes, &; 5,

chrysorrhoea, 9; 6, chrysorrhoea, 6; 7, cestra, Q; 8, delicata, 8; 9, ochroptera, 9; 10, ochroptera, Ô; 11, praslinia, 9;

12, praslinia, €.

Isodontia aurifrons (Smith)

(figs. 1-2, 19, 22)

Sphex aurifrons Smith, 1859: 157. Lectotype 9 (present

designation): Indonesia, Aru (OUM!!)

Sphex morosus Smith, 1860: 122. Holotype @: Indonesia,

Bacan (OUM!!) Syn. n.

Sphex volatilis Smith, 1860: 122. Holotype 9: Indonesia,

Bacan (OUM!!) Syn. n.

Sphex triodon Kohl, 1890: 377. Lectotype @ (present

designation): Indonesia, Java (NMW!!) Syn. n.

Sphex abditus Kohl, 1895: 51. Holotype 9: Sikkim

(NMW!!) Syn. n.

Sphex abditus nugenti Turner, 1910a: 343, 345. Holotype

Q: Australia, Cairns (BMNH!!) Syn. n.

Distribution. - Continental Asia from Southern

India and Sri Lanka to China and Malaysia, Indone-

sia, Philippines, New Guinea, Queensland.

Isodontia formosicola (Strand)

(fig. 23)

Sphex (Isodontia) formosicola Strand, 1913: 82. Lecto-

type Q (present designation): Taiwan: Hoozan (DEI!!)

Distribution. - Continental Asia from China to

Malaysia, Taiwan, Sumatra, Borneo.

Isodontia pilipes sp. n.

(figs. 3-4, 20, 24, 46)

Type material. - Holotype 8, Indonesia, 'S. Sumatra, +

600 m, Res. Benkoelen, Tandjong Sakti, 16-19 VII, M. E.

Walsh leg. 1935’ (RMNH). - Paratypes. Burma: Tenasse-

rim, Ihaungyin Valley, v.1893, ‘Sphex aurifrons Sm., @ (1

2), Tenasserim, Ataran Valley, xii.1891 (1 9), Burma,

Middle Tenasserim, Haundraw Valley, viii.1894 (1 9), all

C. T. Bingham (BMNH). Indonesia: S. Sumatra, Benkulen,

50 m, 19-21.v.1935, M. E. Walsh (1 6, RMNH); W. Suma-

tra, Padangpanjang, 800 m, 1.v.1988, R. Hensen (1 8,

RMNH). Malaysia: Sabah, Ranau 8 mi. N., Paring Hot

Springs, 500 m, 8-11.x.1958, L. W. Quate (1 8, BMNH);

Borneo, , v.1900, P. Cameron coll., ‘Sphex nigripes, Sm.’

(1 9, BMNH); Borneo, Kuching, v.1900 (1 4, BMNH).

Diagnosis

Within the aurifrons-group, pilipes is character-

ized by the first flagellomere being distinctly longer

than the second. Additionally, in the male, the

petiole is as long as the hind basitarsus (shorter in

the other species). I. pzlipes may actually be closest

to the Japanese I. maidli (Yasumatsu). It differs

from that species in the male by the presence of

hair-fringes on the sternites, and in the relative

lengths of the proximal flagellomere (1 : 1 : 1.4 in

matdli). In the female I. pilipes is characterized by

the dark hind tibiae (reddish in maid/1) and the

presence of transverse rugae medially on the prop-

odeum.

HENSEN: Malesian Sphecina

Description

Female. - Body length 17.5-21.0 mm; length of

forewing 12.5-15.0 mm. Black, mandible medially

reddish, tegulae with margin brownish yellow or

reddish, distinctly lighter than the center. Wings

yellowish, apical margins and particularly apex of

forewing infuscated. Erect pilosity of head and

mesosoma brown but yellowish on genae, venter of

mesosoma and propodeum. Face with rather

sparse silvery tomentum, mesosoma without con-

spicuous tomentum. Clypeus 0.9 times as long as

IOC, with median carina on dorsal half; 10V 1.0

times IOC; relative lengths of flagellomeres I-III 1

: 0.8 : 0.85; mandibles slender. Mesoscutum shiny,

sparsely finely punctate, punctures on disc 1-3 di-

ameters apart; propodeal dorsum shiny, rugose-

punctate, with short transverse rugae medially. Pe-

tiole 0.8 times as long as hind basitarsus.

Male. - Body length 15.0-17.0 mm; length of

forewing 11.0-12.5 mm. Like the female, except for

the following. Black; mesoscutum with faint

greyish shine. Wings light brown, with violaceous

shine. Erect pilosity of head brownish to black, of

mesosoma yellowish grey, but darker on mesoscu-

tum. Clypeus 0.85 times as long as IOC; IOV 1.0-1.1

times IOC (1.0 in type); relative lengths of flagel-

lomeres I-III 1 : 1.0 : 1.3; flagellomeres III-VIII

with convex, papillate sensory areas; mandible

with faint third inner tooth near apex; propodeal

dorsum finely punctate-reticulate. Petiole 1.0-1.05

times as long as hind basitarsus; fourth to seventh

sternite with fringes of appressed brown setosity;

gonostyle simple.

Etymology

Pilipes is a noun in apposition, from Latin pilus

= hair and pes = leg.

Group of Isodontia chrysorrhoea (Kohl)

Diagnosis

Body length 9 29-34 mm, @ 20-28 mm; imme-

diately recognized by their large size, and the pres-

ence of yellow bands on the metasoma. Female:

labrum not modified; mandible tridentate; clypeus

rounded; first flagellomere distinctly longer than

second. Male: mandible bidentate; clypeus shal-

lowly emarginate; fourth to sixth flagellomere

with broad, convex papillate sensory area.

Species included

Species treated. - I. chrysorrhoea (Kohl).

Other included species. - I. auripygata Strand

from Taiwan, China and Vietnam.

Isodontia chrysorrhoea (Kohl)

(figs. 5-6, 25)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Sphex apicalis Smith, 1856; 253. Holotype 9: Indonesia,

Sumatra (BMNH!!) [non Smith, 1856: 262].

Sphex (Isodontia) chrysorrhoeus Kohl, 1890: 371 [new

name for S. apicalis Smith, 1856: 253].

Sphex (Isodontia) hewitti Cameron, 1906: 119. Holotype

8: Malaysia, Sarawak, Kuching (BMNH!!).

Distribution. - S. Thailand, Indonesia (Sumatra,

Java, Borneo, Nias), Malaysia (Sarawak); Philip-

pines (Mindanao).

Group of Isodontia diodon (Kohl)

Diagnosis

Body length & 12-18 mm, 9 14-19 mm. Labrum

of female simple; mandible abnormally slender,

nearly straight, bidentate in male, the apex bifid in

female and with small outer tooth; clypeus of fe-

male simple; relative lengths of flagellomeres var-

iable; flagellum of female without sensory areas;

flagellum of male with flat placoids on one or more

flagellomeres or with broad, convex, papillate sen-

sory areas. Scutellum and metanotum convex;

propodeal dorsum reticulate to transversely stri-

gose; outer side of hind tibia with 1-3 spines. Pe-

tiole usually longer than hind basitarsus.

Species included

Species treated. - I. capillata sp. n., cestra sp. n.,

delicata sp. n., diodon (Kohl), egens (Kohl), franz:

(Kohl), maculata sp. n., papua sp. n., permutans

(Turner), severini (Kohl).

Other included species. - I. nıgelloides (Strand)

from Sri Lanka and Southern India and an un-

named Australian species.

Key to species

1. Females (unknown: capillata, delicata, papua)

=) Males (unknown: man) ee 8

2. Forewing with sharply differentiated dark

markrapicallly SS. ri BERE D denti 3)

- Forewing darkened apically, but dark area not

Shacplyidelimitedkt eas. ned ea E 6

SE acialktomentumksolden: obi 4

=MFacialitomentumisilveryietfa st. dere 2: 5

4. Petiole as long as hind basitarsus; first meta-

somal tergite black, laterally reddish translu-

cent SUIAWESITe ra en. ker cestra sp. n.

- Petiole longer than hind basitarsus; first met-

asomal tergite red; Sumatra, Java, Borneo, Phi-

lippines, Nusa Tenggara, Moluccas ............

severini (Kohl)

5. Metasoma, except petiole, entirely red; Borneo

SERA CO Led RE franzi (Cameron)

— Metasoma black, with at most first tergite and

posterior margins of other tergites reddish;

10.

122

15:

14.

157

diodon (Kohl)

. Facial tomentum golden (except specimens

from Gizo Isl.); New Guinea, Solomon Islands

DIS DT NE permutans (Turner)

Facialsıtomentum silyveryas ee dl

. Mesosoma with conspicuous silvery tomentum

on pronotal lobes, metanotum and sides of

propodeum; New Guinea, Bismarck Islands,

BAR ER ee n egens (Kohl)

Mesosoma without silvery tomentum: Bis-

marckilslands Her immaculata sp. n.

. Flagellomeres I-II short, III at least twice as

longsasıl(es- fis220) a ee ee 9

Flagellomeres I-II longer, III at most 1.5 times

asilong:asil6 (ep e 2 14

. Flagellomeres III-VI with broad, flat placoids

over their entire length (figs. 26, 30,31) 10

Flagellomere III at most with small apical pla-

CONIC GS este PS2) TI 13

Posterior tergites with conspicuous, dense yel-

low pilosity; gonostyles apicodorsally with

sharply pointed process (fig. 47); New Guinea

capillata sp. n.

Facial tomentum silvery; posterior tergites

with sparse white or yellowish pilosity; gonos-

tyles apicodorsally rounded (figs. 52, 54) or

Withibroadiangle (fig OE 11

Mesosoma without tomentum; gonostyles

apicodorsally swollen and rounded (fig. 52);

Bismarck Islands... immaculata sp. n.

Mesosoma with conspicuous silvery or golden

tomentum on pronotal lobes, metanotum,

sides of propodeum; gonostyles either angu-

larly raised apicodorsally (fig. 51), or simply

rounded" (fig. 4)" |e RR 12

Gonostyles simply rounded apicodorsally (fig.

54); tomentum of mesosoma golden or (So-

lomon islands) pilosity brown; New Guinea,

Solomon Islands ......... permutans (Turner)

Gonostyles apicodorsally raised into a broad

angle (fig. 51); tomentum of mesosoma

silvery, pilosity pale; New Guinea, Bismarck

IslandsYBiaksy tern egens (Kohl)

Flagellomere III with small apical placoid, IV-

VI with long, narrow, concave placoids (fig.

28); sternite V-VII with fringes of yellow set-

osity; New.Guineab.r.: ein. ae 2 delicata sp. n.

Flagellomere IV with long, narrow, concave

placoid, V with small apical placoid (fig. 32);

sternites without fringes, only with sparse yel-

low pilosity; New Guinea ........ papua sp. n.

Only flagellomeres IV-V with placoids (fig.

2) SEN VSN, PAPER ss cestra sp. n.

Flagellomeres (II-) III-V with broad, convex

Placoidi(tios 129433) Riss See 14

Flagellomere III 1.4 times as long as I (fig. 33);

lower apical angle of gonostyle acute (fig. 50);

facial tomentum usually silvery; Nepal to

China and Singapore .......... diodon (Kohl)

- Flagellomere III 1.1-1.2 times as long as I (fig.

29); lower apical angle of gonostyle a right

angle (fig. 55); facial tomentum golden or pale

golden; Sumatra, Borneo, Java, Nusa Tenggara,

Moluccas, Philippines ........ severini (Kohl)

Isodontia capillata sp. n.

(figs. 26, 47)

Type material. - Holotype @, Indonesia, Irian Jaya,

Jayapura, xii.1936, W. Stüber (RMNH). — Paratypes: In-

donesia: Irian Jaya, Jayapura, vii.1938, L. J. Toxopeus (1

6, RMNH). Papua New Guinea: Madang Province, Duru,

15 km SW. Madang, ca. 5° 20 S, 145° 43 E, 14.111.1987, W.

J. Pulawski (1 8, CAS).

Diagnosis

Differing from all other species of this group by

the dense yellow pilosity covering the posterior

tergites, and by the shape of the gonostyles (fig.

47).

Description

Male. - Body length 15.0-16.5 mm, length of

forewing 11.0-12.0 mm. Black; mesoscutum with

fine oily shine; posterior margins of tergites

broadly reddish to yellowish translucent. Wings

light brownish hyaline, apically infuscated. Erect

pilosity of head and mesosoma yellow; facial to-

mentum golden; mesosoma with conspicuous

golden tomentum on pronotum, sides of mesoscu-

tum, metanotum, mesopleuron and sides of prop-

odeum; metasoma with very conspicuous pale yel-

low pilosity, dense and erect on posterior tergites.

Clypeus 1.0 times as long as 10C, with vague median

carina; IOV 1.1 times IOC; relative lengths of flagel-

lomeres I-III 1: 1.4 : 2.6; third to sixth flagellomere

with broad flat placoid. Mesoscutum densely rather

finely punctate, punctures on disc 0.5-1 diameter

apart; propodeal dorsum shiny, anteriorly trans-

versely strigose, posteriorly very densely punctate.

Petiole 1.2 times as long as hind basitarsus; ster-

nites without fringes, only seventh sternite with

some longer pilosity; eighth sternite apically trun-

cate; gonostyle apically with strong tooth pointing

upward.

Etymology

Capillata is an adjective, from Latin capillatus =

long-haired.

HENSEN: Malesian Sphecina

Isodontia cestra sp. n.

(figs. 7, 27, 48)

Type material. - Holotype Q, ‘Indonesia, N. Sulawesi,

100 m, Tangkoko-Dua Saudara N. R., 1°30’ N, 125° 10’

E, 28.XI.1985, C. v. Achterberg, RMNH’86’ (RMNH). —

Paratypes: Indonesia, Sulawesi: Dumoga-Bone N. P., ca.

220 m, nr. Base Camp Toraut R., 0°34’ N, 123°54 E,

10.xi.1985, C. v. Achterberg (1 9, RMNH); Kendari,

iv.1874, O. Beccari (1 9, McG); North Sulawesi, Toli-Toli,

xi-xii.1895, H. Fruhstorfer (2 4, IRSN, RMNH); 47 km

WSW Kotamobagu, Dumoga Bone N. P., Toraut, 211 m,

vi.1985, G. R. Else (2 9 4 6, BMNH, RMNH); Mt. Sapoetan,

viii.1937, C. T. & B. B. Brues (1 9, MCz).

Diagnosis

Similar to severini and diodon; the female differs

in having the petiole as long as hind basitarsus

(longer in diodon and severini), the male has the

placoids restricted to 4th and 5th flagellomere, and

the gonostyles differ markedly in shape.

Description

Female. - Body length 15.5-18.5 mm, length of

forewing 11.5-13.0 mm. Black; mesoscutum with

oily shine; posterior margins of tergites brown.

Wings yellowish, with sharply defined dark mark

apically. Erect pilosity of head and mesosoma yel-

low; facial tomentum golden; mesosoma with con-

spicuous golden tomentum on pronotum, sides of

mesoscutum, mesopleuron, metanotum, and sides

and declivity of propodeum. Clypeus 1.1 times as

long as IOC, without median carina; IOV 1.1 times

10C; relative lengths of flagellomeres I-III 1: 0.8 :

0.9. Mesoscutum densely rather coarsely punctate,

punctures on disc 0.5-1 diameter apart; propodeal

dorsum shiny, rather coarsely rugose-punctate. Pe-

tiole 0.95-1.05 times as long as hind basitarsus.

Male. - Body length 14.5-17.0 mm, length of

forewing 10.5-12.0 mm. Like the female, except for

the following. Erect pilosity of head and mesosoma

white; facial tomentum silvery; mesosoma without

conspicuous tomentum. ‘Clypeus with vague me-

dian carina; IOV 1.2 times 10C; relative lengths of

flagellomeres I-III 1: 1.0 : 1.15; fourth and fifth

flagellomere with broad convex sensory area. Me-

soscutum sparsely rather coarsely punctate, punc-

tures on disc 1-2 diameters apart. Petiole 1.05 times

as long as hind basitarsus; eighth sternite apically

angularly emarginate; gonostyle in lateral view

strongly constricted in the middle, apically swollen.

Etymology

Cestra is an arbitrary combination of letters, to

be treated as a noun in apposition.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 13-33. 13-18, Sphex, clypeus and mandibles. - 13, muticus, ®; 14, muticus, &; 15, solomon, 2; 16, solomon, 3;

17, praedator, &; 18, walshae, &. 19-33. Isodontia, proximal flagellomeres. - 19, aurifrons, 2; 20, pilipes, 9; 21,

praslinia, 9; 22, aurifrons, &; 23, formosicola, &; 24, pilipes, 8; 25, chrysorrhoea, &; 26, capillata, 3; 27, cestra, 8;

28, delicata, 8; 29, diodon, 8; 30, egens, 6; 31, immaculata, 8; 32, papua, 8; 33, severini, &.

Isodontia delicata sp. n.

(figs. 8, 28, 49)

Type material. - Holotype 8, Indonesia, Irian Jaya,

Jayapura, vii.1938, L. J. Toxopeus, Neth. Ind.-Amer. New

Guinea Exp. 1939-39 (RMNH). — Paratypes: Indonesia:

same data as holotype (5 6, RMNH, BMNH); same locality,

vi.1937 (1 @), xii.1936 (2 &), W. Stüber (RMNH).

Diagnosis

Similar to papza (male, female in both species

unknown), with which it shares the combination of

extremely short flagellomeres I-III, and long nar-

row concave placoid on fourth flagellomere; deli-

cata however has similar placoids on fifth and sixth

flagellomere, and fringes of yellow setosity on the

posterior tergites; these traits are lacking in papa.

Description

Male. - Body length 13.5-15.5 mm, length of

forewing 10.5-11.5 mm. Black; mesoscutum with

oily shine; sides of first tergite and posterior mar-

gins of other tergites and sternites reddish. Wings

slightly yellowish hyaline, apically infuscated. Erect

pilosity of head and mesosoma yellow; facial to-

mentum golden; mesosoma with conspicuous

golden tomentum on pronotum, sides of mesoscu-

tum, mesopleuron, sides of propodeum; fourth to

eighth sternite with fringes of yellow setosity. Cly-

peus 1.0 times as long as IOC, with vague median

carina; IOV 1.0 times IOC; relative lengths of flagel-

lomeres I-III 1: 1.5 : 2.1; third flagellomere with

small apical placoid, fourth to sixth with narrow

concave placoid. Mesoscutum densely rather coar-

sely punctate, punctures on disc 0.5-1 diameter

apart; propodeal dorsum shiny, anteriorly trans-

versely strigose, posteriorly punctate-strigose. Pe-

tiole 1.2-1.3 times as long as hind basitarsus; eighth

sternite truncate; gonostyle simple.

Etymology

Delicata is an adjective, from Latin delicatus =

delicate.

Isodontia diodon (Kohl)

(figs. 29, 50)

Sphex (Isodontia) diodon Kohl, 1890: 377. Lectotype &

(present designation): ‘Celebes, Sumatra’ (NMW!!)

Sphex maia Bingham, 1894: 379. Syntypes, 6, 9: Burma,

Tenasserim (BMNH!!).

Distribution. - This species occurs on the Asian

continent, from Nepal to China and Peninsular

Malaysia.

Notes. - 1. The lectotype of Sphex diodon Kohl

is a male, labelled ‘Stevens, 860’, ‘diodon typ, det

Kohl’ (NMw). Kohl gave as the habitat of the spe-

HENSEN: Malestan Sphecina

cies ‘Celebes, Sumatra’, but this is almost certainly

incorrect. The species that corresponds to both the

type and the description is found only on the Asian

continent.

2. I. diodon has been regarded as a polytypic

species by past workers (Van der Vecht 1957, Bo-

hart & Menke 1976). However, typical I. diodon

differs in several morphological aspects from the

forms associated with it in the past. I. nigelloides

(Strand) is a distinct species. Three other forms are

treated here as belonging to I. severini (Kohl). The

three species are allopatric: I. nigelloides [stat. n.)

is restricted to Southern India and Sri Lanka; I.

diodon occurs on the Asiatic mainland from Nepal

to China and Malaysia, whereas I. severini occurs

in the Archipelago.

Isodontia egens (Kohl)

(figs. 30, 51)

Sphex (Isodontia) egens Kohl, 1898: 335. Holotype &:

New Britain, Kinigunang (TMB!!).

Distribution. - New Britain, Western New Gui-

nea, Biak.

Isodontia franzi (Cameron)

Sphex (Isodontia) franzi Cameron, 1902: 246. Holotype

4: Borneo (BMNH!!).

Distribution. - Only the type specimen from

Borneo is known.

Isodontia immaculata sp. n.

(figs. 31, 52)

Type material. - Holotype d, Papua New Guinea, New

Ireland, Trail to Lelet, Plateau near Danum, 15.xii.1969,

J. E. Tobler (CAs). - Paratypes: Papua New Guinea: same

data as holotype (2 6, CAS, RMNH); Lavongai [New Han-

over], Banatam, 24.iii.1962, Noona Dan Exp. (1 9,

UZMC).

Diagnosis

Within the diodon-group, this species is the

least tomentose one: the facial tomentum is very

sparse and silvery, the mesosoma lacks tomentum,

whereas all other species have conspicuous spots

of silvery or golden tomentum on pronotal lobes,

angles of propodeum, etc. Structurally, the spe-

cies is similar to egens and permutans, males of

these species share the combination of extremely

short proximal flagellomeres, and broad flat pla-

coids on third to sixth flagellomere; immaculata

differs markedly in the shape of the male gono-

style.

197

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Description

Female. - Body length 17.5 mm, length of fore-

wing 12.5 mm. Black. Wings clear, forewing ante-

riorly and apically infuscate, the dark mark not

sharply delimited. Erect pilosity of head and mes-

osoma white; facial tomentum silvery, sparse; mes-

osoma without tomentum. Clypeus 1.1 times as

long as IOC, without median carina; IOV 1.0 times

10C; relative lengths of flagellomeres I-III 1: 0.95

: 1.0. Mesoscutum densely rather coarsely punctate,

punctures on disc 0.5-1 diameter apart; propodeal

dorsum shiny, rather coarsely rugose-punctate. Pe-

tiole 1.1 times as long as hind basitarsus.

Male. - Body length 16.0 mm, length of forewing

11.5 mm.

Like the female except for the following. Clypeus

1.1 times as long as IOC, with vague median carina;

relative lengths of flagellomeres I-III 1 : 1.4 : 2.4

; third to sixth flagellomere with broad flat sensory

area. Mesoscutum densely rather coarsely punctate,

punctures on disc 1-1.5 diameters apart; propodeal

dorsum shiny, rather coarsely rugose-punctate. Pe-

tiole 1.1 times as long as hind basitarsus; eighth

sternite apically not emarginate; gonostyle apico-

dorsally swollen and rounded.

Etymology

Immaculata is an adjective, from Latin zmmacu-

latus = immaculate.

Isodontia papua sp. n.

(figs. 32, 53)

Type material. - Holotype @, Indonesia, New Guinea,

‘Neth. Ind.-American New Guinea Exped., Araucaria

Camp, 800 m, 30.III.1939, L. J. Toxopeus’ (RMNH).

Diagnosis

Similar to delicata (male, female of both species

unknown), with which it shares the combination of

extremely short proximal flagellomeres, and a long

narrow, concave placoid on the fourth flagelio-

mere. However, delicata has similar placoids on

fifth and sixth flagellomere, whereas papua only

has a vestigial one on fifth, and none on sixth.

Additionally, papua lacks fringes of yellow setosity

on the sternites, as found in delicata.

Description

Male. - Body length 15.0 mm, length of forewing

11.5 mm. Black; mesoscutum with oily shine; pos-

terior margins of tergites and sternites reddish to

yellow. Wings light brownish hyaline, apically in-

fuscated. Erect pilosity of head and mesosoma yel-

low; facial tomentum silvery; mesosoma with con-

spicuous golden tomentum on pronotum, sides of

mesoscutum, mesopleuron, etc.; posterior tergites

with appressed pale yellow pilosity. Clypeus 1.0

times as long as IOC, with vague median carina; IOV

1.1 times IOC; relative lengths of flagellomeres I-

II 1: 1.45 : 2.0; fourth flagellomere with long,

narrow, concave placoid, fifth with small apical

placoid. Mesoscutum densely rather coarsely punc-

tate, punctures on disc 0.5-1 diameter apart; prop-

odeal dorsum shiny, anteriorly transversely punc-

tate-strigose, posteriorly reticulate-punctate.

Petiole 1.2 times as long as hind basitarsus; sixth

to eighth sternite with scattered pilosity poste-

riorly, without real fringes; eighth sternite trun-

cate; gonostyle simple.

Etymology

Papua is noun in apposition, it is the common

name for the native inhabitants of New Guinea.

Isodontia permutans (Turner)

(fig. 54)

Sphex permutans Turner, 1912: 198. Lectotype 9 (pres-

ent designation): New Guinea, Wataikwa River

(BMNH!!)

Distribution. - New Guinea, Solomon Islands.

Notes. - 1. The males of permutans and its

closest relatives are all easily recognized using

characters of the genitalia and secondary sexual

characters. The females of permutans, papua, del-

icata, and capillata are apparently inseparable. As

the type of permutans is a female, the identity of

this species is uncertain. I have associated the most

common and widely distributed type of male with

permutans, and described the others as new spe-

cies. I have not been able to associate females with

these. If characters are eventually found to identify

females of these species, my interpretations may

need revision.

2. I. permutans is geographically variable with

respect to the colour of pilosity and tomentum.

Typically (New Guinea) the pilosity is pale yellow,

the tomentum golden. Specimens from Bougain-

ville are similar to the typical form. The pilosity of

head and mesosomal dorsum is partly to entirely

dark in specimens from the other Solomon Islands.

The tomentum is silvery in specimens from Gizo,

Kolombangara, San Cristobal, and Malaita.

Isodontia severini (Kohl) stat. rev.

(figss 3325)

Sphex (Isodontia) severini Kohl, 1898: 337, 9. - Java

(IRSN).

Sphex malayanus Cameron, 1902: 134. Lectotype &

(present designation): Borneo, Kuching (BMNH!!)

Sphex (Isodontia) severini var. philippensis Rohwer,

1921: 669. Holotype 9: Palawan (USNM!!).

Isodontia diodon alemon Vecht, 1957: 367. 4: Sumba

(NMB) Syn. n.

Distribution. — Malaysia, Singapore, Indonesia

(Sumatra, Java, Borneo, Sumba, Halmaheira), Phi-

lippines.

Group of Isodontia ochroptera

Diagnosis

Body length ¢ 16-22 mm, 9 20-22 mm. Labrum

of female with median carina; mandible in both

sexes tridentate, basal inner tooth in male strongly

developed, projecting beyond second inner tooth;

clypeus of female with pair of apical teeth medially;

first flagellomere in female distinctly longer than

second, in male first and second flagellomere

equally long; third to eighth flagellomere of male

with broad, convex, papillate sensory areas; an-

tenna of female without sensory areas; outer side

of hind tibia with five spines; petiole a little shorter

than hind basitarsus.

Species included

Species treated. - I. ochroptera (Kohl).

Other included species. — I. sonani (Tsuneki)

from Taiwan.

Isodontia ochroptera (Kohl)

(figs. 9-10, 34)

Sphex (Isodontia) ochropterus Kohl, 1890: 378. 2: In-

donesia, Sulawesi, Bantimurung (Mus. Dresden, prob-

ably destroyed).

Distribution. - Continental Asia from Fastern

India to Laos and Malaysia, Indonesia (Sumatra,

Borneo, Java, Sulawesi, Aru, Halmaheira), Philip-

pines (Palawan, Samar)

Group of Isodontia praslinia

Diagnosis

Body length & 15-23 mm, 9 16-23 mm. Labrum

of female with a pair of short, parallel, projecting

carinae apically; mandible of both sexes tridentate,

basal inner tooth strongly developed, projecting

beyond second inner tooth; clypeus of female with

median notch; first flagellomere longer than se-

cond; flagellum of male with generally with flat

placoids on flagellomeres IV-VI; antenna of female

without sensory areas. Scutellum and metanotum

convex; propodeal dorsum more or less reticulate;

outer side of hind tibia with 5-10 spines. Petiole

usually shorter than hind basitarsus.

HENSEN: Malesian Sphecina

Species included

Species treated. - I. e/sez sp. n., jaculator (Smith),

nidulans sp. n., praslinia (Guérin-Méneville), se-

picola (Smith).

Other included species. - I. obscurella (Smith)

from Australia, I. xanthognatha (Perez) (= nigella

(Smith), nec Lichtenstein) [stat. n.] from contin-

ental Asia.

Key to species

1. Pilosity of head and mesosoma dark brown to

black; wings dark brown with violaceous shine;

Timor (?), New Guinea and neighbouring is-

la ads praslinia (Guérin-Méneville)

— Erect pilosity of head and mesosoma white to

yellowish; colour of wings variable .......... 2

2. Propodeal dorsum sparsely punctate, interspa-

ces smooth, shiny; interocular distance across

vertex shorter than across clypeus; Philippines

ren. sr cht dal nidulans sp. n.

- Propodeal dorsum punctate-reticulate to stri-

gose-punctate, more or less dull, without

SMOothinters paces ks. "dr Cee 3

3. Facial tomentum sparse, silvery, sculpture of

clypeus easily visible; Moluccas .................

RE RR OA jaculator (Smith)

- Facial tomentum dense, golden or silvery,

sculpture of clypeus largely obscured ........ 4

4. Petiole of female 0.8-0.95, of male 0.9-1.0

times as long as hind basitarsus; IOV 0.85-1.05

times IOC; male: mandibles entirely bright red-

dish; New Guinea and neighbouring islands,

Northern Australia .......... sepicola (Smith)

— Petiole of female 0.95, of male 1.05-1.1 times

as long as hind basitarsus; IOV 1.1 times IOC;

male: basal third of mandibles black; Sulawesi

reen teater ae cai elsei sp. n.

Isodontia elsei sp. n.

(fig. 35)

Type material. - Holotype 8, Indonesia, Sulawesi, ‘Ro-

senberg, Toelabella, Celebes’ (RMNH). — Paratypes. In-

donesia, Sulawesi: Gorontalo (1 &), Kwadang (1 @), both

Rosenberg (RMNH); Patunuang, i.1896, H. Friihstorfer (2

Q, RMNH); 47 km WSW Kotamobagu, Dumoga Bone N.

Pk., Toraut (Forest Edge), 211 m, v-vi.1985, G. R. Else (6

3, 3 9, BMNH, RMNH); Luwuk, Matanyo Forest, N. of

Kayutano, 120 m, 20.x.1989, C. van Achterberg & M.

Tulung (1 9, RMNH).

Diagnosis

Very similar to sepicola, from which it differs

only in the characters mentioned in the key.

Description

Female. - Body length 19.0-20.5 mm; length of

forewing 13.5-15.5 mm. Black; mesoscutum with

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

oily shine. Wings light brownish, apical margins

more strongly infuscated, with violaceous shine.

Erect pilosity of head and mesosoma nearly white;

facial tomentum golden, dense, leaving sculpture of

clypeus visible under a certain angle only; meso-

soma without conspicuous tomentum. Clypeus 0.8

times as long as 10C, without discernable median

carina; IOV 1.1 times 10C; relative length of flagel-

lomeres I-III 1 : 0.8 : 0.8. Mesoscutum shiny, spar-

sely finely punctate, punctures on disc 1-3 diame-

ters apart; propodeal dorsum more shiny than in

I. sepicola, weakly transversely rugose and shal-

lowly punctate. Petiole 0.95 times as long as hind

basitarsus.

Male. - Body length 15.5-20.5 mm; length of

forewing 13.5-15.5 mm. Like the female except for

the following. Black; mandibles with short reddish

zone just before apex. Wings brownish hyaline,

apical margins infuscated. Erect pilosity of head

and mesosoma yellowish to white. Clypeus 0.85

times as long as 10C; relative length of flagello-

meres I-III 1 : 0.8 : 0.85. Petiole 1.05-1.1 times as

long as hind basitarsus; third to seventh sternite

with fringes of brown setosity; gonostyle simple.

Etymology

Elsei is a noun in the genitive case, referring to

Mr. George R. Else, collector of most of the type

series, and many other interesting Sulawesi wasps.

Isodontia jaculator (Smith)

(fig. 36)

Sphex jaculator Smith, 1860: 122. Holotype &: Indonesia,

Bacan (OUM!!).

Distribution. — Exclusively known from the is-

land Bacan, in the Indonesian Northern Moluccas.

Isodontia nidulans sp. n.

Type material. - Holotype 9, Philippines, Luzon,

‘Olongapa, P. I., Subic Bay, V.1967’, ‘Coll. by J. C. Thom-

son’ (USNM).

Diagnosis

Immediately separated from other species of this

group by its very fine sculpture; in particular the

propodeal dorsum is smooth, shiny, with sparse

fine punctuation.

Description

Female. - Body length 20.0 mm; length of fore-

wing 15.5 mm. Black; mesoscutum with oily shine.

Wings brown, with violaceous shine. Erect pilosity

of head and mesosoma white; facial tomentum

silvery, sparse; mesosoma without conspicuous to-

mentum. Clypeus 0.7 times as long as 10C, with

blunt median carina on dorsal half; 10v 0.95 times

10C; relative length of flagellomeres I-II 1 : 0.75 :

0.75. Mesoscutum shiny, obsoletely punctate; prop-

odeal dorsum shiny, along anterior margin with

weak rugae, remaining part smooth, finely, shal-

lowly punctate. Petiole 0.9 times as long as hind

basitarsus.

Etymology

Nidulans is an adjective based on the Latin verb

nidulare = to shine.

Isodontia praslinia (Guérin-Méneville)

(figs. 11-12, 21, 37)

Sphex praslinius Guérin-Méneville, 183 1: pl. 8. Holotype

Q: New Ireland, Pt. Praslin (MCG!).

Sphex (Isodontia) ustulata Kohl, 1890: 378. Holotype 9:

Timor (NMW!!). Syn. n.

Sphex (Isodontia) simplex Kohl, 1898: 334. Lectotype &

(present designation): New Guinea: Erima, Astro-

labe-Bay (TMB!!). Syn. n.

Sphex (Isodontia) insularis Cameron, 1901: 240. Holo-

type 6: New Britain (BMNH!!). Syn. n.

Distribution. - This species occurs on New Gui-

nea and neighbouring islands: New Britain, New

Ireland, Misool Island, the Solomon Islands. The

record from Timor is based only on the type of

ustulata, and it is doubtful whether the species

indeed occurs there. Species whose occurrence is

centered on the Papuan subregion, like I. praslinia,

and also for example I. sepicola and S. cognatus

rarely occur in the Lesser Sunda islands. The two

areas are climatically quite different.

Isodontia sepicola (Smith)

(fig. 38)

Sphex sepicola Smith, 1859: 158. Lectotype 9 (present

designation): Aru (OUM!!)

Sphex albohirtus Turner, 1908: 466. Lectotype (present

designation): Australia: Mackay (BMNH!!). Syn. n.

Distribution. - This species occurs on New Gui-

nea, and neighbouring islands: Aru islands, So-

lomon islands, Misoöl, and in Australia, where it

seems to be restricted to Northern Queensland.

Genus Sphex Linnaeus

Group of Sphex argentatus

Diagnosis

Body length & 20-28 mm, 9 23-34 mm. Mand-

ible of both female and male bidentate; antenna of

male with placoid(s), at least on fifth flagellomere;

scutellum convex, medially impressed; metanotum

medially raised into a pair of tubercles, or into a

single medially impressed tubercle; claw teeth

large, at sharp angle to claw.

Species included

Species treated. — S. alacer Kohl, argentatus Fa-

bricius, diabolicus Smith, finschu Kohl, madasum-

mae Van der Vecht, sericeus Fabricius

Other included species. — carbonicolor Vecht,

decoratus Smith, ephippium Smith, modestus

Smith (all Australian), fumicatus Christ (Africa to

SW. India and Sri Lanka).

Key to species

1. Propodeal dorsum with 4-6 transverse ridges

- Propodeal dorsum without ridges ........ pig

2. Female: apical margin of clypeus with median

tooth; male: 2nd to 6th flagellomere with

broad placoid (fig. 39); New Guinea............

BANE OE SR dara alacer Kohl

- Female: apical margin of clypeus without

tooth; male: 4rd or 4th to 6th flagellomere

with placoid (fig. 42); Sri Lanka to New Guinea

bbl ark aisha ee sericeus (Fabricius)

3. Wings uniformly dark brown, with violaceous

effulgence; Burma to New Guinea, Philippines

A bia teeta melas. madasummae Van der Vecht

- Wings clear or yellowish, only at basis and apex

sometimessdarkenede eh no 4

4. Wings strongly yellowish to yellow-brown; Sri

lanka to Sulawesi ............ diabolicus Smith

- Wings essentially clear, infuscate at base and

Aperen kend Mantes ile. 5

5. Metanotum with single medially impressed

tubercle; male: 4th to 6th flagellomere with

broad placoids (fig. 41); Moluccas, New Gui-

nea, Solomon Islands, Australia finschi Kohl

— Metanotum strongly bituberculate; male: only

Sth flagellomere with narrow placoid (fig. 40);

widespreade nn argentatus Fabricius

Sphex alacer Kohl

(fig. 39)

Sphex alacer Kohl, 1895: 54. Holotype 9: New Guinea

(NMW!).

Distribution. - New Guinea, New Ireland.

Sphex argentatus Fabricius

(fig. 40)

Sphex argentatus Fabricius, 1787: 274. Lectotype 9 (de-

signated by Van der Vecht 1961: 28): India, Coroman-

del (Mus. Lund!).

HENSEN: Malesian Sphecina

Sphex umbrosus Christ, 1791: 293, pl. 29. No locality

given [type lost].

Sphex argenteus Turton, 1800: 485. India, Coromandel

[possible emendation].

Sphex argentifrons Lepeletier, 1845: 337. Lectotype à

(designated by Menke in Bohart & Menke 1975: 114):

Java (Mus. Torino!).

Sphex plumifer Costa, 1864: 112. Philippines, Luzon

(Mus. Naples).

Sphex umbrosus var. nanulus Strand, 1913: 83. Holotype

Q: Taiwan, Taihorin (DEI!!).

Sphex umbrosus var. fumosus Kohl, 1890: 408. Japan

(TMB).

Distribution. - Sri Lanka and India to Indochina

and Japan; Malaysia, Indonesia, Philippines, Papua

New Guinea and NE. Australia.

Note. - Sphex unicolor Fabricius, 1787, listed as

a synonym of argentatus by Bohart & Menke

(1975), was described from Spain where argentatus

does not occur. Van der Vecht (1961) suggested

that the name might refer to S. fumicatus Christ,

1791, but that species has never been reported

from Spain either. The only Spanish species of

Sphex that agrees with the description is S. afer

Lepeletier. Alternatively, the locality may be incor-

rect.

Sphex diabolicus Smith

Sphex diabolicus Smith, 1858: 100. Lectotype 9 (desig-

nated by Van der Vecht 1973): Sarawak (OUM!)

Sphex flammitrichus Strand, 1913: 83. Lectotype 2 (pres-

ent designation): Taiwan (DEI!!)

Sphex umbrosus var. aureopilosus Berland, 1928: 330.

Lectotype @ (designated by Menke in Bohart & Menke

1975): Ba-Cha, Tonkin (MNHN!!)

Sphex fulvohirtus Bingham, 1890: 242. Holotype 9: Sri

Lanka, Pundalaya (BMNH!!). Syn. n.

Types. - The lectotype of Sphex flammitrichus

is a female labelled ‘Kankau (Koshun), Formosa,

H. Sauter, vii.1912’, the paralectotypes are 3 males,

labelled ‘Korosan, Formosa, Sauter, xi.07’ (all DEI).

Distribution. - India and Sri Lanka to Malaysia,

Indonesia (Sumatra and neighbouring islands, W.

Java, Borneo, Sulawesi), Philippines.

Sphex finschii Kohl

(fig. 41)

Sphex finschu Kohl, 1890: 412. Lectotype & (present

designation): New Britain (ZMB!!)

Distribution. - Indonesia (Ambon, Aru Islands,

Irian Jaya), Papua New Guinea, Bismarck Islands,

Solomon Islands, Australia.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Sphex madasummae Van der Vecht

Sphex caerulescens Guillou, 1841: 320. Holotype £: Bor-

neo (MNHN!!) [non Reich, 1795].

Sphex maurus Smith, 1856: 255. Syntypes, 9, &: Sulawesi

(BMNH!!) [non Fabricius, 1787].

Sphex nigerrimus Costa, 1864: 112. Philippines, Luzon

(Naples Museum?) [non Scopoli, 1763].

Sphex madasummae Van der Vecht, 1973: 345 [new

name for Sphex maurus Smith].

Distribution. - Malaysia, Indonesia (not known

from Nusa Tenggara); Philippines, Papua New

Guinea.

Sphex sericeus Fabricius

(fig. 42)

Sphex aurulentus Fabricius, 1793: 201. - India, Tranqu-

ebar (location of type unknown) [non Fabricius,

1787].

Sphex sericeus Fabricius, 1804: 211. Lectotype 9 (desig-

nated by Van der Vecht 1961): ‘in maris pacifici Insu-

lis’ (Mus. Lund!)

Sphex fabricu Dahlbom, 1843: 27. 9: India, Tranquebar

(location of type unknown).

Sphex ferrugineus Lepeletier, 1845: 345. 9: ‘sans patrie,

mais exotique’ (MNHN?).

Sphex lineolus Lepeletier, 1845: 353. &: Java (MNHN?)

[locality probably erroneous].

Sphex ferox Smith, 1862: 55. Syntypes, 9, 4: Ambon,

Sulawesi (BMNH!!, OUM) [non Westwood, 1837].

Sphex lepeletierii Saussure, 1867: 40. Holotype 9: Java

(NMW!!) [locality probably erroneous |.

Sphex godeffroyi Saussure, 1869: 57. Q: Australia,

Queensland, Cape York (Mus. Hamburg).

Sphex aurifex Smith, 1873: 460, pl. xliv, fig. 3. Holotype

Q: NW. Australia, Champion Bay (BMNH!!).

Sphex aurulentus var. pallidehirtus Kohl, 1890: 393. Syn-

types 6: New Guinea: Port Moresby; Ambon (NMW?).

Sphex rugosus Matsumura, 1912: 176, 53, fig. 8. 9: Tai-

wan: Horisha, Koshun.

Sphex sericeus ferocior Van der Vecht & Krombein, 1955:

40 [new name for Sphex ferox Smith].

Sphex sericeus nigrescens Van der Vecht & Krombein,

1955: 39. Holotype ©: Philippines, Luzon, Los Bafios

(USNM).

Sphex sericeus stueberi Van der Vecht & Krombein,

1955: 42. Holotype Q: Indonesia, Irian Jaya, Jayapura

(RMNH!!).

Sphex sericeus wegneri Van der Vecht & Krombein,

1955: 39. Holotype &: Indonesia, Kalimantan, Sama-

rinda, Mura Kaman (RMNH!!).

Distribution. - India and Sri Lanka to China and

Indochina, Malaysia, Indonesia, Philippines, Papua

New Guinea, N. Australia.

Note. - This species occurs in many colour-

forms, which are largely allopatric. Van der Vecht

and Krombein (1955) recognized eight subspe-

cies.

Group of Sphex resplendens Smith

Diagnosis

Body length & 17-24 mm, 9 20-33 mm. Mand-

ible of both female and male bidentate; antenna of

male without placoids; scutellum of female flat or

very weakly convex, conspicuously shiny, of male

convex, medially not or hardly impressed; claw

teeth small, perpendicular to claw.

Species included

Species treated. - S. brachystomus Kohl, con-

frater Kohl, dorycus Guérin-Méneville, mimulus

Turner, muticus Kohl, resplendens Kohl, solomon

sp. n., wilsoni sp. n.

Other included species. - S. fumipennis Smith, S.

gilberti Turner, S. /uctuosus Smith (all from Aus-

tralia), and S. antennatus Smith stat. n. (from New

Caledonia, New Hebrides and Loyalty Islands;

listed as a subspecies of S. fumipennis by Bohart &

Menke 1976).

Key to species

1. Metasoma partly red (sometimes only part of

POSTEIOLICELOILES) REI eee PAPER eee a 2

= 'Metasoma entirely black ist. 4

2. Pilosity of mesosoma white; 8th sternite of

male with narrow apical process; Australia,

News Guinea AR mimulus Turner

- Pilosity of mesosoma dorsally yellow to black,

ventrally brownish to black; 8th sternite of

male triangular, without process ............ 3

3. Metasoma except petiole entirely red; pilosity

of mesosoma dorsally yellow to brownish; pe-

tiole of female longer than wide; gonostyles of

male without inner lobe; New Guinea

DEE, AMI MOREE oe dorycus Guérin-Méneville

— Metasoma with only posterior tergites and

sternites red; pilosity of mesosoma black; pe-

tiole of female as long as wide; gonostyles of

male ventrally with flat, almost circular inner

lobe (fig. 56); Solomon and Bismarck Islands

TUE LEMON O sil rara confrater Kohl

4. Mesonotum densely covered with silvery or

golden tomentum, completely obscuring sculp-

WILLIE 5

- Mesonotum without or with sparse tomentum,

sculpturesvisibleichroushout ame 6

5. Tomentum golden; wings yellowish, apical

margins infuscate; New Guinea ................

ACRI ER OLE wilsoni sp. n.

- Tomentum silvery; wings dark brown, with

violaceous effulgence; New Guinea, Bismarck

Islands on RETE brachystomus Kohl

6." Bemalesıı 1. Mes TALL ed ren 7

MOMENT E NEN a EEE: ©)

7. Mandibles long, strongly curved (fig. 13),

when closed reaching beyond base of opposite

mandible; apical margin of clypeus strongly

arcuate; median carina of labrum projecting

into a small apical tooth; Buru, Ambon

BEER So tri eG I muticus Kohl

- Mandibles short, moderately curved (fig. 15),

when closed not reaching base of opposite

mandible; anterior margin of clypeus weakly

arcuate; median carina of labrum not project-

Inga picallll Ake Australian di ae ithe, Muineent; 8

8. Interocular distance at vertex longer than

combined lengths of flagellomeres II + III;

metanotum without silvery tomentum; Moluc-

cas, New Guinea, Philippines, Australia

MERI Rn resplendens Kohl

— Interocular distance at vertex equal to com-

bined lengths of flagellomeres II + III; meta-

notum densely covered with silvery tomentum;

Solomon Islands ................ solomon sp. n.

9. Mandibles with basal half straight, sharply

curved inward distally (fig. 14); petiole in dor-

sal view more than 2 times as long as wide;

gonostyles not emarginate, the apex simple;

BUGURATIDONERE SR muticus Kohl

— Mandibles smoothly curved, approximately a

quarter-circle (fig. 16); petiole in dorsal view

less than 2 times as long as wide; gonostyles

dorsally emarginate, the apex produced into

tooth pointing upward (fig. 58) ............ 10

10. Pilosity of head and mesosoma dark brown to

black; wings dark brown; Philippines, Moluc-

caseiNewGuineasAustraliad SER CNR

Ans esi vio EK Din resplendens Kohl

- Pilosity of head and mesosoma yellowish,

partly dark brown on head and mesosomal

dorsum; wings brownish hyaline; Solomon Is-

lands. esi esi pal Dal solomon sp. n.

Sphex brachystomus Kohl

Sphex brachystomus Kohl, 1890: 415. Syntypes 9 4: New

Britain (NMW!).

Distribution. - Papua New Guinea, Bismarck

Islands (New Ireland, New Britain).

Sphex confrater Kohl

(fig. 56)

Sphex confrater Kohl, 1890: 414. Syntypes 9: New Bri-

tain (Mus. Hammon’).

? Sphex confrater var. sieberti Strand, 1910: 46. Syntypes

Q: Java, Batavia (location of types unknown).

Distribution. - Solomon Islands (Kolomban-

gara, Guadalcanal), Papua New Guinea, Bismarck

Islands (New Britain, New Ireland).

HENSEN: Malesian Sphecina

Note. — Judging on the type-locality, it seems

very unlikely that steberti Strand is indeed a variety

of confrater, but it is not clear from the description

to which other species it might pertain.

Sphex dorycus Guérin-Méneville

Sphex dorycus Guérin-Méneville, 1838: 262. Holotype 9:

New Guinea, Manokwari (‘Dory’) (MCG).

Sphex errabundus Kohl, 1898: 338. Holotype 2: Indone-

sia, Waigeo ( Waigiu') (TMB!).

Distribution. - Indonesia (Waigeo, Irian Jaya),

Papua New Guinea.

Note. - The synonymy of this species was dis-

cussed by Van der Vecht (1973).

Sphex mimulus Turner

Sphex mimulus Turner, 1910b: 419. Holotype 9: Austra-

lia, Queensland, Cairns (BMNH!).

Distribution. - Australia (Queensland), S. Papua

New Guinea.

Sphex muticus Kohl

(figs. 13-14, 57)

Sphex muticus Kohl, 1885: 199. Lectotype @ (designated

by Van der Vecht 1973): Indonesia, Ambon (NMW!)

Distribution. - Indonesia (Buru, Ambon).

Sphex resplendens Kohl

Sphex nitidiventris Smith, 1859: 158. Holotype 9: In-

donesia, Buru (OUM!) [non Spinola, 1851].

Sphex gratiosus Smith, 1859: 158. Holotype @: Indone-

sia, Aru Islands (OUM!) [non Smith, 1856].

Sphex resplendens Kohl, 1885: 200 [new name for S.

nitidiventris Smith].

Sphex gratiosissimus Dalla Torre, 1897: 424 [new name

for S. gratiosus Smith, 1859].

Sphex wallacei Turner, 1908: 467 [new name for S. nit-

tdiventris Smith].

Sphex lanceiventris Vachal, 1908: 23 [new name for S.

nitidiventris Smith].

Sphex mertoni Strand, 1911: 231. Holotype 2: Indonesia,

Aru Islands, Kobroör, Seltutti (Mus. Senckenberg!!).

Distribution. - Philippines (Negros, Luzon); In-

donesia (Aru Islands, Key Islands, Bacan, Ambon,

Irian Jaya); Papua New Guinea, Australia (Queens-

land). A few specimens from Bangka in the RMNH

are probably incorrectly labelled.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 34-61. Isodontia and Sphex. 34-45, proximal flagellomeres of 8. - 34, I. ochroptera; 35, I. elset, 36, I. jaculator,

37,1. praslinia; 38, I. sepicola; 39, S. alacer; 40, S. argentatus; 41, S. finscht, 42, S. sericeus; 43, S. rex; 44, S. subtruncatus,

45, S. walshae. 46-58. Gonostyle of 4, 46-55 + 57-58, lateral view, 56, ventral view. — 46, 1. pilipes; 47, I. capillata;

48, I. cestra; 49, I. delicata; 50, I. diodon; 51, L egens; 52, I. immaculata; 53, I. papua; 54, I. permutans; 55, I. severint,

56, S. confrater; 57, S. muticus; 58, S. solomon. 59-60, tarsal claw. 59, S. solomon, Q; 60, S. subtruncatus, Q. 61, S. rex,

G, eighth sternite.

Sphex solomon sp. n.

(figs. 15-16, 58-59)

Type material. - Holotype 4, Solomon Islands, Guadal-

canal Is., Lunga, 2.xii.1934, H. T. Pagden (RMNH). — Para-

types: Solomon Islands: Tulagi (3 2), Guadalcanal, Lunga

(1 Q 1 8), Ganonga (1 9 5 8), Nygela (1 ©), all H. T.

Pagden (RMNH); Guadalcanal, Kukum, P. Greenslade (1

©, BMNH); Isabel, Tatamba (1 9, Australian National

Insect Collection, Canberra; 1 9, RMNH); Bougainville,

Buin, G. Daniels (1 9, Australian Museum, Sydney); Gua-

dalcanal, Lunga River Bridge, J. Schenk (1 &, Los Angeles

County Museum); Guadalcanal, Kukum, E. S. Brown (1

& 7 9, BMNH, 2 9, RMNH).

Diagnosis

Similar to resplendens and brachystomus, with

which solomon shares relatively short and weakly

curved mandibles, and an apical lobe on the male

gonostyle. It differs from these species in vestiture:

the silvery tomentum of the mesosoma is restricted

to the pro- and metanotum, particularly conspicu-

ous on the latter, in male, the pilosity of head and

mesosoma is basically pale. Structural features are:

in female, lov equals flagellomere 2 + 3 (longer in

resplendens and brachystomus); in male the apical

lobe of the gonostyle is broad and rounded (narrow,

pointed in the other species).

Description

Female. - Body length 20-25 mm; length of

forewing 15.0-18.5 mm. Black, metasoma with fine

greenish metallic shine. Wings brown. Pilosity of

head and mesosoma dark brown; facial tomentum

brownish to silvery; mesosoma with conspicuous

silvery tomentum on metanotum and sometimes

on pronotum. Clypeus 0.8 times as long as IOC, with

weak median keel on dorsal half, apical margin

slightly raised, with a pair of short teeth medially;

IOV 0.75 times IOC; labrum with median carina,

which does not project apically; relative lengths of

flagellomeres I-III 1 : 0.6 : 0.6. Mesopleuron spar-

sely finely punctate, interspaces punctulate, shiny;

mesoscutum densely, rather coarsely punctate,

punctures 0.5-1 diameter apart, interspaces

smooth, shiny; scutellum slightly convex, sparsely

finely punctate, shiny; propodeum coriaceous, dull.

Fore basitarsus with 8 rake spines, median spines

2 time tarsal width, apical spine as long as second

tarsomere. Petiole 0.3 times as long as hind basi-

tarsus, in dorsal view hardly longer than wide.

Male. - Body length 19-24 mm; length of fore-

wing 13.5-17.0 mm. Like the female except for the

following. Wings brownish hyaline, apical margins

infuscate. Facial tomentum silvery; pilosity of head

and mesosoma basically pale, more or less mixed

with dark hairs or entirely dark on vertex, temples

and mesosomal dorsum. Clypeus 0.95 times as long

HENSEN: Malesian Sphecina

as IOC, apical margin untoothed, medially de-

pressed; IOV 0.85 times IOC; labrum ecarinate; scu-

tellum convex, faintly impressed along midline.

Apical margins of second to seventh sternite with

row of black setae; eighth sternite apically pro-

duced into a tooth, distal third medially keeled;

gonostyle apically with short lobe pointing up-

ward, ventrally in the middle with almost circular

lobe pointing inward.

Etymology

Solomon is a noun in apposition, referring to the

type locality.

Sphex wilsoni sp. n.

Type material. - Holotype 9, Papua New Guinea,

Huon Peninsula, Mongi-Mape Watersheds, Nganduo to

Yunzain, 1000-1500 m, 56.iv.1955, E. O. Wilson (MCz).

Diagnosis

Immediately separated from other species of the

resplendens-group by the conspicuous golden to-

mentum on head and thorax, and the yellow wings.

Structurally, wilsont is similar to confrater, mimu-

lus, dorycus and muticus, with which it shares the

long, strongly curved mandibles.

Description

Female. - Body length 25 mm; length of fore-

wing 19 mm. Black; metasoma with fine metal-

lic shine. Wings yellow, apical margins infuscate.

Pilosity of head and mesosoma dark brown; head,

pro-, meso- and metanotum with dense golden

tomentum, completely obscuring the sculpture of

these parts. Clypeus 0.75 times as long as IOC, with

weak median carina dorsally; apical margin slightly

raised, with a pair of short teeth medially; 10v 0.75

times 10C; labrum with median carina, projecting

apically into a short tooth; mandibles long,

strongly curved, when closed reaching beyond each

other’s bases (cf. fig. 13); relative length of flagel-

lomeres I-III 1 : 0.6 : 0.65. Mesopleuron sparsely

finely punctate, interspaces weakly punctulate; me-

sonotum apparently (obscured by vestiture) den-

sely finely punctate, shiny; propodeum finely cori-

aceous, dull; fore basitarsus with 9 rake spines,

median spines 1.4 times tarsal width, apical spine

shorter than second tarsomere. Petiole 0.25 times

as long as hind basitarsus, in dorsal view hardly

longer than wide.

Etymology

Wilson: is a noun in the genitive case, referring

to E.O. Wilson, collector of the only known spec-

imen of this species.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Group of Sphex subtruncatus Dahlbom

Diagnosis

Body length & 16-28 mm, Q 15-36 mm. Mand-

ible of both male and female bidentate; antenna of

male with placoid(s); scutellum convex, medially

impressed; metanotum simple; claw teeth large, at

sharp angle to claw.

Species included

Species treated. - S. cognatus Smith, formosellus

Van der Vecht, praedator Smith, rex sp. n., staudın-

geri Gribodo, subtruncatus Dahlbom, walshae

sp. n.

Other included species. - S. obscurus (Fabricius)

(Sri Lanka to Thailand, ahasverus Kohl, semifos-

sulatus Van der Vecht, vestitus Smith, ermineus

Kohl, and bilobatus Kohl (all Australian).

Note. - This is obviously a heterogeneous as-

semblage. The species are grouped together only

on basis of the lack of specialized features defining

the other groups. It is undoubtedly paraphyletic,

but I have not been able to find an adequate way of

splitting it up.

Key to species

1. Propodeum with dense golden or silvery to-

mentum, completely obscuring sculpture.... 2

- Propodeum at most with sparse golden or

silvery tomentum, leaving sculpture easily vis-

Iper. ae een el a DI an 3

2. Large species: wing length 26-27 mm; legs

largely reddish; New Guinea ....... rex sp. n.

— Medium-sized species: wing length 13-19 mm;

legs black; Moluccas, New Guinea, Bismarck

and Solomon Islands, Australia …

HR SURNAM ADS ce ERA EN cognatus Smith

3. Female: outer side of mid tibia only with nor-

mal, straight spines apically; male: apical mar-

gin of clypeus straight or slightly concave,

without median lobe (fig. 17) ................ 4

- Female: outer side of mid tibia with stout,

curved spine apically (fig. 64); male: apical

margin of clypeus with short median lobe (fig.

LS) nent ERA urn 6

4. Wings strongly yellowish to dark brown with

violaceous effulgence; India to Moluccas, Phi-

lp pinesRraenana PRIA praedator Smith

- Wings basically clear, at most slightly yellow-

ishiandi'apically infuscate St mr er. 5

5. Pilosity of head and mesosoma dense, yellow-

ish brown; face and mesoscutum with dense

brownish tomentum obscuring the sculpture;

legs reddish; medium-sized species, wing

length 19-20 mm; New Guinea .................

“pin ho Belcher ee staudingeri Gribodo

- Pilosity of head and mesosoma white to pale

yellow; mesoscutum without dense tomentum,

sculpture easily visible; legs black; small spe-

cies, wing length 10.5-13.5 mm; Sumba,

Timor NW AUS tra la ee RE Re

rol BM) PRADA formosellus Van der Vecht

6. Wings dark brown with bluish effulgence, legs

dark; female: sixth tergite with pygidium (fig.

65); male: 3rd to 7th flagellomere with broad

placoids (fig. 45); Burma to Sumatra, Borneo

Lei rara ee DI walshae sp. n.

- Colour of wings and legs variable; female: sixth

tergite without pygidium; male: 3rd flagello-

mere rarely with placoid (only in specimens

from Philippines and Java) (fig. 44); Sri Lanka

to/Philippines and Tlimor tE

ana debe Mione subtruncatus Dahlbom

Sphex cognatus Smith

Sphex cognatus Smith, 1856: 248. Holotype 9: Australia

(BMNH!!).

Sphex amator Smith, 1856: 246. Holotype @: Australia

(OUM!).

Sphex formosus Smith, 1856: 254. Holotype 9: Indone-

sia, Seram (BMNH!!).

Sphex opulentus Smith, 1856: 250. Holotype &: Austra-

lia, Richmond River (BMNH!!).

Distribution. - Indonesia (Maluku, Irian Jaya),

Papua New Guinea, Bismarck Islands, Solomon

Islands, Admiralty Islands, Australia.

Note. - Specimens from the North Moluccas

(Halmaheira etc.) differ from the remaining popu-

lations in having silvery tomentum with a faint

golden tinge, in particular on the propodeum.

Sphex formosellus Van der Vecht

Sphex formosellus Van der Vecht, 1957: 366. Holotype Q:

Timor (RMNH!!).

Distribution. - Indonesia (Timor, Sumba), W.

Australia.

Sphex praedator Smith

(fig. 17)

Sphex praedator Smith, 1858: 14. Holotype &: Indonesia,

Sulawesi (OUM!).

Sphex tyrannicus Smith, 1860: 122. 9: Indonesia, Bacan

(type lost, Van der Vecht 1973).

Sphex luteipennis Mocsary, 1883: 33. Lectotype 9 (desig-

nated by Van der Vecht 1973): Indonesia, Ambon

(TMB!)

Sphex nigripes var. calopterus Kohl, 1890: 168, 422 [as

var. 5]. 6: Indonesia, Sulawesi (NMW).

Sphex nigripes var. kohlianus Strand, 1913: 81 [new

name for S. nigripes var. calopterus Kohl].

Sphex nigripes var. celebesianus Strand, 1913: 81. 9, &:

Indonesia, Sulawesi (NMw) [name for S. nigripes var.

2 of Kohl (1890)].

Sphex nigripes var. melanopodus Strand, 1915: 89. Lec-

totype 9 (designated by Van der Vecht 1973): Sri

Lanka, Nalanda (NMw!)

Distribution. - India and Sri Lanka to China and

Malaysia, Indonesia (Sumatra, Sangihe Islands,

Java, Sulawesi, Moluccas), Philippines.

Sphex rex sp. n.

(figs. 43, 61-62)

Type material. - Holotype 9, Papua New Guinea,

Milne Bay, KBMission, 5.iii.1944, K.V. Krombein

(USNM). — Paratype: Indonesia, Irian Jaya, Ifar, 350 m,

ii.1957, J. v.d. Assem (1 @, RMNH).

Diagnosis

This species is immediately recognized by its

large size (body length 35 mm), reddish legs and

dense golden tomentum on most part of meso-

soma. In these respects it is similar only to S.

basilicus from Australia, from which it differs as

follows: petiole short, 0.35 times as long as hind

basitarsus (0.5 in basilicus); 7th sternite of male

with proximal transverse depression, 8th sternite

trilobate (bilobate in basilicus); anterior femora of

female ventrally covered with tomentum (with

sparse erect pilosity in baszlicus).

Description

Female. - Body length 36 mm, length of forewing

26 mm. Black, the following parts reddish: mand-

ibles except base and apex, femora except basal half,

tibiae, tarsi, tegulae except medially, sixth meta-

somal segment. Wings almost clear, apically infus-

cate. Pilosity yellow, on vertex and mesoscutal disc

infuscate; head and mesosoma with conspicuous

dense golden tomentum, completely obscuring

sculpture of metanotum and propodeum, tomen-

tum on anterior part of mesoscutal disc and scutellar

disc dark brown; fore femora ventrally with tomen-

tum and pilosity. Clypeus 0.85 times as long as IOC,

with blunt median carina on dorsal half, apical

margin with short median lobe; 10v 0.85 times IOC;

labrum with median carina, which projects apically

into a weak tooth; relative lengths of flagellomeres

I-III 1 : 0.6 : 0.6. Mesosoma microsculptured, punc-

tulate to punctulate-reticulate to shagreened (obs-

cured by vestiture almost throughout); scutellum

with median longitudinal impression; fore metatar-

sus with 8 rake spines, median spines 1.5 times

metatarsal width, apical spine as long as second

tarsomere; mid tibia with oblique, slightly curved

spine apically. Petiole 0.35 times as long as hind

basitarsus; sixth tergite without pygidium.

HENSEN: Malesian Sphecina

Figs. 62-65. Sphex. 62, S. rex, Q, fore basitarsus; 63-65,

S. walshae, 2. 63, fore basitarsus; 64, distal part of mid

tibia, outer side; 65, sixth tergite.

Male. - Body length 35 mm, length of forewing

27 mm. Like the female except for the following.

Mandibles black, mid and hind femora nearly en-

tirely black; 7th metasomal segment and 8th ster-

nite reddish. First tergite and sides of 6th and 7th

with conspicuous yellow pilosity. Clypeus 0.95 times

as long as IOC; 10v 0.9 times IOC; labrum ecarinate;

relative lengths of flagellomeres I-III 1: 0.65 : 0.65,

5th flagellomere with narrow concave placoid; sixth

sternite apically shallowly emarginate, seventh

sternite proximally transversely depressed, 8th

sternite trilobate, middle lobe curved upward.

Etymology

Rex is a noun in apposition, from Latin rex =

king.

Sphex staudingeri Gribodo

Sphex staudingeri Gribodo, 1894: 3. Holotype 4: New

Guinea (MCG!).

Distribution. - New Guinea.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Sphex subtruncatus Dahlbom

(figs. 44, 60)

Sphex subtruncatus Dahlbom, 1843: 25. Q: ‘Africa’ (Mu-

seum Lund!) [probably from NE. India, see Van der

Vecht (1973)].

Sphex nigripes Smith, 1856: 254. Syntypes 9: China:

Shanghai (BMNH!), Sumatra [non Fabricius, 1793].

Sphex siamensis Taschenberg, 1869: 413. 9: ‘Siam’ (Mus.

Univ. Halle!) [probably from the Philippines, accord-

ing to Van der Vecht (notes) |.

Sphex erythropodus Cameron, 1889: 108. Syntypes 9:

India (Mus. Calcutta).

Sphex sulciscuta Gribodo, 1894: 2. Holotype 9: Philip-

pines, Mindoro (MCG!).

Sphex nigripes var. tsingtauensts Strand, 1916: 81A (11):

107. Holotype @: China, Tsingtau [Qingdao] (DEI!!).

Sphex subtruncatus coraxus Van der Vecht, 1957: 365.

Holotype 9: Indonesia, Sumba, Pogobina (NMB!).

Sphex subtruncatus orius Van der Vecht, 1957: 366. Holo-

type 6: Indonesia, Flores, Rana Mese (NMB!).

Sphex subtruncatus xuthus Van der Vecht, 1957: 364.

Holotype g: Indonesia, Sumba, Bondo Kodi (NMB!).

Sphex subtruncatus krombeini Van der Vecht in Krom-

bein, 1984: 28. Holotype 9: Sri Lanka, Kanneliya

Jungle (USNM!).

Distribution. - Sri Lanka and India to China and

Malaysia, Indonesia (Sumatra, Bangka, Java, Kali-

mantan, Lombok, Flores, Sumba, Timor), Philip-

pines.

Note. - S. subtruncatus exists in a number of

geographic forms, which differ in colour (legs black

to reddish, wings dark brown to clear), as well as

in the number of placoids on the male flagellum.

Sphex walshae sp. n.

(figs. 18, 45, 63-65)

Type material. - Holotype 9, Indonesia, S. Sumatra,

250 m, Bengkulu, Muara Tenam, 16-23.vii.1935, M. E.

Walsh (RMNH). — Paratypes: India: Sikkim, Ari to Gna-

tong, 7-2500 ft, iv.1894, Bingham (1 9, USNM). Burma:

Tenasserim, Ihaungyin Valley, v.1891, Bingham (1 9,

BMNH). Malaysia: Malay peninsula, Kedah Peak, 3000-

3500 ft, 15.111.1928, H. M. Pendlebury (1 9, BMNH); Sa-

rawak, C. J. Brooks (1 9, BMNH); Sabah, Sandakan, Baker

(2 9, USNM); Sabah, 19 km N. of Kalabakan, 19.xi.1962,

Y. Hirashima (1 ©, BPBM); North Borneo, Frühstorfer (1

Q, TMB). Indonesia. Sumatra: Anei kloof, 500 m, E. Jac-

obson (3 9, RMNH); Bengkulu, 50 m, 10-18.v.1935, M. E.

Walsh (1 9, RMNH); Bengkulu, Bukit Item, 650 m, 11-

15.vi.1935, M. E. Walsh (1 9, RMNH); Bengkulu, Tandjong

Sakti, 1-10.vi.1935, M. E. Walsh (2 & 1 9, RMNH); Lubuk

Sikaping, 450 m, L. Lundeshagen (1 9, RMNH), Aur Ke-

manis, E. Jacobson (1 9, RMNH); no locality (1 9, Zoolo-

gische Staatssammlung, Miinchen). Kalimantan: Keta-

pan, v.1937, M. E. Walsh (2 9, RMNH); Sampagan, 0 m,

1927, M. E. Walsh (1 4, RMNH); South coast (1 9, RMNH);

Kembang Djangut, 75 m, 24-30.xi.1956, A.M.R. Wegner

(5 9, RMNH); no locality, Muller (1 & 5 9, RMNH). ‘Ind.

or.?’ (1 9, RMNH).

Diagnosis

This species is similar to other large, dark-

winged, dark-legged forms: the representative of S.

subtruncatus in the Philippines and the represen-

tative of S. praedator in the Moluccas (apart from

species belonging in other groups). As these forms

are allopatric, S. walshae may be separated from

sympatric forms on the basis of the dark brown

wings with violaceous effulgence. Unlike S. prae-

dator, S. walshae has a stout curved spine at the

outer end of the female mid tibia, and the male

clypeus has a median apical lobe. In these respects

S. walshae is similar to S. subtruncatus, from which

it differs in having a pygidial plate in the female,

and placoids on third to 7th flagellomere in the

male (variable in S. subtruncatus, the mentioned

condition occurring in specimens from the Philip-

pines and Java).

Description

Female. - Body length 23-28 mm, length of fore-

wing 17-20 mm. Black. Wings dark brown with

violaceous effulgence. Pilosity of head and meso-

soma black; clypeus and frons with sparse silvery

tomentum, mesosoma with sparse, inconspicuous

dark brown tomentum; fore femora ventrally with

sparse erect hairs, no tomentum. Clypeus 0.8 times

as long as IOC, without median carina, apical mar-

gin slightly raised and with a pair of short teeth

medially; rov 0.8 times 10C; labrum with median

carina, which projects apically into a tooth; relative

lengths of flagellomeres I-III 1 : 0.7 : 0.75. Meso-

soma microsculptured: punctulate-reticulate to

shagreened, mesonotum and mesopleuron largely

shallowly punctate, punctures 1-2 diameters apart;

propodeum finely rugose; scutellum with median

longitudinal impression; fore basitarsus with 9-10

rake spines, median spines 1.5 times tarsal width,

apical spine as long as second tarsomere; apex of

mid tibia with stout curved spine. Petiole 0.4 times

as long as hind metatarsus; sixth tergite with pyg-

idium.

Male. - Body length 21-24 mm; length of fore-

wing 16-18 mm. Like the female except for the

following. Clypeus 0.9 times as long as IOC, apical

margin with short, depressed median lobe; 10v 0.85

times 10C; labrum ecarinate; length of flagello-

meres I-III 1: 0.75 : 0.75, third to seventh flagello-

mere with broad, flat placoid. Seventh sternite

weakly emarginate; eighth sternite triangular.

Etymology

Walshae is a noun in genitive case, refering to

Mrs. M. E. Walsh, collector of a substantial part of

the type series.

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Received: 3 November 1990

Accepted: 7 January 1991

P. HUEMER

Tiroler Landesmuseum Ferdinandeum Innsbruck

CARYOCOLUM DELPHINATELLA (CONSTANT) SP. REV,

A SENIOR SYNONYM OF C. FIORII (KLIMESCH)

(LEPIDOPTERA: GELECHIIDAE)

Huemer, P., 1991. Caryocolum delphinatella (Constant) sp. rev., a senior synonym of C.

fioru (Klimesch) (Lepidoptera: Gelechiidae). - Tijdschrift voor Entomologie 134: 31-34,

figs. 1-7. [ISSN 0040-7496]. Published 1 July 1991.

Caryocolum delphinatella (Constant) is recognized as good species and revocated from

synonymy with C. interalbicella (Herrich-Schäffer). The female genitalia are described

and figured and data on biology are given. C. fioriti (Klimesch) is synonymised with C.

delphinatella.

P. Huemer, Tiroler Landesmuseum Ferdinandeum, Museumstr. 15, A-6020 Innsbruck,

Austria.

Key words. - Gelechiidae, Caryocolum, synonymy.

In 1890 the well-known French lepidopterist A.

Constant described Lita delphinatella from a single

female collected in July in the Alpes du Dauphiné.

Klimesch (1954) in his revision of European spe-

cies of the former genus Lita feeding on Caryophyl-

laceae, queried delphinatella as a synonym of Ca-

ryocolum interalbicella (Herrich-Schäffer). This

synonymisation was accepted by the present author

(Huemer 1988) although de/phinatella was treated

as a separate species in the French list (Leraut

1980). C. frorız was described from five males col-

lected in the Abruzzi (Klimesch 1953). Later the

species was also recorded from the Swiss and

French Alps.

When re-valuating the original description of

delphinatella, including the colour plate depicting

the holotype, it became clear that this species is not

a synonym of :interalbicella but conspecific with

fioru. This assumption is further supported by rich

material, including two females, collected in the

south-western Alps in July 1990.

Caryocolum delphinatella (Constant) sp. rev.

Lita delphinatella Constant, 1890a (May 28): 6, pl. 1, fig.

2 [moth in colour]. Holotype 9, France: Alpes du

Dauphiné, vii [not traced].

Lita delphinatella Constant, Meess 1910: 370.

Phthorimaea delphinatella (Constant); Meyrick 1925:

95; Gaede 1937: 254; Lhomme [1946]: 632.

Caryocolum delphinatella (Constant); Leraut 1980: 79.

Gnorimoschema fiorii Klimesch, 1953: 277, figs 23, 24.

Lectotype 8 (Designated by Huemer 1988: 481), Italy:

Abruzzen, Gran Sasso, 25.vii.1935 (Fiori) (coll. Kli-

mesch, Linz) [examined]. Syn. n.

Caryocolum fiorii (Klimesch); Huemer 1988: 480.

Description

Adult (Figs 1, 3-5). - Fore wing length & 7.0-7.5

mm, ® 6.5-7.0 mm. Head white; Labial palpus

white, third segment mottled with dark brown.

Thorax white, tegulae dark brown basally. Fore

wing dark brown, mottled with light brown; white

transverse fasciae at one-fifth and one-half, me-

dially narrowed and often reduced towards costa;

fasciae fused and mottled with light brown dor-

sally; separate white subcostal spot at about two-

fifth always well developed; white costal and tornal

spots at four-quarters variable in extension, separ-

ate or fused to fascia; fringes whitish with dark

brown basal line. Hind wing light grey.

Male genitalia (Huemer 1988: Fig. 122).

Female genitalia (Figs 6, 7). - Apophyses poste-

riores 1.60 mm, apophyses anteriores 0.43 mm;

eighth segment without processes; ostium bursae

surrounded by folds; antrum short, funnel-shaped,

posteriorly fused with pair of short lateral sclero-

tizations of ductus bursae which are level with the

tip of the apophyses anteriores; corpus bursae py-

riform with long, hook-shaped signum.

Biology

The immature stages are still unknown. Accord-

ing to personal observations in France the larvae

probably feed on Minuartia sp. and Silene sp. (Ca-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 1-5. Caryocolum spp. - 1, C. delphinatella, original figure by Constant; 2, C. interalbicella; 3, C. delphinatella

(fiorti lectotype); 4, C. delphinatella 9: 5, C. delphinatella 8.

ryophyllaceae) around which the adults have been

collected in numbers in July and early August; most

specimens were attracted to artificial light sources.

The females examined were both collected during

day time resting on rocks. C. delphinatella is an

alpine species which occurs from about 1600 to

2500 m.

Distribution

Only known from the Abruzzi and the south-

western Alps (Alpes Maritimes, Alpes de Haute

Provence, Hautes Alpes, Isère (Dauphiné), Wal-

liser Alpen, Alpi Cozie); according to Lhomme

[1946] also in the Pyrenées (Haute-Garonne).

Remarks

Despite the efforts of several lepidopterists the

holotype of delphinatella could not be traced in

MNHN Paris or any other institution; however,

Constant’s original description and colour illustra-

tion, the latter reproduced here in monochrome

(Fig. 1), leave no doubt about the identity of this

species. It is well characterized by the pure white

head and thorax as emphasised by Constant, and

the white subcostal forewing spot at two-fifths

(Figs 1,3-5). The synonymisation with Caryocolum

interalbicella is certainly incorrect as this species

has a whitish to cream head and tegulae and a dark

brown thorax (Fig. 2). The few French Caryoco-

lum-species with white colour of head and thorax

differ from delphinatella: C. saginella (Zeller) is

distinctly smaller (4.0-5.0 mm); C. cauligenella

(Schmid) has a creamy rather than pure white head

and thorax and also broader fore wings without the

costal spot present in delphinatella; French spe-

cimens of C. peregrinella (Herrich-Schäffer) are

characterized by the larger size (7.5-9.0 mm),

broader fore wings and extended white markings

of the fore wing without separate subcostal spot. C.

leucothoracellum (Klimesch) has different fore-

wing markings with white subcostal streak and a

medial spot. C. frori perfectly matches delphina-

tella and is here placed as a junior subjective syn-

onym. C. fiorii was recently recorded as new to

France (Huemer 1989) because the true identity of

delphinatella, at that time still treated as synonym

of interalbicella, was not recognized.

The female genitalia of delphinatella are similar

to those of other species of the trauniella-group.

The antrum is distinctly longer than in peregri-

nella (Herrich-Schäffer) and trauniella (Zeller).

The lateral sclerotizations of the ductus bursae are

level with the tip of the apophyses anteriores in

delphinatella whereas in other species of the group

they are at most half that length. The differences

in the male genitalia have been discussed pre-

viously (Huemer 1988).

HUEMER: Caryocolum delphinatella

Figs. 6-7. Caryocolum delphinatella, female genitalia. -

6, eighth segment; 7, signum (X 2 magnification of

eighth segment).

Material examined (additional to that previously re-

corded (Huemer 1988, 1989). — Italy: 4 8, Abruzzen,

Prov. Rieti, Terminillo-Paß, 18-2000 m, 5-6.viii.1970,

leg. Groß; 2 4, P. d. Fargno, ob. Bolognola, Mti. Sibillini,

16-1700 m, 14.viii.1970, leg. Groß (Löbbecke Museum,

Düsseldorf); 1 g, Prov. Torino, Alpi Cozie, Colle delle

Finestre, 2150 m, 27.vii.1990, leg. Huemer & Tarmann.

France: 39 4, 1 9, Dep. Alpes-Maritimes, Marguareis W-

Hang, Navela, 2100-2200 m, 21-23.vii.1990, leg. Huemer

& Tarmann; 22 g, 1 9, Dep. Alpes-de-Haute-Provence,

SW Castel de Restfond, Roche Chevaliére, 2480 m,

25.vii.1990, leg. Huemer & Tarmann; 10 4, Dep. Alpes-

de-Haute-Provence, SW Castel de Restfond, Set. de Caire

Brun N-Flanke, 2420 m, 25.vii.1990, leg. Huemer & Tar-

mann (Tiroler Landesmuseum Ferdinandeum, Inns-

bruck).

ACKNOWLEDGEMENTS

I gratefully acknowledge the help received for

this study from Mr O. Karsholt (ZMUC, Copenha-

gen), Dr S. Lòser (Diisseldorf), Dr G. Luquet

(MNHN, Paris) and Dr K. Sattler (BMNH, London).

Furthermore I am indebted to my colleagues M.

Kahlen, B. Plössl and Dr G. Tarmann (Innsbruck)

for their help in the field and valuable comments.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

REFERENCES

Constant, A., 1890a. Séance du 26 Juin 1889. - Bulletin de

la Societe Entomologique de France 1889: 125.

Constant, A., 1890b. Descriptions de Microlepidopteres

nouveaux ou peu connus. - Annales de la Societe

Entomologique de France 69: 5-16.

Gaede, M., 1937. Gelechiidae. - Lepidopterorum Catalo-

gus 79: 1-630.

Huemer, P, 1988. A taxonomic revision of Caryocolum

(Lepidoptera: Gelechiidae). - Bulletin of the British

Museum (Natural History) (Entomology) 57: 439-

Il:

Huemer, P., 1989. Bemerkenswerte Funde von Caryoco-

lum-Arten aus den Südalpen und dem Mediterraneum

(Lepidoptera, Gelechiidae). - Nachrichtenblatt der

bayerischen Entomologen 38: 37-40.

Klimesch, J., 1953-1954. Die an Caryophyllaceen le-

benden europäischen Gnorimoschema Busck

(=Phthtorimaea Meyr.)-Arten. — Zeitschrift der

Wiener Entomologischen Gesellschaft 38: 225-239,

275-282, 311-319, 39: 273-288, 335-341, 357-362.

Leraut, P., 1980. Liste systématique et synonymique des

Lepidopteres de France, Belgique et Corse. - Supple-

ment à Alexanor et au Bulletin de la Société Entomo-

logique de France, 334 pp.

Lhomme, L., [1946-1948]. Catalogue des Lépidoptères de

France et de Belgique 2: 489-648. - Douelle (Lot).

Meess, A., 1910. xxxviii. Familie. Gelechiidae. - In: Hof-

mann, E. & A. Spuler (eds). Die Schmetterlinge Eu-

ropas. 2, 523 pp. - Stuttgart.

Meyrick, E., 1925. Lepidoptera Heterocera. Fam. Gelechi-

adae. - Genera Insectorum 184: 1-290, pls 1-5.

Received: 23 November 1990

Accepted: 26 November 1990

I. LANSBURY

Oxford University Museum

CUTICULAR BLADES AND OTHER STRUCTURES

OF DIAPREPOCORIS KIRKALDY AND

STENOCORIXA HORVATH

(HETEROPTERA: CORIXIDAE)

Lansbury, I., 1991. Cuticular blades and other structures of Diaprepocoris Kirkaldy and

Stenocorixa Horvath (Heteroptera: Corixidae). - Tijdschrift voor Entomologie 134: 35-

46, figs. 1-27. [ISSN 0040-7496]. Published 1 July 1991.

Diaprepocoris Kirkaldy and Stenocorixa Horvath possess cuticular blades and modified

swimming hairs on their hind tibiae and tarsi which resemble those of gyrinids (Coleo-

ptera: Gyrinidae). These have not previously been recorded in the Corixidae. Unusual

macrotrichia are also present on the clavus and corium of all known species of both genera.

The unique characters of these two genera support their separate subfamily placements.

I. Lansbury, Hope Entomological Collections, University Museum, Oxford, OX1 3 PW,

UK.

Keywords. - Australia, Africa, Cuticular blades, modified swimming hairs, respiration,

Corixidae.

A study of the Australasian Diaprepocoris has

revealed the presence of cuticular blades and modi-

fied swimming hairs on the hind tibiae and tarsi.

These structures have not previously been reported

in the Corixidae. A survey of the corixid genera has

shown that a similar combination of cuticular

blades and modified swimming hairs are also pres-

ent in the genus Stenocorixa which is restricted to

tropical Africa. The cuticular blades and associated

specialised swimming hairs are described and their

probable function discussed in relation to the nek-

tonic (submerged) swimming strategies and respi-

ration in both subfamilies.

The double hair pile layer on the forewings of

Diaprepocoris is figured and a dual function sug-

gested for the hair pile. Diaprepocoris is compared

with Stenocorixa, the later apparently having

smooth forewings. The relationship between the

gas stores of Diaprepocoris and the specialised

swimming hairs of the hind legs are discussed. No

data are at present available for Stenocorixa. The

taxonomic position (isolation) of the two subfam-

ilies is briefly discussed.

Diaprepocorinae Lundblad, 1928

Single extant genus Diaprepocoris Kirkaldy,

1897, type species D. barycephala Kirkaldy, 1897.

Within the Diaprepocorinae, Popov (1971) in-

cludes Gazimuria scutellata Popov, from the lower

Jurassic, Akutaev Series, Chitin Province, East

Transbaikals and Karataviella brachyptera Becker-

Migdisova from upper Jurassic, Chayan Region,

Chimkent Province, South Kazakhstan. Walton

(1940) split the Diaprepocorinae into two tribes

using the structure of the female pala:

Diaprepocorini. - Female pala digitform, few

sieve hairs (about 10) and short. Type/Genus spe-

cies Diaprepocoris barycephala.

Corixanectini. - Female pala short and broad,

inner side flattened and fringed with numerous

long sieve hairs. Type/Genus species Corixanecta

zealandiae Hale.

Stenocorixinae Hungerford, 1948

Single extant genus, Stenocorixa Horvath, 1927,

type species S. protrusa Horvath, 1927, monobasic.

Jaczewski (1928) placed Stenocorixa in the

Corixinae; Poisson & Jaczewski (1928) discussing

the systematic position of Stenocorixa concluded

that the poorly developed abdominal asymmetry,

shape of the pala claw and other features suggested

a relationship with Cymatia Flor. They ended by

stating ‘The generic distinctness of Stenocorixa

Horv. is beyond any doubt and even as a separate

genus it seems to stand rather apart of the other

genera of the subfamily Corixinae’.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

SN:

Figs. 1-4. Diaprepocoris barycephala Kirkaldy. - 1, corium, double hair pile, note collapsed microtrichia and longi-

tudinal ridges on macrotrichia X3200; 2, inner lateral margin of clavus, note ‘shephards crook' macrotrichia X400;

3, middle of corium X800; 4, membrane of elytra, note more erect microtrichia and enlarged socket of macrotrichia

X6400. Reduction 84%.

DESCRIPTIVE PART

Diaprepocoris, structure trichia

Macrotrichia of the clavus and corium varying in

shape. The hairs along the inner margin of the

corium are apically curled over forming a ‘shep-

herds crook’ (fig. 2) and are longer and thinner

than those on the inner areas adjacent to the clavus

(fig. 3). The microtrichia are densely packed and

resemble a plastron-like layer however, as they

have collapsed and become matted, they clearly do

not function as a plastron. The distribution of the

microtrichia is fairly uniform over the clavus and

corium. The membrane differs in that the micro-

trichia do not seem to have collapsed as much as

those on the clavus and corium (fig. 1 clavus; fig.

4 membrane). The shape of the sockets within the

microtrichia from which the macrotrichia arise,’

structurally suggests that the latter can be elevated

and depressed (fig. 1). The macrotrichia are rather

more slender and fewer in number on the mem-

brane. Those on the elytra are longitudinally ridged

(fig. 1). The division between the double hair layer

and the edge of the forewing is distinct (fig. 5). The

lateral margin at the nodal furrow shows a scale-

like appearance with no hair layers (fig. 6).

Stenocorixa, structure trichia

Unlike nearly all other genera of corixids, the

pronotum and hemelytra are not rastrate, but

smooth and shining with scattered long strap-like

hairs arising from angled depressions, each with a

complete basal ring (fig. 13). Caudad of each de-

pression, an elongate channel within the clavus-

corium, the strap-like hairs are almost certainly

lodged in the channel(s) when the corixed is swim-

ming.

Diaprepocoris, cuticular blades and swimming

hairs

The hind tarsi of Diaprepocoris have in addition

to the rows of swimming hairs, fringes of cuticular

blades as long or longer than their respective tarsal

segments. When not in use, they overlie each other

like a venetian blind (fig. 7). Fanned out they in-

crease the surface area of the tibiae-tarsi. The

spread of the blades increases the surface area of

the Ist tarsus along the leading edge by 3.5 X 10

(this and following figures approximate) - trailing

edge has a greater density of fine swimming hairs

overlying a row of blades increasing the surface

area by 10 X 2. The leading edge of 2nd tarsus has

a row of blades decreasing in length distally, in-

creasing surface area by 3.0 x 10 — trailing edge has

LANSBURY: Cuticular blades in Corixidae

fewer swimming hairs, the blades are roughly all

the same length, increasing surfaces area by 2.0

10. The cuticular blades thus greatly increase the

surface area (figs. 17, 18). According to Nachtigall

(1974) cuticular blades create about 90% thrust of

that of a solid object of equal size in the Gyrinidae

(Coleoptera).

The blades of Diaprepocoris appear to have a

function similar to those of gyrinids, the latter also

use their middle legs when swimming. The hind

coxae of Diaprepocoris conform to the usual co-

rixid form. The ventral surface of the coxae is

flattened to accommodate the trochanter-femur in

the full recovery position, the femur forming a 33°

angle with the midline of the body. The flattened

area is partially enclosed by the tip of the meta-

xyphus and prevents excessive rotation of the hind

leg (fig. 19). Schenke (1965, 1966) gives an account

of a corixid swimming using Corixa punctata Illiger

as a model. There are striking differences between

the two genera. Corixa has a complex arrangement

of rows of hairs, each with an incomplete basal

ring, this enables the hairs to be folded flat against

the leg during the recovery stroke. The tarsal hairs

are supported by a row or rows of stout flexible

spines which function in a similar fashion. There

are few hind tarsal spines on Diaprepocoris com-

pared with Corixa; D. barycephala \st tarsus has

between 10-13 spines in each of three lateral rows,

one either side of the trailing edge and one along

the leading edge. All the spines are semi-erect and

probably rotate in much the same way as Corixa.

The fringes of swimming hairs are arranged in a

distinct manner. According to Schenke (1965)

there are in excess of 5000 hairs on each rowing leg

of Corixa. In Diaprepocoris, the leading edge of the

lst tarsus has about 30 long hairs, each having an

incomplete basal ring (fig. 8) and a larger number

along the trailing edge. Second tarsus with 8 long

hairs along the leading edge and about 30 along the

trailing edge. Figure 9 shows the incomplete basal

rings of the cuticular blades in the recovery posi-

tion, fig. 10 shows the folding of the blades on the

2nd tarsus partially overlapping the apical claws

and extreme length of some of the blades. The hind

tibiae moderately spinose (fig. 23) interspersed

with hairs, some are expanded distally with a series

of fine incisions (fig. 11). These modified hairs may

reduce drag and loading on the hind legs, another

function may be to move expired gas backwards

over the elytra prior to the corixid surfacing to

renew its gas store. The distribution of the explan-

ate hind tibial hairs and cuticular blades of the tarsi

are not uniform throughout the genus, D. baryce-

phala (fig. 17), D. personata Hale (fig. 18) and D.

pedderensis Knowles (figs. 20, 21). There is also

variation in the development of the swimming

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 5-8. Diaprepocoris barycephala Kirkaldy. - 5, outer lateral margins of corium and edge of double hair pile layer

X3200; 6, lateral margin of forewing at nodal furrow with scale-like surface X800; 7, overlapping cuticular blades on

hind tarsus X200; 8, hind tarsi, note incomplete basal rings X400. Reduction 84%.

LANSBURY: Cuticular blades in Corixidae

Figs. 9-12. Diaprepocoris barycephala Kirkaldy. - 9, base of cuticular blades showing the incomplete basal rings X800;

10, cuticular blades on hind tarsi, note extreme length of blades X50; 11, incised hairs on the hind tibia X800; 12,

immature Diaprepocoris, hind tarsus X50. Reduction 84%.

39)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

14,

ind tarsi, note length and density of blades X50; 16,

’

ith single strap-like hair, note channel in elytra X3200

Figs. 13-16. Stenocorixa protrusa Horvath. - 13, elytra w

explanate hind tarsi with cuticular blades X24; 15, explanate h

hind tarsi X400. Reduction 84%.

d hairs on

INCISE

hairs on the middle legs; in pedderensis none are

visible on a slide mounted leg (fig. 22), personata

has about 20 hairs visible on the tibia and

barycephala has about 50 hairs on the tibia and two

rows of about 60 hairs on the tarsus.

Stenocorixa, cuticular blades and swimming

hairs

The hind tarsi of Stenocorixa (fig. 14) are the

most explanate of all the corixids, they most closely

resemble those of Lethocerus (Belostomatidae, He-

teroptera). The cuticular blades are extremely long

forming a dense matted fringe (fig. 15). It has not

been possible to determine from available prepa-

rations if each of the blades have an incomplete

basal ring q.v. Diaprepocoris (fig. 9). The long

swimming hairs fringing the rows of blades each

appear to have incomplete basal rings. The trailing

edge of the Ist tarsus has about 10 groups of stout

semi-erect spines, between each group, a single

thicker spine. The rows partially enclose several

rows of long fan-shaped hairs which are deeply

incised distally (fig. 16), it is likely that these hairs

have a similar function to those of Diaprepocorts.

The fine structure of the clavus and corium of

Stenocorixa does not suggest that it has a large

supra-alar gas store, its overall streamlined appear-

ance is totally different from that of Diaprepocoris.

The complete basal rings would prevent the hairs

laying flat against the forewings and would pre-

sumably cause minor eddies over the dorsum of the

swimming corixid. The hairs are too few in number

to act as a hydrofuge layer and may have some

sensory function or be used for manouvering nek-

tonically.

There is a rather greater degree of flexibility

between the hind tibiae-tarsi than in Dia-

prepocoris. The hind tibiae of Stenocorixa have a

pair of prominent projections distally. The acu-

minate insertion of the tarsus within the tibia sug-

gests that there is considerable flexibility which

would increase the corixids manouverability. The

2nd tarsus is capable of partial folding as in the

Gyrinidae, this would be advantageous in the re-

covery stroke reducing drag. The hind legs of

Stenocorixa are pro-rata much longer than those of

Diaprepocoris.

Diaprepocoris: hind tibiae not explanate and not

reaching the end of the abdomen.

Stenocorixa: hind femur-tibia-tarsus explanate,

tibia reaching the end of the abdomen.

The power stroke of Stenocorixa may exert a

greater propulsive effort, the folding hind tarsi

reducing drag during the recovery stroke. The rel-

ative shortness of the hind legs of Diaprepocoris

with possible reduction of energy in the power

LANSBURY: Cuticular blades in Corixidae

stroke may be compensated for by the supra-alar

gas store, this would enable the corixid to stay

submerged for longer periods, thus avoiding pre-

dators. The extra bouyancy would however, also

mean that the corixid would need to keep up a high

level of power strokes to maintain depth and atti-

tude when submerged and not resting on sub-

merged objects.

DISCUSSION

Thorpe & Crisp (1949) group aquatic animals

with hydrofuge hairs in four categories, placing

Corixa (Corixinae) in group III, members of which

must surface at intervals to renew their gas stores.

They are bouyant rather than heavier than water

and often have a double hair pile.

Diaprepocoris according to Parsons (1976) has

a much larger exposed dorsal air store covering the

scutellum and forewing; the Micronectinae Jac-

zewski have the supra-alar store restricted to the

anterior part of the embolium and the anterior-

most part of the claval suture; the Corixinae Jac-

zewski have a slightly larger air store extending the

length of the embolium, anterior part of the clavus

suture and that part of the clavus covered by the

pronotum. The mesonotal air store of the Micro-

nectinae and Corixinae covered by the pronotum.

As a form of air bubble respiration, Diaprepocoris

has a much larger gas store than most other corix-

ids, in addition to the supra-alar store, it has sub-

alar gas stores beneath the forewings and over

much of the venter.

The presence of large supra-alar air stores and

sub-alar stores may increase the bouyancy of

Diaprepocoris, the bouyancy levels possibly being

related to the density of swimming hairs and cutic-

ular blades on the hind legs. Possible secondary

functions of the external gas layer may be either as

a flexible skin which would enable the corixid to

swim with less friction. With the exception of D.

pedderensis, all Diaprepocoris species are rela-

tively short and squat, their greatest width being

just over half total length. An alternative function

may be acoustic; Theiss (1982) has shown that in

Corixa, the cervical air store i.e. air stores formed

by the posterior and lateral parts of the head being

flattened forming flanges which overlap the pro-

thorax, are used in the generation and radiation of

sound, thus the dual function of air stores has been

demonstrated.

Taxonomy

Diaprepocoris and Stenocorixa are the most dis-

tinctive genera of the extant Corixidae. Characters

common to both subfamilies are:

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

—

Figs. 17-18. Diaprepocoris hind tarsi. - 17, D. barycephala Kirkaldy, scale line 1 mm, 18, D. personata Hale, scale

line 0.25 mm.

- Hind tarsi fringed with cuticular blades

- Little difference between the male and female

palae

The Diaprepocorinae share a set of characters

which distinguishes the subfamily from all others:

1. Ocelli on vertex between compounds eyes (fig.

24)

Fore tibia-tarsus fused (fig. 25)

3. Tarsal claws of hind legs apical (figs. 17, 18 and

21)

Muscle attachment to male genital capsule dorsal

Female ovipositor with a recognisable blade-

like structure

The Stenocorixinae share a limited set of char-

acters which distinguish the subfamily:

1. Clavus and corium smooth without microtri-

chia (fig. 13)

2. Clavus and corium with scattered long strap-

like hairs

3. Abdominal asymmetry of male 6-7th tergites

hardly differing from female

4. Male left paramere absent

Various proposals have been made regarding the

validity of the corixid subfamilies. Leston (1955)

partially summarised the proposals of China (1943)

and Hungerford (1948) and observed that most

authors treat the Corixidae as a monophyletic

group. Popov (1971) altered this concept to include

the Shurabellidae (Shurabella lepyronopsis Becker-

Migdisova, Liassic fossil). The Corixoidea should be

considered a holophyletic group sensu Ashlock

STAND:

Fig. 19. Diaprepocoris personata Hale: metaxyphus and

ancilliary structures. Abbreviations; e, eye; 11p, lateral

lobe of prothorax; 0, osteole of scent gland; P3, insertion

of 3rd coxa; M, metaxyphus, scale line 1 mm.

Hungerford (1948) split the Corixidae into six

subfamilies:

1. Groups with large exposed scutellum ..........

ee Diaprepocorinae and Micronectinae

- Groups with scutellum hidden ................ 2

2. Hemielytral groove absent ..... Stenocorixinae

= Hemielycralicroovespresentarre ee ee eee 3

3. Rostrum without transverse sulcations or nodal

FUECO Ww ACEP PESI AP TEN Cymatiinae

— Rostrum with transverse sulcations and nodal

furrow ......... Corixinae and Heterocorixinae

LANSBURY: Cuticular blades in Corixidae

Figs. 20-23. Diaprepocoris pedderensis Knowles. - 20, 21, hind tarsi, scale line 1 mm; 22, middle leg, scale line

mm; 23, hind femur and tibia, scale line 0.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Popov (1986) recognised three extant subfami-

lies viz. Corixinae, Diaprepocorinae and Micronec-

tinae. Popov (1971) speculating on the affinities of

Stenocorixa noted that the male genital capsule

differed from all other corixids and that further

study of the aberrant groups (Stenocorixa and

Heterocorixa etc.) would be necesssary before the

status of these groups was fully understood.

Nachtigall (1974) comments ‘The rowing legs of

recent water dwellers are considerably more flat-

tened than those of fossil water dwellers or the legs

of terrestrial relatives’. Within the Corixidae, both

extant and fossil, there is little apparent distinc-

tion. The fossil genus Gazimuria looks like a form

of Diaprepocoris, Stenocorixa has explanate hind

legs, Corixa moderately so, but there does not seem

to be a steady progression throughout the corixid

genera. As no corixids are remotely terrestrial,

Nachtigal’s observations do not seem to be appli-

cable.

Immature stages

Diaprepocoris barycephala 4th-5th instar: The

front leg (fig. 25) closely resembles the adult. As in

all immature corixids the front tibia-tarsus are

fused, this character persists in the adult

Diaprepocoris. The middle leg also closely resem-

bles the adult form, but lacks the swimming hairs

on the tibia-tarsus (fig. 26). The hind leg similar to

the adult with prominent cuticular blades (figs. 12

& 27). The hind tarsus is one-segmented as in all

known immature corixids.

Immature stages of Stenocorixa were not avail-

able for study.

ACKNOWLEDGEMENTS

The Leverhulme Foundation, London; CSIRO

Division of Entomology, Canberra and the Austral-

ian Biological Resource Study, Canberra are

thanked for grants in aid towards a field-trip in

1979. Dr. W. D. Williams, Adelaide University, Dr.

I. A. E. Bayly and Dr. P. S. Lake, Monash University

sent me Diaprepocoris specimens. The SEM fig-

ures were made by Mrs Barbara Luke, Zoology

Department, Oxford University, courtesy of Pro-

fessor D. S. Smith (Hope Professor).

REFERENCES

Ashlock, P. D., 1971. Monophyly and associated terms. —

Systematic Zoology 20: 63-69.

China, W. E., 1943. Some remarks on Walton's classifica-

tion of the British Corixidae (Hemiptera). - Entomol-

ogist's Monthly Magazine 79: 109-111.

Horvath, G., 1927. Zoological results of the Swedish

Expedition to Central Africa 1921, Insecta 25 (aquatic

LE.

===

III:

/ Ht

Dil

N Hal M

SF Fe

——

ZEEE

PI)

un Aly

_ —

Fig. 24. Diaprepocoris pedderensis Knowles, male dorsal

habitus, scale line 1 mm.

and semi-aquatic Hemiptera). - Arkiv för Zoologi

18A (31): 1-4.

Hungerford, H. B., 1948. The Corixidae of the Western

Hemisphere (Hemiptera). — Kansas University

Science Bulletin 32 (25): 1-827.

Jaczewski, T., 1928. Bemerkungen tiber die geographische

Verbreitung der Corixiden. - Annales Musei Zoologici

Polonici 7: 45-67.

Kirkaldy, G. W., 1897. Aquatic Rhynchota: Descriptions

and notes. No. 1. - Annals Magazine Natural History

(6) 20: 52-60.

LANSBURY: Cuticular blades in Corixidae

ER GZ es I

/ AS

i Mil! 7, ene

A = — a — / \

| = =

Il Sas

\ SS m NT

À =

INN) TERN Ä | a /

4%,

Figs. 25-27. Diaprepocoris barycephala Kirkaldy, immature. - 25, front leg, scale line 0.5 mm; 26, middle leg; 27, hind

leg, scale line 0.5 mm.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Leston, D., 1955. Taxonomy of the British Corixidae

(Hem.). - Entomologist's Monthly Magazine 90: 57-

59.

Lundblad, O., 1928. Studien iber schwedische Corixiden

VII. - Entomologisk Tidskrift 49: 9-16, pls 1-3.

Nachtigall, W., 1974. Locomotion: Mechanics and hydro-

dynamics of swimming in aquatic insects. - In: The

Physiology of Insecta, 2nd edition (edited M. Rock-

stein): 382-432, 3, Academic Press New York.

Parsons, M. C., 1976. Respiratory significance of the tho-

racic and abdominal morphology of three Corixidae,

Diaprepocoris, Micronecta and Hesperocorixa

Heteroptera: Hydrocorisae. - Psyche, Journal of En-

tomology, Camb. Mass. 83: 132-179.

Poisson, R. & Jaczewski, T., 1928. Additional notes on the

morphology of Stenocorixa protrusa Horvath

(Heteroptera, Corixidae). - Annales Musei Zoologici

Polonici 7: 115-119.

Popov, Y. A., 1971. Historical development of Hemiptera

Infraorder Nepomorpha (Heteroptera). - Trudy Pa-

leontologicheskogo Instituta Akademiya Nauk SSSR,

129: 1-228 (in Russian).

Popov, Y. A., 1986. Insects in the early Cretaceous eco-

systems of West Mongolia. - Transactions Joint Soviet

Mongolian Palaeontological Expedition 28: 50-83 (in

Russian).

Schenke, G., 1965. Die Ruderbewegungen bei Corixa

punctata Illig. (Cryptocerata). - Internationale Revue

gesamten Hydrobiologie und Hydrographie 50: 73-

84.

Schenke, G., 1966. Schwimmgeschwindigkeit, Schlagfre-

quenze und Stevern bei Corixa punctata Illig. - Zoo-

logischer Anzeiger 176: 5-12.

Theiss, J., 1982. Generation and radiation of sound by

stridulating water insects as exemplified by the corix-

ids. - Behavioral Ecology and Sociobiology 10: 225-

235.

Thorpe, W. H. & Crisp, D. J., 1949. Studies on Plastron

Respiration IV. Plastron Respiration in the Coleo-

ptera. - Journal Experimental Biology 26: 219-260.

Walton, G. A. in Hutchinson, G. E., 1940. A revision of

the Corixidae of India and adjacent regions. - Transac-

tions Connecticut Academy of Arts & Science 33: 339-

476.

Received: 20 March 1991

Accepted: 22 April 1991

NICO NIESER! & PINGPING CHEN?

‘Tiel, The Netherlands & “Organizing Committee XIXth International Congress

Entomology, Beijing, P. R. China

NAUCORIDAE, NEPIDAE AND NOTONECTIDAE,

MAINLY FROM SULAWESI AND PULAU BUTON

(INDONESIA)

Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), I.

Nieser, N. & P. Chen, 1991. Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi

and Pulau Buton (Indonesia). Notes on Malesian aquatic and semiaquatic bugs (Heter-

optera), I. - Tijdschrift voor Entomologie 134: 47-67, figs. 1-21. [ISSN 0040-7496].

Published 1 July 1991.

Apart from new records for many species, mainly for Sulawesi, P. Buton and Sabah (N.

Borneo), eight new species are described. Naucoridae: Aphelocheirus breviculus sp. n.

(Sabah), A. robustus sp. n. (Sulawesi), Coptocatus stereos sp. n. (Sabah); Nepidae:

Ranatra sulawesti sp. n. (Sulawesi); Notonectidae: Enithares caesaries sp. n., E. lansburyi

sp. n., E. phenakismos sp. n. (all from Sulawesi) and E. skutalis sp. n. (P. Buton).

Correspondence: Dr. N. Nieser, Htg. Eduardstraat 16, 4001 RG Tiel, The Netherlands.

Key words. - Sulawesi; Buton; Sabah; Nepomorpha; keys; new species.

During the last decades the study of Nepomor-

pha and Gerromorpha of Indonesia and Malaysia

is making some progress (see e.g. Lansbury 1964-

1985, La Rivers 1970, Polhemus 1986, Polhemus &

Polhemus 1988). New expeditions to formerly un-

explored areas and even rather occasional trips to

already explored regions (see e.g. Nieser & Chen

1991), however, are still adding new species and

new distributional data to our knowledge. As in

many other groups, a high proportion of the spe-

cies in this region seems to have limited ranges. In

water bugs these restricted ranges are usually ap-

parent in stream inhabiting species, whereas many

of the widespread species occur in or on ponds in

cultivated areas.

The species dealt with in this paper were mainly

collected by J. P. Duffels, J. van Tol and N. Nieser

on several expeditions to the eastern part of Su-

lawesi, the nearby island of Buton and Borneo.

Keys to the Malesian genera are included. Keys

to species of Sulawesi and Pulau Buton are given

for most genera, but they should be used with

extreme caution as the fauna of the region is still

poorly known.

Measurements are in millimeters and present

the range or the (arithmetic) mean based on five

randomly chosen specimens, or in case there are

less than five, on all specimens available. The

width of the head is measured across the eyes.

MATERIAL

A list of collecting localities of N. Nieser is

presented in appendix 1. Sampling stations of J. P.

Duffels and J. van Tol are partly connected with

‘Project Wallace’ and with the ‘1989 RMNH Expe-

dition to Sulawesi’. Data on the latter expedition

will extensively described in a separate paper (Van

Tol et al. in prep.). All other sampling stations of

both collectors, including those on Borneo, are

summarized in appendix 2.

In addition to these recent collections, miscel-

laneous Malesian material in the RMNH collection

has been included. Specimens collected by J. P. Duf-

fels are deposited in ZMA, those by J. van Tol in

RMNH, and those by N. Nieser in his own collection,

unless stated otherwise. Deposition of material is

specified only with rare or new species.

The areas in Sulawesi used with the localities

agree with the Indonesian provinces, viz. Sulawesi

Utara for Northern Sulawesi (Celebes), Sulawesi

Tengah for Central Sulawesi and Sulawesi Teng-

gara for Southeastern Sulawesi. Pulau Buton is

mentioned separately although it belongs adminis-

tratively to Sulawesi Tenggara (see Whitten et al.

1988). Localities are in Indonesia, unless stated

otherwise.

Collections from which material has been stu-

died (with abbreviations in brackets): Museum

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Zoologi Bogor, Bogor, Jawa (MBBJ); Bagian Pertan-

ian, Universitas Haluoleo, Kendari, Sulawesi

(BPUH); J. T. Polhemus collection (University of

Colorado Museum), Englewood, Co., U. S. A.

(yrPC); Snow Entomological Museum, University

of Kansas, Lawrence, Kansas, U. S. A. (SEMC); N.

Nieser collection, Tiel, The Netherlands (NC); Ox-

ford University Museum, Oxford, United King-

dom (OXUM); National Museum of Natural His-

tory (Rijksmuseum van Natuurlijke Historie),

Leiden, The Netherlands (RMNH); Zoölogisch Mu-

seum, Afdeling Entomologie, Amsterdam, The

Netherlands (ZMA).

ACKNOWLEDGEMENTS

Thanks are due to Dr J. P. Duffels (ZMA) and Mr.

J. van Tol (RMNH) for putting specimens in their

care to our disposal; to Dr I. Lansbury (OxUM) for

discussing some Notonectidae and to Dr I Made

Ebeneser (BPUH) for showing the senior author

some interesting localities.

SYSTEMATIC PART

NAUCORIDAE

Key to Indonesian and East Malaysian genera

1. Rostrum slender, longer than fore femora,

reaching to middle coxae (Aphelocheirinae)

AASTRA NET Aphelocheirus Westwood

— Rostrum stout, very broad at base, much shor-

ter than fore femora (Naucorinae) .......... 2

2. Rostrum inserted at anterior margin of head,

labrum distinct and well developed ......... 3

- Rostrum inserted in a deep excavation, dis-

tinctly posterior to the apical margin of head,

labrum often greatly reduced (Cheirochelini)

3. Fore tarsi one-segmented, with one single very

small claw (Naucorini) Naucoris Fabricius

- Fore tarsi two-segmented, with two claws,

which are often very inconspicuous (Laccoco-

Tin) MINAS WOR TO NET MAENE de 4

4. Labrum more or less equilaterally triangular

with antacuteltiprs sas eee Laccocoris Stal

- Labrum with sides shorter than base and tip

broadly rounded ............... Heleocoris Stàl

5. Head ventrally with well developed grooves in

which litthe antennaes ER PURE REA

PROTO 07 L'EAU Coptocatus Montandon

= Antennal oroovesfabsentan. MN 6

6. Ventral margin of prothorax with a conspicu-

ous pit covered with amembrane or specialized

pilosity (static sense organ), body elongate,

connexival angles blunt, not spinose, conse-

quently lateral edge of abdomen more or less

CONTINUOUSEN RE CT 7

- Static sense organ on ventral margin of pro-

thorax absent, body oval, connexival angles

sharp, usually spinose, consequently lateral

edge of abdomen distinctly serrate .......... 9)

7. Static sense organ slightly anterior to midway

of lateral margin of prothorax, ventral keel of

head strongly developed with an anterior

spine-like tip, which is at least as long as first

rostral segment ......... Idiocarus Montandon

- Static sense organ near or at antero-lateral an-

gle of prothorax, ventral keel of head normal,

without a prominent anterior spine ......... 8

8. Static sense organ rounded, placed at extreme

antero-lateralangle of prothorax, which is cup-

shaped; anteclypeus short, broadly and evenly

LOUNGEC A Sete Nesocricos La Rivers

— Static sense organ elongate, placed half to

three-quarters of its length caudad to antero-

lateral angles of prothorax; anteclypeus quite

long with sinuate margins producing a medio-

ANLELIOLIODER EP EEE Tanycricos La Rivers

9. Mesotibia distally with five or six decreasingly

sized, transverse rows of dense, short spines

en RN ne Cavocoris La Rivers

- Mesotibia with at most two of such rows of

Spines 10.121092. ADN BORO AGREE ACER SET AUS 10

10. Anteclypeus prolonged, overhanging part be-

ing half to three-quarters the length of labral

baser dE AEEA EM Warisia La Rivers

— Anteclypeus short, overhang at most one

third of the length of labral base ........... JE

11. Abdominal venter showing only four segments

ERICE. WI ANA IA, Quadricoris La Rivers

- Abdominal venter showing the usual five seg-

mentst SINE FE E ROTA RR, 12

12. Larger species, length over 7 mm; abdomen

asymmetrical ........... Sagocoris Montandon

- Smaller species, length slightly less than 7 mm,

AbdoMENisyimmetrica MORE REI ROLL

San Aptinocoris Montandon

Remarks. - Except for Sagocoris, which has also

one representative in the Philippines, the last

mentioned eight genera are restricted to New Gui-

nea.

In the species of Naucoridae described below, the

first abdominal segment is not visible. The

numbers used are the morphological homologous

numbers, thus ‘tergites 2-8° are visible as tergites

1-7.

Aphelocheirus Westwood, 1833

Southeast Asiatic species of the genus were re-

cently revised by Polhemus & Polhemus (1988).

About 18 species are known from Malesia, includ-

ing the Philippines, but more species are to be

discovered. Species of Aphelochetrus are easily

overlooked by the casual collector as most burrow

in the bottom of streams. Thanks to their plastron

respiration they do not need to surface to refill their

air store.

Key to Aphelocheirus of Sulawesi

1. Lateroposterior spines of connexiva 4-5 short

and broad; body length of male 8.9-9.1 mm, of

female 8.5-8.8 mm (Sulawesi Selatan) .........

MERE Te Rue. à A. celebensis Polh. & Polh.

— Lateroposterior spines of connexiva 4-5 rather

long and acute; body length of male c. 10 mm,

oftfemalesmoresthan 92mm. rer een. 2

. 2. Male: Left paramere with a broad hooked apex

(fig. 5). Female: Genital operculum short and

broad with convex caudo-lateral angles (fig. 3)

(Sulawesi Tenggara) ...... A. robustus sp. n.

— Male: Left paramere with a slender curved ap-

ical part. Female: Genital operculum rather

long and slender with apical lateral margins

very slightly concave (Sulawesi Tengah) ......

LIA o EE ALU GF A. lorelindu Polh. & Polh.

Aphelocheirus lorelindu Polhemus & Polhemus

Aphelocheirus lorelindu Polhemus & Polhemus, 1988:

190-192, figs. 18-21, 164.

Material. - Sulawesi Tengah: Palu, 50 km SE of: Lore

Lindu NP, Sopu river near Dongi Dongi, 950 m, SJ 86, 8

Dec 1985, J. van Tol, 3 9, (RMNH).

Distribution. - Known only from the Lore Lindu

N. P. in Central Sulawesi, and surrounding areas.

Aphelocheirus breviculus sp. n.

(figs. 1-2)

Type material. - Holotype 9: Malaysia: S. Sabah: 105

km S of Beaufort, Long Pasia area, Sungai Ritan, 4°24'’N

115° 42’E, 1160 m, undisturbed evergreen tropical rain-

forest, 9.IV.1987, J. van Tol & J. Huisman (RMNH). -

Paratypes: 2 ® with same data (RMNH, NC).

Description

Brachypterous female. - Rather small species,

form elongate oval, length 7.7-8.1 mm, width of

head 1.8-1.9 mm, maximum width across abdomen

4.9-5.1 mm.

Colour greyish brown, head on dorsal side yellow

to light brown, posteriorly darker, with black eyes;

pronotum, hemielytra and tergites greyish to

brown, margins and scutellum paler; legs and ven-

NIESER & CHEN: Malesian Heteroptera

ter yellowish to light brown, central part of tho-

racic and abdominal sternites greyish, pruinose,

with sparse golden pubescence.

Head. Shining, coarsely punctate, produced

ahead of eyes for distance equal to 0.5 the length

of an eye; eyes twice as long as wide, outer margins

sinuate, due to weakly developed antero-lateral

flange; anterior / posterior interoculus = 1.3/1.0

mm.

Thorax. Pronotum shining, roughly punctate,

antero-medially rugose; very sparsely set with in-

conspicuous short fine pale setae, width / length

3.5; lateral margins bearing about twelve stout

minute erect setae. Scutellum shining, rugose,

width / length 2.5, lateral margin hardly sinuate,

shallow transverse sulcus present along anterior

margin (to fit the posterior margin of pronotum).

Hemielytra not touching medially, leaving an area

0.4 mm wide of metanotum uncovered; posteriorly

not reaching the hind margin of abdominal tergite

2, surface punctate-rugose, claval area shining, dull

in remainder, clavus not delineated; embolar mar-

gin evenly curving anteriorly, its posterior angle

strongly produced laterally (fig. 1).

Abdomen. Dull, rugose with narrow shining

margins which bear 5-7 minute erect setae in small

indentations, posterior indentation of each seg-

ment bearing two or three of these minute setae

and a few longer, easily detachable, setae. Tergites

2-8 exposed, bearing very few short thin pale re-

cumbent setae, paired glandular openings present,

but indistinct on posterior margin of tergite 3.

Posterolateral angles of tergites 3-7 bluntly and

broadly spinose, more distinctly produced poste-

riorly.

Ventral side. Length of antennal segments 1-4 is

0.09, 0.23, 0.23 and 0.42 mm. Labrum shining,

anterior margin evenly rounded, rostrum shining,

length 3.2-3.4 mm, reaching middle trochanters.

Prosternum with weak median carina, propleura

with inner projections notched; mesosternum

pruinose with a median carina, tumescent poste-

romedially; metasternum pruinose, metaxiphus

small, apically pointed. Abdomen pruinose, ster-

nites 3 and 4 bearing rounded projections medially

on posterior margins, posterior margins of ster-

nites 4 and 5 lacking rearwardly directed stout setae

medially; genital operculum with posterolateral

angles developed, resulting in a sinuate posterior

margin; narrow tufts of long setae at lateral mar-

gins and intermedially in apical third (fig. 2).

Legs. Set with fine gold setae, fore femur, tibia

and tarsi with thick hair pads on inner surface.

Fore, middle and hind coxae with combs of long

light brown setae on interior margins. Middle leg,

trochanter, femur, tibia and tarsi with thick yellow

hair pads on ventral faces; femur sparsely set with

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 1-5. Aphelocheirus

species. - 1, Apex of em-

bolium of A. minor (left)

and A. breviculus (right);

2, Genital operculum of 9

A. breviculus, 3, idem, A.

robustus; 4, A. robustus,

apex of abdomen of bra-

chypterous 4, dorsal view;

5, A. robustus, parameres,

ventral view. Scales: 1 mm

(1-3), 2 mm (4), 0.5 mm

©):

short stout reddish spines, bearing 6-8 long erect

setae on posterior margin; tibia rather densely set

with short stout reddish spines, bearing about four

long setae on anterior margin and one row of red

spines apically. Hind leg, femur and tibia sparsely

set with short stout reddish spines, tibia also with

an apical transverse row of reddish spines, tibia and

tarsi bearing long swimming hairs on ventral sur-

face. Claws of all tarsi shining, curved, with darker,

reddish brown apices.

Comparative notes

This species runs to A. lahu Polhemus & Polhe-

mus and A. minor Polhemus & Polhemus in the

key of Polhemus & Polhemus (1988). It shares with

A. minor, which is also from Sabah, abdominal

segments 3 and 4 bearing rounded projections me-

dially on the posterior margins. A. breviculus, how-

ever, is one millimeter longer on average, has the

angle of the embolium more produced (fig. 1), and

the ratio width / length of the pronotum greater

than in A. minor, viz. 3.6 and 3.2 respectively. The

eyes of A. minor are narrower, about three times

as long as wide, while A. breviculus has its eyes

twice as long as wide.

Etymology

Breviculus, a Latin adjective, meaning rather

short, refers to the small size of the species.

Aphelocheirus robustus sp. n.

(figs. 3-5)

Type material. - Holotype @ (brachypterous): Su-

lawesi Tenggara, K. Kolaka. Sungai Mowewe, 28. II. 1989,

leg. N. Nieser, N8922 (RMNH). — Paratypes: 9 @ 14 Q

(incl. 1 labelled allotype) brachypterous, 16 @ 7 9 ma-

cropterous, same data as holotype, deposited as follows 3

& 3 Q brach., 3 6 3 Q macr. (NC); 1 9 brach., 1 & macr.

(MBBJ); 1 6 2 2 brach., 2 & 1 9 macr. (JTPC); 1 Q brach.,

2 & macr. (SEMC); 1 & macr., 1 Q brach. (OxUM); 1 9

brach., 1 & 1 9 macr. (RMNH); 1 @ 1 Q brach, 1619

macr. (ZMA).

Additional material. - 18 larvae IV/V, with same data

as holotype.

Description

Brachypterous male. - Large species for this ge-

nus, form elongate oval, length 9.8-10.5 mm; width

men) 5.9-6.2 mm.

Colour. Somewhat variable. Dorsal side: head

yellow with black eyes, pronotum yellow with an-

terior and posterior transverse dull brown band,

scutellum yellow (most specimens) to brown, he-

mielytra brown with variable part posteriorly yel-

low, abdominal tergites dull brown with yellow

posterolateral angles, genital segments yellow.

Ventral side: head, including antennae yellow, ros-

trum pale brown, sternites brown with yellow lat-

eral margins, legs and genital segments yellow.

Head. Shining, finely rugose, produced ahead of

eyes for distance equal to 0.4 X the length of an eye;

eyes length / width = 2.4-2.5, outer margin sinuate

due to weakly developed anterolateral flange; an-

terior / posterior interoculus = 1.35-1.40.

Thorax. Pronotum rugose, sparsely set with

short fine pale setae laterally, apparently glabrous

medially, width / length = 3.7-4.0; lateral margins

bearing c. 10 stout minute erect setae. Scutellum

rugose, appearing glabrous, width / length = 2.3-

2.5, lateral margin hardly sinuate, shallow trans-

verse sulcus present along anterior margin (to fit

the posterior margin of pronotum). Hemielytra

touching medially, reaching posteriorly to or just

beyond base of abdominal tergite 3, surface rugose,

set with sparse fine pale setae, embolar margin

evenly curving anteriorly, bluntly terminated pos-

teriorly in most specimens.

Abdomen. Weakly rugose, set with fine pale se-

tae, tergites 2-8 exposed, paired glandular open-

ings present medially on posterior margin of ter-

gite 3, posterior margin of tergite 5 asymmetrical,

with a medioposterior hump-like projection,

which is delimited more clearly on the left than on

the right (fig. 4), lateral margin of all segments

with short stout setae, posterolateral angles of all

tergites 3-7 spinose, with a few long pale setae at

base of spinose angles.

Ventral surface. Length of antennal segments 1

to 4: 0.10, 0.20, 0.35 and 0.45 mm; rostrum gla-

brous, length 3.2-3.4 mm, reaching to middle

coxae. Labrum yellow, shining, anterior margin

evenly rounded. Prosternum with weak median

carina, propleura with inner projections notched;

mesosternum pruinose with a median carina, tu-

mescent posteromedially; metasternum pruinose,

NIESER & CHEN: Malesian Heteroptera

metaxiphus small and narrowly pointed. Abdomen

pruinose, posterior margins of sternites 4 and 5

with six rearwardly directed stout setae medially;

genital segments glabrous, shining.

Legs. Set with fine golden setae, fore femur, tibia

and tarsi with thick hair pads on inner surface,

claws well developed and curved in most specimens

but in several short, stunted and not curved. Fore,

middle and hind coxae with combs of long light

brown setae on anterior margins. Middle legs, tro-

chanter, femur, tibia and tarsi with thick yellow

hair pads on ventral faces; femur sparsely set with

short stout reddish spines bearing 6-8 long erect

setae ON posterior margin; tibia sparsely set with

short stout reddish spines, bearing three long setae

basally on anterior margin and one row of red

spines apically. Hind leg, femur sparsely set with

short stout reddish spines, tibia with reddish spines

along interior margin, tibia and tarsi bearing long

silvery swimming hairs on ventral surface, claws

shining, curved. In several specimens middle and

hind claws and sometimes tarsi stunted.

Male genitalia. Right paramere longest with

long golden pubescence on right margin, apex

rounded, left paramere hooked with thick golden

pilosity on left margin. (fig. 5).

Brachypterous female. - Length 9.5-10.1 mm;

width of head 2.2-2.3 mm; maximum width 5.8-6.0

mm. Subgenital plate triangular with narrow lat-

eral earlike structure (fig. 3), longer lateral setae

and shorter posterior setae spread out, not in nar-

row tufts.

Macropterous form. - As brachypterous, except

length & 10.6-11.0 mm, 2 10.0-10.6 mm. Prono-

tum well developed, subequal to maximum width

of abdomen; scutellum larger and somewhat in-

flated; mesosternum inflated with broad rounded

carina medially; hemielytra well developed (mem-

brane broken off in some females), blackish in

most fully mature specimens, leaving lateral part of

connexiva uncovered; embolium with a blunt but

distinct angle midway.

Comparative notes

This species runs to A. lorelindu Polhemus &

Polhemus in the key to brachypterous specimens of

Polhemus & Polhemus (1988); both species are

very similar in general shape and size. The female

subgenital plate of A. robustus differs from larger

Southeast Asiatic Aphelochetrus by its small lateral

‘ears’. The right paramere is narrower at base than

A. lorelindu and A. celebensis Polhemus & Polhe-

mus, its closest relatives. See also the key to the

Sulawesi species.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 6-7. Coptocatus stereos. - 6, Left paramere n. Scale

0.5 mm; 7, Tibiotarsus and anterior margin of femur of

fore leg. Scale 1 mm.

Etymology

Robustus, a Latin adjective, refers to the large

size of the species within the genus.

Remarks

The type locality is a lowland stream, 2.5 to 10

m wide, flowing through woodland with fields and

small settlements close by. Aphelocheirus was

found at shallow, partly shaded places, with mod-

erate current and coarse sand bottom. Benthic

fauna, present in rather low density, included

mainly Ephemeroptera, with a few Odonata and

Trichoptera.

The stunted claws and tarsi may be the results of

wear and tear in older specimens, but this pheno-

menon is mentioned in the description, since quite

a few specimens are strikingly affected.

Coptocatus Montandon, 1909

The genus Coptocatus was recently revised by

Polhemus (1986). Including the species described

below, four species are known from the eastern

states of Malaysia (Sabah and Sarawak).

Coptocatus stereos sp. n.

(figs. 6-7)

Type material. - Holotype & (brachypterous): Malay-

sia: S. Sabah. 105 km S of Beaufort: Long Pasia area, Sg.

Maga near confluence Sg. Pasia. 4°26'N 115°40’E, 1210

m asl., larger fast running stream in untouched lower

montane evergreen rain forest, large boulders, rapids. 3

Apr. 1987. Leg. J. van Tol (RMNH).

Additional material. - 3 larvae instar V and 1 instar III

/ IV, same data as holotype (RMNH).

Description

Holotype @. - Large, robust, highly streamlined

and dorsoventrally flattened; general colour me-

dium brown with yellowish spots. Length 20.4 mm,

width of head 5.4 mm; width of pronotum 10.6

mm, maximal width 12.2 mm.

Head. Width / length 5.4 / 4.2 mm. Anteclypeus

greatly produced, apex evenly rounded, projecting

beyond rostrum for a distance greater than exposed

rostral length when viewed laterally; eyes blackish,

convex, roughly rectangular, length / width 1.8 /

1.2 mm, raised above level of vertex and separated

from it by wide shallow sulci, lateral margins bear-

ing a weakly developed flange; anterior / posterior

interoculus 3.2 / 3.2 mm; vertex greatly produced

posteriorly, extending rearwardly about one half

the length of an eye; anteriorly with a pair of broad

shallow sublateral depressions, tapering poste-

riorly.

Thorax. Pronotum width / median length 10.4

/ 3.0 mm; lateral margins with narrow, raised

darker and slightly crenulate ridge, posterolateral

angles strongly produced, rounded. Scutellum

width / length 5.0 / 2.8 mm, anterior margin

strongly reflexed downward. Hemielytra brachy-

pterous, extending to posterior margin of abdom-

inal tergite 3 only, tips broadly rounded, surface set

with fine yellowish granular microstructure (also

present on other parts of body, although far less

dense and less distinct), embolium anteriorly de-

fined by a broad carina, explanate, lateral margin

irregularly and shallowly crenulate, set with long

recumbent light brown setae; hemielytral commis-

sure with a small projecting tooth on left hemie-

lytron distally, fitting into corresponding indenta-

tion of right hemielytron.

Abdomen. Medio- and latero-caudal parts of ter-

gite 3 and all of tergites 4-8 exposed. Lateral mar-

gins with narrow ridges and a dense row of long

light brown setae lying flat on tergites. Medially

some patches of shorter setae at base of tergites,

especially the lobes of tergite 8. Lateroposterior

angles indented with blunt projections and a tuft of

hairs in the indentation.

Ventral surface. Sternites pruinose. Gula and

part of prosternum anterior to fore coxae with a

blunt carina, posterior part of prosternum without

carina. Mesosternum with a shallow median

groove and a distinct xiphus on posterior margin.

Metasternum anteriorly with a strongly dorsally

declivent carina, posterior part flat with a short

broad blunt xiphus. Abdominal sternites with large

stigmata halfway of lateral third; sternites 4, 7 and

8 with small tufts of short golden hairs medially,

sternite six medially produced, sternites 5-8 me-

dially glabrous.

Legs. Fore leg with massive femur, length 6.5

mm, width 4.3 mm; suture between tibia and tarsus

shallow, claw short, blunt, hardly differentiated;

anterior face of tibiotarsus and femur with golden

pubescence distally becoming thicker and longer

and extending onto trochanter; anterior margin of

femur with a median hump and a subapical tooth

(fig. 7). Posterior margin of all femora with a

fringe of long hairs. Middle leg and hind femur

without spines. Middle and hind tibia ventrally

with a thick pad of golden brown pilosity (looks

like a adhesive pad; already fully developed in larva

V, and partly so in larva III/IV). Middle tibia broa-

dened apically. Hind tibia with short spines along

inner and outer faces, inner (posterior) faces with

a dense fringe of long swimming hairs.

Genitalia. Parameres symmetrical, bilobed,

outer lobe swollen (fig. 6). Aedeagus symmetrical,

basally and centrally chitinized, apically and later-

ally fleshy.

Comparative notes

The same size and similar anterior margin of

fore femur as C. kinabalu D. Polhemus, also from

Sabah, which has, however, the inner lobe of para-

mere much shorter and the lateroposterior angles

of abdominal segments not distinctly indented.

The other two species in the genus known from

Sabah and Sarawak do not exceed 15 mm in length.

Etymology

Stereos, a greek adjective, meaning: strong,

solid, cruel, refers to the size and solid build of the

species.

Remark

The holotype has a case, probably of a chiro-

nomid larva on the left hemielytron against the

inner embolar ridge.

Laccocoris Stal, 1856

Seven species are known from Malesia, but none

has been reported from Sulawesi (La Rivers 1971).

The same is true for the genus Heleocoris Stal with

six Malesian species. The last revision of both

genera is Montandon (1897).

NIESER & CHEN: Malesian Heteroptera

Laccocoris staudingeri Montandon

Laccocoris staudingeri Montandon, 1897: 440-442.

Material. - Malaysia. S. Sabah, Long Pasia area, Sg.

Ritan, 4°24'N 115°42’E, 1160 m asl. Leg. J. Huisman &

J. van Tol, 16 1 9 (RMNH).

Distribution. - Brunei and Sabah.

Remarks. - L. staudingeri differs from related

species in northern Borneo, L. horvathi Mont. and

L. maat La Rivers (La Rivers 1970) by having the

interoculus at halfway level slightly more than

twice as wide as an eye.

Naucoris Fabricius, 1775

There are six Malesian species, of which one

occurs in Sulawesi. A revision of the genus is not

available.

Naucoris scutellaris Stal

Naucoris scutellaris Stal, 1859: 266. - Lundblad 1933a:

63-67, fig. 19 (redescription).

Material. - Sulawesi Tenggara: Kendari, N8904, 1 8;

Mowewe, N8922A, 4 & 2 9; 20 km S Pomalaa, N8928

(NC).

Remarks. - A widespread species, occurring

from India trough Sri Lanka and Thailand to Jawa

and Sulawesi. The femur is strongly expanded at its

inner apical two-thirds, giving the inner margin a

deeply dented appearance. Other congeneric spe-

cies have the inner margin of fore femur more or

less straight.

NEPIDAE

Key to Indonesian and Eastern Malaysian

genera

1. Flattened species, parasternites of abdomen

visible, head distinctly narrower than the pro-

notum and partly enclosed by its anterolateral

ANOTESI(INE PINA) RE ERRE E ere 2

- Subcylindrical species, parasternites not vis-

ible, head free from pronotum and both of

subequal width (Ranatrinae) ................. 3

2. Respiratory siphon longer than the inner mar-

gin of the hemielytra ....... Laccotrephes Stàl

- Respiratory siphon not more than half as long

as the distance between apex of scutellum and

the membrane along inner margin of hemi-

elytra (only one very rare species, T. breddini

Montandon, from Northern Borneo) ..........

SPARI STIRIA Telmatotrephes Stàl

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

3. Eyes in lateral view globose, not reflexed

downwards and not obscuring ventral margin

OI NEA ET na neer Ranatra Fabricius

— Eyes in lateral view reflexed downwards, obs-

curing ventral margin of head ..................

PRAIA AI BAE Cercotmetus Amyot & Serville

Laccotrephes Stal, 1865

About eight species recorded from Malesia. A

generic revision is not available and the status of

many species is uncertain, which makes some re-

cords unreliable. Three species are known from

Sulawesi and Pulau Buton.

Key to Laccotrephes of Sulawesi and Buton

1. Body length over 35 mm, usually 40 mm or

MOL ay eet Jenssen dek L. robustus Stal

- Body length less than 35 mm ................ 2

2. Respiratory siphon as long as or somewhat

shorter than body ....... L. occultus Lundblad

- Respiratory siphon distinctly longer than body

MCO ERI E ee Si L. tristis (Stal)

Laccotrephes occultus Lundblad

Laccotrephes occultus Lundblad 1933a: 23-26, fig. 2, tab.

14 (1) [nom. nov. for L. fuscus auctores, nec. L., redes-

cription].

Material. - P. Buton: 16 km E of Baubau, N8944, 1 &

(NC). Maluku: Bacan, Wayauna, c. 50 m alt., logged forest,

hand coll., 23-26 Jun 1985, leg. J. Huijbregts, 2 9 (RMNH).

Distribution. — Sumatera, Borneo, Buton, and

Maluku (Moluccas).

Laccotrephes robustus Stàl

Laccotrephes robustus Stal, 1870: 706. - Lundblad, 1933a:

26-27, fig. 3 [redescription].

Material. — Jawa: Preanger, 1 @ 1 Q; Paluabuan,

3.X1.1911, 1 g. Sumba: Kananggar, 700 m, V.1925, 1 à;

Mao Marroe, 450 m, V.1925, leg. Dammerman, 1 9.

Malaysia (Sabah): Danum valley, 4°48’N 117°48’E, 220

m streamlet, 23.III.1987, leg. J. Huisman, 1 9 (all RMNH).

Distribution. - Widespread species: Indochina

and Taiwan, Philippines, Malacca, Sumatera, Jawa,

Borneo, Sulawesi, Sumba and Bali.

Laccotrephes tristis (Stàl)

Nepa tristis Stàl, 1854: 11.

Laccotrephes tristis; Hale, 1924: 504-507, pl. 34 (figs. 1-

3), pl. 35 (figs. 2, 4, 6), pl. 36 (fig. 18); Lundblad 1933a:

25; Lansbury 1967: 644-646 [description of male ge-

nitalia].

Material. - Sulawesi Utara: Dumoga Bone N. P., To-

raut, c. 200 m, multistr. evergreen forest, creek,

23.V.1985, leg. J. Huijbregts, 1 Q (RMNH).

Distribution. — Previously only known from

Australia and New Guinea.

Remarks. - Identification of this specimen is

doubtful. It differs from typical L. occultus by its

slightly greater size (34 vs. 31-32 mm), its nar-

rower appearance anteriorly (only visible under

direct comparison), and its much longer respira-

tory siphon (40 vs. 30 mm).

Cercotmetus Amyot & Serville

In the excellent revision of the genus by Lans-

bury (1973), six species are recorded from Malesia,

including one from Sulawesi. Most species of this

genus are seldomly collected and then usually only

one specimen at a time. Consequently, distribu-

tional patterns of most species are poorly known.

Key to Malesian species of Cercotmetus

Ea from Lansbury 1973)

. Vertex raised between the eyes, but without a

distinctituberclen sare EIER N Sr 2

— Vertex with a distinct, broadly and tooth-like

tubercle rare Rene RUE 4

2. Less than 43 mm long, middle femora clearly

shorter than prothorax (widespread species:

Sumatera, Jawa, Sarawak) aan

aaa cs C. brevipes Montandon

- More than 47 mm long, middle femora longer

thanyprothorax SERIA AE RR REN. 3)

3. Eyes relatively small, width of vertex halfway

the inner margin of eyes about twice the width

of an eye at that level, vertex with a complete

eireletiof/hairst(Sulawes ep mr

IB AINSI, 43 C. robustus Montandon

— Eyes larger, width of vertex halfway the inner

margin of eyes subequal to the width of an eye,

hairs on vertex few, not forming a complete

circlet posteriorly (Irian Jaya) ...................

TS IRE, PH NO OSG C. dissidens Montandon

4. Distal third of tergite 6 raised and often very

hairy, apex of paramere sinuate, female oper-

culum not or hardly reaching beyond tergite 6

(widespread Sumatera)! war rue eee tae

Hen SSR: OMAN AUS C. compositus Montandon

- Distal third of tergite 6 not distinctly raised

and never hairy, apex of paramere evenly

curved, female operculum reaching beyond ter-

gite 6 for about one third of its length ...... 5

5. Along median line length of anterior lobe (an-

terior to transverse grooves) of pronotum less

than three times the length of posterior lobe

(widespread, Sumatra, Jawa, Borneo) ..........

ae ee eee C. asiaticus Amyot & Serville

- Along median line length of anterior lobe

more than three times the length of posterior

lobei(Kalimantan, Sarawak) 2...

ES C. astaticus var. longicollis Montandon

Cercotmetus robustus Montandon

Cercotmetus robustus Montandon, 1911: 92-93. — Lans-

bury 1973: 95-97, figs. 36-42 [redescription].

Material. — Sulawesi Tenggara: N8922A, 2 Q (NC).

Remarks. — Apparently a rare species, only

known from the female holotype from Central

Sulawesi (‘Posso See’ = Danau Poso) so far. Several

small larvae, possibly of this species, were observed

at locality N8931 (see appendix). Both larvae and

adults are far better swimmers than Ranatra, they

seem to use their fore legs for swimming.

Ranatra Fabricius, 1790

About twelve species recorded from Malesia, of

which three have been reported from Sulawesi

(one with two subspecies). The Oriental species

have been revised by Lansbury (1972). In the fol-

lowing key to the Sulawesi species, also two unre-

corded but widespread species that might turn up

in Sulawesi, were included.

Key to Ranatra of Sulawesi

(Adapted from Lansbury 1972).

1. Head with prominent tubercle between eyes,

metasternum emarginate (fig. 9) ............ 2

- Head at most only slightly raised between eyes,

metasternum not emarginate ................ 3

2. Length of respiratory siphon subequal to

length of body (Sulawesi Tenggara) ...........

PRE R. sulawesti sp. n.

- Length of siphon one-third of body length (not

recorded from Sulawesi, but a widespread spe-

cies, Indochina, Sumatera, Jawa, Nusa Teng-

PALA) Ae Eg R. parmata Mayr

3. Width of fore femora measured from dorsal

margin to apex of large tooth about the same

as widest part of femora proximally ........ 4

- Width of fore femora measured from dorsal

margin to apex of larger tooth clearly greater

than widest part of fore femur proximally (en-

demic to Sulawesi) .... R. malayana Lundblad

4. Larger tooth of fore femora more or less equi-

distant between ends of femora (not recorded

from Sulawesi, but a widespread species: India,

Ceylon, Indochina, Sumatera, Jawa) ............

Re el R. varipes Stàl

— Larger tooth of fore femora clearly nearer to

distallendo Memo ne MEET 3

NIESER & CHEN: Malesian Heteroptera

5. Third episternum with two clumps of small

hairs (on each side) (endemic to Sulawesi Sel-

atan) rer R. longipes celebensis Lansbury

- Third episternum without clumps of small

hairs (Indonesia, incl. Sulawesi Tenggara, E.

Malaysia) no R. longipes longipes

Ranatra longipes longipes Stàl

Ranatra longipes Stàl, 1861: 203.

Ranatra longipes longipes. - Lansbury 1972: 332-334,

figs. 220-229 [redescription].

Material. - Sulawesi Tenggara: N8901, 3 & 3 9; N8909,

3 6; N8910, 2 6; N8917, 2 6; N8919, 1 & 3 9; N8930,

19.

Distribution. - Known from Sumatera, Borneo

(Brunei, Sabah), Jawa, Bali and Sulawesi.

Remarks. - Our specimens were compared with

a series from Bali, of which several were identified

by Lansbury. The specimens mentioned above do

not show any of the diagnostic characters charac-

terizing subspecies celebensis (Lansbury 1972).

Ranatra malayana Lundblad

Ranatra birdi var. malayana Lundblad, 1933a: 40, fig. IA.

Ranatra malayana. - Lansbury 1972: 319-321, figs. 142-

152.

Material. - Sulawesi Tenggara: N8917, 1 @ 1 9 (NC).

Remarks. - Until now only known by three spe-

cimens from Sulawesi Selatan. Respiratory siphon,

when folded back over dorsum, reaching halfway

head in the male, and anterior margin of pronotum

in female.

Ranatra sulawesii sp. n.

(figs. 8-11)

Type material. - Holotype &: Sulawesi Tenggara, K.

Kendari, pond with Nymphaea at Wawonggole,

20.11.1989, N8902, leg. N. Nieser (RMNH). - 10 6 79

(including one labelled allotype), same data as holotype,

deposited as follows: 1 Q (RMNH); 3 6 2 Q (NC), 16 1

9 (MBBJ); 1 & (BPUH); 1& 1 9 (TPC); 1 6 1 9 (SEMC);

1 6 (OxUM).

Description

Adult. - Measurements. Length & 34-38.3 mm;

Q 35-39.5 mm; length of siphon @ 39.0-41.7 mm,

Q 35-45 mm; width of head @ 3.2-3.3 mm, Q 3.2-

3.5 mm; humeral width of pronotum & 3.5-3.9

mm; Q 3.6-4.2 mm. Colour. Dark brown, legs

paler with indistinct annulation.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 8-11. Ranatra sulawe-

si. - 8, antenna; 9, metaster-

num, 10, genital capsule of

male, lateral view; 11, apex

of paramere. Scales: 0.5 mm

(8), 1 mm (9-10), 0.25 mm

(11).

Head with small, but distinct nodule on vertex;

eyes slightly wider than width of interoculus; lora

strongly developed, higher than clypeus, which has

two small nodules apically; second segment of an-

tennae with long, finger-like projection, subequal

to length of segment 3 (fig. 8).

Thorax. Prothorax in lateral view about 1.5 X as

long as fore coxa, about 1.1 X as long as fore femur;

anterior lobe twice as long as posterior lobe in

median line; anterior margin not conspicuously

raised; humeral width / anterior width 1.4; pro-

notal grooves well developed; posterior lobe with-

out tubercles, ventrally with a broad concavity an-

teriorly at insertion of coxae, caused by produced

anteroventral angles, remainder of prothorax flat

with a vague carina in anterior one-third. Scutel-

lum about 1.5 X longer than wide, with a shallow

broad transverse depression in apical third. Meso-

sternum with anterior margin raised, posterior

margin of metasternum deeply incised (fig. 9);

space between middle and hind coxae subequal.

Legs. Fore femur with one median tooth at apical

0.4 of femoral length, no secondary tooth distally.

Length of middle and hind femur subequal. Middle

tibia shorter, hind tibia longer than femur; apical

% of middle and hind tibia with hair fringe. Hind

femur reaching about % along last abdominal ster-

nite in both sexes.

Male genitalia. See figs. 10, 11.

Female. - Genital operculum not reaching the

base of respiratory siphon.

Comparative notes

Based on the tuberculate vertex and emarginate

metasternum this species can be included in the R.

gracilis group (Lansbury 1972). R. sulawesu can be

distinguished by (a) siphon longer than the body

length, (b) distance between the middle and the

hind coxae subequal, and (c) finger-like projection

of antennal segment 2 long. In the key by Lansbury

(1972) it runs to R. parmata Mayr, from which it

differs by its long siphon, and the shape of the pa-

ramere, which is apically more swollen in R. parmata.

Etymology

Sulawesi, a noun in genitive case, refers to the

type locality.

Remark

The type locality is a small pond at the edge of

marshy fields with some water lilies (Nymphaea

sp.), and abundant marginal vegetation flattened

and hanging into the water.

NOTONECTIDAE

Key to Indonesian and Eastern Malaysian

genera

1. Hemielytral commissure anteriorly without a

definite hair-lined pit (Notonectinae) ...... 2

— Hemielytral commissure anteriorly with a def-

inite hair-lined pit (Anisopinae) ...............

nits. Mertens berte den Anisops Spinola

2. Middle femur with an anteapical pointed pro-

tuberance Ara ere Enithares Spinola

- Middle femur without such a protuberance 3

3. Eyes contiguous or forming an ocular commis-

sure near posterior border of head .............

ee etudes Nychia Stal

- Eyes widely spaced near posterior border of

head fm mortes Aphelonecta Lansbury

Anisops Spinola, 1840

Due to insufficient knowledge of the fauna, a

key to the regional species of the genus Anisops

Spinola cannot be presented. The reader is refer-

red to Brooks’ (1951) revision and various papers

by Lansbury (1964, 1965, 1978). Several species,

e.g. A. breddini, A. nasuta and A. paracrinita, can

be found in village ponds and rice fields. Such

species tend to be very widespread in Southeast

Asia.

Anisops batillifrons Lundblad

Anisops batillifrons Lundblad, 1933: 463-464, fig. 8. -

Brooks 1951: 420-423, figs. 71, 101.

Material. - Malaysia, Sabah: 16 km NE Tenom: Agri-

cultural Research Station, Sg. Segalan, 115°59’E 5°12’N,

a. Ll, 23.XI.1987, 1 & 1 Q, leg. J. Huisman & R. de Jong

(RMNH).

Distribution. — A widespread species: India

through Indochina to Hainan, Taiwan, Iriomote

and the Philippines; apparently mainly in subtrop-

ical regions.

Anisops biroi Brooks

Anisops biröi Brooks, 1951: 454-456, fig. 49. - Lansbury

1978: 111, figs. 37-39.

NIESER & CHEN: Malesian Heteroptera

Material. - Sulawesi Tengah: 50 km SE of Palu: Lore

Lindu N.P., Sopu valley near Dongi Dongi, 6 Dec 1985

(sample B), leg. J. van Tol & J. Krikken 3 & 2 9 (RMNH).

Pulau Buton: N8939, 10 @ 4 Q (NC). Maluku: Bacan,

Wayauna, alt. 50 m, sec. growth, eutrophic pool, 5-7 July

1985, HH382, leg. J. Huijbregts, 1 @ 5 ® (RMNH).

Remarks. - Identification of this species some-

what doubtful. The rather short rostral prong sug-

gests the very similar A. rigoensis. None of the

present specimens, however, has a carina on the

posterior part of the vertex, which is reported to

be characteristic for A. rigoensis (Lansbury 1978).

Both species were only known from New Guinea

up to now.

Anisops breddini Kirkaldy

Anisops breddini Kirkaldy, 1901a: 5-6. - Brooks 1951:

439-441, figs. 78, 99; Leong 1962 [life cycle]; Lansbury

1965: 58.

Material. - Sulawesi Tenggara: N8901, 22 @ 27 9, 5

larvae; N8917, 1 6 1 9; N8918, 1 4 19; N8931, 1 6 1

©. Pulau Buton, N8938, 5 & 22 9.

Distribution. - Widespread species: India and

Sri Lanka through Indochina to Jawa, Sulawesi and

Buton.

Anisops nasuta Fieber

Anisops nasuta Fieber, 1851: 484-485. - Lundblad 1933a:

168-171, fig. 58; Brooks 1951: 416-418, figs. 60, 98;

Lansbury 1965: 61.

Material. - Sulawesi Tenggara: N8917, 4 ¢ 4 9 31

larvae; N8931, 3 6 5 Q. Pulau Buton: N8938, 16 & 21 Q.

Distribution. - A very widespread species: E.

India and Sri Lanka through China and Indonesia

to Australia and Samoa.

Anisops occipitalis Breddin

Anisops occipitalis Breddin, 1905: 152. - Brooks 1951:

344-346, fig. 22; Lansbury 1965: 61-67, figs. 3-4, 7-8.

Material. — Sulawesi Tenggara. N8908, 8 @ 15 9;

N8917, 2 6; N8927, 4 & 7 9; N8931, 1 @ 1 9. Pulau

Buton. N8938, 1 8 1 Q.

Distribution. - Widespread through Indonesia

to Northern Australia.

Anisops paracrinata Brooks

Anisops paracrinata Brooks 1951: 329-331, fig. 12. -

Lansbury 1965: 57-58.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Material. - Sulawesi Tengah. SW of Luwuk, Totop

camp along Batui River, 19-21 Oct 1989, Sul. 18, J. P.

Duffels, 1 & (ZMA). Sulawesi Tenggara. Desa Kagunyala,

N8908, 1 & 6 9; Jalan Asera, N8917, 26 & 23 9; Jalan

Asera, N8918, 5 & 1 9; 20 km S Pomalaa, N8927, 4 &;

15 km S Pomalaa, N8931, 17 & 17 9. Pulau Buton. Desa

Gareg-Gareng, N8938, 36 G © 2.

Distribution. - Sumatera, Jawa, Bali, Sumbawa,

Sulawesi, Buton, E. Australia.

Anisops stali Kirkaldy

Anisops stali Kirkaldy, 1904: 113, 132. - Brooks 1951:

319-322, figs. 9, 107; Lansbury 1965: 58.

Material. - Sulawesi Tenggara. Desa Kagunyala,

N8908, 2 9; 15 km S Pomalaa, N8931, 2 9.

Remark. - Identification of these specimens un-

certain since no males were available; they agree

with specimens from Jawa.

Distribution. - Widespread; the Philippines,

Jawa, Sulawesi, Nusa Tenggara, Australia, Okin-

awa.

Anisops tahitiensis Lundblad

Anisops tahitiensis Lundblad, 1934: 121-123, figs. 1-5. —

Brooks 1951: 376-378, fig. 40; Lansbury 1964: 217-

218, fig. 7.

Material. — Sulawesi Tenggara. 20 km E Kolaka,

N8934, 4 6. Malaysia, Sabah. 20 km W of Sandakan,

Sepilok-Laut. ML, 4 Nov. 1987, leg. J. Huisman & R. de

Jong, 1 8 (RMNH).

Remark. - The specimens were compared with

a male from New Guinea and they differ only in the

facial carina being slightly less prominent in the

Sulawesi and Sabah specimens.

Distribution. - Widespread: Andaman Islands,

Borneo, Sulawesi, the Philippines, New Guinea,

New Hebrides, Tahiti, Guadalcanal, Okinawa.

Aphelonecta Lansbury

This genus contains three species, one from

Thailand and two from Borneo (Lansbury 1966).

Aphelonecta alexis Lansbury

Aphelonecta alexis Lansbury, 1965a: 328-330, figs. 2-18.

- Lansbury 1966: 632.

Material. - Malaysia, Sabah. 60 km W of Lahad Datu,

Danum Valley, 14 Mar 1987 (B), leg. J. Huisman, 1 &

(RMNH).

Distribution. - Sabah and Sarawak.

Enithares Spinola

A large, mostly Oriental genus. Thirty-three

species are known from Malesia, of which seven

occur in Sulawesi and one on Buton. There is an

excellent revision of the Oriental species (Lansbury

1968).

Key to the males of Enithares of Sulawesi and

Buton

1. Embolium in ventral view (ventral ridge of

hemielytron) greatly expanded anteriorly (fig.

21); pronotal humeral angles produced into

broad knobs. Length about 9 mm (Sulawesi)

El ahve seh ce i en Bai arr E. producta Lsb.

- Embolium in ventral view not greatly ex-

panded anteriorly; pronotal humeral angles

nouproduced genant RN WAN eh een ae 2

Dinlensthiupito) in 1... eee eee een >

— Length) Stmmyrorsmore m A 4

3. Head width twice the median length or less,

anterior lobe of genital capsule bilobed due to

a deep and broad median incision (widespread

SPECIES) LENIHAN). ARE. E. bakeri Brks.

— Head width just over twice its median length,

anterior lobe of genital capsule not bilobed (P.

Biak, possibly Sulawesi Tenggara) ..............

O Sa VENE ROTTA E. vulgaris Lsb.

4. Length 12.5 mm, median length of head one-

fourth of its length shorter than median length

of pronotum, anterior margin of middle tibia

straight to very slightly concave, posterior lobe

of genital capsule broadly rodlike (Sulawesi

Sélatan) wate. Po bassa: E. horvathi Kirk.

- Length not over 12 mm; if apex of posterior

lobe of genital capsule rodlike, then length not

over 11 mm, anterior margin of middle tibia

moreonlless convex sees Tee cee 5

5. Apex of posterior lobe of genital capsule rod-

like (figs: 13516) G22 RE RARA RON 6

- Apex of posterior lobe of genital capsule

rounded) (figsapl ZAIRE 7

6. Middle tibia not distinctly broadened (fig. 18),

rod-like apices of posterior lobes of genital

capsule rather short with small solid pegs on

inner sides (fig. 13) (Sulawesi Tenggara) ......

AT Sio E. lansburyi sp. n.

- Middle tibia distinctly broadened (fig. 20), rod-

like apices of posterior lobes of genital capsule

more elongate without pegs on inner sides (fig.

16) BMButon) Weeze E. skutalis sp. n.

7. Apical half of middle femora with very long

hairs, covering ventral margin in inner view;

genital capsule with a bilobed posterior lobe

bearing an apicaudal tuft of setae (fig. 12) (Su-

lawesi Tengah) ............. E. caesaries sp. n.

- Middle femora without strikingly long hairs,

posterior lobe of genital capsule not bilobed,

withouttubtor setae (Migone

OI PA on E. phenakismos sp. n.

Enithares bakeri Brooks

Enithares bakeri Brooks, 1948: 40, pl. 1, fig. 3. - Lansbury

1968: 384-385, figs. 91-95; Nieser & Chen, in press.

Material. — Sulawesi Tenggara. Desa Kagunyala,

N8908, 1 4 1 9; Aopa marsh, N8913, 1 9; Jalan Asera,

N8917, 1 g; 15 km S Pomalaa, N8931, 4 9. Pulau Buton.

Desa Gareg-Gareg, N8938, 1 & 1 9. Maluku. Bacan,

Wayaua, eutrophic pool, HH382, 6-7 July 1985, leg. J.

Huijbregts, 2 6 3 ® (RMNH).

Distribution. - One of the few widespread spe-

cies of Enithares in the area, occurring in the Phi-

lippines, Sabah, Sulawesi, Maluku, Flores.

Remarks. - The length of the females from Su-

lawesi and Buton centers around 7.5 mm, which is

slightly less, and of the specimens from the Maluku

around 8.3, which is slightly more than indicated by

Lansbury (1968). This suggests that the variability

in size is greater than known up to now, and that

there is significant variability between popula-

tions.

Enithares bakeri is smaller than most of its con-

geners, and lives e.g. in village ponds together with

the more common species of Anisops.

Enithares hippokleides Kirkaldy

Enithares hippokleides Kirkaldy, 1898: 73. — Lansbury

1968: 400-402, figs. 165-172 [redescription, synon-

ymy |.

Material. — Jawa, S. Muller [19th century], 1 6 2 9; G.

Malang, Aug. 1935, 4 @ 2 © (all det. E. fruhstorferi by

Brooks) (RMNH).

Distribution. - Only known from Jawa.

Enithares intricata Breddin

Enithares intricata Breddin, 1905: 154. - Lundblad 1933a:

174-177, fig. 64, pl. 5, 21; Lansbury 1968: 404-405,

figs. 185-190 [redescription, synonymy ].

Material. — N. Sumatra: Toba plateau, Tigadolok,

2°50°N 99°03’E, c. 150 m, 20 Jun 1972, leg. J. Krikken

(21), 1 9 (RMNH).

Distribution. - Sumatera and Jawa.

Enithares producta Lansbury

Enithares producta Lansbury, 1968: 368-369, figs. 15-21.

NIESER & CHEN: Malesian Heteroptera

Material. - Sulawesi Utara: Dumoga Bone N. P, sev-

eral localities, incl. Tumpah river, Toraut river, Waterfall

Creek, small creek near basecamp, leg. J. P. Duffels, J.

Huijbregts, J. van Tol, total 13 & 14 9 (ZMA, RMNH).

Sulawesi Tengah. 60 km SE Palu: Lore Lindu NP, Danau

Tambing and brooklets, 1600 m, 7 Dec 1985, leg. J. van

Tol, 1 ® (RMNH).

Distribution. — Sulawesi.

Enithares ripleyana Lansbury

Enithares ripleyana Lansbury, 1968: 402-403, figs. 173-

178.

Material. - Maluku. Bacan. Wayaua, logged forest, 5-16

July 1985, leg. J. Huijbregts, 1 @ (NC).

Distribution. - Bacan, which is the present spell-

ing of Batjan, the type locality of this species.

Remarks. - This specimen agrees with the des-

cription of E. ripleyana, except for its smaller size

(7.3 mm), and the apical part of the posterior lobe

of the genital capsule, which is inclined caudad. The

dense tuft of setae on the genital capsule look solid

when glued dry on a card.

Enithares vulgaris Lansbury

Enithares vulgaris Lansbury, 1968: 403-404.

Material. — Sulawesi Tenggara. 20 km E Kolaka,

N8934, 1 9.

Distribution. - So far only known from the type

series from P. Biak, situated east from the NW part

of Irian Jaya.

Remarks. - The size of the Sulawesi specimen is

9.1 mm, which indicates to E. vulgaris rather than

to E. intricata Breddin. Males are needed for defin-

itive identification.

Enithares caesaries sp. n.

(Figs. 12, 17)

Type material. - Holotype @: Sulawesi Tengah. Luwuk

area, Sungai Tikalalang, 20 Oct 1989, 89JvT26, leg. J. van

Tol (RMNH).

Description

The description is based on the holotype only.

Generally a rather large boat-shaped species, grea-

test width at level of apex of scutellum.

Measurements. — Length 10.3 mm, maximal

width 4.03 mm, width of head 3.22 mm, humeral

width of pronotum 3.81 mm, anterior width of

vertex 1.10 mm, synthlipsis 0.70 mm.

Colour. — Pale luteous, eyes dark brown, apical

DO,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 12-16. Genital capsule in male Enithares. - 12, E. caesaries, holotype; 13, E. lansburyi, paratype; 14, E.

hippokleides; 15, E. phenakismos, holotype; 16, E. skutalis, paratype. Scale bar 1 mm.

half of membrane smoky brown to blackish. Ven-

tral side pale with rostrum, stripe along inner (cos-

tal) margin of embolium, pilosity, spines, patches

and stripes dark brown to blackish.

Head and thorax. - Anterior margin of vertex in

dorsal view truncate, hardly produced anterior to

eyes. Greatest width of head 2.3 X its median

length; median length of head and median length

of pronotum subequal, somewhat longer than an-

terior width of vertex. Humeral width of pronotum

3 X its median length, lateral margin slightly di-

verging, posterior margin somewhat sinuate. Dor-

sal margin of pronotal fovea slightly diverging

behind eyes. Embolium only slightly expanded in

anterior third. Nodal furrow about its own length

from membranal suture.

Legs. - Fore trochanter narrow posteriorly, with

some long hairs, without nodule on ventral side,

NIESER & CHEN: Malesian Heteroptera

Figs. 17-20. Middle tibia of

male Enithares. 17, E. cae-

saries, holotype; 18, E. lans-

buryi, paratype; 19, E. phe-

nakismos, holotype; 20, E.

skutalis, paratype. Scale bar 1

mm.

meso-trochanter rounded. Mid-femoral hairs in

apical half very long, covering ventral margin in

inner view; mid tibia not broadened, slightly con-

vex along anterior margin (fig. 17), outer claw of

midleg normal.

Ventral side. - Lateral edges of metaxiphus only

slightly rounded and thickened, apical part with

sinuate margins resulting in a sharply projecting

apex. Connexiva of segments 1-3 with small black

spines, not ridged.

Male genitalia. - Genital capsule as in fig. 12;

posterior lobe strongly sclerotized, its dorsal mar-

gin incised, the distal top with a distinct tuft of

hairs; anterior lobe less sclerotized, parameres very

small, set in depressions in sides of capsule.

Comparative notes

In the key by Lansbury (1968) this species runs

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

1 a

Fig. 21. Base of hemielytra and connexiva in Enithares,

semidiagrammatical, in ventral view. Upper: greatly ex-

panded anteriorly (as in E. producta), lower: not ex-

panded.

to E. alexis Lansb. in view of the midfemoral hairs,

or, ignoring this character, to E. hippokleides Kirk.

or E. timorensis Brooks. The last two species lack

the midfemoral hairs, have also the middle tibia

less hairy and the dorsal margin of the hindlobe of

the genital capsule not incised. This species is also

quite similar to E. lansburyi sp. n. (see the key).

Etymology

Caesaries (Latin: thick strands of hair), a noun

in apposition, refers to the pilosity of the middle

femur.

Enithares lansburyi sp. n.

(figs 13, 18)

Type material. - Holotype &: Sulawesi Tenggara, small

mountain stream 20 km E of Kolaka, N8934, 3 March

1989, leg. N. Nieser (ZMA). - Paratypes: 1 9 (ZMA); 3 &

3 Q (allotype) (NC, 1 & MBBJ).

Description

Shape. - Rather large boat-shaped species, grea-

test width over pronotal humeral angles.

Measurements. - Length @ 9.4-10.1 mm, @ 9.2-

10.0 mm; maximal width 3.7-3.9 mm; width of

head 3.0-3.1 mm; humeral width of pronotum 3.7-

3.9 mm; anterior width of vertex 1.0-1.1 mm; syn-

thlipsis 0.61-0.69 mm.

Colour. - Pale form: Pale luteous, eyes dark

brown, apical half of membrane and a large x-

shaped spot at apex of scutellum smoky brown to

blackish. Ventral side pale with proximal part of

embolium, pilosity, spines, patches and stripes

blackish. Dark form: anterior fourth of pronotum

smoky brown, scutellum shiny black with, in fresh

specimens, light greenish somewhat fluorescent

stripes along lateral margins, in dried specimens

these stripes become pale luteous. Hemielytra hya-

line with apex of clavus, inner angle and outer band

of corium and basal part of membrane black. Dor-

sum of abdomen black, showing through hyaline

parts of hemielytra.

Head and thorax. - Anterior margin of vertex in

dorsal view truncate, slightly produced anterior to

eyes. Greatest width of head 2.5 times its median

length in male, just over two times its length in

female; median length of head and median length

of pronotum subequal, somewhat longer than an-

terior width of vertex. Humeral width of pronotum

just under three times its median length, lateral

margins slightly diverging, posterior margin shal-

lowly sinuate. Dorsal margin of pronotal fovea

slightly diverging behind eyes. Embolium only

slightly expanded in anterior third. Nodal furrow

virtually straight with tip inclined cephalad,

slightly less than its own length removed from

membranal suture in male, distance and length

subequal in female. Legs. - Fore trochanter nar-

row posteriorly, with some long hairs, without

nodule ventrally, middle trochanter rounded. Male

middle tibia not broadened, slightly convex along

anterior margin (fig. 18), outer claw of middle leg

normal.

Venter. - Metaxiphus with rounded and thick-

ened lateral edges, apical part with sinuate margins

resulting in a sharply projecting apex. Connexiva,

segments 1 and 2 with small spines, not ridged.

Male genitalia. - Genital capsule as in fig. 13,

posterior lobe strongly sclerotized, long and stylus-

like distally, the distal part with hairs and apically

some spines, mainly located on the inner faces.

Anterior lobe less sclerotized, parameres very

small, set in depressions in sides of capsule.

Comparative notes

In the key by Lansbury (1968) this species runs

to E. hippokleides Kirk., which lacks the elongate

stylus-like apices of the hind lobes of the genital

capsule. E. lansburyi is very similar to E. skutalis,

but differs in the apex of the metaxiphus, which is

more pronounced, and in the characters mentioned

in the key.

Etymology

Named in honour of Dr. I. Lansbury for his

excellent work on this genus and other Oriental

Notonectidae.

Biological notes

E. lansburyi and E. skutalis were found in similar

habitats, virtually stagnant, shaded pools asso-

ciated with small streams in woodland. In each pool

one or two specimens can be found, while larvae,

especially the smaller ones, may be found in larger

numbers and are of the same or of consecutive

instars. We hypothize, that gravid Enithares fe-

males seek out suitable ponds to deposit a number

of eggs. The larvae start to use other kind of food,

but in the end, when supplies become exhausted,

resort to cannibalism. In this way the species stores

the energy to produce a few adults temporarily in

the larval population, which is also known in sev-

eral species of Notonecta.

Enithares phenakismos sp. n.

(Figs. 15, 19)

Material. - Holotype @: Sulawesi Tengah: Lore Lindu

National Park, 10 km NE Gimpu, lower montane forest,

Rano Rano, 1600 m, 15 Mar 1985, Sta. 42, J. P. Duffels

& M. J. Duffels (ZMA).

Description

Only the holotype is known. Rather large boat-

shaped species, greatest width probably at a level

halfway scutellum (in the type right hemielytron

somewhat broken and folded outward).

Male. - Measurements. Length 11.9 mm, width

of head 3.72 mm, humeral width of pronotum 4.3

mm, anterior width of vertex 1.40 mm, synthlipsis

0.98 mm.

Colour. Pale luteous, eyes black, vertex light

brownish, dark spot on abdominal dorsum at apices

of clavi shining through hemielytra, which are

opaque except for apical, hyaline half of mem-

brane. Ventrally pale with stripes, pilosity and

spines mostly dark brown to blackish.

Head and thorax. Anterior margin of vertex in

dorsal view rounded, hardly produced anterior of

eyes. Greatest width of head twice its median

length; median length of head and median length

of pronotum subequal, only slightly longer than

anterior width of vertex. Humeral width of pro-

notum three times its median length, lateral mar-

gins slightly diverging, posterior margin nearly

straight in median part, gently curved anteriorly in

lateral parts. Dorsal margin of pronotal fovea con-

verging behind eyes. Embolium only slightly ex-

panded in anterior third. Nodal furrow about its

own length removed from membranal suture.

Legs. Fore trochanter posteriorly narrow, with

some long hairs, without nodule ventrally, meso-

trochanter rounded. Mid-femoral hairs normal,

with a row of short black hairs near suture with

trochanter, but lacking the small patch of short

black bristles at its apex, which are visible in most

specimens of E. hippokleides, middle tibia not

broadened, slightly convex along anterior margin

(fig. 19), outer claw of middle leg normal.

Venter. Sides of metaxiphus somewhat rounded

and thicken, apex long and sharply projecting.

Connexiva of segments 1-3 with small black spines,

not ridged.

Genital capsule as in fig. 15, posterior lobe with

sinuate posterior margin.

NIESER & CHEN: Malesian Heteroptera

Comparative notes

This species runs to E. hippokleides Kirk. in the

key by Lansbury (1968). At first sight it looks like

an oversized specimen of E. hippokleides, includ-

ing a similarity of the genital capsule and the pilo-

sity of the legs. On closer inspection there are

several small structural differences, viz. head width

/ synthlipsis about 6 in E. hippokleides and less

than 4 in E. phenakismos, anterior width vertex /

synthlipsis about 2 in E. hippokleides and 1.5 in E.

phenakismos, dorsal margin of pronotal fovea

slightly diverging posteriorly in E. hippokleides

and converging in E. phenakismos, apex of meta-

xiphus longer in E. phenakismos than in E. hippo-

kleides. Characters to distinguish E. phenakismos

from other new species described in this paper can

be found in the key. E. timorensis Brooks, which is

considered to be closely related to E. hippokleides,

is even smaller (length of male up to 9.5 mm) and

has the posterior lobe of the genital capsule not

sinuate.

Etymology

Phenakismos (Gr. deceit), a noun in apposition,

refers to the close similarity to E. hippokleides

Kirk.

Enithares skutalis sp. n.

(figs. 16, 20)

Type material. - Holotype &: Pulau Buton, 16 km E

Baubau, small stream in wet forest, 10.III.1989, N8944,

leg. N. Nieser (ZMA). - Paratypes: 1 @ with same data as

holotype (NC); Buton, small mountain stream, N8941, 2

8 (NC), 1& (MBBJ).

Description

Male. - Shape. Rather large, boat-shaped spe-

cies, greatest width across hemielytra just behind

pronotal humeral angles. ,

Measurements. Length 10.4-10.8 mm, maximal

width 4.10-4.15 mm, width of head 3.25 mm, hu-

meral width of pronotum 4.00-4.05 mm, anterior

width of vertex 1.15-1.20 mm, synthlipsis 0.65-

0.68 mm.

Colour. Pale luteous, eyes dark brown, apical half

of membrane smoky brown, spot just caudally of

scutellum light smoky brown. Ventral side pale

with pilosity, spines, patches and stripes blackish.

Head and thorax. Anterior margin of head in

dorsal view nearly straight. Greatest width of head

2.5 X its median length; median length of head,

median length of pronotum and anterior width of

vertex subequal. Humeral width of pronotum more

than 3 X its median length. Pronotum with lateral

margins diverging, posterior margin straight; dor-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

sal margin of pronotal fovea directed virtually

straight caudad behind eyes. Embolium only

slightly expanded in anterior third. Nodal furrow

virtually straight with tip inclined cephalad,

slightly less than its own length removed from

membranal suture.

Legs. Fore trochanter narrow posteriorly, co-

vered with thick long hairs, without nodule ven-

trally, mesotrochanter rounded. Male middle tibia

slightly broadened, convex along anterior margin

(fig. 20), outer claw of middle leg normal.

Venter. Metaxiphus with distinct rounded and

thickened lateral edges, apical part triangular with

virtually straight lateral margins. Connexiva of

segments 1 and 2 with small black spines, not

ridged.

Genital capsule as in fig. 16, posterior lobe

strongly sclerotized, long, hairy and distally stylus-

like. Anterior lobe less sclerotized, parameres very

small, set in depressions in sides of capsule.

Comparative notes

This species runs to E. hebridensis Lansb. or E.

hippokleides Kirk. in the key by Lansbury (1968),

which, however, both lack the elongate stylus-like

apex of the hind lobe of the genital capsule, and

differ in the shape of the male middle tibia. The

genital capsule of E. skutalis has a structure similar

to E. horvathi and E. lansburyi, but in detail these

are quite different. Besides, E. horvathi is two mm

longer and has the anterior margin of the male

middle tibia slightly concave (convex in E. skuta-

lis).

Etymology

Skutalis (Gr. baton), a noun in apposition, refers

to the rodlike shape of the apex of the posterior

lobe of the genital capsule.

Nychia Stàl

Genus with only a few, mostly poorly known

species. Apparently only one species in Malesia

(Lansbury 1985).

Nychia sappho Kirkaldy

Nychia marshalli var. sappho Kirkaldy, 1901a: 809-810.

Nychia malayana Lundblad, 1933a: 148-155, figs. 49-51.

Nychia sappho; Lansbury 1985: 4-8, figs. 7-21.

Material. - Sulawesi Tenggara: Wawanggole, N8901, 2

8 4 9; Sungai Sampara, N8909, 3 & 3 9; 2 km E Sungai

Sampara, N8910, 1 9; Aopa marsh, N8913, 7 6 3 9; Jalan

Asera, N8916, 1 9. Pulau Buton: Road to Lawele, N8939,

5 & 5 9, all brachypterous.

Distribution. - Malaysia, Indonesia, New Guinea

and Northern Australia.

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(Hemiptera - Heteroptera: Notonectidae) in Austra-

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Received: 12 October 1990

Revised version accepted: 30 March 1991

NIESER & CHEN: Malesian Heteroptera

APPENDIX 1

Localities sampled by N. Nieser, Indonesia, Su-

lawesi Tenggara, 1989.

N8901-N8920: Kabupaten Kendari

N8901-N8902: Kecamatan Wawotobi

N8901. Wawonggole, Sungai Anggoro, 20 Feb 1989.

Quietly flowing stream in open woodland, width 4-5

m, depth up to 1 m, dark brown water; vegetation

hanging from the banks in the water.

N8902. Wawongole. 20 Feb. 1989. Pond with abundant

Nymphaea in marshy fields; shore vegetation hanging

into the water.

N8903-N8912: Kecamatan Kendari

N8903. Eastern road to Lalimboee, 3 km outside Kendari,

21 Feb. 1989. Small stream, width 2-3 m, current

15m/m, slower in ponded sites, bottom sandy, some

detergents, but clear.

N8903A. Same site, puddle at edge.

N8904. Eastern road to Lalimboe, 4 km outside Kendari,

21 Feb. 1989. Turbid, near stagnant stream, width 1.5

m, depth up to 1 m.

N8905. Desa Kagunyala, 21 Feb. 1989. Trench.

N8906. Desa Kagunyala, 21 Feb. 1989. Pond covered with

Azolla and Lemna. 100 X 10 m, depth 0.05 m, bottom

clay and mud.

N8907. Desa Kagunyala, 21 Feb. 1989. Pond with some

Lemna, 15 X 5 m, depth 0.05 m, bottom clay and mud.

N8908. Road outside Desa Kagunyala, 21 Feb. 1989.

Pond, 100 X 50 m, turbid, beige, grass-like vegetation

along one bank. Many Cybister and Dytiscus.

N8909. Sungai Sampara, second bridge along road Ken-

dari-Wawotobi, 22 Feb. 1989. River, sampled along

bank at bridge, water turbid, beige, sand bottom.

N8910. About 2 km E of N8909, 22 Feb. 1989. Puddle in

dry stream bed in savannah-like area at foot of hill

with monsoon forest.

N8911. Small stream about 8 km E of N8909, 22 Feb.

1989. Width 3 m, depth 0.6 m, water turbid, beige,

bottom sand and loam. Current up to 5 m/min.

N8912. Puddle high on banks of N8911, 22 Feb. 1989.

Dimensions 10 X 3 m, depth 0.4 m.

N8913-N8915: Kecamatan Lembuya

N8913. Aopa marsh, 23 Feb. 1989. Bay in marsh at office

building, grass-like vegetation on banks, Ceratophyl-

lum in water, water light brown but clear.

N8914. Road Lembuya-Palangga, desa Lamooso, 23 Feb.

1989. Sungai Simbangi. Open loamy and sand bottom.

Slow current.

N8915. Road Lembuya-Palangga, 23 Feb. 1989. Swampy

banks of stream in savannah.

N8916-N8920. Kecamatan Asera.

N8916. Jalan Asera, first stream drom bridge over S.

Sampara, 24 Feb. 1989. Riffles with current 15-20

m/min. Rhagovelia.

N8917. Pool in meadow near N8916, 24 Feb. 1989. Di-

mensions 4 X 2.5 m, depth up to 0.3 m, water turbid,

beige, grass-like vegetation and Polygonum.

N8918. Flooded banks of second stream, 24 Feb. 1989.

Meadows mainly with Cyperaceae, water turbid, beige.

N8919. Same road, but here named Jalan Paku Jaya, fifth

stream at slightly higher elevation, 24 Feb. 1989. At

bridge, slow current, bottom sand and clay.

N8920. Same road, fourth stream, coming from wood-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

land, 24 Feb. 1989. Ptilomera in shade of trees, Lim-

nogonus in exposed puddle aside of current.

N8921-N8934: Kabupaten Kolaka

N8921. Kecamatan Kolaka

N8921. Sungai Kolaka, 27 Feb. 1989. Upstream of Kol-

aka, river, bottom sand and pebbles, current variable.

Ochterus and Hydrometra on mudflat.

N8921 A. Same site, places with current.

N8921B. Same site, ponded sites.

N8922-N8924: Kecamatan Mowewe.

N8922. Sungai Mowewe, 28 Feb. 1989. Lowland stream,

2.5-10 m wide, in woodland with fields and settle-

ments closeby. Water clear, colourless. Bottom sample

at shallow, partly shaded places with moderate cur-

rent, bottom coarse sand and small pebbles.

N8922A. Same site, pothole with undercut bank, tree

roots, floating plant debris, depth over 1 m.

N8923. Small stream in fields and gardens near N8922,

28 Feb. 1989. Width 0.5-1 m, depth 0.1-0.3 m, bottom

mostly pebbles, rather fast current.

N8924. Puddles on path at N8922-N8923, 28 Feb. 1989.

N8925-N8931: Kecamatan Kolaka.

N8925. Tamborasi, 1 Mar 1989. Subterraneous stream

flowing into small creek.

N8926. Between Tamborasi and Wolo, 1 Mar. 1989.

Mountain stream through cocoa plantation. Ptilomera

and Rhagovelia on more quiet sites behind boulders.

Limnogonus on puddle on banks.

N8927. About 20 km S of Pomalaa, 2 Mar. 1989. Pool at

edge of marsh, 2 X 2 m, depth up to 0.7 m, bottom

loam, water turbid, beige. Juncus, Chara, herbaceous

plants.

N8928. Small stream in same marsh as N8927, 2 Mar.

1989. Water clear, light brown, flowing through dense

marsh vegetation, sample at culvert under road, width

1 m, current c. 10 m/min, depth up to 0.7 m, bottom

sand with pebbles.

N8929. Pond at edge of marshy forest. 2 Mar. 1989.

Nymphaea, Juncus.

N8930. Small stream in marsh, 2 Mar. 1989. Open part

with flooded banks. Grass-like vegetation along banks

with considerable amount of epiphytic algae. Most

species from flooded banks, but Rhagovelia from the

stream.

N8931. C. 15 km S of Pomalaa, 2 Mar. 1989. Sand pits at

edge of marsh. Water clear to somewhat turbid, light

brown, depth up to 0.5 m.

N8932-N8933: Kecamatan Mowewe.

N8932. Road to Mowewe, 3 Mar. 1989. Narrow stream

at edge of marsh, water dark brown, bottom with

much muddy detritus and plant debris, some Azolla.

N8933. Wide stream near N8932, 3 Mar. 1989. Sample

from shallow bay without current, soft loamy bottom,

much debris from palmtree leaves.

N8934. Kecamatan Kolaka.

N8934. Road to Kendari, 20 km E of Kolaka, 3 Mar. 1989.

Small mountain stream in woodland, 1-1.5 m wide,

depth very variable, potholes up to 1 m, current very

variable, small waterfalls and ponded sites.

N8935-N8944. Kabupaten Buton

N8935. First stream crossing road from Baubau to the

north, 8 Mar. 1989. Downstream of man-made water-

fall, width 5 m, depth and current velocity variable,

water colourless, somewhat turbid.

N8936. Desa Gareng-gareng, 8 Mar. 1989. Pool with

many small fish.

N8937. Desa Gareng-gareng, 8 Mar. 1989. Pool with very

few fish.

N8938. Desa Gareng-gareng, 8 Mar. 1989. Dirty pond

without fish.

N8939. Road to Lawele, 9 Mar. 1989. Mangrove swamp

E of road, bottom with thin layer of mud and man-

grove roots, marshy vegetation.

N8940. Lower course of small stream, 9 Mar. 1989. Bot-

tom bare grey sand and mud, some plant debris, slug-

gish.

N8941. Small mountain stream close to the sea, 9 Mar.

1989. Bottom rocky without sand or pebbles, vary

from small waterfalls to nearly stagnant pools, no

water vegetation, but banks with dense cover of

mosses.

N8942. Road to Parjowiro, about 15 km E of Baubau. 10

Mar. 1989. Small cascade-stream, water turbid, and

somewhat milky colour.

N8943. Pool at edge of N8942, 10 Mar. 1989.

N8944. Road to Parjowiro, about 16 km E of Baubau, 10

Mar. 1989. Small stream, anastomosing, ponded sites,

marshy forest, many buttressed trees, hilly country.

N8945-N8949. Kabupaten Kendari.

N8945-N8949. Kecamatan Kendari.

N8945. Teluk Kendari, 13 Mar. 1989. Small stream flow-

ing through coconut plantation into bay, bottom sand

and clay.

N8946. Teluk Kendari, 13 Mar. 1989. Well near coconut

plantation.

N8947. Teluk Kendari, Pulau Bungku Toko, 13 Mar.

1989. Puddles in mangrove.

N8948. Teluk Kendari, close to sea. 13 Mar. 1989. Pool

with seawater behind sand wall overgrown with Ipomoea.

N8949. Teluk Kendari. 13 Mar. 1989. Well protected by

masonry, near N8948.

APPENDIX 2

Localities on Sulawesi and Borneo from recent col-

lections made by J. P. Duffels, J. Huisman, R. de

Jong and J. van Tol.

Borneo

14 Mar 1987 - Sabah. 60 km W of Lahad Datu. Danum

Valley near Danum Valley Field Centre, brooklet at

West 3. Sample B. Alt. 220 m asl. 4°58’N 117°48’E.

(J. Huisman).

23 Mar 1987 - Sabah. 60 km W of Lahad Datu. Danum

Valley near Danum Valley Field Centre, streamlet

near West 11 / North 5. Sample A. Alt. 220 m asl.

4°48’N 117°48E. (J. Huisman).

3 Apr 1987 - S Sabah. 105 km S of Beaufort: Long Pasia

area. Sungai Maga near confluence Sg. Pasia. Alt. 1210

m asl. Larger fast running stream in untouched lower

montane evergreen rain forest. Large boulders, rapids.

4°26'N 115°40’E. (J. van Tol).

8-9 Apr 1987 -S Sabah. 105 km S of Beaufort: Long Pasia

area. Sungai Ritan. Alt. 1160 m asl. Undisturbed ever-

green tropical rain forest. 4°24'N 115°42'E. (J. van

Tol & J. Huisman).

4 Nov 1987 - Sabah. 20 km W of Sandakan. Sepilok Laut

near resthouse. ML light. 5°49’N 118°0G'E. (J. Huis-

man & R. de Jong).

23 Nov 1987 - Sabah. 16 km NE Tenom: Agricultural

Research Station. Sungai Segalan. At light. 5° 12° N

115°59'E. (J. Huisman & R. de Jong).

Sulawesi

23 Feb 1985 — Sulawesi Utara. Dumoga-Bone NP. Project

Wallace. Edward's Camp. Tumpah river. 900 m asl

[recte 600 m]. (J. P. Duffels).

15 Mar 1985 - Sulawesi Tengah. Lore Lindu National

Park. 10 km NE of Gimpu. Lower montane forest.

Rano Rano. Alt. 1600 m asl. Duffels Sta. 42 (J. P. &

M. J. Duffels).

23 Apr 1985 - Sulawesi Utara. Dumoga-Bone NP. Project

Wallace. Waterfall Creek, tributary of Tumpah river.

Sample B. c. 225 m asl. UTM WL9768 [ recte XL0064].

0°35’N 123° 54’E. (J. van Tol).

23 May 1985 — Sulawesi Utara. Dumoga-Bone NP. Pro-

ject Wallace. Tumpah river near confluence Toraut.

Sample A. c. 210 m asl. UTM XL0063. 0°34’N

123°54E (J. van Tol).

3 Jun 1985 - Sulawesi Utara. Dumoga-Bone NP. Project

NIESER & CHEN: Malesian Heteroptera

Wallace. Tumpah river near Edward's subcamp. Sam-

ple A. Alt. c. 600 m asl. UTM WL9365. 0°35’N

123°51E. (J. van Tol).

4 Jun 1985 - Sulawesi Utara. Dumoga-Bone NP. Project

Wallace. Brooklet c. 2 km west of Edward’s subcamp.

Sample A. c. 700 m asl. UTM WL9269. (J. van Tol).

6 Dec 1985 - Sulawesi Tengah. 50 km SE of Palu: Lore

Lindu National Park. Sopu valley near Dongi Dongi.

Sample B. Alt. 950 m asl. Rainwater puddle without

vegetation, near logging road. UTM SJ86. 1°13'S

120° 11'E (J. van Tol & J. Krikken).

7 Dec 1985 - Sulawesi Tengah. 60 km SE of Palu: Lore

Lindu National Park. Danau Tambing and brooklets.

Sample A. Alt. 1600 m asl. Pandanus. 1°20'S

120° 15’E. (J. van Tol).

8 Dec 1985 — Sulawesi Tengah. 50 km SE of Palu: Lore

Lindu National Park. Sopu river near Dongi Dongi.

Sample A. Alt. 950 m asl. UTM=SJ86. 1°13'S

120° 11’E (J. van Tol).

19-21 Oct 1989. - Sulawesi Tengah. SW of Luwuk: Totop

camp along Batui river. Alt. 120 m asl. 1°09'S

122°31’30”E. Lowland rainforest. Sample Sul. 18 (J. P.

Duffels).

20 Oct 1989. — Sulawesi Tengah. Luwuk area. Sungai

Tikalalang. Sample 89JvT026. (J. van Tol).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

BOOK REVIEW

Medvedev, G. S. (editor-in-chief), 1990. Keys to the In-

sects of the European Part of the USSR. Volume IV.

Lepidoptera, Part 2. - Keys to the fauna of the USSR,

130. English Translation [originally published in Rus-

sian, Leningrad, 1981]. - E. J. Brill, Leiden. 1092 pp,

747 figs. [ISBN 90 04 08926 8, price 280 Dutch guilders,

ca US $ 140].

This book is the second volume on Lepidoptera

translated into english. The first was published in

1988 and reviewed by the late A. Diakonoff in this

journal (vol. 132: 162). While the first volume

dealt with the primitive Lepidoptera, the Tortrici-

dae and some primitive Macrolepidoptera, this vo-

lume treats most of the remaining so-called mi-

crolepidopteran families, covering the superfam-

ilies Tineoidea (incl. Gracillariidae and related

families, but excluding the Psychidae, which were

treated in vol. 1), Yponomeutoidea and Gelechioi-

dea (excluding the Coleophoridae). In total about

1280 species from the European part of the USSR

and adjacent countries are covered. Keys are pro-

vided up to species level and genitalia of many

species are illustrated. Also some adults are illus-

trated in line-figures.

Although this is a review of the translation, it

also presents an opportunity, being unable to read

russian, to review the original contents, bearing in

mind that it represents the knowledge of about

1980.

Preparing identification keys for a large order as

the Lepidoptera for the huge territory of the Euro-

pean part of the USSR is in itself a large project,

which is unprecedented outside the USSR. The

keys heavily lean on previously published revisions

and keys, mostly published outside the USSR.

However, gathering such information is worth-

wile, especially for the non-specialist, where the

originals are so scattered over journals and books.

For some families there is also original input, in

particular figures of genitalia of species never illus-

trated before. Regrettably, many male genitalia

have been illustrated in lateral view, where it is

now general practice to illustrate male genitalia in

taxonomic papers in ventral view, with the valvae

spread out. This will make comparison of standard

genitalia preparations with the figures difficult,

and could lead to misidentifications.

The quality of the various families differs

greatly, some are clearly treated by specialists, and

contain original information, others are in fact

almost copies of foreign literature, like the family

Elachistidae, which is merely an abstract of Volume

6 of Fauna Entomologica Scandinavica by Trau-

gott-Olsen & Nielsen (now sold by the same pub-

lisher as this translation!). Important original

treatments are the Gracillariidae by Kuznetsov, the

Momphidae and related families by Zagulyaev and

Sinev and the Oecophoridae by L'vovskii. The

chapter on Gelechiidae by Piskunov is important,

because it is the only currently available treatment

of European Gelechiidae. Unfortunately, the qual-

ity is below expectations, since many species are

excluded from the keys, the genitalia illustrations

are a collection of copies from other sources, in-

cluding some very poor ones, and complex original

figures in lateral view.

Although it is understandable from the point of

costs, what I feel is most missing are colour plates

of adult moths. Identification of moths should start

with comparison of colour patterns, which is so

much easier than to start making preparations of

wing venation and genitalia. With this book, iden-

tification of an unknown genus is often almost

impossible without referring to the genitalia.

I am not able to comment on the quality of the

translation, but would like to make two general

remarks. Firstly, the reproduction of the figures is

poor. Despite the poor quality of paper, the figures

in the Russian text are far superior. Secondly, it is

to be regretted that the translators have chosen to

translate the russian common host-plant names

into common english names. Although there is a

list of these names and scientific names at the end

of the book, it would have been much easier to

include scientific names, where many users will not

be familiar with english common plant names.

In conclusion, this translation is an important

review of European Microlepidoptera, which tem-

porarily will fill the gap between the older plate

books like ‘Spuler’ and the modern treatments of

‘Microlepidoptera Palaearctica’, but it should be

used which much care, since it is definitely not

complete and contains some erroneous informa-

tion. Those, who use the book merely for the illus-

trations of genitalia, are beter off with the original

russian edition, which is also much cheaper.

[E. J. van Nieukerken]

R. PUPLESIS!, S. SEKSJAEVA? & V. SRUOGA!

'Zoologyos katedra VPI, Vilnius

‘Zoological institute, Academy of Sciences, Leningrad

LEAF-MINING LEPIDOPTERA (NEPTICULIDAE,

BUCCULATRICIDAE, GRACILLARIIDAE) FROM

ULMUS IN NORTHERN CASPIYA (KASPIA)

Puplesis, R., Seksjaeva, S. & V. Sruoga, 1991. Leaf-mining Lepidoptera (Nepticulidae,

Bucculatricidae, Gracillariidae) from Ulmus in northern Caspiya (Kaspia). - Tijdschrift

voor Entomologie 134: 69-73, figs. 1-9. [ISSN 0040-7496]. Published 1 July 1991.

Three leaf-mining species from elm (Ulmus carpinifolia Rupp. ex Suckow): Stigmella

kazakhstanica Puplesis sp. n. (Nepticulidae), Bucculatrix caspica Puplesis & Sruoga sp.

n. (Bucculatricidae) and Phyllonorycter schreberella (F.) (Gracillariidae) are recorded

from the northern Caspiyan region (USSR). The new species are described, genitalia and

mines are illustrated. Phyllonorycter schreberella (F.) is diagnosed and its male genitalia

are figured.

Correspondence: Dr. R. Puplesis, Zoologijos katedra VPI, g. Studentu 39, Vilnius 34,

Lithuania.

Keywords. - Nepticulidae; Bucculatricidae; Gracillariidae; leaf-mining Lepidoptera;

northern Caspiya.

The large northern Caspiyan (=Kaspian) region

is generally characterized by a strong continental

climate and by arid landscapes. Except oases by the

rare rivers and settlements, there are only dry

steppes, semideserts and deserts. The abundance of

salt lakes, saline soils (solanchaks and solonetzes)

is very characteristic. The vegetation of most of

these biotopes is extremely poor, usually without

woody plants. Grass cover is thin, the vegetation

consists mainly of different worms-woods

(Artemisia spp.) and cereals, with worm-woods

dominating. The sum of active temperature equals

2800-3400 °C with a wetting coefficient of 0.15-

0.10 (Tushinskij & Davydova 1976).

In oases near the settlements the elm (U/mus

carpinifolia Rupp. ex Suckow) is the dominating

tree. In a few cases it is the only tree present.

Hardly any published data on leaf-mining Lepi-

doptera were available from these regions. Only

one Stigmella (Nepticulidae) specimen deposited

in the Zoological institute (Leningrad), reared by

G. Lindeman from Ulmus sp., from Dzhanibek

(Western Kazakhstan), was known. It belongs to a

new species.

During investigations in early August 1988 in

the localities (fig. 1) Baskunschak (Astrakhan Re-

gion), Gurjew (Gurjew Region), Beyneu (Man-

gyschlak Region), Kara-Kalpakiya and Kungrad

(Kara-Kalpakskaya ASSR), mines of three leaf-

mining Lepidoptera were discovered. Bucculatrix

caspica Puplesis & Sruoga sp. n. was found in the

largest numbers. Numerous cocoons and empty

mines were collected from Ulmus carpinifolia in all

investigated localities, Gurjew and Kungrad. Our

later study of 1988 in neighboring Turkmeniya

showed its absence from the Central Asiatic part of

the USSR. Abundant mines of Phyllonorycter

schreberella (F.), containing larvae or pupae were

collected on Ulmus carpinifolia in Baskuntschak. In

other places of the Caspiyan region, this species

remains yet unrecorded. One empty mine of Nep-

ticulidae was found on Ulmus carpinifolia in Bas-

kuntschak, i. e. about 140 km South of Dzhanibek,

where the above mentioned Stigmella specimen on

Ulmus sp. was collected by Lindeman in 1966. It is

therefore considered to belong to the same species.

Ulmus spp. (including U. carpinifolia Rupp. ex

Suckow) are known to be host plants for Gracilla-

riidae (Kuznetzov 1981), Bucculatricidae (Seks-

jaeva 1981) and Nepticulidae (Johansson &

Nielsen 1990).

No other lepidopterous miners on other plants

have been found in this region.

The type specimens are deposited in the collec-

tion of the Minological research laboratory at the

Department of Zoology of the Pedagogical insti-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Fig. 1. Distribution map of leaf-mining Lepidoptera in

northern Caspiya: Stigmella kazakhstanica (rectangles),

Bucculatrix caspica (dots), Phyllonorycter schreberella

(triangle).

tute, Vilnius, Lithuania (MRL) and in the collection

of the Zoological institute of the USSR Academy

of Sciences in Leningrad (ZIAS).

TAXONOMIC PART

Nepticulidae

Stigmella kazakhstanica Puplesis sp. n.

(figs. 1-3)

Type material. - Holotype &: USSR, western Kazakh-

stan (Kazakh SSR), Dzhanibek, larva on leaf of Ulmus sp.

(probably U. carpinifolia), vi.1966, leg. G. Lindeman

(ZIAS). Leaf-mine (no type material): USSR, Astrakhan

Region, Baskuntschak, on Ulmus carpinifolia, fresh

mine, 4.viii.1988, leg. R. Puplesis et V. Sruoga (MRL).

Diagnosis. - Belongs to the Stigmella ulmivora

group. It is very similar to S. ulmiphaga (Preis-

secker) and S. ulmivora (Fologne). In the male

genitalia it is easily distinguished by the valvae,

which are abruptly broadened at the base. Cornuti

not numerous, lateral lobes of vinculum broad in

contrast to S. ulmivora (Johansson & Nielsen

1990).

Description

Male. - Frontal tuft pale orange. Antennae pale

brown. Eye-caps and collar cream. Palpi cream.

Thorax and forewings uniform, greyish brown. Ci-

lia and hindwings approximately as forewings in

colour.

Fig. 2. Male genitalia of Stigmella kazakhstanica, holo-

type (scale 0.1 mm).

Fig. 3. Mine of Stigmella kazakhstanica on Ulmus sp.,

from which the holotype was reared.

Female unknown.

Male genitalia (fig. 2). - Valva narrow medially

and apically, strongly broadened basally. Transtilla

with short sublateral processes. Juxta present, its

form resembles a triangle apically. Uncus approx-

imately with trapezium shape, with slightly taper-

ing angles. Gnathos with two thin and long poste-

rior processus. Transverse bar of gnathos narrow.

Aedeagus narrowing in middle (in lateral view).

Vesica with two groups of cornuti; one of which is

larger and situated in middle, a smaller group is si-

tuated apically. Cornuti more or less in form of scle-

rotized spines, some cornuti have different forms.

Biology. — Larvae in June and August, mining

leaves of Ulmus carpinifolia Rupp. ex Suckow.

Mine a contorted gallery (fig. 3). Early mine com-

pletely filled with brownish frass; later gradually

widening with frass coiled, leaving wide clear mar-

gins; finally frass in a dark narrow line.

Distribution (fig. 1). - Only known from the

lower Volga valley.

Bucculatricidae

Bucculatrix caspica Puplesis & Sruoga sp. n.

(figs. 1, 4-8)

Type material. - Holotype &: USSR, Astrakhan Re-

gion, Baskuntschak, N 4170, cocoon on leaf Ulmus car-

pinifolia, 4.viii.1988, ex p. 7.viii.1988. Paratypes: 1 6, 1

Q, same data as holotype, ex p. 7-15.viii.1988; 1 8, same

locality, caught on leaf of Ulmus carpinifolia, 4.viii.1988,

leg. R. Puplesis and V. Sruoga (MRL). Leaf-mines (no type

material): Mangyschlak Region, Beyneu; Kara-Kalpaks-

kaya ASSR, Kara-Kalpakiya.

Diagnosis. - This species is similar to the Euro-

pean B. ulmella Zeller (Seksjaeva 1981) (on Quer-

cus) and the neartic B. electa Braun which also

feeds on Ulmus leaves (Braun 1963). B. caspica is

easily recognized by its male genitalia with a long

vinculum, the presence of a transtilla and the form

of the valvae.

Description

Male (fig. 4). - Forewing length 2.9-3.1 mm.

Face creamy white, tuft brown or brownish in cen-

tral part, with creamy white piliform scales later-

ally. Eye-caps creamy white. Antennal flagellum

with alternating white and brown rings of equal

width. Thorax creamy white, slightly mottled, as

some scales have brownish tips. Forewing with

same colour as thorax, but some brown spots are

present. Three or four of these spots are situated

on the costal margin, the two below the fold, in

apical part of forewing are smaller and usually

darker. They include patches of blackish-brown

scales. Dark-tipped scales form a line on the pale

creamy cilia. Hindwings and cilia greyish cream to

pale brownish. Legs cream with little fuscous shad-

ing.

Female. - Forewing length about 3.3 mm. Sim-

ilar to male, but face cream, tuft slightly brownish

centrally. The spots on the forewing may be larger

than in male.

Male genitalia (fig. 5). - Valva with convex mar-

gin medially, suddenly tapering apically. Many

large and some short setae on apical and medial

part of valva. Transtilla present. Anellus forming

an asymmetric ring. Uncus with two large lateral

PUPLESIS ET AL.: Leaf-mining Lepidoptera

setosae lobes. Vinculum ventrally a large, more or

less triangular lobe, rounded anteriorly. Aedeagus

long and slightly bent basally.

Female genitalia (fig. 6). - Apophyses posteri-

ores long. Ductus bursae narrow and very long.

Bursa copulatrix more or less oval with numerous

small spines.

Egg. - Laid on the underside of a leaf of Ulmus

carpinifolia Rupp. ex Suckow, generally close to the

midrib or a lateral vein. In some cases, eggs are laid

away from veins.

Mine (fig. 8). - Gallery very narrow and compar-

atively long (18-20 mm) with black linear frass,

leaving clear margins throughout its whole course.

Cocoon (fig. 7). - Whitish cream with blackish

perpendicular patterns.

Distribution (fig. 1). - Probably widespread in

northern Caspiya, but possibly absent from the

Amudar’ya oases and Turkmeniya.

Gracillaridae

Phyllonorycter schreberella (Fabricius)

(fig. 9)

Diagnosis. — It belongs to the Phyllonorycter

ulmifoliella group, which larvae are usually leaf-

miners of Ulmus (Kuznetzov 1981). It is easily

recognized from all other species in that group by

the silvery shining frons, thorax and base of the

forewings. Tuft on head black. Forewing with two

silver fasciae basally and medially, two silver spots

apically. There is a black spot on base of forewing,

near costal margin. Male genitalia (fig. 9) differs

from all other species of the genus by the narrow

and bent valvae.

Distribution (fig. 1). - P. schreberella is found in

Europe from England and Scandinavia to the Bal-

kans and the European part of the USSR, also in

Asia Minor, the Caucasus and the mountains of

Turkmeniya (Kopet-Dag ridge) (Kuznetzov

1981). The species is here recorded for the first

time from the northern Caspiyan region.

Material examined. - 2 6, 1 9, USSR, Astrakhan re-

gion, Baskuntschak, mines 4.viii.1988, ex p. 7.viii.1988,

leg. R. Puplesis and V. Sruoga (MRL).

ACKNOWLEDGEMENT

We thank Zigmantas Gudzinskas (Vilnius) for

the identification of Ulmus carpinifolia.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Ke 3 : ==

ven

ESS

Ny .

N ae EN

yy Le E

ù Y .

RQ U U

N VEN

} si

Fig. 4-7. Bucculatrix caspica. - 4, forewing (scale 1 mm); 5, male genitalia, holotype (scale 0.1 mm); 6, female genitalia,

paratype (scale 0.1 mm); 7, cocoon, from type locality.

Fig. 8. Mine of Bucculatrix caspica on Ulmus carpinifo-

lia.

REFERENCES

Braun, A., 1963. The genus Bucculatrix in America north

of Mexico. - Memoirs of the American Entomological

Society 18: 1-207.

Johansson, R. & E. S. Nielsen, 1990. Tribus Nepticulini.

- In: R. Johansson et al., The Nepticulidae and Opos-

tegidae (Lepidoptera) of North West Europe. - Fauna

Entomologica Scandinavica 23: 111-238, plates.

Kuznetzov, V. I., 1981. Gracillariidae (Lithocolletidae) —

Moli-pestrianski. - Opredelitel nasekomyh Evropeys-

koy casti SSSR, Leningrad 4(2): 274, 277-278.

PUPLESIS ET AL.: Leaf-mining Lepidoptera

Fig. 9. Male genitalia of Phyllonorycter schreberella (F.)

(scale 0.1 mm).

Seksjaeva, S. V., 1981. Bucculatricidae - Krivousiye kro-

hotki-moli. - Opredelitel nasekomyh Evropeyskoy

casti SSSR, Leningrad 4(2): 136-148.

Tushinskij, G. K. & Davydova, M. I., 1976. Fiziceskaya

geografiya SSSR. - Prosvesceniye, Moscow: 179.

Received: 5 january 1990

Revised version accepted: 22 November 1990

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

BOOK REVIEWS

Emmet, A. M., 1991. The scientific names of the British

Lepidoptera. Their history and meaning. - Harley

Books, Colchester, England. 288 pp., viii plates. [ISBN

0 946589 35 6, paperback edition. Price ca US $ 40].

Many entomologists consider scientific names

simply as labels of the subjects of their studies, and

do not bother about their meaning. Others, how-

ever, are interested in the history of the name and

wonder why they are named as they are. To cite the

author of this book: ‘There is no need to seek the

explanations of scientific names, just as there is no

need to do crossword puzzles. There are, however,

many who derive pleasure from both pursuits.’

For those who are interested in the history of

names in general and nomenclature of Lepidoptera

in particular, Colonel Emmet has provided a

wealth of interesting information, with explana-

tions of names of all British species, genera and

families, including a few synonyms, 3000 names in

total. The author not only tried to give the exact

meaning of the latin or greek words from which

the name supposedly was derived, but also tried to

trace the reasons of the author for giving it this

particular name. This was no small task, when one

realises that very few authors before this century

ever explained the new names they were propos-

ing. In many cases therefore Emmet had to guess

at the thoughts of the name-giving authors, but

was greatly helped by his immense knowledge of

British Lepidoptera. The systematic section of this

book therefore, is by no means a dry dictionary, but

well readable and full of small interesting stories

about authors of the past. Emmet also could rectify

many earlier explanations, by his combined know-

ledge of the classic languages and Lepidoptera.

That with so many names a few misinterpreta-

tions or errors were noted, is a very minor flaw

indeed. To cite some, noted by the reviewer: 34.

Ectoedemia occultella. It is very unlikely that Lin-

naeus was indicating the larval habits of this spe-

cies: he described it from specimens which he

found on his window. The genus Trifurcula (page

45) was named after the three-forked radius in the

hindwing, not forewing. Caradja (species 871a)

was a Rumanian entomologist, not German.

The introduction is a well written history of

nomenclature in Lepidoptera, and gives insight in

the way early authors, like Linnaeus, formed scien-

tific names. It also explains the curious usage of

special endings for particular groups of Lepidop-

tera (such as -ella, -ana, -ata). Emmet follows,

against his classical background, the current lepi-

dopterological practice to use original spellings and

endings, without bothering about gender of the

generic name.

Colonel Emmet has written an invaluable source

of information and fun. The book is warmly recom-

mended to all entomologists interested in names,

and should not be missing in the library of any

lepidopterist.

[E. J. van Nieukerken]

Rubtsov, I. A., 1990. Blackflies (Simuliidae). - Fauna of

the USSR, Diptera, Volume 6, Part 6: p. i-xxviii + 1-

1042, figs. 1-424 [mostly composite]. [ISBN 90 04

088717, distributed outside India by E. J. Brill, Leiden.

Price 300 Dutch Guilders, appr. us $ 150].

According to information provided by the pub-

lishers ‘the purpose of the present volume is to

provide brief descriptions of species and new iden-

tification keys. It primarily incorporates numerous

additions to the first addition of the Fauna of the

USSR. This second edition also includes 18 species

from countries adjoining the Palearctic region,

which have not been recorded to-date in the Soviet

Union, and 30 species described by Enderlein from

Europe (whose description has been improved

upon), which may be discovered later in the Soviet

Union’.

Usually translations of Russian handbooks are

most welcome, but one may doubt whether this is

also true in this particular case. Firstly the original

was published as early as 1956. Secondly, the text

is clearly out of date, since a revised and more

comprehensive work by the same author appeared

between 1959 and 1964 in Lindner's Die Fliegen

der palaearktischen Region’ in the German lan-

guage. Thus, for scientists who understand Ger-

man the information was already available for

more than 25 years. I suppose that most dipterists

not fluent in German or Russian would have pre-

ferred a translation of Lindner rather than one of

the Fauna of the ussr. The translation as such

seems to be done with reasonable care. For in-

stance, the translator notifies the reader about pos-

sible mistakes in the original text, and there is also

a translation of the lettering in the illustrations. As

usual the quality of the Indian printing is far behind

European standards, making some figures difficult

to understand. Comparing the three works it ap-

pears that the figures in Lindner are by far the best.

[J. van Tol]

BERNHARD]. VAN VONDEL

Natuurmuseum Rotterdam

REVISION OF THE PALAEARCTIC SPECIES OF

HALIPLUS SUBGENUS LIAPHLUS GUIGNOT

(COLEOPTERA: HALIPLIDAE)

Vondel, B. J. van, 1991. Revision of the palaearctic species of Haliplus subgenus Liaphlus

Guignot (Coleoptera: Haliplidae). - Tijdschrift voor Entomologie 134: 75-144, figs. 1-312.

(ISSN 0040-7496]. Published 1 July 1991.

The palaearctic species of the subgenus Liaphlus Guignot, genus Haliplus Latreille, are

revised. Thirty species are recognised. The following new synonyms are established : H.

modestus Zimmermann as a junior synonym of H. eximis Clark, H. subguttatus Crotch

and H. salinarius Wallis of H. fulvus (Fabricius), H. pelopis Sahlberg of H. guttatus Aubé,

H. syriacus Wehncke and H. zimmermanni Gschwendtner of H. maculatus Motschulsky

and H. transvolgensis Semenov of H. variegatus Sturm. Eight new species are described:

H. angusi, H. astrakhanus, H. davidi, H. excoffieri, H. gafnyi, H. holmeni, H. jaechi and

H. ortali. A key to the species is provided. Most primary types have been examined.

Lectotypes have been designated for H. abbreviatus Wehncke, H. andalusicus Wehncke,

H. basinotatus Zimmermann, H. dalmatinus Miiller, H. dalmatinus var. weberi Miiller,

H. interpunctatus (Marsham), H. lapponum Thomson, H. fulvus sparreschneideri Muns-

ter, H. guttatus Aubé, H. maculatus Motschulsky, H. pelopis Sahlberg, H. cinereus Aubé,

H. ovalis Sharp, H. sharpi Wehncke, H. syriacus Wehncke, H. zimmermanni

Gschwendtner, H. variegatus ab. pallidior Müller, H. leopardinus Sahlberg and H. trans-

volgensis Semenov.

B. J. van Vondel, Roestuin 78, 3343 CV Hendrik Ido Ambacht, The Netherlands.

Key words. - Liaphlus, palaearctic region, China, new species.

CONTENTS species have been described from the southern and

eastern parts of the region. Various problems in

INCIOGUGHONEL E TRE I DCR dé dent ia e OL cence:

MEEO a nn 76 sa 8 RESSE

SYSTEMALICISECLIO DEEE EE CE A 77 Le noe ESTER IEN

ChecklisttofitreatedispeCie Sto Rn 78 Da Ee

Key to the palaearctic species of Liaphlus … 78

Description of the palaearctic species

O2 DUI assi eri e 81

Unnamed material ............................... 135

Acknowled SEMENTI 125

RÉ LÉLENICES nr AR it Re LE hr 136

MAPS TE I I 140

INTRODUCTION

This revision deals with the palaearctic species

of Liaphlus, a subgenus of Haliplus, one of the five

genera of the waterbeetle-family Haliplidae. The

latest revision of Haliplus in the palaearctic region

was by Zimmermann (1924). The status of some

species remained uncertain. Afterwards some new

for the Haliplus-species lacking the pronotal basal

plicae and possessing a setiferous striole on the

dorsal side of the metatibia. Afterwards Guignot

(1955) proposed a division of Liaphlus into 9 spe-

cies-groups, of which 4 groups are represented in

the palaearctic region as treated in this revision.

Especially outside the palaearctic region his divi-

sion is certainly in some cases based upon misin-

terpretations. For instance the oriental species Ha-

liplus angustifrons Régimbart and H. arrowı

Guignot are placed in different groups. In my opin-

ion however, these species are very closely related.

A similar case is Haliplus pulchellus Clark and H.

diopus Guignot, also placed in different groups, but

also closely related.

The subgeneric status of Haliplus laminatus

(Schaller) is not completely clear. According to the

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Fig. 1. Haliplus flavicollis, dorsal view. - an, antenna; as, apical spur; cl, clypeus; el, elytron; fr, frons; 1, 2, 3, etc, first,

second, third, etc. elytral interval; p, primary puncture-row; pr, pronotum; s, secondary puncture-row; su, suture; ss,

setiferous striole; t, tibia; ve, vertex.

Fig. 2. Haliplus flavicollis, ventral view. - cp, hind coxal plate; ee, elytral epipleuron; mp, metasternal process; pe,

proepisternum; pp, prosternal process; ps, prosternum.

definitions for Liaphlus it should belong to that

subgenus, but the larval stages are distinctly differ-

ent and very much like the larvae found in the

subgenus Haliplus s. str. (sensu Guignot 1928)

(Vondel 1986).

Beutel & Ruhnau (1990) made a phylogenetic

analysis of the genera of Haliplidae based on char-

acters of adults. However, on subgeneric level and

below much more study is needed to elucidate the

phylogenetic relationships within the Haliplidae.

For that reason I refrain from dividing the species

in this revision into species-groups. They are

simply treated in alphabetical order.

For many species the type-material was studied.

For some species, of which the status seemed clear

to me, only few or no attempts have been made to

trace and study the type-material. In some other

cases several attempts to locate the types failed.

About 2800 specimens have been studied, mostly

from Western Europe. Eastern and Southern pa-

laearctic material is relatively scarce in the collec-

tions I had access to.

For all species (re)descriptions are provided.

Female genitalia are not described, because they

usually are very uniform and further research is

necessary to check if there are characters reliable

enough to separate related species.

Descriptions of larvae are not included.

METHODS

Before dissecting the aedeagus the beetles are

softened in water with detergent for about 2 hours.

The aedeagus is pulled out with a small hooked

needle and placed into a drop of alcohol. During

dissection the beetle has to be kept wet, to prevent

the severed aedeagus jumping out of sight. In alco-

hol the left- and right parameres are separated

from the penis by using fine needles. When the

alcohol has almost dried up the three parts are

usually glued with Fauré-mixture on the same piece

of cardboard as the beetle. The penis and the par-

ameres are positioned so that they are seen from

the side. The water-soluble Fauré-mixture makes it

possible to loosen the genitalia to study them in an

other position or to make temporary slide mounts.

In some cases the genitalia are transferred into

Euparalona plastic card, covered by a coverslip and

pinned to the original needle. Genitalia should not

be glued to plastic cards with water-soluble glue as

they are liable to become detached in time.

Drawings of genitalia are made by using a Swift

universal microscope with a camera lucida attach-

ment. The other drawings are made by using a

Zeiss binocular with an ocular-micrometer.

Measurements are made by using the ocular-

micrometer of a Zeiss binocular.

Morphological terms are according to figs. 1 and

_ The palaearctic region as treated in this revision

comprises Europe including Iceland, North Africa

north of the Sahara and Asia north of the line

formed by the northern borders of Pakistan, India,

Nepal, Bhutan, Burma, Laos and Vietnam. For

practical reasons Taiwan and China as a whole are

taken into account, although the south-eastern part

has an oriental fauna.

Material of many institutions and private per-

sons has been studied, including a number of prim-

ary types. Locality-names are given, as far as pos-

sible, according to the Times Atlas of the World

(Comprehensive edition, 1983). Deviating names

on original labels are cited, when necessary, in

parentheses. In a number of cases I was not able to

trace the locality, because of the poor or unreadable

labels. Countries are used with their present-day

boundaries.

Distribution-maps are based on material exam-

ined. Reliable literature-records are included, but

using different symbols.

Data on biology are seldom present on labels, so

notes on the biology are mainly based on literature

and the author's experience. If nothing is menti-

oned in the descriptions about biology, such infor-

mation was unknown to the author.

The institutions from which material has been

examined or which are otherwise referred to in the

text have been abbreviated according to Arnett &

Samuelson (1986) with the addition of:

HUJI - IES Laboratory, Hebrew University of Jer-

usalem, Israel.

MSNT - Museo Civico di Storia Naturale, Trieste,

Italy.

MOG - Museum voor het Onderwijs Museon, 's

Gravenhage, Netherlands.

RIN — Rijks Instituut voor Natuurbeheer, Leersum,

Netherlands.

Further material from the following private col-

lections has been studied: Angus, Englefield Green,

VAN VONDEL: Palaearctic Liaphlus

UK; van Berge Henegouwen, Zoetermeer, Ne-

therlands; Cuppen, Ede, Netherlands; Drost,

Wadenoijen, Netherlands; Giessen, Gaanderen,

Netherlands; Hielkema, Gouda, Netherlands;

Huijbregts, Leidschendam, Netherlands; Jongema,

Wageningen, Netherlands; Kanaar, Leiderdorp,

Netherlands; Leblanc, Troyes, France; Matsui,

Hondo City, Japan; Onder de Linden, Krimpen aan

de IJssel, Netherlands; Rotzer, Gampel, Switzer-

land; Schilthuizen, Leiden, Netherlands; Schreijer,

Alkmaar, Netherlands; Sung Hwa Lee, Taegu, Ko-

rea; Vallenduuk, Lelystad, Netherlands; Vondel,

Hendrik Ido Ambacht, Netherlands.

SYSTEMATIC SECTION

Subgenus Liaphlus Guignot

Haliplus sg. Liaphlus Guignot, 1928: 138. Type species:

Dytiscus fulvus Fabricius, 1801, by subsequent desig-

nation (Guignot 1930: 75).

Haliplus sg. Hoplites Kinel, 1929: 219. Preoccupied. [See

remarks under Haliplus laminatus.| Type species: Dy-

tiscus laminatus Schaller, 1783, by monotypy.

Liaphlus Guignot; Guignot 1933: 228, 1947: 48, 1959: 33,

Balfour-Browne 1936: 76, 1938: 12, Csiki 1946: 558,

Zaitsev 1953: 43, Freude 1971: 15, Galewski 1976: 27,

Franciscolo 1979: 108, Vondel 1986: 132, Holmen

1987: 110, Beutel & Ruhnau 1990: 11.

Diagnosis. - The following character set is diag-

nostic for specimens of this subgenus:

1. Pronotum without basal plicae.

2. Metatibia with a setiferous striole on dorsal face.

3. Males: right paramere with a solid digitus. This

is not the case in H. laminatus (see remarks under

that species).

Remarks. - Liaphlus is represented in all faunal

regions. Beutel & Ruhnau (1990) examined a large

number of characters of nine species of Liaphlus

from different faunal regions, but they did not find

evidence for the monophyly of the subgenus.

Within the Haliplidae Liaphlus can be recog-

nised by means of the following keys.

Key to the genera of world Haliplidae

1. Last segment of palpi longer than penultimate

segment. Metacoxal plates at least covering

part of sixth abdominal sternite.................

enna SV Peltodytes Régimbart

— Last segment of palpi clearly shorter than pen-

ultimate segment. Last three abdominal ster-

mites sreely visible: a. RR 2

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

2. Head as wide as pronotum. Elytra strongly

PUNCLULEKATORMALOWSEL a. We

ARES AS PRESS … Algophilus Zimmermann

- Head not as wide as pronotum. Elytra with

GIEATRFOWSTO{IPUNELULESPRBRARA RE E 3

3. Pronotum almost square, front-corners

strongly rounded, elytra usually with longitud-

INA ALES ARE RENEE Brychius Thomson

— Pronotum widest at base, sides converging an-

teriorly, elytra without longitudinal ridges in

ÉirsusinINtervals PE MNT ES ETAPE STO 4

4. Median part of prosternum and base of pro-

sternal process forming a plateau-like eleva-

tion, at least in part angularly separated from

sides of prosternum......... Haliplus Latreille

- Prosternum evenly rounded from side to side

SONA RE Sc Apteraliplus Chandler

Note: Algophilus is restricted to South Africa,

while Apteraliplus is only known from North

America.

Key to the palaearctic subgenera of Haliplus:

1. Dorsal and ventral side of body with clearly

visible (30 x) micropuncturation, on elytra be-

tween small primary punctures..................

LR INI IA DIREI Haliplidius Guignot

— Elytra smooth or with much weaker micro-

puncturation (visible at 50 x) between strong

PEIMaryapUneture OWS et ere 2

2. Hind tibia with a setiferous striole on dorsal

face. Pronotum without basal longitudinal pli-

Carel, Lie Meng Liaphlus Guignot

- Hind tibia without a setiferous striole on dorsal

face. Base of pronotum on both sides with a,

sometimes very short, rarely absent, longitud-

inaljplicamiparzi na: st ren em. 3

3. Pronotal plicae reaching beyond middle and

strongly bent.......... Neohaliplus Netolitzky

— Pronotal plicae not reaching beyond middle,

straight or slightly curved..... Haliplus s.str.

Checklist of palaearctic Haliplus (Liaphlus)

1. H. abbreviatus Wehncke, 1880

2. H. andalusicus Wehncke, 1874

3. H. angusti sp. n.

4. H. astrakhanus sp. n.

5. H. basinotatus Zimmermann, 1924

6. H. chinensis Falkenström, 1932

7. H. dalmatinus J. Miiller, 1900

8. H. davidi sp. n.

9. H. diruptus Balfour-Browne, 1946

0. H. excoffieri sp. n.

—

30.

. H. eximis Clark, 1863

H. modestus Zimmermann, 1924 syn. n.

H. flavicollis Sturm, 1834

H. fulvus (Fabricius), 1801

H. subguttatus Crotch, 1873 syn. n.

H. salinarius Wallis, 1933 syn. n.

. H. gafnyi sp. n.

. H. guttatus Aubé, 1836

H. pelopis Sahlberg, 1907 syn. n.

H. holmeni sp. n.

. H. jaechi sp. n.

H. kotoshonis Kano & Kamiya, 1931

H. kulleri Vondel, 1988

H. laminatus (Schaller, 1783)

. H. maculatus Motschulsky, 1860

H. syriacus Wehncke, 1880 syn. n.

H. zimmermanni Gschwendtner, 1921

syn. n.

H. maculipennis Schaum, 1864

H. mucronatus Stephens, 1828

H. ortali sp. n.

H. ovalis Sharp, 1884

H. pulchellus Clark, 1863

H. rubidus Perris, 1857

H. sharpı Wehncke, 1880

H. variegatus Sturm, 1834

H. transvolgensis Semenov, 1904 syn. n.

H. villiersi Legros, 1972

Key to the palaearctic species of Liaphlus

. Distance between eyes about eye-width (fig.

1705208244) een ee RE 2

Distance between eyes at least 1.2 X eye-

wıdchs3e. au. ali anna ere ee 4

. Prosternal process furrowed on both sides (fig.

DASEIN TR 26.pulchellus

Prosternal process strongly impressed in mid-

dle, at least in apical half (fig. 174)............ 3

. Elytra with basal black band, eighth puncture-

row with about nine small dark blotches.......

ee brite. corte sestri. A 22.maculipennis

Elytra without basal black band, eighth punc-

ture-row not with dense row of dark blotches

zeide arie see PRIE 18.kotoshonis

. Head relatively wide (fig. 217), distance be-

tween eyes at least 2.3 X eye-width. Metaster-

nal process flat in middle. Elytra without dark

blorches sh pere 23.mucronatus

Head narrower. Metasternal process at least

with small pit or groove in middle........... 5

. Sides of prosternal process before coxae ex-

tending forwards as complete plicae to anterior

edge of prosternum (LR) re eer 8

Sides of prosternal process before coxae not

completely extending forwards as plicae to an-

10.

Male

12.

terior edge of prosternum (fig. 116).......... 6

. Plicae on sides of prosternal process widely

interrupted before coxae (fig. 17). Longer ap-

ical spur of hind tibia usually as long as or

longer than first tarsal segment (fig. 15). Elytra

mostly with darker blotches..... 2.andalusicus

Plicae on sides of prosternal process com-

pletely absent before coxae (fig. 116). Elytra

without clear blotches. Apical spurs of hind

tibia not as long as first tarsal segment....... N

. Punctures of primary elytral rows wide in basal

area, distance between rows not more than

diameter of punctures (fig. 255). Distance be-

tween eyes 1.7-1.9 X eye-width..... 27.rubidus

Punctures of primary elytral rows rather small,

distance between rows more than diameter of

punctures (fig. 113). Distance between eyes

ASG) eye width deli oa 12.flavicollis

. Setiferous striole on dorsal face of hind tibia

stretched over apical half (fig. 195). Elytra us-

ually with more or less confluent and inter-

rupted dark lines following rows of dark punc-

tures, but distinct blotches usually absent. Basal

elytral punctures coalescing to form short

grooves (fig. 188). Male: first tarsal segment of

midlegs strongly dilated ventrally (fig. 192).

Female: elytra almost completely covered with

MICrO-PUNELUTESE te ee 20.laminatus

Setiferous striole on dorsal face of hind tibia

usually not longer than % X tibia length, not

reaching apical point. Male: first 3 tarsal seg-

ments of midlegs weakly dilated ventrally.

Female: elytra without clear micropunctua-

OR IA Ae bore brede 9

. Elytra without dark blotches. Pronotal and ely-

tralipunctures black terme tee ll.eximis

Elytra with dark blotches, or when these

blotches are strongly reduced or absent punc-

tures on disc of pronotum are not black ....10

Elytra with basal transverse black band as dark

asisuturalimaculation #7 e. 34

Elytra without basal black band. Sometimes

some transverse darkening separated from ba-

sal margin or a brown basal band, clearly less

dark than sutural maculation.................. 11

Widened basal punctures on pronotum twice

the width of elytral primary punctures (fig.

141). Body-shape parallel or subparallel. Pro-

sternal process narrow, length 2.6 X width,

slightly narrowed near coxae (fig. 145). Elytra

with mostly vague blotches, one large blotch on

middle of suture and three smaller ones on

CACHE ME ONE RARO 15.guttatus

Basal punctures on pronotum usually not twice

as wide as elytral primary punctures. Body not

clearlygparallel ee ea 12

Weng ches CLS Inn RIINA PITTORI 29

16.

. Metasternal

VAN VONDEL: Palaearctic Liaphlus

Eng: 13

. Elytra with longitudinal distinct blotches in

even intervals, blotches not connected to dar-

kenedisutune stp s-te oe eden nale 14

Elytraliblotchesidifferent +... sr... 17

. Metasternal process flat anteriorly. Vertex dar-

kenedi4& eee azar bea 15

Metasternal process grooved anteriorly. Ver-

tex not darkened

. Setiferous striole weak, about Y, X tibia length

(fig. 33). First elytral primary puncture-row

with about 28 punctures. Pronotum anteriorly

partly bordered. Males: left paramere with

solid digitus (fig. 36). at es. 4.astrakhanus

Setiferous striole strong, about % X tibia

length (fig. 202). First elytral primary punc-

ture-row usually with more than 28 punctures.

Pronotum anteriorly not bordered. Males: left

paramere without solid digitus..................

specimens of 21.maculatus

Distance between eyes less than 1.7 X eye-

width. Males: top of penis short (fig.

ON er specimens of 7.dalmatinus

Distance between eyes more than 1.7 X eye-

width. Males: top of penis long, usually slightly

widenedkapicallya(fig 278) ee 2... ee

REN es specimens of 29.variegatus

Elytra with large maculation, discal blotch and

blotch in apical part connected to darkened

suture sl ae chaos. tases eke ER 18

Elytra with usually vague maculation on first

and even intervals, or only darkening on discal

parts of puncture-rows, blotches on apical part

Nnotconnected to suture: ar. sees oe 21

. Length of prosternal process 2.6 X width (fig.

ZI desde EER 3.angust

Length of prosternal process less than 2.4 X

width: mite Aree Er ea. 19

. Prosternal process hardly narrowed near coxae

(fig. 73). Males: left paramere with short dig-

IUSR 8.davidi

Prosternal process clearly narrowed near

coxae. Males: left paramere without digitus 20

. Distance between eyes more than 1.7 X eye-

Width en specimens of 29.variegatus

Distance between eyes less than 1.7 X eye-

wide oon specimens of 9.diruptus

process grooved anteriorly,

weakivyapunctured nace: tener tes le. Sens 22

Metasternal process flat anteriorly........... 23

. Prosternal process impressed posteriorly (fig.

287). Elytral primary puncture-rows sparse,

sutural punctures weak (fig. 285) 30.villiersi

Prosternal process posteriorly at most with

weak transverse impression (fig. 7). Elytral

primary puncture-rows dense, sutural punc-

turesistrongd(fig-)) hanna. l.abbreviatus

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

GOOG

“|

Cee

Sed è d è 80°

5 oe

, locality unknown. - 3, dorsal view; 4, antenna; 5, elytral punctuation; 6, dorsal side

; 7, prosternal process; 8, lateral view of prosternal process; 9, left paramere; 10, penis; 11, right paramere.

Figs. 3-11. Haliplus abbreviatus

of hind tibia;

DIE

28.

29}

30.

31.

32.

WE Malest SEWER EI RER Silent EHE tie suli 24

Female sso ed e ona 27

24. Left paramere without solid digitus 14.gafnyz

Left paramere with solid digitus.............. 25

. Digitus on left paramere long and parallel-

sided. Base of pronotum with about five

stronger punctures on both sides.... 19.kulleri

Digitus on left paramere tapering. Base of

pronotum with almost continuous row of

SCLONDEM PUNGCULES AR ee. 26

. Left paramere with strongly developed taper-

ing digitus, row of long hairs on inner margin

Ofiparamere Kir ne Oars. ere L 17 jaecht

Left paramere with small tapering digitus, at

most some short hairs on inner margin of

Paramere te ois. MESA PE LA ANN! 24.ortali

Metasternal process flat with shallow pit in

middle, strongly and densely punctured, small

punctures between large ones (fig. 136)........

14.gafnyi

Metasternal process with clear pit in middle,

sparsely and moderately strongly punctured

EEE WEES ER RENT IO NAME, OU 28

Basal punctures on pronotum weak and re-

duced in the middle (fig. 179). Prosternal pro-

cess weakly narrowed (fig. 183)..... 19.kulleri

Base of pronotum with continuous row of

strong punctures (fig. 161). Prosternal process

Stronplysnarrowed! (fig IGS e

17.jaechi or 24.ortali

Distance between eyes less than 1.4 X eye-

WALTER A RAET, OPCs LIL OI 2058 30

Distance between eyes more than 1.4 X eye-

Wit ch MEREN epr: Je SMOLIN RICE, 31

Black suture not reaching first secondary punc-

ture-row in basal part (fig. 91). Body strongly

tapering apically. Penis and parameres figs. 99-

NOME SOR EEE A A MEE BAM 10.excoffieri

Black suture reaching first secondary punc-

ture-row in basal part (fig. 48). Body more

oval. Penis and parameres figs. 54-56..........

SRO SO NI 6. chinensis

Elytral discal blotch connected to suture. Penis

and parameres figs. 241-243........... 25.ovalis

Elytral discal blotch not connected to su-

EUDORA ee een ne 32

Length of hind tarsus including claws more

than 1.3 X tibia length. Prosternal process

slightly narrowed near coxae, narrowed part

0.6 to 0.7 X widest part. Punctures of setiferous

striole not connected in a groove. Male: digitus

of right paramere 1 X length of basal part....

large specimens of 7.dalmatinus

Length of hind tarsus less than 1.3 X tibia

length. Prosternal process strongly narrowed

near coxae, narrowed part 0.5 X width of wid-

ESCHPALELIIR sae een 33

33:

34.

39)

36.

VAN VONDEL: Palaearctic Liaphlus

Distance between eyes less than 1.6 X eye-

width. Setiferous striole on dorsal side of hind

tibia not strongly grooved, separate points us-

ually clearly visible. Male: apical part of penis

very narrow and pointed (fig. 205).............

21.maculatus

Distance between eyes more than 1.6 X eye-

width. Setiferous striole on dorsal side of hind

tibia strongly grooved, separate points not or

hardly visible. Male: apical part of penis wide

and rounded at the top (fig. 127).....13.fulvus

Length 3.5-4.1 mm. Black sutural stripe in an-

terior half at most reaching first secondary

PUNCEUIE LOWER RS 5.basinotatus

Length at most 3.6 mm. Black sutural stripe

reaching first primary puncture-row......... 35

Distance between eyes about 1.2 X eye-width.

Prosternal process wide, impressed apically,

weakly narrowed near coxae (fig. 155). Setifer-

ous striole on dorsal side of hind tibia consist-

ing of about 6 isolated points, not grooved (fig.

154). Male: left paramere with solid digitus

(fig. 158). Between fore-claws a small cushion

(OTR DS de A o al 16.holmeni

Distance between eyes 1.3-1.5 X eye-width 36

Fourth and fifth antennal segment about as

long as wide (fig. 263). Punctures in first to

fourth primary elytral rows weaker than in

other puncture-rows. Prosternal process

broad, hardly narrowed near coxae, hardly or

not impressed apically (fig. 266). Anterior

edge of prosternum clearly and completely bor-

RATER eme er TIERRA 28.sharpt.

Fifth antennal segment clearly longer than

wide. Inner primary puncture-rows of elytra

not weaker than outer ones. Prosternal process

strongly narrowed near coxae, strongly im-

pressed apically. Anterior edge of prosternum

only bordered near process. South-east pa-

laearctic or north oriental specimens

OPOLE, NERE TT 9.diruptus

DESCRIPTION OF THE PALAEARCTIC SPECIES OF

LIAPHLUS

1. Haliplus abbreviatus Wehncke

(figs. 3-11)

Haliplus abbreviatus Wehncke, 1880: 73. Lectotype 9

(here designated), ‘Mesopotamia, Sharp’

(MNHN) [examined].

[Iraq]

Haliplus abbreviatus; Zimmermann 1920: 303, 1924:

136, Alfieri 1976: 38.

Diagnosis

This species is closely related to villzerst or even

conspecific. Specimens of vzlliersi have the proster-

| process more impressed posteriorly, the prim-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

08666

Figs. 12-20. Haliplus andalusicus, lectotype. - 12, dorsal view; 13, antenna; 14, elytral punctuation; 15, dorsal side of

hind tibia; 16, prosternal process; 17, lateral view of prosternal process; 18, left paramere; 19, penis; 20, right paramere.

ary elytral puncture-rows sparser and the sutural

punctures weaker.

Description

Length 3.0 mm, width 1.7 mm. Body oval, widest

in the middle (fig. 3).

Head. - Light brown, rather strongly punctured,

more sparsely punctured between the eyes (fig. 3).

Distance between the eyes 1.6 X eye-width. Anten-

nae (fig. 4) and palpi yellow-brown.

Pronotum. - Yellow-brown, base and front mar-

gin slightly darkened. Lateral borders straight to

slightly convex, with fine dark margins. Disc spar-

sely punctured, front margin behind an unpunc-

tured zone with a strongly punctured band, base

strongly punctured, basal puncture-row partly wi-

dened and darkened (fig. 3).

Elytra. - Yellow-brown with in the anterior half

fairly strong and apically weaker primary punc-

ture-rows, punctures brown to almost black (fig. 5).

First primary puncture-row with about 30 punc-

tures. Secondary puncture-rows fine. Brown, some-

times vague markings on the even intervals, on the

apical point and along the apical % of the suture,

median spot connected to the suture.

Ventral side. - Body yellow-brown to yellow-red,

prosternal process partly dark-brown, elytral epi-

pleura yellow, legs yellow-brown with some dar-

kening around the coxae and trochanteres. Proster-

nal process flat, slightly impressed apically,

narrowed near the front coxae, strongly punctured

(fig.7), laterally completely bordered (fig.8). Meta-

sternal process furrowed, deep pit in the middle,

strongly punctured (fig.7). Setiferous striole on

dorsal face of hind tibia with about 6 punctures,

longer of the two tibial spurs almost as long as the

first tarsal segment (fig.6).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 9-11.

Distribution (fig. 305)

Iraq, Iran?. Alfieri (1976) gives Egypt.

Remarks

This species is described here on the basis of a

single female (lectotype) and two male specimens,

which are considered to belong to this species (figs.

3-11). It is probably a very variable species, which

often may be confused with variegatus. There is a

good chance that a number of the middle eastern

specimens, in literature cited as variegatus belong

to abbreviatus.

Material examined. - 3 ex. - Iraq: 1 Q (lectotype). -

Iran(?): 1 6, Turcom, Kurd., 15943, Millingen, Fry Coll.

(BMNH) — Without locality: 1 @, Reitter (NHMW).

VAN VONDEL: Palaearctic Liaphlus

2. Haliplus andalusicus Wehncke

(figs. 12-20)

Haliplus andalusicus Wehncke, 1874: 135. Lectotype &

(here designated), [Spain] ‘Andalusia, Dieck, Dr.

Guignot visité 1925’ (MNHN) [examined].

Haliplus andalusicus, Marseul 1882: 107, Seidlitz 1887:

30, Escalera 1914: 57, Scholz 1916: 17, Zimmermann

1920: 303, 1924: 136, Guignot 1933: 232, 1947: 52,

1959: 35, Burmeister 1939: 214, Lagar Mascaro 1968:

73, Hoch 1972: 251, Ienistea 1978: 294.

Diagnosis

This species is most easily confused with varte-

gatus from which it can be distinguished by the

interrupted lateral plicae of the prosternal process

and the apical spur of the hind tibia being as long

as or longer than the first tarsal segment.

Description

Length 2.5-3.0 mm, width 1.4-1.7 mm. Body

oval, widest in the middle (fig. 12).

Head. - Yellow-brown with usually brown ver-

tex, sparsely punctured. Distance between the eyes

1.9-2.0 X eye-width. Antennae (fig. 13) and palpi

yellow to yellow-brown.

Pronotum. - Yellow to yellow-red. Lateral

borders straight to slightly convex, finely bordered.

Sparsely to rather strongly punctured, basal punc-

tures slightly widened and darkened (fig. 12).

Elytra. - Yellow to yellow-red, primary punc-

ture-rows rather weak with exception of the

stronger punctures in the anterior half of the first

three rows, 32-36 punctures in the first row (fig.

14). Secondary punctures weak. Darkening of the

punctures not always visible. Brown to almost in-

visible markings on the even intervals, on the ap-

ical point and along the apical % of the suture, the

median spot may be connected to the suture.

Ventral side. - Body yellow to yellow-brown,

elytral epipleura yellow, legs yellow-brown. Pro-

sternal process impressed in the middle, strongly

punctured, narrowed near the coxae (fig. 16), lat-

eral plicae interrupted before the coxae (fig. 17).

Metasternal process grooved with a pit in the mid-

dle to largely impressed, sparsely punctured (fig.

16). Setiferous striole on dorsal face of hind tibia

with 4-8 punctures, longer of the two tibial spurs

as long as or longer than the first tarsal segment

(fig. 15).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 18-20.

Biology

In fresh and especially brackish stagnant water

in pools near the coast and more rarely inland.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 21-29. Haliplus angusi, holotype. - 21, dorsal view; 22, antenna; 23, elytral punctuation; 24, dorsal side of hind

tibia; 25, prosternal process; 26, lateral view of prosternal process; 27, left paramere; 28, penis; 29, right paramere.

Distribution (fig. 298)

South of France, Spain (Burmeister 1939, gives

The Baleares), Morocco, Algeria, Tunisia.

Material examined. — 38 ex. - France: 1 9, La Capelle

[dep. Lozère], 30.vi/iv.1987 (Leblanc); 6 ex., Albaron,

Camargue, A. Chobaut; 1 ex., Montpellier, Et. Laragne,

V.M. Duchon (MNHN). — Spain: 1 @ (lectotype); 1 &

(paralectotype), Malaga, Sammlung Cl. Müller (zsMC); 1

ex., Gibraltar; 1 ex., Palencia (BMNH). - Morocco: 4 8,

Tanger, Rolph (BMNH). - Algeria: 2 ex., Massif des Mou-

zaia (BMNH); 4 ex., Philippeville, A. Thery (NHMw,

MNHN); 1 ex., Biskra, surroundings, de Vauloger; 1 ex.,

Biskra, Mertu[?]; 2 ex., no further data (NHMW).-Tunisia:

2 8, Gafsa (NHMW); 2 6, 1 9, 3 ex, surroundings Gafsa,

de Vauloger (NHMW, MNHN); 2 ex., Southern Tunisia,

ili.iv.1925, ex coll. J. Omer Cooper (BMNH).-Locality un-

known: 2 @ (paralectotypes), ex. coll. Wehncke (MNHN).

3. Haliplus angusi sp. n.

(figs. 21-29)

Type material. - Holotype 6, USSR, Siberia, 28.v-

28.vi.1982, Novosibirsk oblast, Karasuk, 53.5 N, 78 E, R.

B. Angus (ZMAS).

Diagnosis

This species can be distinguished from related

ones by the very narrow prosternal process. Possi-

bly there is material in collections representing this

species, but identified as H. maculatus Mot-

schulsky.

Description

Length 3.3 mm, width 1.7 mm. Body oblong,

subparallel, widest in the middle (fig. 21).

Head. - Yellow-red, dark mark on vertex, weakly

and sparsely punctured. Distance between the eyes

1.6 X eye-width. Antennae yellow (fig. 22), palpi

yellow.

Pronotum. - Yellow to yellow-brown on the disc.

Lateral borders straight, lateral margin clearly

finer anteriorly. Strongly darkened, occasionally

widened punctures along the base, anteriorly

rather weakly punctured, otherwise almost un-

punctured. Along the central part of the base

slightly impressed (fig. 21).

Elytra. - Pale yellow, primary puncture-rows

moderately strong, about 33 punctures in the first

row (fig. 23). Secondary punctures in the first in-

terval almost as strong as the primary punctures,

other secondary punctures rather weak, accompan-

ied by small punctures (fig. 23). Extensive mark-

ings: a large arrow-shaped blotch on the disc, on

the suture and the apex, often connected oblong

blotches in the even intervals. The basal margin,

normally covered by the pronotum, yellow-brown

till the fifth striae (fig. 21).

VAN VONDEL: Palaearctic Liaphlus

Ventral side. - Yellow to yellow-red, elytral epi-

pleura pale-yellow, abdominal segment brown bor-

dered, legs yellow with some darkening near the

coxae. Prosternal process narrow, flat, narrowed

near the coxae, coarsely punctured (fig.25), lateral

plicae complete (fig. 26). Metasternal process flat

with a pit in the middle, sparsely punctured (fig.

25). Coxal plates sparsely and weakly punctured,

near the suture almost unpunctured. Setiferous

striole on dorsal side of hind tibia over % of the

tibia length, with about 10 punctures, longer of the

two tibial spurs % X the length of the first tarsal

segment (fig. 24).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 27-29.

Etymology. - This species is named after the

collector Dr. Robert Angus.

Distribution (fig. 304)

Only known from the type-locality: USSR, Sibe-

ria, Novosibirsk oblast, Karasuk.

4. Haliplus astrakhanus sp. n.

(figs. 30-38)

Type material: Holotype 8, [ USSR, Astrakhan] ‘As-

tracan, Koltze’ (MNHN, Wehncke collection).

Diagnosis

This species is closely related to ortalz, but in the

males the top of the penis is narrower and more

pointed.

Description

Length 3.4 mm, width 1.9 mm. Body oval, widest

in the middle (fig. 30).

Head. - Red-brown, weakly punctured, vertex

little stronger punctured. Distance between the

eyes 1.5 X eye-width. Antennae (fig. 31) and palpi

yellow-red.

Pronotum. - Yellow-red, slightly darker on the

disc and along the base. Lateral borders straight,

finely bordered. Anteriorly partly bordered. Width

1.8 X the length in the middle. The middle of the

base slightly impressed. Weakly punctured, along

the base a row of stronger not or hardly darkened

punctures (fig. 30).

Elytra. - Yellow-red, many longitudinal vague

blotches on the even intervals and along the suture

(fig.30). Primary puncture-rows moderately

strong, about 28 punctures in the first row. Secon-

dary punctures weak, accompanied by very small

ones (fig. 32). All punctures darkened.

Ventral side. - Yellow-red to red-brown, elytral

epipleura yellow, legs yellow-red, darkened to-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

alia da eel aidan ai

> 2888

Figs. 30-38. Haliplus astrakhanus, holotype. - 30, dorsal view; 31, antenna; 32, elytral punctuation; 33, dorsal side

of hind tibia; 34, prosternal process; 35, lateral view of prosternal process; 36, left paramere; 37, penis; 38, right

paramere.

wards the coxae. Prosternal process flat, moder-

ately strongly punctured, slightly narrowed near

the coxae (fig. 34), lateral plicae complete (fig. 35).

Metasternal process with a pit in the middle,

weakly punctured (fig. 34). Setiferous striole '/, X

the tibia length, with about 6 weak punctures,

longer of the two tibial spurs 24 X the length of the

first tarsal segment (fig. 33).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and paramere as in figs. 36-38,

left paramere with a small solid parallel digitus, top

of penis narrow and a little pointed.

Etymology. - Named after the type locality.

Distribution (fig. 304)

Only known from the type-region Astrakhan in

the south-west of the USSR.

5. Haliplus basinotatus Zimmermann

(figs. 39-47)

Haliplus basinotatus Zimmermann, 1924: 137. Lectotype

& (here designated), [USSR] ‘Nikolsk Ussurijsk, Us-

suri geb. Mandl, R. Mouchamps det. 62, Haliplus ba-

sinotatus Zimm., Cotype’ (ISNB) [examined].

Haliplus basinotatus ssp. latiusculus Nakane, 1985a: 63.

Holotype @, Yoshii, Fukuoka Pref., Kyushu, Japan,

10.vii.1957, N. Gyotoku leg. [not examined].

Haliplus basinotatus; Zimmermann 1924: 137, Zaitsev

1953: 75, Nakane 1985a: 63.

Haliplus basinotatus ssp. latiusculus, Nakane 1987: 30.

Diagnosis

This species can be distinguished from related

species by the dark basal band on the elytra and its

large size. The Japanese subspecies latiusculus

should differ from the continental form by having

‘the front margin of pronotum not or scarcely

rounded-produced forwards and the punctuation

and elytral pattern appearing somewhat different’

(Nakane 1985a).

Description

Length 3.5-4.0 mm, width 1.9-2.1 mm. Body

widest just behind the well pronounced shoulders

(fig. 39).

Head. - Yellow to yellow-red, vertex usually dar-

kened and nearly unpunctured, frons punctured.

Distance between the eyes 1.3-1.4 X eye-width.

Antennae (fig. 40) and palpi yellow-brown.

Pronotum. - Yellow to yellow-red, front margin

somewhat darkened or with a small brown blotch.

Lateral borders slightly concave to slightly convex

with very fine margins. In the anterior part with

strong and dense punctures, basal punctures not

wider than elytral punctures (fig. 39).

VAN VONDEL: Palaearctic Liaphlus

Elytra. — Yellow to yellow-red with distinct

brown to nearly black markings on the intervals,

along the suture, on the apex and along the base

until the sixth puncture-row. The median spots are

connected to the suture, but other spots can also

show connections (fig. 39). Primary punctures

strong, 28-31 punctures in the first row. Secondary

punctures fairly strong. All punctures darkened

(fig. 41).

Ventral side. - Body yellow to yellow-red, legs

yellow to yellow-red, darkened towards the brown

trochanteres and coxae. Prosternal process flat,

strongly and densely punctured and narrowed near

the coxae (fig. 43), laterally with complete plicae

(fig. 44). Metasternal process flat, in the middle

behind the coxae with a shallow pit (fig. 43), mod-

erately punctured. Setiferous striole on dorsal face

of hind tibia with 11-20 punctures, sometimes in a

strong groove making the separate punctures

hardly recognizable, longer of the two tibial spurs

about 24 X the length of the first tarsal segment

(fig. 42).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 45-47.

Distribution (fig. 311)

Easternmost parts of the USSR (near Vladivos-

tok), North Korea, Japan (ssp. latiusculus, accord-

ing to Nakane 1985).

Material examined. - 5 ex.- USSR: 1 @, 1 Q (lectotype

and paralectotype) (ISNB). - North Korea: 29,

29.vii.1956, M. Magyar, Ch'ongjin (Tshondtsin) (ISNB); 1

ex., Tumen-ula, Russia/Korea border, 14.vii.1913

(ZMUC).

6. Haliplus chinensis Falkenstròm

(figs. 48-56)

Haliplus chinensis Falkenstròm, 1932: 191. Holotype 9,

‘China, N. O. Sichuan, Sven Hedins Exp. Ctr. Asien,

Dr. Hummel, 22/5, Typus, Hal. chinensis n. sp. Typ.

det. Falkenström’ (NHRS) [examined].

Haliplus chinensis; Falkenstròm 1933: 4 (again described

as n. sp.), Zaitsev 1953: 71 (as ovalis) and 76 (as

chinensis).

Diagnosis

This species is closely related to ovalis. It seems

that ovalis is restricted to Japan, while the contin-

ental specimens, usually considered to belong to

ovalis, seem to be more related to chinensis, al-

though only the female holotype of the latter spe-

cies has the prosternal process slightly impressed.

Despite the fact that all ovalis-like specimens that

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

ee 46

eae

e a 88

Figs. 39-47. Haliplus basinotatus, lectotype. - 39, dorsal view; 40, antenna; 41, elytral punctuation; 42, dorsal side of

hind tibia; 43, prosternal process; 44, lateral view of prosternal process; 45, left paramere; 46, penis; 47, right paramere.

I have seen have a flat prosternal process I think

that the continental specimens belong to chinensis.

The division between the two is mainly based on

the smoothly curved penis in the continental spe-

cimens, while the specimens from Japan have the

penis abruptly bent in the apical part.

Description

Length 3.9-4.2 mm, width 2.1-2.2 mm. Body

oval, widest in the middle (fig. 48).

Head. - Yellow to yellow-red, dark marking on

vertex, weakly punctured, on the vertex with

strong punctures. Distance between the eyes 1.4-

1.5 X eye-width. Antennae (fig. 49) and palpi yel-

low-red.

Pronotum. - Yellow, lateral borders straight,

finely bordered. Densely, on the disc sparsely punc-

tured, basal punctures widened and darkened,

sometimes only slightly.

Elytra. - Yellow. Suture, apical point and 9 or 10

blotches dark, central blotch on first and third in-

terval sometimes connected to the suture (fig. 48).

Primary punctures moderately strong, about 32-36

punctures in the first row. Secondary punctures

fairly strong, apically almost as strong as the prim-

ary punctures. All punctures darkened (fig. 50).

Ventral side. - Body yellow to yellow-red, elytral

epipleura yellow, legs yellow-red, slightly darkened

towards the coxae. Prosternal process flat to clearly

depressed in the middle of the apical part, strongly

and densely punctured, narrowed near the coxae

(fig. 52), lateral plicae complete (fig. 53). Metaster-

nal process flat with a pit in the middle, rather

weakly punctured (fig. 52). Setiferous striole on

dorsal face of hind tibia about half of the tibia

length, consisting of about 14 punctures, longer of

the two tibial spurs % X the length of the first

tarsal segment (fig. 51).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 54-56.

Distribution (fig. 309)

China, provinces of Sichuan, Shanxi, Jiangsu,

Yunnan.

Remarks

Sato (1984) mentioned a subsp. sinicus Balfour-

+ Browne, but the description is unknown to me.

Material examined. — 18 ex.- China: 1 9, Holotype 9.

(NHRS); 1 8, 1 9, Nanjing, v.1928, J. G. Needham (ISNB);

1 6, 1 9, (Kiangau), C. E Wu (OLML); 1 6, 1 9, Lex.

Shanghai; 1 ex., Yunnan Sen, Excoffier, 1898 (MNHN); 4

8, Kelan Pt. (1 ex. labelled as type of subsp. sinicus

Balfour-Browne) (BMNH, MNHN); 5 ex., Xinjiang (Chin-

kiang) (SMFD).

VAN VONDEL: Palaearctic Liaphlus

7. Haliplus dalmatinus Miller

(figs. 57-69)

Haliplus dalmatinus Müller, 1900: 113. Lectotype & (here

designated), [ Yugoslavia, Knin] ‘Kistanje, Müller D.,

Museo Civico di Trieste’ (MSNT) [examined].

Haliplus dalmatinus var. Weberi Miiller, 1900: 114. Lec-

totype @ (here designated), [Yugoslavia] 'Mostarsko

blato, TYPUS, Museo Civico di Trieste’ (MSNT)

[examined ].

Haliplus dalmatinus, Zimmermann 1920: 307, 1924: 37,

Bertrand 1928: 87, Scholz 1929: 16, Guignot 1933:

241, Burmeister 1939: 212, Zaitsev 1953: 71, Gueor-

guiev 1958: 44, Franciscolo 1972: 63, Ienistea 1978:

294.

Haliplus dalmatinus var. Webert, Zimmermann 1920:

308, 1924: 37, Bertrand 1928: 87, Scholz 1929: 16,

Guignot 1933: 241, 1947: 51.

Diagnosis

This species is closely related to fulvus, but the

latter has shorter metatarsi, a more narrowed pros-

ternal process, in the males a longer digitus on the

right paramere and generally a larger size.

Description

Length 3.2-3.9 mm, width 1.8-2.2 mm. Body

oval, sides somewhat parallel, widest in the middle

(fig. 57).

Head. - Yellow-red with dark vertex, weakly and

sparsely, behind the eyes stronger and more den-

sely punctured. Distance between the eyes 1.4-1.7

X eye-width. Antennae (fig. 58) and palpi yellow-

red.

Pronotum. - Yellow, lateral borders straight to

slightly convex, finely bordered, except in the mid-

dle fairly strongly punctured, basally with a row of

widened punctures, basal and discal punctures dar-

kened.

Elytra. - Yellow with variable pattern of brown

blotches in the even intervals, sometimes con-

nected to each other, most of the suture and apex

darkened (figs. 57, 66-69). Fairly strong primary

puncture-rows, about 30 punctures in the first row.

Secondary punctures relatively strong. All punc-

tures darkened (fig. 59).

Ventral side. - Body yellow-red, elytral epipleura

yellow, legs yellow-red to brown towards the coxae.

Prosternal process flat, rather strongly punctured,

narrowed near the coxae (fig. 61), lateral plicae

complete (fig. 62). Metasternal process flat or

slightly grooved, shallow pit in the middle, sparsely

punctured (fig. 61). Setiferous striole on dorsal face

of hind tibia with 4-7 punctures, longer of the two

tibial spurs % to 24 X the length of the first tarsal

segment (fig. 60). Length of hind tarsus 1.3-1.5 X

the tibia length, first tarsal segment 1.6-1.7 X the

length of the second segment.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 48-56. Haliplus chinensis (48-53, holotype; 54-56, from Nanjing). - 48, dorsal view; 49, antenna; 50, elytral

punctuation; 51, dorsal side of hind tibia; 52, prosternal process; 53, lateral view of prosternal process; 54, left

paramere; 55, penis; 56, right paramere.

VAN VONDEL: Palaearctic Liaphlus

UU GOA

159

Figs. 57-65. Haliplus dalmatinus, lectotype. - 57, dorsal view; 58, antenna; 59, elytral punctuation; 60, dorsal side of

hind tibia; 61, prosternal process; 62, lateral view of prosternal process; 63, left paramere; 64, penis; 65, right paramere.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

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Figs. 66-69. Haliplus dalmatinus, elytra. - 66, from Kistanje; 67, from Zara; 68, from Knin; 69, from Skutari.

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 63-65, length of distal process of right para-

mere |. X the length of the basal part (fig. 65).

Distribution (fig. 301)

Along the coast of Yugoslavia and Albania, on

the island Corfu (Greece), Turkey, Italy? (identifi-

cation not sure).

Remarks

Since the description of this species various au-

thors have regarded it as a variety of fulvus. Some

authors considered dalmatinus a variety or subspe-

cies of variegatus. Although in some cases dalma-

tinus is very hard to distinguish from small spe-

cimens of fulvus I consider dalmatinus a separate

species and I regard the var. weberi as an occasional

aberration. The elytral maculation is very variable

and ranges from almost absent to very extended.

One of the syntypes of Haliplus zimmermanni

from Italy might belong to this species.

Specimens from Zadar (Zara), labelled as type, are

not considered to belong to the syntype-series as

this locality is not mentioned in the original des-

cription.

Material examined. - 36 ex. - Yugoslavia: 34, 29 (lec-

totype and paralectotypes); 3 ex., surr. of Zadar (Zara),

Müller, (1 labelled as Typus); 2 ex., Krk (Veglia), Neto-

litzky; 1 ex., Dalmatia, Rab (Arbe), 20.ix.1910 (MSNT); 1

ex. (paralectotype), Knin (Kistanje), Miiller D., Dal-

matin det. J. Müller m, Cotypus (ZSMC); 1 & (paralecto-

type), Knin D., Müller, Cotypus; 1 ex., surr. Zadar (Zara),

Müller, Cotypus; 1 6, Dalmatia; 1 @, Dalmatia, Omis

(Almissa), v.1911, Schatzm.; 3 ex. (lectotype and paralec-

totypes of dalmatinus var. weberi), Mostarsko blato, Ty-

pus, Haliplus v. Weberi (MSNT); 1 ex., Sinj, Dalmatia

(NHMW). — Albania: 2 4, Shkodér (Skutari), I. See-Ufer;

2 &, Vraka near Shkodër (NHMW). - Greece: 1 ex., Kérkira

(Corfu), J. Sahlb., 1035, syntype Haliplus leopardinus

Sahlberg (1900) (zsMc). - Turkey: 3 6, 2 9, Yüksekova,

3.vi.1987, M. Jäch; 5 6, Kastamonu, 25.vi.1989, M. Jäch

(NHMW). — Italy : 1 9 (syntype of Haliplus zimmermanni

Gschwendtner, 1921), Salo, 23.v.1909, Type 1921, det. L.

Gschw., Type, Haliplus ?dalmatinus Müller det. B. J. v.

Vondel 1987 (ZSMC).

8. Haliplus davidi sp. n.

(fig. 70-78)

Type material: Holotype &, [China, Beijing] ‘Nord

Pekin, A. David, 1865, Museum Paris, Haliplus Sharpi

Westw., M. Régimbart det. 1898, 227.65 [round label]

(MNHN); Paratype 9, China, Yunnan (MNHN); Paratype

8, Beijing (Peping), CF. Wu (OLML); Paratype 4, Hei-

longjiang (Mandsch.), Harbin, 28.ix.1930, v. Jettmer

(ZMHB).

Diagnosis

Male specimens can be distinguished from di-

ruptus by the presence of a digitus on the left

paramere. I am not always able to separate the

females of these species.

VAN VONDEL: Palaearctic Liaphlus

Figs. 70-78. Haliplus davidi, holotype. - 70, dorsal view; 71, elytral punctuation; 72, dorsal side of hind tibia; 73,

prosternal process; 74, lateral view of prosternal process; 75, hind coxal plate; 76, left paramere; 77, penis; 78, right

paramere.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 79-87. Haliplus diruptus, holotype. - 79, dorsal view; 80, antenna; 81, elytral punctuation; 82, dorsal side of hind

tibia; 83, prosternal process; 84, lateral view of prosternal process; 85, left paramere; 86, penis; 87, right paramere.

Description

Length 3.2 mm, width 1.7 mm. Body oval, widest

in the middle (fig. 70).

Head. - Yellow-brown, brown blotch on vertex,

densely punctured, unpunctured band between the

hind parts of the eyes. Distance between the eyes

1.3 X eye-width. Antennae yellow (missing in ho-

lotype), palpi yellow-brown.

Pronotum. - Yellow-brown. Lateral borders

straight, clearly bordered. Width at the base 2.0 X

the length in the middle and 1.6 X the width in

front. Weakly and densely, along the base more

strongly punctured, some basal punctures dar-

kened. Beside the middle of the base slightly im-

pressed (fig. 70).

Elytra. - Yellow-brown, with darkened suture

and apex and several blotches (fig. 70). Primary

puncture-rows moderately strong, wider near the

base, basal punctures in the second to fourth row

widened, about 35 punctures in the first row. Se-

condary punctures moderately strong, in some in-

tervals very sparse, accompanied by very small

punctures (fig. 71), all punctures darkened.

Ventral side. - Yellow-red to yellow-brown, ely-

tral epipleura yellow, legs yellow-brown, some dar-

kening towards the coxae. Prosternal process at the

apex strongly impressed, coarsely punctured, pos-

teriorly little narrower than anteriorly, slightly

narrowed near the coxae (fig. 73), lateral plicae

complete (fig. 74). Metasternal process grooved

and strongly impressed in the middle, rather

strongly punctured (fig. 73). Coxal plates strongly

and densely punctured, small punctures between

the normal ones, which are provided with hairs,

along the suture with a weakly punctured area,

apical parts of the plates dilated (fig. 75). Setiferous

striole 4 of the tibia length, with about 5 punc-

tures, longer of the two tibial spurs % the length

of the first tarsal segment (fig. 72).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 76-78,

left paramere with a short solid digitus.

Etymology. — Named after the collector A.

David.

Distribution (fig. 312)

Only known from China: Yunnan, Beijing and

Harbin.

9. Haliplus diruptus Balfour-Browne

(figs. 79-90)

[Haliplus simplex sensu Régimbart 1899, nec Clark 1863.

Misidentification].

Haliplus diruptus Balfour-Browne, 1946: 436. Holotype

VAN VONDEL: Palaearctic Liaphlus

8 ‘China, Tientsin’ [Tianjin] ‘EF. M. Thomson, 1904-

229, Haliplus simplex Rég.-Zimm., Haliplus diruptus

nom. nov. J. Balfour-Browne det., Type’ (BMNH)

[examined].

Haliplus simplex; Zimmermann 1924: 136, Takizawa

1931: 142, Zaitsev 1953: 74, Sato 1960: 252, 1984: 2,

Cho 1969: 167, Yoon 1988: 621.

Haliplus diruptus; Guignot 1954: 563.

Diagnosis

A very variable species. Specimens with and

without black blotches or bands on the base of the

elytra and specimens with weak or with very strong

and close punctures on the head are considered to

belong to this species. Specimens with a very re-

duced and weakened maculation resemble davidi,

from which males can be distinguished by the lack

of a small solid digitus on the left paramere, but

distinguishing females can be problematic.

Description

Length 2.8-3.5 mm, width 1.6-2.0 mm. Body

oval, widest in the middle (fig. 79).

Head. - Yellow to yellow-brown, vertex slightly

darkened, puncturation weak and fairly sparse to

strong and dense. Distance between the eyes 1.3-

1.5 X eye-width. Antennae (fig. 80) and palpi yel-

low-red.

Pronotum. - Yellow to yellow-red. Lateral

borders straight to slightly convex, finely bordered.

Punctures moderately strong to strong along the

base and in the anterior central part, basal and

anterior punctures darkened, remaining punctures

sometimes darkened.

Elytra. - Yellow to yellow-brown, variable mac-

ulation: darkening along suture and on apex, a

central blotch on the suture and small blotches on

the even intervals; inextreme forms the maculation

can be very extended and includes a large blotch or

even a complete band along the base (figs. 88-90).

Primary punctures moderately strong, first row

slightly weaker than second to fourth or even fifth

row, 30-40 punctures in the first row. Secondary

punctures moderately strong, rows consisting of

normal and very small ones, sometimes very strong

along suture. All punctures darkened (fig. 81).

Ventral side. - Body yellow to yellow-brown,

elytral epipleura yellow, legs yellow to yellow-

brown, slightly darkened near the coxae. Proster-

nal process clearly impressed in the apical part,

strongly narrowed near the front coxae (fig. 83),

lateral plicae complete (fig. 84), fairly strongly

punctured. Metasternal process flat with a pit in

the middle, sparsely punctured (fig. 83). Setiferous

striole on dorsal face of hind tibia about % X the

tibia length, containing about 8 punctures, longer

of the two tibial spurs about % X the length of the

first tarsal segment (fig. 82).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 88-90. Haliplus di-

ruptus, elytra. - 88, from

Beijing; 89,90, from Ton-

kin.

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Se Os

Male sexual characters. — First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 85-87.

Distribution (fig. 311)

Easternmost part of China, provinces of Hebei,

Heilongjiang, Hunan and Jiangsu. Extreme south-

eastern part of the USSR: Maritime Territory.

Vietnam. Guignot (1954) gives Burma, Shan State.

Material examined. — 81 ex. - China: 1 &, Holotype; 1

Q (paratype), Tianjin (Tientsin), vi.1902, F. M. Thom-

son; 1 6, Harbin, 25.viii.1952 (BMNH); 1 Ó, 1 9, Beijing

(Peping), C. E Wu (oLML); 1 6, 1 9, Tianjin (Tientsin)

(SMED); 1 ex., Tianjin (Tien-Tsin); 1 ex., Beijing (Nord

Pekin), A. David 1865; 4 ex., Shanghai; 1 ex., Hengshan

(Hontsang), Weber; 1 @, (Zi-ka-wei) (MNHN); 1 ex,

Yangzhou (Yang Chow Ku), 20.vi.1947, Sunabricotier

(ISNB); 1 &, Beijing (Peping), C. F Wu (OLML). - Korea:

1 ex., 11.vii.1956, M. Magyar, Sariwon, Type, Haliplus

sharpi var. homologus n. var. cf. texte, R. Mouchamps

det. 63 [unpublished name] (ISNB). — Vietnam: 62 ex.,

Tonkin, Hoa Binh region, vii.1940 (part of material), A.

de Cooman (ISNB, MNHN). — Without locality: 1 ex.

(MNHN).

10. Haliplus excoffieri sp. n.

(fig. 91-101)

Type material: Holotype g, [China] ‘Yunnan-Sen, Mgr

Excoffier, 1898, Dr. Régimbt vidit 1898, Museum Paris

ex coll. Oberthiir’. Paratypes: 1 6,3 9, same data (MNHN).

Diagnosis

This species is closely related to chimensis and

ovalis, but the distance between the eyes is usually

less than 1.4 X eye-width, the darkened suture is

only reaching to the first secondary puncture-row

and in the males the penis is clearly widened before

the apex.

Description

Length 3.7-3.8 mm, width 1.9-2.0 mm. Body

oblong oval, tapering apically, widest in or just

before the middle (fig. 91).

Head. - Brown, dark marking on vertex, rather

densely punctured, unpunctured band between the

hind parts of the eyes. Distance between the eyes

1.2-1.4 X eye-width. Antennae yellow to yellow-

brown (fig. 92), palpi yellow-brown.

Pronotum. - Yellow to yellow-brown, lateral

borders straight, clearly bordered. Width at the

base about 2.0 X the length in the middle and 1.6

X the width in front. Sparsely, anteriorly densely,

basally strongly punctured, on the disc almost un-

punctured. About ten of the darkened basal punc-

tures widened, sometimes slightly impressed along

the central part of the base (fig. 91).

Elytra. - Yellow to yellow-brown, suture dar-

kened until the secondary puncture-row, dark

blotches on the apex and in mainly the even inter-

vals. Primary puncture-rows moderately strong,

second to fourth puncture-row getting stronger

basally, 32-36 punctures in the first row. Secondary

puncture-row along the suture rather weak, but

very dense in the apical half, on the other intervals

sometimes relatively strong (fig. 93). All punctures

darkened.

Ventral side. - Yellow to yellow-red, elytral epi-

pleura yellow, legs yellow-brown, darkened to-

wards the coxae. Prosternal process slightly con-

vex, strongly narrowed near the coxae, densely

punctured (fig. 96), lateral plicae complete (fig.

97). Metasternal process flat, strong impression in

the middle, weakly punctured (fig. 96). Coxal plates

strongly punctured, along the suture weakly punc-

tured (fig. 98). Hind margins of fifth and sixth

abdominal segment with complete double punc-

ture-rows (fig. 98). Setiferous striole almost half of

the tibia length, with about 10 groove-like con-

nected punctures, longer of the two tibial spurs %

the length of the first tarsal segment, tibia 3 X the

length of the first tarsal segment (fig. 95).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

the ventral side. Penis and parameres as in figs. 99-

101.

Etymology. — This species is named after the

collector Mgr J. C. Excoffier.

Distribution (fig. 312)

Only known from China, Yunnan-province.

11. Haliplus eximis Clark

(figs. 102-110)

Haliplus eximis Clark, 1863: 418. Syntypes, Korea, Clark

(depository unknown; probably in SAMA or MAMU).

Haliplus modestus Zimmermann, 1924: 139. Holotype

6, ‘Sumatra, Type, Holotype Staatssamml. Miinchen’

(ZSMC) [examined]. Syn. n.

Haliplus hiogoensis Kano & Kamiya, 1931: 1. Types

burned during World War II (Dr. T. Nakane, pers.

comm.) [not examined].

Haliplus emmerichi Falkenstròm, 1936: 79. Holotype 9,

[Kangdin, Sichuan] ‘Nitou Tatsienlu, Szechuan China

, Typus, Haliplus Emmerichi Falk. 9, det. Falken-

ström’ (NHRS) [examined].

[Haliplus flavicollis; auctt., nec Sturm 1834: Wu 193:342.

Misidentification |.

Haliplus eximis; Zimmermann 1920: 305, 1924: 132,

Takizawa 1931: 140, Guignot 1950: 97, Zaitsev 1953:

77, Cho 1969: 166, Sato 1984: 2, Nakane 1985a: 63,

1985b: 40, 1987: 29, Yoon 1988: 623.

Haliplus modestus, Zimmermann 1927: 2.

Haliplus hiogoensis; Sato 1984: 2, Nakane 1985b: 40.

Haliplus emmericht, Guignot 1950: 97, Zaitsev 1953: 69,

Sato 1984: 2.

Diagnosis

This species can be distinguished from related

species in the East Palaearctic and Oriental region

by the total lack of maculation between the elytral

puncture-rows. It resembles the West Palaearctic

flavicollis, which however lacks the lateral plicae in

the anterior part of the prosternal process.

Description

Length 3.4-3.9 mm, width 1.9-2.1 mm. Body

oval, widest in or just before the middle (fig. 102).

Head. - Yellow-brown to rust-coloured, mode-

VAN VONDEL: Palaearctic Liaphlus

rate weakly punctured. Distance between the eyes

1.3-1.4 X eye-width. Antennae (fig. 103) and palpi

yellow-brown.

Pronotum. - Yellow to rust-coloured. Lateral

borders straight to slightly convex, bordered, ba-

sally wider than base of elytra, strongly tapering

anteriorly. Width 2.1-2.2 X the length in the mid-

dle. With strong punctures except on the unpunc-

tured disc, basal punctures partly widened, basal

and anterior punctures darkened.

Elytra. - Yellow to rust-coloured. Strong primary

puncture-rows, about 32 punctures in the first row.

Secondary punctures relatively strong, as dense and

apically almost as strong as the primary punctures

(fig. 104). Except for the partly darkened suture, no

markings present.

Ventral side. - Yellow, sternites yellow-brown,

legs yellow-brown to rust-coloured. Prosternal

process broad and flat, slightly narrowed near the

coxae, strongly punctured (fig. 106), lateral plicae

complete (fig. 107). Metasternal process flat with

a pit in the middle, sparsely punctured (fig. 106).

Setiferous striole on dorsal face of hind tibia with

about 12 punctures, longer of the two tibial spurs

% X the length of the first tarsal segment (fig. 105).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 108-110.

Distribution (fig. 308)

China (prov. Sichuan, Xinjiang, Zhejiang, Zait-

sev (1953) gives Shanghai), Korea, Japan (Hon-

shu, Kyushu), Vietnam, Sumatra.

Remarks

The specimen from China, present in BMNH and

labelled as Type of eximis, does not belong to

Clark’s two syntypes, described from Korea. The

holotype of modestus, described from Sumatra is

here considered conspecific with exzmzs.

Material examined. - 16 ex. - China: 1 4, Type

(BMNH); 1 ex., Kiangau, C. FE Wu (OLML); 1 9, Holotype

Haliplus emmerichi (NHRS); 4 ex., North-west. China,

Xinjiang (Chinkiang), Cotypus (SMFD); 1 6, Zhejiang,

Zhoushan (Chekiang, Chusan), 15.iv.1931 (MNHN). — Ja-

pan: 10 ex., Aobadai, Aira I, Kagoshima Pref., 26.ix.1985.

E. Matsui (Vondel). - Korea: 1 ex., Tshou-Bon San,

1.vii.1956, M. Magyar (ISNB). - Vietnam: 6 ex., Tam Dao,

Tonkin, H.Perrot; 1 ex., Tonkin, Hoa Binh, iii.1940, A. de

Cooman (ISNB). - Indonesia: 1 9, holotype of modestus

(ZSMC).

12. Haliplus flavicollis Sturm

(figs. 111-119)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

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punctuation; 94, fore claws; 95,

prosternal process; 98, hind coxal plate and sternites;

91, dorsal view; 92, antenna; 93, elytral

; 97, lateral view of

; 96, prosternal process

; 100, penis; 101, right paramere.

Figs. 91-101. Haliplus excoffieri, holotype. -

dorsal side of hind tibia

99, left paramere

VAN VONDEL: Palaearctic Liaphlus

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, elytral punctuation; 105, dorsal side of

; 108, left paramere; 109, penis; 110, right

110. Haliplus eximis, China. - 102, dorsal view; 103, antenna; 104

; 106, prosternal process; 107, lateral view of prosternal process

Figs. 102-

hind tibia

paramere.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 111-119. Haliplus flavicollis, Hendrik Ido Ambacht, Netherlands. - 111, dorsal view; 112, antenna; 113, elytral

punctuation; 114, dorsal side of hind tibia; 115, prosternal process; 116, lateral view of prosternal process; 117, left

paramere; 118, penis; 119, right paramere.

100

Haliplus flavicollis Sturm, 1834: 150. Types not located,

probably lost as most of the Sturm-types [not exam-

ined].

(Haliplus ferrugineus, sensu auctt., nec Linnaeus, 1767.

Misidentification |.

(Haliplus impressus, sensu auctt., nec Fabricius,1787.

Misidentification].

Haliplus flavicollis var. pallida Semenov, 1904: 202.

Types probably in ZMAS [not examined].

Haliplus flavicollis var. expallidus Zaitsev, 1907: 122

(Replacement name for pallida Semenov).

Haliplus unicolor Fiori, 1914: 188. Types not located [not

examined |.

Haliplus flavicollis, Aubé 1836: 24, 1838: 12, Schaum

1848: 42, Thomson 1854: 183, Seidlitz 1887: 31,

Miiller 1900: 115, Apfelbeck 1904: 361, Fiori 1904:

196, Zimmermann 1920: 305, 1924: 132, Guignot

1928: 143, 1933: 230, 1947: 50, Kinel 1929: 219,

Scholz 1929: 15, Kolossov 1931: 116 (var. russkyi

Stange, nom. nud.), Balfour-Browne 1936: 75, 1938:

12, 1940: 148, 1951: 1, 1953: 5, FE Burmeister 1939:

212, E. Burmeister 1982: 94, 1987: 184, Csiki 1946;

559, Zaitsev 1953: 68, Gueorguiev 1960: 23, Lindroth

1960: 42, Freude 1971: 15, Seeger 1971a: 409, 1971b:

532, 197 1c: 165, Franciscolo 1972: 61, 1979: 117; Jack-

son 1973: 253, Ienistea 1974: 208, 1978: 294, Mino-

ransky & Dzumailo 1974: 26, Galewski 1976: 31,

Konev 1976: 56, Mateleshko 1977: 68, Fichtner 1981:

327, Holmen 1981: 2, 1987: 114, Leblanc 1984: 8,

Lundberg 1986: 11, Zakharenko & Moroz 1988: 283,

Belyashevskiy 1989: 124, Vondel 1989a: 58, 1989b: 17.

Haliplus flavicollis var. expallidus; Zimmermann 1920:

305, Guignot 1933: 239, Zaitsev 1953: 68.

Haliplus unicolor; Burmeister 1939: 214.

Diagnosis

This species is related to eximis and rubidus.

From eximzs it can be distinguished by the lack of

lateral plicae in the anterior part of the prosternal

process. In rubidus the basal elytral punctures are

larger than the distance between the punctures of

the first and second row in the basal part, while in

flavicollis these punctures are clearly smaller than

that distance. In specimens from Morocco, how-

ever, that difference is less clear. The males of

flavicollis have the left paramere narrower at the

base than in rubidus.

Specimens with vague blotches on the elytra can

be distinguished from fulvus by the lack of the

lateral plicae in the anterior part of the prosternal

process.

Description

Length 3.6-4.1 mm, width 1.8-2.1 mm. Body

fairly slender, sometimes subparallel, widest before

the middle (fig. 111).

Head. - Yellow to rust-coloured, sparsely punc-

tured. Distance between the eyes 1.4-1.6 X eye-

width. Antennae (fig. 112) and palpi yellow to

yellow-red.

Pronotum. — Yellow to rust-coloured. Lateral

VAN VONDEL: Palaearctic Liaphlus

borders straight, finely bordered. Sparsely, ante-

riorly and basally more densely punctured, basal

irregular puncture-row about as strong as basal

elytral punctures. All punctures usually black (fig.

bly

Elytra. - Yellow to rust-coloured, strong primary

puncture-rows, about 32 punctures in the first row.

Secondary rows with relatively strong and between

them very weak punctures (fig. 113). All punctures

black. Suture weakly darkened, no clear markings

on the intervals, sometimes vague blotches on the

apical half.

Ventral side. - Body yellow to yellow-brown,

elytral epipleura yellow, legs yellow-brown, femora

darkened basally, trochanteres dark. Prosternal

process flat, wide apically, strongly narrowed near

the coxae (fig. 115), lateral border without plicae

before the coxae (fig. 116), strongly and densely

punctured. Metasternal process flat with a strong

impression in the middle, sparsely punctured (fig.

115). Setiferous striole on dorsal face of hind tibia

long and grooved, separate punctures hardly recog-

nizable, longer of the two tibial spurs about % X

the length of the first tarsal segment (fig. 114).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs hardly widened later-

ally, slightly widened ventrally, scaly hairs on ven-

tral side. Penis and parameres as in figs. 117-119.

Female: Elytra completely and very densely co-

vered with micropunctures.

Biology

In clean and clear stagnant or very slowly run-

ning water of lakes, ditches, canals and pools with

rich vegetation, especially Characeae (Seeger

1971c), Najadaceae (Burmeister 1939) or Ca-

bomba-beds in Kazakhstan (Konev 1976). Also

recorded from brackish water. The adults feed on

eggs and larvae of Chironomidae (Seeger 1971c),

aquatic macro-invertebrates as Asellus (only when

dead or dying), Stalis larvae (probably predated),

dead waterbeetles, even of their own species and

probably Characeae or filamentous algae. Seeger

(1971a, b, c) provided much information on the

life-cycle. The egg has been described by Seeger

(1971a), the 2nd and 3rd-stage larvae and the pupa

have been described by Bertrand (1928). This spe-

cies is able to fly (Jackson 1973, taken at light).

Distribution (fig. 295)

All Europe except the north of Norway, the

utmost north of Sweden, the north of Finland,

Iceland, Spain and Portugal. Turkey. USSR: Euro-

pean part, Western Siberia except the extreme

north, Transcaucasia, Kazakhstan, China (Guignot

1947), Mongolia. Records from Morocco probably

concern specimens of rzbidus, which is not always

101

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

e .

’

Austria. - 120, dorsal view; 121, antenna; 122, elytral punctuation; 123, dorsal

; 125, lateral view of prosternal process

Figs. 120-128. Haliplus fulvus, Gars,

: 126, left paramere; 127, penis; 128,

side of hind tibia; 124, prosternal process

right paramere.

102

easy to distinguish from flavicollis. Egypt? (Hol-

men 1987). Ali (1976) records specimens of fulv1-

collis from Iraq, but according to the characters he

used in his identification-key he obviously meant

flavicollis.

Material examined. — 881 ex. — Austria: 97 ex. (CNCI,

NHMW, ZMAN, RMNH). — Belgium: 7 ex. (RIN, RMNH, Jon-

gema). — Czechoslovakia: 20 ex. (NHMW, RMNH). —

France: 23 ex. (CNCI, NHMW, RIN, RMNH, Jongema, Von-

del). - Germany: 24 ex. (NHMW, ZMAN, RMNH). — Great

Britain: 9 ex., (CNCI, BMNH, ZMAN, Vondel). — Greece: 2

ex. (ZMAN). — Hungary: 2 ex. (CNCI). - Ireland: 1 ex.

(BMNH). — Italy: 4 ex. (CNCI, Jongema, RMNH, NHMW). —

Luxembourg: 1 ex. (ZMAN). - Mongolia: 1 &, Uiir Gol,

28.ix.1897, Frenzel (ZMHB). — Netherlands: 649 ex.

(ZMAN, MOG, FNML, NHMR, RIN, RMNH, Berge Hene-

gouwen, Cuppen, Drost, Hielkema, Huybregts, Jongema,

Onder de Linden, Schilthuizen, Vallenduuk, Vondel). —

Poland: 13 ex. (NHMW, RMNH).- Rumania: 1 ex. (RMNH).

- Switzerland: 8 ex. (ZMAN, Rotzer). - USSR: 1 ex.,

Panfilov, Kazakhstan; 1 ex., Kiyev, Ukraine, 27.ix.1988,

M. Jäch (NHMw).- Locality not traced or unknown: 17 ex.

(BMNH, NHMW, RMNH).

13. Haliplus fulvus (Fabricius)

(figs. 120-131)

Dytiscus fulvus Fabricius, 1801: 271. Types not located,

probably lost (Zimsen 1964: 70) [not examined].

Dytiscus interpunctatus Marsham, 1802: 429. Lectotype

Q (here designated), interpunctatus (BMNH)

[examined].

Haliplus lapponum Thomson, 1854: 184. Lectotype (here

designated), H. ferrugineus, coll. Lund, H.ferrugineus

Zett., ?syntype (MZLU, Zetterstedt-collection)

[examined |.

Haliplus pyraeneus Delarouzée, 1857: xcv. Types from

Lac Noire near Cauterets (Hautes Pyrénées) are prob-

ably in MNHN [not examined]. Material from NHMW,

labelled as TYPUS and COTYPUS is collected in 1881

and thus can not belong to the syntype-series.

Haliplus lapponum var. niger Seidlitz, 1887: 31. Types

not located [not examined].

Haliplus fulvus var. carlittensis Régimbart, 1901: 323.

Types probably in MNHN [not examined].

Haliplus subguttatus Crotch, 1873: 384. Syn. n.

Haliplus fulvus a. multistriatus Scholz, 1915: 232. Types

not located [not examined].

Haliplus fulvus Sparre-schneideri Munster, 1922: 15.

Lectotype @ (here designated), [ Norway, Malselv]

‘Bjerkeng, 12.6.97, H. Schneider: i. lit fulv. Schneideri

det. Ganglbauer, COTYPUS’ (NHMw) [examined].

Haliplus fulvus v. unicolor Munster, 1922: 14. Types not

located [not examined].

Haliplus salinarius Wallis, 1933: 56. Holotype 6,

[Canada] ‘Oyosoos, B.C., V-4-1929° (CNCI) [exami-

ned]. Syn. n.

[Haliplus ferrugineus; sensu auctt., nec Linnaeus, 1767.

Misidentification].

Haliplus fulvus; Clairville 1806: 220, Sturm 1834: 148,

Babington 1836: 177, Erichson 1837: 183, Thomson

1854: 183, Westhoff 1881: 43, Seidlitz 1887: 31, Ap-

VAN VONDEL: Palaearctic Liaphlus

felbeck 1904: 360, Escalera 1914: 57, Zimmermann

1920: 307, 1924: 134, Bertrand 1928: 87, 1949: 25,

Guignot 1928: 144, 1933: 230, 1947: 51, Scholz 1929:

16, Balfour-Browne 1936: 74, 1938: 11, 1940: 146,

1951: 4, 1953: 5, Pearce 1939: 34, F. Burmeister 1939:

212, Horion 1941: 365, Brinck 1942: 5, Strand 1944:

122, Csiki 1946: 560, Jackson 1952: 66, 1956: 87, 1973:

253, Zaitsev 1953: 70, Riha 1955: 57, Gueorguiev

1958: 44, 1960: 23, 1981: 401, Lindroth 1960: 42,

Lagar Mascaro 1968: 72, Freude 1971: 15, Seeger

1971b: 558, 1971c: 165, Ienistea 1974: 208, 1978: 294,

Minoransky & Dzumailo 1974: 26, Alfieri 1976: 38,

Galewski 1976: 29, Konev 1976: 56, Gislason 1977:

159, Mateleshko 1977: 68, Franciscolo 1979: 119,

Fichtner 1981: 327, Holmen 1981: 2, 1987: 115, Leb-

lanc 1984: 40, Lundberg 1986: 11, E. Burmeister 1987:

184, Zakharenko & Moroz 1988: 283, Belyashevskiy

1989: 124, Vondel 1989a: 58, 1989b: 17.

Haliplus interpunctatus, Zimmermann 1920: 307.

Haliplus lapponum; Seidlitz 1887: 31, Zimmermann

1920: 308, Bertrand 1928: 87, 1949: 25, Guignot 1928:

144, 1930: 73, 1933: 231, 1947: 51, Kinel 1929: 219,

Scholz 1929: 16, Balfour-Browne 1936: 74 1938: 11,

1940: 147, Burmeister 1939: 213, Pearce 1939: 34,

Horion 1941: 365, Brinck 1942: 5, Strand 1944: 122,

Zaitsev 1953: 71, Riha 1957: 57, Lagar Mascaro 1968:

72, Ienistea 1978: 294, Lundberg 1986: 11.

Haliplus pyraeneus, Marseul 1882: 106, Zimmermann

1920: 308, Bertrand 1928: 87, Kinel 1929: 219, Scholz

1929: 16, Burmeister 1939: 213.

Haliplus subguttatus; Blatchley 1910: 203, Roberts 1913:

101, Zimmermann 1920: 139, Wallis 1933: 54, 1973:

100, J. Balfour-Browne 1948: 158, Hatch 1953: 192,

Ruette 1970: 57, Hilsenhoff & Brigham 1978: 21,

White et al. 1985: 357, Larson 1987: 107.

Haliplus lapponum var. niger; Scholz 1929: 16, Guignot

1933: 242.

Haliplus fulvus var. carlittensis, Zimmermann 1920: 308,

Bertrand 1928: 87, Guignot 1928: 144, 1930: 73, 1933:

242, Kinel 1929: 219, Scholz 1929: 16, Burmeister

1939: 213, Pearce 1939: 34.

Haliplus fulvus a. multistriatus, Zimmermann 1920: 308,

Bertrand 1928: 87, Guignot 1933: 241.

Haliplus fulvus sparre-schneideri, Guignot 1933: 241,

Strand 1944: 122, Zaitsev 1953: 71, Döbler 1976: 182

(as schneideri), Ienistea 1978: 294.

Haliplus fulvus v. unicolor; Falkenstròm 1941: 88.

Haliplus salinarius; Hatch 1953: 192, Ruette 1970: 56,

Larson 1987: 107.

Haliplus ferrugineus; Gyllenhal 1808: 546, Zetterstedt

1828: 236, Stephens 1829: 40, Aubé 1836: 22, 1838:

10, Babington 1836: 176, Zimmermann 1920: 308.

Diagnosis

This species is very variable in size, maculation

and coloration. Of all Haliplidae it has the largest

distribution, covering most of the palaearctic and

the north of the nearctic region. Mainly because of

the variation in elytral maculation a number of

varieties, subspecies and separate species such as

lapponum (fig. 129) and pyraeneus (fig. 130) have

been described. A range of intermediate forms be-

tween the latter species and fulvus has been exam-

ined and I can only confirm what other authors like

103

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

ole sel She cheb?

ERS ee

Figs. 129-131. Haliplus

fulvus, elytra. - 129, lec-

totype of H. lapponum,

130, from Cauterets,

France, the type locality

of H. pyraeneus; 131, lec-

totype of H. fulvus

sparre-schneideri.

Balfour-Browne (1936) and Holmen (1987) al-

ready concluded, that they belong to the same spe-

cies.

Description

Length 3.6-4.5 mm, width 2.0-2.4 mm. Body

long oval, widest in the middle (fig. 120).

Head. - Yellow-red, brown or rust-coloured,

puncturation weak to moderately strong, between

the eyes sparser, behind the eyes stronger and

denser punctured. Distance between the eyes 1.6-

1.9 X eye-width. Antennae (fig. 121) and palpi

yellow.

Pronotum. - Yellow to rust-coloured. Lateral

borders straight to slightly convex, finely bordered.

Punctures moderately strong, on the disc almost

absent, the basal row widened and slightly to

strongly darkened.

Elytra. - Yellow to rust-coloured, very variable

pattern of light to dark brown blotches in the even

intervals, sometimes the blotches are hardly or not

present or they are very strong and then covering

nearly the entire elytra (described as sparre-

schneideri) (fig. 131). Primary punctures moder-

ately strong, sometimes stronger in the basal part.

Secondary rows irregular and containing two types

of punctures, the usual ones and very fine ones (fig.

122). All punctures darkened.

Ventral side. - Body yellow, yellow-red or rust-

104

9.0 0.0 0.0 9.9

Bag geen

coloured, elytral epipleura yellow, legs yellow to

rust-coloured, darkened towards the coxae. Pros-

ternal process flat to slightly convex, narrowed

near the coxae (fig. 124), lateral plicae complete

(fig. 125), with moderately strong to very strong

and dense punctures. Metasternal process flat with

a pit in the middle (fig. 124), with moderately

strong and dense punctures. Setiferous striole on

dorsal face of hind tibia grooved, stretched over

about % of the length, consisting of 10 to 20 often

hardly recognizable punctures, longer of the two

tibial spurs % X the length of the first tarsal seg-

ment (fig. 123). Length of hind tarsus, including

claws, about 1.2 X the tibia length, first tarsal

segment 1.5-1.9 X the length of the second seg-

ment.

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 126-128, length of the distal process of the

right paramere % X the length of the basal part

(fig. 128).

Biology

In clean clear or peaty stagnant or slowly run-

ning water of lakes, pools, canals, rivers and ditches

with rich vegetation of especially Characeans. Even

recorded from brackish water. The larvae probably

feed on Characeans such as Nitella, while the adults

are omnivorous with a diet of algae, pollen and

invertebrates like Oligochaeta (Seeger 1971b). The

third-stage larva was first described by Schigdte

(1864). The beetle has been seen flying and has

been caught at light.

Distribution (fig. 292)

All of Europe, including Iceland. USSR: Euro-

pean part and Siberia as far as Yakutia in the east

and Transcaucasia and Kazakhstan in the south.

Turkey. North Africa: Morocco, Algeria. The re-

cords of Syria (Burmeister 1939) and Iraq (Ali

1976) possibly concern maculatus, which is cer-

tainly a different species. Canada (states of British

Columbia, Alberta, North West Territories, Saskat-

chewan, Manitoba, Ontario, Quebec, Nova Scotia

and New-Foundland) and northern parts of the

United States from west to east, southwards to

about 45° N (states of Washington, Idaho, South

Dakota, Minnesota, Wisconsin, Indiana and Mas-

sachusetts) (until now referred to as salinarius and

subguttatus).

Remarks

Type-material of the nearctic species subguttatus

sensu Roberts and salinarius Wallis has been exam-

ined and in my opinion both are conspecific with

fulvus. For salinarius that possibility was already

noticed by Holmen (1987). Crotch (1873) menti-

oned subguttatus referring to a Leconte manus-

cript, but he did not regard it as a separate species,

but as a form of cribarius Leconte. Blatchley (1910)

described it as a subspecies of cribarius. Roberts

(1913) discovered that subguttatus was not conspe-

cific with cribarius and described it as a separate

species. His holotype is probably in AMNH. Two of

his paratypes have been examined (CNCI).

Material examined. - 712 ex. - Austria: 36 ex. (RMNH,

NHMW, ZMAN, OLML). — Belgium: 7 ex. (RMNH, RIN, Jon-

gema). — Canada: 3 ex. (holotype and 2 paratypes of

salinarius); 16, (paratype of subguttatus), Fraz. Val. B.C.;

1 8, (paratype of subguttatus), Antigonish, N.S. (CNCI);

44 ex. (CNCI, Vondel). - Czechoslovakia: 2 ex. (NHMW,

OLML). — Denmark: 1 ex. (NHMW). - Finland: 10 ex.

(RMNH, ZMAN, RMNH, NHMW). — France: 30 ex. (RMNH,

ZMAN, RMNH, NHMW, BMNH, OLMI, Vondel). - Germany:

39 ex. (RMNH, NHMW, ZMAN, BMNH, OLML). - Great Bri-

tain: 94 ex. (CNCI, ZMAN, BMNH, Vondel). - Greece: 1 ex.

(OLML). - Hungary: 2 ex. (NHMW, ZMAN).-Ireland: 18 ex.

(BMNH). - Italy: 3 ex. (CNCI). — Netherlands: 228 ex.

(BMNH, ZMAN, FNML, NHMR, NHMW, RIN, RMNH, Cuppen,

Drost, Hielkema, Jongema, Onder de Linden, Vondel). —

Norway: 38 ex. (RMNH, ZMAN, BMNH, OLML, MZLU,

RMNH); 1 8, (lectotype of fulvus var. sparre-schneidert),

19, (paralectotype of fulvus var. sparre-schneidert), Bjer-

keng, ix.1878, Schneider, Haliplus Sparre-Schneideri n.

sp., fulv. Schneideri det. Ganglbauer, Typus (NHMW). —

Poland: 12 ex. (NHMW). — Rumania: 2 ex. (NHMW). —

Spain: 1 ex. (Cuppen). - Sweden: 11 (NHMW, ZMAN,

VAN VONDEL: Palaearctic Liaphlus

RMNH, BMNH). — USA: 1 ex., Larive Lake, South Dakota,

22.vi.1940 (CNCI). - USSR: 1 ex. (BMNH). — Yugoslavia:

3 ex. (CNCI, OLML). - Locality not traced or unknown: 118

ex. (NHMW, BMNH, MZLU, RMNH, OLML), 1 ex., (9 lecto-

type of interpunctatus), (BMNH); 1 ex. (lectotype of lap-

ponum) (MZLU).

14. Haliplus gafnyi sp. n.

(figs. 132-140)

Type material: Holotype 4, Israel, Nesher quarry,

27.1.1989, R. Ortal, IES 5344 (Huy). Paratypes: 1 6,

Israel, Berekhat, Nesher quarry, 26.xii.1989, R. Ortal, IES

5502 (Angus); 1 &, Cyprus, Zakaki Marshes, ii.1948,

G. Mavromoustaki (Angus); 1 6, Syria, Damascus

(MNHN).

Diagnosis

This species resembles abbreviatus and villiersi

in a number of characters, but can be distinguished

from them by the flat, strongly and densely punc-

tured metasternal process.

Description

Length 2.9-3.1 mm, width 1.5-1.7 mm. Body

oblong oval, widest in the middle (fig. 132).

Head. - Yellow-brown, vertex brown, moderate

strongly punctured, on the vertex strong darkened

punctures. Distance between the eyes 1.4-1.6 X

eye-width. Antennae (fig. 133) and palpi yellow to

yellow-brown.

Pronotum. - Yellow to yellow-brown, anterior

margin darkened in the middle. Lateral borders

straight, finely bordered. Moderate strongly punc-

tured, along the base with a partly double row of

strong punctures, discal punctures brown, basal

punctures black.

Elytra. - Yellow, darkened suture narrow, at

most reaching the secondary punctures on the disc,

vague brown blotches on the shoulders and in the

apical part (fig. 132). Primary punctures strong, 28

punctures in the first row. Secondary punctures

strong. All punctures darkened (fig. 134).

Ventral side. - Yellow to yellow-brown, elytral

epipleura yellow, legs yellow-brown, some darken-

ing towards the coxae. Prosternal process flat,

moderate strongly punctured, narrowed near the

coxae (fig. 136), lateral plicae complete (fig. 137).

Metasternal process flat with a small pit in the

middle, strongly and densely punctured (fig.136).

Setiferous striole % X the tibia length, with about

10 punctures, longer of the two tibial spurs 24 X the

length of the first tarsal segment (fig. 135).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 138-

140.

105

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 132-140. Haliplus gafnyi, holotype. - 132, dorsal view; 133, antenna; 134, elytral punctuation; 135, dorsal side

of hind tibia; 136, prosternal process; 137, lateral view of prosternal process; 138, left paramere; 139, penis; 140, right

paramere.

106

Etymology. - Named after the late Mr. Gabriel

Gafny, Hydrologist of the Nature Reserve Author-

ity of Israel.

Biology

In temporary pool.

Distribution (fig. 303)

Only known from Israel.

15. Haliplus guttatus Aubé

(figs. 141-149)

Haliplus guttatus Aubé, 1836: 27. Lectotype & (here de-

signated), unlabelled (MNHN, coll. SEE Aubé)

[examined].

Haliplus pelopis Sahlberg, 1907: 21. Lectotype & (here

designated), Elis, J. Sahlb., Spec. typ., 2284, Mus. Zool.

H.fors, Spec. typ. No. 852, Haliplus pelopis J. Sb.

[aedeagus missing] (UZMH) [examined]. Syn. n.

Haliplus guttatus; Aubé 1838: 15, Miiller 1900: 113, Ap-

felbeck 1904: 359, Fiori 1904: 195, Escalera 1914: 57,

Zimmermann 1920: 308, 1924: 135, Guignot 1928:

145, 1933: 232, 1947: 52, 1959: 36, Scholz 1929: 15, E.

Burmeister 1939: 213, E. Burmeister 1987: 177, Csiki

1946: 560, Zaitsev 1953: 72, Lagar Mascaro 1968: 73,

Ienistea 1978: 294, Franciscolo 1979: 122.

Haliplus pelopis; Zimmermann 1920: 314, 1924: 135,

Burmeister 1939: 214, Zaitsev 1953: 72, Ienistea 1978:

294.

Diagnosis

Very variable in shape, from oval to strongly

parallel, maculation not always clearly visible.

Description.

Length 3.6-4.1 mm, width 1.7-2.1 mm. Body

parallel to subparallel (fig. 141).

Head. - Yellow-red to rust-coloured, dark blotch

on vertex, weakly, behind the eyes more strongly

punctured. Distance between the eyes 1.7-1.9 X

eye-width. Antennae (fig. 142) and palpi yellow-

red.

Pronotum. - Yellow to rust-coloured. Lateral

borders convex to nearly straight, sometimes

slightly concave before the basal corners, finely

bordered. Weakly and sparsely, in the basal part

densely punctured, basal row of darkened punc-

tures twice as wide as the elytral punctures.

Elytra. - Yellow with a variable pattern of vague,

sometimes hardly visible blotches, in most cases a

large blotch on the suture (fig. 141). Fine primary

punctures basally stronger, about 37 punctures in

the first row. Secondary punctures weak. All punc-

tures darkened (fig. 143).

Ventral side. - Body yellow to yellow-red, elytral

epipleura yellow, legs yellow-red, some darkening

towards the coxae. Prosternal process flat, slightly

VAN VONDEL: Palaearctic Liaphlus

narrowed near the coxae (fig. 145), lateral plicae

complete (fig. 146), coarsely and densely punc-

tured. Metasternal process flat with a shallow pit

in the middle, weakly and sparsely punctured (fig.

145). Setiferous striole on dorsal face of hind tibia

about 4 X the tibia length with 5-10 punctures,

longer of the two tibial spurs % to % X the length

of the first tarsal segment (fig. 144). Length of hind

tarsus including claws 1.3-1.4 X the tibia length,

first tarsal segment about 1.3 X the length of the

second segment.

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 147-149.

Biology

In stagnant water, fresh or brackish, even in

brooks.

Distribution (fig. 297)

Mediterranean, south and south-west of France,

Corsica, Spain including Baleares, Italy including

Sardinia and Sicily, Malta, Yugoslavia, Greece,

North Africa: Morocco, Algeria, Tunisia. All re-

cords from Poland and the southern states of the

USSR are suspect and likely concern other species

such as fulvus, variegatus or maculatus.

Remarks

The types of guttatus and pelopis have been

examined. They do not show all differences used in

earlier identification-keys. I regard both conspe-

cific.

Material examined. - 67 ex. - Algeria: 5 ex.; 2 ex.,

Algiers, v.1861, H. Clark; 3 ex., Kraatz; 3 ex., lac des

Serpents, Djijellé, Massif des Mouzaia (BMNH); 1 ex.

Philippeville (MNHN); 2 9, Taguin; 4 ex., Prov. d’Alger,

Taguin, 1895, de Vauloger (NHMW). — France: 1 ex.,

Nizza, (NHMW); 1 ex., Montpellier, v.1861, H. Clark

(BMNH); 1 ex., Camargue, nr. Salin de Giraud, 12.vi.1980

(Huijbregts); 1 ex., Albaron (B. du R.), L. Schaefer

(RMNH); 1 ex., Albaron (B. du R.); 1 ex., Pyrenées; 1 ex.,

Montfort, Landes (ZMAN); 2 ex., Beragne (MNHN); | ex.,

Les Angles, Gard. iv.1928; 1 ex., Camargue; 1 ex., Corsica,

1905, Aleria; 4 ex., Hyères (BMNH). - Greece: 1 & (Lec-

totype of pelopis) (UZMH); 1 ex. (MNHN); 2 ex., Acarnan,

8.1.1969, Forster (OLML); 1 ex., Attika (ZMAN); 4 ex.,

Zante, Kalamaki, 1909, M. Hilf; 1 ex., Khalkidhiki, Sitho-

nia, 22.vii.1988, M Jäch; 1 ex., Kérkira, Stephanides

(NHMW). — Italy: 1 ex., Sardinia, Assuni, Krausse

(NHMW); 1 À, Sardegna, Gag., 7 km W. of Carbonia, alt.

20 m., 6.v.1984 (Huijbregts); 1 ex., Lucania, Policorof

M.T., 3.xi.1972, Angelini; 1 ex., Toscane, Dahl (RMNH);

1 6, Florenz, Brozzi (OLMI); 1 ex., Etruria, (BMNH). —

Malta: 1 ex., Chadwick Lakes, 31.1.1987, M. Borg, fresh

water (NHMW). - Spain: 2 ex., Palencia (MNHN, ZMAN).

— Yugoslavia: 1 &, Zadar (Zara), Sturany, 1891 (NHMW).

- Locality not traced: 1 ex., S. Sparato, Dodero (MNHN)

107

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 141-149. Haliplus guttatus, Zante, Greece. - 141, dorsal view; 142, antenna; 143, elytral punctuation; 144, dorsal

side of hind tibia; 145, prosternal process; 146, lateral view of prosternal process; 147, left paramere; 148, penis; 149,

right paramere.

108

- Without locality: 1 6, Lectotype of guttatus; 1 9, 2 8,

Paralectotypes of guttatus (MNHN); 2 ex., 67-56 (BMNH);

1 ex., UlIr.; 2 ex., Dhl., aa 44 (NHMW)

16. Haliplus holmeni sp. n.

(figs. 150-160)

Type material: Holotype g, [China] “Yunnan, Museum

Paris, coll. Maurice Régimbart, 1908’ (MNHN).

Diagnosis

Looks related to sharpi, but the male has a left

paramere with a solid digitus. The fore tarsi have

a cushion between the claws.

Description

Length 3.3 mm, width 1.7 mm. Body oval, widest

in the middle (fig. 150).

Head. - Brown with yellow labrum, dark blotch

on vertex, moderately strongly punctured, unpunc-

tured band between the hind parts of the eyes.

Distance between the eyes 1.2 X eye-width. Anten-

nae yellow, base of eighth till eleventh segment

slightly darkened, top of ultimate segment pale

(fig. 151), palpi yellow.

Pronotum. - Yellow-brown to brown anteriorly,

lateral borders somewhat irregularly convex,

brown and clearly bordered. Width at the base 2.0

the length in the middle and 1.9 X the width in

front. Sparsely, anteriorly densely, basally strongly

punctured. Basal punctures black and partly wi-

dened. Slightly impressed along the central part of

the base (fig. 150).

Elytra. - Yellow-brown with extensive black

markings: wide band along the base till the fifth

puncture-row, suture, arrow-shaped central blotch

and blotches on the apical half, the apex and three

large blotches on the fifth to ninth interval. Prim-

ary puncture-rows strong, about 24 punctures in

the first row, third to fifth row close to each other

in the central part. Secondary punctures moder-

ately strong (fig. 152). Primary and secondary

punctures tending to form transverse impressions

in the basal part. All punctures darkened.

Ventral side. - Rust-coloured to brown, elytral

epipleura yellow-brown, legs yellow-red, darkened

towards the coxae. Prosternal process slightly im-

pressed, stronger impressed apically, densely punc-

tured, slightly narrowed near the coxae (fig. 155),

lateral plicae complete (fig. 156). Metasternal pro-

cess flat, impressed apically and a shallow pit in the

middle, sparsely punctured (fig. 155). Coxal plates

rather strongly punctured, along the suture weakly

punctured, short hairs in the punctures, only few

small punctures between the normal ones (fig.

157). Hind margins of the fifth and sixth abdom-

inal sternite with complete puncture-rows (fig.

VAN VONDEL: Palaearctic Liaphlus

157). Setiferous striole consisting of about 6 separ-

ate punctures, longer of the two tibial spurs about

% the length of the first tarsal segment (fig. 154).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Between the fore-claws a small cu-

shion (fig. 153). Penis and parameres as in figs.

158-160, left paramere with a parallel solid digitus.

Etymology. - This species is named after my

colleague specialist of Haliplidae, Mogens Hol-

men.

Distribution (fig. 312)

Only known from south-west China (Yunnan)

17. Haliplus jaechi sp. n.

(fig. 161-169)

Type material: Holotype 6, [Turkey], ‘TR, Bismil 6,

Diyarbakir, 29.v.1987 (35), Jäch’ (NHMW).

Diagnosis

This species is related to kulleri and ortali. The

left paramere has a strongly developed solid digitus

and on the inner margin a well developed row of

setae.

Description:

Length 3.3 mm, width 1.6 mm. Body oblong

oval, widest in the middle (fig. 161).

Head. - Brown, puncturation moderately strong

and dense, unpunctured band between the hind

parts of the eyes. Distance between the eyes 1.4 X

eye-width. Antennae yellow-brown, second to

fourth segment yellow, third segment 1.5 X the

length of the fourth segment (fig. 162). Palpi yel-

low-brown.

Pronotum. - Yellow-brown. Lateral borders

slightly convex, darkened and clearly bordered.

Width at the base 1.8 X the length in the middle

and 1.5 X the width in front. Sparsely punctured,

on the disc almost unpunctured, along the base

some slightly darkened larger punctures. Along the

base weakly impressed (fig. 161).

Elytra. - Yellow to yellow-brown, darkened su-

ture and apex, vague blotches on the even intervals.

Near the base a transverse slightly darkened band.

Primary puncture-rows moderately strong, 28

punctures in the first row. Secondary punctures

moderately strong, accompanied by very small

punctures (fig. 163), all punctures darkened.

Ventral side. - Yellow to yellow-brown, elytral

epipleura yellow, legs yellow-brown, darkened to-

wards the coxae. Prosternal process slightly con-

vex, strongly narrowed near the coxae, rather

weakly punctured (fig. 165), lateral plicae complete

109

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 150-160. Haliplus holmeni, holotype. - 150, dorsal view; 151, antenna; 152, elytral punctuation; 153, fore claws;

154, dorsal side of hind tibia; 155, prosternal process; 156, lateral view of prosternal process; 157, hind coxal plate;

158, left paramere; 159, penis; 160, right paramere.

110

VAN VONDEL: Palaearctic Liaphlus

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, holotype. - 161, dorsal view; 162, antenna; 163, elytral punctuation; 164, dorsal side

, prosternal process; 166, lateral view of prosternal process; 167, left paramere; 168, penis; 169, right

Figs. 161-169. Haliplus jaechi

of hind tibia; 165

paramere.

iat

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 170-178. Haliplus kotoshonis, Okinawa. - 170, dorsal view; 171, antenna; 172, elytral punctuation; 173, dorsal

side of hind tibia; 174, prosternal process; 175, lateral view of prosternal process; 176, left paramere; 177, penis; 178,

right paramere.

112

(fig. 166). Metasternal process flat with a shallow

impression in the middle, sparsely punctured (fig.

165). Coxal plates sparsely punctured. Setiferous

striole on dorsal side of hind tibia about ‘4 of the

tibia length, with about 7 punctures, longer of the

two tibial spurs % the length of the first tarsal

segment (fig. 164).

Male sexual characters. — First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 167-

169, left paramere with a solid digitus and a well

developed row of long bristles on the inner margin.

Etymology. — Named after the collector Dr.

Manfred Jäch.

Distribution (fig. 302)

Only known from south-east Turkey, Bismil.

18. Haliplus kotoshonis Kano & Kamiya

(figs. 170-178)

Haliplus kotoshonis Kano & Kamiya, 1931: 2. Types

burned during World War II (Nakane pers. comm.).

Haliplus kotoshonis; Sato 1984: 2, Nakane 1985a: 64,

1987: 30.

Diagnosis

This species resembles pulchellus and maculi-

pennis by having the eyes close to each other, but

it can be distinguished by the simply impressed

prosternal process and the lack of a basal black

elytral band.

Description

Length 3.5-3.6 mm, width 1.9-2.0 mm. Body

oval, middle part subparallel, widest in the middle

(fig. 170).

Head. - Brown, along the eyes dark brown.

Puncturation strong and dense. Clypeus impressed

in the middle. Distance between the eyes 0.8-1.0 X

eye-width. Antennae (fig. 171) and palpi brown-

red.

Pronotum. - Brown-red. Lateral borders slightly

convex, finely bordered. Hind corners protruding

and strongly rounded. Punctures very strong, ex-

cept on the disc dense and along the base often

coarse. Base strongly impressed.

Elytra. - Yellow-brown, dark brown maculation

on sutural interval and on the apex, a central blotch

on the disc and small sometimes connected

blotches on the other intervals. Suture slightly bor-

dered in the basal part and at the apex. All primary

punctures strong to very strong, basal punctures

coarse, 32-34 punctures in the first row. Secondary

punctures strong and dense. All punctures dar-

kened and with a small yellow centre (fig. 172).

VAN VONDEL: Palaearctic Liaphlus

Ventral side. - Body brown, elytral epipleura

yellow-brown, legs yellow-brown to brown,

slightly darkened near the coxae. Prosternal pro-

cess clearly impressed in the middle, narrowed near

the front coxae (fig. 174), lateral plicae complete

(fig. 175), strongly and partially coarsely punc-

tured. Metasternal process impressed and with a

pit in the middle, strongly punctured (fig. 174).

Setiferous striole on dorsal face of hind tibia about

4, X the tibia length, containing about 8 punctures,

longer of the two tibial spurs about 24 X the length

of the first tarsal segment (fig. 173).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 176-178.

Distribution (fig. 310)

Taiwan (type-locality: isle of Kotosho), Japan

(Okinawa).

Remarks

Although this species seems to belong to the

oriental fauna, it is treated here for practical rea-

sons.

Material examined. - Japan: 1 8, 1 9, Kimibaru, Oh-

gimi V., Okinawa-Hontô, 18.viii.1989, leg. E. Matsui (in

authors coll.).

19. Haliplus kulleri Vondel

(figs. 179-187)

Haliplus kulleri Vondel, 1988: 97. Holotype @, Israel,

25.3.81, Lower Galilee, Parod Res., N. Ammud Basin,

35 km W. of Akko, IES 2350, Col. 1673 (NHMW)

[examined].

Diagnosis

Closely related to jaechi and ortali, but in the

males characterised by the long parallel-sided solid

digitus on the left paramere.

Description

Length 2.9-3.0 mm, width 1.6 mm. Body fairly

narrow, subparallel, widest in the middle (fig. 179).

Head. - Brown, puncturation behind the eyes

rather dense, anteriorly more sparse, leaving an

unpunctured shiny area between the eyes. Distance

between the eyes 1.4-1.5 X eye-width. Antennae

(fig. 180) and palpi yellow.

Pronotum. - Yellow, slightly darkened on the

disc. Lateral borders nearly straight, finely bor-

dered. Sparsely and weakly punctured, leaving an

unpunctured shiny area on the disc, two short basal

rows of slightly widened and darkened punctures

(fig. 179).

113

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

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Figs. 179-187. Haliplus kulleri, holotype. - 179, dorsal view; 180, antenna; 181, elytral punctuation; 182, dorsal side

of hind tibia; 183, prosternal process; 184, lateral view of prosternal process; 185, left paramere; 186, penis; 187, right

paramere.

114

Elytra. — Yellow with a pattern of brown

blotches, one triangular on the middle of the suture

and some smaller ones around. Moderately strong

darkened primary puncture-rows, about 30 punc-

tures in the first row. Secondary punctures small,

alternated with very fine punctures (fig. 181).

Ventral side. - Body yellow-brown, elytral epi-

pleura yellow, legs yellow. Prosternal process

slightly convex, narrowed near the coxae, covered

with strong punctures (fig. 183), lateral plicae con-

tinuous (fig. 184). Metasternal process flat with a

shallow pit in the middle, sparsely punctured (fig.

183). Legs half shiny, with fine micro-punctuation.

Setiferous striole on dorsal face of hind tibia with

about 9 punctures, longer of the two tibial spurs

about % X the length of the first tarsal segment

(fig. 182). Length of hind tarsus including claws 1.3

X the tibia length, first tarsal segment 1.5 X the

length of the second segment.

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs hardly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 185-187, left paramere with a long parallel

digitus (fig. 185).

Female: Unknown.

Biology

In temporary pool.

Distribution (fig. 302)

Only known from Israel, Lower Galilee.

Material examined: 2 ex. - Israel: 1 & (holotype); 1 4,

Berekhat Shosh, 23.1.1989, IES 5343 (HUJI).

20. Haliplus laminatus (Schaller)

(figs. 188-198)

Dytiscus laminatus Schaller, 1783: 314. Types not located,

Schaller’s material has been lost (Horn & Kahle 1936)

[not examined].

Haliplus cinereus Aubé, 1836: 30. Lectotype (here desig-

nated), Gallia, 2645, coll. Chevrolat, Det. Sharp. 82,

Haliplus cinereus Aubé p18, affinis Steph., Type, cine-

reus Aubé, Type d’Aube, TYPE (pink label) (ISNB)

[examined].

Haliplus laminatus ?var. ater Redtenbacher, 1842: 8.

Types not located [not examined].

Haliplus laminatus Q var. punctulatus Penecke, 1901: 11.

Types not located [not examined].

Haliplus lombardus Fiori, 1904: 198. Types not located

[not examined].

Haliplus laminatus; Seidlitz 1887: 31, Apfelbeck 1904:

361, Fiori 1904: 197, Zimmermann 1920: 310, 1924:

131, Guignot 1928: 142, 1933: 229, 1947: 48, Kinel

1929: 218, Scholz 1929: 15, Balfour-Browne 1938: 12,

1940: 154, 1951: 5, 1953: 5, Burmeister 1939: 213,

Horion 1941: 365, Csiki 1946: 559, Zaitsev 1953: 67,

Gueorguiev 1958: 44, 1960: 23, Lindroth 1960: 42,

VAN VONDEL: Palaearctic Liaphlus

Lagar Mascaro 1968: 71, Freude 1971: 15, Seeger

1971a: 430, 1971b: 558, 1971c: 166, Ienistea 1974: 207,

1978: 294, Ali 1976: 92, Galewski 1976: 30, Mate-

leshko 1977: 68, Franciscolo 1979: 113, Fichtner

1981:327, Holmen 1981: 2, 1987: 110, Leblanc 1984:

8, Lundberg 1986: 11, Vondel 1986: 128, 1989a: 57.

Haliplus cinereus, Aubé 1836: 30, 1838: 18, Erichson

1837: 185, Zimmermann 1920: 310, Balfour-Browne

1936: 75, 1938: 12, Burmeister 1939: 213.

Haliplus laminatus ?var. ater; Zimmermann 1920: 310.

Haliplus Q var. punctulatus; Zimmermann 1920: 310.

Haliplus lombardus, Zimmermann 1920: 312, Burmeis-

ter 1939: 213.

Diagnosis

This species can be distinguished from all other

palaearctic species of Liaphlus by the dark lines on

the elytral puncture-rows and in the males by the

strongly dilated first mesotarsal segment and the

lack of a digitus on the right paramere.

Description

Length 2.5-3.6 mm, width 1.5-1.9 mm. Body

with subparallel sides in the central part (fig. 188).

Head. - Yellow-brown with brown collar, with

dense and on the collar strong and darkened punc-

tures. Distance between the eyes 2.0-2.1 X eye-

width. Antennae (fig. 189) and palpi yellow.

Pronotum. - Yellow. Basally not as wide as base

of elytra. Lateral borders slightly convex, finely

bordered. Densely, on the disc more sparsely punc-

tured.

Elytra. - Yellow to yellow-red, strong primary

puncture-rows with widened and usually groove-

like basal punctures, about 42-46 punctures in the

first row. Secondary punctures rather weak. All

punctures darkened (fig. 190). Primary rows and

suture partly, sometimes completely covered with

dark lines, not connected to each other (fig. 188).

Ventral side. - Yellow to yellow-brown, elytral

epipleura yellow, legs yellow to yellow-brown,

coxae red-brown. Prosternal process grooved, nar-

rowed near the coxae, widened towards the apex,

punctures moderately strong and dense (fig. 193),

lateral plicae complete (fig. 194). Metasternal pro-

cess slightly impressed in the middle, moderately

strongly punctured (fig. 193). Setiferous striole on

dorsal side of hind tibia stretched over the apical %,

longer of the two tibial spurs % X the length of the

first tarsal segment (fig. 195).

Male sexual characters. - First three protarsal

segments slightly widened, scaly hairs on ventral

side, first segment ventrally with a complete row

of strong spines (fig. 191). First mesotarsal seg-

ment strongly delated ventrally (fig. 192). Penis

and parameres as in figs. 196-198, right paramere

without a digitus as characteristic for Liaphlus (fig.

198).

Female. - Elytra with strong micropuncturation.

115

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

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process; 195, dorsal side of hind tibia

tarsal segments of forelegs

116

Biology

Usually in slow running water of rivers, canals

and brooks, but also found in silt ponds. The adults

feed on filamentous algae and invertebrates like

larvae of Chironomidae and Oligochaeta (Seeger

1971b) and freshly killed Asellus aquaticus (Vondel

1986). The larvae (3rd and 2nd instar) were des-

cribed by Vondel (1986).

Distribution (fig. 293)

Central Europe from Denmark to the east coast

of Spain, central Italy and Albania and from south-

east England to the European part of the USSR.

The species is also recorded from Iraq (Ali 1976).

The records from Ireland (Burmeister 1939, Fran-

ciscolo 1979) are doubtful.

Remarks

This species is usually treated as belonging to the

subgenus Liaphlus, although Kinel (1929) erected

a separate subgenus Hoplites for this species. The

name Hoplites is preoccupied (see: International

Commission on Zoological Nomenclature 1955:

opinion 353), but never replaced. There is reason

to believe that this species is more closely related

to the subgenus Haliplus s. str. (Vondel 1986; Hol-

men 1987). Before erecting new subgeneric names

it seems better to me to await the results of syste-

matic studies on a world basis.

Material examined. - 481 ex. - Albania: 2 ex. (NHMW).

— Austria: 81 ex. (ZMAN, CNCI, NHMW, NHMR). - Belgium:

2 ex. (RMNH). — Czechoslovakia: 18 ex. (ZMAN, CNCI,

RMNH, MHNG, NHMW, RMNH). - France: 12 ex. (NHMW,

RMNH, Jongema, BMNH, Vondel). — Germany: 20 ex.

(ZMAN, NHMW, RMNH). — Great Britain: 5 ex. (BMNH). —

Hungary: 3 ex. (NHMW). - Italy: 12 ex. (RMNH, NHMW).

— Luxembourg: 1 ex. (ZMAN). - Netherlands: 291 ex.

(ZMAN, RIN, RMNH, Cuppen, Drost, Giessen, Huijbregts,

Jongema, Vallenduuk, Vondel). - Poland: 11 ex. (ZMAN,

RMNH, NHMW). — Spain: 2 ex. (NHMW). - Locality not

traced or unknown: 22 ex. (BMNH, NHMW, RMNH); 1 ex.

(lectotype of cinereus) (ISNB).

21. Haliplus maculatus Motschulsky

(figs. 199-207)

Haliplus maculatus Motschulsky, 1860: 100. Lectotype 9

(here designated), [USSR, n.w. of Caspian sea], ‘Bet-

scho-Ko’ [red label, hardly readable] ‘Haliplus macu-

latus Motsch., Des K. occ., 1295’ [red label] (ZMUM)

[examined ].

Haliplus syriacus Wehncke, 1880: 73. Lectotype 9 (here

designated) ‘Syria, Sharp’ (MNHN) [examined]. Syn.

Haliplus zimmermanni Gschwendtner, 1921: 56. Lecto-

type & (here designated) ‘Oberösterreich, L.

Gschwendtner, Umg. Linz, 5.5.1909, Type, Coll.

Gschwendtner, Haliplus Zimmermanni Gs, det.

VAN VONDEL: Palaearctic Liaphlus

Gschwendt, Type’

[examined]. Syn. n.

Haliplus maculatus; Zimmermann 1920: 317, Kolossov

1931: 116, Zaitsev 1953: 75, Ienistea 1978: 294.

Haliplus syriacus; Zimmermann 1920: 317, 1924: 136,

Guignot 1933: 244, Zaitsev 1953: 74, Burmeister

1985: 41.

Haliplus zimmermanni; Zimmermann 1924: 134, Müller

1926: 167, Guignot 1933: 241, Horion 1941: 365,

lenistea 1978: 295, Franciscolo 1979: 120.

[aedeagus missing] (OLML)

Diagnosis

This species closely resembles small specimens

of fulvus. The male can be distinguished from fu/-

vus by the narrow and somewhat pointed top of the

penis.

Description

Length 3.4-4.0 mm, width 1.8-2.2 mm. Body

oval, widest in the middle (fig. 199).

Head. - Yellow-brown to rust-coloured, vertex

usually slightly darkened, weakly to moderately

punctured, unpunctured band between the eyes.

Distance between the eyes 1.4-1.6 X eye-width.

Antennae yellow, first segment brown (fig. 200),

palpi yellow.

Pronotum. - Yellow to rust-coloured. Lateral

borders straight, finely bordered. Sparsely and

weakly, along the base strongly punctured, basal

punctures usually slightly darkened (fig. 199).

Elytra. - Yellow to rust-coloured, rather weak to

moderately strong primary puncture-rows, 28-38

punctures in the first row. Secondary punctures

moderately strong, accompanied by very small

punctures (fig. 201), usually all punctures dar-

kened. Markings variable, oblong blotches along

the suture, on the apex and on the even intervals,

the central blotches on the second and fourth inter-

val often connected anteriorly, the markings are

often hardly visible or even absent (fig. 199).

Ventral side. - Yellow-brown to rust-coloured,

elytral epipleura yellow to yellow-red, legs yellow-

red to rust-coloured, darkened towards the coxae.

Prosternal process slightly convex, rather sparsely

punctured, narrowed near the coxae (fig. 203), lat-

eral plicae complete (fig. 204). Metasternal process

flat with a shallow pit in the middle, sparsely punc-

tured (fig. 203). Coxal plates with moderately

strong and in between fine punctures, the strong

punctures absent on a wide band along the suture.

Setiferous striole on dorsal side of hind tibia about

% of the tibia length, with 7-10 punctures, longer

of the two tibial spurs about % the length of the

first tarsal segment (fig. 202).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 205-

207.

117

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

e e

Figs. 199-207. Haliplus maculatus, Vienna, Austria. - 199, dorsal view; 200, antenna; 201, elytral punctuation; 202,

dorsal side of hind tibia; 203, prosternal process; 204, lateral view of prosternal process; 205, left paramere; 206, penis;

207, right paramere.

118

Biology

In temporary pools, in brooks.

Distribution (fig. 300)

Austria, Hungary, Czechoslovakia, Rumania,

Yugoslavia, Italy? (identification uncertain), Po-

land?, Israel, Syria, Afghanistan, USSR near border

with Iran east of the Caspian sea and north of the

Black and the Caspian sea. All European specimens

I examined are from more than 60 years ago, so the

species might be extinct in Europe.

Remarks

The types of maculatus, syriacus and zimmer-

manni have been compared. Although the first two

are females and the important character of the

pointed top of the penis cannot be checked, I con-

sider the three conspecific.

Material examined. — 41 ex. - Afghanistan: 1 9, W.

Afghanistan, Hari Rud, Herat, 20.vii.1949, Danish Cen-

tral Asian Expedn, St. no. 4, Ex stomach of Lobivanellus

indicus, Haliplus fulvus, J. Balfour-Browne det. 1964

(BMNH). — Austria: 1 & (lectotype zimmermanni); 1 9,

Wien, Prater, Kniz (OLML); 2 6, Neusiedler See; 1 4,

Wien surr., Smolik; 2 9, Lavant Tal, Car (RMNH); 1 @,

Gars, Minarz; 5 6, 1 9, Wien, Prater; 1 @, surr. Wien; 1

6, 2 9, Steiermarken (Styria), Kars; 1 6, Klosterneuburg,

Pinker (NHMW). — Czechoslovakia: 1 &, Veseli (Moravia,

surr. Ostra), Th. v. Wanka (NHMW). - Hungary: 2 6,

Ujpest (RMNH); 1 @, Szekesfehervar (Szfehervar), 1928,

(OLML). — Israel: 1 6, Berekhat Sasa, 10.viii.1985, Ortal

& Jäch (Hujr); 2 9, Jerusalem, Sauliy (MNHN). — Italy: 1

©, Toscana, Dahl (RMNH). - Rumania: 1 4, Bocsa Vasovei

(N-Bogsan, Banat) (RMNH). - Syria: 1 9 (lectotype)

(MNHN). — Yugoslavia: 1 9, Fruska Gora (NHMW). -

USSR: 1 Q (lectotype of maculatus) (ZMUM); 1 9, Kopet-

Dagh, Germab (OLML); 2 9, Geok Tepe (Caucase, Guéox-

Tapa), L. Mesmin (Angus); 1 6, Geok Tepe (Caucase,

Gok Tepe), L.Mesmin (OLMI). - Locality not traced: 1 4,

Munganast, Type,1921, coll. Gschwendtner, Haliplus

Zimmermann Gschw., det. L. Gschw., Paralectotype

[aedeagus missing] (OLML). - Without locality: 2 3,

(OLML); 1 @, Ullr. (NHMW).

22. Haliplus maculipennis Schaum

(figs. 208-216)

Haliplus maculipennis Schaum, 1864: 107.

Haliplus maculipennis; Régimbart 1877: 354, Sahlberg

1913: 40, Zimmermann 1920: 312, 1924: 138, Guignot

1959: 37, Alfieri 1976: 38.

Diagnosis

This species can be distinguished from other

palaearctic species by the distance between the eyes

being about equal to eye-width and the row of dark

blotches on the elytral marginal parts.

VAN VONDEL: Palaearctic Liaphlus

Description

Length 3.0-3.3 mm, width 1.6-1.7 mm. Body

wide with subparallel sides (fig. 208).

Head. - Yellow-red with darker vertex, weakly to

moderately strongly punctured. Distance between

the eyes about eye-width. Antennae (fig. 209) and

palpi yellow-red.

Pronotum. - Yellow-red. Lateral borders

straight, strongly bordered, posterior corners usu-

ally protruding. Strong punctures along base (fig.

208).

Elytra. — Yellow-red, primary puncture-rows

weak except in the basal part of the first 3 or 4

rows, basal part of third row usually impressed,

about 30 punctures in the first row. Secondary

punctures weak (fig. 210). Extensive elytral mark-

ings: complete suture, a basal band until the 5th

puncture-row, triangular apically hooked central

blotch and several smaller blotches, 8th puncture-

row with a row of small blotches (fig. 208).

Ventral side. - Body yellow-red, legs yellow-red,

darkened towards the coxae. Prosternal process

wide, slightly narrowed near the coxae, strongly

impressed towards the apex, sparsely but strongly

punctured (fig. 212), lateral plicae complete (fig.

213). Metasternal process flat with a pit in the

middle, weakly punctured (fig. 212). Setiferous

striole about Y, X the tibia length, longer of the two

tibial spurs about *4 X the length of the first tarsal

segment (fig. 211).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 214-

216.

Distribution (fig. 306)

Egypt. The variety abyssinicus Régimbart in

Egypt, Sudan, Ethiopia, Zaire, Ivory Coast and

Bénin. Subspecies capensis Gueorguiev in South

Africa.

Remarks

In the collection of München (ZSMC) 1 Q is pres-

ent indicated as type: Egypt, Schaum, Typ, Samm-

lung Cl. Miiller, Haliplus maculipennis Schaum

[examined]. According to Guignot (1959) the type

should be in ZMHB. [not examined]. Also because

this species originates from the Ethiopian fauna I

postponed the designation of a lectotype and the

examination of the described varieties or subspe-

cies until a revision of the Ethiopian fauna is un-

dertaken.

Material examined. - 4 ex. - Egypt: 1 9, Schaum, Typ

(ZSMC); 1 g, Cairo, J.Sahlb. (NHMW).- Without locality: 1

9, 18 (ZSMC).

119

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 208-216. Haliplus maculipennis (208, 212-216, locality unknown; 209 -211, Cairo, Egypt). - 208, dorsal view;

209, antenna; 210, elytral punctuation; 211, dorsal side of hind tibia; 212, prosternal process; 213, lateral view of

prosternal process; 214, left paramere; 215, penis; 216, right paramere.

120

23. Haliplus mucronatus Stephens

(figs. 217-225)

Haliplus mucronatus Stephens, 1829: 40. Types probably

in BMNH [not examined].

Haliplus badius Aubé, 1836: 25. Types probably in MNHN

[not examined].

Haliplus parallelus Babington, 1836: 178. Types possibly

in CUMZ [not examined].

Haliplus siculus Wehncke, 1883: 145. Types probably in

MNHN [not examined].

Haliplus mucronatus, Schaum 1848: 42, Miiller 1900:

113, Apfelbeck 1904: 359, Fiori 1904: 193, Escalera

1914: 57, Zimmermann 1920: 312, 1924: 131, Guignot

1928: 143, 1933: 229, 1947: 49, 1959: 33, Scholz 1929:

14, F. Burmeister 1939: 213, Balfour-Browne 1940:

149, 1953: 5, Horion 1941: 365, Bertrand 1949: 25,

Zaitsev 1953: 67, Gueorguiev 1958: 44, 1981: 400,

Lagar Mascaro 1968: 72, Freude 1971: 15, Franciscolo

1972: 65, 1979: 115, Ali 1976: 92, Ienistea 1978: 294,

Fichtner 1981: 327, Leblanc 1984: 40, Vondel 1985:

33, E. Burmeister 1987: 177, Holmen 1987: 115.

Haliplus badius; Aubé 1838: 13, Schaum 1848: 42, Séidlitz

1887: 29, Zimmermann 1920: 312.

Haliplus parallelus, Schaum 1848: 42, Zimmermann

1920: 312.

Haliplus siculus; Seidlitz 1887: 29, Apfelbeck 1904: 359,

Scholz 1916: 165, Zimmermann 1920: 312.

Diagnosis

This species can be distinguished from related

species by the large distance between the eyes

(more than 2.3 X eye-width) and the metasternal -

process, not being grooved or pitted in the middle.

Description

Length 4.0-4.4 mm, width 1.9-2.2 mm. Body

subparallel (fig. 217).

Head. - Yellow-red to brown, punctures moder-

ately strong. Distance between the eyes 2.3-2.8 X

eye-width. Antennae (fig. 218) and palpi yellow-

red.

Pronotum. - Yellow-red. Lateral borders slightly

convex, weakly tapering anteriorly, finely bor-

dered. Densely and fairly strongly punctured, ba-

sally with a row of widened punctures, on the disc

with an unpunctured transverse band. Punctures

usually darkened (fig. 217).

Elytra. - Yellow-red without markings, all prim-

ary puncture-rows about equal in strength, 45-50

punctures in the first row. Secondary puncture-

rows well developed, number of punctures about

the same as in the primary rows. All punctures

darkened (fig. 219). Basal punctures of 3rd to 5th

row usually groove-like.

Ventral side. - Body yellow to yellow-red, legs

yellow to yellow-red. Prosternal process flat to

slightly impressed, strongly, apically even coarsely

punctured, gradually narrowed towards the coxae

(fig. 221), lateral plicae complete (fig.222). Metas-

ternal process slightly convex, sparsely punctured

VAN VONDEL: Palaearctic Liaphlus

(fig. 221). Setiferous striole on dorsal side of hind

tibia long, more than half of the tibia length, longer

of the two tibial spurs about two thirds the length

of the first tarsal segment (fig. 220).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 223-

225.

Biology

In fresh or brackish water, particularly in clay

and gravel pits with Characeans. The larva was

described by Bertrand (1928, as guttatus).

Distribution (fig. 294)

Central and southern Europe: south and sou-

theast of England, the Netherlands, Belgium,

France including Corsica, Spain including Mallorca,

the south of Germany, Switzerland, Italy including

Sardinia and Sicily, Yugoslavia, Albania, Greece,

Turkey, USSR along the Black Sea, Iraq, North

Africa along the coast from Morocco till Libya. The

record from the Harz mountains in East Germany

was confirmed by Fichtner (1981), but he is not

sure that the labelling is correct. The records from

Leningrad and the Kirghiz steppes are incorrect

(Zaitsev 1953).

Material examined. - 116 ex. - Albania: 4 ex. (NHMW).

- Algeria: 1 ex. (BMNH). - France: 26 ex. (NHMW, RMNH,

Jongema, ZMAN, Vondel). - Greece: 8 ex. (NHMW, Kanaar,

ZMAN). — Italy: 23 ex. (NHMW, CNCI, Jongema, ZMAN,

RMNH). - Netherlands: 21 ex. (ZMAN, RMNH, Vondel). —

Spain: 22 ex. (NHMW, CNCI, RMNH, ZMAN, Cuppen). —

Yugoslavia: 10 ex. (NHMW, RMNH, ZMAN). — Locality not

traced or unknown: 5 ex. (RMNH, NHMW).

24. Haliplus ortali sp. n.

(fig. 226-234)

Type material: Holotype 8, [Turkey] ‘TR, Bismil 6,

Diyarbakir, 29.5.1987 (35), Jach’ (NHMw). Paratypes (15

ex.): Israel: 1 @, 15.2.1982, Nesher quarry, rainpool,

Haifa, 15.ii.1982, R. Ortal & A. Sofer, IES 2575, Col. 1751-

01; 1 4, Nesher quarry, rainpool near Haifa, 27.1.1989, R.

Ortal, IES 5344; 1 8, Golan, Birket sinndiane, 11.ii.1981,

R. Ortal, IES 2502, Col. 2744; 1 9, Hula, Shosh pool,

12.1.1982, Z. Kuller & G. Gafny, IES 2599, Col. 1758; 1

6, 2 9, Berekhat, Nesher quarry, 26.xii.1989, R. Ortal, IES

5502; 1 g, Ein Abu Mashag, 12.ix.1989, IES 5401 (HUJI).

- Cyprus: 2 6, 4 9, Zakaki Marshes, ii.1948, G. Mavro-

moustakis (Angus). — Syria: 1 8, Damascus, brooklet

(MNHN).

Diagnosis

This species resembles very much species like

kulleri, jaechi or even abbreviatus, gafnyi and vil-

lierst. The males can be distinguished by the

121

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Netherlands. - 217, dorsal view; 218, antenna; 219, elytral punc-

Figs. 217-225. Haliplus mucronatus, Oostvoorne,

: 222, lateral view of prosternal process; 223, left

tuation; 220, dorsal side of hind tibia; 221, prosternal process

paramere; 224, penis; 225, right paramere.

122

VAN VONDEL: Palaearctic Liaphlus

. se eeoses

0 0 ec oe oe?

Figs. 226-234. Haliplus ortali, holotype. - 226, dorsal view; 227, antenna; 228, elytral punctuation; 229, dorsal side

of hind tibia; 230, prosternal process; 231, lateral view of prosternal process; 232, left paramere; 233, penis; 234, right

paramere.

123

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

‘Se e e oe o ©

lio a 0 200 S-

Figs. 235-243. Haliplus ovalis, holotype. - 235, dorsal view; 236, antenna; 237, elytral punctuation; 238, dorsal side

of hind tibia; 239, prosternal process; 240, lateral view of prosternal process; 241, left paramere; 242, penis; 243, right

paramere.

124

pointed digitus on the left paramere. In the

Turkish holotype that digitus is at the top provided

with 3 setae, while the Israeli paratypes have only

1 seta.

Description

Length 2.9-3.4 mm, width 1.5-1.8 mm. Body

oval, widest in the middle (fig. 226).

Head. - Yellow-brown to brown, slightly dar-

kened on the vertex, weakly punctured, unpunc-

tured band between the hind parts of the eyes.

Distance between the eyes 1.4-2.0 X eye-width.

Antennae yellow (fig. 227), palpi yellow.

Pronotum. - Yellow to yellow-brown on the disc,

lateral borders straight, darkened and finely bor-

dered. Width at the base 2.0 X the length in the

middle and 1.6 X the width in front. Weakly and

sparsely punctured, on the disc almost unpunc-

tured, along the base some larger darkened punc-

tures, along the base weakly impressed (fig. 226).

Elytra. - Yellow to yellow-brown, apical % of

suture darkened, clear discal blotch connected to

the dark suture or vague darkening and darkened

puncture-rows on the disc and vague blotches in

the apical half in the even intervals. Primary punc-

ture-rows fairly strong, about 32 punctures in the

first row. Secondary punctures moderately strong,

accompanied by very small punctures (fig. 228), all

punctures strongly darkened.

Ventral side. - Yellow to yellow-brown, pro- and

metasternal process brown, elytral epipleura yel-

low, legs yellow-brown, darkened towards the

coxae. Prosternal process slightly convex, nar-

rowed near the coxae, moderately strongly punc-

tured (fig. 230), lateral plicae sometimes inter-

rupted along the front coxae (fig. 231). Metasternal

process flat with a shallow impression in the mid-

dle, sparsely punctured (fig. 230). Coxal plates

rather densely punctured, small punctures between

the larger ones, sutural area only weakly and spar-

sely punctured. Hind margins of the fifth and sixth

abdominal sternite with only few punctures, coxal

plates reaching the fifth sternite. Setiferous striole

about ‘4 of the tibia length, with about 6 coarse

punctures, longer of the two tibial spurs % the

length of the first tarsal segment (fig. 229).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side, penis and parameres as in

figs. 232-234, left paramere with a pointed solid

digitus, at the top provided with 1 to 3 setae.

Etymology. - Named after Dr. R. Ortal who

collected part of the type-material.

Biology

In temporary pools and in brooklet.

VAN VONDEL: Palaearctic Liaphlus

Distribution (fig. 303)

South-east Turkey, Cyprus, Syria and Israel,

25. Haliplus ovalis Sharp

(figs. 235-243)

Haliplus ovalis Sharp, 1884: 440. Lectotype & (here de-

signated), ‘Japan, Jokohama, 29.10.81, Lewis, Type,

Haliplus ovalis Type D. S., Sharp Coll., 1905-313’

(BMNH) [examined].

Haliplus ovalis; Zimmermann 1920: 314, 1924: 134, Ta-

kizawa 1931: 141, Wu 1932: 343, J. Balfour-Browne

1946: 436, Sato 1960: 252, 1984: 3, Ohkura 1962: pl.

64, Nakane 1963: 55, 1985a: 63, 1985b: 40, 1987: 30,

Cho 1969: 166, Yoon 1988: 622.

Diagnosis

This species differs from the closely related chz-

nensis in having the distance between the eyes

more than 1.4 X the eye-width and in the male by

the abruptly bent top of the penis.

Description

Length 4.1-4.3 mm, width 2.1-2.2 mm. Body

oval, widest in the middle (fig. 235).

Head. - Yellow-brown, dark spot on vertex,

weakly punctured. Distance between the eyes 1.5 X

eye-width. Antennae (fig. 236) and palpi yellow-

red.

Pronotum. - Yellow, lateral borders straight,

finely bordered. Densely, on the disc very sparsely

punctured, basal punctures widened and darkened,

sometimes only slightly.

Elytra. - Yellow, moderately strong primary

punctures, about 40 in the first row. Secondary

punctures numerous. All punctures darkened (fig.

237). Suture, apical point and 9 or 10 spots dar-

kened, central spots on first and third interstriae

sometimes connected with suture (fig. 235).

Ventral side. - Body yellow to yellow-red, elytral

epipleura yellow, legs yellow-red, gradually dar-

kened towards the coxae. Prosternal process flat,

strongly and densely punctured, narrowed near the

coxae (fig. 239), lateral plicae complete (fig. 240).

Metasternal process flat with a pit in the middle,

rather weakly punctured (fig. 239). Setiferous stri-

ole on dorsal side of hind tibia almost half of the

tibia length, with about 14 punctures, longer of the

two tibial spurs two thirds the length of the first

tarsal segment (fig. 238).

Male sexual characters. - First three tarsal seg-

ment of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 241-

243.

Distribution (fig. 309)

Japan: Kyushu, Honshu. The record of Korea

(Takizawa 1931) probably concerns chinensis. The

125

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 244-252. Haliplus pulchellus, lectotype. - 244, dorsal view; 245, antenna; 246, elytral punctuation; 247, dorsal

side of hind tibia; 248, prosternal process; 249, lateral view of prosternal process; 250, left paramere; 251, penis; 252,

right paramere.

126

records of China (Zaitsev 1953) likely concern cht-

nensis.

Remarks

Types of the very closely related ovalis and chi-

nensis have been examined and compared. Unfor-

tunately the holotype of chinensis is a female. The

shape of the penis of the lectotype and other exam-

ined Japanese specimens of ovalis is different from

that of the continental specimens I have seen. So

far I consider the Japanese specimens to belong to

ovalis and the continental specimens to chinensis.

J. Balfour-Browne (see Sato 1984) already noticed

a difference in continental material and obviously

planned to describe Haliplus ovalis ssp. sinicus

from Djalantun, but that name is not known to be

published and is thus unavailable.

Material examined. — 11 ex. — Japan: 1 @ (lectotype)

(BMNH); 1 9, Tokio, Ahlwarth (ZMHB); 1 9, surr. Tokio,

Bodemeyer (RMNH); 1 @, Murakami, Niigata Pref., 6-

1l.viii.1961, M. Sato; 1 9, Kurokawa, N-Echigo,

S.viii.1961, M. Sato (ZMUC); 1 ex, Settsu, 5.v.1931; 1 ex.,

surr. Tokio, E. v. Bodemeyer; 1 ex., Lewis (SMFD); 1 ex.,

Yokohama, 28.ix-3.xi.1881, G. Lewis (MNHN); 2 ex., Ichi-

nokawa, Aso T., Kumamoto Pref., 28.viii.1986, E. Matsui

(Vondel).

26. Haliplus pulchellus Clark

(figs. 244-252)

Haliplus pulchellus Clark, 1863 : 418.

Haliplus oceanicus Régimbart, 1886 : 139.

Haliplus pulchellus var. indicus Régimbart, 1899 : 189.

Diagnosis

This species differs from other species in the

area by the small distance between the eyes and the

double furrowed prosternal process.

Description

Length 2.5-3.4 mm, width 1.5-2.0 mm. Body

oval, widest just before the middle (fig. 244).

Head. - Brown to dark brown, darkened along

the eyes and near the antennae, sometimes with a

weak arrow-like darkening in the middle. Sparsely

and weakly, behind the eyes more strongly punc-

tured. Distance between the eyes 0.9-1.1 X eye-

width. Antennae yellow to brown-yellow (fig. 245),

palpi yellow-red to brown.

Pronotum. - Yellow to yellow-red, large median

dark blotch that can be reduced to a round blotch

on the anterior half with a partly reduced or vague

or completely absent blotch on the posterior half

(fig. 244). Lateral borders straight to slightly con-

vex, anterior corners concave, clearly, vaguely or

not visibly bordered, margin never reaching the

VAN VONDEL: Palaearctic Liaphlus

anterior corner. Apical corners rounded, base near

the scutellum usually finely impressed. Strongly,

anteriorly weaker punctured, near the basal corners

with one or two stronger punctures in a slight

depression. Basal punctures darkened.

Elytra. - Yellow to yellow-red, clear and ex-

tended markings along the suture and along the

base until the fifth primary puncture-row, a central

blotch in the third interval connected to the suture

and various small blotches in the lateral part (fig.

244). Primary punctures of average strength,

stronger towards the base, about 30 punctures in

the first row. Secondary punctures relatively strong

accompanied by some very small punctures. All

punctures darkened except the central hole (fig.

246). Lateral borders sometimes slightly serrulate

in the basal part.

Ventral side. - Body yellow-brown to dark

brown, elytral epipleura yellow to yellow-brown,

legs yellow-brown with brown to dark-brown

femora and coxae. Prosternal process with two

strong grooves, bordered anteriorly, punctured,

sometimes strongly, especially in the grooves,

weakly narrowed near the coxae, slightly diverging

towards the apex (fig. 248), lateral plicae complete

(fig. 249). Metasternal process flat with two pits,

these pits sometimes very strong and than almost

forming one confluent pit, sparsely punctured (fig.

248). Coxal plates rather sparsely, near the suture

weakly punctured, sutural corners more or less rec-

tangular. Sternites with complete puncture-rows.

Last sternites almost completely covered with

strong punctures, apical point with a very short

ridge. Setiferous striole on dorsal side of hind tibia

very short, very close to the lateral row of setae and

often hardly to distinguish from it, longer of the

tibial spurs about % X the length of the first tarsal

segment (fig. 247).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 250-252.

Distribution (fig. 307)

China along the coast north to Xiamen; Thai-

land; Vietnam; Indonesia: Sumatra, Java, Sulawesi;

India; Sri Lanka.

Remarks

This species from oriental origin is included in

this revision because it penetrates into China, es-

pecially along the coast. The species will be treated

completely, however, in a revision of the Oriental

Haliplidae.

Material examined from the treated area: 2 ex., China,

127

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

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Figs. 253-261. Haliplus rubidus, France medit. - 253, dorsal view; 254, antenna; 255, elytral punctuation; 256, dorsal

side of hind tibia; 257, prosternal process; 258, lateral view of prosternal process; 259, left paramere; 260, penis; 261,

right paramere.

128

Amoy [Xiamen], Galatea (zMUC). The other material

will be treated in the revision of oriental Haliplidae.

27. Haliplus rubidus Perris

(figs. 253-261)

Haliplus rubidus Perris, 1857: 117. Types probably in

Ecol. Agricult. Montpellier, France [not examined].

Haliplus perforatus Schaum, 1859: 48. Depository of

types unknown [not examined].

Haliplus rubidus, Marseul 1882: 107, Seidlitz 1887: 31,

Zimmermann 1920: 314, 1924: 132, Guignot 1928:

143, 1933: 230, 1947: 50, 1959: 35, Scholz 1929: 15,

Burmeister 1939: 214, Lagar Mascaro 1968: 72, Ienis-

tea 1978: 294, Franciscolo 1979: 119.

Haliplus perforatus, Zimmermann 1920: 314.

Diagnosis

Closely related to flavicollis, from which it

differs in the larger punctures on the elytral base,

the more pronounced shoulders, the wider distance

between the eyes and the usually larger size.

Description

Length 3.9-4.4 mm, width 2.0-2.3 mm. Body

with strong shoulders, tapering strongly behind

the middle (fig. 253).

Head. - Yellow-red to rust-coloured, weakly

punctured. Distance between the eyes 1.7-1.9 X

eye-width. Antennae (fig. 254) and palpi yellow-

red to rust-coloured.

Pronotum. - Yellow-red to rust-coloured. Lateral

borders straight, clear darkened margins, narrowed

anteriorly. Densely and weakly, basally strongly

punctured, some basal punctures widened (fig. 253).

Elytra. - Yellow-red to rust-coloured, very strong

primary puncture-rows, near the base punctures

wider then the distance from each other (fig. 255),

about 30 punctures in the first row. Secondary

punctures moderately strong, 20 in the second in-

terval.

Punctures weakly darkened. No markings present.

Ventral side. - Yellow-red to rust-coloured, ely-

tral epipleura yellow-red, legs yellow-red to rust-

coloured, darkened towards the coxae. Prosternal

process flat or slightly impressed and very wide

apically, coarsely punctured, narrowed strongly

near the coxae (fig. 257), lateral plicae absent in

front of the coxae (fig. 258). Metasternal process

flat, strong pit in the middle, coarsely punctured

(fig. 257). Setiferous striole on dorsal side of hind

tibia more than half of the tibia length, longer of

the two tibial spurs little more than half the length

of the first tarsal segment (fig. 256).

Male sexual characters. — First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 259-

261.

VAN VONDEL: Palaearctic Liaphlus

Biology

Stagnant water in pools, dune-pools.

Distribution (fig. 299)

South of France along Atlantic ocean and along

the Mediterranean Sea, Italy in the province Ligu-

ria, Spain, Portugal, Morocco, Canary Islands.

Material examined. - 31 ex. - France: 1 ex., Lac d’Hour-

tin, Medoc, 28.ix.1973 (Jongema); 1 ex., Biarritz; 1 ex.,

Pyrenaei (ZMAN); 2 ex., Gallia m.; 2 ex., Gironde, Vaul-

oger; 1 ex., France Merid, Gironde, de Vauloger; 2 ex., F.

de la Teste de Vauloger (NHMW); 3 6, 2 9, Pyrenees,

Biarritz; 1 @, 2 9, Sauveterre, Gard, v.1928, Guignot; 1

ex.; 1 ex., Landes; 2 ex., Pyrenees (BMNH). - Portugal: 2

ex., Alentejo prov. Rio Mira nr. S. Ana da Serra,

21.ix.1976, no. 60, 37°38'N 8°13'W, v. Berge Hene-

gouwen (RMNH); 1 ex., Algarve, Silves, Ribeira de Arade,

10.11.1982; 1 ex, Algarve, Porto de Lagos, barrage,

12.11.1982 (Cuppen). - Morocco: 1 9, Tanger, Haliplus

fulvus (MHNG); 1 &, Tanger, vii.1895 (MNHN). — Locality

unknown: 1 ex. (BMNH).

28. Haliplus sbarpi Wehncke

(figs. 262-270)

Haliplus sharpi Wehncke, 1880: 74. Lectotype 9 (here

designated), ‘China, Sharp, Dr. Régimbart vidit 1898’

(MNHN) [examined]

Haliplus tsukushiensis Yoshimura, 1932: 102. Depository

of types unknown [not examined].

Haliplus sharpi; Sharp 1884: 440, Zimmermann 1920:

316, 1924: 137, Takizawa 1931: 142, Wu 1932: 343,

Zaitsev 1953: 76, Nakane 1963: 55, 1985a: 63, 1985b:

40, 1987: 30, Sato 1984: 3.

Haliplus tsukushiensis, Ohkura 1962: pl. 64, Nakane

1963: 55, 1985b: 40, Sato 1984: 3.

Diagnosis

This species could be confused with specimens of

diruptus in which a basal dark band is present on

the elytra. The prosternal process in sharp: is wide

and hardly narrowed near the coxae, while diruptus

has a narrow prosternal process, which is strongly

narrowed near the coxae. In the male the penis of

both species is clearly different.

Description

Length 3.6 mm, width 1.9 mm. Body oval, widest

before the middle (fig.262).

Head. - Yellow-brown with darkened vertex and

front-margin, moderately punctured. Distance be-

tween the eyes 1.3 X eye-width. Antennae yellow,

first segment yellow-brown, third segment clearly

longer than wide, fifth segment slightly longer

than fourth (fig. 263). Palpi yellow.

Pronotum. - Yellow. Lateral borders straight,

finely bordered. Sparsely, basally and anteriorly

more densely punctured, on each side of the base

129

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

„eens,

Figs. 262-270. Haliplus sharpi (262-267, lectotype; 268-270, from Korea). — 262, dorsal view; 263, antenna; 264,

elytral punctuation; 265, dorsal side of hind tibia; 266, prosternal process; 267, lateral view of prosternal process; 268,

left paramere; 269, penis; 270, right paramere.

130

with a row of partly widened and darkened punc-

tures (fig. 262).

Elytra. - Yellow to yellow-brown, rather strong

primary puncture-rows, about 36 punctures in the

first row. Secondary punctures in the first interval

strong and dense, in the other intervals with nor-

mal and very small punctures (fig. 264). All punc-

tures darkened. Distinct markings along the base,

suture and apical point, connected with blotches in

the second interval, blotches in the fourth and sixth

interval (fig. 262).

Ventral side. - Body yellow-red to brown-red,

abdominal segments darkened basally, elytral epi-

pleura yellow, legs brown-red, darkened towards

the coxae. Prosternal process flat, slightly im-

pressed in the apical part, densely punctured,

slightly narrowed near the coxae, widened towards

the apex (fig. 266), lateral plicae complete (fig.

267), anterior edge of prosternum strongly bor-

dered, pro-episternum unpunctured. Metasternal

process grooved and with a pit in the middle, den-

sely punctured, basally as wide as the prosternal

process (fig. 266). Coxal plates strongly punctured,

width of punctures less than the distance to each

other, accompanied by some very small punctures,

area along the suture impressed and only weakly

and sparsely punctured. Setiferous striole on dorsal

side of hind tibia about 4% X the tibia length, longer

of the two tibial spurs about % of the length of the

first tarsal segment (fig. 265).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs slightly widened, scaly

hairs on ventral side. Penis and parameres as in

figs. 268-270.

Biology

Stagnant pools. Caught at light and in a hotel

room.

Distribution (fig. 310)

China, prov. Jiangsu; Japan, Hokkaido and Shi-

koku; Korea.

Material examined. — 8 ex. - China: 1 9 (lectotype)

(MNHN ex coll. Wehncke); 1 ex., Jiangsu, Shanghai,

summer 1943, Marist Brothers (ISNB). — Japan: 1 9,

Japon, Heller, Dr. Régimbart vidit 1898, © Haliplus

sharpi Wehncke, 1880 (paralectotype) (MNHN ex coll.

Wehncke); 1 ex., Shikoku, Matsuyama, vii.1959, Light

trap, (ZMUC). - Korea: 28, 1 9, Sariwon,12-22.vii.1956,

M. Magyar, (ISNB); 1 9, Prov. South Pyongan, Pyongyan,

room of Hotel Te-dong, 30.vii.1975, J. Papp and A. Voj-

nits (CNCI); 1 @, 1 9, Kyongpuk-do, Yongpung-kun,

Hwayon-pond, 2.viii.1988, Sung Hwa Lee (Vondel).

29. Haliplus variegatus Sturm

(figs. 271-282)

VAN VONDEL: Palaearctic Liaphlus

Haliplus variegatus Sturm, 1834: 157. Syntypes probably

lost [as most Sturm-types] [not examined].

Haliplus subnubilis Babington, 1836: 177. Syntypes not

located [not examined].

Haliplus variegatus var. obsoletus Westhoff, 1881: 43.

Syntypes not located [not examined].

Haliplus variegatus ab. pallidior Müller, 1900: 115. Lec-

totype 8 (here designated) [Yugoslavia] ‘Castelnuovo,

Humler, TYPUS, Museo Civico di Trieste,

H.variegatus ab. pallidior M.’ (MSNT) [examined].

Haliplus pantherinus Sahlberg, 1900? (preoccupied).

Haliplus leopardinus Sahlberg, 1900: 183 (replacement

name for H. pantherinus). Lectotype @ (here desig-

nated) [Greece, Kérkira] ‘Corfu, J. Sahlb., Haliplus

pantherinus Sahlb. Type, Collect. Hauser, v. leopardi-

nus det. Ganglbauer’ (NHMW) [examined]

Haliplus transvolgensis Semenov, 1904: 216. Lectotype

(here designated) [USSR] ‘Nicolajevskij-Gorodok,

Saratov, 431.V.[19]00, Sacharov, Haliplus transvol-

gensis mTyp., A. Seminov det. (ZMAS)[examined].

Syn. n.

Haliplus variegatus subsp. galilea Sahlberg, 1913: 40.

Types not located [not examined].

Haliplus variegatus; Aubé 1836: 28, 1838: 16, Erichson

1837: 184, Schaum 1848:42, Thomson 1854: 485, Ré-

gimbart 1877: 354, Westhoff 1881: 43, Seidlitz 1887:

30, Miiller 1900: 114, A pfelbeck 1904: 360, Fiori 1904:

196, Sahlberg 1913: 39, Zimmermann 1920: 317,

1924: 136, Guignot 1928: 145, 1933: 231, 1947: 51,

Scholz 1929: 17, F. Burmeister 1939: 214, Balfour-

Browne 1940: 152, 1953: 5, Horion 1941: 366, Csiki

1946: 560, Zaitsev 1953: 73, Gueorguiev 1958: 44,

1960: 23, 1981: 401, Lindroth 1960: 46, Lagar Mascaro

1968: 73, Freude 1971: 15, Franciscolo 1972: 60, 1979:

121, Hosseinie 1974: 241, 1978: 167, Ienistea 1974:

208, 1978: 295, Ali 1976: 92, Galewski 1976: 29, Mate-

leshko 1977: 68, Fichtner 1981: 328, Holmen 1981: 2,

1987: 116, E. Burmeister 1982: 94, 1987: 184, Leblanc

1984: 8, Vondel 1985: 33, 1989a: 58, Lundberg 1986:

11, Zakharenko & Moroz 1988: 283, Nilsson 1988: 6.

Haliplus subnubilis; Schaum 1848: 42, Zimmermann

1920: 317.

Haliplus variegatus var. obsoletus; Guignot 1933: 244,

Zaitsev 1953: 73, Gueorguiev 1960: 23.

Haliplus variegatus ab. pallidior, Zimmermann 1920:

317, Scholz 1929: 17, Guignot 1933: 244, Zaitsev

1953: 74, Gueorguiev 1958: 44, 1960: 23, 1963: 216,

Ienistea 1978: 295, Leblanc 1984: 8.

Haliplus leopardinus, Apfelbeck 1904: 360, Sahlberg

1913: 39, Zimmermann 1920: 317, Döbler 1976: 182.

Haliplus transvolgensts; Zaitsev 1905: 37, Zimmermann

1920: 316 1924: 135, Kinel 1929: 219, Scholz 1929: 18,

Burmeister 1939: 214, Csiki 1946: 559.

Diagnosis

A very variable species in size, colour and mac-

ulation. The distance between the eyes, 1.7-2.0 X

eye-width, usually distinguishes it from related

species of the same size, especially in the Middle-

East.

Description

Length 2.6-3.4 mm, width 1.4-1.8 mm. Body

oval, widest before the middle (fig. 271).

131

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 271-279. Haliplus variegutus, Texel, Netherlands. - 271, dorsal view; 272, antenna; 273, elytral punctuation, 274,

dorsal side of hind tibia; 275, prosternal process; 276, lateral view of prosternal process; 277, left paramere; 278, penis;

279, right paramere.

132

Head. - Yellow to rust-coloured, darkened ver-

tex, rather weakly punctured, on the vertex with

stronger darkened punctures, between the eyes

with an unpunctured band. Distance between the

eyes 1.7-2.0 X eye-width. Antennae yellow to yel-

low-brown, third segment longer than the fourth

or fifth segment (fig. 272), palpi yellow to rust-

coloured.

Pronotum. - Yellow to rust-coloured, sometimes

anterior and basal margins slightly darkened. Lat-

eral borders straight, finely bordered. Sparsely, on

the disc almost not, basally strongly punctured,

basal punctures darkened (fig. 271).

Elytra. - Yellow to rust-coloured, rather strong

primary puncture-rows, 26-32 punctures in the

first row. Secondary punctures in the first interval

fairly strong, in the remaining intervals moder-

ately strong, accompanied by very small punctures.

All punctures darkened (fig. 273). Extensive mark-

ings on the disc, along the suture, on the apex and

on the even intervals with extensions to the odd

intervals. The central blotches are not always con-

nected to the suture (figs. 271, 280-282).

Ventral side. - Yellow-red to rust-coloured, ely-

tral epipleura yellow, legs yellow-red to rust-co-

loured, darkened towards the coxae. Prosternal

process flat or slightly convex, narrowed near the

coxae (fig. 275), lateral plicae complete (fig. 276),

rather coarsely punctured. Metasternal process

slightly to strongly and widely grooved, coarsely

punctured (fig. 275). Coxal plates sparsely, laterally

more densely punctured, with a few very small

punctures, apical point broadly rounded. Setiferous

striole about ‘4 of the tibia length, with about 5

punctures, longer of the two tibial spurs % the

length of the first tarsal segment (fig. 274). Hind

tibia about 3 X as long as the first tarsal segment.

VAN VONDEL: Palaearctic Liaphlus

Figs. 280-282. Haliplus

variegatus, elytra. — 280,

lectotype of H. variega-

tus var. pallidior, 281,

from Castelnuovo, Yu-

goslavia; 282, from

Trenton, England.

Male sexual characters. -- First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 277-

279)

Biology

In stagnant waterbodies with fresh water with

often a vegetation of Characeans, Sphagnum or

Hypnum, in temporary dune-pools, in peat bogs.

The third-stage larva was first described by

Schigdte (1864).

Distribution (fig. 296)

Europe: From Ireland and England to the central

and southern part of European USSR, northwards

to the south of Norway, Sweden and Finland and

southwards to Spain, Italy and Greece. Algeria,

Israel, Turkey, Syria. The records from Iraq (Ali

1976), Iran (Hosseinie 1974, 1978) and Afghanis-

tan (Gueorguiev 1963) most likely concern other

species like abbreviatus or villiersi. The record

from West Siberia (Zaitsev 1905, as transvolgen-

sis) probably concerns H. maculatus or angust.

Remarks

Examination of the type of transvolgensis

showed that it is conspecific with varzegatzs. In the

Mediterranean area small specimens are present,

described as pallidior or leopardinus, but in my

opinion these are conspecific with variegatus.

Material examined. - 425 ex. - Algeria: 2 ex. (BMNH).

— Austria: 49 ex. (ZMAN, RMNH, NHMW, RMNH). — Bel-

gium: 1 ex. (NHMW). - Denmark: 2 ex. (BMNH, ZMAN). —

England: 31 ex.BMNH, ZMAN, RMNH). — France: 69 ex.

(MNHN, ZMAN, RMNH, NHMW, Huijbregts, Vondel). - Ger-

many: 25 ex. (ZAMAN, RMNH, NHMW, BMNH). - Greece: 1

8 (lectotype leopardinus) (NHMW), 9 ex. (paralectotypes

155

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 283-291. Haliplus villiersi (283-288, holotype; 289-291, from type-locality). - 283, dorsal view; 284, antenna;

285, elytral punctuation; 286, dorsal side of hind tibia; 287, prosternal process; 288, lateral view of prosternal process;

289, left paramere; 290, penis; 291, right paramere.

134

leopardinus) (2 6,3 Q NHMW; 1 9 UZMH; 1 ex. OLML; 1

GB, 1 Q zsMC); 38 ex. (BMNH, ZMAN, NHMW, RMNH, Huij-

bregts). - Hungary: 7 ex. (ZMAN, NHMW). — Ireland: 3 ex.

(BMNH). - Israel: 2 6, Berekhat Zemorot nr. Ashquelon,

15.1.1981 (NHMW). — Italy: 15 ex. (BMNH, CNCI, NHMW,

Huijbregts, Jongema). — Netherlands: 108 ex (RMNH,

Schilthuizen, Cuppen, ZMAN, BMNH). — Poland: 6 ex.

(BMNH, ZMAN). — Spain: 4 ex. (ZMAN, NHMW). - Sweden:

2 ex. (BMNH). — Switzerland: 1 ex. (BMNH). — Turkey: 1

Q (MNHN). — USSR: 1 9, (lectotype transvolgensis)

(ZMAS). — Yugoslavia: 8 ex. (lectotype and 4 paralecto-

types of pallidior) (MSNT); 10 ex. (NHMW, CNCI). — Local-

ity not traced or unknown: 30 ex. (BMNH, ZMAN, MHNG,

RMNH, NHMW).

30. Haliplus villiersi Legros

(figs. 283-291)

Haliplus villiersi Legros, 1972: 573: Holotype @, ‘Iran,

Zabol, IV.1965, Museum Paris, Mission Franco-Ira- ‘

nienne 1965, A, Type, Haliplus (Liaphlus) villiersi n.

sp., C. Legros det.’ (MNHN) [examined].

Diagnosis

A variable species, that is closely related to ab-

breviatus and variegatus, from which it can be

distinguished by the usually impressed prosternal

process. Maybe abbreviatus and villiersi are con-

specific.

Description

Length 3.0-3.4 mm, width 1.7-2.0 mm. Body

oval, sides subparallel, shoulders a little pro-

nounced (fig. 283).

Head. - Yellow-brown, weakly punctured. Dis-

tance between the eyes about 1.6 X eye-width.

Antennae (fig. 284) and palpi yellow..

Pronotum. - Yellow-red, lateral borders straight,

narrowed anteriorly, finely bordered. Weakly,

along the base more strongly, on the disc more

sparsely punctured, basal punctures slightly dar-

kened (fig. 283).

Elytra. - Yellow to yellow-brown, rather weak to

moderately strong primary puncture-rows, 30 to 35

punctures in the first row. Punctures of 8th and 9th

row with long hairs. Secondary punctures in two

types: moderately strong or very weak (fig. 285).

All punctures darkened. Usually brown markings

on the suture, apex, even intervals and sometimes

in the first interval (fig. 283), the markings can be

very weak.

Ventral side. - Body yellow to yellow-red, elytral

epipleura yellow, legs yellow-red. Prosternal pro-

cess flat or slightly impressed apically, coarsely

punctured, slightly narrowed near the coxae (fig.

287), lateral plicae complete (fig. 288). Metasternal

process flat or grooved in the middle and with a

central pit (fig. 287), weakly punctured. Setiferous

VAN VONDEL: Palaearctic Liaphlus

striole on dorsal side of hind tibial about one third

of the tibia length, longer of the two tibial spurs %

the length of the first tarsal segment (fig.286).

Male sexual characters. - First three tarsal seg-

ments of fore- and midlegs widened, scaly hairs on

ventral side. Penis and parameres as in figs. 289-

291.

Distribution (fig. 305)

Iran.

Material examined. - 6 ex. - Iran: 1 & (holotype); 1 6,

4 ©, Zabol, 19.iv.1965, Mission Franco-Iranienne 1965 (1

Q allotype) (MNHN)

Unnamed material

The specimens I failed to determine are listed

below. Some of them might belong to still undes-

cribed species.

Species near H. diruptus. - [China] ‘Yunnan

Fou, Yunnan, R. Mouchamps det. 63, Haliplus di-

ruptus B.Br. (1 9, 1 @ without aedeagus) (ISNB).

Belonging to Haliplus diruptus or H. davidi or to

an undescribed species. - [China, Beijing] ‘Peping,

Collector C. F. Wu, Col. Gschwendtner, simplex

Clark Gschwendtner’ (2Q)(OLML). Possibly be-

longing to H. diruptus or H. davidi. - [China]

‘Yunnan Fou, Yunnan, H. Perrot; ® (MNHN). Pos-

sibly belonging to H. diruptus or H. davidi.

Species near H. variegatus. - [USSR] ‘Ross. O.

(2 92) (BMNH). Possibly belonging to an undes-

cribed species.

Species near H. ortali. - [Iran] ‘Perse, Susa,

iv.1899, Escalera, Haliplus sp.x., Dr. Régimbart

vidit 1900 (1 2) (MNHN). Possibly belonging to H.

ortali.

ACKNOWLEDGEMENTS

I am very grateful to Dr. R. Angus (Englefield

Green) for his valuable help and for critically read-

ing the manuscript.

I wish to express my sincere thanks to the fol-

lowing persons for placing material or information

at my disposal: Dr. G. Alberti (Trieste, MSNT), Dr.

L. Baert (Brussels, ISNB), Dr. A. L. van Berge Hene-

gouwen (The Hague, MOG), Dr. O. Bistròm (Hel-

sinki, UZMH), Dr. M. Brancucci (Basel, NHMB), Dr.

M. J. D. Brendell (London, BMNH), mr. B. Brugge

(Amsterdam, ZMAN), Dr. J. G. M. Cuppen (Ede),

Dr. R. Danielsson (Lund, MZLU), Dr. K. Desender

(Brussels, ISNB), mr. M. B. P. Drost (Wadenoijen),

Dr. J. P. Duffels (Amsterdam, ZMAN), Dr. Th. G.

Giessen (Gaanderen), Dr. F. Gusenleitner (Linz,

OLML), Dr. F. Hieke (Berlin, ZMHB), mr. M. Hiel-

kema (Gouda), Dr. S. J. Hine (London, BMNH), mr.

135

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

M. Holmen (Copenhagen, ZMUC), Dr. J. Huijbregts

(Leiden, RMNH), Dr. M. Jäch (Vienna, NHMW), mr.

Y. Jongema (Wageningen), Dr. B. Korotyaev (Le-

ningrad, ZMAS), Dr. J. Krikken (Leiden, RMNH), Dr.

P. Leblanc (Piney), Dr. Sung Hwa Lee (Taegu), Dr.

L. LeSage (Ottawa, cncı), Dr. P. Lindskog (Stock-

holm, NHRS), Dr. I. Löbl (Genève, MHNG), Dr. K.

Majer (Brno, MMBC), Dr. E. Matsui (Hondo City),

Dr. T. Nakane (Chiba), Dr. R. Ortal (Jerusalem,

HUJI), mrs. H. Perrin (Paris, MNHN), Dr. R. Poggi

(Genua, MCSN), mr. EF F. Repko (Leersum, RIN), Dr.

G. Scherer (Munich, zsMc), Dr. M. Schilthuizen

(Leiden), Dr. A. Smetana (Ottawa, CNCI), mr. G.

Stobbe (Leeuwarden, FNML), Dr. R. zur Strassen

(Frankfurt am Main, sMFD), mr. H. Vallenduuk

(Boxtel). The Uyttenboogaart-Eliasen Foundation

is acknowledged for financial support.

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Received: 10 October 1990

Accepted: 3 December 1990

139

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Fig. 292. Distribution of Haliplus fulvus (dots: material examined, stars: literature records).

140

VAN VONDEL: Palaearctic Liaphlus

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141

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 297-299. Distribution of Haliplus spp. 297, H. guttatus (dots: material examined, stars: literature records); 298,

H. andalusicus (dots: material examined, stars: literature records); 299, H. rubidus (dots: material examined, stars:

literature records).

142

VAN VONDEL: Palaearctic Liaphlus

Figs. 300-306. Distribution of Haliplus spp. 300, H. maculatus (dots: material examined, stars: literature records);

301, H. dalmatinus (dots: material examined); 302, H. jaechi (dot) and H. kulleri (star: material examined); 303, H.

gafnyi (stars) and H. ortali (dots); 304, H. angusi (dot) and H. astrakanus (star); 305, H. abbreviatus (dots: material

examined, stars: literature records) and H. villiersi (cross: material examined); 306, H. maculipennis (dot: material

examined).

143

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 134, 1991

Figs. 307-312. Distribution of Haliplus spp. 307, H. pulchellus (dots: material examined); 308, H. eximis (dots:

material examined, stars: literature records); 309, H. chinensts (dots: material examined) and H. ovalis (cross: material

examined); 310, H. kotoshonis (dot: material examined, star: literature records) and H. sharpi (cross: material

examined); 311, H. diruptus (dots: material examined, stars: literature records) and H. basinotatus (cross: material

examined, square: literature records); 312, H. excoffieri (star), H. davidi (dot) and H. holmeni (cross).

144

Tijdschrift voor Entomologie

Volume 134, no. |

Articles

| C. Gielis

Pterophorus Schaffer in the Ethiopian region (Lepidoptera: Pterophoridae:

Pterophorinae): new species and checklist.

9 R. V. Hensen

Review of Malesian Sphecina (Hymenoptera, Sphecidae, Sphecinae).

31 P. Huemer

Caryocolum delphinatella (Constant) sp. rev., a senior synonym of C. fiorii

(Klimesch) (Lepidoptera: Gelechiidae).

35 I. Lansbury

Cuticular blades and other structures of Diaprepocoris Kirkaldy and Stenocorixa

Horvath (Heteroptera: Corixidae).

47 N. Nieser & P. Chen

Naucoridae, Nepidae and Notonectidae, mainly from Sulawesi and Pulau

Buton (Indonesia). Notes on Malesian aquatic and semiaquatic bugs

(Heteroptera), |.

69 R. Puplesis, S. Seksjaeva & V. Sruoga

Leaf-mining Lepidoptera (Nepticulidae, Bucculatricidae, Gracillariidae) from

Ulmus in northern Caspiya (Kaspia).

75 B.J. van Vondel

Revision of the palaearctic species of Haliplus subgenus Liaphlus Guignot

(Coleoptera: Haliplidae).

Book reviews

68 Medvedev, G. S. (ed), 1990. Keys to the Insects of the European Part of the

USSR. Volume IV. Lepidoptera, Part 2. English Translation. [E. J. van

Nieukerken].

74 Emmet, A. M., 1991. The scientific names of the British Lepidoptera. Their

history and meaning. [E. J. van Nieukerken].; Rubtsov, |. A., 1990. Blackflies

(Simuliidae). - Fauna of the USSR. Diptera, Vol. 6, Part 6. [J. van Tol].

Published | July 1991 ISSN 0040-7496

Volume 134, no. 2, 1991 a): NEEN 0040-7494

Tijdschrift

voor

Entomologie

A journal of systematic and evolutionary

entomology since 1858

Published by the Nederlandse Entomologische Vereniging o