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HARVARD UNIVERSITY

Ernst Mayr Library

of the Museum of

Comparative Zoology

Volume 140, no. I, 1997 AL UNS ia 0040-74

Tijdschrift

voor

Entomologie

A journal of systematic and evolutionary

entomology since 1858

Netherlands Journal of Entomology

Published by the Netherlands Entomological Society -

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), N. E. Stork (London) and M. R. Wilson (Cardiff).

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 9517, 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

c/o Instituut voor Taxonomische Zodlogie

Plantage Middenlaan 64

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The Netherlands

Subscription price per volume Hfl. 300,— (postage included).

Special rate for members of the society. Please enquire.

Instructions to authors

Published with index of volume 139 (1996).

Graphic design

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

PETER F. BELLINGER & WixcLem N. ELLIS

California State University, Northridge California & Institute for Systematics en M | Pepto

L IBR ARY

Biology, Amsterdam

re OM © ME ET?

U 4 17 7/

GENERIC NAMES OF COLLEMBOLA:

SUPPLEMENT 1984 — 1996

H ARVARD

UNIVE = RSITY

Bellinger, P. F. & W. N. Ellis, 1997. Generic names of Collembola: supplement 1984-1996. —

Tijdschrift voor Entomologie 140: 1-11 [rssn 0040-7496]. Published 31 October 1997.

This is a second update to the list of Collembolan generic names published by the authors in

1973, covering names and nomenclatural actions that have come to our attention since 1984

and through 1996. Some 134 new generic names are included, with bibliographic citations and

indications of type species. Some corrections to the earlier lists and some comments on previ-

ously described genera are included. The new name Linasa is proposed to replace Silana Yoshii

& Suhardjono, preoccupied.

Correspondence: Peter F. Bellinger, California State University, Northridge, 18111 Nordhoff

Street, Northridge, California 91330, U.S.A.

Key words. — Collembola; nomenclature.

Our previous lists of generic names of Collembola

were published in 1973 and 1984. Proposal of new

names has continued, and a few earlier entries require

modification for various reasons. All new names have

again been checked against Neave's Nomenclator

Zoologicus and its supplements, and against the lists

of new generic and subgeneric names appearing an-

nually in part 20 of the Zoological Record; as a con-

sequence, we propose one new name here to replace a

junior homonym.

We take this opportunity to acknowledge the assis-

tance of our colleagues who have sent copies of their

papers. We particularly wish to thank Miguel Alonso-

Zarazaga for calling our attention to a potentially se-

rious problem in nomenclature, to Dr. Marcia

Edwards for comments on this matter, and to Dr.

Ryozo Yoshii for authorizing the renaming of his pre-

occupied name Silana.

As in our first supplement, we have marked new

taxa with an asterisk * and the gender of the names is

abbreviated as F(emale), M(ale) and N(euter).

Acherongia Massoud & Thibaud, 1985*

Annls Soc. ent. Fr.(2)21: 40.

Type species: A. minima Massoud & Thibaud,

1985, l.c., by original designation. F

Albanura Deharveng, 1982

Trav. Lab. Ecobiol. Arthr. edaph. Toulouse 3(4): 13.

Type species: Neanura (Deutonura) nana

Cassagnau & Peja, 1979, by original designation. F

Note: This is the valid proposal of the name

Albanura, incorrectly credited to Cassagnau in the

first supplement.

Allocyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44, ut subgenus

Ascocyrtus Yosii, 1963.

Type species: Acanthurella lepidornata Handschin,

1930, by original designation. M

Note: the junior author's name is given as ‘Yayuk

R. S. in the table of contents of the issue.

Allonychiurus Yoshii, 1995*

Ann. spel. Inst. Japan

Onychiurus Gervais, 1841.

Type species: Onychiurus flavescens Kinoshita,

1916, by original designation. M

13: 8, ut subgenus

Americabrya Mari Mutt & Palacios-Vargas, 1987*

J. N. Y. ent. Soc. 95(1): 100.

Type species: Janetschekbrya arida Christiansen &

Bellinger, 1980, by original designation. F

Antarctophorus Potapov, 1992*

Revue Ecol. Biol. Sol 28(4): 492.

Type species: Anurophorus subpolaris Salmon,

1962, by original designation. M

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Anurachorutes Kuznetsova & Potapov, 1988*

Zool. Zhurn. 67 (12): 1836.

Type species: A. martynovae Kuznetsova &

Potapov, 1988, l.c., by original designation. M

[ Archifurca Richards, 1956* ]

Thesis: 89.

Type species: A. coachellae Richards, 1956, l.c., by

original designation. F

Note: since Richards' thesis is technically unpub-

lished these names have no standing in nomenclature.

Austrocyrtus Yoshii, 1989*

Contr. biol. Lab. Kyoto Univ. 27: 234, ut sub-

genus Pseudosinella Schäffer, 1897.

Type species: P. (A.) speciosa Yoshii, 1989, l.c., by

original designation. M

Austrodontella Ellis & Bellinger, 1973

Note: Possible junior objective synonym of

Tridontella Stach, 1949; see below.

Austroseira Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2:23, ut subgenus Seira

Lubbock, 1869.

Type species: Lepidocyrtus schaefferi Schott, 1901,

by original designation. F

Bagnallophorus Weiner, 1996*

Annls Soc. ent. Fr. (N. S.) 32 (2): 183.

Type species: Onychiurus normalis Gisin, 1949, by

original designation. M

Batikphysa Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 74, ut subgenus

Callyntrura Börner, 1906.

Type species: C. (B.) batik Yoshii & Suhardjono,

1989, L.c., by original designation. F

Bessoniella Deharveng & Thibaud, 1989*

Bull. Mus. natl Hist. nat. Paris (4) 11A (2): 398.

Type species: B. procera Deharveng & Thibaud,

1989, L.c., by original designation. F

Blasconurella Deharveng & Bedos, 1992*

Trop. Zool. 5 (2): 300.

Type species: B. arcuata Deharveng & Bedos,

1992, l.c., by original designation. F

Blissia Rusek, 1985*

Can] Zoo! 6310):2077

Type species: B. glabra Rusek, 1985, l.c., by origi-

nal designation. F

Botryanura Chelnokov, 1987*

Ent. Obozr. 66 (1): 100, ut subgenus Entomobrya

Rondani, 1861.

Type species: E. (B.) oleniensis Chelnokov, 1987,

l.c., by original designation. F

Boudinotia Weiner & Najt, 1991*

Mem. Mus. natl Hist. nat. Paris (A) 149: 121.

Type species: B. prima Weiner & Najt, 1991, l.c.,

by original designation. F

Caledonura Deharveng, 1988*

Mem. Mus. natl Hist. nat. Paris (A) 142: 49.

Type species: C. tillierae Deharveng, 1988, l.c., by

original designation. F

Calvinura Cassagnau, 1988*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 5(4):

23.

Type species: C. besucheti Cassagnau, 1988, l.c., by

original designation. F

Camerounura Cassagnau, 1991*

Revue Ecol. Biol. Sol 28 (2): 221.

Type species: C. delamarei Cassagnau, 1991, l.c.,

by original designation. F

Cansilianura Dallai & Fanciulli, 1983*

Redia 66: 235.

Type species: C. malatestai Dallai & Fanciulli,

1983, l.c., by original designation. F

Carocyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44, ut subgenus

Acrocyrtus Yosii, 1959.

Type species: Lepidocyrtus (Acrocyrtus) carosus

Yosii, 1959, by original designation. M

See note on authorship under Allocyrtus, above.

Catalanura Deharveng, 1982*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 3(4): 15.

Type species: Neanura (Deutonura) catalana

Deharveng, 1979, by original designation. F

Note: this is the valid proposal of the name incor-

rectly credited to Cassagnau in the first supplement.

Caucasanura Kuznetsova & Potapov, 1988*

Zool. Zhurn. 67 (12): 1837.

Type species: C. stebaevae Kuznetsova & Potapov,

1988, l.c., by original designation. F

Caufrenyllodes Greenslade & Deharveng, 1984*

J. Aust. ent. Soc. 23 (3): 224.

Type species: C. bacilloformis Greenslade &

Deharveng, 1984, L.c., by original designation. M

Cephalachorutes Bedos & Deharveng, 1991*

Tijdschr. Ent. 134 (2): 145.

Type species: C. asiaticus Bedos & Deharveng,

1991, l.c., by original designation. M

Chaetobella Cassagnau, 1983*

Nouv. Revue Ent. 13 (1): 16, 19.

Type species Lobella (Propeanura) numatai Yosii,

1966, by original designation. F

Chaetophorura Rusek, 1976

Note: This name was misspelled ‘Chaetaphorura in

the first supplement (p.4).

Chihuahuachorutes Palacios-Vargas, 1990*

Folia ent. mex. 79: 7.

Type species: C. escobarae Palacios-Vargas, 1990,

l.c., by original designation. M

Chirolavia Deharveng, 1991*

Raffles Bull. Zool. 39 (1): 54.

Type species: C. murphyi Deharveng, 1991, l.c., by

original designation. F

Chribellphorura Weiner, 1996*

Annls Soc. ent. Fr. (N. S.) 32 (2): 174.

Type species: Onychiurus (Archaphorura) allanae

Christiansen & Bellinger, 1980, by original designa-

tion. F.

Christobella Fjellberg, 1985*

Ent. scand. Suppl. 21: 116.

Type species: Neanura ornata Folsom, 1902, by

original designation. F

Cinctocyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44, ut subgenus

Ascocyrtus Yosii, 1963.

Type species: Lepidocyrtus cinctus Schäffer, 1898,

by original designation. M

See note on authorship under Allocyrtus, above.

Coenaletes Bellinger, 1985*

Carib. J. Sci. 21 (3/4): 118.

Type species: Actaletes vangoethemi Jacquemart,

1980, by original designation. M

Cordobaia Simön, Bach, & Gaju, 1987*

Eos 62 (1/4): 303.

Type species: C. bembezarensis Simön, Bach, &

Gaju, 1987, l.c., by original designation. F

Coreanura Deharveng & Weiner, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 4(4):

Type species: C. filiae Deharveng & Weiner, 1984,

l.c., by monotypy. F

BELLINGER & ELLIS: Collembola

Cribrochiurus Weiner, 1996

Annls Soc. ent. Fr. (N.S.) 32(2): 189.

Type species: Onychiurus cribrosus Gisin, 1957, by

original designation. M

Note: as mentioned here by Weiner, and by Ellis &

Bellinger (1973), the name Cribrochiurus was pro-

posed by Palissa (1964) without a type species; the

name was therefore unavailable at that time.

Cyprania Bretfeld, 1992*

Mitt. zool. Mus. Kiel, Suppl. 4: 45.

Type species: C. gisae Bretfeld, 1992, l.c., by origi-

nal designation. F

Dableyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44,55, ut subgenus

Ascocyrtus Yosii, 1963.

Type species: Lepidocyrtus dahlii Schäffer, 1898, by

original designation. M

See note on authorship under Allocyrtus, above.

Deboutevillea Murphy, 1965

Incorrect original spelling (genus named for C.

Delamare Deboutteville) of Debouttevillea Murphy,

1965.

Unavailable name.

Deharvengiurus Weiner, 1996*

Annls Soc. ent. Fr. (N. S.) 32 (2): 183.

Type species: Onychiurus argus Denis, 1924, by

original designation. M

Deuteraphorura Absolon, 1901

Note: Weiner, 1996, Annls Soc. ent. Fr. (N. S.) 32

(2): 184, states that the type of this genus is

‘Onychiurus fimetarius (L.) (auct.) Stach, 1934’, by

Stach's designation [in 1954]. This is contrary to the

selection by Ellis & Bellinger, 1973, of Onychiurus

ambulans (L.). The implications of this statement are

complex and will be treated elsewhere.

Deuterobella Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 5, ut subgenus Lobella

Börner, 1906.

Type species: L. imadatei Yosii, 1976, by original

designation. F

Digitanura Deharveng, 1987*

Bull. Annls Soc. r. belge Ent. 123(4/6): 125.

Type species: D. quadrilobata Deharveng, 1987,

l.c., by original designation. F

Elgonura Cassagnau, 1984*

Trav. Lab. Écobiol. Arthr. édaph. Toulouse 4(3): 5.

Type species: E. erinacea Cassagnau, 1984, l.c., by

original designation. F

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fissuraphorura Rusek, 1991*

Acta ent. bohemoslov. 88: 145.

Type species: F. cubanica Rusek, 1991, l.c., by

original designation. F

Folsomia Willem, 1902

Add to entry:

The genus and its type species, Æ candida, were

placed on the respective official lists by Opinion

1421, in connection with the suppression of the

name Entomobrya cavicola Banks, 1897, a senior sub-

jective synonym of candida.

Formosanochiurus Weiner, 1986*

2“internat. Sem. Apterygota: 93.

Type species: Onychiurus formosanus Denis, 1929,

by original designation. M

Ghirkanura Kuznetsova & Potapov, 1988*

Zool. Zhurn. 67 (12): 1839.

Type species: G. chernovae Kuznetsova & Potapov,

1988, Lc. by original designation. F

Gnathofolsomia Deharveng & Christian, 1984*

Verh. zool.-bot. Ges. Osterreich 122: 97.

Type species: G. palpata Deharveng & Christian,

1984, L.c., by original designation. F

Haloxenylla da Gama & Deharveng, 1984*

Bull. Soc. Hist. nat. Toulouse 120: 131.

Type species: Xenylla affiniformis Stach, 1929, by

original designation. F

Hazaranura Cassagnau, 1991*

Trav. Lab. Écobiol. Arthr. édaph. Toulouse 6(4): 16.

Type species: H. elongata Cassagnau, 1991, l.c., by

original designation. F

Hemilobella Deharveng & Greenslade, 1992*

Invert. Taxon. 6 (3): 729.

Type species: H. rounsevelli Deharveng & Green-

slade, 1992, l.c., by original designation. F

Himalmeria Cassagnau, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 4(3):

Type species: H. (H.) gurung Cassagnau, 1984, l.c.,

by original designation. F

Hyperlobella Cassagnau, 1988*

Trav. Lab. Écobiol. Arthr. édaph. Toulouse 5(4):

22, 24.

Type species: Protanura kraepelini Börner, 1906,

by original designation. F

Intermediurida Najt, Thibaud & Weiner, 1990*

Bull. Mus. natl Hist. nat. Paris (4)12A (1): 95,

114.

Type species: Z anophthalma Najt, Thibaud &

Weiner, 1990, l.c., by original designation. F

Irianella Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 41, ut subgenus

Pseudoparonella Handschin, 1925.

Type species: P. (4) halmaherae Yoshii &

Suhardjono, 1992, l.c., by original designation. F

Isotogastrura Thibaud & Najt, 1992*

Bonner zool. Beitr. 43 (4): 545.

Type species: 1. arenicola Thibaud & Najt, 1992,

l.c., by original designation. F

Jailolaphorura Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 9, ut subgenus

Onychiurus Gervais. 1841.

Type species: O. (7) jailolonis Yoshi &

Suhardjono, 1992, l.c., by original designation. F

Javaphysa Yoshii, 1992*

Mem. Mus. Victoria 53:

Callyntrura Borner, 1906.

Type species: C. (/.) javana Yoshii, 1992, l.c., by

original designation. F

131, ut subgenus

Koreanurina Najt & Weiner, 1992*

Pan-Pacif. Ent. 68 (3): 200.

Type species: K. szeptyckii Najt & Weiner, 1992,

Le, by original designation. F

Lanocyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44, 58, ut subgenus

Lepidocyrtus Bourlet, 1839.

Type species: Podura lanuginosa Gmelin, 1790, by

original designation. M

Lanzhotia Rusek, 1985*

Acta ent. bohemoslov. 82 (3): 175.

Type species: L. brachycera Rusek, 1985, l.c., by

original designation. F

Lasofinius Ireson & Greenslade, 1990*

J. Aust. ent. Soc. 29: 205.

Type species: L. willi Ireson & Greenslade, 1990,

Lc. by original designation. M

Lawrenceana Mitra, 1993*

Rec. zool. Survey India, Occ. Pap. 154: 34.

No species included.

Unavailable name.

Leenurina Najt & Weiner, 1992*

Pan-Pacif. Ent. 68 (3): 203.

Type species: L. jasii Najt & Weiner, 1992, l.c., by

original designation. F

Linasa nom. nov.*

New name for Si/ana Yoshii & Suhardjono, 1992,

nec Spaeth, 1914.

Type species: Salina (Silana) moluccana Yoshii &

Suhardjono, 1992. F

Madecassiella Betsch & Waller, 1996*

Bull. Soc. ent. Fr. 101 (4): 416.

Type species: M. ibityensis Betsch & Waller, 1996,

by original designation. F

Marginobrya Yoshii, 1992*

Acta zool. Asiae orient. 2:

Entomobrya Rondani, 1861.

Type species: E. (M.) marginifera Yosii, 1992, l.c.,

by original designation. F

100, ut subgenus

Merapicyrtus Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 82, ut subgenus

Ascocyrtus Yosii, 1963.

Type species: A. (M.) merapicus Yoshii & Suhar-

djono, 1992, l.c., by original designation. M

Mexicanura Palacios-Vargas, 1984

An. Esc. nac. Cienc. biol. Mex. 27: 69.

Lapsus calami for Americanura Cassagnau, 1983.

Unavailable name.

Monolamellus Lee, 1979*

Korean J. Ent. 9 (2); 5, ut subgenus Zomocerus

Nicolet, [1842].

Type species: Tomocerus laxalamella Lee, 1975, by

monotypy.

Published without diagnosis in the combination 7.

(M.) laxalamella, nomen nudum.

Unavailable name.

Mucrella Fjellberg, 1985*

Ent. scand. Suppl.

Hypogastrura Bourlet, 1839.

Type species: H. (M.) denali Fjellberg, 1985, Lc.

by original designation. F

21:12, ut subgenus

Multivesicula Rusek, 1982*

Vestn. cesk. Spol. zool. 46(1): 33.

Type species: M. columbica Rusek, 1982, l.c., by

original designation. F

Nahuanura Palacios-Vargas & Najt, 1986*

Folia ent. mex. 68: 18.

Type species: N. ce Palacios-Vargas & Najt, 1986,

BELLINGER & ELLIS: Collembola

l.c., by original designation. F

Najtia Arlé & Mendonga, 1986*

Revista bras. Ent. 30(1): 1.

Type species: /sotoma vicaria Arlé, 1959, by origi-

nal designation. F

Najtiaphorura Weiner & Thibaud, 1991*

Bull. Mus. natl Hist. nat. Paris (4) 13A (1-2): 113.

Type species: N. sandrinae Weiner & Thibaud,

1991, l.c., by original designation. F

Najtnella Yosii, 1989*

Contr. biol. Lab. Kyoto Univ. 27 (3): 249, ut sub-

genus Pseudoparonella Handschin, 1925.

Type species: P. novaecaledoniae Yosii, 1960, by

original designation. F

Najtsira Yosii, 1989*

Contr. biol. Lab. Kyoto Univ. 27 (3): 236, ut sub-

genus Lepidosira Schött, 1925.

Type species: L. (N.) najtae Yoshii, 1989, l.c., by

original designation. F

Narisa Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 37, 38, ut subgenus

Salina MacGillivray, 1894

Type species: S. pallens Yoshii, 1984, by original

designation. F

[ Nasopus Richards, 1956* ]

Thesis: 86.

Type species: Smynthurus spinatus MacGillivray,

1893, by original designation. M

Unpublished objective synonym of Pseudo-

bourletiella Stach, 1956. Unavailable name.

Neokatianna Snider, 1989*

Florida Ent. 72 (3): 541, ut subgenus Sminthurinus

Börner, 1901.

Type species: S. (N.) acantholaema Snider, 1989,

Le, by original designation. F

Nepalimeria Cassagnau, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 4 (3):

ls Bs Oe

Type species: N. dal Cassagnau, 1984, L.c., by orig-

inal designation. F

Nusasira Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 38, ut subgenus

Lepidosira Schott, 1925.

Type species: Pseudosira (Mesira) javana Borner,

1913, by original designation. F

See note on authorship under Allocyrtus, above.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Oceaniella Yoshii, 1989*

Contr. biol. Lab. Kyoto Univ. 27 (3): 261, ut sub-

genus Pseudoparonella Handschin, 1925.

Type species: P. (O.) najtae Yoshii, 1989, l.c., by

original designation. F

Octodontophora Chelnokov, 1990*

Ent. Obozr. 69 (2): 343.

Type species: O. ornata Chelnokov, 1990, l.c., by

original designation. F

Onerocyrtus Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 44, 45, ut subgenus

Acrocyrtus Yosii, 1959.

Type species: Lepidocyrtus onerosus Yoshii, 1982, by

original designation. M

See note on authorship under A//ocyrtus, above.

Ongulogastrura Thibaud & Massoud, 1983*

Mém. biospéol. 10: 317.

Type species: O. longisensilla Thibaud & Massoud,

1983, L.c., by original designation. F

Ongulonychiurus Thibaud & Massoud, 1986*

Bull. Mus. natl Hist. nat. Paris (4) 8A (2): 327.

Type species: O. colpus Thibaud & Massoud,

1986, L.c., by original designation. M

Orchesella Templeton, 1835

Trans. ent. Soc. London 1 (2): 92.

Type species: O. filicornis Templeton, 1835, by

subsequent designation by Rothschild, 1878, Musée

entomologique illustré 3: 365. F

Note: the type designation by Rothschild antedates

that of O. cincta by Börner, 1903, cited in the List, p.

37. Fortunately O. filicornis and O. cincta are general-

ly regarded as synonymous. In reply to an inquiry to

the International Commission we are informed that

the citation of type species for Orchesella on the

Official List has been altered to read ‘Orchesella fil-

icornis Templeton, 1835 = O. cincta (Linnaeus,

158)

Placed on the Official List of Generic Names in

Zoology: 723, Opinion 291.

Palaeosminthurus Pierce & Gibron, 1962

Note: according to Najt, 1987, Revue fr. Ent. (N.

S.) 9: 152, the type and only included species, P. juli-

ae, belongs to Formicidae rather than Collembola.

Paracerura Deharveng & Oliveira, 1994*

Revue suisse Zool. 101 (2): 442.

Type species: P. virgata Deharveng & Oliveira,

1994, l.c., by original designation. F

Paralobella Cassagnau & Deharveng, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 5 (1): 9.

Type species: P. orousseti Cassagnau & Deharveng,

1984, Lc. by original designation. F

Paramanura Cassagnau, 1986*

2" int. Sem. Apterygota: 315.

Type species: P. najtae Cassagnau, 1986, l.c., by

original designation. F

Paranurida Skarzynski & Pomorski, 1994*

Genus 5 (4): 363.

Type species: P. kopetdagi Skarzynski & Pomorski,

1994, L.c., by original designation. F

Parectonura Deharveng, 1988*

Mém. Mus. natl Hist. nat. Paris (A) 152: 46.

Type species: P. ciliata Deharveng, 1988, l.c., by

original designation. F

Parthenaphorura Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 31, ut subgenus

Onychiurus Gervais, 1841.

Type species: O. (P.) bogoriensis Yoshii &

Suhardjono, 1989, I. c., by original designation. F

See note on authorship under Allocyrtus, above.

Pectenisotoma Gruia, 1983*

Rés. Expéd. biospéol. cubano-roum. 4: 193.

Type species: P. theodori Gruia, 1983, l.c., by orig-

inal designation. F

Penelopella Cassagnau, 1986*

2“int. Sem. Apterygota: 315.

Type species: P. pacifica Cassagnau, 1986, l.c., by

original designation. F

Philotella Najt & Weiner, 1985*

Annls Soc. ent. Fr. 21 (1): 29.

Type species: P. deharvengi Najt & Weiner, 1985,

l.c., by original designation. F

Phradmon Greenslade & Deharveng, 1991*

Invert. Taxon. 5 (4): 840.

Type species: P. australasiae Greenslade &

Deharveng, 1991, l.c., by original designation. M

Plumosinella Gruia, 1984*

Trav. Inst. Spéol. “Emil Racovitza’ 23: 21, ut sub-

genus Pseudosinella Schäffer, 1897.

Type species: Pseudosinella (P.) kenchristianseni

Gruia, 1984, l.c., by original designation. F

Pongeiella Rusek, 1991*

Acta Soc. zool. bohemoslov. 55: 65.

Type species: Tullbergia (T.) falca Christiansen &

Bellinger, 1980, by original designation. F

Probrachystomellides Weiner & Najt, 1991*

Bonner zool. Beitr. 42 (3/4): 376.

Type species: P. nicolaii Weiner & Najt, 1991, l.c.,

by original designation. M

Psammisotoma Greenslade & Deharveng, 1986*

Proc. R. Soc. Queensland 97: 89.

Type species: P. kingae Greenslade & Deharveng,

1986, l.c., by original designation. F

Psammophorura Thibaud & Weiner, 1994*

Polskie Pismo ent. 63: 8.

Type species: P. gedanica Thibaud & Weiner,

1994, l.c., by original designation. F

Pseudoxenyllodes Kuznetsova & Potapov, 1988*

Zool. Zhurn. 67 (12): 1833, ut subgenus

Xenyllodes Axelson, 1903.

Type species: X. (P.) macrocanthus Kuznetsova &

Potapov, 1988, l.c., by original designation. M

Rambutanura Deharveng, 1988*

Can. J. Zool. 66 (3): 714.

Type species: R. yoshiiana Deharveng, 1988, l.c.,

by original designation. F

Rambutsinella Deharveng & Bedos, 1996*

Raffles Bull. Zool. 44 (1): 280.

Type species: À honchongensis Deharveng &

Bedos, 1996, |. c., by original designation. F

Raponella Najt, 1988*

Nouv. Revue Ent. (N. S.) 5 (3): 205.

Type species: Probrachystomella dodecophthalma

Najt, 1988, l.c., by original designation. F

Rotundiphorura Rusek, 1991*

Acta ent. bohemoslov. 88: 152.

Type species: R. habanica Rusek, 1991, l.c., by

original designation. F

Rusekella Deharveng, 1982*

Bull. Soc. Hist. nat. Toulouse 118: 235.

Type species: À. cantabrica Deharveng, 1982, l.c.,

by original designation. F

Sahacanthella Potapov & Stebaeva, 1994*

Misc. zool. 17: 135.

Type species: S. kele Potapov & Stebaeva, 1994,

l.c., by original designation. F

Salvarella Greenslade & Najt, 1987*

Revue fr. Ent. (N.S.) 9(3): 115.

Type species: S. wallacei Greenslade & Najt, 1987,

l.c., by original designation. F

BELLINGER & ELLIS: Collembola

Secotomodes Potapov, 1988*

Zool. Zhurn. 67 (1): 144.

Type species: S. sibiricus Potapov, 1988, l.c., by

original designation. M

Sericeotoma Potapov, 1991*

Acta zool. cracov. 34 (1): 287, ut subgenus /sotoma

Bourlet, 1839.

Type species: Z. (S.) knissi Potapov, 1991, l.c., by

original designation. F

Siamanura Deharveng, 1987*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 5 (2):

1922

Type species: S. clavata Deharveng, 1987, l.c., by

original designation. F

Sibiracanthella Potapov & Stebaeva, 1994*

Misc. zool. 17: 130.

Type species: S. nuda Potapov & Stebaeva, 1994,

l.c., by original designation. F

Sibirisotoma Rusek, 1991*

Acta ent. bohemoslov. 88: 33.

Type species: S. stebajevae Rusek, 1991, l.c., by

original designation. F

Silana Yoshii & Suhardjono, 1992*

Acta zool. Asiae orient. 2: 28, ut subgenus Salina

MacGillivray, 1894; nec Spaeth, 1914, Deutsche ent.

Ztg. 1914: 563 (Coleoptera).

Type species: Salina (Silana) moluccana Yoshii &

Suhardjono, l.c., by original designation. F

Note: the spelling Sirana on p. 40 is an obvious

lapsus calami, but in any case is preoccupied by Sirana

Griveaud, 1975, Bull. Soc. ent. Fr. 807/8): 232

(Lepidoptera).

Junior homonym; replaced by Linasa nom. nov.

Singalimeria Cassagnau, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 4(3):

16%;

Type species: S. pachyderma Cassagnau, 1984, l.c.,

by original designation. F

Sinnamarides Betsch, 1991*

Revue Ecol. Biol. Sol 28 (2): 230.

Type species: S. delamarei Betsch, 1991, l.c., by

original designation. F

Speleophysa Bagnall, 1969*

In Glennie, Trans. Cave Res. Group Great Britain

11(2): 107. Nomen nudum.

Type species: S. varioseta Bagnall, 1969, l.c., by

monotypy. Nomen nudum. F

Unavailable name.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Spinactaletes Soto Adames, 1989*

Carib. J. Sci. 24 (3/4): 174.

Type species: S. calcalectoris Soto Adames, 1989,

l.c., by original designation. M

[ Spinaphorura Arbea, 1987* |

Thesis [not seen]; Diss. Abs. int. C 49(2): 214

(1988).

Type species: ? Spinaphorura clavata Arbea, 1987. F

Note: according to Jordana et al., 1990, Publ. biol.

Univ. Navarra, Ser. Zool.21:129, Spinaphorura clava-

ta is a species of Neonaphorura.

Unpublished. Unavailable name.

Spinonychiurus Weiner, 1996*

Annls Soc. ent. Fr. (N. S.) 32 (2): 188.

Type species: Onychiurus edinensis Bagnall, 1935,

by original designation. M

Stenaphorurella Luciänez & Simén, 1992*

Eos 68(2): 108.

Type species: Stenaphorura quadrispina Börner,

1901, by original designation. F

Stenomeria Cassagnau, 1990*

Annls Soc. ent. Fr. (N. S.) 26 (1): 29.

Type species: S. deharvengi Cassagnau, 1990, Lc.

by original designation. F

Sturmius Bretfeld, 1994*

J. Zool. syst. Evol. Res. 32: 264.

Type species: S. epiphytus Bretfeld, 1994, l.c., by

original designation. M

Sundasira Yoshii & Suhardjono, 1989*

Acta zool. Asiae orient. 1: 38, ut subgenus

Lepidosira Schött, 1925.

Type species: L. (S.) sundana Yoshii & Suhardjono,

1989, l.c., by original designation. F

See note on authorship under Allocyrtus, above.

Tamulmeria Cassagnau, 1988*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 5(5):

3}, AG.

Type species 7° callipygos Cassagnau, 1988, l.c., by

original designation. F

Tasphorura Greenslade & Rusek, 1996*

SystrEnt 21-28;

Type species: 7. vesiculata Greenslade & Rusek,

1996, l.c., by original designation. F

Thibaudichiurus Weiner, 1996*

Annls Soc. ent. Fr. (N. S.) 32 (2): 189.

Type species: Onychiurus mariangeae Thibaud &

Lee, 1994, by original designation. M

Tillieria Weiner & Najt, 1991*

Mem. Mus. natl Hist. nat. Paris (A) 149: 123.

Type species: T. insularis Weiner & Najt, 1991,

l.c., by original designation. F

Tridontella Stach, 1949

Apterygotan fauna of Poland 2, Neogastruridae

and Brachystomellidae: 317.

Type species: Odontella trispina Womersley, 1935,

by original designation.

This name was originally proposed, on p. 261 of

the same volume, as Triodontella; the misspelling or

variant spelling 7ridontella appears on p. 317 and

340, as noted in the original List (p. 56). Since

Triodontella Stach is a junior homonym of

Triodontella Reitter, 1919, we proposed (1973, p. 7)

the replacement name Austrodontella. However, in

the Nomenclator Zoologicus vol. 6 (1966), p. 300,

the spelling Zridontella is treated as valid, with

Triodontella noted as a variant spelling. According to

one of the authors of this volume of the Nomenclator

(M. Edwards, in litt.), any action taken beyond bibli-

ographical listing was unintentional. The question of

the validity of this and perhaps other names appar-

ently validated in the Nomenclator has been called to

the attention of the International Commission.

Tritosminthurus Snider, 1988*

Ent. News 99(5): 260.

Type species: 7. schuhi Snider, 1988, l.c., by origi-

nal designation. M

[ Trochopilosa Richards, 1956* |

Thesis: 96, ut subgenus Allacma Börner, 1906.

Type species: A. (7°) allacma Richards, l.c., by orig-

inal designation. F

Note: since Richards’ thesis is technically unpub-

lished, these names have no standing in nomencla-

ture.

Troglobius Palacios-Vargas & Wilson, 1990*

Int. J. Speleol. 19: 68.

Type species: 7. coprophagus Palacios-Vargas &

Wilson, 1990, l.c., by original designation. M

Venezuelida Diaz & Najt, 1995*

Bull. Mus. natl Hist. nat. Paris (4) 16A (2-4): 428.

Type species: V. polylepiana Diaz & Najt, 1995,

l.c., by original designation. F

Vietsira Yoshii, 1994*

Acta zool. Asiae orient. 3: 27.

Type species: V. squamigera Yoshii, 1994, l.c., by

original designation. F

Weinera Thibaud, 1993*

Revue fr. Ent. (N. S.) 15 (2): 73.

Type species: W. ghislaineae Thibaud, 1993, l.c.,

by original designation. F

Willemgastrura Oliveira & Thibaud, 1988*

Amazoniana 10 (3): 300.

Type species: W. coeca Oliveira & Thibaud, 1988,

l.c., by original designation. F

Yetimeria Cassagnau, 1984*

Trav. Lab. Ecobiol. Arthr. édaph. Toulouse 4 (3):

154

Type species: Y. lama Cassagnau, 1984, L.c., by

original designation. F

Zelandanura Deharveng & Wise, 1987*

Rec. Auckland Inst. Mus. 24: 143.

Type species: Z. bituberculata Deharveng & Wise,

1987, L.c., by original designation. F

REFERENCES

Ellis, W. N. & P. F. Bellinger, 1973. An annotated list of

the generic names of Collembola (Insecta) and their type

species. — Monographieën Nederlandsche Entomo-

logische Vereeniging 7: 1-74.

Ellis, W. N. & P. F. Bellinger, 1984. Generic names of

Collembola: supplement 1973-1983. — Tijdschrift voor

Entomologie 127 (1): 1-15.

INDEX TO SPECIES

acantholaema Snider, 1989 (Sminthurinus

(Neokatianna)) Neokatianna Snider, 1989

affıniformis Stach, 1929 (Xenylla). — Haloxenylla da

Gama & Deharveng, 1984

[allacma Richards, 1956 (Allacma (Trochopilosa)). —

Trochopilosa Richards, 1956]

allanae Christiansen & Bellinger, 1980 (Onychiurus

(Archaphorura)). — Chribellphorura Weiner, 1996

anophthalma Najt, Thibaud, & Weiner, 1990 (Inter-

mediurida). — Intermediurida Najt, Thibaud, &

Weiner, 1990

arcuata Deharveng & Bedos, 1992 (Blasconurella). —

Blasconurella Deharveng & Bedos, 1992

arenicola Thibaud & Najt, 1992 (Isotogastrura). —

Isotogastrura Thibaud & Najt, 1992

arida Christiansen & Bellinger, 1980 (Janet-

schekbrya). — Americabrya Mari Mutt & Palacios-

Vargas, 1987

asiaticus Bedos & Deharveng, 1991 (Cephalacho-

rutes). — Cephalachorutes Bedos & Deharveng,

1991

australasiae Greenslade & Deharveng, 1991 (Phrad-

mon). — Phradmon Greenslade & Deharveng, 1991

bacilloformis Greenslade & Deharveng, 1984. —

BELLINGER & ELLIS: Collembola

(Caufrenyllodes). — Caufrenyllodes Greenslade &

Deharveng, 1984

batik Yoshii & Suhardjono, 1989 (Callyntrura (Batik-

physa). — Batikphysa Yoshii & Suhardjono, 1989

bembezarensis Simon, Bach, & Gaju, 1987 (Cordo-

baia). — Cordobaia Simén, Bach, & Gaju, 1987

besucheti Cassagnau, 1988 (Calvinura). — Calvinura

Cassagnau, 1988

bituberculata Deharveng & Wise, 1987 (Zelanda-

nura). — Zelanadanura Deharveng & Wise, 1987

bogoriensis Yoshii & Suhardjono,1989 (Onychiurus

(Parthenaphorura)). — Parthenaphorura Yoshii &

Suhardjono, 1989

brachycera Rusek, 1985 (Lanzhotia). — Lanzhotia

Rusek, 1985

calcalectoris Soto Adames, 1989 (Spinactaletes). —

Spinactaletes Soto Adames, 1989

callipygos Cassagnau, 1988 (Tamulmeria). — Tamul-

meria Cassagnau, 1988

candida Willem, 1902 (Folsomia). — Folsomia Willem,

1902

cantabrica Deharveng, 1982 (Rusekella). — Rusekella

Deharveng, 1982

carosus Yosii, 1959 (Lepidocyrtus (Acrocyrtus)). — Caro-

cyrtus Yoshii & Suhardjono, 1989

catalana Deharveng, 1979 (Neanura (Deutonura)). —

Catalanura Deharveng, 1982

cavicola Banks, 1897 (Entomobrya). — Folsomia

Willem, 1902

ce Palacios-Vargas & Najt, 1986 (Nahuanura). —

Nahuanura Palacios-Vargas & Najt, 1986

chernovae Kuznetsova & Potapov, 1988 (Ghirka-

nura). — Ghirkanura Kuznetsova & Potapov, 1988

ciliata Deharveng, 1988 (Parectonura). — Parectonura

Deharveng, 1988

cincta Linnaeus, 1758 (Podura). — Orchesella Temple-

ton, 1835

cinctus Schäffer, 1898 (Lepidocyrtus). — Cinctocyrtus

Yoshii & Suhardjono, 1989

[clavata Arbea, 1987 (Spinaphorura). — Spinaphorura

Arbea, 1987]

clavata Deharveng, 1987 (Siamanura). — Siamanura

Deharveng, 1987

[coachellae Richards, 1956 (Archifurca). — Archifurca

Richards, 1956]

coeca Oliveira & Thibaud, 1988 (Willemgastrura). —

Willemgastrura Oliveira & Thibaud, 1988

colpus Thibaud & Massoud, 1986 (Ongulonychiurus).

— Ongulonychiurus Thibaud & Maddoud, 1986

columbica Rusek, 1982 (Multivesicula). — Multi-

vesicula Rusek, 1982

coprophagus Palacios-Vargas & Wilson, 1990 (77o-

globius). — Troglobius Palacios-Vargas & Wilson,

1990

cribrosus Gisin, 1957 (Onychiurus). — Cribrochiurus

Weiner, 1996

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

cubanica Rusek, 1991

Fissuraphorura Rusek, 1991

dahlii Schäffer, 1898 (Lepidocyrtus). — Dahleyrtus

Yoshii & Suhardjono, 1989

dal Cassagnau, 1984 (Nepalimeria). — Nepalimeria

Cassagnau, 1984

deharvengi Cassagnau, 1990

Stenomeria Cassagnau, 1990

deharvengi Najt & Weiner, 1985 (Philotella). —

Philotella Najt & Weiner, 1985

delamarei Betsch, 1991 (Sinnamerides). — Sinna-

merides Betsch, 1991

delamarei Cassagnau, 1991

Camerounura Cassagnau, 1991

denali Fjellberg, 1985 (Hypogastrura (Mucrella)). —

Mucrella Fjellberg, 1985

dodecophthalma Najt, 1988 (Raponella). — Raponella

Najt, 1988

edinensis Bagnall, 1935 (Onychiurus). — Spinony-

chiurus Weiner 1996

elongata Cassagnau, 1991

Hazaranura Cassagnau, 1991

epiphytus Bretfeld, 1994 (Sturmius). — Sturmius

Bretfeld, 1994

erinacea Cassagnau, 1984 (Elgonura). — Elgonura

Cassagnau, 1984

escobarae Palacios-Vargas, 1990 (Chihuahuachorutes).

— Chihuahuachorutes Palacios-Vargas, 1990

falca Christiansen & Bellinger, 1980 (Tullbergia

(Tullbergia)). — Pongeiella Rusek, 1991

filiae Deharveng & Weiner, 1984 (Coreanura). —

Coreanura Deharveng & Weiner, 1984

filicornis Templeton, 1835 (Orchesella). — Orchesella

Tempeton, 1835

fimetarius Linnaeus, 1767, nec 1758 (Podura). —

Deuteraphorura Absolon, 1901

flavescens Kinoshita, 1916 (Onychiurus). — Allony-

chiurus Yoshii, 1995

formosanus Denis, 1929 (Onychiurus). — Formosa-

nochiurus Weiner, 1986

gedanica Thibaud & Weiner, 1994 (Psammophorura).

— Psammophorura Thibaud & Weiner, 1994

ghislaineae Thibaud, 1993 (Weinera). — Weinera

Thibaud, 1993

gisae Bretfeld, 1992 (Cyprania). — Cyprania Bretfeld,

1992

glabra Rusek, 1985 (Blissia). — Blissia Rusek, 1985

gurung Cassagnau, 1984 (Himalmeria). — Himalmeria

Cassagnau, 1984

habanica Rusek, 1991 (Rotundiphorura). — Rotun-

diphorura Rusek, 1991

halmaherae Yoshii & Suhardjono, 1992 (Pseudo-

paronella (Irianella)). — lrianella Yoshii &

Suhardjono, 1992

honchongensis Deharveng & Bedos, 1996 (Rambut-

sinella). — Rambutsinella Deharveng & Bedos, 1996

(Fissuraphorura). = —

(Stenomeria). —

(Camerounura). —

(Hazaranura). -

ibityensis Betsch & Waller, 1996 (Madecassiella). —

Madecassiella Betsch & Waller, 1996

imadatei Yosii, 1976 (Lobella (Lobella)). — Deutero-

bella Yoshii & Suhardjono, 1992

insularis Weiner & Najt, 1991 (Tillieria). — Tillieria

Weiner & Najt, 1991

jailolonis Yoshii & Suhardjono, 1992 (Onychiurus

(Jailolaphorura)). — Jailolaphorura Yoshii &

Suhardjono, 1992

jasii Najt & Weiner, 1992 (Leenurina). — Leenurina

Najt & Weiner, 1992

javana Börner, 1913 (Pseudosira (Mesira)). — Nusasira

Yoshii & Suhardjono, 1989

javana Yoshii, 1992 (Callyntrura (Javaphysa)). —

Javaphysa Yoshii, 1992

juliae Pierce & Gibron, 1962 (Palaeosminthurus). —

Palaeosminthurus Pierce & Gibron, 1962

kele Potapov & Stebaeva, 1994 (Sahacanthella). —

Sahacanthella Potapov & Stebaeva, 1994

kenchristianseni Gruia, 1984 (Pseudosinella (Plumo-

sinella)). — Plumosinella Gruia, 1984

kingae Greenslade & Deharveng, 1986 (Psammi-

sotoma). — Psammisotoma Greenslade & Dehar-

veng, 1986

knissi Potapov, 1991 (Sericeotoma). — Sericeotoma

Potapov, 1991

kopetdagi Skarzynski & Pomorski, 1994 (Paranu-

rida). — Paranurida Skarzynski & Pomorski, 1994

kraepelini Börner, 1906 (Protanura). — Hyperlobella

Cassagnau, 1988

lama Cassagau, 1984 (Yetimeria). — Yetimeria Cas-

sagnau, 1984

lanuginosa Gmelin, 1790 (Podura). — Lanocyrtus

Yoshii & Suhardjono, 1989

[laxalamella Lee, 1979 Tomocerus (Monolamellus). —

Monolamellus Lee, 1979]

lepidornata Handschin, 1930 (Acanthurella). —

Allocyrtus Yoshii & Suhardjono, 1989

longisensilla Thibaud & Massoud, 1983 (Ongulo-

gastrura). — Ongulogastrura Thibaud & Massoud,

1983

macrocanthus Kuznetsova & Potapov, 1988

(Xenyllodes (Pseudoxenyllodes)). — Pseudoxenyllodes

Kuznetsova & Potapov, 1988

malatestai Dallai & Fanciulli, 1983 (Cansilianura). —

Cansilianura Dallai & Fanciulli, 1983

marginifera Yoshii, 1992 (Entomobrya (Margino-

brya)). — Marginobrya Yoshii, 1992

martynovae Kuznetsova & Potapov, 1988 (Anura-

chorutes). — Anurachorutes Kuznetsova & Potapov,

1988

merapicus Yosii & Suhardjono, 1992 (Ascocyrtus

(Merapicyrtus)). — Merapicyrtus Yosii & Suhar-

djono, 1992

minima Massoud & Thibaud, 1985 (Acherongia). —

Acherongia Massoud & Thibaud, 1985

moluccana Yoshii & Suhardjono, 1992 (Salina

(Silana)). — Linasa nom. nov. — Silana Yoshii &

Suhardjono, 1992

murphyi Deharveng, 1991 (Chirolavia). — Chirolavia

Deharveng, 1991

najtae Cassagnau, 1986 (Paramanura). — Paramanura

Cassagnau, 1986

najtae Yoshii, 1989 (Lepidosira (Najtsira)). — Najtsira

Yoshii, 1989

najtae Yoshii, 1989 (Pseudoparonella (Oceaniella)). —

Oceaniella Yoshii, 1989

nana Cassagnau & Peja, 1979 (Neanura (Deuto-

nura)). — Albanura Deharveng, 1982

nicolaii Weiner & Najt, 1991 (Probrachystimellides).

— Probrachystomellides Weiner & Najt, 1991

novaecaledoniae Yosii, 1960 (Pseudoparonella). —

Najtnella Yoshii, 1989

nuda Potapov & Stebaeva, 1984 (Sibiracanthella). —

Sibiracanthella Potapov & Stebaeva, 1984

numatai Cassagnau, 1983 (Lobella (Propeanura)). —

Chaetobella Cassagnau, 1983

oleniensis Chelnokov, 1987 (Entomobrya (Botrya-

nura)). — Botryanura Chelnokov, 1987

onerosus Yoshii, 1982 (Lepidocyrtus). — Onerocyrtus

Yoshii & Suhardjono, 1989

ornata Chelnokov, 1990 (Octodontophora). — Octo-

dontophora Chelnokov, 1990

ornata Folsom, 1902 (Neanura). — Christobella Fjell-

berg, 1985

orousseti Cassagnau & Deharveng, 1984 (Paralo-

bella). — Paralobella Cassagnau & Deharveng, 1984

pachyderma Cassagnau 1984 (Singalimeria). — Singa-

limeria Cassagnau, 1984

pacifica Cassagnau, 1986 (Penelopella). — Penelopella

Cassagnau, 1986

pallens Yoshii, 1984 (Salina). — Narisa Yoshii &

Suhardjono, 1992

palpata Deharveng & Christian, 1984

(Gnathofolsomia). — Gnathofolsomia Deharveng &

Christian, 1984

polylepiana Diaz & Najt, 1995 (Venezuelida). —

Venezuelida Diaz & Najt, 1995

prima Weiner & Najt, 1991

Boudinotia Weiner & Najt, 1991

procera Deharveng & Thibaud, 1989 (Bessoniella). —

Bessoniella Deharveng & Thibaud, 1989

quadrilobata Deharveng, 1987 (Digitanura). —

Digitanura Deharveng, 1987

quadrispina Börner, 1902. — Stenaphorurella Luciänez

& Simon, 1992

rounsevelli Deharveng & Greenslade, 1992

(Hemilobella). — Hemilobella Deharveng & Green-

slade, 1992

(Boudinotia).

BELLINGER & ELLIS: Collembola

sandrinae Weiner & Thibaud, 1991 (Najtiaphorura).

— Najtiaphorura Weiner & Thibaud, 1991

schaefferi Schött, 1901 (Lepidocyrtus). — Austroseira

Yoshii & Suhardjono, 1989

schuhi Snider, 1988 (Tritosminthurus). — Tritosminth-

urus Snider, 1988

sibiricus Potapov, 1988 (Secotomodes). — Secotomodes

Potapov, 1988

speciosa Yoshii, 1989 (Pseudosinella (Austrocyrtus)). —

Austrocyrtus Yoshii, 1989

[spinatus MacGillivray, 1893 (Smynthurus). — Naso-

pus Richards, 1956]

squamigera Yoshii, 1994 (Vietsira). — Vietsira Yoshii,

1994

stebaevae Kuznetsova & Potapov, 1988 (Caucasa-

nura) — Caucasanura Kuznetsova & Potapov,

1988

stebajevae Rusek, 1991 (Sibirisotoma). — Sibirisotoma

Rusek, 1991

subpolaris Salmon, 1942 (Anurophorus). — Antarcto-

phorus Potapov, 1992

sundana Yoshii & Suhardjono, 1989 (Lepidosira

(Sundasira)). — Sundasira Yoshii & Suhardjono,

1989

szeptyckii Najt & Weiner, 1992 (Koreanurina). —

Koreanurina Najt & Weiner, 1992

theodori Gruia, 1983 (Pectenisotoma). — Pectenisotoma

Gruia, 1983

tillierae Deharveng, 1988 (Caledonura). — Caledonura

Deharveng, 1988

trispina Womersley, 1935 (Odontella). — Tridontella

Stach, 1949

vangoethemi _ Jacquemart,

Coenaletes Bellinger, 1985

[varioseta Bagnall, 1969 (Speleophysa). — Speleophysa

Bagnall, 1969]

vesiculata Greenslade & Rusek, 1996 (Tasphorura). —

Tasphorura Greenslade & Rusek, 1996

vicaria Arlé, 1959 (Isotoma). — Najtia Arlé & Men-

donca, 1986

virgata Deharveng & Oliveira, 1994 (Paracerura). —

Paracerura Deharveng & Oliveira, 1994

wallacei Greenslade & Najt, 1987 (Salvarella). —

Salvarella Greenslade & Najt, 1987

willi Ireson & Greenslade, 1990 (Lasofinius). —

Lasofinius Ireson & Greenslade, 1990

yoshiiana Deharveng, 1988 (Rambutanura). —

Rambutanura Deharveng, 1988

1980 (Actaletes). —

Received: 5 September 1996

Accepted: 12 February 1997

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

MikHaiz V. KOZLOV

Laboratory of Ecological Zoology, University of Turku, Finland

EE NEW AURORASPECIES-GROUPTOERWHE GENUS

NEMOPHORA HOFFMANNSEGG, WITH TWO NEW

SPECIES FROM TAIWAN (LEPIDOPTERA, ADELIDAE)

Kozlov, M. V., 1997. The new aurora species-group of the genus Nemophora Hoffmannsegg,

with two new species from Taiwan (Lepidoptera, Adelidae). — Tijdschrift voor Entomologie

140: 13-16, figs. 1-12 [1ssn 0040-7496]. Published 31 October 1997.

Nemophora aurora sp. n. and N. uncella sp. n. described from specimens collected in Taiwan

form a monophyletic aurora species-group defined by two apomorphies: (1) dark brown later-

al bands of forewing fascia with glossy silver-grey internal zone, (2) presence of small swelling

bearing a tuft of long setae on oval depigmented spot at the base of valva in male genitalia. The

aurora species-group is closely related to degeerella species-group.

Mikhail V. Kozlov, Laboratory of Ecological Zoology, University of Turku, Turku 20014,

Finland.

Key words. — Nemomorpha; Adelidae; taxonomy; Taiwan.

Most of the East Palaearctic and Oriental species of

the genus Nemophora Hoftmannsegg, 1798, have

never been critically examined since Meyrick's (1912)

catalogue or their original description. This explains

the fact that only three of ca 20 species of this genus

collected in Taiwan are identified yet (Heppner

1992). Two further species of Nemophora, which pre-

sumably form a monophyletic unit introduced here as

aurora species-group, are described below.

The male genitalia were investigated and figured as

described by Kozlov (1993). The interocular index

was measured according to Davis (1975), being the

ratio between the vertical diameter of the compound

eye and the interocular distance measured at a point

of the frons midway between the base of the antennal

sockets and the anterior tentorial pits. Recent locality

names are added in square brackets, whenever possi-

ble.

The type specimens are deposited in the U. S.

National Museum of Natural History, Smithsonian

Institution, Washington, D. C., U.S.A. (usNM) and

the National Museum of Natural Science in

Taichung, Taiwan, China (NMNST).

Taxonomic position of N. auroraand N. uncella

Two species described below share the following

characters with degeerella species-group: (1)

Background forewing colour yellow to pale ochreous;

(2) Dark forewing pattern with predominance of lon-

gitudinal stripes; (3) Forewing fascia consists of tree

bands, the medial one being of the same colour as

forewing background; (4) Basal part of forewing with

characteristic pattern formed by silver-grey, dark

brown bordered longitudinal stripes along costa, M

and CuP veins, with some additional dark brown

lines between these stripes.

The phylogenetic significance of these traits is not

completely clear yet, and it can hardly be refined be-

fore the majority of Nemophora species is revised.

However, characters 1 to 4 are regarded as apomor-

phies; among them, character 4 is presumably aut-

apomorphic for the aurora and degeerella species-

groups.

Monophyly of the aurora species-group is support-

ed by the following synapomorphies: (5) Lateral

bands of forewing fascia dark brown, with internal

zone formed by silver-grey glossy scales; (6) Valva me-

dially with small basal swelling bearing a tuft of long

setae on small oval depigmented spot. Independent

appearance of character (6) was reported for N. ichno-

desma (Meyrick) (Kozlov 1995).

115)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Key to the species based on external characters and

male genitalia

1. External silver-grey band in forewing fascia com-

plete; vinculum short (< 3 X length of valva); me

dial margin of valva distally without prominent

lobe appearing laterally as a triangular protuber-

ACER RE ENR nr I aurora

— External silver-grey band in forewing fascia bro-

ken in the middle, and part adjacent to the costal

margin is hook-shaped in such a way that its dis-

tal part is parallel to the costal margin; this distal

part in some specimens is separated from the base

and forms a distinct spot; vinculum long (> 3 X

length of valva); medial margin of valva distally

with prominent lobe appearing laterally as a tri-

anoulanproruberaneer.. em nee uncella

Nemophora aurora sp. n.

(figs. 1, 3-7)

Type material. - Holotype d : Taiwan, Tyokakurai,

26.11.1944, S. Issiki (Issiki Collection, USNM). -

Paratypes: 34, same data as holotype. 2d, 19,

Taiwan, Baibara [? Nantou Co., Hsinseng],

24.11.1943, S. Issiki. 19, labelled as previous, except

for date ‘25.11.1943’. 19, labelled as previous, except

for date ‘27.11.1943’. 19, Taiwan, Tattaka [Nantou

Co., Tatachia], 7.VI.1943, S. Issiki. 26, Taiwan,

Musya [? Nantou Co., Lenai], 27.11.1943, S. Issiki.

1d, Taiwan, E.VII.1947, Chingsing Li (all in Issiki

Collection, UsNM). 16, 29, Taiwan, Natou Co.

Jean-ai, Songgang, 14.1V.1996, S. H. Yen (NMNST).

Description

Male. Forewing length 6.7-7.4 mm; wing expanse

14-15 mm. Vertex yellow, with admixture of brown

hair-like scales; frons yellow. Compound eyes not en-

larged (interocular index 0.6). Proboscis brownish

yellow, base covered with yellow scales. Labial palpus

long (about 1.6 X vertical eye diameter), yellow,

with brown tip and brown raised hair-like scales.

Antenna 3.5 X forewing length, with simple inward-

ly directed pegs on basal flagellar segments; scapus

and base of flagellum brown, then the colour of fla-

gellum changes clinally to grey. Tegula and thorax

(dorsum) bronze. Forewing (fig. 1) yellowish ochre-

ous, glossy bronze; basal area with three silver stripes

(along costal, medial and postcubital veins) and some

four or five dark brown lines. Apical quarter of

forewing with numerous (10 to 14) narrow dark

brown radial lines; in melanic specimens dark lines

wide, creating impression of narrow yellow lines on

dark brown background. Medial band of fascia yel-

low, bordered on both sides by wide ochreous bands

with diffuse margins; central zone of these bands with

Figs. 1-2. Nemophora, forewing pattern. — 1, N. aurora; 2,

N. uncella. Scale 1.0 mm.

glossy silver-grey zone. Internal margin of the proxi-

mal ochreous band straight, almost indistinct from

the ochreous background; external margin of the dis-

tal ochreous band convex in middle of forewing. Cilia

bronze. Hindwing base grey, apex brown, glossy pur-

plish; costal area light grey; cilia grey to bronze. Legs

yellowish-brown to brown; tibia and all tarsomeres

apically with yellow rings. Epiphysis at 1/2 of fore tib-

ia, almost reaching its tip. Abdomen brown.

Female. Forewing length 6.0-6.2 mm; wing ex-

panse 12.5-13.2 mm. Antenna 1.4 X forewing,

brown; scapus and base of flagellum glossy purplish.

Otherwise similar to male.

Male genitalia (figs. 3-7). Tegumen dome-shaped,

with sharp medial ridge. Socii elongate, about same

diameter as medial part of aedeagus. Vinculum with

straight lateral margins, tip rounded. Length of valva

c. 0.35 X length of vinculum. Medial margin of val-

va ventrally with emargination at 0.4 valvar length;

tip narrowly rounded. Valva medially with small (1/3

X length of valva, in lateral view); basal swelling

bearing tuft of long setae on small oval depigmented

spot. Valvae not accreted basally. Arrow-head of jux-

ta narrow (width about 1/2 X length), tip of arrow-

head pointed. Distal half of aedeagus membranous,

with two slightly curved narrow sclerotized bands;

aedeagus basally of the same width as medially.

Biology

Moths were collected in the evergreen forest domi-

nated by oak (Quercus spp.), with admixture of Litsea

acuminata, Cinnamomum spp., Castanopsis for-

mosana, Ilex spp., Acer kawakamii and Lithocarpus

spp. Moths fly very high above the canopy layer; max-

imum activity occurred at 9 - 10 a. m. (S. H. Yen,

pers. comm.).

Kozrov: Nemophora from Taiwan

Figs. 3-12. Nemophora, male genitalia. — 3, N. aurora, genital complex, ventral view; 4, same, genital complex, lateral view; 5,

same, juxta; 6, same, aedeagus, ventral view; 7, same, aedeagus, lateral view; 8, N. uncella, genital complex, ventral view; 9,

same, genital complex, lateral view; 10, same, juxta; 11, same, aedeagus, ventral view; 12, same, aedeagus, lateral view; scale

0.25 mm.

Nemophora uncella sp. n.

(figs. 2, 8-12)

Type material. — Holotype d : Taiwan, Hassenran

[? Taichung Co., Pahsienshan], 24.x.1929, S. Issiki

(Issiki Collection, USNM). Paratypes: 2d, 19, la-

belled as holotype (Issiki Collection, usnm). 1d,

Taiwan, Syôrei, 12.viii.1943, A. Mutuura (Issiki

Collection, USNM). 26, Taiwan, Hnalien Co.

Tayulin-Tzer-en, 15.vii.1995, S. H. Yen (NMNST).

1d, Taiwan, Natou Co., Jean-ai, Songgang,

14.iv.1996, S. H. Yen (NMNST). 29, Taiwan,

Taichung Co., Hoping, An-ma-shan, 1800 m,

23.41.1996, S. H. Yen (NMNST). 19, Taiwan,

Kaohsiung Co., Maoling, Teng-Jur Area, 2200 m,

15-15.iv.1996, W. Y. Chou (NMNST).

Description

Male. Forewing length 8.2-8.7 mm; wing expanse

17-18 mm. Vertex yellow, with sparse brown hair-

like scales; frons straw-yellow. Compound eyes not

enlarged (interocular index 0.6). Proboscis yellow,

with two lines of brown scales on the base of frontal

surface. Labial palpus long (about 1.8 X vertical eye

diameter), straw-yellow, tip dorsally brown. Antenna

3.8-4.1 X forewing length, with simple inwardly di-

rected pegs on basal flagellar segments; scapus and

basal half of flagellum dark brown, glossy purplish,

then colour of flagellum changes clinally to bronze.

Tegula and thorax (dorsum) bronze, glossy gold.

Forewing (fig. 2) straw-yellow; basal area with silver

stripes (along costal and medial veins) and some four

or five dark brown lines; dorsal margin dark in the

basal one-third of the wing. Apical quarter with 7 to

9 suffuse dark brown radial lines; these lines are much

shorter near costal margin then near ventral one.

Medial band of fascia yellow, bordered on both sides

by wide ochreous bands with diffuse margins; central

zone of these bands with glossy silver-grey zone.

Internal margin of the proximal ochreous band slight-

ly convex, almost indistinct from the ochreous back-

ground. The silver-grey zone of distal band consists of

two parts: the costal one turns towards the apex of the

wing, forming a hook, whereas the dorsal part is

slightly turned towards the wing base. Distal part of

the dorsal hook-like strip in some specimens is isolat-

ed in a form of the elongate spot. Cilia bronze.

Hindwing brown, slightly darker apically; costal area

grey; cilia greyish-brown to brown. Legs yellow ven-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

trally and brown dorsally, except for apical parts of

tibia and tarsomeres which are yellow on both sides.

Epiphysis at 1/2 of fore tibia, not reaching its tip.

Abdomen brown dorsally, yellow ventrally.

Female. Forewing length 8.0 mm; wing expanse

16.5 mm. Antenna 1.5 X forewing, brown; scapus

and basal one-third of flagellum with appressed dark

brown, glossy purplish scales. Otherwise similar to

male.

Male genitalia (figs. 8-12). Tegumen dome-

shaped, with sharp medial ridge. Socii elongate, about

same diameter as medial part of aedeagus. Vinculum

with straight lateral margins, tip almost pointed.

Length of valva c. 0.30 X length of vinculum.

Medial margin of valva ventrally with prominent lobe

appearing laterally as a triangular protuberance; tip of

valva widely rounded. Valva medially with small

(< 1/3 X length of valva, in lateral view) basal

swelling bearing tuft of long setae on small oval de-

pigmented spot. Valvae accreted basally. Arrow-head

of juxta narrow (width about 1/2 X length), tip of ar-

row-head pointed. Distal one-third of aedeagus

membranous, with two slightly curved narrow sclero-

tized bands; aedeagus basally of the same width as

medially.

Biology

For habitat characteristics, see N. aurora. Moths fly

in the canopy and subcanopy layers, 7 - 20 m above

the ground; maximum activity occurred at 9-10 a. m.

and 15-16 p. m. Moths were observed feeding on

flowers of Lithocarpus formosana, L. kawakamii and

Castanopsis longicaudata (S. H. Yen, pers. comm.).

ACKNOWLEDGEMENTS

The financial support of the Smithsonian

Institution, Washington, for a three-week fellowship

in the National Museum of Natural History, is grate-

fully acknowledged. I also would like to thank

Donald Davis and Vichai Malikul for their help dur-

ing my stay in Washington, and Ebbe S. Nielsen for

his constructive criticism. I am very much indebt to

S. H. Yen who loaned specimens from NMNST and

provided biological information on the described

species.

REFERENCES

Davis, D. R., 1975. West Indian moths of the family

Psychidae with descriptions of new taxa and immature

stages. — Smithsonian Contributions to Zoology 188: 1-

66.

Heppner, J. B., 1992. Adelidae, p. 63. In: Heppner, J. B. &

Inoue, H. Lepidoptera of Taiwan, v. 1, pt. 2: Checklist.

Association for Tropical Lepidoptera, Gainesville,

Florida, USA. 276 pp.

Kozlov, M. V., 1993. New species of Cauchas Zeller

(Lepidoptera: Adelidae) from the Altai and Tianshan

Mountains. — Nota lepidopterologica 16: 113-123.

Kozlov, M. V., 1995. A taxonomic revision of the askoldella

species-group of the genus Nemophora Hoffmannsegg

(Lepidoptera, Adelidae). — Entomologica Scandinavica

26: 459-472.

Meyrick, E., 1912. Lepidoptera Heterocera (Tineae). Fam.

Adelidae. — Genera Insectorum 133: 1-12.

Received: 10 September 1995

Accepted: 20 October 1996

Nico NIESER', HERBERT ZETTEL’ & Pinc Ping CHEN!

' Tiel, The Netherlands &° Naturhistorisches Museum Wien, Austria.

TEN NEW SPECIES OF RHAGOVELIA

(HETEROPTERA: VELIIDAE) FROM

SULAWESI (INDONESIA).

Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), VII

Nieser, N., H. Zettel & P. P. Chen, 1997. Ten new species of Rhagovelia (Heteroptera:

Veliidae) from Sulawesi. Notes on Malesian aquatic and semiaquatic bugs (Heteroptera), VII.

— Tijdschrift voor Entomologie 140: 17-41, figs. 1-45 [ıssn 0040-7496]. Published 31 October

1997.

Ten new species of Rhagovelia from the province of Sulawesi are described: À. abra, R. gyrista,

R. krama, R. ochra, R. plychona, R. sterea from the main island of Sulawesi, R. abra from Togian

Island, R. cylindros and R. skoura from Sangihe Island and À. tsouloufi from Salibabu and

Taulaud Islands. Two new species groups, the R. gyrista- and R. plychona-groups are erected,

some species are transferred to other groups and the species groups in the area are redefined.

Some additional records for several species, a check-list and keys to species groups and species

of Rhagovelia for the Sulawesi area are also given.

Correspondence: Dr. Nico Nieser, Htg. Eduardstr. 16, 4001 RG Tiel, The Netherlands.

Key words. — Sulawesi, Sangihe I.; Talaud I.; Togian I.; Veliidae, Rhagovelia, key, new species,

new species groups.

The ten species of Rhagovelia Mayr, 1865, de-

scribed below have partly been collected on islands

belonging to the provinces of Sulawesi Utara and

Sulawesi Tengah. Three species belong to a small col-

lection of aquatic bugs made by the first author in the

Sangihe-Talaud group. The fourth is based on a few

specimens collected on the Togian group which be-

longs to Sulawesi Tengah which were in the NHMw.

In addition there are six new species and additional

records from the main island of Sulawesi collected by

J.P. & M. J. Duffels (ZMAN) and J. van Tol (RMNH).

So all localities are within the administrative entity

Sulawesi of the Republic of Indonesia. As there is at

present great interest in the taxonomy and biogeogra-

phy of Australasian Rhagoveliinae (Polhemus &

Polhemus 1988, Lansbury 1993, Nieser & Chen

1994, Yang & Polhemus 1994, Zettel 1994, 1995) it

was decided to publish the descriptions of these

species in advance of a report on the Sangihe-Talaud

collection. The Sulawesi area seems to be especially

rich in species, in ten years of collecting (1985-1994)

29 species have been discovered.

Measurements are in mm, angles in radians.

Measurements have been based, when sufficient spec-

imens available, on five specimens of each sex and

morph, preferably from the sample containing the

holotype. Measurements refer to the maximum value

and have been taken with the axis along which the

body or body part is measured in a horizontal plane.

The width of a leg segment is its maximum width

measured to the bases of eventual teeth or spines,

length of a leg segment is measured to the tip of even-

tual apical teeth or spurs. The presentation of mea-

surements may vary in the number of digits behind

the decimal point. This is on purpose, the same part

of the body may in one species display more variation

than the next so 0.2 (0.15-0.25) can be found next to

0.20 (0.195-0.205). Distinction is sometimes made

between macropterous (with complete wings) and

dealate (with wings partly or nearly entirely torn off)

as the frequency of the dealate condition differs be-

tween species and species groups. However, dealate

specimens are cited as macropterous in the records of

material.

Locality indications between braces, { }, are not on

the labels but have been added by the authors as ex-

planation.

Specimens have been deposited in the following

collections: 1) registered in Arnett et. al. (1993):

BMNH (London, U. K.), MBBJ (Bogor, Indonesia),

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

rec (Englewood, Co. U.S.A.), MUDH (The Hague,

The Netherlands), NHMw (Vienna, Austria), OXUM

(Oxford, U. K.), RMNH (Leiden, The Netherlands),

semc (Snow Entomological Museum, Lawrence,

Kansas, U.S.A.), uscp (Cebu, Philippines), ZMAN

(Amsterdam, The Netherlands). Not registered:

NCTN (Nieser Collection, Tiel, The Netherlands).

ACKNOWLEDGEMENTS

Thanks are due to Drs. J. P. Duffels (ZMAN), and ].

Margerison Knight (BMNH) to study specimens in

their care and J. van Tol (RMNH) for the permission to

study specimens in his care and help with the distrib-

ution maps. J.T. Polhemus (yrec) for the loan of the

unique holotype of Rhagovelia grayi, J. T. Polhemus

and L. van den Hoek Ostende (RMNH) for criticism

on the first draft of the manuscript.

SYSTEMATIC PART

Species groups in Indo-Pacific Rhagovelia

Polhemus & Polhemus (1988) divided the Indo-

Pacific Rhagovelia species in eight species groups. A

few species are reassigned below to other, partly new-

ly erected, groups due to the increased knowledge of

Indo-Pacific Rhagovelia. As a rule, however, the

‘Polhemi’-groups seem to be monophyletic with the

exception of the large R. papuensis-group. Our con-

cept of the papuensis-group is mainly based on its

Sulawesian and Philippine species which may consti-

tute a restricted section of the group as compared to

the interpretation of this group by Polhemus &

Polhemus (1988). Apparently the largest number of

species of this group is found in Sulawesi (Lansbury

1993, Nieser & Chen 1993, this paper). Nevertheless

we decided against an evaluation of the entire group

mainly for two reasons, this paper was meant to re-

strict its scope to Sulawesi and J.T. Polhemus (JTPc)

informs us that he still has a number of undescribed

species of this group from more eastern regions. It

seems better to wait for a description of these addi-

tional species before attempting a final evaluation of

the papuensis-group.

Five species were found to be sufficiently aberrant

that they are considered to constitute two new species

groups: the gyrista- and plychona-groups (with three

and two species respectively) which, at least superfi-

cially seem to be related to each other. Since the study

by Matsuda (1956) it is known that the knowledge of

macropterous and dealate specimens is very impor-

tant for the delimitation of species groups in the

genus Rhagovelia, so these two new groups are mainly

based on characteristics of the macropterous morph.

The occurrence of the À. sarawakensis-group in

Sulawesi is doubtful. There is only one record, a

macropterous female of the Bornean species À.

samarinda Polhemus & Polhemus (1988).

Keys to the species-groups of Sulawesi

Apterous specimens

1. Pronotum shorter than length of an eye,

mesonotum exposed .... À. sarawakensis-group

— Pronotum much longer than length of an eye

(fig. 21, mesonotum covered by pronotum, at

mosuits;hindmarein visible. nn. 2

DENE Ae 3

MR ITA ESRI EB. NT ENCRES CORP 6

3. Hind femur with teeth or granules in proximal

third (figs. 2, 4)

— Hind femur without teeth or granules in proxi-

mal third (figs. 1, 13)

4. Body dorsally and antennae with dense ap-

pressed golden pubescence, inner side of hind

tibia regularly and sharply indented (fig. 11)

ESE I N R. gyrista-group

— Body dorsally and antennae without conspicu-

ous golden pubescence, inner side of hind tibia

irreeularlyandentedi(fies22, 78). mar.

EE OEREN Mae R. papuensis-group

5. Anterior trochanter with a tuft of hairs, body

dorsally with conspicuous golden pubescence,

hind femur basally yellow .. À plychona-group

— Anterior trochanter without a tuft of hairs, body

dorsally without conspicuous golden pubes-

cence, hind femur basally black in most species

IVA be ES ÉRIC OMAR SA 1, R. orientalis-group

6. Hind femur curved, its anterior face (partly) flat-

tened and densely pubescent, hind femur with-

out teeth or with some small teeth on the poste-

Fiomiacci (ieee 2) ewe. Ma R. gyrista-group

— _ Hind femur without these combination of char-

acters, in most species with teeth on the ventral

margin, only rarely without teeth

7. Dorsum covered with golden pubescence, hind

femur without teeth ventrally (fig. 14)

ATA EE an OR Ot IN AN R. plychona-group

— _ Dorsal pubescence sometimes yellow but with-

out golden metallic shine, hind femur usually

with some teeth or spines

8. Legs slender, with metallic shine, middle and

hind femur basally black (except in A. kas-

tanoparuphe), connexiva black (except in À. kas-

tanoparuphe), abdomen short, its outlines con-

vergent (fig. 21), tergite 7 frequently with a

process (fig. 34), jugae and proepisterna without

blackispicules RP Eee PRE R. orientalis-group

— Legs stouter (figs. 3, 8, 9), without metallic ...

shine, middle and hind femur basally yellow,

connexiva yellow (except: brown in À. pruinosa),

tergite 7 without process, abdomen longer, its

outlines (not that of the connexiva) less conver-

gent (figs. 17, 20), jugae and proepisterna with

or without black spicules .. À. papuensis-group

Macropterous specimens

Macropterous specimens of the À. gyrista-group are

unknown.

1. Hairs on fore wing restricted to costal margin

— Hairs on fore wing covering a large area in the

anterior part of its base ..... R. plychona-group

2. Closed cells of fore wing reaching the apical

third

— Closed cells of fore wing reaching only the apical

half, abdominal carinae reaching tergite ......

legen elles boues R. sarawakensis-group

3. Fore wing brown or black, without a longitudi-

nal yellowish streak basally, abdominal carinae

teachinogtersite > an wer... R. orientalis-group

— Fore wing brown or black, with a longitudinal

white or yellowish streak basally, abdominal

carinae reaching tergite 3 .. À. papuensis-group

(See also characteristics used for identification of

apterous specimens of these two groups.)

Dealate specimens

Dealate specimens of the À. plychona- and the R.

papuensts-group are unknown.

1. Fore wing broken at middle, behind its cells . . .

re adipe pe dans ge R. sarawakensis-group

= Eoreiwing/brokentaubasen "Pret en 2

2. Hind femur in male with a row of small teeth in

proximal third, in female curved, flattened and

anteriorly pubescent ........ R. gyrista-group

— Hind femur in male without teeth in proximal

third, not curved flattened, or pubescent in fe-

male tenet. bike R. orientalis-group

The Rhagovelia papuensis-group

Diagnosis: Variable in size, length 2.6 - 5.3 mm,

generally slender. Colour variable, from black to yel-

lowish, in most species at least connexiva yellowish or

reddish. Legs extensively yellowish in most species.

Pronotum of apterous specimens long, covering the

mesonotum completely or nearly so. Fore wing with

three or four closed cells reaching its apical third. Fore

wing with a longitudinal white or yellowish streak

basally. Dorsal abdominal carinae of winged speci-

mens extending posteriorly to tergite 3. Dealate spec-

imens unknown. Hind femur of males strongly thick-

ened in most species.

NIESER ET AL.: Rhagovelia from Sulawesi

India,

Moluccas,

Taiwan,

New

Distribution: Sri Lanka,

Philippines, Borneo, Sulawesi,

Guinea, Australia, Solomon Islands.

This is the largest species-group in Sulawesi and in

the Philippines, whereas from the other parts of its

distribution area only a few species have been record-

ed.

Key to the species of Sulawesi and adjacent islands

(apterous specimens)

Male of R. unica unknown.

1. Females

= Malest era ehe. area 20

2. Connexiva strongly convergent, folded over ab-

domen and touching or nearly touching each

other posteriorly part, consequently at least part-

ly covering tergite 7 (figs. 18, 19)

— Connexiva, although strongly convergent in

some species, distinctly separated in posterior

part, tergite 7 largely uncovered (figs. 15-17, 20)

3. Hind femur petiolate, outline from its base to

the base of the first tooth concave (fig. 9), outer

rim of connexiva, sometimes indistinctly, nar-

rowly shining on segments 4 and 5

— _ Hind femur not petiolate, outline from its base

to the base of the first tooth convex or, rarely,

straight (fig. 3)

4. Connexiva of segment 6 with many long hairs,

process of connexiva directed upward (figs. 18,

DO) RAA SN EME UE Rp R. tsouloufi sp.n.

— Connexiva of segment 6 without long hairs,

process of connexiva directed more or less hori-

zontally posteriorly

5. Mesopleurae anteriorly rugulose, but without pit

DUNCTUTES AMAR Rn R. minahasa

— Mesopleurae anteriorly with distinct pit punc-

TÜTE RS AE pila Sin 5. )

6. Length 3.5 - 3.8 mm, sternite 7 laterally on hind

margin with a tuft of long black hairs (fig. 27) .

wieren ea os RET Se tirs R. krama sp.n.

— Length 2.8 - 3.0 mm, sternite 7 laterally on hind

margin without long black hairs, but gonocoxa

withlongenhaurser. en er. R. blogiokommena

7. Connexiva on segments 2 and 3 with long, erect

hairs and on segment 7 with a long and slender

process, predominately black species ........

— Connexiva of segments 2 and 3 without long

hairs, process on segment 7 different, predomi-

nately yellowish to brown species

8. Process of connexiva very long and flattened,

middle and hind femora flattened

— Process of connexiva shorter, triangular, middle

13:

14.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

and hind femora not flattened R. grayi

Tergite 8 horizontal, in the same plane as tergite

7 (figs. 25, 26)

Tergite 8 vertical or obliquely directed down-

ward, not in the same plane as tergite 7 (figs. 28,

30)

. Connexiva of segment 4 with a distinct tuft of

[ong Re ER 11

Connexiva of segment 4 without long hairs (fig.

DOI ian VARO ARCS PRI 12

. Hind femur without teeth proximally of the

long spine in the middle, connexiva of segment 7

internally and posterior corners of tergite 8 with

distinct tufts of quite long hairs .. À. tropidata

Hind femur with at least one spine-like smaller

tooth proximal of the long spine in the middle

(fig. 8), connexiva of segment 7 internally and

tergite 8 with some hairs which are not grouped

IU ESA (Gute, WG) 2 9.08 R. achna sp.n.

Connexiva dark, on segment 4 with a swollen

shining area, and with longer hairs internally on

segments 6 and 7, tergite 8 with many long hairs

ineaudalkthirden 2 ma. ote es R. pruinosa

Connexiva yellow, without swollen shining area

on segment 4, with or without longer hairs on

segment 6 and 7, tergite 8 at most with longer

hairs in the posterior corners

Body completely yellow, connexiva without

longer hairs (except serial setae, fig. 25)

RP ee IO R. ochra sp.n.

Body partly dark, connexiva with longer hairs on

segments 4, 6 or 7

Connexiva on segment 4 strongly convergent,

on segment 7 internally with long hairs, metan-

otum with a pair of tubercles laterally, tergites 4

- 5 with a distinct carina along middle

SE FRI DOTARSI ele LR ra Lu eg R. robina

Connexiva more or less straight, gradually con-

vergent from base to tip, on segment 7 internal-

ly without long hairs, metanotum without a pair

of tubercles, tergites without carina . À. kalami

. Connexiva densely set with many long erect

hairs, especially on segments 6 and 7 . R. unica

Connexiva without erect hairs or at most with 1

- 5 bristle-like hairs on each segment ..... 16

Connexiva of segment 7 produced in an elongate

spiniform process (figs. 27, 28, 30), hind femur

slightly petiolate and with a few small denticles

tnsbasalkthindts REA SD E RO 17

Connexiva of segment 7 produced in a short tri-

angular process, hind femur not petiolate and

without small denticles in basal third ....... 19

. Sternite 7 laterally on hind margin with a tuft of

lonsiblackihairs(fiss 02 7, 30) EE 18

Sternite 7 laterally without long black hairs (fig.

DEE ENE I R. abra sp.n.

18.

19}

Sternites black, outer margin of connexiva dull

brownish, little contrasting; tergites 1-3 with a

few scattered bristles (fig. 17), connexiva with 1-

4 bristles per segment (fig. 30) . R. skoura sp.n.

Sternites partly yellow, outer margin of connexi-

va of a well contrasting medium brown to dull

orange; tergites 1-3 without bristles (fig. 19),

connexiva on each side with two bristles on seg-

ment Wond A ir n R. krama sp.n.

Hind femur strongly incrassate, less than 3.5

times as long as wide, laterotergites 3 and 4 with

longer hairs than laterotergite 5 ... R wallacei

Hind femur more slender, more than 4.0 times

as long as wide (without teeth), laterotergites 3 -

5 with equally short hairs ......... R. horaia

Males

20.

23.

24.

Hind femur with a long spine in the middle of

its length, followed by a row of spines of gradu-

ally decreasing length distally, proximally of the

long spine with a single row of short black gran-

ules of which the last may be slightly longer and

tooth-like (figs. 2, 4-6, 10); hind tibia variable .

Hind femur ventrally with two larger spines of

subequal length, one in proximal third and one

distally of the middle, each followed by a row of

spines of decreasing length (fig. 7), proximal

third with a row of short black granules; hind

tibia sinuate and with a larger tooth in distal

third R. achna sp.n.

Sternite 7 with a distinct tuft of hairs medially

ee N N 22

Sternite 7 without a tuft of hairs, but sometimes

with longer hairs on a median carina ..... 23

. Tuft of hairs near to the anterior margin of ster-

nite 7, hairs on tergites short ..... R. pruinosa

Tuft of hairs about halfway on sternite 7, hairs

on tergites longer, slightly shorter than length of

tergite 4 R. kalami

Tergites with numerous erect black hairs which

are distinctly longer than length of tergite 4,

connexiva reddish to dark brown, body length

usually over 2.9 mm

Tergites without hairs or with shorter, more

oblique hairs which are much shorter than

length of tergite 4, only in one small species

(body length up to 2.92 mm) with yellow con-

nexiva the hairs more erect and only slightly

shortentnaniterpite ARRONE IAS SR 26

Hind femur relatively slender, more than 3.5

times longer than wide, proximal row with more

than ten granules, tip of paramere slender

PROLE, MICI te ee eae Sn R. trichota

Hind femur stouter, less than 3.0 times longer

than wide, basal row with less than ten granules,

25:

iif

29.

30.

Ilo

92°

Bi PIONPArAMErE WI CERRI I 25

Proepisterna with small black denticles in anteri-

or part, sides of pronotum black R. skoura sp.n.

Proepisterna without small black denticles, sides

of pronotum yellow ....... R. tsouloufi sp.n.

Hind tibia distinctly curved, with a larger tooth

in the distal third, colour of pronotum com-

pletely yellow, orange brown, or brown, para-

meres apically with a ventral broadening (fig. 40)

Hind tibia straight or only very slightly curved,

without larger tooth, colour of pronotum partly

black, at least in the middle of posterior half,

apex of paramere not ventrally widened (figs. 35,

37)

Proximal row on hind femur with more than 25

black ernest e R. grayi

Proximal row on hind femur with less than 23

Blacks sra nesta Rn 28

Lateral parts of tergites with scattered small

black denticles, extending at least in low num-

bers to laterotergites

Tergites and laterotergites without small black

denticles rer. TE R. lorelinduana

Laterotergites with numerous black denticles,

connexiva of segments 2 - 6 without any longer

NÉS LEE AR EST R. ochra sp.n.

Laterotergites with very few scattered black den-

ticles, connexiva with at least one longer hair in

the posterior corner of each segment AR. horaia

Sternites 5 and 6 medially with tufts of long dark

hairs, black denticles scarce on the lateral parts of

the thorax and lacking or very scarce on lateral

parts of sternites, on laterotergites and tergites .

REI) er no R. wallacei

Sternites 5 and 6 medially only with slightly

longer, yellow hairs, if with a small tuft on stern-

ite 5, then black denticles numerous in these

DÉCORER AND AEN ee ee a

Black denticles restricted to proepisterna, small

species (body length up to 2.92 mm) with rather

long suberect hairs on tergites R. blogiokommena

Black denticles extending to mesopleura, meta-

pleura, sternites, laterotergites and the lateral

parts of tergites, hairs on the tergites shorter 32

Abdomen and metathorax completely yellowish

to orange brown, tergites only with short ap-

pressed hairs, hind femur slender, about 3.5

times as long as wide, paramere with broadened

apex and sternite 7 with a blunt carina .......

GA CE VEREINE COSTOSO R. robina

Abdomen extensively black, tergites with scat-

tered longer, suberect hairs, hind femur stouter,

at most 3.0 times as long as wide, parameres with

simple tip or sternite 7 with a strongly developed

narrow keel

NIESER ET AL.: Rhagovelia from Sulawesi

33. Sternite 7 laterally compressed, with a strongly

developed median carina, which is accentuated

by a well developed fringe of hairs, paramere

withibroadenediapexe "17777777 R. tropidata

— Sternite ventrally flattened, with a low median

carina, paramere with simple tip ......... 34

34. Black denticles all over the anterior margin of

the pronotum, numerous in the median yellow-

ish area, median carina of sternite 7 very faintly

indicate se eye cra Apa ONE R. abra sp. n.

— _ Black denticles lacking in the middle part of the

anterior margin of the pronotum, median carina

in posterior part of sternite 7 low, but distinct,

and accentuated by shallow lateral impressions .

35 Pro-, meso- and metapleura densely set with

black denticles, hind margin of pronotum black

ET ae | 2 R. minahasa

— Pro-, meso- and metapleura set with at most a

few black denticles, hind margin of pronotum

VEllowat SPM cle ee Nets SRI R. krama sp.n.

Rhagovelia blogiokommena Nieser & Chen

Rhagovelia blogiokommena Nieser & Chen 1993: 268, figs.

2-6, 82.

Material. — Sulawesi Selatan, Batusitanduk {2°48'S

120°10'E} N of Palopo, narrow river, 2.X1.1993, leg.

J. P. & M. J. Duffels 18 109 apterous, 26 39

macropterous (ZMAN, 1d 19 macr. NCTN); Onang

{3°07'S 118°49'E}, Sungai Parabaya, 19.X1.1993,

leg. J. P. & M.J. Duffels, 48 72 apt. (ZMAN, 16 19

NCIN, 1d 19 NHMw).

Previously known from three localities in the S half

of Sulawesi all under 100 m asl.

Rhagovelia kalami Nieser & Chen

Rhagovelia kalami Nieser & Chen 1993: 273-274, figs. 24-

POMEZIA

Material. — Sulawesi Selatan, Zuid Celebes, Nang-

galay 2800myeRantepaos VII 19375. les Fa €:

Drescher, 1d apt. (BMNH)

This specimen has a much darker colour than the

type series from Buton Island, however, structurally

we have found no differences. Previously known by

the type series from Buton only.

Rhagovelia pruinosa Polhemus & Polhemus

Rhagovelia pruinosa Polhemus & Polhemus 1988: 195-195,

figs. 149-157, 229.

Material. — Sulawesi Selatan, SW Celebes, 1100m,

Mt. Lompobatang area, Malino {ca 5°15'S

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 1-14. Hind legs or hind femur of apterous specimens of Rhagovelia: 1 R. cylindros 8,2 R ochra 3,3 R. ochra 2,4 R.

skoura 3,5 R abra 3,6 R. krama 3,7 R achna 8,8 R achna 2,9 R tsouloufi 3,10R. tsouloufi Qi Ie gyrista DIR

gyrista © (inset: femur in frontal view), 13 R plychona G,14R plychona OF

119°48'E}, 2, 8-10.VI.1982, M. A. Lieftinck, 18

(RMNH); Sulawesi Selatan, C. Sulawesi, Batusitanduk,

NW of Palopo {ca 120°09'E 2°50'S}, narrow river

2.X1.1993, leg. J. P. & M. J. Duffels, 18 (zman).

Widely distributed in South and Fast Sulawesi and

Buton (Nieser & Chen 1993)

Rhagovelia abra sp. n.

(figs. 5, 15, 28, 35, 45)

Type material. — Holotype apterous d (ZMAN): IN-

DI.

DONESIA: Sulawesi Selatan, E side lake Matana, Sg.

Salura, 450m, 20 Oct 1993, narrow tributary to lake

Matana, 2°32'S, 121°28'E, J. P. & M. J. Duffels (fig.

45). — Paratypes, same data as holotype 35 59 all

apterous (16 39 including allotype ZMAN, 1d 19

NCIN, 16 19 NHMW).

Description. — Apterous form. Dimensions. Length

298-3055 5 1023.23 width MOST

1.18-1.21; width of head 3 0.72-0.74, 2 0.75-0.79;

width of pronotum d 0.90-0.98, 2 0.98-1.03.

Colour generally dark grey-brown to blackish, an-

Table 1

NIESER ET AL.: Rhagovelia from Sulawesi

Measurements of leg segments in Rhagovelia. Tarsal segments 1 and 2 of fore and hind legs have not been measured for

most species. When not given, they measure together 0.05-0.07 and 0.12-0.15 respectively and do not seem to provide

specific characters.

femur tibia tars] tars2 tars3

R. abra 3

fore leg 0.85 0.90 - - 0.22

middle leg 1.40 1.14 0.06 0.42 0.67

hind leg 1254 Wasizeen È 3 0.25

R. abra 9

fore leg 0.85 0.88 - - 0.25

middle leg 1.42 1.09 0.07 0.41 0.68

hind leg 1.19 1.17 - - 0.26

R.achna 3

fore leg 0.90 0.98 - - 0.27

middle leg 1.54 al 0.08 0.41 0.67

hind leg 1.56 1.42 0.07 0.10 0.33

R. achna 2 *

fore leg 0.83 0.91 - = 0.25

middle leg 1.43 1.06 0.07 0.37 0.65

hind leg 1.28 1.22 0.06 0.08 0.30

R. cylindros 3

fore leg 0.93 1.01 = = 0.23

middle leg 1.59 1.18 0.09 0.57 0.71

hind leg 1.34 1.31 0.06 0.09 0.26

R.cylindros 9

fore leg 0.92 0.97 - - 0.22

middle leg 1.52 15112 0.07 0.51 0.73

hind leg 1.23 1.36 0.07 0.10 0.27

R. krama 3

fore leg 0.93 0.98 - - 0.24

middle leg 1.57 1.20 0.08 0.50 0.68

hind leg 1.34 1.24 0.07 0.10 0.30

R. krama 9

fore leg 0.95 1.00 = = 0.23

middle leg 1.54 1.23 0.08 0.50 0.71

hind leg 1.36 1.36 0.06 0.09 0.28

R. ochra &

fore leg 1.08 1.11 = = 0.24

middle leg 1.64 1.42 0.08 0.50 0.72

hind leg ie eS Stokes 012038

terior quarter of pronotum orange-yellow, connected

with yellowish to light brown propleura and proster-

num. Dorsal half (males) to two thirds (females) of

connexivum brownish, not very contrasting. Venter

castaneous, medial area darker, juga, prosternum, ab-

dominal sternite 7 and genital segments of male

lighter. Basal half of first antennal segment, basal part

of rostrum, basal three quarters of fore femur, basal

third of hind femur, acetabula, coxae and trochanters

(continued)

femur tibia tars] tars2 tars3

R. ochra ©

fore leg 0.96 0.96 - - 0.21

middle leg 15% 1.27 0.07 0.44 0.71

hind leg oe 0.10 0.40

R. plychona 3

fore leg 1.21 1.35 - - 0.31

middle leg 2.01 1.64 0.09 0.77 0.86

hind leg 1.53 11070) 0.07 0.18 0.40

R. plychona ©

fore leg 1.28 137 - - 0.33

middle leg 1:95 1.67 0.07 0.79 0.88

hind leg 1.50 1.86 0.08 0.18 0.42

R. skoura &

fore leg 0.90 1.00 = = 0.27

middle leg 1.51 1317 0.08 0.40 0.70

hind leg 1.34 1.23 - 0.10 0.33

R. skoura 9

fore leg 0.91 0.96 - = 0.26

middle leg 1.46 1.10 0.08 0.43 0.72

hind leg 1732 1.30 0.07 0.10 0.31

R. sterea 9 *

fore leg 1.10 112 - = 0.30

middle leg 1872 1.47 0.08 0.60 0.78

hind leg 1.30 1.65 0.07 0.12 0.40

R. tsouloufi 3

fore leg 0.82 0.88 = = 0.23

middle leg 1.35 1.12 0.07 0.40 0.66

hind leg 1.26 1.20 0.06 0.09 0.28

R. tsouloufi 2

fore leg 0.84 0.90 = = 0.23

middle leg 1.36 til 0.08 0.41 0.68

hind leg 1.16 1.22 0.07 0.10 0.24

* based on a single specimen

pale; hind femur in ventral view medium brown.

Paler parts of fore femur variable between specimens,

ventrally more extensive than dorsally.

Minute black denticles spread over propleura and

the orange band on pronotum, on mesosternum

reaching to mesoacetabula but not much on meso-

pleura. Mesopleuron in front of acetabulum with a

group of 6-8 coarse punctures more distinct in fe-

males than in males. Dorsum, especially pronotum

TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 140, 1997

rather bare, punctures on posterior three quarters of

pronotum distinct. Vertex interoculus, sides of thorax

and abdomen, antennae and legs with the usual pu-

bescence and setae.

Length of antennal segments (6 and 9) I: II: II

: IV 0.72 : 0.37 : 0.50 : 0.43. Pronotum long, cover-

ing mesonotum, length : width 0.78 : 0.96. Length of

metanotum on midline, 0.10. Abdominal tergites 1-5

subequal in length (0.15-2.0), tergite 6 slightly longer

(0.30) in female, tergite 7 0.30 in male, 0.38 in fe-

male.

Male. Posterior trochanter with 2-3 small but dis-

tinct blunt teeth. Posterior femur three times as long

as wide (1.25/0.4), ventral margin in proximal third

with about eight minute and closely set teeth fol-

lowed by a row of about eight larger spines decreasing

in size, of which the first (proximal) is half as long as

the width of femur (fig. 5). A row of five smaller

spines located in distal part ventrally of the main row.

Larger spines black tipped, smaller spines and teeth

entirely black or nearly so. Posterior tibia virtually

straight, armed beneath with a row of about 15 rather

coarse pegs. Connexiva more or less horizontal, grad-

ually converging posteriad. Basal width of abdominal

tergite 7 and 8 subequal to their median lengths.

Abdominal sternites 2-6 of subequal length (0.15-

0.20), sternite 7 longer (0.3) with a pair of shallow

impressions separated by a narrow and low keel, ster-

nite 8 with an indistinct ventral keel. Parameres as in

fig. 35.

Female. Posterior trochanter without small teeth.

Posterior femur nearly three and a half times as long

as wide (1.2/0.35), proximal row of minute teeth re-

placed by 4-5 coarser teeth, larger spines except for

the first shorter than in male. Posterior tibia with

somewhat smaller and less distinct pegs as in male.

Connexiva more or less vertical, on tergites 4 and 5

somewhat sinuate, otherwise gradually converging

caudally, caudally leaving most of tergites uncovered

(fig. 15, apices pointed, accentuated by a tuft of cau-

dally directed bristles. Tergite 8 slanting ventrad at an

angle of about 0.257. Proctiger vertical. Sternite 7

about half as long as remaining abdominal sternites

(0.5/1.0). Gonocoxa hidden in segment 7 (fig. 28).

Etymology. — Abros, greek adjective meaning

coarse, referring to the extensive cover of fine denti-

cles.

Comparative notes. — Similar to À. blogiokommena,

R. tropidata, R. wallacei and, in males, R. minahasa.

Of these À. blogiokommena has the black denticles re-

stricted to the posterior part of jugum and the an-

teroventral part of propleura alongside the rostrum.

The others have the minute black denticles distinctly

present on pro- and meso-pleura. À. minahasa lacks

mesothoracic punctures and has the connexiva of the

female folded over the abdomen on segments 5-7, R.

tropidata has a prominent ventral keel on sternite 7 in

males, R. wallacei has characteristic tufts of bristles

abdominal sternites 5 and 6 and the connexiva slant-

ing upward in males. The latter two species have the

minute black denticles not extending onto the orange

trans verse band dorsally on pronotum as in R. abra.

Females of R. tropidata and R. wallacei have the apices

of connexiva more truncate than R. abra, only 0-2

small teeth on proximal ventral margin of hind femur

and, moreover, in R. wallacei females the connexiva

are straight.

Rhagovelia achna sp. n.

(figs. 7, 8, 16, 26, 36, 45)

Type material. — Holotype apterous d (NHMW) IN-

DONESIA: Sulawesi, Togian Inseln, 3 Feb 1995 leg.

Seifert & Greindl (59) (fig. 45). — Paratypes same

data as holotype 364 29 (NHMw, 16 NCTN).

Description. — Apterous form. Medium sized rela-

tively slender and pale species. Dimensions. Length

S 3.10-3.15, 2 3.26; width d 1.15-1.18, 9 1.18;

width of head d 0.76-0.78, £ 0.75; width of prono-

UM ONU O7 221502:

Colour generally brownish. Anterior quarter of

pronotum orange-yellow, connected with equally

pale propleura and prosternum, meso- and meta-

pleura orange-brown. Disk of pronotum medium to

light brown, with an ill-defined lighter median longi-

tudinal stripe. Abdomen dorsally and laterally medi-

um brown, outer half of connexiva orange-yellow.

Venter orange-brown. Antenna grey-brown basal half

to three quarters of first segment pale. Acetabula, cox-

ae, trochanters and most of anterior and posterior fe-

mur sordid pale yellowish. Middle femur except for

base, tibiae and tarsi grey-brown. Apex of anterior fe-

mur and variable dorsal and ventral stripe on hind fe-

mur greyish.

Minute black denticles prominent on thoracic and

abdominal venter to sternite 6, spreading on propleu-

ra and bases of jugae. A line of pits anteriorly and pos-

teriorly on orange-yellow band of pronotum, pitting

of posterior three quarters of pronotum and meso-

pleuron (fig. 26) distinet. Body clothed with short

pubescence with some sparse fine longer hairs, thick-

er and more prominent laterally and ventrally (espe-

cially caudally in males), disc of pronotum rather

bare. Most of abdominal tergite 7 (and genital tergites

of male) free of pubescence, shining. Vertex, inte-

roculus, sides of thorax and abdomen, antennae and

legs with the usual pubescence and setae.

Length of antennal segments (d and 2) I: II: Ill

: IV 0.75 : 0.40 : 0.56 : 0.43. Pronotum long, cover-

ing mesonotum, shorter than wide (0.8/1.0). Length

of metanotum on midline, 0.1. Abdominal tergites 1-

5 subequal in length (0.2), tergite 6 subequal in male

slightly longer (0.25) in female, tergite 7 0.35 in 6,

0.40 in 9. Abdominal sternites 2-6 subequal (0.2)

and 7 longer (0.30) in male. In female sternites 2-5

subequal (0.2), 6 slightly (0.25) and 7 distinctly

(0.45) longer.

Male. Posterior trochanter with about four small

but distinct blunt teeth. Posterior femur strongly in-

crassate, on average two and a half times as long as

wide (1.55/0.62). Ventral margin in proximal quarter

with about twelve very small closely set teeth followed

by an irregular row of about ten larger spines of which

the larger ones are about as long as one fifth the width

of femur (fig. 7) dorsally and a row of about four

smaller spines (of the same size as the apical ones in

the dorsal row) ventrally in distal third. Posterior tib-

ia distinctly curved, armed beneath with a double row

of about 30 small teeth in proximal three quarters,

followed by 4-6 larger teeth of which the first, mark-

ing distal quarter is the most prominent. Spines and

teeth brown tipped, the smaller ones nearly entirely

brown. Connexiva slanting upward about 0.257

gradually converging posteriad leaving tergite 7 ex-

posed. Basal width of abdominal tergite 7 less than its

median length (0.30/0.35). Venter with a low broad

keel, abdominal sternite 7 flattened with median keel

more prominent. Sternite 8 shorter than sternite 9

(0.25/0.30), sternite 8 with an indication of a median

keel basally. Parameres rather long (fig. 36).

Female. Posterior trochanter without small teeth.

Posterior femur about 3.5 times as long as wide

(1.28/0.37), with an irregular row of spines about

seven in apical two thirds only, consisting of one larg-

er spine (about half as long as the width of femur)

preceded by one and followed by five smaller, distally

decreasing, teeth dorsally and a row of about four

smaller spines (of the same size as the apical ones in

the dorsal row) ventrally in distal third (fig. 8).

Posterior tibia with a double row of about 30 small

but distinct teeth in proximal three fourth.

Connexiva more or less vertical, curved inward and

with a somewhat thickened rim accentuated by some-

what stronger developed pilosity on paratergite 4,

caudal part virtually parallel, converging slightly on

last segment only, leaving the tergites uncovered (fig.

16), apices truncate with a few bristles only. Tergite 8

and proctiger horizontal. Gonocoxa distinctly pro-

truding (fig. 26).

Macropterous form unknown.

Etymology. — Achnos, greek adjective meaning sal-

low, referring to the somewhat indefinite general

colour.

Comparative notes (see key). — Separated from all

other Sulawesi species in this group by the armature

NIESER ET AL.: Rhagovelia from Sulawesi

of the hind femur in both sexes. This characteristic

and the shape of the parameres set this species closer

to some Philippine species (R. cotabatoensis Hunger-

ford & Matsuda and related species).

Rhagovelia krama sp. n.

(figs. 6, 19, 27, 37, 45)

Type material. — Holotype apterous 4 (RMNH) IN-

DONESIA: Sulawesi Selatan: C. Sulawesi, 30 km N

Wotu, Sg. Anoa, river through primary forest above

waterfall near bridge Wotu-Tentena, upstream pond-

ed sites, small tributaries, boulder-sand, w=4-8m,

d=0.2-0.8m, 650m (asl), 20°20'30"S, 120°47'45"E,

26 Oct. 1993, leg. J. van Tol (fig. 45). — Paratypes,

same data as holotype 3d 29 apterous, distributed as

follows, 16 19 (allotype) RMNH, 1d 19 ncrn, 1d

NHMW; C. Sulawesi, Sg. Anowah, 41 km N of Wotu

along Trans-Sulawesi Highway, narrow streams in

undisturbed rainforest above waterfall, 650m asl.

24.X.1993, J. P. & M. J. Duffels 16 19 apterous

(ZMAN)

Description. — Apterous form. Medium sized gen-

erally dark species. Dimensions. Length d 3.20-3.38,

550378 width 1kT221,. 255 27177502138;

width of head & 0.77-0.80, 2 0.82-0.83; width of

pronotum d 0.97-1.04, 2 1.11-1.12.

Colour generally dark grey, anterior quarter of

pronotum yellow connected with yellow propleurae

and prosternum. Basal three quarters of antennal seg-

ment 1, acetabula, coxae, trochanters, basal four fifth

of anterior femur, ventral surface of middle and hind

femur, basal quarter of hind femur dorsally, ventral

surface of hind tibia, abdominal sternite 7 and genital

sternites yellow to light brown. Connexiva with an

orange-brown rim about half as wide as connexiva in

male, three quarters as wide in female.

Minute black denticles distinct laterally and latero-

ventrally on thorax and basal half of abdomen, ex-

tending to jugae, propleurae, along lateral part of an-

terior margin of pronotum, acetabulae, connexiva

and the basal abdominal tergites. Posterior three

quarters of pronotum punctured, in addition a few

punctures laterally near anterior margin of pronotum

and at posterior margin of propleurae. Vertex, inte-

roculus, sides of thorax and abdomen, antennae and

legs with the usual pubescence and setae. Body

clothed with sordid yellowish pubescence, dorsally

more distinct in males than in females. Caudal seg-

ments of abdomen with some more distinct pilosity.

Length of antennal segments I : 11: II: IV d 0.76

: 0.40 : 0.51 : 0.45, 2 0.80 : 0.41 : 0.50 : 0.45.

Pronotum long, covering mesonotum, shorter than

wide (4 0.8/1.0, 2 0.9/1.1). Length of metanotum

on middle line, 0.10-0.12. Abdominal tergites 1-4

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 15-24. Dorsal view of abdomen of apterous females of Rhagovelia: 15 R abra, 16 R. achna, 17 R. skoura, 18 R. tsouloufi,

19 R. krama, 20 R. ochra, 21 R. cylindros (body without extremities), 22 R. gyrista, 23 R. sterea, 24 R. plychona.

subequal in length (0.2), tergite 5 subequal, 6 slightly subequal, 0.2 in both sexes; sternite 2 equal (0.2), 6

(0.23) and 7 distinctly (0.4) longer in male. Tergite 5 (0.25) and 7 (0.35) slightly longer in male. Sternites

slightly (0.25), 6 clearly (0.33) and tergite 7 distinct- 2 (0.25) and 6 (0.33) slightly, sternite 7 distinctly

ly (0.6) longer in female. Abdominal sternites 3-5 (0.65) longer in female.

Male. Posterior trochanter with 2-4 small but dis-

tinct blunt teeth. Posterior femur incrassate, on aver-

age three times as long as wide (1.37/0.46). Ventral

margin in proximal half with a somewhat irregular

row of 10-15 very small closely set teeth followed by

a double row of about ten larger spines decreasing in

size distally, of which the first (proximal) is two fifth

as long as the width of femur (fig. 6) dorsally and a

row of 1-6 small spines (of the same size as the apical

ones in the dorsal row) ventrally. Posterior tibia soft-

ly curved to virtually straight, armed beneath with an

irregular double row of about 20 small teeth, distal

teeth very slightly larger and sharper than proximals.

Larger spines brown tipped, smaller ones nearly en-

tirely brown. Connexiva more or less horizontal,

gradually converging posteriad. Basal width of ab-

dominal tergite 7 three quarters its median length

(0.3/0.4). Tergite 8 basally as wide as long (0.35).

Abdominal venter with a, rather indistinct, median

keel on sternites 4-7, accentuated in sternites 4 and 5

by pilosity, on 7 by flattening of the sternite. Sternite

8 laterally compressed basally, suggesting a continua-

tion of the ventral keel. Parameres rather long (fig.

37).

Female. Posterior trochanter without small teeth.

Posterior femur ‘petiolate’ proximally, nearly four

times as long as wide (1.35/0.35), a few small teeth in

proximal part only, row of spines in apical half con-

sisting of one large spine (over half as long as the

width of femur) followed by about 8 much smaller,

distally decreasing, teeth. Posterior tibia with about

20 small but distinct teeth. Connexiva more or less

horizontal on segments 1 and 2, on segment 3 and 4

torsion to vertical, on tergites 5 and 6 curved back

outward and converging caudally, the prolonged

apices nearly meeting over tergite 8 which is horizon-

tal (fig. 19). Gonocoxa distinctly protruding, with a

very shallow lateral impression (fig. 27).

Etymology. — Krama, Greek noun meaning mix-

ture referring to this species combining various char-

acteristics of related species.

Comparative notes. — This species belongs to a sub-

group within the papuensis group consisting of rela-

tively squat species with orangeish anterior band of

pronotum connected with similarly coloured pro-

pleura, in males rather strongly inflated hind femur

and virtually straight tibia without larger subapical

tooth. In Sulawesi this set is represented by R. blo-

giokommena, R. tropidata and R. wallacei, the females

of these three have tergite 8 more exposed, males can

be separated by the characteristics mentioned in the

key.

NIESER ET AL.: Rhagovelia from Sulawesi

Rhagovelia ochra sp. n.

(figs. 2, 3, 20, 25, 40, 45)

Type material. — Holotype apterous d (RMNH) IN-

DONESIA: C. Sulawesi, 30 km N Wotu, Sg. Anoa, riv-

er through primary forest above waterfall near bridge

Wotu-Tentena, upstream ponded sites, small tribu-

taries, boulder-sand, w=4-8m, d=0.2-0.8m, 650m

(asl), 20°20'30"S, 120°47'45"E, 26 Oct 1993, leg. ].

van Tol (fig. 45). — Paratypes, same data as holotype

66 29 apterous, 2d macropterous distributed as fol-

lows: 48 29 (1 apt. allotype, 1 macr.) RMNH, 19 1d

apt. 16 macr. NcTN, 1d NHMw.

Description. — Apterous form. Medium sized slen-

der and pale species. Dimensions. Length d 3.55-

3.58, 2 3.52-3.55; width & 1.08-1.15, £ 1.16-1.19;

width of head d 0.74-0.80, 2 0.76-0.79; width of

pronotum d 0.98-1.00, 2 1.03-1.04.

Colour generally pale, yellow and light orange.

Dorsal side mainly orange, eyes dark grey to brown,

interoculus somewhat darker than rest of dorsal side,

apex of rostrum brown. Anterior quarter of prono-

tum pale yellow, connected with equally pale pro-

pleura and prosternum. Lateral third to half of con-

nexivum yellow. Venter mostly orange. First antennal

segment yellowish, segments 2-4 brownish. Legs

mostly yellowish, tarsi and apices of femurs and tibi-

ae infuscated in most specimens.

Minute black denticles spread over body surface

except anterior and dorsal parts of head and the pitted

posterior three quarters of pronotum. Punctures on

posterior three quarters of pronotum distinct. Vertex,

interoculus, sides of thorax and abdomen, antennae

and legs with the usual pubescence and setae. Body at

first sight looking rather bare as its pubescence is

short, appressed and in the same colour as body.

Caudal segments of abdomen with some more dis-

tinct pilosity.

Length of antennal segments I : II: IM : IV d 0.81

: 0.49 : 0.69 : 0.54, ® 0.80 : 0.45 : 0.68 : 0.52.

Pronotum long, covering mesonotum, shorter than

wide (0.8/1.0). Length of metanotum on middle line,

0.12. Abdominal tergites 1-5 subequal in length

(0.2), tergite 6 slightly longer 0.25 in male 0.30 in fe-

male, tergite 7 0.40 in both sexes. Abdominal stern-

ites 3-5 subequal 0.15-0.20 in male, 0.20 in female;

sternites 2 and 6 slightly (0.22) sternite 7 distinctly

(0.40) longer in male. In female abdominal sternites 2

and 6 0.30 and sternite 7 0.50.

Male. Posterior trochanter with 4-6 small but dis-

tinct blunt teeth. Posterior femur variably incrassate,

on average two and a half times as long as wide

(1.68/0.69). Ventral margin in proximal half with

about 20 very small closely set teeth followed by a

double row of about 10 larger spines decreasing in

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

size distally, of which the first (proximal) is one third

as long as the width of femur (fig. 2) dorsally and a

row of 4-7 small spines (of the same size as the apical

ones in the dorsal row) ventrally. Posterior tibia soft-

ly distinctly curved, armed beneath with a double row

of about 40 (25 in inner, 15 in outer row) small teeth,

most specimens with a somewhat larger tooth at api-

cal third. Both curvature and development of larger

tooth on tibia correlated with thickness of femur.

Spines and teeth brown tipped, those on tibia nearly

entirely brown. Connexiva more or less horizontal,

gradually converging posteriad. Basal width of ab-

dominal tergite 7 less than its median length

(0.35/0.40). Abdominal sternite 7 flattened medially,

flat area diverging posteriorly in most specimens,

with an indication of a broad, low median keel, later-

al margins of flattened area accentuated in posterior

half by tufts of bristles. Parameres rather long (fig.

40).

Female. Posterior trochanter without small teeth.

Posterior femur slightly over five times as long as wide

(1.39/0.27), with a single row of spines in apical half

only, consisting of one large spine (about 3/4 as long

as the width of femur) followed by 4-6 much smaller,

distally decreasing, teeth (fig. 3). Posterior tibia with

a single row of about 12 small but distinct teeth.

Connexiva more or less vertical, on tergites 4 and 5

curved inward and with a somewhat thickened rim,

otherwise gradually converging caudally, leaving most

of tergites uncovered (fig. 20), apices only slightly

pointed (fig. 25), more or less triangular. Tergite 8

and proctiger about horizontal in allotype, slanting

ventrad at an angle of over 0.257 in the other female.

Gonocoxa distinctly protruding, with a shallow later-

al impression in apical half.

Macropterous specimens (26 only). — Essentially

as apterous form except for the development of tho-

rax, wings and some details mentioned below. Length

3.70-3.75, humeral width of pronotum 1.40-1.41,

median length of pronotum subequal to humeral

width 1.33-1.40. General colour slightly darker than

in apterous specimens, hemielytra dark, smoky

brown-grey, proximal two thirds of lateral (anterior)

proximal cell sordid white. Hemielytra reaching

halfway to just over proctiger, with four cells two

proximal elongate, reaching nearly halfway

hemielytron. Outer (anterior) apical cell narrow

reaching distal three quarters of hemielytron, central

cell broad reaching distal two thirds. Hind wings dark

smoky grey. Carinae on basal part of abdomen reach-

ing to caudal margin of tergite 3. Connexiva horizon-

tal, virtually parallel in the basal two thirds, curved

medially in caudal two thirds

Etymology. — Ochros, greek adjective meaning

pale, referring to the general colour.

Comparative notes. — Its pale orange colour sets

this species apart from anything seen in Sulawesi and

the Philippines. Other comparatively light reddish

species such as R. robina Nieser & Chen or R. horaia

Nieser & Chen are, nevertheless, much darker and

have the third antennal segment relatively shorter. R.

ochra belongs to a subgroup of the R. papuensis-group

characterized by a strongly inflated hind femur and a

curved hind tibia with one or more larger subapical

teeth in males and a relatively long third antennal seg-

ment (Nieser & Chen 1993). However, the third an-

tennal segment in R. ochra is much longer than in the

above mentioned species.

Rhagovelia skoura sp. n.

(figs. 4, 17, 30, 38, 44)

Type material. — Holotype apterous d (RMNH): IN-

DONESIA: Sulawesi Utara, P. Sangihe, Bowokulu,

19.X1.1994, leg. N. Nieser, N9477 (fig. 44). Upper

reaches of mountain stream, not much water, boul-

ders, narrow stretches with strong current alternating

with quiet pools. Rhagovelia on quiet stretch up and

down stream of bridge. — Paratypes (NCTN unless oth-

erwise stated): Same data as holotype 1806 1062

apt, 2d macropterous, distributed as follows: 24

3@ (including allotype) RMNH; JTPC, NHMW, ZMAN

each 28 29 apt.; MBBJ, MUDH, SEMC each 1d 19

apt. — Additional paratypes (adults only), all Pulau

Sangihe and leg. N. Nieser: Naha, Sungai Laine,

27.V1.1994, N9454. Lower reaches of stream, about

10m wide, mostly about 0.5 m deep, water slightly

turbid (after a night of rain), bottom, coarse sand and

pebbles. Rhagovelia at the edges between vegetation,

126 109 apt. (26 29 uscp); Naha, Sungai Laine, at

last bridge upstream, 27.V1.1994, N9456. Slowly

flowing river partly filled with Hydrilla, water faintly

bluish (probably soap from washing cloth), 38 39

apt; Akembawu, ford & bridge in Sungai Dali,

16.X1.1994, N9470. Open terrain with some xero-

phytic plants, stones & boulders (the ford is an outlet

in case of banjir). Sample from mouth at sea to 1.5

km inland, 276 379 apt., 26 12 macr., 1 lvV; Desa

Simuang (near Malahu), Sungai Simuang,

28.VI.1994 (for details see À. cylindros), 248 279

apt., 1® macr.; Desa Laine, Sungai Laine, pothole at

water fall, 12.XI.1994, N9463 (for details of habitat

see under R. cylindros). Rhagovelia at open to moder-

ately open spots, 48d 569 apt., 34d 302 macr., 5

lvv. (JTPC, MBBJ, NHMW, OXUM, SEMC, ZMAN each 1d

19 macr.); Lelepu, Sura (=Sungai) Lelepu,

13.X1.1994, leg. N. Nieser, N9464. Mainly lowland

stream aspect, some Rhagovelia also from stretch with

mountain stream aspect, 594 539 apt., 7d 82

macr., 40 lvv; Desa Utaurano, Sungai Apanu-

kang, 14.X1.1994, N9465, (for details see R. cylindros)

196 429 apt., 126 149 macr.; same, pothole,

N9465A 86 7% apt, 26 1 ? macr.; Sungai

Masalihe, 16.XI.1994, N9468, mountain stream in

rather deep (c. 20m) gorge. Main stream rather

rough, mainly boulders and rapids, some quieter bays

with pebbles and sand. Most Rhagovelia from

N9468A, narrow parallel stream with sand bottom,

mostly c. 2-5m wide, 546 459 apt., 5d 89 macr.,

5 lvv.; Sungai Kendahe, bridge at Poto, mountain

stream, 17.XI.1994, N9471. Medium sized, fast

flowing stream, bottom coarse pebbles, width 3m, at

bridge much wider, and slower (spot extensively used

for washing cloth), specimens taken from quiet bays

at edges of stream, 168 109 apt, 11d 79% macr.; N.

side, near Poto, stream, 17.X1.1994, N9472. Small

mountain stream through overgrown coconut grove,

some boulders, bottom pebbles, some plant debris,

water clear, hyaline, 126 129 apt. (38 3% oxum,

26 22 uscr); N. side, near Poto, stream,

17.XI.1994, N9473. Mountain stream, narrower and

quieter than N9471, some boulders, bottom small

pebbles, coarse sand, 154 89 apt., 2d 19 macr., 6

lvv.; Small stream at Kampung Lapango-Hakadele

(near Desa Sawaeng), 18.X1.1994, N9474. Mountain

stream, sample from slowly flowing rather flat part.

Bottom with some mud. /pomoea growing into the

water which has a bluish tinge (soap ?), 76d 929

apt., 138 172 macr., 2 lvv.; Sungai Makariahe, near

Gunung, 19.XI.1994, N9478. Upper reaches of

mountain stream, more or less a trickle of water be-

tween boulders, some small pools from which the

sample was taken, 1d 19 apt, 1% macr.; E. side,

Sungai Miulu, 20.XI.1994, N9480, 7d 99 apt., 19

macr., 2 lvv.

Description. — Apterous form. A medium sized

rather broad and generally dark species. Dimensions.

Length d 3.20-3.28, 9 3.38-3.50; width (at base of

abdomen) & 1.23-1.28, 2 1.35-1.38; width of head

3 0.80-0.82, 2 0.82-0.85; width of pronotum d

LEE ARE al

Colour generally dark grey to black, juga and basal

part of rostrum lighter, narrow anterior transverse

band anteriorly on pronotum (reaching laterally

halfway eyes) orange-yellow, covered with grey pru-

inosity which becomes more distinct laterally. Outer

rim of connexiva brownish, not very contrasting, in

most specimens best visible on segments 3-6. Basal

half of first antennal segment and fore femur, basal

third ventrally, basal sixth dorsally of hind femur, ac-

etabula, coxae and trochanters pale.

Minute black denticles spreading from prosternum

to propleura reaching just behind eyes, moreover

some on jugae and acetabula especially dorsally on

metacetabulum. Meso- and metapleuron in an-

terodorsally of acetabulum with distinct punctures

NIESER ET AL.: Rhagovelia from Sulawesi

(fig. 30). Dorsum, apart from the normal fine pubes-

cence clothed with long erect bristles, sparse or absent

on disk of pronotum and, especially in female, on

caudal abdominal segments. Punctures on posterior

three quarters of pronotum indistinct (with ill-de-

fined margins and covered by relatively dense pubes-

cence). Vertex, interoculus, sides of thorax and ab-

domen, antennae and legs with the usual pubescence

and setae.

Length of antennal segments (male and female) I:

II: III: IV 0.81 : 0.40 : 0.49 : 0.50. Pronotum long,

covering mesonotum, length : width 0.90 : 1.13.

Length of metanotum on middle line, 0.10.

Abdominal tergites 1-5 subequal in length (0.20), ter-

gite 6 slightly longer, 0.25 in male to 0.30 in female,

tergite 7 0.40.

Male. Posterior trochanter with 2-5 small but dis-

tinct blunt teeth. Posterior femur 2.7 times as long as

wide (1.34/0.50), Ventral margin in proximal third

with about 10-15 minute and closely set teeth fol-

lowed by a row of 9-11 larger spines decreasing in

size, of which the first (proximal) is nearly half as long

as the width of femur (fig. 4). In addition two or three

smaller spines located in distal part ventrally of the

main row. Larger spines black tipped, smaller spines

and teeth entirely black or nearly so. Posterior tibia

virtually straight, armed beneath with a double row of

about 30 rather coarse pegs. Connexiva usually hori-

zontal, gradually converging posteriad. Basal width of

abdominal tergite 7 less than its median length

(0.30/0.40); (exposed part of) tergite 8 as wide as long

(0.30) Abdominal sternites 2-6 of subequal length

(0.2), sternite 7 longer (0.35) ventrally flattened with

a low keel, sternites 5-6 with median tufts of bristles.

Parameres as in fig. 38.

Female. Posterior trochanter without small teeth.

Posterior femur three and a quarter times as long as

wide (1.32/0.4), proximal row of minute teeth or re-

placed by up to three coarser teeth, larger spines

about eight in number and except for the first shorter

than in male, two to three more ventral spines pre-

sent. Pegs on posterior tibia nearly covered by the tib-

ial pilosity. Connexiva more or less horizontal to

slanting upward to 0.157. Converging caudally in a

nearly straight line (fig. 17), leaving most of tergites

uncovered, apices pointed, accentuated by a tuft of

caudally directed bristles in addition a smaller tuft of

bristles more ventrally on hind margin of segment 7

(fig. 30). Position of tergite 8 and proctiger slanting

nearly vertically ventrad. Sternite 7 about twice as

long as sternite 6 (0.30/0.60) median length of re-

maining abdominal sternites about 0.2. Gonocoxa

hidden in segment 7.

Macropterous form. — Essentially as apterous form

except for development of hemielytra and thorax.

Length 3.3-3.4, humeral width of pronotum 1.38-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

0,5 mm

Figs. 25-34. Lateral view of abdomen (27-28 caudal segments only) of apterous females of Rhagovelia: 25 R. ochra, 26 R achna,

27 R. krama, 28 R. abra, 29 R. tsouloufi, 30 R. skoura, 31 R. sterea, 32 R gyrista, 33 R. plychona, 34 R. cylindros.

1.42, median length of pronotum 1.20-1.30.

Pronotum black with a sharply contrasting orange-

yellow transverse band at anterior margin, sometimes

covered with whitish pubescence. Hemielytra reach-

ing 0.2 beyond apex of abdomen, dark brown with

black veins. A pair of elongate proximal cells marked

with sordid white bands, reaching slightly beyond

halfway hemielytron, a pair of shorter central cells,

outer one (near costal margin) reaching to one fourth

from apex, inner one to one third from apex of

hemielytron. Keels on base of abdomen strongly de-

veloped, reaching to caudal margin of tergite 3.

Brownish rim of connexiva (partly not covered by

hemielytra) more strongly developed than in apterous

form, a shining brownish patch on tergite 7.

Etymology. — Skouros, greek adjective meaning

dark, referring to the generally dark colour compared

to its nearest relatives.

Comparative notes. — See key, general shape simi-

lar to À. wallacei, including the tufts of bristles on

sternites 5 and 6 of male, but this species has the yel-

lowish transverse band of pronotum broad, reaching

to propleurae and the long erect bristle-like hairs dor-

sally restricted to connexiva.

Rhagovelia tsouloufi sp. n.

(figs. 9, 19, 18, 29, 39, 44)

Type material. — Holotype apterous d (RMNH) IN-

DONESIA: Sulawesi Utara, Pulau Salibabu, Sg. Musi,

downstream stretch in agricultural area, quiet, shal-

low, mostly 2-3m wide and 0.05-0.1m deep, bottom

pebbles, stones and a few boulders, water clear, hya-

line, N9489, 26 Nov. 1994, leg. N. Nieser (fig. 44).

— Paratypes (NCTN unless otherwise indicated): Same

data as holotype 358 139 apterous, 74 59

macropterous distributed as follows: 38 29 apt., 1d

19 macr. NHMw; 29 (including allotype) apt., 1d

macr. RMNH. — Additional paratypes, all leg. N.

Nieser: P. Salibabu, Lirung, narrow stream on rocks,

very steep, draining small marsh on top of hill (c. 350

m asl), various shallow pools with much plant debris

connected by trickles of water, shaded by remnants of

(? primary) forest, N9483, 24. XI. 1994, 6d 49 apt.,

16 macr. (ld macr. yrec, 36 32 oxum, 2d 19

uscr); Pulau Karakelong, D(esa) Ambela, at inlet of

irrigation system, 30.X1.1995, N9493. Sungai

Ambela, broad river bed with sand & pebbles, with

little water. One side still forested, other side dis-

turbed, 138 49 apt, 16 macr. (jrpc 36 19, MBBJ

26 19 apt. 1d macr., MuDH 16 19, semc 2d,

ZMAN 2d 19. P. Karakelong, Sungai Ambela at wa-

terfall, 2.xii.1994, pool slightly downstream in low-

land rain forest, N9497A, 456 26% apt. P.

Karakelong, Sungai (at) Sawang, 3.xii.1994, N9498.

Lowland stream trough agricultural (small scale) area,

light brown very turbid (after rain), 28 49 apt., 2d

22 macr., 1 lvv.

Description. — Apterous form. Rather small gener-

ally dark species. Dimensions. Length d 2.89-3.08,

2 3.28-3.48; width (across base of abdomen) &

1.08-1.12, £ 1.15-1.23; width of head & 0.72-0.75,

2 0.75-0.80; width of pronotum d 0.93-0.98, 2

1.01-1.10.

Colour generally dull dark brown to grey, eyes dark

castaneous, yellowish transverse band anteriorly on

pronotum distinctly contrasting, narrow, reaching

slightly beyond the inner margins of eyes. Basal half

of antennal segment 1, acetabula, coxae, trochanters,

proximal three quarters of anterior femur and proxi-

NIESER ET AL.: Rhagovelia from Sulawesi

mal one third of hind femur pale. Connexiva with a

narrow orange-brown rim. Venter and sides and lat-

eral parts of first tergites in female somewhat pruinose

grey. Minute black denticles present on anterolateral

angles of pronotum, due to grey back ground rather

indistinct. Row along anterior margin and posterior

three quarters of pronotum distinctly punctured, pro-

and meso-pleura with well-developed coarse punc-

tures. Vertex, interoculus, sides of thorax and ab-

domen, antennae and legs with the usual pubescence

and setae. Body clothed with short pubescence, virtu-

ally absent laterally in females, superimposed by

sparse longer bristles in males which are hardly devel-

oped in females.

Length of antennal segments (4, ©) I: II: II:

IV 0.72 : 0.39 : 0.49 : 0.42. Pronotum long, cover-

ing mesonotum, shorter than wide (4 0.80/0.96, 9

0.85/1.05). Length of metanotum on midline, 0.1.

Abdominal tergites 1-6 in male, 1-5 in female sube-

qual in length (0.15-0.20), tergite 7 in male 0.40 ter-

gite 6 in female 0.35, tergite 7 0.50. Abdominal ster-

nites 2-5 in male, 3-5 in female subequal, about 0.15

long. In male sternite 6 slightly longer 0.20, sternite 7

distinctly longer, 0.35; in female sternite 2 0.20, 6

0.30 and 7 0.6.

Male. Posterior trochanter with 1-4 small black

teeth. Posterior femur incrassate, on average slightly

over two and a half times as long as wide (1.26/0.48).

Ventral (inner) margin with 8-10 very small black

teeth in proximal third, followed distally by a double

row of spines, the posterior (dorsal) row consisting of

about 12 spines of apically decreasing spines, the larg-

er spine about one third as long as the width of femur.

The anterior (ventral) row consisting of about six

smaller spines (fig. 10). Posterior tibia virtually

straight, armed beneath with a, proximally double

distally single row of 24-30 small teeth which are

partly covered by the pilosity of the femur.

Connexiva more or less horizontal to slanting slight-

ly upward (about 0.157), gradually converging poste-

riad. Abdominal tergite 7 bare, shining, its basal

width three quarters its median length (0.30/0.40).

Tergite 8 basally as wide as long (0.30). Abdominal

sternite 7 brownish flattened, with a faint indication

of a median keel only. Sternites 8 and 9 medium

brown, sternite 8 flattened without keel. Parameres

stout (fig. 39).

Female. Middle femur distinctly and broadly

dorsoventrally compressed halfway its length.

Posterior trochanter without small teeth. Posterior fe-

mur with a narrow part in proximal third (‘petiolate’)

remainder less incrassate than in male, slightly over

three times as long as wide (1.17/0.37); its armament

restricted to a single row of about seven distally de-

creasing spines in distal two thirds, of which the

largest is about half as long as the width of femur (fig.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

9). Posterior tibia armed with an irregular inner row

of about 25 small spines largely hidden in the pilosity

of the tibia. Connexiva more or less vertical, curving

inward over segments 4-5, strongly converging on

segments 6-7, meeting over abdomen on posterior

half of segment 7 (fig. 18) including the upwards

pointing caudal connexival angles which are accentu-

ated by some bristles (fig. 29). The dorsal part of the

caudal margin of segment 7 accentuated by a dense

fringe of large bristles. Tergites 8 and 9 vertical, only

visible in caudal view. A tuft of erect somewhat cau-

dally pointed golden brown bristles on laterosternite

4. Sternites 5-7 flattened.

Macropterous form. — Essentially as apterous form

except for development of wings and thorax. Length

3 3.20-3.38, 2 3.48-3.55, humeral width of prono-

tum d 1.30-1.34, 9 1.37-1.42, median length of

pronotum d 1.20-1.30, 2 1.26-1.40. Pronotum

black, orange-yellow transverse band at anterior mar-

gin reaching slightly beyond inner margins of eyes,

rather indistinct due to cover by whitish pubescence.

Propleura with a distinct row of punctures near ven-

tro-posterior margin. Hemielytra reaching apex of ab-

domen in males and about 0.1 beyond apex of ab-

domen in females, brown to dark grey with darker

veins. A pair of elongate proximal cells marked with

sordid white (especially the one at costal margin)

reaching halfway hemielytron, a pair of distal cells of

which the one near costal margin is distinctly smaller

reaching two thirds the length of hemielytra. Dorsum

of abdomen castaneous, brownish rim of connexiva

broader than in apterous form, keels on base of ab-

domen well developed and long, reaching to the base

of tergite 4. Connexiva of female sinuate, slanting up-

ward nearly 0.47.

Etymology. — Tsouloufi (tsoulouphi), Greek noun

meaning tress (of hair), referring to various tufts of

bristles on abdomen in female.

Comparative notes. — Apparently related to R. tri-

chota Nieser & Chen and R. minahasa Polhemus &

Polhemus females of which have the caudal part of

connexiva more or less folded over abdomen, the

characteristics in the key will serve to separate these

species.

The À. gyrista-group

Diagnosis: Medium sized to rather large Rhagovelia,

length 3.6-4.1 mm, ground colour dark grey to black,

body dorsally and antennae with conspicuous golden

pubescence. Posterior femur in males slender, proxi-

mal granulate teeth rather large. Posterior femur in fe-

males dorsoventrally flattened. Dealate specimens rel-

atively common, basal abdominal carinae (only

observed in À. gyrista) in macropters broad and low,

reaching to posterior margin of tergite 2.

Distribution: Endemic to Sulawesi.

R. hamjadi Polhemus & Polhemus (1988) was

originally placed in the R. papuensis-group. In view of

the above mentioned characteristics which it shares

with À gyrista we consider these two to form a sepa-

rate group within the genus. R. sterea sp. n. so far only

known by the unique apterous female holotype is in-

cluded in this group because of its similarity to À.

hamjadı.

Key to species of the R. gyrista group

(apterous specimens)

l. Orange-yellow transverse band anteriorly on

pronotum wide, connected with the similarly

coloured pleuract A nor. nen. R. sterea sp. n

— Orange-yellow transverse band anteriorly on

pronotum narrow, reaching to inner margins of

eyes, propleurae dark

2. Male, posterior femur proximally with a row of

about 16 granulate teeth, distally with a row of

six spines. Female, abdomen strongly curved up-

wards caudally (fig. 32), connexiva bent inward

and folded over lateral parts of abdominal ter . .

gites 2-01 22) EINE: R. gyrista

— Male, posterior femur proximally with a row of

about 20 granulate teeth, distally with a row of

8-9 spines. Female, abdomen straight, connexiva

not folded over abdomen, their margin only

slightlyiconcave mann RARE R. hamjadi

Rhagovelia gyrista sp. n.

(figs. 11, 12, 22, 32, 41, 45)

Type material. — Holotype, apterous 9 (ZMAN),

INDONESIA: Sulawesi Selatan: SW Sulawesi, Mamasa,

Sg. Loko, 1400m, 17 Nov 1993, leg. J. P. & M. J.

Duffels (fig. 45). — Paratypes, same data as holotype,

14 (allotype) 29 apt., 2d 19 macropterous ZMAN

(1d macr., 19 apt. NctN). The macropters have the

wings entirely broken off except for some remnants at

the humeral angles of pronotum (dealate); Sulawesi

Selatan: Zuid Celebes, Nanggala, 800m, Rantepao,

VII. 1937, leg. F. C. Drescher, 16 apt. (BMNH)

Description. — Apterous specimens. Dimensions

length d 3.93, 2 4.07-4.11; width (across base of

connexiva) 6 1.30, 2 1.45-1.51; width of head 6

0.90, 2 0.90-0.91; width of pronotum d 1.12, 2

1.29-1.30. Colour generally dark grey to blackish.

Orange-yellow transverse band anteriorly on prono-

tum narrow, reaching the inner margins of eyes. Basal

third of first antennal segment, acetabula, coxae and

trochanters, basal half to two thirds of anterior femur

and basis of hind femur, yellow. Outer rim of con-

nexiva narrowly brown in females, not obvious in

male. Venter dull dark grey to black, sternites 7-10

brownish. Minute black denticles restricted to jugum

and inner faces of fore and middle acetabula.

Interoculus, thoracic and abdominal tergites clothed

with dense appressed golden-yellow pilosity much

more developed and distinct in male than in fe-

males.In addition dorsum of thorax and abdomen

with sparse long erect dark cilia. Vertex sides of tho-

rax, antennae and legs with the usual pubescence and

setae.

Length of antennal segments (no apparent differ-

ences between male and female) I: II: II]: IV 0.97:

0.45 : 0.74 : 0.62. Pronotum long, covering mesono-

tum. Disk of pronotum with some indistinct punc-

tures, obscured by pilosity, in posterior part. Length:

width of pronotum about 1.2 (6 1.28 / 1.05, 1.50

/ 1.17). Length of metanotum on midline, 0.10.

Abdominal tergites 1-6 in male subequal in length

(0.20), in female becoming gradually longer posteri-

orly, (from 0.22-0.30) but mostly covered by connex-

iva. Tergite 7 over twice as long as preceding tergites

in male (0.45); 1.5 times as long as preceding in fe-

male (0.45/0.30).

Male. Posterior trochanter length: width 0.4 : 0.2,

without teeth or warts. Posterior femur slender, five

times as long as wide (1.5/0.3), on ventroposterior

margin about halfway with a rather small spine, its

length about one sixth the width of femur, with a row

of about seven spines of gradually decreasing length

distally (fig. 11). Posterior tibia straight, armed be-

neath with a row of about 20 short stout teeth giving

a serrate impression (fig. 11). Connexiva slanting up-

ward about 0.257 or somewhat less, virtually parallel

on segments 1-5 strongly converging posteriad on

segments 6-7, no caudal points. Basal width of ab-

dominal tergite 7 less than 1.5 times its median

length (0.55/0.45). Abdominal sternite 7 compara-

tively short, about 1.2 times as long as sternite 6

(0.30/0.25), sternite 6 somewhat, sternite 7 distinctly

flattened medially, without a carina. Genital seg-

ments prominent (fig. 7), fusiform, tergite 8 longer

tan tergite 7 (0.55/0.45). Sternite 8 laterally com-

pressed at base, resulting in a low broad carina ven-

trally. Median length of pygophore equal to median

length of sternite 8 (0.30). Parameres as in fig. 41.

Female. Posterior femur six times as long as wide

(1.5/0.25). Posterior femur and tibia without teeth or

spines (fig. 12). Connexiva strongly curved inwards,

folded over abdomen leaving on tergites 3-6 only a

narrow median strip uncovered (fig. 22). Caudal apex

of connexiva truncate with distinct tufts of pilosity;

gonocoxae | (sternite 8) laterally with well-developed

pilosity which anterolaterally forms additional tufts

(fig. 22). Tergite 8 nearly horizontal, as long as ter-

gite 7 (0.45) (fig. 32). Sternite 7 large, about two fifth

NIESER ET AL.: Rhagovelia from Sulawesi

the length of the preceding abdominal sternites to-

gether (0.6/1.5). Gonocoxa clearly visible, about half

the length of sternite 7, ventrally compressed, the

sides reaching further ventrally than the flattened to

somewhat impressed ventral part, forming a pair of

lateral ridges. Proctiger large.

Macropterous form. Mostly as apterous except for

modifications of the thorax and presence of wings.

Dimensions, length d 4.80-5.08, 9 4.92-5.18,

hemielytra reaching 0.55-0.67 beyond the apex of ab-

domen; humeral width of pronotum d 1.75-1.80, 2

1.85-1.90, median length of pronotum d 1.71-1.78,

? 1.75-1.85. Hemielytra dull dark grey to blackish,

veins only slightly darker, forming two elongate basal

and two more or less squared apical cells. Hind wings

dark smoky brown-grey. Laterobasal quarter of

hemielytra with golden-yellow pubescence, golden-

yellow pubescence on abdominal dorsum restricted to

a median band. Pronotum with obtuse humeral an-

gles and a rounded caudal apex. Metanotum with a

pair of sublateral pits. Second abdominal tergite with

a transverse pitted groove, longitudinal carinae not

developed. Caudal tufts of bristles on female ab-

domen less prominent than in apterous form.

Etymology. — Gyristos, a Greek adjective meaning

curved refers to the general body form in females.

Comparative notes (see keys). — In view of its size

combined with striking golden pubescence dorsally

the Sulawesi species which look similar belong to the

R. gyrista and plychona groups. The curved abdomen

of the female is at once characteristic (actually the

characteristic body form of females was the reason to

choose a female as holotype). R. plychona females

have, apart from the leg characteristics separating the

groups, the abdomen less distinctly curved and very

distinct tufts of bristles on the gonocoxae. Males of À.

gyrista can be recognized by the characteristics men-

tioned in the keys. In the key by Nieser & Chen

(1993) R. gyrista runs to R. chrysomalla which, how-

ever, belongs to the R. plychona-group.

Rhagovelia sterea sp. n.

(figs. 23, 31, 45)

Type material. — Holotype (unique specimen),

apterous 2 (ZMAN), INDONESIA: SW Sulawesi,

Mamasa, Sg. Loko, 1400m, 17 Nov 1993, leg. J. P. &

M.J. Duffels (fig. 45).

Description. — Apterous female. Dimensions

length 3.92, width (across base of connexiva) 1.40,

width of head 0.88, width of pronotum 1.25.

Colour generally dark brown-grey to blackish.

Anterior fifth of pronotum, propleura and proster-

num orange-yellow anterior margin of pronotum

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 35-43. Parameres of Rhagovelia: 35 R. abra, 36 R. achna, 37 R. krama, 38 R. skoura, 39 R. tsouloufi, 40 R. ochra, 41 R

gyrista, 42 R. plychona, 43 R. cylindros.

narrowly dark, reaching halfway eyes. Jugae, basal

fourth of first antennal segment, acetabula, coxae and

trochanters, basal third of anterior femur laterally and

most of its anterior (‘under’) surface orange-yellow.

Anterior surface of middle and hind femur brownish,

depending on incidence of light accentuated by pale

pilosity. Outer rim of connexiva, abdominal sternite

7 and most of gonocoxae brown. Venter dull dark

grey to black. Minute black denticles spreading from

jugum and prosternum through propleurae to antero-

lateral parts of pronotum. Body dorsally clothed with

short, not very dense golden pilosity, ventral pilosity

somewhat longer and denser, yellowish. Vertex, sides

of thorax, antennae and legs (except middle and hind

femur) with the usual pubescence and setae.

Length of antennal segments I : II : IN : IV 0.90:

0.43 : 0.70 : 0.50. Pronotum long, covering mesono-

tum. Disk of pronotum with many relatively small

punctures. Length: width of pronotum about 0.9

(1.10/1.25). Length of metanotum on midline, 0.10.

Abdominal tergites 2-5 subequal in length (0.20), ter-

gites 1 and 6 0.30, tergite 7 0.40.

Posterior trochanter length : width 0.35 : 0.16,

without teeth or warts. Posterior femur slender,

slightly over five times as long as wide (1.3/0.25), on

posterior margin in apical part three small teeth.

Posterior tibia straight without teeth. Connexiva ver-

tical, strongly converging on segments 1-4, curved in-

ward on segments 4-5 and slightly convex on seg-

ments 6-7, without caudal points or pilosity (fig. 23).

Tergites 8 and 9, however, with some bristles partly

combined in a tuft. Basal width of abdominal tergite

7 1.25 times its median length (0.50/0.40). Tergite 8

horizontal, as long as tergite 7 (0.40), proctiger fully

visible, pointing caudally. Sternite 7 large, slightly less

than half as long as the preceding abdominal sternites

together (0.6/1.3). Gonocoxa extended, clearly visi-

ble, about half the length of sternite 7, ventrally

slightly compressed, caudally distinctly hairy (fig.

331)

Male and macropterous form unknown.

Etymology. — Stereos, a Greek adjective meaning

solid or stout, refers to the stout build of this species

compared to most other Rhagovelia.

Comparative notes (see keys). — In the key to

groups this species runs to the gyrista-group by the

structure of the female hind femur which is similar to

that of À. hamjadi Polhemus & Polhemus, which has

also a similar general shape. R. hamjadi has, however,

the anterior orange band on pronotum narrow,

reaching inner eye margins, and is slightly smaller. À.

sterea has the dorsal golden pubescence less developed

than related species.

The À. plychona-group

Diagnosis: Large Rhagovelia, length 3.8-4.4 mm,

ground colour dark grey to blackish, body dorsally

with golden pubescence. Posterior legs slender, femur

of male without proximal granulate teeth, femur of

female not distinctly flattened. Hemielytra unicolor-

ous, hairy in latero-proximal quarter, with four closed

cells. Basal dorsal carinae of abdomen in macropter-

ous form not developed.

NIESER ET AL.: Rhagovelia from Sulawesi

Distribution: Endemic to Sulawesi.

Remarks. In general habitus, golden pubescence

dorsally and slender hind legs the À. gyrista and R. ply-

chona groups are quite similar. The structural charac-

teristics of the hind legs and basal carinae on ab-

domen of macropterous form are, however, quite

different. Unfortunately no specimen with complete

hemielytra is known from the R. gyrista-group. R.

chrysomalla Nieser & Chen (1993) provisionally

placed with the R. papuensis group by the authors has

to be transferred to the R. plychona-group.

Key to species of the R. plychona group

(apterous specimens)

1. Yellow anterior transverse band of pronotum

connected with the yellowish propleurae (in

some specimens dark spots behind the eyes),

connexiva broadly yellowish. Connexiva of fe-

male straight, not folded over tergites, abdomen

not curved upward, gonocoxae without tufts of

longbristles. el. are: R. chrysomalla

— Yellow anterior transverse band on pronotum

narrow, at most reaching halfway eyes, propleu-

ra dark, connexiva narrowly yellowish brown.

Connexiva of female curved inward and folded

over tergites (fig. 24), abdomen curved upward

(fig. 33), gonocoxae with distinct tufts of long

buistlest( us NAAN AAN Bee. R. plychona

Rhagovelia plychona sp. n.

(figs. 13, 14, 24, 33, 42, 45)

Type material. — Holotype, apterous d (ZMAN),

INDONESIA: SW Sulawesi, Karangan, ca. 30 km NE of

Enrekang, 1450m, gardens, 10-11 Nov. 1993, leg. J.

P. & M. J. Duffels (fig. 45). — Paratypes (adults only)

with same data as holotype: 348 202 apt., 206

209 macr., 22lvIV/V distributed as follows: 1d

ape dg ne, ‚nunes Il) 1% pe O Mmaer

MBBJ; 30 39 apt., 3d 39 macr. NCTN; 26 29 apt.,

26 22 macr. NHMw; 1d 19 apt., 19 macr. oxuM;

1d 19 apt, 1d 1% macr. SEMC, remaining speci-

mens including allotype apt. ® in ZMAN. — Ad-

ditional paratypes (adults only): SW. Sulawesi, ca. 30

km NE of Enrekang, Gowa camp. Pos 2. 6-9 Nov.

1993, 3°24'15"S 120°00'30"E. Gunung Rantemario,

Sg. Gowa Sarumpa'pa. River through undistd. lower

montane forest, w=3-4m, 1800 m asl. J. van Tol, 34

29 apt. 12 IvIV/V (RMNH).

Description. — Apterous form. Dimensions. Length

3 4.05-4.16, 2 4.30-4.42; width d 1.40-1.48, 2

1.62-1.68; width of head & 0.93-0.97, 2 0.96-1.01;

width of pronotum d 1.25-1.31, 2 1.48-1.52.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 44.

Localities of new species of

Rbagovelia: The more im-

portant islands berween

Sulawesi (Sul), Indonesia

and Mindanao (Min),

Philippines: Si Siau, Sa

Sangihe, type locality of R.

cylindros and R. skoura; Sb

Salibabu, type locality of R.

tsouloufi, Ka Karakelong ad-

ditional localities of À.

tsoulouft.

Colour dark grey to blackish, eyes castaneous.

Orange-yellow transverse band anteriorly on prono-

tum narrow, reaching the inner margins of eyes. Basal

third of first antennal segment, acetabula, coxae and

trochanters (middle trochanter slightly infuscated),

basal half to two thirds of anterior femur and in some

specimens basis of hind femur, pale. Outer rim of

connexiva brown, more distinctly in female than in

male. Venter dull dark grey to black, sternites 7-10

brownish in most specimens.

Minute black denticles restricted to jugum and

proepisternum. Interoculus, thoracic and abdominal

tergites clothed with dense appressed golden-yellow

pilosity (this golden pilosity is also present in larva V).

In addition dorsum of thorax and abdomen with

sparse long erect dark cilia. Vertex sides of thorax, an-

tennae and legs with the usual pubescence and setae

except for hind femur and tibia in male which are

clothed with long hairs.

Length of antennal segments (no apparent differ-

ences between male and female) I: Il : III: IV 0.97:

0.45 : 0.74 : 0.62. Pronotum long, covering mesono-

tum. Disk of pronotum with some indistinct punc-

tures, obscured by pilosity, in posterior part. Length:

width of pronotum about 1.2 (4 1.28/1.05, ?

1.50/1.17). Length of metanotum on midline, 0.10.

Abdominal tergites 1-6 in male subequal in length

(0.20), in female becoming gradually longer posteri-

orly, (from 0.22-0.30) but mostly covered by connex-

iva. Tergite 7 over twice as long as preceding tergites

in male (0.45); 1.5 times as long as preceding in fe-

male (0.45/0.30).

Male. Posterior trochanter length: width 0.4: 0.2,

without teeth or warts. Posterior femur slender, five

times as long as wide (1.5/0.3), on ventroposterior

margin about halfway with a rather small spine, its

length about one sixth the width of femur, with a row

of about seven spines of gradually decreasing length

distally (fig. 13). Posterior tibia straight, armed be-

neath with a row of about 20 short stout teeth giving

a serrate impression. Connexiva slanting upward

about 0.257 or somewhat less, virtually parallel on

segments 1-5 strongly converging posteriad on seg-

ments 6-7, no caudal points. Basal width of abdomi-

nal tergite 7 less than 1.5 times its median length

(0.55/0.45). Abdominal sternite 7 comparatively

short, about 1.2 times as long as sternite 6 (0.3/0.25),

sternite 6 somewhat, sternite 7 distinctly flattened

medially, without a carina. Genital segments promi-

nent (fig. 7), fusiform, tergite 8 longer than tergite 7

(0.55/0.45). Sternite 8 laterally compressed at base,

resulting in a low broad carina ventrally. Median

length of pygophore equal to median length of stern-

ite 8 (0.3). Parameres as in fig. 42.

Female. Posterior femur six times as long as wide

(1.5/0.25). Posterior femur and tibia without teeth or

spines (fig. 14). Connexiva strongly curved inwards,

folded over abdomen leaving on tergites 3-6 only a

narrow median strip uncovered (fig. 24). Caudal apex

of connexiva truncate with little pilosity at the tip,

however, gonocoxae 1 (sternite 8) distinct tufts of

rather long bristles (fig. 33). Tergite 8 nearly hori-

zontal, as long as tergite 7 (0.45). Sternite 7 large,

about two fifth the length of the preceding abdominal

sternites together (0.6/1.5). Gonocoxa clearly visible,

about half the length of sternite 7, ventrally com-

pressed, dorsally with a distinct tuft of bristles.

Proctiger distinctly visible.

Macropterous form. Mostly as apterous except for

modifications of the thorax and presence of wings.

Dimensions, length d 4.80-5.08, 9 4.92-5.18,

hemielytra reaching 0.55-0.67 beyond the apex of ab-

domen; humeral width of pronotum d 1.75-1.80, 9

1.85-1.90, median length of pronotum d 1.71-1.78,

2 1.75-1.85. Hemielytra dull dark grey to blackish,

veins only slightly darker, forming two elongate basal

and two more or less squared apical cells. Hind wings

dark smoky brown-grey. Laterobasal quarter of

hemielytra with golden-yellow pubescence, golden-

yellow pubescence on abdominal dorsum restricted to

a median band. Pronotum with obtuse humeral an-

gles and a rounded caudal apex. Metanotum with a

pair of sublateral pits, second abdominal tergite with

a transverse pitted groove, longitudinal carinae not

developed. Abdomen of female not curved upwards

and with caudal tufts of bristles less prominent than

in apterous form.

Etymology. — Plychonos, greek adjective meaning

pleated, refers to the connexiva folded over dorsum

in apterous females of this species.

Comparative notes (see keys). — In view of their

general shape and dorsal golden pilosity the species of

the À. gyrista and R. plychona groups are superficially

similar. Especially females of À. plychona and R.

gyrista look somewhat similar at first glance but are

easily distinguishable by inspection of the leg charac-

teristics separating the groups. In addition the curva-

ture of the abdomen and the position of the connexi-

va are also somewhat different (figs. 22, 24, 32, 33).

Other species can also be separated by the characteris-

tics mentioned in the group and species keys. In ad-

dition the parameres of R. chrysomalla are very slight-

ly broader than those of R. plychona but this is only

recognizable when specimens are compared. The

swollen caudal part of the proctiger is relatively longer

in R. chrysomalla than in R. plychona (length flat basal

part/ length swollen apical part 0.34/0.42 and

0.40/0.36 respectively).

The Rhagovelia orientalis-group

Diagnosis: Small or medium sized, length up to 3.7

mm, body generally squat, outlines of abdomen

strongly convergent caudally*. Females not or only

NiesER ET AL.: Rhagovelia from Sulawesi

slightly larger than males. Body colour completely

black (except in À. kastanoparuphe), except for a small

orange spot on anterior part of pronotum. Legs black,

usually with a metallic shimmer’, only basal parts yel-

low in some species, middle and hind femur com-

pletely black in most of the species. Juga and proepis-

terna without black spicules. Pronotum long in

apterous specimens, covering the mesonotum. Fore

wing of macropterous specimens with three or four

closed cells, the distal cells reaching the apical third of

the wing. Dealate specimens common*. Abdominal

carinae of macropterous specimens reaching the hind

margin of tergite 2 at most*. Hind trochanter without

teeth or granules, with dense pilosity*. Hind femur of

male slightly to moderately thickened, generally with

a distal row of teeth, proximally without teeth or

granules. Parameres short, crescent-shaped in most of

the species. Abdomen with modifications in most of

the species, dorsally in females and ventrally in males.

Discussion: This species group is a clearly mono-

phyletic group based on several diagnostic characters

(proposed synapomorphies marked with * in diagno-

sis). It shows the close zoogeographical relations be-

tween the Philippines and Sulawesi. Twenty species

are known, 14 from the Philippines (Zettel 1995),

five from the main island of Sulawesi (Nieser & Chen

1933), and one from Sangihe Isle.

Distribution: Philippines (except Palawan), Sula-

wesl.

Key to the species of Sulawesi and adjacent islands

(apterous specimens and macropterous males)

Is Eem alert BE TN 9)

ANSE AO ENEN EE NEE 8

Females

2. Hind margin of tergite 7 with a long, finger-like

process (fig. 34)

— Hind margin of tergite 7 straight or with a short,

triangular process

3. Process on tergite 7 directed backward, tergite 8

with two distinct tufts of hairs in its posterior

Comets) Ws Jatten R. sulawesiana

— Process on tergite 7 directed upward, tergite 8

without distinct tufts of hairs

4. Hind margin of sternite 7 laterally with long

bhckihairs(fie 524) Eee R. cylindros

— _ Hind margin of sternite 7 laterally without long

Rates mer demen ae R. daktylophora

5. Hind margin of tergite 7 straight . À celebensis

— Hind margin of tergite 7 with a small hairy

process in middle or broadly protruding ....6

6. Connexiva and middle of sternite 7 brown,

process of connexivum very long, hind margin of

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

tergite 7 broadly triangular . À. kastanoparuphe

Connexiva and middle of sternite 7 in fully de-

veloped specimens black

Process of connexivum short, but with long

hairs, hind margin of tergite 7 with a very short,

hairy process in middle .... À. pseudocelebensis

Process of connexivum long, acute, process of

tergite 7 generally longer ..... R. daktylophora

Males

Sternites without conspicuous long pilosity,

middle and hind femur basally yellow, connexi-

va brown, paramere distally truncate ........

ROAN EEE da SEO R. kastanoparuphe

Sternites with conspicuous long hairs, middle

and hind femur black, connexiva black, para-

meres distally rounded or pointed

Sternite 6 carinate, long pilosity restricted to this

Carin Drs II R. daktylophora

12%

Fig. 45.

Sulawesi proper: 1 R. achna,

2 R. abra, 3 R. krama and

R. ochra, 4 R. gyrista and R.

sterea, 5 R. gyrista, R. ply-

chona.

Sternite 6 not carinate, pilosity different ... 10

. On sternite 5 and 6, pilosity laterally longer than

along median line R. cylindros

On sternite 5 and 6, pilosity along median line

as long as or longer than laterally ........ 11

. Sternite 7 weakly carinate, ventral margin

strongly concave in lateral view

arte ete RE AN a R. pseudocelebensis

Sternite 7 strongly carinate, ventral margin

straight in lateral view

Tergite 7 about 1.5 times as long as its basal

width er R. sulawesiana

Tergite 7 about 1.1 times as long as its basal

We DER R. celebensis

Rhagovelia celebensis Polhemus & Polhemus

Rhagovelia celebensis Polhemus & Polhemus 1988: 199-200,

figs. 179-186, 229.

Material. — Sulawesi Selatan, E side Lake Matana,

Kg. Salura, 2°32'S 121°28'E, 450m, narrow tributary

Lake Matana, 15.X.1993, 14; same, 20.X.1993, 4d

49 apterous, leg. J. P. & M. J. Duffels (zman).

Sulawesi Utara, Mt. Ambang nr. Kotamobagu, 20

km E of alt. m 1210, 8.X1.1985, leg. J. Krikken, mul-

tistr. evergreen forest, soil litter sifted, 19 apt.

(RMNH).

Rhagovelia kastanoparuphe Nieser & Chen

Rhagovelia kastanoparuphe Nieser & Chen, 1993: 274-275,

figs. 30-34,84.

Nieser & Chen (1993) stated a similarity of À. kas-

tanoparuphe and ‘R. celebensis and related forms’.

Because of some characters (especially in colour and

shape of parameres) which are differing from the oth-

er species of the R. orientalis-group, this species was

provisionally placed in the À papuensis-group, which

was regarded polyphyletic. The occurrence of a

dealate specimen and most of the distinctive charac-

teristics mentioned in the diagnosis show that À. kas-

tanoparuphe can best be placed in the À orientalis-

group, although it seems not to be closely related with

any other described species in this group.

Rhagovelia pseudocelebensis Nieser & Chen

Rhagovelia pseudocelebensis Nieser & Chen 1993: 275, figs.

35-40, 80-81.

Material. — Sulawesi Selatan, Batusitanduk {2°48'S

120°10'E} N of Palopo, narrow river, 2.XI.1993, leg.

J.P. & M. J. Duffels, 45 39; 15 km W of Palopo,

2°57'10"S 120°07'30"E, gardens and disturbed rain

forest a30X1993x legen]. Pace MJ Duftels, 12

(ZMAN).

Rhagovelia sulawesiana Polhemus & Polhemus

Rbagovelia sulawesiana Polhemus & Polhemus 1988: 198-

199, figs. 158-164, 230.

Material. — Sulawesi Selatan, SW Celebes, 1100m,

Mt. Lompobatang area, Malino {ca 5°15'S

119°48'E}, 2, 8-10.VI.1982, M. A. Lieftinck, 58 29

apt., 1d 19 macr. (RMNH).

Although Polhemus & Polhemus (1988) stated the

similarities between À. sulawesiana and R. celebensis,

they decided to class them within different groups

without explanation. Rhagovelia celebensis was cor-

rectly regarded as a species of the À. orientalis-group,

but À sulawesiana as a species of the À. papuensis-

group. The classification of apterous specimens of

these groups may be difficult using the characters giv-

en by Polhemus & Polhemus (1988), but the

macropterous specimens studied show clearly that À.

NIESER ET AL.: Rhagovelia from Sulawesi

sulawesiana also belongs to the R. orientalis-group. In

addition the body shape, the teeth on hind femur,

and the lack of spicules on proepisterna and juga are

typical for this group. The shape of the paramere of

male and the long process of the tergite 7 of female

show a close relationship with À. cylindros sp.n. and

R. daktylophora.

Rhagovelia cylindros sp. n.

(figs. 1, 21, 34, 43, 44)

Type material. — Holotype apterous 9 (ZMAN) IN-

DONESIA: Sulawesi Utara, Pulau Sangihe, Desa Laine,

Sungai Laine, 12. Nov. 1994, N9463, leg. N. Nieser

(fig. 44). Sungai Laine [different from N9454/56

which is on the other side of the island!], rather large

pothole at water fall used for swimming (and picnics

on the banks ‘proyek wisata = tourist spot’), a lot of

Hydrilla. Rhagovelia at open to moderately open

spots. Paratypes (NCTN unless otherwise indicated),

same data as holotype, 248 102 apterous, 3d 59

macropterous, distributed as follows: 12 apt. (allo-

type) ZMAN, 16 19 apt., 16 macr. NHMw, 36 39

OXUM. Additional paratypes (adults only) all P.

Sangihe and leg. N. Nieser: Desa Simuang (near

Malahu), Sungai Simuang, 28.VI.1994, N9457.

Mountain stream, water clear, hyaline, boulders,

stones, stretches with sand, 1d 39 apt., 12 macr.

(RMNH); Desa Utaurano, Sungai Apanukang,

14.X1.1994, N9465. Upstream tributary of S. Laine

(N9454 &-56). Mountain stream, bottom boulders,

large stones and conglomerate rock, 5-10m wide,

depth variable (up to >1m, mostly about 0.3 m), ve-

locity of current variable, 336 329 apt., 2d macr.

(MUDH, SEMC 1d 19 apt. each); N9465A: Rather

large pothole, 40x15m c. 1m deep in mountain

stream, virtually stagnant, bottom rock covered with

fine silt. Used e.g. for rearing goldfish in cages in the

ipond3:39ig 15. Ssapt ISA maer (Ods 2 9uapr,

16 macr. numw, 46 39 prec, 2d 29 mBBJ, 2d

22 uscp).

Description. — Apterous form. Rather small gener-

ally dark species. Dimensions. Length d 2.88-3.00,

® 3.15-3.40; width (across mesoacetabula) d 1.20-

1.30, 2 1.32-1.40; width of head & 0.76-0.80, 2

0.76-0.80; width of pronotum d 0.98-1.00, 2 0.92-

1.04.

Colour generally dark grey to black, yellowish

transverse band anteriorly on pronotum narrow,

reaching to inner margins of eyes, indistinct due to a

cover of grey pubescence. Basal third of antennal seg-

ment 1, distal part of anterior and posterior acetabu-

la, apical rim of mesoacetabula, anterior and posteri-

or coxae, anterior and part of posterior trochanters

and in most specimens variable part proximally of an-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

terior femur yellow to light brown. Connexiva entire-

ly grey to black. Pro- and mesopleura except for its

dorso-posterior quarter, pruinose, light grey.

Minute black denticles absent. Posterior three

quarters of pronotum indistinctly punctured, pleura

smooth without recognizable rows of punctures.

Vertex, inter oculus, sides of thorax and abdomen,

antennae and legs with the usual pubescence and se-

tae. Body clothed with short pubescence, virtually ab-

sent laterally in females, superimposed by longer yel-

low bristles which are especially strongly developed

ventrally in males and caudally in both sexes but

much more prominent in females (figs. 21, 34).

Length of antennal segments I: II: III : IV d 0.81

210/421 0 48110 46 210.7892 0:40) ACID AT

Pronotum long, covering mesonotum, shorter than

wide (c. {0.8-0.9}/1.0). Length of metanotum on me-

dian line, 0.1. Abdominal tergites 1-4 subequal in

length (0.15), remaining tergites in female covered by

connexiva, in male tergite 5 subequal, 6 slightly (0.2)

and 7 distinctly (0.4) longer. Abdominal sternites 3-5

subequal, about 0.15 long, sternites 2 and 6 slightly

longer 0.20, sternite 7 distinctly longer, 0.35 in male.

Sternite 2-5 subequal, 0.20; 6 slightly (0.25) and 7

distinctly (0.55) longer in female.

Male. Hind femur slightly incrassate, on average

slightly over five times as long as wide (1.34/0.26).

Ventral margin slightly distally of halfway with a sin-

gle larger spine, half as long as the width of femur, fol-

lowed by a double row of about seven small teeth (fig.

1). Posterior tibia virtually straight, armed beneath

with a row of about 15 small teeth which are shorter

than and covered by the pilosity of the tibia.

Connexiva more or less horizontal to slanting slight-

ly upward (less than 0.157), gradually converging

posteriad. Basal width of abdominal tergite 7 three

quarters its median length (0.3/0.4). Tergite 8 basally

as wide as long (0.35). Abdominal sternites 5 and 6

flattened with an indication of a keel and pilosity lat-

erally longer than medially, sternite 7 ventrolaterally

compressed with a distinct though low and compara-

tively broad, median keel. Parameres as in fig. 43.

Female. Posterior femur as in male, about five

times as long as wide (1.23/0.25). Posterior tibia un-

armed except for a small apical spine. Connexiva

more or less vertical strongly converging on segments

1-5, nearly meeting over abdomen on segment 6 and

anterior part of 7, then diverging to make room for

the erect finger-like projection on tergite 7 (fig. 21).

Apex of connexiva with a triangular caudal projec-

tion, caudal margin accentuated by a dense fringe of

large bristles (fig. 34). Tergites 8 and 9 vertical, only

visible in caudal view.

Macropterous form. — Essentially as apterous form

except for development of hemielytra and thorax.

Length d 3.40-3.46, 9 3.58-3.62, humeral width of

pronotum d 1.28-1.42, © 1.37-1.41, median length

of pronotum d 1.18-1.22, 9 1.23-1.28. Pronotum

black orange-yellow transverse band at anterior mar-

gin reaching to inner margins of eyes, rather indis-

tinct due to cover by whitish pubescence. Propleura

with a distinct row of punctures near ventro-posterior

margin. Hemielytra reaching 0.4 beyond apex of ab-

domen, brown with darker veins. A pair of elongate

proximal cells not reaching halfway hemielytron, an

elongate outer apical cell (near costal margin) reach-

ing to one fourth from apex, central cell absent. Keels

on base of abdomen well-developed but short, reach-

ing only to about halfway tergite 2. Connexiva of fe-

male vertical, not bent over abdomen.

Etymology. — Cylindros (kulindros), Greek noun

meaning cylinder, referring to the form of caudal part

of abdomen in female.

Comparative notes. — This species is closely related

with À. sulawesiana and R. daktylophora, with which

it shares the long process on tergite 7 of females.

Females differ from À. sulawesiana in the upwardly

directed process on tergite 7 (posteriorly directed in

R. sulawesiana), from R. daktylophora in the blunt

process of connexiva, which are nearly touching each

other (but clearly separated in À daktylophora), and

from both in the long black hairs laterally on hind

margin of sternite 7. Males differ from all the other

species of the group in the hairs on sternites 5 and 6

which are laterally longer than medially.

The Rhagovelia sarawakensis-group

Diagnosis: Small and slender species, length less

than 3 mm. Colour black with a short orange brown

band on pronotum. Pronotum in apterous specimens

shorter than length of an eye, about one third as long

as the exposed mesonotum. Fore wing with three or

four closed cells slightly reaching distal half.

Abdominal carinae of winged specimens extending

posteriorly to tergite 3. In dealate specimens fore

wing broken in middle, behind closed cells. Tergite 8

of female horizontal.

Distribution: Sri Lanka, Southeast Asia and South

China, Ryukyu Islands, Taiwan, Palawan, Borneo,

Sumatera, Java, Bali, Nusa Tenggara.

Only one species, R samarinda, is reported from

Sulawesi by the record of a single macropterous fe-

male (Polhemus & Polhemus 1988). There are no

further records from Sulawesi, and the species group

is also not known from other parts of Wallacea in-

cluding the Philippines (except Palawan). Therefore

the authors regard this single record as doubtful.

A single male of À. sumatrensis Lundblad, which

according to Polhemus (1990) belongs to this group,

from Lombok (new record) is deposited in RMNH. As

this species was previously recorded from Flores and

Sumbawa (Nieser & Chen 1992) it can probably be

found in most of the Nusa Tenggara.

Check list of species groups and species of

Rhagovelia Mayr known from the province of

Sulawesi.

R. papuensis-group

abra sp. n.

achna sp. n.

blogiokommena Nieser & Chen, 1993

grayi Polhemus & Polhemus, 1988

horaia Nieser & Chen, 1993

kalami Nieser & Chen, 1993

krama sp. n.

lorelinduana Polhemus & Polhemus, 1988

minahasa Polhemus & Polhemus, 1988

ochra sp. n.

pruinosa Polhemus & Polhemus, 1988

robina Nieser & Chen, 1993

skoura sp. n.

tsouloufi sp. n.

trichota Nieser & Chen, 1988

tropidata Nieser & Chen, 1993

unica Polhemus & Polhemus, 1988

wallacei Polhemus & Polhemus, 1988

R. gyrista-group

gyrista sp. n.

hamdjadi Polhemus & Polhemus, 1988

R. plychona-group

chrysomalla Nieser & Chen, 1993

plychona sp. n.

R. orientalis-group

celebensis Polhemus & Polhemus, 1988

cylindros sp.n.

daktylophora Nieser & Chen, 1993

kastanoparuphe Nieser & Chen, 1993

pseudocelebensis Nieser & Chen, 1993

sulawesiana Polhemus & Polhemus, 1988

R. sarawakensis-group

? samarinda Polhemus & Polhemus, 1988

NIESER ET AL.: Rhagovelia from Sulawesi

REFERENCES

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1993. The insect and spider collections of the world. —

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Lansbury, I., 1993. Rhagovelia of Papua New Guinea,

Solomon Islands and Australia (Hemiptera-Veliidae). —

Tijdschrift voor Entomologie 136: 23-54.

Matsuda, R., 1956. A supplementary study of the genus

Rhagovelia (Hemiptera, Veliidae) of the Western

Hemisphere. A deductive method. — University of Kansas

Science Bulletin 38(I): 915-1017.

Nieser, N. & P. P. Chen, 1992. Notes on Indonesian water-

bugs (Nepomorpha & Gerromorpha). — Storkia 1: 30-

40.

Nieser, N., & P. P. Chen, 1993. The Rhagovelia

(Heteroptera: Veliidae) of Sulawesi (Indonesia). —

Tijdschrift voor Entomologie 136: 259-281.

Polhemus, J. T., 1990. Miscellaneous studies on the genus

Rhagovelia Mayr (Heteroptera: Veliidae) in Southeast

Asia and the Seychelles Islands, with keys and descrip-

tions of new species. — Raffles Bulletin of Zoology 38: 65-

75.

Polhemus, J. T., & D. A. Polhemus, 1988. Zoogeography,

ecology, and systematics of the genus Rhagovelia Mayr

(Heteroptera: Veliidae) in Borneo, Celebes, and the

Moluccas. — Insecta Mundi 2: 161-230.

Yang, C.M. & Polhemus D. A., 1994. Notes on Rhagovelia

Mayr (Hemiptera: Veliidae) from Singapore, with de-

scription of a new species. — The Raffles Bulletin of

Zoology 42: 987-993.

Zettel, H., 1994. Revision der philippinischen Arten der

Gattung Rhagovelia Mayr, 1 Teil (Heteroptera: Veliidae).

— Entomological Problems 25: 33-48.

Zettel, H., 1995. Revision der philippinischen Arten der

Gattung Rhagovelia Mayr (Heteroptera: Veliidae). 2 Teil

— Entomological Problems 26: 43-78.

Received: 30 May 1996

Accepted: March 1997

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

JoHN T. POLHEMUS

Colorado Entomological Museum, Englewood, Colorado, USA.

SEVEN NEW SPECIES OF HYDROTREPHES CHINA

(HELOTREPHIDAE: HETEROPTERA) FROM SULAWESI

Polhemus, J. T., 1997. Seven new species of the genus Hydrotrephes China from Sulawesi.

Tijdschrift voor Entomologie 140: 43-54, figs. 1-55 [issn 0040-7496]. Published 31 October

1997.

A definite locality in Sulawesi is established for Hydrotrephes bouvieri (Kirkaldy). The following

new species are described; Hydrotrephes celebensis Polhemus sp. n., H. kamarora Polhemus sp. n.,

H. marana Polhemus sp. n., H. nieseri Polhemus sp. n., H. taweli Polhemus sp. n., H. variegatus

Polhemus sp. n., and H. viriosus Polhemus sp. n. Some morphological features of Helotrephidae

are discussed.

Dr. J. T. Polhemus, Colorado Entomological Museum, 3115 S. York, Englewood , Colorado

80110 USA.

Key words. — Indonesia; Sulawesi; Helotrephidae; Hydrotrephes new species.

The following descriptions of new taxa and notes on

Hydrotrephes bouvieri (Kirkaldy) are needed for a forth-

coming publication on the Nepomorpha of Sulawesi

by Nieser and Chen. These descriptions have been pre-

pared for some time, intended to be part of a compre-

hensive revision of the genus Hydrotrephes, however the

above intended publication by Nieser and Chen and a

revision of the Helotrephidae of the Philippines in-

tended by Herbert Zettel have persuaded me to instead

deal with the genus on a regional basis. The material

described here was mostly collected by D. A. Polhemus

and myself during a 1985 expedition to the Malay

Archipelago and Southeast Asia supported by the

National Geographic Society. We have collected many

species (mostly undescribed) of Hydrotrephes on Leyte,

Luzon, Mindanao, Mindoro, and Palawan in the

Philippines; Bali, Java, Sulawesi, north and south

Sumatra, Sumba, and Timor in Indonesia; East

Malaysia (Sabah) and West Malaysia. Additional

species of the genus were collected by M. Sat6 on

Bohol and Cebu in the Philippines, and by Taylor and

Messer on Moratai, North Moluccas. Zettel (in litt.)

possesses other species from Thailand, Laos, East

Malaysia (Sarawak), and Indonesia (Nias). Thus the

genus Hydrotrephes, with only six previously described

species from Luzon, Sri Lanka, Sulawesi and Sumatra

(all represented in the J. T. Polhemus Collection), is

actually speciose and widespread; the general distribu-

tion, by region, was given by Polhemus & Polhemus

(1990), along with a pattern of endemism exhibited by

Rhagovelia (Veliidae) and Ptilomera (Gerridae) that is

similar to some Hydrotrephes species on Sulawesi.

The last systematic reviews of the major classifica-

tion of Helotrephidae were given by Esaki and China

(1927, 1928) and China (1935), however Papäcek,

Stys and Tonner (1988) have provided an extensive re-

view and morphological analysis in connection with

the description of a new subfamily. The genus

Hydrotrephes (type species, Helotrephes bouvieri

Kirkaldy) was established by China (1935), after previ-

ously being carefully described by Esaki and China

(1928, as Helotrephes) based on syntypes of bouvieri.

Various authors have dealt with other genera of

Helotrephidae and described new subfamilies, tribes,

genera and species, but no new species of Hydrotrephes

have been described for 66 years.

All measurements are given in millimeters. The

holotypes and some paratypes will be placed in the

Smithsonian Institution (USNM); all other specimens

are in the Polhemus collection (JTPC), except some du-

plicate material that will be distributed to at least the

Nieser Collection (NcTN), American Museum of

Natural History (AMNH), Naturhistorisches Museum

Wien (NHMW) and the Museum Zoologicum

Bogoriense (MBBJ).

SPECIES DISCRIMINATION

A number of characters were evaluated as potential-

ly useful in discriminating species of the genus

Hydrotrephes, some previously used by other authors.

The morphological nomenclature follows Esaki and

China (1928) and Papácek, Stys and Tonner (1988)

unless otherwise noted. I have concluded that that

species discrimination should primarily rely on the

morphology of the male genitalia, and to a lesser degree

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Zan

Figs. 1-4. Hydrotrephes bouvieri Kirkaldy. — 1, Male ventral paramere; 2, Male dorsal paramere; 3, Aedeagus; 4, Female ab-

dominal sternite VII. Internal phragma indicated by dashed line.

Figs. 5-11. Hydrotrephes celebensis sp. n. — 5, Cephalonotum, anterior view; 6, Pronotal and genal plates, meso- and proepis-

terna. g - pronotal and genal plates, m - mesoepisternum, p - proepisternum; 7. Ventral abdominal carinae II, III; 8, Male

ventral paramere; 9, Male dorsal paramere; 10, Aedeagus; 11, Female abdominal sternite VII. All scale bars = 0.5 mm.

the female abdominal sternite VIT and pattern of dark

markings on the cephalonotum; all other characters are

variable and must be used in conjunction with the for-

mer, with caution. For this reason a key has not been

provided, but figures are presented instead to aid in the

separation of species.

Lateral pronotal and genal plates: These structures

are well developed in all Helotrephini (Esaki & China

1928, fig. la) but lacking in all Limnotrephini. The

shape, particularly of the distinct ‘notch’ below the eye,

is variable, and although sometimes helpful, should

usually not be considered as diagnostic by itself.

Overall body size: The overall size is useful in sepa-

rating species, and seems remarkably constant for indi-

vidual populations of each species, except for the spec-

imens of the sympatric variety of H. nieseri from

northern Sulawesi which are much smaller than the

nominate form (see discussion below). The macropter-

ous form (with a claval suture) is usually darker and

dorsoventrally larger than the brachypterous form

(without a claval suture; hemelytra brachypterous,

hindwings micropterous), and these two morphs may

appear to be different species at first glance.

Male terminalia and genitalia: The morphological

details of the complex terminal abdominal segments

and genitalia of Hydrotrephes males are the most reli-

able characters for species separation. This includes the

ventral laterotergites of segments VII and VIII, the

posterior margin of abdominal segment IX, the para-

meres, and the aedeagus.

Mesopleural and propleural plates: The shapes of

these plates are sometimes useful in discriminating be-

tween closely related sympatric species.

Ventral abdominal carinae: This character was used

by Esaki and China (1927, 1928) and China (1935),

but has proved to be variable and generally unreliable

except for gross features. A few species have extreme

modifications which can be diagnostic.

Prosternal carina: The prosternal carina is variable,

but sometimes helpful when extreme differences are

evident between species.

Female abdominal sternite VII. This structure, often

called the subgenital plate, is sometimes diagnostic and

often helpful, but the differences between species are

often subtle, and closely related species may exhibit a

similar shape.

Color pattern: The pattern of dark markings on the

cephalonotum, in particular on the frons, are helpful in

discriminating between closely related species. While

the pattern may vary considerably in a given popula-

tion, it varies in a predictable way, with certain ‘mark-

ers constant (analogous to the widely used hemelytral

‘eunomy’ in the family Saldidae). The maculations of

the hemelytra are also sometimes useful in separating

sympatric species.

Stridulatory mechanisms: All species of the genus

PoLHEMUS: Hydrotrephes of Sulawesi

Hydrotrephes possess a stridulatory mechanism, as de-

scribed by Polhemus (1990). These appear to be mo-

notonous and not species diagnostic. It is conceivable

that an examination by SEM would reveal diagnostic

fine structure not visible at 80 X.

Hydrotrephes bouvieri (Kirkaldy)

(figs.1-4)

Helotrephes bouvieri Kirkaldy 1904: 129. Syntypes 4 males, 2

of unknown sex, Celebes, de la Savinière, National

Museum of Natural History, Paris [not examined]; Esaki

and China 1927: 281; Esaki and China 1928: 143 (re-

description, figures); Lundblad 1933: 114.

Hydrotrephes bouvieri — China 1935: 594 (new genus, key);

Miyamoto 1952: 2; Poisson 1960: 333; Polhemus 1990:

60 (world checklist).

Material examined. — 46 brachypterous adults, 3

nymphs, Sulawesi Utara, Kab. Bolaang Mongondow,

Lake Mala (Moat), E of Kotamobagu, 0°44' N,

124°27' E, 1000 m el., CL 2113, 10 Sept: 1989, Joa

& D. A. Polhemus (JTPC, NCTN, USNM, MBBJ, NHMW).

Discussion. — The males at hand match in every de-

tail the exhaustive description given by Esaki & China

(1928). The male aedeagus and parameres are shown

in figs. 1-3. Because the female of this species was not

previously known, sternite VII is figured here (fig. 4).

No definite locality was previously established for this

species, as the labels on the syntypes simply gave

‘Celebes’. The habitat of Hydrotrephes bouvieri is un-

usual for the genus, as it was collected in a lake rather

than a stream. The habitat of bouvieri was in tangled

root masses, along the wave undercut edges of isolated

emergent clumps of reeds that formed ‘pedestals’ in

shallow water along the lake shore. Hydrotrephes corpo-

raali China has been collected in both lakes and

streams (Lundblad 1933; Polhemus unpubl.), so it is

likely that H. bouvieri also inhabits streams.

Hydrotrephes celebensis Polhemus sp. n.

(figs. 5-11)

Type material: Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Selatan Prov., Marana

River, nr. Camba, 50 km E of Maros, CL 2167, 450 m

el., 14 Oct. 1985, J. T. & D. A. Polhemus (usnm).

Paratypes, 7 brachypterous adults, 8 macropterous

adults, same data as holotype (JTPC, NCTN, USNM,

MBBJ).

Size. — Brachypterous form: length 2.40 - 2.42 mm,

width across base of cephalonotum 2.02 - 2.08 mm.

Colour. — Brachypterous form: ground color yellow-

ish, heavily marked with brown. Cephalonotum with

complex pattern of deep brown markings (fig. 5);

pronotum with irregular transverse light band across

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 12-18. Hydrotrephes kamarora sp. n. -12, Cephalonotum, anterior view; 13, Pronotal and genal plates; 14, Ventral ab-

dominal carinae II, III; 15, Male ventral paramere; 16, Male dorsal paramere; 17, Aedeagus; 18, Female abdominal sternite

VII.

middle. Base of scutellum with a transverse deep

brown stripe. Hemelytra, scutellum with strong pat-

tern of deep brown irregular, randomly scattered mark-

ings covering about half of dorsum. Venter brown.

Legs, antennae yellowish, legs darker basally, rostrum

brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, set with small alveoli, in dorsal view

(of entire insect) broader than long (2.08: 1.26), later-

al margins carinate behind eyes. Mesopleural and pro-

pleural plates notched (fig. 6). Eye length/width,

0.54/0.29. Interocular distance 0.86. Rostral segments

1, 2 extremely short; lengths of rostral segments 3: 4;

0.18: 0.43.

Scutellum shining, finely rugulose, set with tiny

alveoli, cach with one slender setae, each surrounded

by a roughly circular transparent region; in some spec-

imens alveoli very clearly delineated due to underlying

structure or pigmentation visible through translucent

cuticle; length: width; 1.15: 1.15.

Hemelytra opaque, faintly rugulose, set with alveoli

each bearing a slender seta; distal locking tab (pseudo-

membrane) on right hemelytron of usual form. Hind

wings reduced to small membranous strips. Ventral

carinae of abdominal segments II, III as in fig. 7.

All femora with pectinate bristles beneath, longer

basally. Claws of each leg about half as long as distal

tarsal segment. Male genital segments modified, twist-

ed strongly to the left; aedeagus and parameres as

shown in figs. 8-10. Female abdominal segments sym-

metrical, subgenital plate (sternite VII) as shown in fig.

Wl.

Macropterous form: similar in size, color and struc-

ture to brachypterous form, except for presence of

claval suture and well developed hind wings.

Discussion. — Hydrotrephes celebensis sp. n. is so far

known from a single population. The body size of both

sexes and wing morphs is unusually uniform. The pat-

tern of dark markings on the cephalonotum, and male

genitalia are diagnostic.

Etymology. — The name celebensis refers to the island

of origin.

Hydrotrephes kamarora Polhemus sp. n.

(figs. 12-18)

Type material: Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Tengah Prov., stream 10

km SE of Kamarora, Lore Lindu National Park, CL

2156, 950 mel., 8 Oct. 1985, J. T. & D. A. Polhemus

(usnm). Paratypes (nymphs not paratypes), Indonesia,

Celebes: 4 brachypterous adults, 6 macropterous

adults, 8 nymphs, same data as holotype (JTPC, NCTN,

USNM); 2 macropterous males, 1 macropterous female,

Sulawesi Tengah Proy., stream 9 km E of Taweli, CL

ICO MIS OEM el tO Oct MOSS AIN SDAL

Polhemus; 1 brachypterous male, 2 brachypterous fe-

males, 1 nymph, Sulawesi Utara Prov., forest stream S

of Lake Mala, CL 2118, 1200 m el., 11 Sept. 1985, J.

T. & D. A. Polhemus (yrPc).

Size. — Brachypterous form: length 2.88 - 3.42 mm,

width across base of cephalonotum 1.94 - 2.38 mm.

Colour. — Brachypterous form: ground color yellow-

ish, heavily marked with brown. Cephalonotum often

completely dark anterior to suture, except along eyes

and a small median light streak; lighter colored speci-

mens with a more complex pattern of deep brown

markings (fig. 12); pronotum with irregular transverse

light band across middle. Scutellum basally dark,

lighter posteriorly, with several yellowish areas medial-

ly. Hemelytra dark on basal fourth, lighter posteriorly,

with scattered lighter regions; some specimens almost

completely dark. Venter yellowish brown. Legs, anten-

nae yellowish, rostrum brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, set with alveoli, in dorsal view (of en-

tire insect) broader than long (2.34 : 1.44), lateral mar-

gins carinate behind eyes. Mesopleural and propleural

plates shallowly notched (fig. 13). Eye length/width,

0.58/0.29. Interocular distance 1.19. Rostral segments

1, 2 extremely short; lengths of rostral segments 3: 4;

0.22 : 0.54.

Scutellum shining, not rugulose, set with tiny alve-

oli, each with one slender setae, each surrounded by a

roughly circular transparent region; in some specimens

alveoli very clearly delineated due to underlying struc-

ture or pigmentation visible through translucent cuti-

cle; length: width, 1.48: 1.33.

Hemelytra opaque, shining, set with alveoli each

PoLHEMUS: Hydrotrephes of Sulawesi

bearing a slender seta; distal locking tab (pseudomem-

brane) on right hemelytron of usual form. Hind wings

reduced to small membranous strips. Ventral carinae of

abdominal segments II, III as in fig. 14.

AIl femora with pectinate bristles beneath, longer

basally, densest on anterior femora, sparsest on posteri-

or femora. Claws of each leg about one-third as long as

distal tarsal segment. Male genital segments modified,

twisted strongly to the left; aedeagus and parameres as

shown in figs. 15-17. Female abdominal segments

symmetrical, subgenital plate (sternite VII) as shown in

fig. 18.

Macropterous form: similar in size, color and struc-

ture to brachypterous form, except for presence of

claval suture and well developed hind wings.

Discussion. — Hydrotrephes kamarora sp. n. is so far

known from three populations. The body size of both

sexes and wing morphs is quite uniform within each

population, but varies considerably between popula-

tions. Elevation does not seem to be correlated with

size, as the largest specimens are from the type locality

at 950 m, the smallest from near Lake Mala at 1200 m,

and the intermediates from east of Taweli at 150 m. In

all specimens the scutellum is shining and not rugu-

lose. The pattern of dark markings on the cephalono-

tum, and male genitalia are diagnostic.

Etymology. — The name kamarora is a noun in ap-

position, referring to the village nearest the type locali-

ty.

Hydrotrephes marana Polhemus sp. n.

(figs. 19-26)

Type material: Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Selatan Prov., Sungai

Pattanuang at Biseang Labboro Recreation Area, 7 km

SW of Bantimuring, CL 2165, 0-100 m el., 13 Oct.

1985, J. T. & D. A. Polhemus (usnm). Paratypes

(nymphs not paratypes), Indonesia, Celebes: 7

brachypterous males, 1 brachypterous female, 9

nymphs, same data as holotype (JTPC, NCTN); 2

brachypterous males, 1 brachypterous female, Sulawesi

Selatan Prov., Marana River nr. Laiya, CL 2167, 0-100

m el., 14 Oct. 1985, J. T. & D. A. Polhemus (yrec).

Size. — Brachypterous form: length 2.99 - 3.09 mm,

width across base of cephalonotum 2.09 - 2.16 mm.

Colour. — Brachypterous form: ground color yellow-

ish brown to fuscous, moderately marked with brown.

Cephalonotum with complex pattern of deep brown

markings (fig. 19); pronotum largely yellowish brown,

with rather regularly spaced maculations. Base of

scutellum with a broad transverse brown stripe.

Hemelytra, scutellum with numerous small deep

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 19-26. Hydrotrephes marana sp. n. -19, Cephalonotum, anterior view; 20, Pronotal and genal plates, meso- and proepis-

terna; 21, Ventral abdominal carinae II, III, CL 2167; 22, Ventral abdominal carina III, CL 2165; right, male; left, female;

23, Male ventral paramere; 24, Male dorsal paramere; 25, Aedeagus; 26, Female abdominal sternite VIT.

brown irregular, randomly scattered markings, not

dense, often anastomosing. Venter brown. Legs, anten-

nae yellowish, rostrum yellowish to brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, densely set with small alveoli, in dor-

sal view (of entire insect) broader than long (2.12 :

1.37), lateral margins carinate behind eyes.

Mesopleural and propleural plates deeply notched (fig.

20). Eye length/width, 0.58/0.29. Interocular distance

1.01. Rostral segments 1, 2 extremely short; lengths of

rostral segments 3: 4; 0.25 : 0.54.

Scutellum faintly shining, finely rugulose, set with

tiny alveoli, each with one slender setae, each sur-

rounded by a roughly circular transparent region; in

some specimens alveoli very clearly delineated due to

underlying structure or pigmentation visible through

translucent cuticle; length: width, 1.30: 1.33.

Hemelytra opaque, faintly shining, finely rugulose,

set with alveoli each bearing a slender seta; distal lock-

ing tab (pseudomembrane) on right hemelytron of

usual form. Hind wings reduced to small membra-

nous strips. Ventral carinae of abdominal segments II,

III as in figs. 21-22.

Fore and middle femora with pectinate bristles be-

PoLHemus: Hydrotrephes of Sulawesi

Figs. 27-35. Hydrotrephes nieseri sp. n. — 27, Cephalonotum, anterior view, typical variety; 28, Cephalonotum, anterior view,

variety minutus, 29, Pronotal and genal plates; 30, Ventral abdominal carinae II, III; 31, Male abdominal segment IX, ven-

tral view; 32, Male ventral paramere; 33, Male dorsal paramere; 34, Aedeagus; 35, Female abdominal sternite VII.

neath, longer basally; hind femora mostly bare. Claws of

each leg about one-third to half (on posterior) as long as

distal tarsal segment. Male genital segments modified,

twisted strongly to the left; aedeagus and parameres as

shown in figs. 23-25. Female abdominal segments sym-

metrical, subgenital plate (sternite VII) as shown in fig. 26.

Macropterous form: unknown.

Discussion. — H. marana is so far known only from

a limited area of Sulawesi Selatan Province. The pat-

tern of dark markings on the cephalonotum, and male

genitalia are diagnostic.

Etymology. — The name marana is a noun in appo-

sition, referring to the river where part of the type series

was collected.

Hydrotrephes nieseri Polhemus sp. n.

(figs. 27-35)

Type material. — Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Utara Prov., tributary to

Tumpah River, 0° 35' N, 123° 54' E, 62 km SW of

Kotamobagu, CL 2101, -235 m el. 4 Sept. 1985, J. T.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

& D. A. Polhemus (usnm). Paratypes (all collected by

J. T. & D. A. Polhemus; nymphs not paratypes),

Indonesia, Celebes, Sulawesi Utara Prov.: 144

brachypterous adults, 14 macropterous adults, 17

nymphs, same data as holotype (JTPC, NCTN, AMNH,

USNM, MBBJ, NHMW); 54 brachypterous adults, 4

macropterous adults, 15 nymphs, Tumpah River, 0°

35' N, 123° 54' E, 60 km SW of Kotamobagu, CL

2100, 211 m el., 3 Sept. 1985; 2 macropterous fe-

males, stream, trib. of Sungai Ongaak Mongondow, 4

km S of Inobonto, CL 2106, 6 Sept. 1985; 24

brachypterous adults, 13 macropterous adults, 2

nymphs, upper Sungai Metelanga and tributary, 10

km Sof sWoloduas Gl 2103, 72 Seper 1985;76

brachypterous males, 2 brachypterous females, warm

stream, 8 km S of Dolodua on Malibagu rd., CL 2110,

7 Sept. 1985; 27 brachypterous adults, 1 macropterous

female, 3 nymphs, Sungai Metelanga, 5 km S of

Dolodua, CL 2111, 7 Sept. 1985; 13 brachypterous

adults, 1 nymph, cave spring and stream at

Komangaan, NW of Kotamobagu, CL 2120, 14 Sept.

1985; 4 brachypterous males, 2 brachypterous females,

1 macropterous female, Pononontuna River at

Tapakulintang, 200 m el., CL 2121, 15 Sept. 1985; 14

brachypterous adults, 2 macropterous females,

Tondano river tributary, S of Airmididi, CL 2127, 20

Sept. 1985; 1 macropterous male, stream W of

Danowudu, E of Manado, CL 2129, 20 Sept. 1985

(ITPE).

Size. — Brachypterous form: length 2.38 - 2.95 mm,

width across base of cephalonotum 1.66 - 2.02 mm.

Colour. - Brachypterous form: ground color yellow-

ish brown to fuscous, extensively marked with brown.

Cephalonotum with complex pattern of deep brown

markings (fig. 27, 28); pronotum largely yellowish

brown, broadly embrowned anteriorly along suture,

narrowly dark posteriorly, with a few median macula-

tions in a broad sordid yellowish transverse band. Base

of scutellum with a broad transverse brown stripe.

Hemelytra, scutellum with numerous large brown ir-

regular, randomly scattered markings, not well de-

fined, often anastomosing, more numerous basally.

Venter brown. Legs, antennae yellowish, rostrum

brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, densely set with small alveoli, in dor-

sal view (of entire insect) broader than long (2.02 :

1.08), lateral margins carinate behind eyes.

Mesopleural and propleural plates deeply notched (fig.

29). Eye length/width, 0.50/0.27. Interocular distance

0.97. Rostral segments 1, 2 extremely short; lengths of

rostral segments 3: 4; 0.22 : 0.43.

Seutellum faintly shining, finely rugulose, set with

tiny alveoli, each with one slender setae, each sur-

rounded by a roughly circular transparent region; in

some specimens alveoli very clearly delineated due to

underlying structure or pigmentation visible through

translucent cuticle; length: width, 1.30 : 1.22.

Hemelytra opaque, faintly shining, finely rugulose,

set with alveoli each bearing a slender seta; distal lock-

ing tab (pseudomembrane) on right hemelytron of

usual form. Hind wings reduced to small membranous

strips. Ventral carinae of abdominal segments II, III as

in fig. 30.

All femora with pectinate bristles beneath, longer

basally; on hind femora sparse, shorter. Claws of each

leg about one-third as long as distal tarsal segment.

Male genital segments modified, twisted strongly to

the left; ventral view of segment IX shown in fig. 31;

aedeagus and parameres as shown in figs. 32-34.

Female abdominal segments symmetrical, subgenital

plate (sternite VII) as shown in fig. 35.

Macropterous form: similar in size and structure to

brachypterous form, except for presence of claval su-

ture and well developed hind wings, and color usually

darker.

Discussion. — Hydrotrephes nieseri sp. n. has two dis-

tinct varieties, easily separable in most samples on the

basis of size and coloration of the frons, that were ini-

tially considered to be separate species. The male and

female terminalia and all other characters are the same,

however, so that these forms can be considered only as

varieties, not distinct species-group taxa. The larger

specimens with a distinctly separated dark spot on the

lower frons (fig. 27) are the typical variety, and the

smaller forms with the frons completely dark or with a

continuous dark vertical marking on the frons (fig. 28)

are given the varietal name Hydrotrephes nieseri var.

minutus. In a sample from CL 2108 these two varieties

intergrade and exhibit the complete range of coloration

and size, but in most populations the two varieties are

easily separable.

H. nieseri may be separated from its congenors by

the male genitalia (e. g. posterior margin of abdominal

segment IX straight, details of the parameres, shape of

the ‘bird head’ of the aedeagus), and from most species

by the shape of female sternite VII. The latter, howev-

er, is similar in females of H. taweli, which also have a

similar body coloration: females of these species may be

separated by the coloration of the frons. In H. nieseri

the frons is almost always completely dark between the

eyes, almost to the lower eye level (figs. 27, 28), with-

out light areas mesad of the postero-mesal eye margin,

but rarely with a small light region mesad of the antero-

mesal eye margin; in H. taweli there is almost always a

light region next to each inner eye margin (fig. 36),

rarely with the mesal dark region reaching the inner eye

margins, but always with a light area (fig. 36, arrow)

mesad of each postero-mesal eye margin.

è |

40 39

PoLHEMUS: Hydrotrephes of Sulawesi

Figures 36-43. Hydrotrephes taweli sp. n. — 36, Cephalonotum, anterior view; consistently light region, arrow; 37, Pronotal

and genal plates; 38. Ventral abdominal carinae II, IH; 39. Male abdominal segment IX, ventral view; 40, Male ventral para-

mere; 41, Male dorsal paramere; 42, Aedeagus; 43, Female abdominal sternite VII.

Etymology. — The name nieseri honours Dr. Nico

Nieser in recognition of his many contributions to the

study of aquatic Heteroptera.

Hydrotrephes taweli Polhemus sp. n.

(figs. 36-43)

Type material: Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Tengah Prov., stream 9

km E. of Taweli, NE of Palu, CL 2160, 150 m el., 10

Oct. 1985, J. T. & D. A. Polhemus (USNM). Paratypes

(nymphs not paratypes), Indonesia, Celebes: 78

brachypterous adults, 75 macropterous adults, 10

nymphs, same data as holotype (JTPC, NCTN, AMNH,

USNM, MBBJ, NHMW);18 brachypterous adults, 1

macropterous male, Sulawesi Utara Prov., Tondano

River tributary, S of Airmididi, CL 2127, 20 Sept.

1985, J. T. & D. A. Polhemus (rec); 5 brachypterous

males, 3 brachypterous females, 38 macropterous

males, 35 macropterous females, 24 nymphs, Sulawesi

Utara Prov., Gunung Klabat, Sg. Giriam, N9452, 23

June 1994, Nico Nieser (JTPC, NCTN).

Size. — Brachypterous form: length 2.52 - 2.74 mm,

width across base of cephalonotum 1.84 - 1.91 mm.

Colour. — Brachypterous form: ground color yellow-

ish brown, heavily marked with brown.

Cephalonotum with complex pattern of deep brown

markings (fig. 36), variable, but always with a light area

(arrow) mesad of each postero-mesal eye margin;

pronotum with broad irregular transverse light band

across middle. Scutellum with a basal transverse brown

stripe; basal and medial markings often coalescing to

form a large irregular triangle extending posteriorly be-

yond middle, plus additional dark markings.

Hemelytra extensively marked with large brown irreg-

ular markings, denser basally, often anastomosing, cov-

ering more than half of hemelytral area. Venter brown.

Legs, antennae yellowish, rostrum brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, densely set with small alveoli, in dor-

sal view (of entire insect) broader than long (1.84 :

1.15), lateral margins carinate behind eyes.

Mesopleural and propleural plates deeply notched (fig.

37). Eye length/width, 0.50/0.25. Interocular distance

0.83. Rostral segments 1, 2 extremely short; lengths of

rostral segments 3: 4; 0.14 : 0.47.

Scutellum shining, finely rugulose, set with tiny

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

OTE

CES

Figs. 44-51. Hydrotrephes variegatus sp. n. — 44, Cephalonotum, anterior view; 45, Pronotal and genal plates; 46, Ventral ab-

dominal carinae II, III; male right; III, female, left; 47, Male abdominal segment IX, ventral view; 48, Male ventral paramere;

49, Male dorsal paramere; 50, Aedeagus; 51, Female abdominal sternite VII.

alveoli, each with one slender setae, each surrounded

by a roughly circular transparent region; in some spec-

imens alveoli very clearly delineated due to underlying

structure or pigmentation visible through translucent

cuticle; length: width, 1.01: 0.97.

Hemelytra opaque, faintly shining, finely rugulose,

set with alveoli each bearing a slender seta; distal lock-

ing tab (pseudomembrane) on right hemelytron of

usual form. Hind wings reduced to small membranous

strips. Ventral carinae of abdominal segments II, III as

in fig. 38.

Fore and middle femora with pectinate bristles be-

neath, longer basally; on hind femora only on basal

half. Claws of each leg about one-half to one-third (on

posterior) as long as distal tarsal segment. Male genital

segments modified, twisted strongly to the left; ventral

view of segment IX shown in fig. 39; aedeagus and

parameres as shown in figs. 40-42. Female abdominal

segments symmetrical, subgenital plate (sternite VII) as

shown in fig. 43.

Macropterous form: similar in size, color and struc-

ture to brachypterous form, except for presence of

claval suture and well developed hind wings.

Discussion. — H. taweli may be separated from its

congenors by the morphology of the male and female

abdominal terminalia, and the coloration of the frons.

It is closest to H. nieseri, and the female sternite VII of

the two species are similar, but they may be separated

by the differently formed dark pattern of the frons; see

discussion under A. nieseri.

Etymology. — The name taweli is a noun in apposi-

tion, referring to the village nearest the type locality.

Hydrotrephes variegatus Polhemus sp. n.

(figs. 44-51)

Type material: Holotype, brachypterous male:

Indonesia, Celebes, Sulawesi Selatan Prov., Sungai

Pattanuang at Biseang Labboro Recreation Area, 7 km

SW of Bantimuring, CL 2165, 0-100 m el., 13 Oct.

1985, J. T. & D. A. Polhemus (usnm). Paratypes

(nymphs not paratypes), Indonesia, Celebes: 33

brachypterous adults, 20 nymphs, same data as holo-

type (JTPC, NCTN, AMNH, USNM, MBBJ, NHMW); 1

brachypterous male, 2 brachypterous females, Sulawesi

Selatan Prov., Marana River nr. Laiya, CL 2167, 0-100

m el., 14 Oct. 1985, J. T. & D. A. Polhemus GTPC).

Size. — Brachypterous form: length 2.52 - 2.63 mm,

width across base of cephalonotum 1.66 - 1.87 mm.

Colour. — Brachypterous form: ground color yellow-

ish brown to fuscous, moderately marked with brown.

Cephalonotum with complex pattern of deep brown

markings (fig. 44); pronotum largely yellowish brown,

with rather regularly spaced maculations. Base of

scutellum with a transverse brown stripe. Hemelytra,

scutellum with numerous small deep brown macula-

tions, often oval, rather regularly distributed, often

anastomosing. Venter brown. Legs, antennae yellow-

ish, rostrum yellowish to brown.

Structure. — Brachypterous form: cephalonotum

shining, convex, densely set with small alveoli, in dor-

sal view (of entire insect) broader than long (2.12 :

1.37), lateral margins carinate behind eyes.

Mesopleural and propleural plates deeply notched (fig.

45). Eye length/width, 0.58/0.29. Interocular distance

1.01. Rostral segments 1, 2 extremely short; lengths of

rostral segments 3: 4; 0.25 : 0.54.

Scutellum faintly shining, finely rugulose, set with

tiny alveoli, each with one slender setae, each sur-

rounded by a roughly circular transparent region; in

some specimens alveoli very clearly delineated due to

underlying structure or pigmentation visible through

translucent cuticle; length: width, 1.30: 1.33.

Hemelytra opaque, faintly shining, finely rugulose,

set with alveoli each bearing a slender seta; distal lock-

ing tab (pseudomembrane) on right hemelytron of

usual form. Hind wings reduced to small membranous

PoLHEmus: Hydrotrephes of Sulawesi

strips. Ventral carinae of abdominal segments II, III as

in fig. 46.

All femora with pectinate bristles beneath, longer

basally; hind femora with fewest. Claws of each leg

about one-half as long as distal tarsal segment. Male

genital segments modified, twisted strongly to the left;

ventral view of segment IX shown in fig. 47; aedeagus

and parameres as shown in figs. 48-50. Female abdom-

inal segments symmetrical, subgenital plate (sternite

VII) as shown in fig. 51.

Macropterous form: unknown.

Discussion. — The dorsal dark markings of H. varie-

gatus are much more regularly spaced than in other

species from Sulawesi, and are often absent from the

hemelytra near the base of the scutellum. The pattern

of dark markings on the cephalonotum, and male gen-

italia are diagnostic.

Etymology. — The name variegatus refers to the dor-

sal color pattern of this insect.

Hydrotrephes viriosus Polhemus sp. n.

(figs. 52-55)

Type material: Holotype, macropterous female:

Indonesia, Celebes, Sulawesi Tengah Prov., stream 10

km SE of Kamarora, Lore Lindu National Park, CL

2156, 950 mel, 8 Oct. 1985, J. T. & D. A. Polhemus

Grec; to be placed in the usNM when more material is

available).

Figs. 52-55. Hydrotrephes viriosus sp. n. — 52, Cephalonotum, anterior view; 53, Pronotal and genal plates; 54, Ventral ab-

dominal carinae II, III; 55, Female abdominal sternite VII.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Size. — Macropterous female: length 3.09 mm,

width across base of cephalonotum 2.82 mm.

Colour. — Macropterous female: ground color yel-

lowish brown to fuscous, heavily marked with brown.

Cephalonotum with complex pattern of deep brown

markings (fig. 52); pronotum largely fuscous, with

rather regularly spaced maculations covering all but a

few small irregular yellowish areas near suture. Scu-

tellum with a broad basal transverse brown stripe;

heavily marked with deep brown anastomosed mark-

ings, similar to those on pronotum. Hemelytra with

numerous small brown circular markings; almost all of

these encompass one alveolus, and most alveoli are pig-

mented, but the markings rarely anastomose. Venter

brown. Legs, antennae yellowish, rostrum yellowish to

brown.

Structure. — Macropterous form: cephalonotum

shining, convex, densely set with small alveoli, in dor-

sal view (of entire insect) broader than long (2.82:

1.44), lateral margins carinate behind eyes.

Mesopleural and propleural plates deeply notched (fig.

53). Eye length/width, 0.72/0.47. Interocular distance

1.22. Rostral segments 1, 2 extremely short; lengths of

rostral segments 3: 4; 0.29: 0.58.

Scutellum shining, not rugulose, set with tiny alve-

oli, each with one slender setae, each surrounded by a

roughly circular transparent region; in some specimens

alveoli very clearly delineated due to underlying struc-

ture or pigmentation visible through translucent cuti-

cle; length: width, 1.73: 1.73.

Hemelytra opaque, faintly shining, finely rugulose,

set with alveoli each bearing a slender seta; distal lock-

ing tab (pseudomembrane) on right hemelytron of

usual form. Hind wings well developed, reaching tips

of hemelytra. Ventral carinae of abdominal segments

II, II as in fig. 54.

Fore and middle femora with pectinate bristles be-

neath, longer basally; hind femora mostly bare. Claws

of each leg about one-third as long as distal tarsal seg-

ment. Female abdominal segments symmetrical, sub-

genital plate (sternite VII) as shown in fig. 55.

Brachypterous form: unknown.

Discussion. — It is with some reservations that a sin-

gle female is proposed as a new species, however I am

unable to place this specimen in any of the other

species before me. It is almost twice as large as H.

celebensis, which seems to be its closest congener. The

female sternite VII is different than any other, and

while the shape is superficially similar to A. celebensis,

the internal phragma is in a different location; the

mesopleural and propleural plates have a slightly differ-

ent shape; and the pattern of dark markings on the

cephalonotum and the hemelytra do not match any

other species.

The single specimen is macropterous, suggesting

that a more robust population exists somewhere in the

vicinity, and the specimen is a ‘stray.’

Etymology. — The name viriosus, L., robust, strong,

refers to the appearance of this insect.

ACKNOWLEDGEMENTS

My special thanks to Dan A. Polhemus for assistance with

field work and reviewing the manuscript. I am also indebted

to Herbert Zettel and Nico Nieser for helpful reviews and in-

formation, and to the latter for the exchange of specimens. I

wish to thank Dr. Mohammed Amir, and other personnel

from the Indonesian Institute of Sciences (LIPI), Bogor, who

provided assistance in obtaining permits for our field work in

Sulawesi.

This research was based in part on material collected dur-

ing the author's participation in Project Wallace, a joint sci-

entific undertaking sponsored by the Entomological Society

of London and the Indonesian Institute of Sciences (LIPI).

Further support was provided in part by grant 3053-85 from

the National Geographic Society, Washington, D. C., and by

grant BSR-9020442 from the National Science Foundation,

Washington, D. C. This research was carried out as an ad-

junct faculty member of Colorado State University. I thank

these organizations for their continued support of research

into the systematics and zoogeography of aquatic

Heteroptera.

REFERENCES

China, W. E., 1935. New and little-known Helotrephidae

(Hemiptera, Helotrephidae). — Annals and Magazine of

Natural History (10) 15: 593-614.

Esaki, T. & W. E. China, 1927. A new family of Heteroptera.

— Transactions of the Entomological Society of London

TER 279295,

Esaki, T. & W. E. China, 1928. A monograph of the Helo-

trephidae, subfamily Helotrephinae (Hem. Heteroptera). —

EOS, Revista Española de Entomologia 4: 129-172.

Kirkaldy, G. W., 1904. Über Notonectiden (Hemiptera). —

Wiener Entomologische Zeitung 23: 93-135.

Lundblad, O., 1933. Zur Kenntnis der aquatilen und semi-

aquatilen Hemipteren von Sumatra, Java und Bali. —

Archiv für Hydrobiologie, 1933, Supplement-Band 12,

Tropische Binnengewasser 4: 1-195, 263-498.

Miyamoto, S., 1952. Biology of Helotrephes formosanus Esaki

et Miyamoto, with descriptions of larval stages. - Sieboldia

1: 1-10, 3 pls.

Papäcek M., P. Stys & M. Tonner, 1988. A new subfamily of

Helotrephidae (Heteroptera, Nepomorpha) from South-

east Asia. — Acta Entomologica Bohemoslovaca 85: 120-

152, 1 pl.

Poisson, R., 1960. Deux Helotrephes nouveau de l'Afrique

éthiopienne (Insectes-Hétéroptères). — Revue de Zoologie

et de Botanique Africaines 61: 333-341.

Polhemus, J. T., 1990. A new tribe, a new genus and three

new species of Helotrephidae (Heteroptera) from southeast

Asia, and a world checklist. - Acta Entomologica Bohemo-

slovaca 87: 45-63.

Received: 20 November 1996

Accepted: 17 February 1997

R. PUPLESIS, A. DISKUS' & E. J. van NIEUKERKEN’

‘Department of Zoology, Vilnius Pedagogical University, Vilnius

“National Museum of Natural History, Leiden

STIGMELLA DIVINA SP. N., A REMARKABLE SPECIES

FROM TURKMENISTAN AND TURKEY

(FELD © PAER AINE PGW AE)

Puplesis, R, A. Diskus & E.J. van Nieukerken, 1997. Stigmella divina sp. n., A remarkable

species from Turkmenistan and Turkey (Lepidoptera, Nepticulidae). — Tijdschrift voor

Entomologie 140: 55-58, figs. 1-11 (1ssN 0040-7496]. Published 31 October 1997.

Stigmella divina sp. n. is described from the Kopet Dag mountains in Turkmenistan and

Central Anatolia (Turkey). The species cannot be placed in any known species group, because

of its remarkable genitalia. The male possesses a unique androconial patch on the forewing un-

derside.

Correspondence: R. Puplesis, Department of Zoology, Vilnius Pedagogical University,

Studentu 39, Vilnius 2034, Lithuania, e-mail: [email protected].

Key words. — Nepticulidae; Palaearctic; Central Asia; Stigmella.

The Nepticulidae of Turkmenistan were recently

revised, together with those from other parts of

Central Asia (Puplesis 1994). Since then a few more

species have been described, and a checklist was pub-

lished (Puplesis & Diskus 1995, 1996a, 1996b,

1996c, Puplesis et al 1996). Some of the species col-

lected by the first two authors during the 1993 expe-

dition to the Kopet Dag ridge remain to be described.

One of these is a peculiar species of Stigmella, which

does not fit in any of the recognised species-groups.

The same species was encountered by the third au-

thor amongst unidentified material from Turkey.

The nepticulid fauna of Turkey is still imperfectly

known, there is as yet no general review available, but

a few species were mentioned in several taxonomic

papers (Klimesch 1978, van Nieukerken 1985, 1990,

van Nieukerken & Puplesis 1991).

We describe this species here to point to its unique

set of characters, not fitting into any known species

group, awaiting further revisionary work on western

Palaearctic Stigmella.

Type material is deposited in the collection of the

Department of Zoology of Vilnius Pedagogical

University, Lithuania (VPU), the National Museum of

Natural History, Leiden (RMNH) and the Zoological

Museum, Copenhagen (ZMUC). Terminology of ex-

ternal features and genitalia follows van Nieukerken

et al. (1990) and Puplesis (1994). Genitalia structures

of the Turkmenian specimens were examined and fig-

ured in glycerine.

Stigmella divina sp. n.

(figs. 1-11)

Type material. — Holotype d : Turkmenistan, wes-

tern Kopet Dag, 40 km E Kara Kala (= Garrygala),

800 m, 7.viii.1993, leg. R. Puplesis & A. Diskus

(vpu). Paratypes: 9d, 159 same data as holotype,

25.vi-18.viii.1993 (vru, RMNH); 1 d Turkey, Sivas,

10 km W Giiriin, 1650 m, 27.vii.1989, leg. Fibiger

& Esser, Genitalia slide EJVN 3180 (zmuc).

Diagnosis. — Females can be recognised by the

combination of a yellowish cream thorax and fore-

wing base and the wide fascia. Males, which have a

dark thorax, can easily be recognised by the elongate

yellow androconial patch on the forewing underside.

The male genitalia are characterised by the shape of

the valva with the bifid apex and the dagger shaped

juxta.

Description

Male (fig. 1). — Forewing length 2.3-2.7 mm.

Head: frontal tuft from ferruginous to brown, occa-

sionally fuscous; scape and collar cream; antenna

cream-grey to grey-brown, with 30-31 segments.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 1. Stigmella divina, male Paratype, Turkey. Left: dorsal view; right: underside of left wings, showing androconial patch.

Painted by Roland Johansson.

Thorax, tegulae and forewing grey-brown, with some

purplish reflection; area posterior to fascia with inten-

sive reflections, darker. Fascia creamy white, rather

wide, but slightly varying in shape. Underside of

forewing with large cream to yellow elongate spot of

androconial scales, further dark grey. Cilia pale grey

to grey-cream, lustre. Hindwing totally or partly cov-

ered by pale cream scales, underside grey. Abdomen

fuscous, valval lobes and anal tufts brown and lustre.

Female. — Forewing length approximately 2.2-2.4

mm. Head: frontal tuft pale ferruginous to pale

brown; scape and collar cream; antenna grey-brown

to brownish-cream, with 27-28 segments. Thorax,

tegulae and small area of forewing base yellowish

cream. Hindwing grey. Further as male, except an-

droconial scales.

Male genitalia (figs. 2-5, 7-10). — Capsule 285-

315 um long. Vinculum with shallow anterior exca-

vation. Tegumen narrow. Uncus relatively large,

trapezoid, slightly bilobed, surface somewhat papil-

late dorsally. Gnathos with long posterior processes

and short and broad anterior processes. Valva 180-

185 um long, slender triangular, basally joined with

juxta, at apex curved medially (inwards), ending in

two distinct teeth-like processes. Transtilla with long

sublateral processes; transverse bar narrowed in mid-

dle. Aedeagus (figs. 3, 4, 5, 9, 10) 275-310 um long,

basally widened; vesica with several large cornuti,

varying in shape and size; basally or medially with a

group of many spinelike cornuti. Manica absent.

Juxta between valvae, joined to valvae and aedeagus,

apex bifid, resembling the juxta in Stigmella viscerella

(Stainton) (Johansson & Nielsen 1990: figs. 247-

249). Distally the scerite with well sclerotised, slight-

ly bifurcated process.

Female genitalia (fig. 6). — Apophyses long, poste-

riores slightly wider than usually. Accessory sac small,

but distinct. Ductus spermathecae slightly longer

than corpus bursae, but very slender, with some nar-

row convolutions. Corpus bursae relatively small and

pectinations on bursae obscure.

Biology

Host-plant unknown. Adults fly from late June to

August.

Distribution (fig. 11)

Turkmenistan (western part of Kopet Dag) and

Turkey (Central Anatolia, Sivas province).

Figs. 2, 3. Male genitalia of Stigmella divina, sp.n., holotype

(western Turkmenistan). - 2, Capsule; 3, Aedeagus. Scale

0.1 mm.

PUPLESIS, DISKUS & NIEUKERKEN: Stigmella divina sp. n.

Ù \ er 5

\ =

Figs 4, 5. Aedeagus of Stigmella divina, sp.n., paratype, Fig. 6. Female genitalia of Srigmella divina, sp.n., paratype

(western Turkmenistan). — 4, Ventral view; 5, Lateral view (western Turkmenistan). Scale 0.1 mm.

(same specimen, figured in glycerine). Scale 0.1 mm.

Figs 7-10. Male genitalia of Stigmella divina, sp. n., paratype (Turkey), slide Evy 3180.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

f \ 7 Fier

) RR U REN È

(SE EEEN ay Distribution of Stigmella

N Cs l | a divina.

SIE 85

? i Fr bp

< IN

SA N \ =

IA ne NC

II SZ zes WA EN

da œ PR } D EN

Ur, Ne SL

IE Ÿ

wae

2 He!

= s o

se in \

[LES Sr —c We

i |

| )

2 ECN à

Discussion

We have not been able to determine the position of

Stigmella divina in any of the currently recognised

species groups. It resembles somewhat species in the

S. ulmivora group, but particularly the valvae and an-

droconial scales are very different. Without a phylo-

genetic framework for Stigmella, we prefer to leave S.

divina tentatively unplaced within the genus.

ACKNOWLEDGEMENTS

We are particularly indebted to Roland Johansson,

for preparing the water-colours and allowing us to use

them. We thank Ole Karsholt for lending us the ma-

terial collected by Fibiger & Esser. The research pre-

sented in this publication was made possible in part

by Grant LAO000&LHXxX100 from the International

Science Foundation.

REFERENCES

Johansson, R. & E.S. Nielsen, 1990. Tribus Nepticulini. —

In Johansson, R. et al.: The Nepticulidae and Oposte-

gidae (Lepidoptera) of North West Europe. — Fauna

Entomologica Scandinavica 23: 239-321.

Klimesch, J., 1978. Beitrag zur Kenntnis der Nepticuliden-

fauna von Anatolien und der Insel Rhodos (Lepidoptera,

Nepticulidae). — Tijdschrift voor Entomologie 121: 239-

278.

Nieukerken, E. J. van, 1985. A taxonomic revision of the

Western Palaearctic species of the subgenera Zimmer

mannia Hering and Ectoedemia Busck s. str. ( Lepidopte-

ra, Nepticulidae), with notes on their phylogeny. —

Tijdschrift voor Entomologie 128: 1-164.

Nieukerken, E. J. van, 1990. The Trifurcula subnitidella

group (Lepidoptera: Nepticulidae): taxonomy, distribu-

tion and biology. — Tijdschrift voor Entomologie 133:

205-238.

Nieukerken, E.J. van, E.S. Nielsen, R. Johansson & B.

Gustafsson, 1990. Introduction to the Nepticulidae. — In

Johansson, R. et al. The Nepticulidae and Opostegidae

(Lepidoptera) of North West Europe. — Fauna Entomo-

logica Scandinavica 23:11-109.

Nieukerken, E. J. van & R. Puplesis, 1991. Taxonomy and

distribution of the Trifurcula (Glaucolepis) raikhonae

group (Lepidoptera: Nepticulidae). — Tijdschrift voor

Entomologie 134: 201-210.

Puplesis, R., 1994. The Nepticulidae of Fast Europe and

Asia. — Leiden, Backhuys Publishers. 201 pp + 840 figs.

Puplesis, R. & A. Diskus, 1995. Acalyptris argyraspis sp. n., a

remarkable species from Tadzhikistan (Lepidoptera:

Nepticulidae). — Phegea 23: 51-54.

Puplesis, R. & A. Diskus, 1996a. First record of the genus

Etainia Beirne from Central Asia with description of two

new species and some provisional notes on the world fau-

na (Lepidoptera: Nepticulidae). — Phegea 24: 41-48.

Puplesis, R. & A. Diskus, 1996b. A review of the Stigmella

sorbi species group with descriptions of two new species

from Turkmenistan and Tadzhikistan (Lepidoptera:

Nepticulidae). — Phegea 24: 171-182.

Puplesis, R. & A. Diskus, 1996c. Five new mining

Lepidoptera (Nepticulidae, Bucculatricidae) from Central

Asia. — Tijdschrift voor Entomologie 139: 181-190.

Puplesis, R., A. Diskus, R. Noreika, & N. Saparmamedova,

1996. Revised check-list of mining Lepidoptera (Nep-

ticuloidea, Tischerioidea and Gracillarioidea) from Central

Asia. — Tijdschrift voor Entomologie 139: 191-200.

Received:1 May 1997

Accepted: 30 June 1997

Opetre ROHFRITSCH

Institut de Biologie moleculaires des Plantes, C.N.R.S., Strasbourg

MORPHOLOGICAL AND BEHAVIOURAL

ADAPTATIONS OF TEE GATEMIDGE/ZASIOPTERA

ARUNDINIS (SCHINER) (DIPTERA, CECIDOMYIIDAE)

NOIGOLEE Gi-AND TRANSPORT CONIDIATOF Tis

FUNGAL SYMBIONT

Rohfritsch, O., 1997. Morphological and behavioural adaptations of the gall midge Lastoptera

arundinis (Schiner) (Diptera, Cecidomyiidae) to collect and transport conidia of its fungal sym-

biont. — Tijdschrift voor Entomologie 140: 59-66, figs. 1-13. [issN 0040-7496]. Published 31

October 1997.

The gall midge Lasioptera arundinis attacks lateral shoots of the stems of the reed (Phragmites aus-

tralis Trin.) with the help of its fungal symbiont (of the genus Macrophoma). Together with the

fungus, the larvae move up the shoot and penetrate into the stem. Thus allowing both organisms

access to vascular tissues of the host. The larvae feed on the host tissue and on the fungus and the

overwintering larvae are not harmed by the fungal mycelium. This narrow relationship between a

gall midge and fungus is an example of obligate mutualism. The female imago and the first-larval

instar have evolved specific behavioral traits and stuctural adaptations to disseminate the fungus.

The fungal conidia are collected on upper internodes of galled shoots by the female just before

oviposition. The female transports the conidia to an oviposition site in specialized structures on

the cercus (tip of the ovipositor). These structures, called mycangia, are composed of a sclerotized

plate covered with large spoon-like spines which collect the conidia of the right size and shape.

The conidia slide down the collecting spines and enter laterally situated pouches beneath the

spines. The mycangia join the extremity of the oviduct such that eggs and conidia are deposited

together. The first larval stage with its long bristles and numerous spines carries the fungal mate-

rial and disseminates it along the feeding route.

O. Rohfritsch, I. B. M. P., 12, rue du Général Zimmer, 67084 Strasbourg cedex, France.

Key words. — Lasioptera arundinis, Phragmites australis, fungus, mycangia, gall, symbiosis.

There are approximately 4300 described species

within the family Cecidomyiidae (Diptera:

Nematocera); about half are gall inducers (Felt 1940)

and the others are detritus feeders. The ancestral feed-

ing biology of gall midges was thought to be myce-

tophagy (Mamaev 1968, Gagné 1986, Roskam 1992)

whereas the derived taxa belonging to the tribes

Cecidomyiinae and Oligotrophini induce galls, as do

most of the Lasiopterini and all of the Asphondyliini.

Many of the Asphondyliini and Lasiopterini exhibit a

peculiar feature that is reminiscent of the ancestral

mycetophagous condition -the surface of their gall

chamber is covered with a layer of fungal hyphae on

which the larvae feed. These are ambrosia gall midges

which are analogous to ambrosia beetles (Neger

1913) that also feed on symbiotic fungi.

Gall midges of the genus Lasioptera which often oc-

cur on stems of reeds and grasses (Gagné 1989) are

usually associated with a fungal symbiont. Meyer

(1952) observed that the gall of Z.rubi (Schrank) on

Rubus caesius L. lacked the characteristic layer of nu-

tritive cells found in most galls, and instead a mycelli-

um was present along the surface of the larval cham-

ber with intracellular haustories within tissues of the

gall. In contrast, the larvae of some Lasioptera, such as

L. berlesiana Paoli, stay in preformed galleries and

feed on a fungal associate that grows in tunnels which

begin as tephritid oviposition holes on olives (Solinas

1967). L. donacis (Coutin and Faivre-Amiot 1981)

feeds on the fungus Aspergillus niger V. Thieg. grow-

ing in old galleries of a chloropid fly in leaf sheaths of

Arundo donax L.(Coutin and Faivre-Amiot 1981).

Hermann et al. (1993) described the association of L.

ephedricola Cockerell with the fungus Aureobasidium

pullulans (de Bary) Arnaud which forms a dark ring

around gall chambers on Ephedra trifurca Torr.

Skuhravä and Skuhravy (1981, 1992) described the

life cycle of L. arundinis Shinner and L. hungarica

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Möhn which induce galls within stems of the com-

mon reed Phragmites australis Trin. in central Europe

and found that galls of both contain a fungus consid-

ered to be of the genus Sporothrix. Developmental

morphology of the gall of L.arundinis has also been

described (Rohfritsch 1992) and at the end of this

study, it was suggested that the fungus has a narrow

and obligatory relationship with the inducer. It was

suggested that without the larvae, the fungus would

be unable to attack the stem of reeds and that the lar-

vae need the fungus to penetrate to the stem medulla

and to obtain food. It is known that the first-instar

larvae of L.arundinis carry the conidia upwards with-

in the stem and that all nearby larvae follow the same

entrance route. However, the question remains as to

how the first-instar larvae come in contact with the

fungus.

There have been several suggestions as to the mech-

anism by which fungi are brought into the galls in-

cluding non-specific air borne inoculation (Batra and

Lichtwardt 1963), contamination of the eggs by adult

feces (Haridass 1987), and the specific transfer of fun-

gal spores by the ovipositor or abdominal mycangia

(Bissett and Borkent 1985). Mycangia are pockets on

the bodies of insects that selectively collect and trans-

port fungi (Batra and Lichtwardt 1963). According to

Borkent and Bissett (1985), the conidia carried by

Lasiopterini are entrapped by two dorso-lateral

groups of strong setae on uromerVIII. These authors

also observed conidia among the setae of the cercus;

however, in another study, Tastäs-Duque and Sylvén

(1989) were not able to find conidia on the setae of

uromerVIII of L. rubi. Hermann et al.(1993) found

that adults of L. ephedricola have structures on their

ovipositors which could serve as mycangia, but were

unable to find fungal propagules on the ovipositors of

newly emerged females. Hermann et al. (1993) con-

cluded that host leaves were the source of fungi for ei-

ther newly hatched larvae, or females prior to oviposi-

tion, or that oviposition occurs on or near an

inoculum source.

The purpose of the present paper is to demonstrate

that: 1) females emerging from their pupae do not

ROHFRITSCH: Mycangia of Lasioptera arundinis

carry the fungus, 2) females collect conidia on the up-

per internodes of attacked shoots, 3) mycangia are

present on the cercus which selectively collect conidia

from the epidermis of the leaf sheath, 4) conidia and

eggs are deposited simultaneously, 5) larvae dissemi-

nate the fungus along the dispersal route and aid its

progress into the stem.

MATERIAL AND METHODS

Insect rearing

Lateral shoots of common reed attacked by L.

ae 2

arundinis were collected in January from the ‘Parc de

Pourtales’ near Strasbourg, France. The whole shoots,

about 25cm long, were kept at room temperature

(22°) in a glass jar covered with muslin under natural

daylight.

Preparation for light microscopy

Non-fixed material. — The fungal flora present on

the leaf sheaths was observed in one of the staining so-

lutions described below either separately from the

plant or on a peel of the epidermis. The appendages

of the ovipositor and the cercus were observed in a

similar way. Stains used were: (1) coton blue which

selectively stains the fungal mycelium, (2) lactic

reagent (Gazet du Chatelier 1948, Rohfritsch

1992b), which makes the tissue partially transparent

and stains starch, oils, cutin and lignin, (3) toluidine

blue which stains nucleus, cytoplasma and cell walls.

Fixed material. — Fixative: Formaldehyde-Alcohol-

Acetic acid (ethanol 70% 90 ml, glacial acetic acid

5ml, formaldehyde 5ml). Portions of the internodes

(about 3cm long) were fixed over a period of 24h at

room temperature. Smaller pieces of the fixed mater-

ial (1cm) were embedded in paraffin and sectioned in

3-5 um slices with a microtome (Jung, Heidelberg)

and transferred to glass slides. After paraffın removal,

the sections were stained with toluidine blue or with

safranine-light green.

Dissections were photographed with a Zeiss

Tessovar while histological sections were pho-

tographed with a Leitz microscope.

Figs. 1-7: Lateral shoots of common reed attacked by Lasioptera arundinis and its associated fungus. Bar scale: 0.5 cm for Figs. 1-

4. 100 pm for figs. 5-7.— 1, Attacked shoot with mature larvae, basal internodes. The fungal mycelium has plugged the entrance

le)

split; 2, Upper internode of the attacked shoot. The fungus has grown through the leaf sheath and conidia are present on the epi-

dermis; 3, Dissection of an internode filled with fungus, which contains numerous larval chambers (arrows); 4, Dissection of an

upper internode. The fungus (arrow) is only present in the stem cortex, without the larva it does not reach the medulla (M). —

Figs. 5-7: Cross sections of galled stems; 5, Section through the cortical tissue of a young reed stem. Both larvae and fungus have

penetrated the medulla. The entrance cavity is covered by the fungal mycelium. The vascular bundles (vb) are accessilble to the

feeding larva; 6, Section through a maturing gall. All stem cells are hypertrophied. The entrance split is filled with a mat of black

mycelium (arrow). The fungus has attacked the most inward vascular bundles (vb). The larva is feeding on young fungal hyphae

growing on the plant tissue; 6a, Detail of fig.6 showing the larval feeding site; 7, Section through a mature gall. The larva has

stopped feeding, the black fungal mycelium fills the whole medullar cavity.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

{le

RESULTS AND DISCUSSION

Biology of L.arundinis and gall development

Larvae of L.arundinis and its associated fungus pro-

duce a thickening and shortening of lateral shoots of

common reed. Galls are made from hypertrophied

stem tissue and hypertrophied medulla (figs. 1 and 3).

There are 1 to 25 larvae per internode, with most lar-

vae found in the central part of the shoot. Each inter-

node constitutes one larval chamber in young galls.

Later, the proliferating fungus isolates each larva in an

individual chamber, so that the mature gall appears as

multi-chambered. The uppermost internodes are free

oflarvae and fungi but between galled and normal tis-

sues | or 2 internodes are superficially attacked by the

fungus (figs. 2, 4). Overwintering larvae use their

spatula to bore an exit hole in mid-May. Adults exit

the gall at the end of June. Only lateral shoots dam-

aged at their tops by other invertebrates are attacked.

Each female lays from 60 to 110 eggs under the basal

leaf sheath (Skuhrava & Skuhravy 1981). All freshly

hatched larvae move along the same route towards the

apex. The larvae carry conidia and disseminate the

fungus along their route. In addition, the larvae at-

tack the plant tissue with their mandibles, creating a

stress and a discrete wounding on the plant epider-

mis. The fungus is now able to penetrate into the epi-

dermis and invade the cortical tissue of the stem

(Rohfritsch 1992). With the help of the fungus, the

larvae enter the stem and move upwards; the fungus

produces a longitudinal gallery in the cortical tissue

which allows the larvae to progress through the nodes

of the stem upwards. The fungus produces cell disso-

ciation and the larval activity enlarges the cavity (fig.

5). Both organisms enhance proteosynthesis of ad-

joining cells, with some cells developing the cytologi-

cal characteristics of nutritive cells. When the third-

instar larvae reach the medulla, the fungus invades the

medullar parenchyma and the innermost vascular

bundles (figs.6, 6a). The mycelium soon covers the

larval cavity and fills the medulla once the larvae stop

feeding (figs. 3, 7) (Rohfritsch 1992).

ROHFRITSCH: Mycangia of Lasioptera aründinis

Emergence of the gall midge and conidia collection

Attacked shoots were collected at the end of

January and observed regularly each week. Within 2

weeks, the larvae bored exit holes through the fungal

mycelium and the leaf sheaths, except the outermost

sheath. Adults exited the galls 3 weeks later. Males

emerged first and newly emerged females were free of

fungal material. The ovipositor was retracted after

mating, then the females flew about and finally

alighted on the leaf sheath of an upper shoot intern-

ode. The ovipositor was extruded and the surface of

the leaf sheath was palpated. The visited internode

was not galled but the fungus was present along a nar-

row strip in the stem cortical tissue. From this super-

ficially located infection, the fungus had grown

through the leaf sheath and produced mycelium and

conidia on the epidermis (figs. 2, 11). As was general-

ly observed with ambrosia gall midges (Borkent &

Bissett 1988), we never observed L. arundinis picking

up fungal material directly from the gall surface. The

observed Lasioptera gall midge found the fungal coni-

dia on the galled host plant but not on the gall direct-

ly. It is not known how the females distinguish the 5

or more different fungi present on the surface of the

same leaf sheath.

Mycangia

Borkent & Bissett (1988) reported that female

Lasioptera species use specialized structures located on

uromerVIII of the female abdomen to carry conidia

and transfer them to the host plant during oviposi-

tion. Like Tastas-Duque & Sylvén (1989), I was un-

able to find conidia on uromer VIII, even after the fe-

male had visited the upper internodes of the galled

shoot (fig. 9). However by examining the whole

ovipositor in lactic reagent, I found distinct mycangia

on the cercus near the tip of the ovipositor, close to

the outlet of the oviduct (figs. 8, 10, 12). The paired

mycangia were made up of pockets located beneath

the sclerotized plate which carried large spoon-like

spines. These spines have been previously observed

Figs. 8-13: Structural adaptations of Lasioptera arundinis for carrying fungal symbiont. Bar scale : SOum in figs. 8 and 9,10um

in all other figures. — 8: Posterior part of the ovipositor with the mycangial structures on uromer X. The superior lamella of

uromer X is covered with spines(sp) and spoon-like sensilla (ss). The apical portion of the superior lamella is covered with sen-

sory hairs (sh). Arrow indicates the entrance into the mycangia. Asterisk indicates the outlet which joins the oviduct. The ovi-

positor was processed in lactic reagent, and photographed in ethanol; 9, Portion of ovipositor showing spines (sp) on uromer

VIII; 10, Mycangial pouch on uromer X. The conidia are collected by the spoon-like spines on the sclerotized plate. The coni-

dia glide down the spines and enter the pouch (the entrance is indicated by an arrow). During oviposition, the conidia leave the

pouch via the outlet (asterisk) which joins the oviduct (ov). Stain: lactic reagent. Insert: Detail of the conidia present in the my-

cangial pouch; 11, Conidia observed on the leaf sheath of the upper internode of the attacked shoot. This is the same intern-

ode shown in fig.2. Stain: lactic reagent; 12, Same mycangia as in fig.10, mycangia were observed in ethanol after treatment

with lactic reagent, (the conidia have been removed) ss: spoon-like spines, ov: oviduct. Entrance into the mycangia is indicated

by an arrow; outlet indicated by asterisk; 13, Section of first- instar larva (L) showing long bristles and numerous spines.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

on other Lasioptera species (Skuhravá & Skuhravy

1981, Tastäs-Duque & Sylvén 1989); however, the

pockets just beneath the spines have not been seen. I

found a few conidia on the spoon-like spines and at

least 30 conidia of the same type in each pocket be-

neath the spines. The shape and the size of conidia

corresponded with the shape of the spines. The spines

evidently pick-up the conidia when the ovipositor lat-

erally palpates the surface of the leaf sheath, retaining

only conidia of the right size and shape. The conidia

then slip to the base of the spine and, with the help of

other spines, are guided into the cavity opening at the

top of the two laterally situated pouches, beneath the

sclerotized plate (fig. 10). Five females were examined

and only one type of conidia was present in their my-

cangia. Small fragments of mycelium were sometimes

present. An exit hole at the bottom of the mycangia

joins the extremity of the oviduct (figs. 10, 12) such

that during oviposition both eggs and conidia are de-

posited.

The mycangia of L. rubi are similar to the struc-

tures described above. Because the female could not

find its fungal symbiont on the gall surface or along

nearby plant surfaces, the mycangia were empty (un-

published results).

Fungus

The fungus previously associated with this gall was

described as belonging to the genus Sporothrix of the

Deuteromycetes (Skuhravá & Skuhravy 1981); how-

ever, the size and shape of the conidia are similar to

those of the genus Macrophoma (Borkent & Bissett

1988). Borkent & Bissett suggested that the fungal

genus Macrophoma was specifically associated with

the ambrosia gall midges. The conidia carried by

L.arundinis had a distinct terminal structure, a sort of

dense rim at their top (fig. 11). Otherwise, the coni-

dia shared the characteristic features of Macrophoma

conidia, as described in Borkent & Bissett (1988)

with similar shape, a truncated base, indicative of

holoblastic ontogeny, they were first hyaline and

aseptated, and at later stages developed brownish pig-

ments and one septa. According to Borkent & Bissett

these ambrosia fungi may be referred to the

coelomycetes anamorphous genera and all the

anamorphous observed in the gall midge mycangia

could be considered congeneric.

The nutrition of the fungus was biotrophic and it

was feeding inter- and intracellularly without ever

killing the cells. The fungus had long slender

‘prospektiv’ hyphae which dispersed intercellularly,

and were directed straight to the vascular bundles.

These hyphae produced cell wall dissociation. Along

this open route and especially near the larvae, large

cytoplasma-enriched hyphae developed intercellular-

ly with haustoria inside the cells. Near the end of lar-

val development, the old, highly chitinized mycelium

formed a dense mat along the medullar cavity and,

once the larvae stopped feeding, they filled the cavity.

This mycelium was unable to fruit on the gall surface.

Along the entrance canal, the plant reacted by lignify-

ing its cell walls such that the fungus was stopped in

its lateral progression.

Thus the fungus can only develop towards the cen-

ter of the stem. From the medulla the fungus attacked

only the innermost vascular bundles and fructifica-

tion did not occur. No larvae were observed along the

youngest attacked internodes; the fungus invaded

only the cortical parenchyma and produce a superfi-

cial gallery along the internode. From here the fungus

was able to produce fructifications by growing

through the leaf sheath (figs. 2,4, 11).

Fungal dispersion via the larva

First-instar larvae have unusually long bristles and

many spines (fig. 13), between which fungal material

was found. It is via these bristles and spines that the

fungus comes in contact with the young epidermis of

the host. Larval activity also results in cell wounding

(Rohfritsch & Shorthouse 1982, Rohfritsch 1992)

which allows the fungus to invade the reed stem.

Larvae later help the fungus to reach the different lay-

ers of vascular bundles and finally the medulla. The

fungus is apparently unable to join the medulla with-

out the larvae and can not progress up the stem (fig.

4).

Galls of L. arundinis on the stems of reed appear to

be ambrosia galls. As in the case of ambrosia beetles,

the gall midge collects and carries its symbiont in spe-

cialized structures: the mycangia. This is the first re-

ported observation of fungal collecting and transport-

ing behaviour by gall midges and also the first

description of mycangial pouches in the genus

Lasioptera. These fungus collecting structures are lo-

calized on the cercus. They are composed of two lat-

eral pouches, associated with spoon-like setae. These

large hooked setae have also been observed on L. hun-

garica (Skuhrava & Skuhravy 1981) and L. rubi

(Tastás-Duque & Sylvén 1988) where they were

thought to serve as sensilla which register chemical

stimuli both by olfaction and contact. My observa-

tions have shown that the sensilla are able to discrim-

inate the fungal conidia not only by olfaction but also

by their shape and size and thus they may help the

midge to find its symbiont on the leaf sheath of the

right internode. It is possible that chemical signals are

involved in recognizing the correct fungal symbiont.

Thus it is incorrect to conclude that the ovipositor

picks up fungal conidia similar to the way in which a

finger picks up objects when it is run across a surface.

The gall of L. arundinis is therefore a model of mu-

tualistic association between a gall midge and its fun-

gal symbiont where both organisms feed and develop

on the stems of reed. Together they are able to enter

into the stem tissues and feed on vascular tissues with

the midge larvae also feeding on the fungus; they first

feed on young, tender hyphae budding in the larval

chamber and later on the older mycelium (Rohfritsch

1992). The larvae do not only use the mycelium to

enter the reed stem and locate food, but to attain pro-

tection as well. In this association, the fungus is dis-

persed into a highly specialized and protected niche.

With the help of gall-midge larvae, it gains access to

young tissues of the elongating shoot.

It also is apparent that coevolution between gall

midges and fungi has taken place as evidenced by

physical traits such as the mycangia of the imago and

the long bristles and spines of the larvae. There are

also behavioural traits of the insect for collecting and

carrying specific fungal spores and to inoculate them

in a specific manner in the plant tissue along a line

which is the larval dispersal route. Success of this

plant-fungus-larva interaction is mainly dependent

on larval behavioral traits. The fungus is controlled by

plant defense reactions and by the larvae and, the in-

sect appears to control physiological processes and

morphogenetic expression of the fungus including

conidia germination, stimulation of mycelial growth

and control over this growth.

Ambrosia gall midges have evidently evolved the

means of using and manipulating the fungus, and the

fungus may have reciprocated by evolving similarly

accommodating traits such as fructifications on upper

internodes of attacked shoots, along with a particular

size and structure of conidia. The fungus in turn

shifted from a saprophytic to a biotrophic mode of

life and has avoided plant defense reactions. The fun-

gus now remains under the control of the gall midge

to avoid competition for food and space and the pro-

liferating dense mycelium never destroys the overwin-

tering larvae.

Thus ambrosia galls such as this result from the ac-

tivity of both the insect and the fungus. Cell wall

maceration is produced by the fungus but cell activa-

tion to high proteosynthesis which characterize nutri-

tive tissues of most gall-inducing insects can also be

induced by biotrophic fungus. Furthermore, the hy-

pertrophic gall growth may result from the activity of

both organisms. The fungus does not reach the

medulla in the upper internodes where it develops

without the larva; it does not develop a thick myceli-

um and has less influence on growth of the internode.

Consequently, both organisms stay in a narrow mu-

tualistic relationship. The fungus is collected, trans-

ported and deposited via highly evolved mycangia.

According to Bissett & Borkent (1988), all Zasioptera

carry the same kind of conidia, all from the genus

ROHFRITSCH: Mycangia of Lasioptera arundinis

Macrophoma. It thus can be speculated that all

Lasioptera female imagoes have mycangia beneath the

spoon-like spines on their cerci. To observe the coni-

dia in the mycangia, it is necessary to give the midge

the opportunity to collect them and it is essential to

avoid ethanol for material preservation or observa-

tion.

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Received: 2 September 1996

Accepted: 3 March 1997

Louis M. ROTH

Museum of Comparative Zoology, Harvard University, Cambridge, U. S. A.

THE COCKROACH GENERA PSEUDOTHYRSOCERA

SHELFORD, HAPLOSYMPLOCE HANITSCH,

AND EPISYMPLOCE BEY-BIENKO

(BEAT TARA) BEAIRELLIDAE BEAT TELLINAE)

Roth, L.M. 1997. The cockroach genera Pseudothyrsocera Shelford, Haplosymploce Hanitsch, and

Episymploce Bey-Bienko (Blattaria: Blattellidae, Blattellinae). — Tijdschrift voor Entomologie 140:

67-110, figs. 1-123 [rssn 0040-7496]. Published 31 October 1997.

Twelve species of Pseudothyrsocera are redescribed and one new species, P. perkinsi, is described. A

key is given to identify the males. The genus is divided into two species groups. Leptothyrsocera is

a junior synonym of Pseudothyrsocera. Ischnoptera sinensis Walker (= Episymploce sinensis) is trans-

ferred to Pseudothyrsocera.

Eight species of Haplosymploce are redescribed, and a key to males is given. New synonyms are:

Symploce bicolorata Roth is Haplosymploce montis (Shelford), and Symploce ferruginea Roth is

Haplosymploce nigra (Hanitsch). Mopsera andamanica Princis, Pseudomops pica Walker, Pseudo-

thyrsocera ruficollis Shelford, and P. moultoni Hanitsch are transferred to Haplosymploce. Haplo-

symploce guttifera (Walker) is Beybienkoa guttifera [= its junior synonym Beybienkoa papuensis

(Roth)], and Haplosymploce walkeri Princis is Hemithyrsocera walkeri.

Two new species of Episymploce, E. simmonsi n. sp. and E. parafissa n. sp., are described and sev-

eral species of Symploce are transferred to that genus. Several new records of Episymploce are given.

Lectotypes are designated for the following species: Pseudothyrsocera montana Shelford,

Pseudomops pica Walker, Thyrsocera circumcincta Stal, T. lugubris Stal, T. rufiventris Stal, T. semi-

cincta Stal, Ischnoptera montis Shelford, and /. nigra Hanitsch.

Dr. L. M. Roth, 81 Brush Hill Road, P.O. Box 540, Sherborn, Ma 01770, U.S.A.

Key words. — Pseudothyrsocera; Haplosymploce, Episymploce, Blattellidae; cockroaches; taxonomy;

redescriptions; new species; new combinations; new records.

This work is a continuation of my studies of cock-

roaches from the Indo-Australian regions. Two gen-

era, namely, Pseudothyrsocera Shelford and

Haplosymploce Hanitsch, are revised. A third genus,

Episymploce Bey-Bienko, is included because some of

its diagnostic characters are similar to these two gen-

era.

In this paper I redescribe 12 species of

Pseudothyrsocera, describe one new species, and pro-

vide a key to distinguish the males. Leptothyrsocera

Princis is a synonym of Pseudothyrsocera. I have divid-

ed Pseudothyrsocera into two species groups, one of

them having characters annectant to Episymploce.

I recognize ten species (one with a query) of

Haplosymploce and redescribe eight of them;

Pseudothyrsocera andamanica (Princis) belongs in

Haplosymploce. Symploce bicolorata Roth is

Haplosymploce montis (Shelford), and Symploce ferrug-

inea Roth is Haplosymploce nigra (Hanitsch).

Two new species of Episymploce are described, and

several species of Symploce are transferred to that

genus.

The following museums and their curators or col-

lection managers loaned me specimens: ANSP -

Academy of Natural Sciences of Philadelphia, PA,

U.S.A; Mr. Donald Azuma; BPBM - Bernice P.

Bishop Museum, Honolulu, Hawaii, U.S.A.; HECO -

Hope Entomological Collections, Oxford University,

England; Dr. George C. McGavin and Mr. I

Lansbury; Mczc - Museum of Comparative Zoology,

Harvard University, Cambridge, mA, U.S.A.; MNHG -

Museum für Naturkunde, Humboldt-Universitàt zu

Berlin, Germany; Dr. Kurt Günther; NMWA -

Naturhistoriches Museum Wien, Vienna, Austria;

Dr. U. Aspöck & Dr. A.P. Kaltenbach; Nrss -

Naturhistoriska Riksmuseet, Stockholm, Sweden; Per

Inge Persson; RMNH - National Museum of Natural

History (Rijksmuseum van Natuurlijke Historie),

Leiden, The Netherlands; Mr. J. van Tol; zis -

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Zoological Institute, Lund, Sweden; Dr. Roy

Danielsson; zman - Zoologisch Museum, Univer-

siteit van Amsterdam, The Netherlands; Mr. Willem

Hogenes; zmuc - Zoological Museum, Copenhagen,

Denmark; Soren Langemark.

SYSTEMATIC PART

Key to genera Pseudothyrsocera, Haplosymploce,

and Episymploce

I Radial vein of hind wing simple (figs. 8, 10) or

sometimes with a pseudoposterior branch (fig.

41). Intercercal processes present (fig. 82) or

absent (fig.3)

— Radial vein of hind wing with a distinct poste-

rior branch (fig. 112). Intercercal processes ab-

ont (i WG). TA II Episymploce

DI Hind wing cubitus vein with one or two com-

plete and usually no (rarely 1) incomplete

branches (figs. 8, 10, 16. Intercercal processes

absent (figs LS AR Pseudothyrsocera

= Hind wing cubitus vein with four to seven

complete and two to four incomplete branches

(fig. 89). Intercercal processes present (fig. 79)

RE EE le ee le erge Haplosymploce

Genus Pseudothyrsocera Shelford

Pseudothyrsocera Shelford, 1906: 250.- Princis 1951: 54

(footnote, selected type species, Blatta xanthophila

Walker), 1969: 953 (literature).

Mopsera Hebard, 1929: 78. Type species: Phyllodromia rec-

tangulariter-vittata Brunner, by monotypy.

Leptothyrsocera Princis, 1965: 152. Type species:

Pseudothyrsocera fulva Hebard, by monotypy. Syn. n.

Diagnosis. — Some of the antennal segments may

or may not be plumose. Tegmina and wings narrow,

fully developed, the former with simple, straight radi-

al vein, discoidal sectors longitudinal (fig. 32). Hind

wing with radial vein simple, or questionably

branched, straight, media vein simple or forked, cubi-

tus vein with mostly one or two, and usually no or

rarely one incomplete branches, apical triangle absent

or subobsolete; anterior field comparatively narrow

(figs. 8, 10, 16). Front femur usually Type B, (fig.

20d), rarely Type A, (fig. 12), B, or intermediate be-

tween A and B (figs. 20a-d); pulvilli on four proximal

tarsomeres of all legs, tarsal claws simple, symmetri-

cal, arolia present, may be greatly reduced. Male: first

and seventh abdominal terga specialized (figs. 15,

17), or only the seventh with a tergal gland (fig. 28).

Supraanal plate symmetrical, paraprocts dissimilar,

intercercal processes absent (fig. 11). Subgenital plate

(visible portion in the pinned specimen) usually is

asymmetrical, or nearly so, its shape variable (e.g.,

figs. 6, 14, 26, 37, 42). One (figs. 26, 35, 37) or usu-

ally two styles are present, and if the latter they are

similar (figs. 29, 47) or dissimilar (fig. 14). Genitalia

with three distinct phallomeres (e.g., fig. 37): genital

hook on the left side, with a preapical incision (fig. 6);

median phallomere simple (e.g., fig. 14), or bifurcate

on distal half (figs. 6, 19, 37); right phallomere con-

sisting of two or more sclerites, one of them often a

clac st, 19):

Remarks.- Princis (1965: 152) erected the mono-

typic genus Leptothyrsocera, and designated

Pseudothyrsocera fulva Hebard as its type species. He

selected the following characters given by Hebard in

his description of fulva as the diagnostic features of

the genus: Tegmen with the cubitus vein at the distal

end of the anal field geniculate (abruptly bent in an

obtuse angle) and there giving off a vein. Radius of

hind wing with a bifurcation. Front femur Type B,

[the front femur of fulva is Type B,, Type B,, or in-

termediate between Types A and Type B (figs. 20a-

d)]; four proximal tarsomeres of the hind tarsi with

very small pulvilli, tarsal claws symmetrical, unspe-

cialized, arolia very small. Princis stated that

Leptothyrsocera can be separated from Pseudothyrsocera

by bifurcation of the radius vein of the hind wing.

The geniculate cubitus of the tegmen may be found

in some species of Pseudothyrsocera (e.g., fig.32).

Sometimes it is difficult to decide whether or not the

radius is bifurcated. Rehn (1951: 20) states that the

radius usually has an apical posterior branch which

may be either simple or show secondary divisions,

and at times it has become indistinguishable or lost;

his drawing (fig. 2) shows the branch distinctly pro-

jecting posteriorly between the radius and media

veins. There is no question that the radius is bifurcat-

ed if the branch arises somewhere along its length and

extends posteriorly between it and the media vein, as

is characteristic of most Episymploce (fig. 112),

Hemithyrsocera (fig. 108) and many other genera. The

radial veins of species of Pseudothyrsocera are essential-

ly unbranched. In Pseudothyrsocera fulva the so-called

bifurcation continues as a straight line from the ra-

dius and does not project distinctly posteriorly be-

tween the radius and media; the penultimate apical

ramus with its five small branches (in the &) is wide-

ly separated from the small terminal vein (Hebard

1929: pl. 6, fig. 2) and both Hebard and Princis con-

sider the vein to be bifurcated; I interpret this type of

radial vein as having a pseudoposterior branch (figs.

41, 45). The lengths and branching of the apical rami

of the radius (Rehn's 1951 terminology) may vary

among species (e.g. figs. 8, 16, 31, 41, 45), and the

branch may protrude very slightly posteriorly. The

presence of one additional small terminal spine on the

anteroventral margin of the front femur (Type B,) in

Leptothyrsocera is of no generic importance here be-

cause these small stout spines may or may not be pre-

sent or are about the same length as piliform spinules

(figs. 20a-d). I can see no distinct differences between

Leptothyrsocera and Pseudothyrsocera and I consider

the former a junior synonym. Hebard (1929) includ-

ed only the type species in Mopsera and suggested that

Phyllodromia castanea Brunner (listed with a query in

Princis's catalog) might also belong in this genus.

Princis (1969: 954) listed Mopsera as a synonym of

Pseudothyrsocera and I agree with him. The genus

Mopsella Hanitsch (1936: 394) which Bruijning

(1947: 226) renamed Hanitschia, because the original

name was preoccupied, is nearest to Mopsera accord-

ing to Hanitsch (1936: 394): ‘Points of agreement be-

tween Mopsella and Mopsera are: small size and deli-

cate structure; broad interocular space; simple, not

plumose, antennae; narrow and elongate tegmina and

wings; radial vein of tegmina and wings simple (i.e.,

not forked); discoidal sectors of the tegmina longitu-

dinal; ulnar [= cubitus] vein of hind wings with com-

plete branches; apical triangle absent. The chief dif-

ferences are: Mopsella has a broad, not elongate

pronotum; its hind femora are weakly spined (front

femora missing); the costal and axillary veins of the

wings are much fewer in number than in Mopsera.’

The so-called differences between these two genera

do not clearly separate them and they are most likely

synonyms (of Pseudothyrsocera). Unfortunately, the

type species, Mopsella toxopei Hanitsch, is known

only from a unique female and the male should be ex-

amined before a final decision can be made regarding

its correct generic placement.

The male genital hook of Pseudothyrsocera is on the

left side which places it in the Blattellinae of

McKittrick's (1964) system. I divide Pseudothyrsocera

into the following two species groups and their sub-

groups:

1. fulva-species-group. — Subgenital plate variable

in shape, the exposed portion (in the pinned speci-

men) not distinctly trigonal, without a process mid-

way on the thickened left lateral margin (figs. 14, 19).

One (fig. 37) or two styles (fig. 19) present.

Subgroup (a). Only the seventh abdomninal ter-

gum specialized. — Species: perkinsi; rubronigra; xan-

thophila.

Subgroup (b). Abdominal terga one and seven spe-

cialized.- Species: fulva; rectangularitervittata; scutig-

era.

2. lugubris-species-group. — Visible portion of the

subgenital plate trigonal, usually with a process mid-

way on the lateral margins, left side incrassate, with a

pair of similar or dissimilar or similar styles usually

close together at the apex (figs. 47, 56), or sometimes

ROTH: Blattaria, Blattellidae, Blattellinae

more widely separated (fig. 67).

Subgroup (a). Only the seventh abdominal tergum

specialized. — Species: circumcincta; henrici; lugubris;

rufiventris.

Subgroup (b). Only the first abdominal tergum

specialized. — Species: sinensis.

Subgroup (c). First and seventh abdominal terga

specialized. — Species: montana.

The /ugubris-species-group has subgenital plates

and styles similar to those of many species of

Episymploce and they may be a link between

Pseudothyrsocera and Episymploce.

Key to males of Pseudothyrsocera

1. Shape of the exposed portion of the subgenital

plate variable, not trigonal, left side not thick-

ened and without a process midway on the lat-

eral margin, with one or two styles (Figs. 14,

19, 26) (fulva-species-group)

= Shape of the exposed portion of the subgenital

plate trigonal, the left side thickened and with

a lateral process near the middle of the margin

(except in henrici), two styles present(figs. 42,

47, 49, 56, 67) (lugubris-species-group) .. 7

. Tergal gland on segments one and seven (figs.

15 sel Deer er ann mens nt an Gaia 3

— Tergal gland only on segment seven (figs. 9,

28, 34)

3 (2). Apex of the subgenital plate with a very large

process that is upturned obliquely to the left

and bearing a minute terminal style, second

style absent (figs. 5-7). Pronotal disk with a

broad, dark, round, O-shaped macula (fig.2)

Ke ERS. rectangularitervittata

= Subgenital plate, styles, and pronotum not as

above ned cent Net 4

4 (3). Apex of supraanal plate with a pair of small

contiguous styles (fig. 19). Supraanal plate as

in fig. 18. Pronotal disk without a dark macula

We ATROCE IO EL eS fulva

= Apex of subgenital plate roundly produced,

with a minute spinelike style at the apex (fig.

26). Pronotal disk with a large, dark macula

(Ag DR ia ora scutigera

. Subgenital plate rounded apically, with a small

spinelike style on the right posterolateral cor-

ner and a minute hidden style medially (figs.

13, 14). Supraanal plate as in figs. 9, 11 ....

Ru er be che den av Ses xanthophila

= Subgenital and supraanal plates not as above

. Hind margin of subgenital plate rounded, not

produced, medially with a pair of similar, al-

most contiguous styles (fig. 29) .. rubronigra

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 1-8. Pseudothyrsocera rectangularitervittata (Brunner), males from Kuching, Sarawak: 1, head; 2, pronotum; 3, supraanal

plate and paraprocts (ventral); 4, abdominal terga five to ten and subgenital plate (dorsal); 5, supraanal and subgenital plates

(end view); 6, subgenital plate and genitalia (dorsal); 7, elongated process of the subgenital plate, and penultimate sternum

(left lateral view); 8, left hind wing [a = simple (unbranched) radial vein].

— Apex of subgenital plate produced and bearing

at its apex a large, broad style, second style ab-

SCHALEN eee aomeos ooo Ok perkinsi

7 (1). Abdominal terga one and seven specialized.

Supraanal plate and paraprocts as in fig. 55

LE es PEN eee nt montana

= Only abdominal tergum one or seven special-

ized. Supraanal plate and paraprocts not as

ADOVE eR TTC M LA 8

. Only abdominal segment one specialized (fig.

1B in Roth 1987c). Styles similar, very large,

close together (fig. 1D in Roth 1987c)

SN vn A AR PRE he ent i le LS sinensis

= Only abdominal segment seven specialized.

Stylesinotas DOS repre rei tia. Lee 9

9 (8). Left style elongated, spinelike, erect, located at

the apex of the subgenital plate, right style well

separated from it; midway on the left and right

margins of the subgenital plate without a

membranous process (fig. 67) ...... henrici

= Left style not as above, both styles close togeth-

er at the apex of the plate; a small membranous

process is located midway on the left margin of

the subgenital plate (a process occurs on the

right side, but it may be greatly reduced or sub-

obsoleto)i(fies 47 49) DO 10

10 (9) Pronotum completely dark or with a very nar-

row yellowish line along the anterior margin

(fig. 38). Supraanal plate and paraprocts as in

AO a rufiventris

= Pronotum dark, with a distinct pale ring

around the entire border of the disk (figs. 43,

LS) one A RR ME RA ci 11

11(10) Supraanal plate with median region roundly

produced, the curved hind margin with a row

of dark setae (fig. 50). Abdominal terga and

STEEMATO LAN OI SIAM erst ere D circumcincta

- Supraanal plate as in fig. 46. Abdominal terga

reddish brown or black, sterna reddish

fulva-species-group

Pseudothyrsocera rectangularitervittata (Brunner)

(figs. 1-8)

Phyllodromia rectangulariter-vittata Brunner, 1898: 203, pl.

XVI, fig. 3 (female) (p. 107, recte vittata Br., lapsus cala-

mi). Holotype @ [not examined], Baram, N.W. Borneo

(Sarawak); probably in the Genoa Museum, Italy. —

Kirby 1904: 91; Shelford 1908a: 13; Hanitsch 1923: 411.

Mopsera rectangularitervittata (Brunner). — Hebard 1929:

78, pl. 5, fig. 6, pl. 6, fig. 1 (male and female); Bruijning

1948: 98; Princis 1969: 954.

Pseudothyrsocera rectangularitervittata (Brunner). — Princis,

1969: 956.

ROTH: Blattaria, Blattellidae, Blattellinae

Specimens examined. — Borneo. NMWA: los

Grabowsky, coll. Br. v. W. Sarawak. ansp: Kuching

[1°32'N 110°20'E], Sarawak, 16 (terminalia slide

454), 9.v.1900, Dyak coll. [all that remains of this

specimen is one badly damaged wing, and the termi-

nal abdominal segments in a vial attached to the pin.

Hebard's description of the genus apparently was

based only on this one specimen, presumably before

it was badly damaged. I prepared a slide of the termi-

nal segments and genitalia, see figs. 3, 6.]. HECO: N.

Borneo, Kuching, 36, 1899, Dyak coll., pres. 1899

by R. Shelford.

Redescription. — Male: Head with interocular

width greater than the distance between large round

ocellar spots (fig. 1). Pronotum suboval widest be-

hind the middle (fig. 2). Tegmina and wings fully de-

veloped extending beyond end of abdomen, the for-

mer with simple radial vein and longitudinal

discoidal sectors. Hind wing with simple, straight ra-

dial vein, medial vein simple, or bifurcate distally,

weakly concave, cubitus vein weakly concave with

one or two complete and no incomplete branches,

apical triangle absent (fig. 8). Front femur Type B,

with seven or eight long and short stout proximal

spines that decrease strongly in length distad succeed-

ed by a short row of five or six piliform spinules ter-

minating in three large spines; pulvilli on four proxi-

mal tarsomeres, tarsal claws symmetrical, simple,

arolia present. First abdominal tergum with a tuft of

setae anteromedially; fifth abdominal tergum with a

deep, wide, inverted U-shaped excavation; seventh

abdominal segment with a tuft of setae on an antero-

medial elevation (fig. 4). [According to Hebard

(1929: 78) the male is specialized on abdominal seg-

ment one (‘median segment’) and segments five to

seven (‘fourth to sixth’}. Although the fifth tergum is

deeply excavated (fig. 4), there are no setae, depres-

sions, or clear areas on it and I do not consider this

segment and the sixth specialized. Only segments one

and seven (fig. 4) have a group of medial setae.

Supraanal plate symmetrical, with hind margin

broadly V-shaped (may be deflexed so that the V is

not seen from above), right and left paraprocts small,

dissimilar sclerites; medially between the cerci is a

membranous shelf, intercercal processes absent (fig.

3). Subgenital plate elongate, strongly convex, at its

apex a huge upturned process obliquely to the left, at

its apex a small style (figs. 5-7). Genitalia as in fig. 6:

hook on the left side, elongate, with a preapical inci-

sion; median phallomere divided near the middle, the

distal half with a two-pronged asymmetrical struc-

ture; right phallomere consisting of three sclerites,

one of which is a reduced cleft and another is a plate

bearing setae along one margin.

Colour. — Head with occiput and vertex dark

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

©) 1 mm

Figs. 9-14. Pseudothyrsocera xanthophila (Walker), males. 9-13, from holotype: 9, abdominal terga seven to ten (supraanal

plate); 10, hind wing; 11, supraanal plate and paraprocts (ventral); 12, front femur (anterior view); 13, subgenital plate and

ninth laterotergites (ventral); 14, subgenital plate (from holotype), and genitalia (from Forsten Belang) (dorsal; arrow indi-

cates the vestigial left style).

brown, remainder brownish yellow, labrum dark (fig.

1). Pronotal disk with a broad, black ring surround-

ing a yellowish macula, lateral region yellowish, sub-

hyaline (fig. 2). Tegmina brownish-hyaline, anterior

margin yellowish. Wings infuscated, darker in the

costal region and near the apical margin in both ante-

rior and posterior fields (fig. 8). Abdominal terga

brownish with lateral zones brownish yellow, seg-

72)

ments seven to nine darker except for brownish yel-

low lateral maculae, supraanal plate completely dark

(fig. 4). Abdominal sterna brownish yellow. Cerci

dorsally with proximal cercomeres dark brown, re-

maining segments and ventral surface, yellowish. Legs

brownish yellow.

Female: Not seen.

Measurements (mm). Length, 9.6-11.0; pronotum

length X width, 2.8-3.1 X 3.3-3.5; tegmen length,

10.1-10.5; interocular width 0.7-0.8.

Pseudothyrsocera xanthophila (Walker)

(figs. 9-14)

Blatta xanthophila Walker, 1868: 230 (male).

Thyrsocera xanthophila (Walker). — Kirby 1904: 78.

Pseudothyrsocera xanthophila (Walker). — Shelford 1906:

250; 1907: 489, pl. 30, figs. 1, 2 (male); 1908a: 5; Princis

1969: 954.

Specimens examined. — Holotype d (terminalia

slide 270) of Blatta xanthophila, Menado, Celebes,

Wallace, W.W. Saunders colln., purchased and pres.

'73 by Mrs. F.W. Hope; Type Orth. 51, in HECO.

Additional specimens. — Sumatra. NMWA: Sumatra,

Deli, Bandar Baroe, 19, 23.1.22, Fulmek 1921-26,

coll. R. Ebner (det. Hanitsch, 1931). RMNH: Belang,

Forsten, 16 (terminalia slide 207) (labelled xan-

thophila by Bruijning).

Redescription. — Male: Head with interocular

width slightly greater than the distance between an-

tennal sockets. Pronotum suboval. Tegmina and

wings fully developed extending beyond end of ab-

domen, former with simple radial vein and longitudi-

nal discoidal sectors. Hind wing with straight, simple

radial vein, media and cubitus veins weakly curved,

former simple, the latter with one to three complete

and no incomplete branches, apical triangle absent

(fig. 10). Front femur intermediate between Type A,

and B, with the row of preterminal spines very short

and stout, not piliform-like (fig. 12); pulvilli on four

proximal tarsomeres, tarsal claws simple, symmetri-

cal, arolia small. First abdominal tergum unspecial-

ized; hind margin of fifth tergum straight; seventh

tergum with a pair of deep fossae separated by a pale

medial elevation, setae absent (fig. 9); ninth lateroter-

gites dissimilar, the right one with a spinelike process

(fig. 13); supraanal plate produced, the apex narrow-

ly concave with a dark downwardly directed spine in

each corner (fig. 11; the spines are not seen in dorsal

view, fig. 9); paraprocts dissimilar the left one with

one and the right with two spinelike sclerotizations;

intercercal processes absent (fig. 11). Subgenital plate

narrow, elongate, densely covered with long slender

setae on the posterior half, hind margin rounded, left

style subobsolete, the right one small spinelike arising

within the margin on the dorsal surface (figs. 13, 14;

both styles cannot be seen in the pinned specimen

which made Shelford conclude that they are absent;

his 1907: pl. 30, fig. 2 drawing of the subgenital plate

shows an elongated process on the left side of the

plate which I did not see in my specimen.). Genitalia

as in fig. 14: hook on the left side, with a preapical in-

cision; median phallomere a simple curved rod; right

ROTH: Blattaria, Blattellidae, Blattellinae

phallomere consisting of three sclerites one of them a

cleft.

Colour. — Yellowish without distinctive markings.

Tegmina brownish yellow hyaline. Hind wing with

proximal or more portion of anterior field yellowish,

remainder darkly infuscated (fig. 10). Cerci yellowish

on both surfaces.

Female (previously undescribed): Head with inte-

rocular space greater than the distance between an-

tennal sockets. Pronotum suboval. Front femur is dis-

tinctly Type B, with four to six stout spines

decreasing in length distad, succeeded by seven pili-

form spinules, terminating in three larger terminal

spines [the male front femur is intermediate between

Types A and B, with the preterminal spines short and

stout which are similar to piliform spinules in length

(fig. 12)]. Tegmina and wings fully developed, ex-

tending beyond the end of the abdomen, the former

with longitudinal sectors. Hind wing with simple ra-

dial and media veins, cubitus vein with two complete

and no incomplete branches, apical triangle absent.

Supraanal plate trigonal, apex rounded. The coloura-

tion is yellowish, essentially unicolorous, and similar

to the male.

Measurements (mm) (® in parentheses). — Length,

10.0 (9.5); pronotum length X width, 3.2 X 3.5-

3.7 (3.2 X 3.7); tegmen length, 10.0-11.0 (11.5); in-

terocular width, 0.9-1.0 (1.0).

Remarks. — Princis (1969: 954) questioned the

Sumatra record of xanthophila but the species does

occur on that island.

Pseudothyrsocera fulva Hebard

(figs. 15-20)

Pseudothyrsocera fulva Hebard, 1929: 79, pl. 6, fig. 2 (habi-

tus) (female). — Hanitsch 1929a: 269 (incorrectly syn-

onymized with Pseudothyrsocera rubronigra Hanitsch);

1932a: 64 (incorrectly synonymized with rubronigra).

Pseudothyrsocera xanthophila (nec Walker). — Hanitsch

1929a: 269; Bruijning 1948: 93, fig. 42 [misidentifica-

tion and incorrectly synonymized fulva with xanthophila

(Walker) ]}.

Leptothyrsocera fulva (Hebard). — Princis 1965: 152, 1969:

953.

Specimens examined. — Holotype 2, Fort de Kock

[0°19'S 100° 22'E], Sumatra, 920 m, 1.1922, E.

Jacobson; Type no, 1147, in ansp. Additional speci-

mens. — ZMAN: same locality and collector as holo-

type, 12, vi.1922, 16 (terminalia slide 51), 29 (one

labelled Pseudothyrsocera rubronigra Hanitsch, by

Hanitsch), 1924, 16, 49, 1925, 22 (one labelled

Pseudothyrsocera xanthophila Walker, by Hanitsch),

1926. One retained in MCZC. RMNH: Fort de Kock,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 15-20. Pseudothyrsocera fulva Hebard, from Fort de Kock, Sumatra. 15-19, males: 15, first abdominal tergum; 16, hind

wing; 17, abdominal terga five to seven; 18, supraanal plate and paraprocts (ventral); 19, subgenital plate and genitalia (dor-

sal); 20, front femora (anterior views) (a, male, b-d, females).

Sumatra, 920 m, 1d, 1926, E. Jacobson; 14, 19,

Sumatra Exp., 1877-1878; Suban Ajam, Sum., 19,

vii.1916; Forsten Belang, 1d. One retained in Mczc.

HECO: Fort de Kock, Sumatra, 920 m, 18,1%, 1925,

29, 1926, E. Jacobson (misidentified as xanthophila

by Hanitsch).

Redescription. — Male: Head slightly or distinctly

exposed, eyes wide apart, interocular space greater

than distance between ocellar spots and antennal

sockets. Pronotum suboval, widest behind middle.

Tegmina and wings fully developed extending be-

yond end of abdomen, former with longitudinal dis-

coidal sectors. Hind wing with straight, simple, radi-

al vein, media and cubitus veins straight or weakly

curved, the latter with one bifurcated or two com-

plete and no incomplete branches, apical triangle ab-

sent (fig. 16). Front femur Type B, or intermediate

between A, and B,, the large proximal spines decreas-

ing sharply in length so that they become very short

stout spines about the same length as the small num-

ber of piliform spinules (fig. 20a); pulvilli on four

proximal tarsomeres, tarsal claws symmetrical, sim-

ple, arolia small. First abdominal tergum with setae

on anterior half, these densest anteromedially (fig.

15); fifth abdominal tergum without setal tufts or

clear areas (not specialized), hind margin deeply, con-

cavely excavated; sixth segment not specialized, sev-

enth tergum with a pair of depressions anteromedial-

ly, separated by a longitudinal ridge bearing setae (fig.

17); supraanal plate with hind margin broadly exca-

vated, paraprocts dissimilar, the right one with a ser-

rated sclerotization, the left one with a spinelike

process; intercercal processes absent (fig. 18).

Subgenital plate asymmetrical, the apex produced, di-

rected dorsad and bearing a pair of small dissimilar

contiguous styles, the right one with a large terminal

spine (fig. 19). Genitalia as in fig. 19: hook on the left

side, with a preapical incision; median phallomere

apically acute, with a distal branch whose apex is spic-

ular; right phallomere consisting of two sclerites, one

of them a reduced cleft, distal margin of the other se-

tose.

Female: Front femur Type B, (fig. 20d) or B,, or in-

termediate between Types A and B, the large stout

proximal spines decreasing sharply in length, these

followed by a few minute piliform spinules (some

small stout spines about the same length as spinules

may occur among the slender spinules; fig. 20c); at

least 2 females have no piliform spinules and the short

row of spines that preceed the 3 terminal spines are

stout and about the same length as piliform spinules

(Type A; fig. 20b). Supraanal plate symmetrical,

rounded, apex reaching slightly beyond margin of

subgenital plate.

ROTH: Blattaria, Blattellidae, Blattellinae

Colour. — Head, pronotum, cerci, and legs brown-

ish yellow; basal antennal segments brownish yellow,

remainder dark brown to black. Tegmina reddish

brown-hyaline gradually becoming pale along the an-

terior margin. Hind wing with yellowish tinge, apex

of anterior field and margin of posterior field infus-

cated. First abdominal segment and terga six to ten

dark brown, remainder brownish yellow. The tegmi-

na and dark markings on the abdomen usually are

lighter in the female.

Measurements (mm) (® in parentheses). — Length,

10.0-11.1 (9.0-11.3); pronotum length X width,

3.1-34 X 3.6-3.9 (3.3-3.5 X 3.6-4.1); tegmen

length, 10.0-11.6 (10.0-11.6); interocular width, 1.1

(182);

Remarks. — Although rectangularitervittata and ful-

va have been misidentified, their subgenital plates,

styles, and genital phallomeres are distinctly different.

Hanitsch (1929a: 268, 269) reported 11 specimens

of Pseudothyrsocera xanthophila and five examples of

rubronigra from Fort de Kock (same data as speci-

mens examined here) and stated that the two species

differed in colour; because Hebard's fulva also came

from Fort de Kock, Hanitsch synonymized it with

xanthophila. Bruijning (1948: 94) accepted

Hanitsch's identifications and synonymy stating that

the only difference between these two species is

colour and concluded that rubronigra is a dark morph

of xanthophila and synonymized the two species.

Clearly, rubronigra (figs. 27-31) differs from fulva

(figs. 15-19), as well as xanthophila (figs. 9-14) and

the three species are distinct. Bruijning's (1948: fig.

42) drawing of the subgenital plate and styles is that

of fulva and not xanthophila as he indicated.

Pseudothyrsocera scutigera (Walker)

(figs. 21-26)

Pseudomops scutigera Walker, 1868: 212 (male).

Pseudothyrsocera scutigera (Walker). — Kirby 1904: 77;

Shelford1906: 250, 1907: 489, 1908a: 5; Hanitsch 1915:

34, 1923: 397, 1931: 393; Hebard 1929: 10 (scutiger);

Bruijning 1948: 95; Princis 1969: 954 (literature).

Specimens examined. — Holotype, d [abdomen

missing], Sarawak, Wallace in Mr. Saunders collec-

tion; Type Orth. 52 in HECO. Additional specimens.

— Sabah. zits: Malaysia, Sabah, Sipitang [5°05'N

115°33'E] Mendolong, 14, 8.xii.1987, T1B/W4,

14 (terminalia slide 423), 25.iv.1988, 16,

11.v.1988, 16, 14.iii. 1988, S. Adebratt. One re-

tained in Mczc. Unknown country. RMNH: Orut

Ratuna, 18, v.1865, Van Hasselt (reported by

Bruijning, 1948: 95).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 21-26. Pseudothyrsocera scutigera (Walker), males from Sabah: 21, pronotum; 22, supraanal plate and paraprocts (ven-

tral); 23, first abdominal tergum; 24, seventh abdominal tergum; 25, hind wing; 26, subgenital plate and genitalia (dorsal).

Redescription. — Male: Head exposed, interocular

width slightly greater than distance between ocellar

spots. Antennae not plumose. Pronotum suboval,

sides deflexed, widest behind middle (fig. 21).

Tegmina and wings fully developed extending be-

yond end of abdomen, former with longitudinal dis-

coidal sectors. Hind wing radial and media veins sim-

ple, straight, cubitus straight with a single complete

and no incomplete branches, apical triangle

absent (fig. 25). Front femur Type B, with four or five

large or stout proximal spines; pulvilli on four proxi-

mal tarsomeres, tarsal claws simple, symmetrical, aro-

lia well developed. First abdominal tergum special-

ized with setal groups anteromedially, the densest in a

raised middle region (fig. 23). Seventh abdominal ter-

gum with a pair of shallow depressions separated by a

longitudinal ridge bearing two rows of setae (fig. 24).

Supraanal plate with sides of hind margin oblique,

0.5 mm

gato 2 mm

ROTH: Blattaria, Blattellidae, Blattellinae

Figs. 27-31. Pseudothyrsocera rubronigra (Hanitsch), male lectotype. 27, supraanal plate and paraprocts (ventral); 28, seventh

abdominal tergum; 29, subgenital plate and genitalia (dorsal); 30, left tegmen; 31, left hind wing.

apical region rounded, weakly indented medially;

paraprocts dissimilar, left one with a small hooklike

sclerotization, right one with two dissimilar spinelike

sclerotizations; intercercal processes absent (fig. 22).

Subgenital plate practically symmetrical, hind margin

with a rounded protruding structure medially, which

may represent a nonarticulated style bearing a minute

apical seta; to the left of this structure is a long slender

seta or one that is swollen medially (fig. 26). Genitalia

as in fig. 26: hook on the left side with a preapical in-

cision; median phallomere slender, simple; right

phallomere consisting of two sclerites of which one is

a cleft.

Female: Unknown.

Colour. — Head black, labrum lighter, maxillary

palpomeres one to three and basal half of the fourth

pale, remainder black; antennae with about the first

30 antennomeres black succeeded by about 30 white

segments, remainder black. Pronotum with a large

black macula surrounded by yellow (fig. 21);

mesonotum dark, metanotum yellow. Tegmina with

part of anterior region reddish, remainder black. First

abdominal tergum black except for yellow lateral

zones, segments two to five yellow, the sixth black ex-

cept for yellow lateral areas, seven to ten black except

for pale apex on supraanal plate. Abdominal sterna

yellow, subgenital plate dark. Front coxae black on

basal half, rest white, mid and hind coxae mostly

black their distal parts and outer margins, and

trochanters white, femora, tibiae and tarsae dark red-

dish brown. Cerci white, their proximal segments

with yellowish tinge.

Measurements (mm) (the holotype is smaller than

the others and is shown in parentheses). — Length,

8.6-10.5 (abdomen missing); pronotum length X

width, 2.8-3.0 X 3.1-3.2 (2.5 X 2.8); tegmen

length, 10.6-11.2 (10.0); interocular width, 0.9-1.0

(0.8).

Remarks. — The distinctive colour markings (par-

ticularly the pronotal macula) separates this species

from others in the genus.

Pseudothyrsocera rubronigra (Hanitsch)

(figs. 27-31)

Phyllodromia rubro-nigra Hanitsch, 1923: 412, fig. 11 only,

(male only, not female). — Bruijning 1948: 93 (incorrect-

ly synonymized with xanthophila).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 32-37. Pseudothyrsocera perkinsi sp. n., male holotype: 32, left tegmen; 33, left hind wing. 34, seventh abdominal ter-

gum; 35, single style at the apex of the subgenital plate (ventral); 36, supraanal plate and paraprocts (ventral); 37, subgenital

plate and genitalia (dorsal; a, left phallomere; b, median phallomere; c, sclerites of the right phallomere; d, style).

Pseudothyrsocera rubro-nigra (Hanitsch). — Hanitsch 1928:

14, 1929a: 269, 1932a: 64; Hebard 1929: 10 (exclusive

OL LIL

Pseudothyrsocera fulva (nec Hebard). — Hanitsch 1929a: 269

(misidentification).

Mopserina rubronigra (Hanitsch). — Princis 1965: 152 (ex-

clusive of 9°).

Hemithyrsocera rubronigra (Hanitsch). — Princis 1971: 1127

(literature, exclusive of ‘9 ’).

Specimens examined. — Lectotype d (selected by

Hebard 1929: 79) (terminalia slide 267), Gunong

Angsi, Negri Sembilan, Malaya, [2°45'N 102°10'E]

2000'-2790', iv.1918, Hanitsch; Type Orth. 263'/, in

HECO. Additional specimens. — Malaya. HECO: same

data as lectotype, 1, Type Orth. 2637/, (see remarks,

below).

Redescription. — Male: Head exposed, eyes bul-

bous, somewhat reduced, interocular space greater

than the distance between antennal sockets and ocel-

lar spots; proximal antennal segments weakly

plumose. Pronotum suboval. Tegmina and wings ful-

ly developed extending beyond end of abdomen, the

former narrow with longitudinal discoidal sectors

(fig. 30). Hind wing with simple, thickened costal

veins, media and radial veins straight, simple, cubitus

vein straight, with one long complete and no incom-

plete branches, apical triangle absent (fig. 31).

Anteroventral margin of front femur Type B,, with

five to seven large proximal spines; pulvilli on four

proximal tarsomeres, tarsal claws simple, symmetri-

cal, arolia small. First abdominal tergum unspecial-

ized. Seventh abdominal tergum with a small group

of setae anteromedially (fig. 28). Supraanal plate hind

margin with oblique sides and broadly subtruncate

apex, right and left paraprocts dissimilar, intercercal

processes absent (fig. 27). Subgenital plate weakly

asymmetrical with a pair of similar, almost contigu-

ous styli mesad on the hind margin; each style con-

sists of a small base arising within the margin of the

plate succeeded by a rectangular segment and termi-

nating in an articulated spine (fig. 29). Genitalia as in

fig. 29: hook on the left side, with a preapical inci-

sion, median phallomere, slender, simple, apex acute,

right phallomere consisting of two sclerites one of

which is a cleft, and near it are two small spines.

Colour. — Head orangish, unicolourous, maxillary

palpomeres dark brown, proximal antennal segments

and apex blackish, remainder yellowish. Pronotum

orangish without markings. Tegmina dark reddish

brown-hyaline, veins very dark. Hind wing weakly in-

fuscated, anterior region and apex of anterior field

darker (fig. 31). Abdominal terga light brown, termi-

nal segments darker. Abdominal sterna light brown,

subgenital plate darker. Coxae and femora orangish,

the latter apically dark, tibiae and tarsi blackish

brown. Cerci yellowish white.

Female. Unknown.

Measurements (mm). — Length, 7.8; pronotum

length X width, 2.2-2.3 X 2.6; tegmen length, 9.4-

9.5; interocular width, 1.0-1.1.

Remarks. — Hanitsch (1923) reported only two

syntypes, one male and one female, in his original de-

scription. These two specimens have handwritten

species labels. The ‘female’ actually is a male and is a

new species described below as perkinsi. There is a

third male (listed above as additional specimens) la-

belled Type Orth. 263°/,); it has the same data as the

lectotype and is clearly rubronigra but is not a syntype

and does not have a handwritten identification label.

Hanitsch noted that the ‘2’ of rubronigra was larg-

ROTH: Blattaria, Blattellidae, Blattellinae

er and the shape of the pronotum differed from the

male but he considered them to be the same species.

Hebard (1929: 79) disagreed and correctly stated that

the two sexes were different species. Princis (1965:

152) actually examined the “Typen (d 2)’ of rubron-

igra and concluded that they belonged to the same

species, were not related to ‘Pseudothyrsocera’ fulva

Hebard and placed it in Mopserina Princis which he

later (1971: 1124) synonymized with Hemithyrsocera,

and incorrectly listed (p. 1127) rubronigra under that

genus.

Pseudothyrsocera perkinsi sp. n.

(figs. 32-37)

Phyllodromia rubro-nigra Hanitsch (in part, ‘paralectotype’

male, incorrectly determined as ‘9°), 1923: 412, fig. 12

only.

Specimen examined. — d holotype (terminalia

slide 268), Gunung Angsi, Negri Sembilan, Malaya,

[2°45'N 102°10'E] 2000'-2790', iv. 1918, Hanitsch

(Type Orth. 2637, of Phyllodromia rubro-nigra

Hanitsch, 6, not ® as indicated); in HECO.

Description. — Male: Head almost completely hid-

den, eyes bulbous, somewhat reduced, wide apart, in-

terocular space greater that the space between ocellar

spots and antennal sockets; antennae missing.

Pronotum suboval. Tegmina and wings fully devel-

oped, the former narrow with longitudinal discoidal

sectors (fig. 32). Hind wing with thickened costal

veins, all but the preterminal one simple; radial vein

simple, media curved, cubitus vein curved with two

complete and no incomplete branches, apical triangle

absent (fig. 33). Front femur Type B, with six large

proximal spines; pulvilli on four proximal tarsomeres,

tarsal claws simple, symmetrical, arolia small. First

abdominal segment unspecialized. Seventh abdomi-

nal tergum with two small groups of setae medially

(fig. 34). Supraanal plate subtrapezoidal, paraprocts

dissimilar, the right one terminating in several small

spines, intercercal processes absent (fig. 36).

Subgenital plate distinctly asymmetrical, the distal

half of the left margin excavated, with a single large

style at the apex (figs. 35, 37). Genitalia as in fig. 37:

hook on the left side, with a preapical incision; medi-

an phallomere slender with a distal branch that termi-

nates in several small spines; right phallomere consist-

ing of two sclerites one of which terminates in a

narrow row of minute setae.

Colour. — Head and pronotum unicolorous,

orangish; maxillary palpomeres three and four pale,

terminal segment light brown. Tegmina blackish

with a narrow orangish stripe along most of the ante-

rior margin (fig. 32). Wings darkly infuscated, anteri-

or and apical regions darker (fig. 33).

79,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 38-42. Pseudothyrsocera rufiventris (Stàl), male from Surigao, Mindanao, Philippine Islands: 38, pronotum (the very nar-

row yellow anterior margin is asymmetrical and is missing on the right side); 39, abdominal terga seven to ten and subgeni-

tal plate (dorsal); 40, supraanal plate and paraprocts (ventral); 41, hind wing (a, pseudobranch of the radial vein); 42, sub-

genital plate and genitalia (dorsal).

Female: Unknown.

Measurements (mm). — Pronotum length X

width, 3.3-3.7; tegmen length, 11.3; interocular

width, 1.1.

Etymology. — The species is dedicated to my friend

Dr. Philip Perkins, Collection Manager, Entomology

Department, Museum of Comparative Zoology,

Harvard University.

Remarks. — See above remarks under rubronigra.

The single huge style at the apex of the subgenital

plate is unique for the genus.

lugubris-species-group

Pseudothyrsocera rufiventris (Stàl)

(figs. 38-42)

Thyrsocera rufiventris Stal, 1877: 33 (male). — Sjöstedt 1933: 3.

Hemithyrsocera rufiventris (Stal). — Kirby 1904: 77.

Pseudothyrsocera rufiventris (Stal). — Shelford 1908a, 5;

1908b: 469; Princis 1969: 956 [listed as a synonym of P.

signata (Brunner); see remarks, below].

Specimens examined. — Lectotype, d (here desig-

nated; with an unpublished lectotype label written by

Princis, 1968), Ins. Phillipp.; in Nrss. Philippine

Islands. Ansp: Surigao [9°45'N 125°30'E],

Mindanao, 2d (one labelled Pseudothyrsocera lugubris

(Stal) by Hebard, 1928) (one with terminalia slide

455); N.W. Panay [?7°20'N 124°14'E], 1d, Baker

(labelled Pseudothyrsocera rufiventris (Stal) by Hebard,

1928).

Redescription. — Male: Head exposed, interocular

distance greater than the distances between ocellar

spots and antennal sockets; antennae not plumose.

Pronotum suboval, widest behind the middle (fig.

38). Tegmina and wings fully developed, narrow, ex-

tending beyond end of abdomen, the former with

simple, straight radial vein and longitudinal discoidal

sectors. Hind wing with radial and media veins sim-

ple, straight, cubitus vein straight with two complete

and one long incomplete branch, apical triangle ab-

sent (fig. 41). Front femur Type B, with four large

proximal spines; pulvilli on four proximal tarsomeres,

tarsal claws symmetrical, simple, arolia small. First

abdominal tergum unspecialized. Seventh abdominal

tergum with a pair of shallow medial depressions with

minute spaced setae (which may be white), setal

groups absent; the depressions are separated by a lon-

gitudinal ridge (fig. 39) and there is a clear membra-

nous zone in the anterior part of the ridge which is

hidden by the overlapping sixth tergum in the pinned

specimen. Supraanal plate symmetrical, trigonal, apex

shallowly notched; paraprocts dissimilar darkly scle-

rotized, the right one with two spinelike processes, in-

tercercal processes absent (fig. 40). Subgenital plate

convex, trigonal, the sides upturned, asymmetrical,

with a large spinelike process about midway on the

left margin and a small dark seta on the opposite side;

a pair of small similar styles occur apically on the dor-

sal surface within the margin (not visible in ventral

view) (figs. 39, 42). Genitalia as in fig. 42: hook phal-

lomere large, on the left side with a preapical incision

and three groups of setae on the shaft just before the

curved region; median phallomere with a small acute

sclerite apically; right phallomere consisting of two

sclerites one of which is a reduced cleft.

Colour. — Very dark. Head with reddish eyes, ocel-

lar spots yellowish, antennae and maxillary palpi

black, clypeus and labrum somewhat lighter.

Pronotum usually solidly black or with a narrow in-

complete yellowish line along the anterior margin

(fig. 38). Tegmina black with a hyaline spot (very in-

distinct in lectotype) in the subcostal area. Hind

ROTH: Blattaria, Blattellidae, Blattellinae

wings very dark brown. Abdominal terga black,

supraanal plate dark reddish brown or black.

Abdominal sterna reddish, apex of subgenital plate

with a dark blotch. Legs black, outer margins of cox-

ae yellow. Cerci black.

Female: Not seen.

Measurements (mm). — Length, 12.0-14.5; prono-

tum length X width, 3.6-4.1 X 4.2-4.7; tegmen

length, 12.0-14.5; interocular width, 1.0-1.1.

Remarks. — Shelford (1908a: 5) listed P. semicincta

as a synonym of rufiventris, and Princis (1969: 956)

listed rufiventris as a synonym of P. signata. However,

the pronontum of rufiventris is virtually all dark ex-

cept for a subobsolete pale anterior margin (fig. 38);

the pronotum of signata (= semicincta) is shiny black

with yellow very narrow on the anterior, and lateral

margins (may be absent), and broad posteriorly (figs.

57, 59). Until the male of signata is found and com-

pared with rufiventris, I am listing them separately.

The male subgenital plate, styles, and genitalia of

rufiventris (fig. 42), lugubris (fig. 47) and circumcinc-

ta (fig. 49) are very similar. These taxa can be separat-

ed by the differences in the pronotal markings and

the shapes of the supraanal plates (cf. figs. 39, 46, 50).

Pseudothyrsocera lugubris (Stal)

(figs. 43-47)

Thyrsocera lugubris Stal, 1877: 33 (male). — Sjöstedt 1933: 3.

Hemithyrsocera lugubris (Stal). — Kirby 1904: 77.

Pseudothyrsocera lugubris (Stal). — Shelford 1908a: 5; Princis

1969: 955 (literature exclusive of synonymy).

Specimens examined. — Lectotype, d (here desig-

nated; with an unpublished lectotype label written by

Princis 1968), no exact locality, Ins. Phillip.

[Philippine Islands], Semper; in NRss. — Para-

lectotypes. Phillipine Islands. Nrss: same data as lec-

totype, 3d (1 with terminalia slide 72).

Redescription. — Male: Head with interocular

space slightly greater than distance between ocelli-

form spots and antennal sockets; antennae not

plumose. Pronotum suboval, widest behind middle

(fig. 43). Tegmina and wings fully developed extend-

ing beyond end of abdomen, former with longitudi-

nal discoidal sectors. Hind wing with simple, straight

radial and media veins, cubitus vein straight with two

long complete and no incomplete branches, apical tri-

angle absent (fig. 45). Front femur Type B, with three

large proximal spines, or with an additional one or

two small spines; pulvilli on four proximal tar-

someres, tarsal claws symmetrical, simple, arolia pre-

sent. First abdominal tergum unspecialized. Seventh

abdominal tergum with a pair of nonsetose depres-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 43-47. Pseudothyrsocera lugubris (Stäl), males from the Philippines: 43, lectotype, pronotum; 44-47, paralectotype: 44,

seventh abdominal tergum; 45, hind wing; 46, supraanal plate and paraprocts (ventral); 47, subgenital plate and genitalia

(dorsal; a, spinelike process on the left side; b, thickened left margin; c, small spinelike process on the right side).

sions separated by a longitudinal ridge; a narrow clear

zone anteromedially is seen in a slide preparation but

is hidden by the overlapping sixth segment in the

pinned specimen (fig. 44). Supraanal plate with me-

dian portion roundly produced or with a shallow me-

dial indentation (visible in pinned specimens), the

hind margin with some small spines, right and left

paraprocts dissimilar, intercercal processes absent (fig.

46). Subgenital plate asymmetrical, trigonal, the left

side with a large spinelike process and a broad round-

ly thickened border covered with fine setae, the right

margin with a minute spinelike process; a pair of sim-

ilar closely spaced serrated styli located at apex of

plate (fig. 47). Genitalia as in fig. 47: elongated geni-

tal hook on left side with a preapical incision; median

phallomere a curved rod with an acute membrane en-

closing the apex; right phallomere consisting of two

small sclerites, one a cleft.

Colour. — Head black, clypeus lighter, genae and

ocelliform spots yellow; antennae and maxillary palpi

black. Pronotum completely black or with a narrow

yellowish band along posterior margin and indistinct

dull reddish narrow band around the remaining mar-

gin (lectotype) (fig. 43), or dull reddish on hind mar-

gin only. Tegmina reddish brown or black with a yel-

lowish-hyaline mark on distal half of subcostal

region. Hind wings dark brown. Abdominal terga

dark reddish brown or black. Abdominal sterna red-

dish, subgenital plate dark brownish. Legs black, cox-

ae with outer margin yellow. Cerci black.

Female: Not seen.

Measurements (mm). — Length, 11.5-14.0; prono-

tum length X width, 3.3-4.0 X 3.9-4.5; tegmen

length, 12.0-14.0; interocular width, 1.0-1.1.

Remarks. — The species is closest to rufiventris (see

remarks under that species).

Pseudothyrsocera circumeincta (Stal)

(figs. 48-52)

Thyrsocera circumcincta Stàl, 1877: 33 (male). — Kirby 1904:

77 (synonymized under signata); Shelford 1908a: 5,

1908b: 469; Sjöstedt, 1933: 3.

Pseudothyrsocera circumcincta (Stal). — Shelford 1908a: 5;

1908b: 469; Princis 1969: 955 (listed as a synonym of

lugubris).

Specimens examined. — Philippine Islands. nass:

Holotype d (terminalia slide 73), of Thyrsocera cir-

cumcincta Stäl, Ins. Philipp. no exact locality,

Semper [Princis wrote ‘Pseudothysocera lugubris (Stal),

K. Princis, 1968’ on back of holotype label]; same

data as holotype, 19 (with handwritten label, ‘... =

circumcincta, teste Shelford’). Ansp: Imugan

OMAN Ono 2E NE zon Sr labelled

Pseudothyrsocera circumelusa (Stal) 9, by Hebard

1928].

Redescription. — Male: Head exposed, interocular

width greater than space between ocellar spots and

antennal sockets; antennae not plumose. Pronotum

suboval, widest behind middle (fig. 48). Tegmina and

wings fully developed, discoidal sectors of former lon-

gitudinal. Hind wing with radial and medial veins

simple, straight, cubitus vein straight with two long

complete and no incomplete branches, apical triangle

absent (similar to fig. 45). Front femur Type B, with

four long or three long and three short stout spines;

pulvilli on four proximal tarsomeres, tarsal claws sym-

metrical, simple, arolia small. First abdominal tergum

unspecialized. Seventh abdominal tergum with a pair

of shallow depressions separated by a longitudinal

nonsetose ridge (similar to fig. 44). Supraanal plate

medially broadly produced, the rounded hind margin

with a dense row of small uniform spines, paraprocts

dissimilar, intercercal processes absent (fig. 50).

Subgenital plate trigonal, with a spinelike process

midway on the left side, the margin posterior to it

thickened and covered with small setae; right and left

styles similar, spinelike (fig. 49). Genitalia as in fig.

49: hook on the left side with a preapical incision;

median phallomere a slender rod; right phallomere

consisting of two sclerites one of them a reduced cleft.

ROTH: Blattaria, Blattellidae, Blattellinae

Colour. — Head black, genae and ocellar spots

white; maxillary palpi and antennae black. Pronotum

black, completely surrounded by a broad yellow band

(fig. 48). Tegmina light reddish brown-hyaline, sub-

costal region lighter. Abdominal terga and sterna

orangish. Cerci dark brown. Legs dark reddish

brown, outer margins of coxae, and trochanters pale.

Female: Head slightly exposed, interocular width

greater than space between ocelliform spots; antennae

not plumose. Pronotum suboval (fig. 51). Tegmina

and wings fully developed extending beyond end of

abdomen, former with longitudinal discoidal sectors.

Hind wing with straight, simple, radial and media

veins, cubitus vein straight with two long complete

branches, apical triangle absent. Front femur Type B,

with four large proximal spines, tarsal claws simple,

symmetrical, arolia present. Supraanal plate broadly

trigonal, apex rounded (fig. 52).

Colour. — Head black, ocelliform spots and genae

yellow, maxillary palpi and antennae black.

Pronotum black completely surrounded by a broad

yellow margin (fig. 51). Tegmina dark reddish

brown, subcostal region yellow-hyaline. Wings dark

brownish. Abdominal terga black with triangular yel-

low marks on the posterolateral corners of segments

two to seven or two to six, supraanal plate completely

dark (fig. 52). Abdominal sterna black, lateral and

hind margins yellowish, basal half of subgenital plate

dark, distal half reddish. Legs black, outer margins of

coxae yellow.

Measurements (mm) (® in parentheses). — Length,

10.0 (10.5-13.4); pronotum length width, 3.1 X

4.0 (3.4-3.8 X 4.3-4.5); tegmen length, 10.5 (10.0-

11.7); interocular width, 0.9 (1.1).

Remarks. — Pseudothyrsocera circumcincta is clearly

closely related to /ugubris because of the similarities of

their subgenital plates. Princis (1969: 955) consid-

ered the former to be a synonym of the latter.

However, there is a distinct difference between their

sizes and supraanal plates; also their colours are strik-

ingly different and I prefer to treat them as distinct

taxa until additional specimens are studied to show

the extent of variation.

Kirby (1904: 77) synonymized P. circumcincta

(Stal) with szgnata but I am treating them separately.

Unfortunately, the latter was based on females only.

Brunner incorrectly described the posterolateral cor-

ners of the first five abdominal terga as having trian-

gular yellow spots whereas the signata holotype has

yellow lateral corners on segments three to five (as in

semicincta, fig. 58). The two females that I have de-

termined as circumcincta have a black abdomen with

yellow posterolateral corners on terga two to seven

(fig. 52), or two to six.

TUDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 48-52. Pseudothyrsocera circumcincta (Stal). 48-50, male holotype: 48, pronotum; 49, subgenital plate and genitalia (dor-

sal); 50, supraanal plate and paraprocts (ventral); 51, 52, female from Imugan, Luzon, Philippine Islands: 51, pronotum; 52,

abdomen (dorsal) (first tergum not shown).

Eee

È;

RER

à a

IR Bean

Ton

ROTH: Blattaria, Blattellidae, Blattellinae

Figs. 53-56. Pseudothyrsocera montana (Shelford), males from Kuching, Sarawak. 53, lectotype, pronotum and proximal re-

gion of the tegmina; 54-56, paralectotype: 54, hind wing; 55, supraanal plate and paraprocts (ventral); 56, subgenital plate

and genitalia (dorsal).

Pseudothyrsocera montana Shelford

(figs. 53-56)

Pseudothyrsocera montana Shelford, 1906: 251 (male). —

Shelford 1908a: 5; Hanitsch 1915: 35, 1923: 463;

Hebard 1929: 10; Bruijning 1948: 95.

Specimens examined. — Lectotype (here designat-

ed), d, Mt. Matanga, nr. Kuching, N.W. Borneo

[Sarawak], about 3000', vi.1900, pres. 1906 by the

Sarawak Museum; Type Orth. 49'/, in Heco.

Paralectotype. Sarawak. HECO: 14 (terminalia slide

299), same data as lectotytpe, Type Orth. 49°/,.

Redescription. — Male: Head with vertex exposed,

interocular space greater than the distance between

antennal sockets; antennae slight thickened, not

plumose. Pronotum suboval (fig. 53). Tegmina and

wings fully developed extending beyond end of ab-

domen, the former with longitudinal discoidal sec-

tors. Hind wing with simple radial and media veins,

cubitus vein with two complete and no (left wing) or

one (right wing) incomplete branches (paralecto-

type), apical triangle absent (fig. 54). Front femur

Type B, with seven stout proximal spines; pulvilli on

four proximal tarsomeres, tarsal claws symmetrical,

simple, arolia present. First abdominal tergum with a

group of setae anteromedially. Seventh abdominal

terga with a pair of shallow fossae containing sym-

metrically spaced small setae (seen in slide prepara-

tion). Supraanal plate trigonal, sides weakly concave,

apex slightly indented, intercercal processes absent,

right and left paraprocts with dissimilar spinelike

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

2mm

processes (fig. 55). Subgenital plate asymmetrical, the

posterior half trigonal (that portion visible in the

pinned specimen), left side roundly thickened, with

an elongated process near the middle of the lateral

margins, the one on the right side larger; a pair of nar-

rowly separated dissimilar spinelike styles, the right

one slightly longer, are located at the apex of the plate

(fig. 56; Shelford did not see the right style which was

upturned and hidden in the pinned specimen, and he

incorrectly stated that there is only one style).

Genitalia as in fig. 56: hook on the left side, very

large, apex with a preapical incision; median phal-

lomere a curved rod with a very small preterminal

process; right phallomere consisting of three sclerites

one of them a small cleft.

Female: Unknown.

Colour. — Head, maxillary palpi, antennae, prono-

tum, meso-, metanotum, and abdominal terga black.

Tegmina black with a yellowish spot on the distal half

Figs. 57-60. Pseudothyrso-

cera spp. 57-59, P. signata

(Brunner): 57-58, female

lectotype of Thyrsocera

semicincta Stal, pronotum,

and abdomen; 59, female

holotype of P. signata,

pronotum. 60, P. circum-

clusa (Stal), female lecto-

type, pronotum.

2mm

of the subcostal field (fig. 53).

Hind wings infuscated (fig. 54). Abdominal sterna,

coxae and femora reddish; cerci and apices of femora,

tibiae, and tarsi, black.

Measurements (mm). Length, 11.5; pronotum

length X width, 4.4 X 5.4; tegmen length, 14.2;

interocular width, 1.4.

Pseudothyrsocera signata (Brunner)

(figs. 57-59)

Thyrsocera signata Brunner, 1865: 119 (female).

Pseudomops signata (Brunner). — Walker 1869: 135.

Hemithyrsocera signata (Brunner). — Kirby 1904: 77.

Pseudothyrsocera signata (Brunner). — Shelford 1908a: 5.

Thyrsocera semicincta Stal, 1877: 33 (female). — Sjöstedt

193333:

Hemithyrsocera semicincta (Stal). — Kirby 1904: 77.

Pseudothyrsocera semicincta (Stäl). — Princis 1969: 956 (listed

as a synonym of signata).

Specimens examined. — Holotype 2 of Thyrsocera

signata Brunner, Philippines (no exact locality),

Thorey, coll. Br. v. W.; in NMWA. Lectotype (here

designated; with an unpublished lectotype label writ-

ten by Princis, 1968), ® of P. semicincta, Ins.

Philipp., no exact locality (Philippine Islands) [with

Princis's handwritten ‘Pseudophoraspis rufiventris

(Stal), K. Princis, 1968’ on back of his ‘lectotype’ la-

bel]; in Nrss. Paralectotype. NRSS: same data as lecto-

type, 12 (with handwritten label, rufiventris, teste

Shelford). Additional specimens. — ANSP: no exact lo-

cality, 12, Acc. No. 11998, Bu. of Sci., P.I. (det. as

semicincta by Hebard 1928).

Redescription. — Female: Head slightly exposed,

interocular width greater than distance between ocel-

lar spots and antennal sockets; antennae not plumose.

Pronotum suboval (fig. 57). Tegmina and wings fully

developed extending beyond end of abdomen, dis-

coidal sectors of former longitudinal. Hind wing with

radial and media veins straight, simple, cubitus vein

straight with two long complete and no incomplete

branches, apical triangle absent. Front femur Type B,

with three or four large proximal spines; pulvilli on

four proximal tarsomeres, tarsal claws symmetrical,

simple, arolia present. Supraanal plate trigonal, apex

rounded.

Colour. — Head black, genae black; maxillary palpi

and antennae black. Pronotum black with a narrow

yellow band on anterior margin and on posterior and

distal parts of lateral margins (fig. 57, 59). Tegmina

black with a yellowish-hyaline mark on distal half of

subcostal zone. Hind wing black-hyaline. Abdominal

terga black with a large triangular yellow macula in

posterlolateral corners of segments three to five (fig.

58). Abdominal sterna reddish, or reddish with

brown areas medially on proximal half or more. Legs

black or brownish black, coxae with yellowish white

outer margins. Cerci black.

Male: Unknown.

Measurements (mm). — Length, 13.0-14.0; prono-

tum length X width, 4.2-4.3 x 4.8-5.1; tegmen

length, 13.2-15.0; interocular width, 1.2.

Remarks. — Shelford (1908a: 5) listed P. semicincta

as a synonym of rufiventris but the pronotum of the

former is incompletely or completely ringed by yel-

low (figs. 57, 59) which is lacking in rufiventris (fig.

38) and I provisionally list the two taxa separately un-

til males are studied. I agree with Princis in consider-

ing semicincta a synonym of signata; the posterolater-

al corners of three abdominal terga are yellow (fig. 58)

in both taxa. There is a slight difference in the yellow

border of the pronotum, where it is narrowly com-

plete in signata and lacking laterally in semicincta.

ROTH: Blattaria, Blattellidae, Blattellinae

Pseudothyrsocera circumclusa (Stäl)

(fig. 60)

Thyrsocera circumclusa Stal, 1877: 34 (female). — Shelford

1908b: 469 (synonimized under circumcincta), 1908a: 5

(synonimized under circumcincta); Sjöstedt 1933: 3.

Hemithyrsocera circumclusa (Stal). — Kirby 1904: 77.

Pseudothyrsocera circumclusa (Stal). — Princis 1969: 955 (list-

ed as a synonym of lugubris).

Specimens examined. — Lectotype, ® (here desig-

nated; with an unpublished lectotype label written by

Princis, 1968), (abdomen missing), Ins. Philipp., no

exact locality, (Philippine Islands) (with handwritten

‘Pseudothyrsocera lugubris (Stal), K. Princis, 1968’ on

back of his lectotype label); in NRss.

Redescription. — Female: Head exposed, interocu-

lar width greater than distance between ocellar spots

and antennal sockets. Pronotum suboval, widest be-

hind middle (fig. 60). Tegmina and wings (damaged)

fully developed, former with longitudinal discoidal

sectors. Front femur Type B, with four large proximal

spines, pulvilli on four proximal tarsomeres, tarsal

claws symmetrical, simple, arolia present (only one

front leg present, all others missing). All but first two

abdominal terga missing.

Colour. — Head black, ocellar spots and genae yel-

low, with two small red dots between antennal sock-

ets. Pronotal disk black completely surrounded by a

broad yellow band, and with one round and two

smaller red maculae on each side within the dark mar-

gins of the disk anteriorly (fig. 60). Only the first two

abdominal terga are present and these are dark brown

without yellow lateral corners. Legs dark brown, out-

er margins of coxae whitish.

Male: Unknown.

Measurements (mm). — Pronotum length X

width, 3.7 X 4.3; tegmen length, 11.8; interocular

width, 1.1.

Remarks. — Shelford (1908a: 5) synonymized cir-

cumclusa with circumcincta. Princis (1969: 955) listed

circumclusa as a synonym of lugubris, perhaps because

of Shelford's synonymy and because Princis believed

that circumcincta is conspecific with lugubris.

However, none of the other species of

Pseudothyrsocera have the small red markings on the

pronotum present in circumclusa and I am listing it

separately until the male is discovered.

Pseudothyrsocera henrici Hanitsch

(figs. 61-67)

Pseudothyrsocera henrici Hanitsch, 1935:

Bruijning 1947: 226; Princis 1969: 954.

17 (male). —

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0.5 mm

Figs. 61-67. Pseudothyrsocera henrici Hanitsch, male holotype: 61, head and maxillary palps; 62, pronotum; 63, hind wing;

64, supraanal plate and paraprocts; 65, seventh abdominal tergum; 66, ninth abdominal tergum and laterotergites (dorsal; the

laterotergites are turned under the tergum); 67, subgenital plate and genitalia (dorsal).

Specimen examined. — Holotype, & (terminalia

slide 5), Latimondjong Mountains, Uru, Sulawesi,

800 m, viii-ix.1930, G. Heinrich; in MNHG.

Redescription. — Male: Head exposed, interocular

space about the same as the width between antennal

sockets; fifth maxillary palpomere enlarged and

longer than the fourth (fig. 61); antennae not

plumose (Hanitsch incorrectly described them as

plumose). Pronotum suboval (fig. 62). Tegmina and

wings fully developed, extending beyond end of ab-

domen, the former with simple radial vein, and longi-

tudinal discoidal sectors. Hind wing with simple radi-

al and media veins, cubitus vein with two complete

and no distinct incomplete branches (one small,

curved, venule), apical triangle absent (fig. 63). Front

femur Type A, (as in fig. 12); pulvilli on four proxi-

mal tarsomeres, tarsal claws simple, symmetrical, aro-

lia present. First abdominal tergum unspecialized.

Seventh abdominal tergum with a pair of shallow de-

pressions anteromedially (hidden by the overlapping

sixth tergum) separated by a clear, nonsetose region

(fig. 65). Ninth laterotergites weakly dissimilar, the

ventral margin of the left one acute at the posterior

end, the same margin of the right plate rounded (fig.

66). Supraanal plate produced, symmetrical, hind

margin with concave sides, apex shallowly indented;

intercercal processes absent; paraprocts dissimilar, the

right one with a small spinelike process basally (fig.

64). Subgenital plate, trigonal, asymmetrical, the left

side roundly thickened and setose, without a process

(also absent on the right side) near the middle; locat-

ed apically is a long, slender, sclerotized, articulated,

erect, spinelike left style; well separated to the right

and lying against the surface of the plate, is the small-

er, dissimilar right style which is dark on its proximal

half and becomes membranous and colourless distally

(this style is completely hidden in the pinned speci-

men; Hanitsch incorrectly stated that styles are absent

even though the large left one is exposed in the

pinned specimen.) (fig. 67). Genitalia as in fig. 67:

hook on the left side with a small preapical structure;

median phallomere a simple, apically blunt rod; right

phallomere much smaller than the genital hook, con-

sisting of three sclerites, one of them a small cleft.

Colour. — Head with occiput and vertex black,

shading into dark reddish brown (fig. 61); palps and

antennae dark. Pronotum brownish yellow, lateral

borders subhyaline, hind border whitish (fig. 62).

Tegmina hyaline, reddish brown, weakly infuscated

in the apical region. Wings darkly fuscated, region

between the media and cubitus veins mostly colour-

less (fig. 63). Abdominal terga dark brown, medial

gland area on segment seven pale (fig. 65).

Abdominal sterna light brownish. Legs dark reddish

brown, coxae weakly infuscated. Cerci dark brown on

both surfaces, apical segment pale.

Female: Unknown.

Measurements (mm). — Length, 17.0; pronotum

length X width, 2.9 X 3.2; tegmen length, 10.0; in-

terocular width, 0.8.

Remarks. — Hanitsch (1935) stated that henrici is

closely allied to xanthophila, the other

Pseudothyrsocera from Sulawesi, but differs in colour.

However, although there are some similarities be-

tween the two taxa, (e.g. Type A, front femur, only

the seventh abdominal tergum specialized, widely

separated styles), I place henrici in the lugubris

ROTH: Blattaria, Blattellidae, Blattellinae

species-group, and xanthophila in the fulva-species-

group because of differences in the shapes of their

subgenital plates.

Pseudothyrsocera sinensis (Walker) comb. n.

Ischnoptera sinensis Walker, 1869: 148 (male).

Phyllodromia sinensis (Walker). — Shelford 1908a: 13.

Periplaneta apicalis Shiraki (in part), 1931: 181 (female not

male as indicated; female = Hebardina sp.). — Asahina

19794339:

Symploce sinensis (Walker). — Bey-Bienko 1950: 155, figs.

54-56 (male & female); Princis 1959: 134 (male).

Episymploce sinensis (Walker). — Asahina 1979: 339, figs. 1A-

C, 3A, 4A-E (male & female); Roth 1987c, 130, figs. 1A-

I, 2A-C (redescriptions).

The shape and markings of the tegmina and wings

(Asahina 1979, fig. 3A; Roth 1987c, figs. 1H, I) are

characteristic of Pseudothyrsocera. The trigonal shape

of the subgenital plate and styles (Roth 1987c, fig.

1C) place the species in the /ugubris-species-group.

Distribution of Pseudothyrsocera by species:

circumcincta: Philippines

circumclusa: Philippines

fulva: Sumatra

henrict: Sulawesi

lugubris Philippines

montana: Sarawak

perkinsi: Malaya

rectangularitervittata: Borneo; Sarawak

rubronigra: Malaya

rufiventris: Philippines

scutigera: Sabah; Sarawak

signata: Philippines

sinensis. Hong Kong; Taiwan

xanthophila: Sulawesi; Sumatra

Distribution by country or regions:

Borneo: rectangularitervittata

Hong Kong: sinensis

Malaya: perkinsi; rubronigra

Philippines: circumcincta; circumclusa; lugubris;

rufiventris; signata

Sabah: scutigera

Sarawak: montana; rectangularitervittata, scutigera

Sulawesi: henrici; xanthophila;

Sumatra: xanthophila

Taiwan: sinensis

Genus Haplosymploce Hanitsch

Haplosymploce Hanitsch, 1933b: 236. — Hanitsch 1933a:

128; Princis 1951: 56 (footnote: selected /schnoptera ni-

gra Hanitsch as the type species).

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0.5 mm

Figs. 68-73. Haplosymploce spp. 68-71. H. pica (Walker): 68, pronotum and proximal part of the tegmina (male lectotype);

69, hind wing (® paralectotype from Sumatra); 70, 71, male paralectotype from Singapore: 70, supraanal plate, intercercal

processes, and paraprocts (ventral); 71, subgenital plate and genitalia (dorsal). 72, 73, H. ruficollis (Shelford), male holotype:

72, supraanal plate, intercercal processes, and paraprocts (ventral); 73, subgenital plate and genitalia (dorsal).

Princis (1969: 875) listed six species of

Haplosymploce, namely: nigra Hanitsch, montis

Shelford, reversa Walker, guttifera Walker, walkeri

Princis, and curta Hanitsch. Symploce bicolorata Roth

is a junior synonym of H. montis, and Symploce fer-

ruginea Roth is a junior synonym of Haplosymploce

nigra.

Pseudothyrsocera pica Walker, P. ruficollis Shelford,

P. moultoni Hanitsch, and P. andamanica (Princis)

are transferred to Haplosymploce. Blatta guttifera

Walker belongs in the genus Beybienkoa Roth, and

Beybienkoa papuensis (Roth) originally described as a

Symploce is a junior synonym of Beybienkoa guttifera.

Haplosymploce walkeri Princis (= Blatta elegans

Walker) is Hemithyrsocera walkeri. | have not seen

Haplosymploce curta Hanitsch (male unknown) but

am retaining it in that genus until the male is de-

scribed.

Rediagnosis. — Hanitsch's diagnosis of this genus

is: ‘Front femur armed after Type A; radial vein both

of tegmina and wings simple, ulnar [= cubitus] vein

sigmoid, with several complete and one or more in-

complete branches, no apical triangle’. A study of the

species of Haplosymploce listed in Princis's catalogue

shows that variation of the above characters makes it

difficult to place some of the species in this genus and

a rediagnosis follows: Antennae not plumose.

Tegmina and wings fully developed, the former with

simple radial vein and longitudinal or weakly oblique

discoidal sectors. Hind wing with simple radial vein,

cubitus vein oblique, or subsigmoid, with three to

five complete and zero to four incomplete branches,

apical triangle absent. Front femur variable: Type A

(fig. 85), or B (figs. 81, 84), (more than one type of

femur may occur in the same species); pulvilli on four

proximal tarsomeres, tarsal claws simple, symmetri-

cal, arolia small. Male: First and seventh abdominal

terga specialized (figs. 87, 88), or only the first seg-

ment with a medial gland. Paraprocts dissimilar, large

intercercal processes present on the ventral surface of

the subgenital plate near the bases of the cerci (e.g.,

figs. 70, 74, 82, 90 ). Subgenital plate asymmetrical,

generally similar in shape in all the species (e.g., figs.

71, 73, 83, 91); styles small, similar in shape, equal or

slightly different in length, cylindrical, (figs. 71, 73,

83, 91, 100). Genital hook on the left side, with a

preapical incision; median phallomere a stout or slen-

der rod, sometimes with a setal patch near the acute

apex (figs. 80, 83, 100).

Remarks. — The shapes of the male subgenital plate

and styles, are similar in all species of Haplosymploce.

I place eight of the ten species of Haplosymploce,

whose males are known, in the following two species

groups:

ROTH: Blattaria, Blattellidae, Blattellinae

1. pica-species-group. — Only the first abdominal

tergum specialized. Front femur Type A. Cubitus

vein of hind wing with three or four complete and no

incomplete branches. Species: bicolor; moultoni; pica;

ruficollis.

2. nigra-species-group. — First and seventh abdom-

inal terga specialized. Front femur Type A or B.

Cubitus vein of hind wing with four or five complete

and two or three incomplete branches. Species: an-

damanica; montis (= bicolorata); nigra (= ferruginea);

reversa.

Key to known males of Haplosymploce

1. Only the first abdominal tergum specialized.

Front femur Type A. Cubitus vein of the hind

wing with three or four complete and no in-

complete branches. (pica-species-group) ..2

= Abdominal terga one and seven specialized.

Front femur Type A or B. Cubitus vein of hind

wing with four or five complete and two or

three incomplete branches. (nigra-species-

group)

Tegmina yellowish, with the basal and apical

regions black. Wings with base and apex black

(MESZ) ONE RL Re bicolor

+ Tegmina with pale spots in the subcostal zone

and bases of the anal zone (figs. 68) ...... 3

Pronotumtblack eee nein. ieee pica

— Pronotum red

Pronotal disk with a solid dark macula sur-

rounded by pale orange-yellow (fig. 93)

ROMEU AGL OURS SE ne DEAN rene andamanica

Pronotal disk completely dark

Tegmina pale yellowish-white, transparent,

strikingly lighter than the dark pronotum (fig.

8A in Roth 1985b) .... montis (= bicolorata)

Tegmina and pronotum dark

Femora pale with their apices dark brown (figs.

84-86). Supraanal plate, paraprocts, and inter-

gercaliprocessestas insfig:19 ON. iti reversa

= Femora completely dark (as in fig. 81).

Supraanal plate, paraprocts, and intercercal

processes as in fig. 82, and figs. 10F, G, in

Roth 985 ba BANNER nigra (= ferruginea)

pica-species-group

Haplosymploce pica (Walker) comb. n.

(figs. 68-71)

Pseudomops pica Walker, 1868: 213 (male).

Thyrsocera pica (Walker). — Kirby 1904: 78.

Pseudothyrsocera pica (Walker). — Shelford, 1906: 250, 1907:

488; 1908a: 5, pl.1, fig. 3 [habitus]; Hanitsch 1915: 34,

pl. 7, fig. 37 (male) [habitus], 1919: 72, 1923: 463;

Hebard 1929: 10; Bruijning 1948: 95.

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Figs. 74-77. Haplosymploce spp. 74-76, H. bicolor (Shelford), male holotype: 74, supraanal plate, intercercal processes, and

paraprocts (ventral); 75, subgenital plate and genitalia (dorsal); 76, left tegmen (top) and hind wing (bottom). 77, H. moul-

toni (Hanitsch), female holotype, left tegmen (top) and hind wing (bottom).

Specimens examined. — Lectotype, d (here desig-

nated), (terminalia slide 296), Singapore, Wallace,

Mr. Saunder's collection; Type Orth. 47°/, in HECO.

Paralectotypes. Singapore. HECO: 1d (terminalia

slide 297), same data as lectotype, Type Orth. 47'/..

Sumatra. Heco. 19, Wallace, in Mr. Saunder's

Collection, Type Orth. 47/..

Redescription. — Male: Head with interocular

space greater than the width between antennal sock-

ets; fifth maxillary palpomere longer than the fourth;

antennae with proximal and distal segments not

plumose, the antennomeres between them thickened

and densely plumose. Pronotum subparabolic,

smooth, hind margin rounded (fig. 68). Tegmina and

wings fully developed extending beyond the end of

the abdomen, the former with longitudinal discoidal

sectors. Hind wing with simple radial vein, cubitus

vein with three or four complete and no incomplete

branches, apical triangle absent (fig. 69). Front femur

Type A, with four long proximal spines succeeded by

a row of short stout equal spines (stouter than spin-

ules), pulvilli on four proximal tarsomeres, tarsal

claws symmetrical, simple, arolia small. First abdom-

inal tergum with a median setose gland. Seventh ab-

dominal tergum unspecialized. Supraanal plate sym-

metrical, transverse, median region broadly produced

and setose; intercercal processes large, paraprocts dis-

similar, the left one sclerotized with three terminal

spines (fig. 70). Subgenital plate asymmetrical, left

side of hind margin shallowly excavated, with a pair

of cylindrical styles, the left one larger (fig. 71).

Genitalia as in fig. 71: hook on the left side with a

preapical incision and with a minutely spicular clear

membrane; median phallomere a curved, apically

acute rod; right phallomere consisting of three scle-

rites one a cleft, another with four or seven small

spines.

Female: Front femur with five or six long proximal

spines succeeded by a row of slender spinules and ter-

minating in three heavy spines (Type B.). Supraanal

plate trigonal, sides weakly concave, apex rounded,

reaching hind margin of subgenital plate.

Colour. — Head black, shiny; maxillary palpomeres

and antennae black. Pronotum black. Tegmina black

with subcostal (mediastine) area white, and with a

large, white macula at the base of the anal field (fig.

68). Hind wing darkly infuscated except for a colour-

less proximal region in the anterior field (fig. 69).

Abdomen black. Coxae with basal region black, dis-

tally and part of the outer border whitish; femora, tib-

iae and tarsi black.

Measurements (mm) (@ in parentheses). Length,

9.8 (11.2); pronotum length X width, 3.3-3.6 X

3.6-3.9 (3.9 X 4.2); tegmen length, 11.5-12.0

(13.3); interocular width, 1.1-1.2 (1.4).

Remarks. — The subgenital plate, styles and geni-

talia are similar to those of Pseudothyrsocera ruficollis

(see remarks below, under that species).

Haplosymploce ruficollis (Shelford) comb. n.

(figs. 72-73)

Pseudothyrsocera ruficollis Shelford, 1906: 251, pl. 14, fig. 6

(habitus) (male). — Shelford 1908a: 5; Hanitsch 1915:

35, pl. 7, fig. 36 (habitus), 1923: 463; Hebard 1929: 10;

Bruijning 1948: 95.

Specimen examined. — Holotype, ¢ (terminalia

slide 298), Penang [Pinang, Malaysia], Cantor; Type

Orth. 48 in HECO.

Redescription. — Male: Head with interocular

space greater than the distance between antennal

sockets; antennae missing. [When Shelford described

the species, the antennae were mutillated. He stated

that when intact specimens are found the basal half of

the antennae probably would be plumose, as in pica.

Hanitsch (1915: 35) did see two additional speci-

mens from Sarawak and Selangor, and found that the

antennae are plumose]. Pronotum subparabolic.

Tegmina and wings fully developed extending be-

yond end of abdomen, the former with longitudinal

discoidal sectors. Hind wing with simple radial and

media veins, cubitus vein with three complete and no

incomplete branches, apical triangle absent. Legs

missing. First abdominal tergum with a setose gland.

Seventh abdominal tergum unspecialized. Supraanal

plate with the median region produced, intercercal

processes large, similar, paraprocts dissimilar, the left

ROTH: Blattaria, Blattellidae, Blattellinae

one with three terminal spines (fig. 72). Subgenital

plate asymmetrical, the hind margin on the left side

weakly excavated, with a pair of similar, widely sepa-

rated styles (fig. 73). Genitalia as in fig. 73: hook on

the left side with a preapical incision; median phal-

lomere a slender, curved, apically acute rod; right

phallomere consisting of four sclerites one of which is

a small cleft.

Colour. — Head and pronotum red; maxillary palpi

and antennae black. Tegmina black with a yellowish-

white macula on the subcostal (mediastinal) region

and at the base of the anal field. Wings dark brown,

pale in the proximal region, veins mostly dark (yel-

lowish at the bases of the veins in the anterior and

posterior fields). Coxae with their outer borders and

distal ends yellowish-white.

Female: Unknown.

Measurements (mm). — Length, 12.0; pronotum

length X width, 3.7 X 4.2; tegmen length, 13.2; in-

terocular width, 1.3.

Remarks. — Shelford (1906: 251) was correct in

saying that ruficollis is most closely allied to pica, a

conclusion supported by the strong similarity in the

two species of the subgenital plate, styles, genitalia,

supraanal plate, paraprocts, and intercercal processes

(cf. figs 70, 71 with 72, 73). In fact, ruficollis may

eventually prove to be a colour variant of pica. The

tegminal colour and markings are similar (the medial

macula is much larger and more distinct in moultoni),

but the colour differences of the head and pronotum

are strikingly different and readily separate the two

taxa.

Haplosymploce bicolor (Shelford) comb. n.

(figs. 74-76)

Pseudothyrsocera bicolor Shelford, 1909: 612 (male). —

Hanitsch 1915: 36 (male), 1923: 463; Hebard 1929: 10;

Bruijning 1948: 94; Princis 1969: 955.

Specimen examined. — Holotype, d (terminalia

slide 300), Mt. Matang, N.W. Borneo [Sarawak],

vi.1900; Type Orth. 50 in HECO.

Redescription. — Male: Head with interocular

space greater than the distance between the antennal

sockets; about the basal two thirds of the antennae

densely plumose. Pronotum suboval. Tegmina and

wings fully developed extending beyond end of ab-

domen, the former with longitudinal discoidal sectors

(fig. 76, top). Hind wing with simple radial and me-

dia veins, cubitus vein with four complete (not five as

stated by Shelford) and no incomplete branches, api-

cal triangle absent (fig. 76, bottom). Front femur

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 139, 1996

Figs. 78-83. Haplosymploce spp. males. 78-80, H. montis (Shelford), holotype: 78, seventh abdominal tergum; 79, subgenital

plate, intercercal processes, and paraprocts (ventral); 80, genitalia (dorsal). 81-83, Haplosymploce nigra Hanitsch: 81, (lecto-

type) front femur (anterior surface); 82, 83 (paralectotype): 82, supraanal plate, intercercal processes, and paraprocts (ventral;

a, left intercercal process; b, left paraproct); 83, subgenital plate and genitalia (dorsal).

Type A,, or A, (the first of the four terminal spines is

only slightly longer than the one preceding it); pulvil-

li on four proximal tarsomeres, tarsal claws symmetri-

cal, simple, arolia small. First abdominal tergum with

a setal specialization. Seventh abdominal tergum un-

specialized. Supraanal plate transverse, symmetrical,

midregion of the hind margin produced, intercercal

processes, large, similar, right and left paraprocts dis-

similar (fig. 74). Subgenital plate almost symmetrical,

styli widely separated (fig. 75; Shelford (1909) stated

that the left style is much longer than the right one;

however, the slide preparation shows that the left

style is damaged and is missing the distal region.).

Genitalia as in fig. 75: hook on the left side with a

preapical incision; median phallomere a slender, api-

cally acute rod and near the apex, but separated from

it, is a small, dark, dense group of setae; right phal-

lomere consists of three sclerites one of which is a

cleft.

Colour. — Head black; maxillary palpi black; basal

two thirds of the antennae black, the distal third be-

yond the plumose region, yellow. Pronotum black.

Proximal and apical region of the tegmina black, the

intermediate zone yellowish with a small dark spot in

the middle, and with a yellowish spot in the subcostal

region (fig. 76, top). Wings with base and apex black-

ish, remainder hyaline (fig. 76, bottom). Abdominal

terga and sterna black. Legs black except for the cox-

ae whose apical halves and outer margins are yellow-

ish.

Female: Unknown.

Measurements (mm). — Length, 11.5; pronotum

length X width, 4.4 X 5.4; tegmen length, 14.2;

interocular width, 1.4.

Remarks. — The striking colour of the tegmina and

wings readily identify this species.

Haplosymploce moultoni (Hanitsch) comb. n.

(fig. 77)

Pseudothyrsocera moultoni Hanitsch, 1915: 36, pl. 3, fig. 16

(female). — Hanitsch 1923: 463; Hebard 1929: 10;

Bruijning 1948:95; Princis 1969: 955.

Specimen examined. — Holotype 9, Mt. Merinjak,

Sarawak, 2200 feet, 19.v.1914, Moulton; Type Orth.

260 in HECO (The specimen is fragmented and parts

are mounted on cards.)

Redescription. — Female: Head with interocular

space greater than the distance between antennal

sockets; antennae missing (according to Hanitsch, the

antennae are plumose). Pronotum _ parabolic.

Tegmina and wings fully developed extending be-

yond end of abdomen, the former with longitudinal

discoidal sectors (fig. 77, top). Hind wing with sim-

ple radial and media veins, the cubitus vein with four

complete and one incomplete branches, apical trian-

gle absent (fig. 77, bottom). Front femur Type A;

pulvilli on four proximal tarsomeres, tarsal claws sym-

metrical, simple, arolia small. Supraanal plate dam-

aged.

ROTH: Blattaria, Blattellidae, Blattellinae

Colour. — Head reddish brown with a large black

macula on the genae; antennae black, except for a few

distal white segments, terminal antennomeres black

(from Hanitsch). Tegmina reddish brown with three

large maculae, one occupying almost the entire anal

field, a second medially, and the third at the apex (fig.

77, top). Hind wing with apical region of the anteri-

or field dark brown and black, remaining region yel-

lowish; posterior field infuscated, darker basad and

posteriorly (fig. 77, bottom). Abdomen reddish

brown. Legs with coxae black basally, yellowish dis-

tally and along the outer border, femora black with a

yellowish band along the ventral margin, tibiae, tarsi,

and cerci black.

Male: Unknown.

Measurements (mm). — Length, 12.5; pronotum

length X width, 4.6 X 5.3; tegmen length, 14.3;

interocular width, 1.5.

Remarks. — Based on the colour and pattern of the

tegmina and wings, moultoni and bicolor may prove to

be synonyms when the male of the former is found,

and larger series are examined to determine the extent

of colour variation.

Haplosymploce montis (Shelford)

(figs. 78-80)

Ischnoptera montis Shelford, 1906: 266, pl. XVI, fig. 10

(male). —Shelford 1908a: 7; Hanitsch, 1915: 39, 1923:

463.

Parasymploce montis (Shelford). — Hebard 1929: 10.

Haplosymploce montis (Shelford). — Bruijning 1948: 76, figs.

13, 37; Princis 1950a: 178.

Symploce bicolorata Roth, 1985b: 148, figs. 8, 9 (male not fe-

male, see remarks). Syn. n.

Specimens examined. — Holotype d of Jschnoptera

montis Shelford (terminalia slide 258), N.W. Borneo,

Sarawak, Mt. Matang nr Kuching, 2.v.1902, pres.

1905 by the Sarawak Museum; Type Orth. 57, in

HECO. Additional specimens. — Sabah. RMNH: N.

Borneo, 12.5 km S. Nabawan, Kg. Pamuntaria, nr.

river, 116.27E 5.02N, 400 m, 16, 16.xi. 1987, J.

Huisman & R. de Jong. BPBM: Tawau, Quoin Hill,

Cocoa Res. Sta., light trap, d holotype No. 13472

(wing on slide 214), 17.ix.1962, and 1d paratype

(terminalia slide 213), 3.ix.1962, Y. Hirashima, of

Symploce bicolorata Roth; Tawau, Quoin Hill, Cocoa

Res. Sta., Malaise, 225 m, 26, 20.ix.1962, Y.

Hirashima, in jungle, light trap, 26, 3-7.vii.1962, H.

Holtmann; Forest Camp, 19 km north of Kalabakan,

IG, MIDP EO mis le sl C2 RICE

Kuncheria. zits: Malaysia, Sabah, Sipitang,

Mendolong, 1d, 25.xi.1987, AlL, 1d, 5.iv.1988,

18, 30.iv.1988, 16, 6.1v.1988, 34, 5.v.1988, 16,

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 84-92. Haplosymploce reversa (Walker). 84-85, front femora (anterior surfaces): male and female respectively, from

Kedah Peak; 86-91, male holotype: 86, front femur (anterior surface); 87, first abdominal tergum (pinned specimen); 88, sev-

enth abdominal tergum (slide preparation); 89, hind wing; 90, supraanal plate, intercercal processes, and paraprocts (ventral);

91, subgenital plate and genitalia (dorsal); 92, female from Kuala Lumpur, supraanal plate (dorsal).

i11.1989, S. Adebratt. Two retained at mczc. Malaya.

BPBM: Malay Pen., S.E. Pahang, Rompin Mining

Co., Railway Track, 50 km, 1d, 9.iv.1961, KJ.

Kuncheria. Sumatra. RMNH: Tanangtalu, 14,

v.1915, E. Jacobson. ZMAN: Bandar Baru, 850 m,

Ld, 11.1921, J.B. Corporaal (Sumatran specimens re-

ported by Bruijning 1948: 76).

Remarks. — The distal parts of the supraanal and

subgenital plates (and styles) are missing from the

type of montis but the intercercal processes and para-

procts (fig. 79), genital phallomeres (fig. 80), and the

specializations on the seventh (fig. 78) and first ab-

dominal terga are intact. The front femur is Type A,

but the row of preterminal spines are small and simi-

lar in length but stouter than piliform spinules; other

specimens of montis have Type B, or intermediate be-

tween A, and B, (a mixture of piliform spinules and

stout spines of the same length). These structures, as

well as the distinctive colour pattern of dark reddish

brown pronotum and pale yellowish-white semi-

transparent tegmina (although there is some variation

in the pronotal colour, some being lighter than oth-

ers) are so similar to those of Symploce bicolorata Roth

(1985c: fig. 8) that I am synonymizing the two

species. The measurements (mm) of the present male

specimens are as follows: Length, 11.0-16.0; prono-

tum length X width, 3.5-4.0 X 4.1-4.6; tegmen

length, 14.0-15.0; interocular width, 0.8.

The female paratype of S. bicolorata from Pelawan

besar, Kalimantan, is not Haplosymploce montis but is

possibly a species of Symploce.

Haplosymploce nigra (Hanitsch)

(figs. 81-83)

Ischnoptera nigra Hanitsch, 1928: 15, pl. 1, fig. 6 (male).

Haplosymploce nigra (Hanitsch), 1933b: 237; Bruijning

1948: 76; Princis 1969: 875.

Symploce ferruginea Roth, 1985b: 152, figs. 10, 11 (male and

female). Syn. n.

Specimens examined. — Lectotype, d (here desig-

nated), Mentawe[a]i Island, Siberoet, 15.ix.1924,

ERE Karny; | Type Orth. 3251/2) in ECO:

Paralectotype: Sumatra. HECO: N. Pagi Island, West

Sumatra, Type Orth. 325/, 1d (terminalia slide

256), x.1924, C.B.K. & N.S. Additional specimens.

— West Malaysia. HECO: Pahang, Fraser's Hill, 4000

ft., d holotype (terminalia slide 193) of Symploce fer-

ruginea Roth (Type Orth. 970), 1.vi.1932, H.M.

Pendlebury. Sabah. BPBM: Forest Camp, 19 km north

of Kalabakan, 60 m, light trap, ® paratype of

Symploce ferruginea Roth, 29.x.1962, Y. Hirashima.

zits: Malaysia, Sabah, Sipitang, Mendolong, 19,

29.x11.1987, AIL, 19, iii.1988, 19, 5.iv.1988, 1988,

ROTH: Blattaria, Blattellidae, Blattellinae

19, 6.iv.1988, 19, 5.v.1988, W5L, 12, 19.iv.1988,

S. Adebratt. Two retained in Mczc. RMNH: Malaysia-

SE. Sabah, nr. Danum Valley Field C., c. 150 m,

WO Mal. trap 5, 19, 20.vi. —12.vii.1987, C. v.

Achterberg & D. Kennedy; N. Borneo, Sabah, 24 km

on rd. Keningau-Kimanis (N. side), 116.03E 5.27N,

1350 m, 12, 19.xi.1987, J. Huisman & R. de Jong.

Redescription. — Male: Head hidden, interocular

space about the same as the distance between anten-

nal sockets. Pronotum suboval. Tegmina and wings

fully developed extending beyond end of abdomen,

the former with simple radial vein and weakly oblique

discoidal sectors. Hind wing with simple radial vein,

cubitus oblique, weakly sigmoid, with four or five

complete and two or three incomplete branches, api-

cal triangle absent (Hanitsch 1928: pl.1, fig. 6). Front

femur with eight or nine stout proximal spines, the

basal five or six about equal in length, the others

much smaller, succeeded by five to eight piliform

spinules and three large terminal spines (fig. 81)

[Type B,; Hanitsch (1928: 15) stated that the anteri-

or edge of the front femur of nigra has ‘... 4 large

spines, followed by 5 shorter ones (type A).’; actually

the stout spines are succeeded by a short row of pili-

form spinules which classify the femur as Type B);

pulvilli on four proximal tarsomeres, tarsal claws sym-

metrical, simple, arolia small. First abdominal tergum

with a large depression and a dense group of setae me-

dially. Seventh abdominal tergum with a depression

and large medial tuft of setae. Supranal plate, para-

procts, and intercercal processes as in fig. 82.

Subgenital plate and genitalia as in fig. 83.

Female (illustrated in Roth, 1985b, figs. 11A-C).

Front femur Type A,. Cubitus vein of hind wing

oblique or subsigmoid, with six or seven complete

and three incomplete branches, apical triangle absent.

Supraanal plate trigonal.

Colour. — Varies from dark reddish brown to

blackish brown. Legs (femora, tibiae, tarsi), and cerci

dark. Wings dark brown.

Measurements (mm) ( in parentheses). — Length,

10.2-10.7 (12.0-18.0); pronotum length X width,

3.3-3.4 X 4.0-4.1 (4.3-4.7 X 5.2-5.8); tegmen

length, 11.5-12.5 (16.5-19.5); interocular width, 0.9

(0.9-1.0).

Remarks. — I am synonymizing ferruginea with ni-

gra because the differences between their males are

minor. Similarities between the intercercal processes,

paraprocts, and genitalia suggest a close relationship

between Haplosymploce nigra and H. montis (cf., figs.

79, 80, with 82 and 83).

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

<<< S

Coma

Figs. 93-101. Haplosymploce andamanica (Princis), types. 93-100, male holotype: 93, pronotum; 94, subgenital plate (ven-

tral); 95, supraanal plate, paraprocts, and intercercal processes (ventral); 96, glandular region on first abdominal tergum; 97,

gland on seventh abdominal tergum (slide preparation); 98, hind wing; 99, abdominal terga seven to ten (supraanal plate);

100, subgenital plate and genitalia (dorsal); 101, female paratype, supraanal plate (dorsal).

Haplosymploce reversa (Walker)

(figs. 84-92)

Ischnoptera reversa Walker, 1869: 147 (male). — Kirby 1904:

81; Shelford 1907: 489, 1908a: 7; Hanitsch 1915: 37,

1923: 398.

Parasymploce reversa (Walker). — Hebard 1929: 10.

Haplosymploce reversa (Walker). — Bruijning 1948: 78, fig.

38; Princis 1969: 875.

Specimens examined. — Holotype, d (terminalia

slide 257), Singapore, Wallace, W.W. Saunders coll.,

purch. and pres. '73 by Mrs. F.W. Hope; Type Orth.

54 in HECO [Mr. I. Lansbury informed me that the

holotype was almost totally destroyed in the return

mail; he placed the remnants in a ‘container’ and at-

tached it to the type pin]. Additional specimens. —

West Malaysia. HECO: Kuala Lumpur, Malay Pen.,

19, ex coll. Dept. Agric.; Kedah Peak, 3300 ft., 1d,

EAO DEE IL Sp ZEIGE

Redescription. — Male: Interocular space slightly

less than distance between antennal sockets.

Pronotum subparabolic. Tegmina and wings fully de-

veloped extending beyond end of abdomen, the for-

mer with simple radial vein and longitudinal dis-

coidal sectors. Hind wing with simple radial vein,

cubitus vein with five complete and three or four in-

complete branches, apical triangle absent (fig. 89),

the triangular region folded when at rest. Front femur

Type A, (fig. 86, holotype; six spines preceding the

three long terminal spines are small and about the

same length but are stouter than piliform spinules)

(the femur of a male from Medan, Sumatra, also is

Type A,, see Bruijning 1948: 78, fig. 38), or Type B,

with nine stout proximal spines decreasing in length

distad, succeeded by six piliform spinules and termi-

nating in three large spines (fig. 84, d from Kedah

Peak, West Malaysia); pulvilli on four proximal tar-

someres, tarsal claws symmetrical, simple, arolia

small. First abdominal tergum with a pair of deep fos-

sae divided medially by a longitudinal ridge bearing a

dense group of setae and with setae along the anterior

margins of the fossae (fig. 87). Seventh abdominal

tergum with two, almost contiguous groups of setae

in a shallow depression (fig. 88; this gland may be

hidden by the overlapping sixth tergum). Supraanal

plate symmetrical hind margin convexly rounded, en-

tire; intercercal processes weakly dissimilar, spinelike,

right and left paraprocts dissimilar, the right one with

a curved, spinelike process and a dark sclerite bearing

three small, dark, spines (fig. 90). Subgenital plate

asymmetrical with dissimilar styles, the left one more

slender (fig. 91). Genitalia as in fig. 91: hook on the

left side with a preapical incision; also on the left side

is a setose sclerite; median phallomere a dark, curved,

apically acute rod with a small setose sclerite near the

apex; right phallomere consisting of at least two scle-

rites one of which is a reduced cleft.

Female: Interocular space about the same as dis-

tance between antennal sockets. Cubitus vein of hind

wing with five or six complete and two incomplete

branches, apical triangle absent. Front femur Type A,,

(sometimes the last four or five spines are longer than

the preceding ones thus appearing to be Type A,, or

A,, as in fig. 85). Supraanal plate subtrigonal, sides of

the hind margin concave, apex rounded (fig. 92).

Colour. — Head reddish brown without markings;

proximal segments of antennae dark brown, remain-

der light brown; segments four and five of maxillary

palpi and last segment of labial palpi dark brown,

other segments pale. Pronotum reddish brown, with-

out markings, lateral borders opaque. Tegmina hya-

line, reddish brown. Hind wing with costal vein re-

gion yellowish, its margin darkened (fig. 89), veins

yellowish. Abdominal terga and sterna reddish

brown. Basal half of dorsal surface of the cerci light

ROTH: Blattaria, Blattellidae, Blattellinae

brown, distal half and ventral surface dark brown.

Coxae and most of the femoral surface reddish

brown, apexes of the femora (figs. 84-86), tibiae and

tarsi dark brown.

Measurements (mm) (® in parentheses). — Length,

13.5 (13.5-14.7); pronotum length X width, 3.6-4.0

X 4.3-4.6 (4.0-4.1 X 4.6-5.0); interocular width,

OFZO)

Remarks. — The shape of the female's subgenital

plate of reversa resembles that of the female of

Haplosymploce nigra (= ferruginea; see Roth, 1985b:

nee)

Haplosymploce andamanica (Princis) comb. n.

(figs. 93-101)

Mopsera andamanica Princis, 1951: 59, pl. 6, fig. 64 (male

& female).

Pseudothyrsocera andamanica (Princis). — Princis 1969: 956.

Specimens examined. — Holotype d (terminalia

slide 111), Port Blair, Andaman Islands, Roeppstorff;

in zMuc. Paratype. Andaman Islands. zmuc: 19,

same data as holotype.

Redescription. — Male: Head surface rough and

with small setae, interocular width the same as the

distance between antennal sockets; fifth maxillary

palpomere distinctly longer than the fourth.

Pronotum suboval, widest behind the middle, surface

densely punctate and punctulate, and covered with

small setae (fig. 93). Tegmina and wings fully devel-

oped extending beyond the end of the abdomen, the

former with longitudinal discoidal sectors. Hind

wings with subcosta extending to about the middle of

the anterior margin, costal veins thickened, not

clubbed, radial and media veins simple, cubitus vein

weakly curved, with four complete and two incom-

plete branches, apical triangle absent (fig. 98). Front

femur Type B, with 7 large proximal spines; pulvilli

on four proximal tarsomeres, tarsal claws simple,

symmetrical, arolia very small. First abdominal ter-

gum medially with a pair of raised ridges and a deep

fossa, and two groups of setae originating along the

anterior margin and overlapping the fossae; a pair of

filamentous membranes, apparently arising from the

posterior margin of the metanotum extends to the

first abdominal tergum (fig. 96). Seventh abdominal

tergum with a deep medial fossa in which are a dense,

group of curved setae (figs. 97). Supraanal plate ap-

pears to be trapezoidal, in the pinned specimen, be-

cause the hind margin is deflexed ventrad (fig. 99); in

the slide preparation the hind margin is convexly

rounded (fig. 95) with long setae along the hind mar-

gin; a pair of similar intercercal processes are spinelike

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

0.5 mm

Figs. 102-104. Beybienkoa guttifera (Walker). 102-103, male holotype: 102, subgenital plate and genitalia (dorsal; the left

hooklike phallomere and right style are missing); 103, supraanal plate and paraprocts (ventral); 104, male subgenital plate and

genitalia, from near Mt Tozer, Queensland, Australia (dorsal) (from Roth, 1991, fig. 64A, as Beybienkoa papuensis).

and have a small spine apically; right and left para-

procts are dissimlar, each consisting of a stout curved

sclerite the right one bearing seven stout spines api-

cally and the left one with three apical spines (fig. 95).

Subgenital plate asymmetrical, the hind margin to the

left of the midline excavated, with a pair of dissimilar

styles separated by the width of the excavation, the

right style larger (figs. 94, 100). Genitalia as in fig.

100: hook on the left side with a preapical incision,

and surrounded by a spicular membrane; near the

hook is a dense, dark group of setae; median phal-

lomere a slender, apically acute rod, above it is a dense

group of dark setae near the apex; right phallomere

with three sclerites one of which is a cleft.

100

Female: Interocular width slightly greater than in

male. Front femur Type B.. Supraanal plate strongly

trigonal, apex acute (fig. 101).

Colour. — Head black, only the clypeus and

mouthparts partly brownish; maxillary palpomeres

blackish; basal segments of the antennae blackish, re-

mainder brownish yellow. Pronotal disk with a solid

dark macula competely surrounded by pale orange

yellow (fig. 93). Tegmina dark reddish brown. Hind

wing hyaline, without infuscation, veins brownish.

Abdominal terga with middle segments orange yel-

low, distal terga dark brown. Abdominal sterna with

proximal segments orange yellow, distal segments

blackish brown. (Female hind margin of subgenital

plate orangish). Coxae and legs blackish brown.

Measurements (mm) (2 in parentheses). — Length,

13.5 (12.0); pronotum length X width, 3.4 X 4.3

(3.7 X 4.4); tegmen length, 14.0 (14.0); interocular

width, 0.5 (0.7).

Remarks. — Princis (1969: 956) placed this species

in Pseudothyrsocera. However, the hind wing vena-

tion, subgenital plate, styles, and intercercal process-

es, are similar to those structures in species of

Haplosymploce, and I am transferring andamanica to

this genus.

Haplosymploce curta (Hanitsch)

Symploce curta Hanitsch, 1932c: 264 (female).

Haplosymploce curta (Hanitsch). — Hanitsch 1933a: 128;

Bruijning 1947: 218; Princis 1969: 876.

Holotype (not examined), 2, Mampoegrot

(Mampoe cave), S. Celebes, S. Leefmans, ix.1929; the

type is supposed to be in the Leyden Museum, but ac-

cording to Bruijning (1947: 218) it is not there and

probably is in the collection of the Buitenzorg

Museum (now Museum Bogor).

Description (from Hanitsch 1932c). — Female:

Head exposed, interocular space as wide as the dis-

tance between antennal sockets. Pronotum sub-orbic-

ulate, anterior margin straight to weakly emarginate,

hind margin obtusely produced. Tegmina short,

reaching to the middle of the supraanal plate: 11

costals, the ninth and eleventh forked, the tenth four-

branched; radial vein simple, discoidal sectors longi-

tudinal. Wings as long as the tegmina, subcostal vein

bifurcate; 8 costals, radial and media veins straight,

simple, cubitus vein with three complete and two in-

complete branches, apical triangle absent. Front fe-

mur Type A. Supraanal plate trigonal, keeled.

Colour. — Head dull orange, antennae rusty red-

dish brown. Pronotal disk dull orange, margins dark-

er. Tegmina golden amber. Wings faintly orange,

costal and apical regions dark orange. Cerci dull or-

ange. Abdominal sterna dull reddish orange. Legs dull

brownish yellow.

Male: Unknown.

Measurements (mm). — Total length, 11.0; prono-

tum length X width, 4.0 X 4.3; tegmen length, 8.8.

Remarks. — Until the male is found I am provi-

sionally retaining the species in Haplosymplce because

its cubitus vein of the hind wing has three complete

branches and its front femur is Type A.

? Haplosymploce castanea (Brunner) comb. n.

Phyllodromia castanea Brunner, 1898: 204, pl. 16, fig. 7

ROTH: Blattaria, Blattellidae, Blattellinae

(male). — Kirby 1904: 93; Shelford 1908a: 13; Hanitsch

1923: 402.

? Mopsera castanea (Brunner). — Hebard 1929: 78; Bruijning

1948: 98.

? Pseudothyrsocera castanea (Brunner). — Princis 1969: 956.

Holotype (not examined), 4 , Brunei, Borneo; sup-

posedly in Brunner's collection in NMWA but it is

not there (Kaltenbach, personal communication).

Description (from Brunner). — Male: Reddish

brown. Vertex exposed, the front rusty red brown.

Pronotum elongate, lateral margins pale. Marginal

field of the tegmen brownish yellow. Wing not very

infumate. Cubitus vein of hind wing with four

branches. Legs and abdomen reddish brown.

Measurements (mm). — Body length, 11.0; prono-

tum length X width, 3.0 X 3.3; tegmen length,

10.5.

Remarks. — Brunner's meagre description is not

sufficient to place castanea in Pseudothyrsocera un-

equivocally. I am provisionally placing it in

Haplosymploce because the cubitus vein of the hind

wing has four (?complete) branches.

Distribution of Haplosymploce by species:

andamanica: Andaman Islands

bicolor. Sarawak

? castanea: Brunei

curta: Sulawesi

montis: Sabah; Sarawak; Sumatra

moultoni: Sarawak

nigra: Mentawai Island; Sabah; Sumatra; West

Malaysia

pica: Singapore, Sumatra

reversa: Singapore; West Malaysia

ruficollis. Malaysia

Distribution of Haplosymploce by country or

region:

Andaman Islands: andamanica

Brunei: ? castanea

Malaysia: ruficollis

Mentawai Islands: nigra

Sabah: montis, nigra

Sarawak: bicolor, montis, moultoni

Singapore: pica; reversa

Sulawesi: ? castanea; curta

Sumatra: nigra; pica

West Malaysia: nigra; reversa

The following two species, Blatta guttifera and

Blatta walkeri, which were listed by Princis (1969:

875, 876) under Haplosymploce belong in two differ-

ent genera and are redescribed here as new combina-

tions:

101

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 105-109. Hemithyrsocera walkeri (Princis), male holotype of Blatta elegans Walker: 105, pronotum; 106, seventh ab-

dominal tergum showing the medial gland and lateral lobes; 107, supraanal plate, subobsolete intercercal processes, and para-

procts (ventral); 108, hind wing; 109, subgenital plate and genitalia (dorsal).

Beybienkoa guttifera (Walker) comb. n.

(figs. 102-104)

Blatta guttifera Walker, 1868: 230 (male).

Phyllodromia guttifera (Walker). — Kirby 1904: 93;

Shelford1907: 493, 1908a: 14.

Haplosymploce guttifera (Walker). — Princis 1969: 875.

Symploce papuensis Roth 1985d: 300, figs. 1A-J (male). Syn.

Beybienkoa papuensis (Roth). — Roth 1991: 656, figs. 64A-C

(male & female).

Specimen examined. — Holotype, d (terminalia

slide 302), Aru Island [Moluccas], Mr. Saunders coll.;

Type Orth. 97 in zeco. [the type is in very poor con-

dition].

102

Remarks. — Unfortunately the hind wings are bad-

ly damaged, the right style and hook left phallomere

are missing from the type (fig. 102). However, this

species so closely agrees with the description of

Symploce papuensis (Roth 1985d: 300) and later its

new combination Beybienkoa papuensis (Roth 1991:

656) (from Papua New Guinea and Queensland,

Australia), that I am synonymizing the two species.

Similarities are found in the pronotum and head

markings (see figs. 1A, B in Roth, 1985d), the

supraanal plate and paraprocts (fig. 103), subgenital

plate and left style (right style broken off); cf. figs.

102 and 104. There is only a minor difference in the

right genital phallomere, one of whose sclerites in

papuensis has four spines (fig. 104), whereas there are

nine in guttifera (fig. 102), but I consider this in-

traspecific variation. The hind wing cubitus vein of

papuensis has two long complete and zero to one in-

complete branches, apical triangle distinctly devel-

oped (fig. 1J in Roth, 1985d).

Hemithyrsocera walkeri (Princis) comb. n.

(figs. 105-109)

Blatta elegans Walker (nec Eschscholtz, 1822) 1868: 226

(male).

Phyllodromia elegans (Walker),- Kirby 1904: 92; Shelford

1907: 492, 493, 1906: 13; Hanitsch 1915: 55, 1923:

463.

Parasymploce elegans (Walker). — Hebard 1929: 10.

Haplosymploce walkeri Princis 1969: 876.

Specimen examined. — Holotype, d (terminalia

slide 303), Sarawak, Wallace, in Mr. Saunder's coll.;

Type Orth. 95 in HECO.

Redescription. — Male: Head with interocular

space distinctly less than the distance between anten-

nal sockets. Pronotum suboval, hind margin pro-

duced (fig. 105). Tegmina and wings fully developed

extending beyond end of abdomen, the former with a

branched radial vein and longitudinal discoidal sec-

tors. Hind wing with simple costal veins, radial vein

branched before the middle, media and cubitus veins

deeply concave, the latter with two complete and no

incomplete branches, apical triangle well developed

(fig. 108). Front femur Type A,; pulvilli on four prox-

imal tarsomeres, tarsal claws symmetrical, apparently

with subobsolete serrations, arolia present. First ab-

dominal tergum unspecialized. Seventh abdominal

tergum with a medial setose gland and dark, well de-

veloped lateral lobes (fig. 106). Supraanal plate hind

margin convexly rounded, entire; intercercal process-

es greatly reduced; right and left paraprocts dissimilar

(fig. 107). Subgenital plate asymmetrical with a small

cylindrical right style, and on the left side a large,

darkly sclerotized, curved structure (possibly a left

style) (fig. 109). Genitalia as in fig. 109: hook on the

left side, without a preapical incision; median phal-

lomere a stout rod with a slender filament arising

from its upper third, right phallomere consisting of

two large sclerites one of which is a cleft structure.

Female: Unknown.

Colour. — Head brownish yellow with a dark band

on the vertex; last maxillary palpomere, and antennae

dark. Pronotum with a pair of large dark bands very

narrowly separated medially, and with two large

brownish yellow maculae on the posterior half, later-

al and anterior borders brownish yellow (fig. 105).

Tegmina reddish brown. Hind wings infuscated,

costal area darker (fig. 108). Legs brownish yellow.

Cerci brownish.

Measurements (mm). — Length, 10.6; pronotum

ROTH: Blattaria, Blattellidae, Blattellinae

length width, 3.1 3.7; tegmen length, 13.0; in-

terocular width, 0.4.

Remarks. — Hebard (1929: 10) listed this species

(and three others) in a table as … elegans. Princis

(1969: 876) assumed he placed them in Parasymploce

because several taxa listed before them were placed in

that genus. If he meant the dots to represent

Parasymploce, he was correct because Parasymploce is a

junior synonym of Hemithyrsocera (Roth 1995: 962).

The gland and lateral lobes of the seventh tergum,

wing venation and distinct apical triangle, subgenital

plate and genitalia of walkeri are typical of

Hemithyrsocera (= Parasymploce), and the species

clearly is not a Haplosymploce as claimed by Princis.

The subobsolete serrations on the tarsal claws is un-

usual for the genus.

New combinations, new species, and new records

of Episymploce

Genus Episymploce Bey-Bienko

Episymploce Bey-Bienko, 1950: 157; Asahina, 1979: 352;

Roth, 1986b,: 355; 1986c: 173; 1987a: 455; 1987b: 111;

1987c: 125; 1987d: 143 (generic revision, new species).

Diagnosis. — Tegmina and wings fully developed

(rarely with reduced tegmina and wings in the fe-

male), the former with longitudinal discoidal sectors.

Hind wing with radial vein usually branched before

or after the midpoint; cubitus weakly or distinctly

curved with one to seven (rarely none) complete

branches, and none to seven incomplete rami, apical

triangle subobsolete or absent (fig. 112).

Anteroventral margin of front femur Type A, B,

(rarely B), or intermediate between the two types;

pulvilli on four proximal tarsomeres, tarsal claws sym-

metrical, simple, arolia present. Male: First abdomi-

nal tergum with or without a gland. Seventh abdom-

inal tergum always specialized (figs. 111, 113).

Supraanal plate often asymmetrical, the hind margin

may be invaginated, excised (fig. 116), sometimes en-

tire, or with a symmetrical or asymmetrical process

(fig. 120); right and left paraprocts dissimilar; inter-

cercal processes absent (figs. 116, 120). Subgenital

plate asymmetrical, often trigonal with the left (figs.

111, 115), or right and left sides setose and incrassate

(the thickened region may vary in length, and some-

times is absent); usually there is a spinelike process on

each side about midway on the lateral margin, these

differing in length on the right and left sides (these

are similar to subgenital plates of the /ugubris-species-

group of Pseudothyrsocera; see fig. 47). Two similar or

dissimilar styles usually occur on the apical end of the

subgenital plate (fig. 118). Genitalia consisting of

103

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

SSS

Figs. 110-118. Episymploce simmonsi sp. n., male paratype: 110, pronotum; 111, abdominal terga seven to ten, and subgeni-

tal plate (dorsal); 112, hind wing (a, posterior branch of the radial vein); 113, seventh abdominal tergum; 114, ninth ab-

dominal tergum and laterotergites (dorsal; the laterotergites are turned under the tergum); 115, eighth abdominal sternum,

ninth laterotergites, and subgenital plate (ventral); 116, supraanal plate and paraprocts (ventral); 117, left style and thickened

corner of the subgenital plate; 118, subgenital plate and genitalia (dorsal).

three phallomeres, the hook occurring on the left side

(fig. 118), placing the genus in the Blattellinae.

Remarks. — The hind wing venation of

Haplosymploce is similar to that of Episymploce except

that its radial vein is unbranched (or has a pseudopos-

terior branch). Also the former genus has intercercal

104

processes which are lacking in Episymploce.

In my revision of Episymploce (Roth, 1987a: 150) I

described and redescribed 53 species. In this paper I

describe two new species and transfer some species of

Symploce, raising the total number in the genus to

about 70.

Episymploce simmonsi sp. n.

(figs. 110-118)

Specimens examined. — Holotype, d, Indonesia,

Sulawesi, Tengah, Gng., Sinsing, above Sinsing

Camp, SW. of Luwuk, understory/canopy lowland

rainforest at light, 200 m, Sample Sul. 16, 16.x.1989,

J.P. Duffels; in ZMAN. Paratypes: Sulawesi. ZMAN:

Indonesia, Sulawesi Tenggara, Centipede Camp, nr.

Gng. Watowila, NE. of Kolaka, c. 3°49'S 121°40'E,

canopy undisturbed hilly rainforest at light, 1100 m,

Sample Sul. 31, 1d (terminalia slide 47), 2.xi.1989,

Sample Sul. 34, 12, 5.xi.1989, J.P. Duffels.

Description. — Male: Head slightly exposed, inter-

ocular space the same as distance between ocellar

spots. Pronotum suboval (fig. 110). Tegmina and

wings fully developed extending beyond end of ab-

domen, the former with longitudinal discoidal sec-

tors. Hind wing with five subcostal veins and ten sim-

ple costal veins, radial vein with a branch originating

before the middle, terminating in one forked and one

simple branch; media and cubitus veins concave the

former simple, the latter with four complete and two

small incomplete branches, apical triangle subobso-

lete (fig. 112). Front femur Type A,; pulvilli on four

proximal tarsomeres of all legs, tarsal claws simple,

symmetrical, arolia small. First abdominal tergum

with a row of setae along an anterior ridge and a few

setae anteromedially. Seventh abdominal tergum

with a pair of deep, dark fossae anteromedially, with a

row of setae along the anterior margins of the depres-

sions; between the fossae is a triangular elevation

bearing a group of setae (figs. 111, 113). Right and

left ninth laterotergites similar, ventral margins with-

out dark setae (fig. 114). The supraanal plate hind

margin has a U- or V-shaped medial excavation form-

ing a pair of lobes that do not reach the hind margin

of the subgenital plate if their apexes are deflexed

(holotype, fig. 111); in the paratype the lobes are not

deflexed and their tips reach the hind margin of the

plate; each lobe has a minute dark terminal spine; in-

tercercal processes absent; right and left paraprocts

dissimilar, the right one with a small sclerotized, api-

cally rounded process and a large, more lightly sclero-

tized plate, left paraproct with a darkly sclerotized

plate (figs. 111, 116). Subgenital plate asymmetrical

the hind margin truncate, left side roundly thickened

or swollen and bearing minute spicules or spines;

styles, which are not visible in ventral view (fig. 115),

are strongly dissimilar, the right one large, spinelike,

lying pressed against the right side of the plate, left

one much smaller, corkscrew shaped, arising near the

base of the swollen margin (figs. 111, 115, 117, 118).

Genitalia as in fig. 118; hook on the left side with a

preapical incision, median phallomere a slender rod,

ROTH: Blattaria, Blattellidae, Blattellinae

its apex darkly sclerotized and acute, right phallomere

consisting of about four sclerites one of which is a

small cleft.

Female: Supraanal plate convexly rounded. Branch

of radial vein of hind wing terminally forked.

Colour. — Head yellowish brown without mark-

ings, labrum darker; maxillary palpomeres two and

three pale, segments four and five dark brown.

Pronotum with a pair of irregular, oblique blackish

stripes that reach the anterior and posterior borders,

background colour yellowish brown (fig. 110).

Tegmina without markings, hyaline, light brown,

humeral area lighter. Hind wing infuscated (fig. 112).

Abdominal terga brown with yellowish lateral bor-

ders, fossae on abdominal tergum seven black; in the

holotype, the supraanal plate has a pair of dark round

maculae anteromedially (fig. 111), which is absent in

the paratype, remainder yellowish brown. Abdominal

sterna and legs light brown. Cerci light brown dorsal-

ly, darker ventrally.

Measurements (mm) (® in parentheses). Length,

13.5-14.0 (12.8); pronotum length X width, 3.1-3.4

X 3.8-4.1 (3.0 X 3.7); 14.0-14.2 (12.7); interocular

space, 0.4-0.6 (0.8).

Etymology. — The species is dedicated to Dr.

Emory Simmons, mycological taxonomist, friend,

and former colleague at the U.S. Army Research and

Development Laboratories, Natick, MA.

Remarks. — Compared with the other known

species on Sulawesi, the subgenital plate, and styles of

simmonsi come closest to those of Episymploce sulawe-

siensis Roth (cf. figs. 117-118 with figs. 6C, E, in

Roth 1986c). The two species differ strongly in their

pronotal markings, and shapes of the supraanal plate

and ninth laterotergites (cf. figs. 110, 111, 114, 116,

with figs. 6A, B, G in Roth, 1986c).

Episymploce parafissa sp. n.

(figs. 119-123)

Specimen examined. — Holotype 4 (terminalia

slide 50), Indonesia, Sulawesi, Tengah, Totop camp

alonguBatuieriverm 0922 SES OLE RS WAMOL

Luwuk, Sample Sul. 21, understory/canopy lowland

rainforest, at light, 120 m, 21.x.1989, J.P. Duffels; in

ZMAN.

Description. — Male: Head hidden, interocular

space greater than distance between ocellar spots.

Pronotum suboval (fig. 119). Tegmina and wings ful-

ly developed extending beyond end of abdomen, the

former with longitudinal discoidal sectors. Hind wing

with simple, thickened costal veins, radial vein with

an apically forked branch arising near the middle,

105

TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 119-123. Episymploce parafıssa sp. n., male holotype: 119, pronotum; 120, supraanal plate and paraprocts (ventral); 121,

seventh abdominal tergum; 122, ninth abdominal tergum and laterotergites (dorsal; the laterotergites are folded under the

tergum); 123, subgenital plate and genitalia (dorsal; the arrow indicates the subobsolete right style).

media vein simple, cubitus vein with two complete

and three incomplete branches, apical triangle small.

Front femur Type A; pulvilli on four proximal tar-

someres, tarsal claws symmetrical, simple, arolia

small. First abdominal tergum unspecialized.

Abdominal tergum seven with a pair of shallow, non-

setose depressions separated by a clear longitudinal

zone (fig. 121). Supraanal plate strongly asymmetri-

cal, hind margin deeply excavated forming two lobes,

the right one terminating as a stout spinelike process

and two dissimilar spinelike processes arising on the

ventral surface; intercercal processes absent; para-

procts dissimilar sclerotized plates, the left one larger

(fig. 120). Ninth laterotergites dissimilar, the left one

larger and with a fringe of minute spicules along the

ventral margin, these absent from the smaller right

106

plate (fig. 122). Subgenital plate asymmetrical with a

small apically spined structure protruding from the

hind margin and to its right a very small left style and

a subobsolete right style; midlateral margins of the

plate without processes (fig. 123; both styles are not

visible ventrally in the pinned specimen and the right

style can be detected under high magnification in the

slide preparation). Genitalia as in fig. 123: hook on

the left side, with a preapical incision; median phal-

lomere a curved simple rod, apex blunt; right phal-

lomere consisting of at least two distinct sclerites one

of which is a reduced cleft.

Colour. — Head black, ocellar spots whitish,

clypeus and labrum pale; maxillary palpi and anten-

nae black. Pronotum blackish with a broad yellow

border laterally and posteriorly and a small yellow

mark anteriorly (fig. 119). Tegmina dark reddish

brown with a yellow band along the subcostal region

and along about two thirds the length of the costal

zone. Abdominal terga dark brown, the lateral bor-

ders yellow. Supraanal plate reddish brown with a yel-

lowish spot on the distal margin of each lobe on either

side of the excavation. Abdominal sterna brown, yel-

low along the lateral margins. Legs dark brown.

Female. Unknown.

Measurements (mm). — Length, 9.7; pronotum

length X width, 3.0 X 3.5; tegmen length, 10.5;

interocular width, 0.8.

Remarks. — The pronotal markings of parafissa are

similar to those of fissa (cf. fig. 119, with fig. 9A in

Roth 1986c) and both have the hind margin of the

the supraanal plate deeply excavated (cf. fig. 120,

with fig. 9B in Roth, 1986c). Although there are dis-

tinct specific differences in the styli, detailed structure

of the supraanal plate and paraprocts, and the ninth

laterotergites, indicate that these two species are close-

ly related.

The following species of Symploce are transferred to

Episymploce, I had placed these in Symploce because

they have symmetrical supraanal plates, without

modified hind margins. One of the characters used to

separate Episymploce from Symploce was an asymmet-

rical plate, or if symmetrical its hind margin was ex-

cised or showed some modification. However, I now

believe that some species may have symmetrical, un-

modified supraanal plates, and belong in Episymploce,

provided other features (especially the subgenital

plate, styles, and wing venation) are characteristic of

the genus.

Episymploce falcifera (Hanitsch) comb. n.

Ischnoptera falcifera Hanitsch, 1925: 81, figs. 4, 5.

Symploce falcifera (Hanitsch). — Hebard 1929: 10; Roth

1985a: 456, figs. 1A-J. [Sarawak].

Episymploce juxtafaleifera (Roth) comb. n.

Symploce juxtafalcifera Roth, 1985a: 459, figs. 2A-J, 3A-1.

(Borneo, Philippines, Singapore].

Episymploce tibangensis (Roth) comb. n.

Symploce tibangensis Roth, 1985a: 460, figs. 4A-J. [Sarawak.

Episymploce busuangensis (Roth) comb. n.

Symploce busuangensis Roth, 1985a: 461, figs. 5A-K

[Philippines].

ROTH: Blattaria, Blattellidae, Blattellinae

Episymploce talinasensis (Roth) comb n.

Symploce talinasensis Roth, 1985a: 464, figs. GA-I.

[Philippines].

Episymploce torrevillasi (Roth) comb. n.

Symploce torrevillasi Roth, 1985a: 464, figs. 7A-J, 8A-G

[Philippines].

Episymploce telephoroides (Walker) comb. n.

Blatta telephoroides Walker, 1871: 23.

Allacta telephoroides (Walker). — Kirby 1904: 100.

Phyllodromia telephoroides (Walker). — Shelford 1908a: 12.

Gislenia telephoroides (Walker). — Princis 1959: 125 (incor-

rectly synonymized with Blatta brevipes Walker).

Symploce telephoroides (Walker). — Roth 1985a: 468, figs.

9A-I [India].

Episymploce dispar (Princis) comb. n.

Symploce dispar Princis, 1957: 147, fig. 14. — Roth 1985a:

468, figs. 10A-M. [Flores].

Episymploce forficula (Bey-Bienko) comb. n.

Symploce forficula Bey-Bienko, 1957: 906. — Roth 1986a:

379, figs. 3A-I. [China, Thailand].

Episymploce perakensis (Roth) comb. n.

Symploce perakensis Roth, 1986a: 379, figs. 4A-G. [West

Malaysia].

Episymploce quadripunctata (Hanitsch) comb. n

Phyllodromia quadri-punctata Hanitsch, 1915: 57, pl.1, fig.

‘Phyllodromia quadripunctata Hanitsch. — Hebard 1929:

Symploce quadripunctata (Hanitsch). — Princis 1969: 886;

Bruijning 1948: 83; Roth 1986a: 386, figs. 8A-G, 9A-E

(redescription).

Episymploce bipinnacula (Roth) comb. n.

Symploce bipinnacula Roth, 1986a: 389, figs. 10A-I. [South

Vietnam].

Episymploce ligulata Bey-Bienko

Episymploce ligulata Bey-Bienko, 1957: 911, fig. 12.

Symploce ligulata (Bey-Bienko). — Roth 1985c: 214, 1986a:

391, figs. 12A-J. [China, Thailand].

Episymploce popovi Bey-Bienko

Episymploce popovi Bey-Bienko, 1957: 910.

Symploce popovi (Bey-Bienko). — Roth 1985c: 214, 1986a:

394. [China].

107

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Episymploce marginata Bey-Bienko

Episymploce marginata Bey-Bienko, 1957: 911.

Symploce marginata (Bey-Bienko). — Roth 1985c: 214,

1986a: 394.

NEW RECORDS

Episymploce sundaica (Hebard)

Symploce sundaica Hebard, 1929: 64, pl. 4, fig. 5, pl. 5, fig.

2 (male & female).

Episymploce sundaica (Hebard). — Roth 1985c: 214, 1986b:

365, figs. 7-13 (synonymy and redescriptions male & fe-

male).

Specimen examined. — Sabah. RMNH: N. Borneo,

16 km NE. of Tenom, orchid garden surr. by planta-

tions, 180 m, 1d, 06-1986, J. Huisman. Philippine

Islands. ansp: Zambalea Province, Luzon, 2d, 19,

x11.1917, W. Boettcher.

Remarks. — Episymploce sundaica is a very widely

distributed species with at least seven synonyms. The

type is from Sumatra but the species is found in Java,

Kalimantan (East Borneo), Laos, Okinawa, Papua

New Guinea, possibly Sulawesi (Celebes), Thailand,

and Taiwan; in Sabah, it was previously reported

from Sensuron (Roth 1986b: 371-373).

Episymploce malaisei malaisei (Princis)

Symploce malaisei Princis, 1950b: 217, figs 14, 15 (male &

female).

Episymploce malaisei malaisei (Princis). — Roth 1987c: 136,

figs. 3A-K (redescription).

Specimen examined. — Burma. ansp: Myitkyina,

Upper Burma, at tent lights or environs of town, 14,

10.vi.1945, J.W.H. Rehn.

Remarks. — This subspecies also is found in China.

Episymploce suknana Roth

Episymploce suknana Roth, 1987b: 119, figs. GA-J (male).

Specimens examined. — India. mczc: Alagar hill,

about 16 km from the ancient temple city of

Madurai, rainforest, about 900 m, 34 (one with ter-

minalia in vial), 49, reared from nymphs (killed

26.vii.1994), A. Padmanabhan.

Measurements (mm) (& holotype in brackets; 9

in parentheses). — Length, 18.5-20.0 [19.0] (19.4-

22.5); pronotum length X width, 4.3-4.6 X 5.5-5.8

[4.6 X 6.1] (4.8-5.3 X 6.0-6.5); tegmen length,

17.5-18.1 [20.0] (19.0-20.5); interocular width, 0.6

(0.9).

108

Remarks. — This species was previously known

from the unique male holotype from Sukna, 55 km

south of Darjeeling. The specimens have a large black

macula on the pronotum and the head is black except

for the pale clypeus. The male supraanal and subgen-

ital plates and genitalia agree well with those of the

holotype.

According to Padmanabhan (personal communica-

tion), the nymphs of suknana are shiny black and

rarely have a pair of orange spots on the pronotum.

The immatures live in holes under stones. Adults also

occur under stones and litter and rarely are seen on

shrubs. Females carry their oothecae for two to three

days before depositing them. Eggs hatch in 34 to 36

days.

ACKNOWLEDGEMENTS

I am indebted to Dr. Horst Bohn of the

Zoologisches Institut der Universitit München,

Germany, for critically reviewing the manuscript and

his suggestions. I thank the Australian Biological

Resources Survey (ABRS) for partial support, and the

museums, curators and collection managers indicated

in the introduction, who loaned me specimens.

REFERENCES

Asahina, S., 1979. Taxonomic notes on Japanese Blattaria.

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Princis, Parasigmoidella Hanitsch and Robshelfordia

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Invertebrate Taxonomy 5: 553-716.

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of the Entomological Society of London (1907): 455-

470.

109

TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 140, 1997

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— British Museum: London, pp. 1-43.

Received: July 1996

Accepted: December 1996

Index to genera and species

Valid names of genera and species in italics, syn-

onyms in roman type

AllactaSaussure&@Zehntnen en aan vee oie 107

andamanica Princis (Haplosymploce) .. 67, 91, 98, 99, 101

apicalis Shiraki (Pseudothyrsocera)

Bey bienodiothe rs ke ee eee eee 91

bicolor Shelford (Haplosymploce) ..... ND OON

bicolorata Roth (Haplosymploce) … 6791953977

bipinnacula Roth (Episymploce) .................. 107

AG Altin acus 4660000000808 00 68, 73, 91, 102, 107

brevipes Walker (Episymploce) ................... 107

busuangensis Roth (Episymploce) .................. 107

castanea Brunner (? Haplosymploce) ............ 69, 101

circumcincta Stal (Pseudothyrsocera)

TS ace yard ED 69, 71, 83, 84, 86, 87-89

circumclusa Stal (Pseudothyrsocera) ........ 83, 86, 87, 89

curta Hanitsch (Haplosymploce) ............... 91, 101

Aisparnincisy EDP ONE CCE TOO EEE 107

dlezing Wallac (GG) RP ee 91, 102, 103

Episymploce Bey-Bienko ....... 67-69, 89, 103, 104, 105

falcifera Hanitsch (Episymploce)

ferruginea Roth (Symploce)

forficula Bey-Bienko (Episymploce) ...............- 107

fulva Hebard (Pseudothyrsocera) ....... 68, 73, 74, 75, 89

juluzspecieseroup EEE CE CC LE CCC 69, 71, 89

Gislenia PHI ORIO ee 107

guttifera Walker (Beybienkoa) ............. 91, 100, 102

anssen naine EE 69

Haplosymploce Hanitsch … 67, 68, 89, 101-103, 105

110

Elebardina Bey. Bienkon ser 89

Hemithyrsocera Saussure ............. 68, 78, 81, 87, 91

henrici Hanitsch (Pseudothyrsocera) ....... 69, 71, 87, 89

Ischnoptera,Bunmeisten nr EEE SINE G5), 97 1107

juxtafalcifera Roth (Episymploce) .................. 107

EeptothyrsoceralDrineis REPARER EEE ET 67, 68, 69, 73

ligulata Bey-Bienko (Episymploce) ................ 107

lugubris Stal (Pseudothyrsocera) . . 69, 71, 81, 82, 83, 87-89

leu brie SPECIES groups rr er 69, 81, 89, 104

malaisei malaisei Princis (Episymploce) ............. 108

marginata Bey-Bienko (Episymploce) … 108

montana Shelford (Pseudothyrsocera) … 69, 71, 85, 89

montis Shelford (Haplosymploce) 67, 91, 94, 95, 97, 98, 101

Mopsellalanitsche On PME RE EE Er 69

Mopseravlele DATA EEE rier EEE 68, 69, 71, 99, 101

Moon Ines MEER 78

moultoni Hanitsch (Haplosymploce) ...... 91-93, 95, 101

nigra Hanitsch (Haplosymploce) 67, 91, 94, 97, 98, 99, 101

aM 6.00.0,0.0000 0 8.0.0.00.000 0000006000 91

papuensis Roth (Beybienkoa) ......... 91, 100, 102, 103

parafissa Roth (Episymploce) ............. 105, 106, 107

BarasymploceiHlebard res ee 95, 98, 103

Perakensi Roum EPTO 107

baan amd saas III E 89

perkinsi Roth (Pseudothyrsocera) … … 69, 71, 79, 89

Phyllodromia Serville . .. 69, 71, 77, 79, 89, 101-103, 107

pica Walker (Haplosymploce) ........ 9091519293 101

DICASPÉCIESSIOUPI EE ETTI TIE 91

popovi Bey-Bienko (Episymploce) ................. 108

IESCUGOMOPISENI MEER RETI STAI

Pseudothyrsocera Shelford . . 67, 68, 69, 92, 93, 95, 99, 101

quadripunctata Hanitsch (Episymploce) ............ 107

rectangularitervittata Brunner (Pseudothyrsocera)

RR, ah aloe salata 69, 70, 71, 85, 89

reversa Walker (Haplosymploce) ......... 91, 96, 98, 101

rubronigra Hanitsch (Pseudothyrsocera)

Cp a Re cot et RR LALA

ruficollis Shelford (Haplosymploce) ....... 90, 91, 93, 101

rufiventris Stal (Pseudothyrsocera) 69, 71, 80, 81, 83, 87, 89

scutigera Walker (Pseudothyrsocera) ....... 69, 75, 76, 89

semicincta Stal (Pseudothyrsocera) . .............. 85-87

signata Brunner (Pseudothyrsocera) … . . 83, 85, 86, 87, 89

simmonsi Roth (Episymploce) ................ 104, 105

sinensis Walker (Pseudothyrsocera) ........... 69, 71, 89

suknana RotkEdisymploce) ECO EEE RENE 108

sulawesiensis Roth (Episymploce) .................. 105

sundaica Hebard (Episymploce)

Symploce Hebard . 89, 91, 95, 97, 101, 102, 105, 107, 108

talinasensis Roth (Episymploce) ................... 107

telephoroides Walker (Episymploce) ................ 107

NhyrsoceralBurmeist ee 81, 83, 86, 87, 92

tibangensis Roth (Episymploce) nace II 107

torrevillasi Roth (Episymploce) ................... 107

toxopeu laniesch/(Mopsella)ea ern TICO 69

walkeri Princis (Hemithyrsocera)

xanthophila Walker (Pseudothyrsocera) . ................

Re AS CHOUIMEN

Museon, Department of Biology, Den Haag

REVISION OF THE GENUS

METAEUCHROMIUS BLESZYNSKI

(LEPIDOPTERA: PYRALIDAE: CRAMBINAE)

Schouten, R. T. A., 1997. Revision of the genus Metaeuchromius Bleszynski (Lepidoptera:

Pyralidae: Crambinae). — Tijdschrift voor Entomologie 140: 111-127, figs. 1-28. [issn 0040-

7496]. Published 31 October1997.

The species of the genus Metaeuchromius Bleszynski, 1960 are revised. Metaeuchromius is a se-

nior synonym of Pseudeuchromius Bleszynski, 1965. Pseudeuchromius latus (Staudinger, 1870)

and Diptychophora euzonellus Hampson, 1896 are transferred to Metaeuchromius. Three new

species are described: M. latoides (Turkey), M. changensis (China) and M. inflatus (Nepal).

Newly discovered scent organs on the male abdominal sternite are figured and described. A key

is provided together with full (re)descriptions of all species, with notes on distribution and bi-

ology.

R. È A. Schouten, Museon, Department of Biology, Stadhouderslaan 41, 2517 HV Den

Haag, The Netherlands. E-mail [email protected]

Keywords. — Pyralidae; Crambinae; Metaeuchromius, key; new species; checklist; tympanal or-

gans; abdominal scent organs.

Metaeuchromius was described as a monotypic

genus for Eromene yuennanensis Caradja, 1937.

Bleszynski (1965) described the second species of the

genus, Metaeuchromius circe. Eromene latus

Staudinger, 1870 was transferred to the genus

Pseudeuchromius by Bleszynski (1965) and is now in-

cluded in the genus Metaeuchromius. Diptychopora eu-

zonellus Hampson, 1896 is here transferred to

Metaeuchromius. Metaeuchromius flavofascialis Park,

1990 is the most recently described species. At the

start of this study the genus comprised three species.

In this article three new species are described and two

species are transferred to Metaeuchromius.

Thus, the genus now contains eight species.

Metaeuchromius has its main distribution in the east-

ern Palaearctic. Only M. latusand M. latoides occur in

the more western part of the Palaearctic.

Bleszynski (1965) erected the monotypic genus

Pseudeuchromius for Euchromius latus (Staudinger,

1870). Comparing the diagnoses given by Bleszynski

(1965) of Pseudeuchromius and Metaeuchromius the

following characters to distinguish Pseudeuchromius

from Metaeuchromius are found: (1) The more or less

closed cell of the hind wing; (2) The elongated

apophyses.

Ad. (1) Upon checking various specimens of

Pseudeuchromius latus 1 found that the cell of the hind

wing in this species is also more or less closed. Ad (2)

In my opinion the elongation of the apophyses is not

a very strong character at the generic level. The elon-

gated apophyses of P. latus can be regarded as an sec-

ondarily derived character to facilitate the deposition

of eggs in narrow and relatively deep locations.

This together with the newly discovered abdominal

scent organs, which are found in P. latus, M. yuenna-

nensis (type species of Metaeuchromius) and several

other species of Metaeuchromius, have made me de-

cide to synonymize Pseudeuchromius with Meta-

euchromius.

Material and methods

The formula for the black terminal dots mentioned

in the species descriptions is read as follows; 2-2-3-2

stands for: the black terminal dots closest to the apex

start with a group of two, followed by a second group

of two dots, a group of three dots and finally a group

of two dots near the tornus (fig. 7).

The spelling of the localities is taken literally from

the labels as to make tracing of the material easier. For

the Chinese material the equivalent in Pinyin tran-

scription is given.

Distribution maps are based on personally exam-

ined material only.

The purpose of the key is to make identification as

easy as possible, starting with external characters and,

101

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

- 4, RR 5

yf ft,

Figs. 1-2. M. euzonellus. — 1, Specialized scales covering the opening of the abdominal scent organ. 2, ‘Pits’ (see arrow) in ab-

dominal scent organ.

when unavoidable, followed by characters of the gen-

italia. Identification can be difficult or even impossi-

ble if the specimen is not in a good condition. In such

cases the genitalia have to be checked.

The terminology of the tympanal organs follows

Maes (1985).

Abbreviations for specimen depositories follow

Arnett et al. (1993).

Metaeuchromius Bleszynski, 1960

Metaeuchromius Bleszynski, 1960: 217. Type species:

Eromene yuennanensis Caradja, 1937: 151.

Pseudeuchromius Bleszynski, 1965: 90. Type

Eromene lata Staudinger, 1870: 204. Syn. n.

species:

Characteristics

The genus Metaeuchromius is externally very simi-

lar to Euchromius Guenée, 1845 and Miyakea

Marumo, 1933. Metaeuchromius is in my opinion

more closely related to Miyakea than to Euchromius.

This in the light of Miyakea and Metaeuchromius

sharing the rounded not forwardly protruding frons,

the ‘broad’ fore wings, the absence of a sclerotized

spinula and the apex of the gnathos directed upward.

Both genera also share the closed cell of the hindwing.

Landry (1995) sets Euchromius and Platytes sepa-

rate from the other Crambini as the most plesiomor-

phic group. This based on the synapomorphic char-

acter-state 11 (0): RS of the forewing free from other

radial veins. Both Metaeuchromius and Miyakea share

this synapomorphy with them.

The character-states by which Euchromius and

Platytes are placed in the Crambini are: character 15

(0): cell of the hindwing open; 27 (1): apex of the

gnathos directed downward; 41 (1): papillae anales

not coalesced.

112

Character-state 41 (1) is also found in Meta-

euchromiusand Miyakea. Character-state 15 (0) is not

found in Metaeuchromius and Miyakea (the cell of the

hind wing is closed), but I agree with Landry (1995:

37) this apomorphic character may have arisen inde-

pendently. As regarding to character-state 27 (1) the

same arguments may apply for Metaeuchromius and

Miyakea as for Ancylolomia (Landry 1995: 44): ‘the

most parsimonious solution regarding character-state

changes in this character is that the apomorphic state

occurs once in the Crambini except Ancylolomia and

independently in Calamotropha, Evergestis and

Scoparia. My conclusion based on the former para-

graphs is: Metaeuchromius is more closely related to

Miyakea than to Euchromius. A phylogenetic analysis

is not within the scope of the present study. Therefor

I make no comment of the relationship of Meta-

euchromius within the Crambinae. To make a more

scientifically thorough decision whether Meta-

euchromius and Miyakea form a monophyletic group

with Euchromius and Platytes two conditions must be

fulfilled: A) both genera must be included in the

cladistic analysis. B) The cladistic analysis must be

carried out with a monophyletic group and not by us-

ing a geographical area with taxa whose monophylet-

ic origin is highly disputable.

External characters

Metaeuchromius has no character at genus level to

distinguish it from Miyakea and Euchromius. For the

description of wings patterns etc. refer to Schouten

(1988).

Male genitalia

The uncus is broad (except in M. euzonellus), end-

ing in a sharp-pointed tip. Gnathos is simple hook-

shaped as long as the uncus. The tegumen has no spe-

cial shapes or structures. Sacculus present or incon-

spicuous. Processus of sacculus absent. Processus infe-

rior valvae absent. Processus basalis present or not,

usually rounded. Cucullus oblong to elongated usual-

ly ending in a sharp-pointed tip. Juxta variable, v-

shaped, rectangular or slender. Vinculum without

any special structures. Aedeagus varying from short to

long and slender, armed with many minute to one

cornutus and cornuti small to large.

Female genitalia

Papillae analis normal to small. Apophyses posteri-

ores and anteriores short to very long. Ostium simple.

Ductus bursae short or long, with or without scleroti-

zation, with or without a pouch-shaped enlargement.

Ductus seminalis slender. Bursa copulatrix rounded

or oblong, with or without signa, or sclerotization.

Tympanal organs

In Metaeuchromius the tympanum and conjunc-

tivum make an angle and the bulla tympani are of the

‘open’ type. Consequently, Metaeuchromius has tym-

panal organs of the ‘Crambidae’ type, as defined by

Minet (1982) and Maes (1985).

Description of the tympanal organs in

Metaeuchromius (figs. 3, 4). — The praecinctorium is

simple sac-shaped, no thorn-like sclerotization at the

connection with the pons tympani. The bulla tympa-

ni are bean-shaped. The fornix tympani, supporting

the conjunctivum, are well developed and clearly vis-

ible. The rami tympani are not connected, not form-

ing a well defined semi-circle as in Euchromius and

Miyakea. The sacci tympani are open and not very

deep. The processi tympani vary in size, but are small

to almost absent in /ata. The processi tympani are

dorsal-ventrally orientated and thus it is not always

possible to measure their size. The spinula is not scle-

rotized. The venula prima, situated lateral to the

fornix tympani, is always present. The venula secun-

da, posterior of the fornix tympani, is more or less an

extension of the venula prima and not very well de-

veloped in some species.

Abdominal scent organs

The presence of abdominal scent organs in

Crambinae species was up to now unknown. Several

species of Metaeuchromius i.c. yuennanensis, changen-

sis, circe, lata and euzonellus have abdominal scent or-

gans on sternite III of the males. The scent organs can

be classified as type 5 as used by Scoble (1992). The

type 5 scent organs consist of a group of scales which

cover a separately located glandular area. The scales

can be differentiated in various ways. The type 5 or-

gans are found on various parts of the body e.g. legs,

wings, abdomen. The type 5 scent organs have been

SCHOUTEN: Metaeuchromius

discovered on several families, Noctuidae,

Nymphalidae and Sphingidae (Scoble 1992). The

scent organs in males are generally believed to be used

when a male is in close contact with a female.

Diagnosis

(1) The dorsal insertion of the ductus ejaculatorius

is subterminal; (2) The gnathos is simple, hook-

shaped; (3) Sternite III of the males bear a more or

less developed pair of ‘scent organs’ sometimes with

specialized scales (except in flavofascialis and inflatus);

(4) The frons is rounded; (5) M1 of the hindwing is

located in the lower angle of the more or less closed

cell; (6) The ramus tympani is not connected, thus

not semicircular; (7) The saccus tympani is ‘open

and small; (8) The spinula is not sclerotized.

Checklist of Metaeuchromius Bleszynski

changensis sp. n.

circe Bleszynski, 1965

euzonellus (Hampson, 1896) comb. n.

flavofascialis Park, 1990

inflatus sp. n.

latoides sp. n.

latus (Staudinger, 1870) comb. n.

yuennanensis (Caradja, 1937)

Key to the species of Metaeuchromius Bleszynski

The key can only be used when specimens are in a

good condition. For a definite identification always

check the genitalia.

1. Six, seven or eight black dots at termen of the

forewing MON AS, CENT. to, be ER... D

— Four to five black dots at termen of the forewing

N RAE nn Weder Ae SE euzonellus

2. Groundcolour of the forewing pure white, usu-

ally a clear dark brown to black spot in posterior

ACI TINI NOI inflatus

— Groundcolour of the forwing dirty to creamy

WIEN RI I IT 3

SAM edialfasca wish 4

—22:Medialkraseiayellowishe Se ne ee 6

4. Males (male of /atoides unknown) ...... latus

Female ARE se re Rh 5

5. Bursa copulatrix with one minute signum latus

— Bursa copulatrix without a signum_.. . latoides

GAMElindwine darkibrown ere wed 20 oe circe

— Hindwing creamy white to grey or light grey-

brown. 0 IV. REL Rn SPER ER en 7

7. Medial fascia convex

— Medial fascia straight

8. Females (female of flavofascialis unknown)

ES IHREN. IR Wen: changensis

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 3-4.— 3, M. euzonellus. Abdomen, ventral aspect showing the tympanal organs and the abdominal scent organs on stern-

ite III. 4, M. changensis. Abdomen, ventral aspect showing the tympanal organs and the rounded scent organs on sternite III.

AMIE Re er Vath ae om rd de 9

9. Large group of cornuti on the vesica occupying

two-thirds of the length of aedeagus (fig. 16) ..

DSP Oe PRON, PMC ede changensis

— vesica with a group of cornuti occupying less

than one-third of the aedeagus (fig. 20)

ao dt dei RS flavofascialis

Metaeuchromius changensis sp. n.

(figs. 4, 6,9, 12, 16)

Type-material. — Holotype: d Chang Yang A.E.

Pratt Coll. May 1888’, GS 17849. Paratypes: 2

4, Chang Yang A.E. Pratt Coll. June 1888’, GS

17854, other male without abdomen. 2d 19 ,'Siang-

yang-fu [=Xiangfan] China W. 5000ft /93’, without

abdomen. All type-material in BMNH.

Diagnosis. — Externally very similar to the other

species in the genus. Differs in male genitalia (female

genitalia unknown) in having asymmetrical valvae in

114

combination with the aedeagus as long as or longer

than the length of the larger valve.

External characters (fig. 12). — Wingspan 14-17

mm. Frons not produced forward, creamy white,

dark brown centre; vertex creamy white to brown; the

length of the labial palp is two to two and a half times

the eye diameter, sides brown becoming lead-grey,

grey from above, creamy white from below; maxillary

palp brown, terminal part creamy white; on antenna

creamy white, inconspicuous darkly ringed. Thorax

brown; patagia brown; tegulae brown, inner side

lined creamy white. Forewing, groundcolour creamy

white densely suffused with ochreous to dark brown

scales, posterior area with one, more or less clear, dark

brown spot; medial fascia straight, running to one-

third of the dorsum; subterminal line brown-grey,

about midway between terminal dots and termina-

tion of posterior area; area adjacent to terminal black

dots white; seven black terminal dots, formula 2-3-2;

fringes shiny evenly grey. Hindwing light to dark

grey, subterminal fascia present, termen darkly bor-

dered; fringes creamy white with darker line.

Abdomen (figs. 4, 6). — Scent organs incorporated

in the abdomen, roundish, kidney-shaped, small

opening laterally directed. Wall of scent organs with

pits. No special scale formation observed in relation

to the scent organs.

Male genitalia (fig. 16). — Uncus normal, tapering

to sharp pointed tip; gnathos equal in length, dorsal

thorns absent; valvae asymmetrical, sacculus slender

in one valve, normal in the other, processus basalis

small somewhat rectangular, cucullus ending in bent

pointed tip, unequal in size; juxta oblong; vinculum

normal; aedeagus as long as or longer than length of

larger valve, with many cornuti, total length of cor-

nuti shorter than length of aedeagus.

Female genitalia. — Unknown, the only available

female specimen lacks an abdomen.

Biology. — Unknown. Specimens caught in May

and June. Found at an altitude of 1650 m.

Distribution (fig. 9). — Eastern part of China.

Metaeuchromius circe Bleszynski, 1965

(figs. 10, 14, 18, 23)

Metaeuchromius circe Bleszynski, 1965: 87. Holotype:

3 ,'Kwansien F.7.30', GS 7251 BM. BMNH.

Material. — 1d 49. China: Changyang, 19

(BMNH); Kwansien, 16 19 (BMNH); Siang-yang-fu

[=Xiangfan], 29 (BMNH).

Diagnosis. — Differs from all other species in hav-

ing dark brown wings in combination with a

wingspan of 14-16 mm.

External characters (figs. 14). — Wingspan 14-16

mm. Frons not produced forward, mingled creamy

white with brown, vertex creamy white to brown; the

length of the labial palp is two to two and a half times

the eye diameter, sides creamy white at base then

brown, creamy white and terminally brown, light

brown to brown from above, creamy white from be-

low; maxillary palp creamy white, darkly ringed at

base of last segment; antenna creamy white to brown.

Thorax brown; patagia creamy white with two broad

longitudinal dark brown stripes; tegulae dark brown,

inner side clearly lined creamy white. Forewing,

groundcolour creamy white very densely suffused

with ochreous to dark brown scales; medial fascia sin-

gle, yellow, nearly straight to slightly convex, inner

side whitish bordered, sometimes with small yellow-

ish spot, fascia running to ca. one-third of the dor-

sum; subterminal line dark brown, about midway be-

tween terminal black dots and termination of

posterior area; area adjacent to terminal black dots

white; seven black terminal dots, formula 2-3-2;

SCHOUTEN: Metaeuchromius

fringes shiny, evenly grey. Hindwing dark brown,

subterminal fascia absent; fringes light brown with

dark brown line.

Abdomen. — Sternite III with elongated groove,

without pits. The slide is too badly made to describe

scale formation if present.

Male genitalia (fig. 18). — Uncus stout, tapering to

sharp pointed tip; gnathos equal in length, dorsal

basal part bluntly enlarged; sacculus inconspicuous;

processus basalis small, inconspicuous; costa stronger

sclerotized ending in pointed tip; cucullus normal;

juxta broad triangular; vinculum short, rectangular;

aedeagus with circa four large cornuti.

Female genitalia (fig. 23). — Papillae anales indent-

ed; tergite VIII semi-connected forming inconspicu-

ous projection-like sclerotization; ostium simple;

ductus bursae short, sclerotized, terminal part broad-

ened, sac-like; ductus seminalis not found in slide;

bursa copulatrix oblong, without signum.

Biology. — Unknown. Specimens have been caught

in July and August. Found at an altitude of 1650 m.

Distribution (fig. 10). — East and central China.

Remarks. — Bleszynski (1965) quotes the holotype

label as ‘Kwansien F.7.31°, but upon checking, it

reads ‘F.7.30’.

Metaeuchromius euzonellus (Hampson, 1896) comb. n.

(Hes 212259015212)

Diptychophora euzonella Hampson, 1896: 18. Lectotype: d,

‘Khasis June, 1894 Nat. Coll’, GS 7481 (BMNH).

Material. — 206 39. India: Assam: Cherrapunji,

46 29 (BMNH); Khasi Hills, 7& (BMNH); Khasis, Id

19 (BMNH).

Diagnosis. — Resembles species with forewing

groundcolour white, differs in having five or four

black terminal dots. Differs in male genitalia in hav-

ing a fish-hook shaped gnathos, strongly bent down-

ward with upward curved tip. Differs in female geni-

talia from all other species in having tergite VIII

armed with sharp pointed tips.

External characters (fig. 13). — Wingspan 12-13

mm. Frons slightly produced forward, white; vertex

white; the length of the labial palp is three times the

eye diameter, with sides white at base, becoming

brown-grey, white from above and below; maxillary

palp white, brown at base; on antenna white, brown-

ish on upper half. Thorax, patagia and tegulae brown-

ish white. Forewing groundcolour white, densely suf-

fused with dark-brown scales, anterior part with

yellow-brown patch at costa, sometimes a second

brown patch near the medial fascia; medial fascia sin-

gle, slightly convex, running to halfway of the dor-

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 5-6. — 5, M. latus. Specialized scales covering the shallow, elongated scent organ. Arrow points to the terminal edge of

the glandular groove. 6, M. changensis. Abdominal scent organ with small opening (arrow) and "pits" in the wall of the ab-

dominal scent organ.

sum; subterminal line diffuse, brown, about midway

between terminal dots and termination of posterior

area; area adjacent to terminal dots white; five or four

black terminal dots, formula 3-2 or 3-1; fringes shiny,

evenly lead-grey, two white patches. Hindwing

creamy white to light brown, subterminal fascia pre-

sent, termen darkly bordered; fringes white with

brownish line.

Abdomen (figs. 1-3). — Scent organs incorporated

in the abdomen, round, large opening laterally direct-

ed. Wall of scent organs with large pits. Scent organs

covered with relatively broad scales. Tips of scales

touch scent organ pit. Scales can be put at a right an-

gle from the abdomen, thus spreading the scent or-

gans' contents (fig. 3).

Male genitalia (fig. 21). — Uncus strongly bent, tip

curved upward; gnathos proceeding beyond uncus,

dorsal thorns elongated; sacculus inconspicuous; cos-

ta armed with very stout double pointed projection,

smaller one with curved tip, cucullus slender, tip

pointed; juxta narrow; aedeagus slender, armed with

tooth-like cornutus.

Female genitalia (fig. 27). — Papillae anales small,

indented; membrane of tergite VIII without scleroti-

zation, tergite VIII with sharp pointed tips; ostium

simple with broadly rounded projection; ductus bur-

sae long; ductus seminalis narrow; bursa copulatrix

roundish, with spined area, without signum.

Biology. - Unknown. Caught in April, May, June,

July and September.

Distribution (fig. 9). — India: Assam.

116

Fig. 7. Wing pattern. aa = anterior area; d = dorsum; df =

double fascia; f = fringe; fl = fringe line; Pa = posterior area;

sf = subterminal fascia; sl = subterminal line; t = termen; td

= terminal dots.

f Pa df aa

sl Pa d

Metaeuchromius flavofascialis Park, 1990

(fig. 10, 20)

Metaeuchromius flavofascialis Park, 1990: 139. Holotype: à,

Chuncheon, GW, 30.V11.1988 (K.T. Park), GS 1840.

Kangweon National University, Chuncheon.

Material. — 66. China: West Tien-mu-shan

[=Tianmu Shan], 1d (BMNH). Korea: Chuncheon,

2d (kNuc), 1d (Rras); Sogumgang, 1d (kNuc); Mt.

Samag, 1d (KNUC).

Diagnosis. — Externally very similar to the other

species of the genus. Differs in male genitalia (female

unknown) in having the cucullus ending in a slender

tip, in combination with the group of cornuti occu-

pying less than one-third of the length of the aedea-

gus.

External characters. - Wingspan 13 mm. Frons not

produced forward, creamy white; vertex creamy

white; the length of the labial palp is one and a half

times the eye diameter, with the sides creamy white at

base becoming brown, creamy white to brown from

above and below; maxillary palp creamy white, dark-

ly ringed at base of last segment; on antenna creamy

white with some inconspicuous darker scales. Thorax

creamy white to light brown; patagia creamy white

with two broad longitudinal brown stripes; tegulae

brown, inner side lined creamy white. Forewing

groundcolour creamy, white densely suffused with

ochreous to dark brown scales, posterior area with

one or two small inconspicuous dark brown spots;

medial fascia nearly straight, running to circa one-

third of the dorsum; subterminal line brown, about

midway between terminal dots and termination of

posterior area; area adjacent to terminal black dots

white; seven black terminal dots, formula 2-3-2;

fringes shiny, evenly grey. Hindwing creamy white to

light grey, subterminal fascia very inconspicuous, ter-

men darkly bordered; fringes creamy white with dark-

er line.

Abdomen. — No indication of scent organs or spe-

cial formation of scales on sternite III.

Male genitalia (fig. 20). — Uncus tapering to sharp

pointed tip; gnathos about equal in length to uncus,

dorsal thorns absent; sacculus inconspicuous, proces-

sus basalis slender, fold reaching to the middle of the

tip of the valvae, cucullus ending in sharp pointed tip;

juxta elongated, more or less rectangular; aedeagus

relatively large, vesica with one group of cornuti.

Female genitalia. — Female unknown.

Biology. — Unknown. Caught in July, August and

September. The locatity of Mt. Samag was at 650 m

altitude.

Distribution (fig. 10). — Eastern part of China and

Korea.

SCHOUTEN: Metaeuchromius

Fig. 8. Distribution map. M. latus (dots) and M. latoides (tri-

angle).

Fig. 9. Distribution map. M. yuennanensis (squares), M.

changensis (dots), M. inflatus (triangle) and M. euzonellus

(diamond).

Fig. 10. Distribution map. M. circe (triangles), M. flavofas-

cialis (diamond).

119072

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 11-14. — 11, Metaeuchromius yuennanensis. 12, Metaeuchromius changensis. 13, Metaeuchromius euzonellus. 14,

Metaeuchromius circe.

Fig. 15. Metaeuchromius latus

118

Metaeuchromius inflatus sp. n.

(figs. 9, 22, 28)

Type-material. — Holotype d ‚Nepal Prov. Chisa-

pani Garhi Bhainse Dobhan 730m 16.-20.VII.1967

leg. Dierl-Schaft Staatsslg. München’, GS RS 483.

Paratype 9, same data as holotype, GS RS 501, both

in ZSMC.

Diagnosis. — Resembles species with groundcolour

of forewing white. Differs in male genitalia in having

the processus basalis strongly swollen, rounded. Can

be distinguished in female genitalia in having the os-

tium lip-shaped in combination with broad, strongly

sclerotized ductus bursae continued into a sac-like

part.

External characters. - Wingspan 11-13 mm. Frons

not produced forward, white to creamy white; vertex

creamy white, the length of the labial palp is one and

a half to two times the eye diameter, with the sides

white to creamy white, tip brown-grey, light brown

from above, white to creamy white from below; max-

illary palp white, dark brown ringed at base of last

segment; antenna white-brown ringed. Thorax, pata-

gia and tegulae white with yellow scales. Forewing

groundcolour pure white, densely suffused with

brown scales; medial fascia single more or less

straight, running to halfway to one-third of the dor-

sum; subterminal line brown, closer to termination of

posterior area than to terminal black dots; area adja-

cent to terminal black dots broad, white; seven black

terminal dots, formula 2-3-2; fringes shiny, white

with brown lines in upper half, brown at base and

white terminally in lower half. Hindwing white, sub-

terminal fascia absent, termen with inconspicuous

dark spot at inner corner; fringes white.

Abdomen. — No indication of scent organs or spe-

cial formation of scales on sternite III.

Male genitalia (fig. 22). — Uncus long, slender, ta-

pering to sharp pointed tip; gnathos equal in length-

to the uncus, without dorsal thorns; processus basalis

large, roundish, swollen, cucullus ending in inwardly

bent tip; juxta somewhat rectangular; vinculum long,

pointed; aedeagus relatively large, with two large cor-

nuti.

Female genitalia (fig. 28). — Papillae anales more or

less fused; ostium relatively large, with small lip; duc-

tus bursae broad, strongly sclerotized in upper part,

followed by enlarged sac-like part, ending slightly

swollen above bursa copulatrix; ductus seminalis nar-

row; bursa copulatrix oblong, without signa.

Biology. — Unknown. The specimens were caught

in July at 730m altitude.

Distribution (fig. 9). — Only known from Nepal.

SCHOUTEN: Metaeuchromius

Metaeuchromius latoides sp. n.

(fig. 8, 26)

Type-material— Holotype: 9 ‘Turquie Acigol

U.V., 31-7-1973 U. Dall'Asta GS R.S.470 (in

MAES).

Diagnosis. — Differs from most species in having a

whitish medial fascia. Differs from M. latus in having

eight black terminal dots (this character may turn out

to be invalid when more material is collected). Differs

in female genitalia from M. /atus in lacking the small

roundish signum in the oblong bursa copulatrix.

External characters. - Wingspan 19 mm. Frons not

produced forward, creamy white mottled with

brown; vertex creamy white mottled with brown; the

length of the labial palp is two and a half times the eye

diameter, creamy white mottled with brown; maxil-

lary palp lost on holotype; on antenna creamy white

to brown. Thorax and patagia creamy white mottled

with dark brown; tegulae creamy white, coarsely

brown mottled. Forewing groundcolour creamy

white, densely suffused with ochreous to dark brown

scales, posterior area with two , more or less clear,

dark brown spots; medial fascia single, convex, white,

running to about halfway of the dorsum, subterminal

line brown, about midway between terminal dots and

termination of posterior area; area adjacent to termi-

nal dots yellow to white; eight or nine black terminal

dots, formula 2-2-3-1 or 2-2-3-2; fringes too worn to

be described. Hindwing brownish, light subterminal

fascia present, termen darkly bordered; fringes

brownish.

Male genitalia. - Unkown.

Female genitalia (fig. 26). — Papillae anales small;

membrane and tergite VIII very long, tergite very

lightly sclerotized, apophyses very long; ostium very

simple, roundish; ductus bursae normal swollen un-

der ostium, finely wrinkled; ductus seminalis narrow;

bursa copulatrix weakly sclerotized, oblong, without

signa.

Biology. — Unknown. the holotype was caught at

the end of July.

Distribution (fig. 8). — Turkey.

Metaeuchromius latus (Staudinger, 1870) comb. n.

(ites, 5p, 5925)

Eromene lata Staudinger, 1870: 204. — Holotype 9,

‘Graecia Kr.’, GS 1147 (in ZMHB).

Pseudeuchromius latus (Staudinger, 1870) — Bleszynski,

1965: 90.

Pseudochromius latus (Staudinger). — Maes 1986: 79 [lapsus

calami].

Material. — 56, 139. Greece: Graecia Kr. 19

119

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 16-17. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 16, Metaeuchromius

changensis, 17, Metaeuchromius yuennanensis. Scale bar 0.5 mm.

120

(zMHB); 2 km SW of Anoyia, 18 (gras); Mt.

Taygetos, 1d (rras), 38, 29 (zmuc); Zachlorou,

16,22 (BMNH), 19 (RTAS), 72 (zmuc).

Diagnosis. — Differs from most species in having a

whitish medial fascia. Differs from M. latoides in hav-

ing six or seven black terminal dots (this character

may turn out to be invalid when more material be-

comes available). Differs in female genitalia from M.

latoides (male unknown) in having a small signum in

the roundish bursa copulatrix.

External characters (fig. 15). — Wingspan 18-22

mm. Frons not produced forward, creamy white to

light brown; vertex creamy white; the length of the

labial palp is three times the eye diameter, with the

sides light brown to creamy white at base, becoming

lead-grey, brown from above, creamy white from be-

low; maxillary palp brown; on antenna creamy white,

darkly ringed. Thorax light brown to brown; patagia

brown; tegulae brown evenly mottled. Forewing

groundcolour creamy white, densely suffused with

ochreous to dark brown scales, posterior area with

two more or less clear dark brown spots; medial fascia

single, convex, white, running to halfway to one-third

of the dorsum; subterminal line dark-brown, about

midway between terminal dots and termination of

posterior area; area adjacent to terminal dots yellow

to white; six or seven black terminal dots, formula 2-

3-1 or 2-3-2; fringes evenly lead-grey. Hindwing light

grey, subterminal fascia present, termen darkly bor-

dered; fringes grey to creamy white.

Abdomen (fig. 5). — Scent organs only indicated by

minute fold of sternite III, pits absent. Sternite III

armed with elongated broad scales.

Male genitalia (fig. 19). — Uncus stout, tapering to

sharp double pointed tip; gnathos equal in length or

slightly shorter than uncus, dorsal thorns absent; sac-

culus inconspicuous, sclerotized, processus basalis in-

conspicuous, costa stronger sclerotized to halfway,

stronger sclerotized part ending pointed, cucullus

broad, rounded; juxta elongated; vinculum long,

pointed; aedeagus very large, posterior part swollen,

with one elongated group of short cornuti.

Female genitalia (fig. 25). — Papillae anales small;

membrane and tergite VIII very long, tergite very

lightly sclerotized, apophyses very long; ostium very

simple, round; ductus bursae widening proximal to

the ostium, then wrinkled, distal half with contorted

slightly sclerotized lining; ductus seminalis narrow;

bursa copulatrix roundish, with one very small

roundish signa.

Biology. — Habitat at 600 m altitude: shrubby and

hilly area grazed occasionally by sheep. Specimens

were caught in the last week of June, July and August

at an altitude of 600 to 1000 m.

SCHOUTEN: Metaeuchromius

Distribution (fig. 8). — So far only found in Greece;

records from Turkey (e.g. Maes 1986) may refer to

M. latoides.

Remarks. — Bleszynski (1965) does not mention a

signum in his description nor is one present in the

drawing of the female genitalia. Upon checking the

holotype, it turned out to have a small signum.

Perhaps due to not removing the genitalia from the

abdomen and not colouring the slide Bleszynski

missed seeing the small signum.

Metaeuchromius yuennanensis (Caradja, 1937)

(figs. 9, 11, 17, 24)

Eromene yuennanensis Caradja, 1937: 151. Lectotype: 5, Li-

kiang. (China). Provinz Nord-Yuennan, 4.8.1934. (not

10.6.1935 as stated in Bleszynski 1965) H. Hone’.

Muzeul de Istoria Naturala ‘Grigore Antipa’, Bucuresti.

Metaeuchromius yuennanensis (Caradja, 1937) Bleszynski,

1960: 217. — Bleszynski 1963: 112.

Material. — 96, 72. China: Likiang, 2d, 19

(BMNH), 16, 19 (RTAS), 34, 39 (zemk), 2d, 29

(Bucu); West Tianmu Shan, 16 (ZEMK).

Diagnosis. — Differs in male genitalia from all

species in having the valvae asymmetrical and the cu-

cullus ending in a bent, slender tip, in combination

with the total length of the group of cornuti about

equal to that of the aedeagus. Differs in female geni-

talia (female of M. changensis is unknown) in having

the ductus bursae split in two parts, of which one is

strongly sclerotized and partly armed with spines,

ending less sclerotized and somewhat wrinkled.

External characters (fig. 11). — Wingspan 16-20

mm. Frons not produced forward, creamy yellow-

white; vertex creamy yellow-white; the length of the

labial palp is twice that of the eye diameter, with the

sides creamy white at base becoming brown, light

brown from above, creamy white from below; maxil-

lary palp light brown, darkly ringed at base of last seg-

ment, terminal part light brown; on antenna creamy

white, inconspicuously ringed. Thorax brown; pata-

gia yellow-brown; tegulae brown, inner side clearly

lined creamy white. Forewing groundcolour creamy

white, densely suffused with grey and ochreous to

dark brown scales, posterior area with more or less

clear dark spot near medial fascia; medial fascia single,

slightly convex, running to one-fourth to one-third of

the dorsum; subterminal line brown, about midway

between terminal dots and termination of posterior

area; area adjacent to terminal black dots white; seven

black terminal dots, formula 2-3-2; fringes shiny,

evenly grey. Hindwing creamy white to grey, subter-

minal fascia clearly present, termen darkly bordered;

fringes creamy white with faint darker line.

121

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 18-20. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, lateral aspect. — 18, Metaeuchromius circe,

19, Metaeuchromius latus, 20, Metaeuchromius flavofascialis. Scale bar 0.5 mm.

122

SCHOUTEN: Metaeuchromius

Figs. 21-22. Male genitalia; ventro-caudal aspect, valvae spread; aedeagus separated, 21 ventral aspect, 22 lateral aspect. — 21,

Metaeuchromius euzonellus, 22, Metaeuchromius inflatus. Scale bar 0.5 mm.

Abdomen. — Two scent organs incorporated in the

abdomen, roundish, kidney-shaped, with small open-

ing laterally directed. Wall of scent organs with pits.

No special scale formation observed in relation to the

scent organs.

Male genitalia (fig. 17). — Uncus tapering to sharp

pointed tip; gnathos equal in lengthto uncus, dorsal

thorns absent; valvae asymmetrical, sacculus slender

in one valve, normal in the other, processus basalis

somewhat rectangular, cucullus asymetrical, ending

in bent pointed tip unequal in size; juxta large some-

what oblong; vinculum normal; aedeagus shorter

than length of smaller valve, with many very long cor-

nuti, total length of cornuti as long as length of aedea-

gus.

123

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 23-25. Female genitalia; ventral aspect. — 23, Metaeuchromius circe, scale bar 0.5 mm; 24, Metaeuchromius yuennanensis,

scale bar 0.5 mm; 25, Metaeuchromius latus. Scale bar 0.85 mm.

124

SCHOUTEN: Metaeuchromius

26,

aspect. —

ventral

Metaeuchromius latoides, scale

bar

DIA

euzonellus,

mm;

0.5 mm;

0.85

Metaeuchromius

scale

28,

scale

bar

Metaeuchromius inflatus,

bar 0.5 mm.

125

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Female genitalia (fig. 24). — Papillae anales idened;

ostium simple; ductus bursae split in two parts, one

very long strongly sclerotized part with many spines

which ends sac-like, wrinkled and less sclerotized, sec-

ond part; ductus seminalis narrow; bursa copulatrix

oblong, without signum.

Biology. — Unknown. The specimens have been

caught in August and September at an altitude of up

to 3000 m.

Distribution (fig. 9). — China. See also under re-

marks.

Remarks. — There may be another species very sim-

ilar to M. yuennanensis occurring in Japan, or M.

yuennanensis itself inhabits Japan. A specimen was

found identified as M. yuennanensis in the BMNH. The

lack of abdomen makes it impossible to decide

whether it belongs to a new species or not.

Metaeuchromius yuennanensis tibetanus Bleszynski

Metaeuchromius yuennanensis tibetanus Bleszynski, 1965: 87.

Holotype: d, Batang. (Tibet). Im Tal des Yangtze (ca.

2800 m) 13.9.1936. H. Hône. Paratypes 1d, 29

22.9.1936, 18.9.1936, 1.9.1936. All in Bucu.

Diagnosis. — The slightly lighter colouration of the

fore wings is the only distinguishing character for this

subspecies.

Biology. — Unknown. The specimens have been

caught in September at an altitude of 2800 m.

Discussion

The males of Metaeuchromius yuennanensis, M.

changensis, M. latus, M. circe and M. euzonellus have

ventrally located abdominal scent organs which pro-

vide these species with a synapomorphic character.

M. latoides is only known from the unique female.

Comparing this species with the other females of

Metaeuchromius it can be seen to be most closely re-

lated to M. latus. This conclusion is based on the al-

most similar features of the genitalia. The only differ-

ence is the presence of a signum in M. latus and the

presence of two extra black terminal dots in M. /a-

toides. This very close relationship allows the conclu-

sion that the up to now unknown male of /atoides

must also have abdominal scent organs.

The males of M. flavofacialis and M. inflatus lack

these male scent organs. However, the placement by

Dr. Park of flavofascialis in the genus Metaeuchromius

is in my opinion justifiable considering the other

characters it shares with the other members of the

genus. M. inflatus also lacks the abdominal scent or-

gans, but in light of other characters (wing pattern,

gnathos and valvae) it most logically belongs in

126

Metaeuchromius. M. euzonellus has the Sc and R, of

the forewing partly fused, but the presence of the ab-

dominal scent organs are regarded as being more im-

portant. Also, within several other Crambinae genera,

and even within a single species, the fusion of Sc and

R, are known to be variable.

ACKNOWLEDGEMENTS

I wish to thank the following persons and institutes

for loans of material without which this study would

not have been possible. Dr. W. Dierl (zsmc,

München, Germany), Dr. O. Karsholt (zmuc,

Kopenhagen, Denmark), Dr. K. Maes, (MRAC,

Tervuren, Belgium), Dr. M. Mey (zmx8, Berlin,

Germany), Prof. Dr. K. T. Park (kNuc, Kangweon,

Korea), Dr. D. Stuning (ZFMK, Bonn, Germany).

Dr. B. Landry is acknowledged for his comments

on the manuscript.

As always I am greatly indebted to Mr. M. Shaffer

(BMNH, London, United Kingdom) for his encour-

agements, help and great expertise on which I can al-

ways rely and ask for.

Visits to the BMNH, ZSMC and zmuc have been

made possible by grants awarded by the

Uyttenboogaart-Eliasen Stichting.

REFERENCES

Bleszynski, S., 1960. Studies on the Crambidae (Lepi-

doptera). Part XXIX. Species of the genus Euchromius

Gn. — Acta Zoologica Cracoviensia 5: 203-247.

Bleszynski, S., 1963. Studies on the Crambidae

(Lepidoptera). Part 40. A review of the genera of the fam-

ily Crambidae with data on their synonymy and types. —

Acta Zoologica Cracoviensia 3: 91-132.

Bleszynski, S., 1965. Crambinae. In: H.G. Amsel, H.

Reisser und F. Gregor, Microlepidoptera Palaearctica.

Vol. 1: i-xlvii, 1-533. Wien.

Caradja, A. & E. Meyrick, 1937. Materialen zur einer

Mikrolepidopterenfauna des Yülingshanmassivs (Provinz

Yiinnan). — Iris 51: 137-182.

Gaskin, D. E., 1974. The species of Pareromene Osthelder

(Pyralidae: Crambinae: Diptychophorini) from Malaysia,

Indonesia and New Guinea. — Journal of Entomology (B)

43(2): 185-208.

Gaskin, D. E., 1985. Morphology and reclassification of

the Australasian, Melanesian and Polynesian

Glaucocharis Meyrick (Lepidoptera: Crambinae: Dipty-

chophorini). — Australian Journal of Zoology, Suppl.

Series No. 115: 1-75.

Hampson, G. F., 1896. On the classification of the

Schoenobiinae and Crambinae, two subfamilies of moths

of the Pyralidae. — Proceedings of the Zoological Society

of London 1895: 897-974.

Landry, B., 1995. A phylogenetic analysis of the major lin-

eages of the Crambinae and of the genera of Crambini of

North America (Lepidoptera: Pyralidae). - Memoirs on

Entomology, International 1, 242 pp. Associated

Publishers, Gainesville.

Maes, K., 1985. A comparative study of the abdominal tym-

panal organs in Pyralidae (Lepidoptera) I. Description,

terminology, preparation technique. — Nota Lepi-

dopterologica 8(4): 341-350.

Maes, K., 1986. Pyralidae from Turkey. — Nota Lepi-

dopterologica 9(1-2): 78-80.

Meyrick, E., 1930-1936. Exotic Microlepidoptera. 4.: 1-

642. London.

Minet, J., 1982. Les Pyraloidea et leur principales divisions

systématiques. — Bulletin de la Société entomologique de

France 86 (1981): 262-280.

Park, K. T., 1990. Two new species of Pyralidae

SCHOUTEN: Metaeuchromius

(Lepidoptera from Korea). — The Korean Journal of

Entomology 20 (3): 139-144.

Schouten, R.T.A., 1988. Revision of the species of the genus

Euchromius Guenée, 1845 (Lepidoptera: Pyralidae:

Crambinae) occurring in the Afrtropical region. —

Zoologische Verhandelingen Leiden 244: 1-64.

Scoble, M. J., 1992. The Lepidoptera: form, function and

diversity. — Oxford University Press, New York, 404 pp.

Staudinger, O., 1870. Beitrag zur Lepidopterenfauna

Griechenlands. — Horae Societatis Entomologicae

Rossicae 7: 3-304.

Received: 1995

Accepted: March 1997

127

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

BOOK REVIEW

Bengt A. Bengtsson, 1997. Scythrididae. — Micro-

lepidoptera of Europe (P. Huemer, O. Karsholt &

L. Lyneborg eds.) vol. 2, 301 pp., 14 colour plates,

419 figs. Hardback, 2417 cm. Apollo Books,

Stenstrup. [ISBN 87-88757-11-0]. Price DKK 500.—

excl. postage.

The second volume of this new series (see this jour-

nal, vol. 139: 84 for a review of the first volume),

treats 237 species of this relatively unknown family of

micro-moths. These moths, often day-flying, are

most abundant in the mediterranean region. This is

the first time that this family is revised for this large

geographic area, resulting in 40 (!) new species (three

of which are described by Passerin d'Entreves) and in

many other nomenclatural changes. The speciose

genus Scythris is divided into 64 informal species-

groups.

There is a short general introduction and a check-

list. All species are briefly described with the follow-

ing entries: diagnosis, male genitalia, female genitalia,

distribution, biology, and often: remarks. Most are il-

lustrated in colour, as water-colours, made by the au-

thor, with the right pair of wings depicted only. The

male and female genitalia are illustrated in simple

line-drawings.

The author is to be congratulated for bringing to-

gether this overwhelming amount of information in a

relatively short time. The descriptions are adequate,

and the water-colours look nice. It is unfortunate that

they are not printed bigger. They are 4-5 X natural

size (measured from size in the description: I have not

been able to find an indication of the magnification),

but particularly some of the smaller ones show very

little detail. The white species are almost invisible

against the white paper background, why haven’t

they been brought together, and printed on a darker

background? The black-and-white drawings are rela-

tively bold, and do not compare with the quality of

the colour-plates.

This series is announced as an identification series.

128

In this light it is to be regretted that the book does not

provide any help to the inexperienced user, to get fa-

miliar with the overwhelming number of similarly

looking moths. The author believes that leafing

through colour plates and genitalia-illustrations is the

quickest way to arrive at a species identification (page

11). However, one needs to have an enormous mem-

ory for forms, when comparing 237 species. With all

the problems which keys can provide, I would have

much preferred to have some sort of key, to bring

structure in this large amount of variation. That

should not necessarily be a dichotomous key, but

could have been a table of distribution of some im-

portant character-states. Also the species-groups do

not really help, because there is no indication how to

identify a species into a group. With modern com-

puter technology, the design of a key would have

been quite well possible, even with this amount of in-

formation. Another omission is, in my opinion, an

index to hostplants. Even with the little knowledge

about this topic for the scythrids, it would have facil-

itated the search for certain hosts.

My final comments are on the design of this book.

Although this may be partly affected by personal

taste, the design of this book is very unbalanced.

There has apparently been no professional graphic

designer, the used font is too big, there are too many

different font types, and there is often too much

white on the page. By using a smaller font and two

colums, many pages could have been spared, and used

to print the colour-illustrations somewhat larger. The

distribution catalogue remains (see review vol. 1) also

rather unpractical, especially because the countries are

listed alphabetically and not geographically.

This work is, in conclusion, an important contribu-

tion to the taxonomic knowledge of European and

Mediterranean Microlepidoptera, and, despite the

somewhat disappointing design, printing, and lack of

keys, to be recommended for all serious lepidopterists.

[Erik J. van Nieukerken]

Braprey J. SINCLAIR

Department of Environmental Biology, University of Guelph

A NEW SPECIES OF ORNITHOICA RONDANI FROM

NEW CALEDONIA (DIPTERA: HIPPOBOSCIDAE)

Sinclair, B.J. 1997. A new species of Ornithoica Rondani from New Caledonia (Diptera:

Hippoboscidae). — Tijdschrift voor Entomologie 140: 129-132, figs. 1-6 [1ssN 0040-7496].

Published 31 October 1997.

Ornithoica caledonica sp.n. (pusilla group) is described from New Caledonia, collected from the

endemic flightless bird Rhinochetos jubatus. In addition, Ornithoctona plicata (von Olfers) was

collected from the endemic pigeon Ducula goliath. A total of six species of Hippoboscidae are

now recorded from New Caledonian birds.

Bradley J. Sinclair. Present address: Biosystematics Laboratory, Graduate School of Social and

Cultural Studies, Kyushu University 01, Ropponmatsu 4-2-1, Fukuoka-shi, 810 Japan.

Key words. — Diptera, Hippoboscidae, Ornithoica, new species, New Caledonia.

Ectoparasites including louse flies (Diptera:

Hippoboscidae) were collected as part of a survey of

the parasites and their impact on two birds endemic

to New Caledonia; the kagu (Rhinochetos jubatus

Verreaux & Des Murs) and the notou (Ducula goliath

(Gray)). Field work was conducted by Dr Frederic

Beugnet in association with the CIRAD agency.

The louse fly, Ornithoica caledonica sp. n. (assigned

to the O. pusilla (Schiner) group) was collected from

the kagu. The kagu is a pale grey, mid-sized (58 cm),

flightless bird (Delacour 1966). Although flightless,

the kagu has large, brightly marked, flexible wings,

believed to be used for sexual displays (Lindsey

1993). It is a diurnal bird, inhabiting dense rain-

forests, and forages on insects, snails, and earthworms

(Lindsey 1993). The kagu is an endangered species,

with an estimated 200 breeding pairs. The wide-

spread louse fly, Ornithoctona plicata (von Olfers) was

collected from the notou, a large (50 cm) endemic pi-

geon, which inhabits montane forests (Delacour

1966).

Prior to this study five species of Hippoboscidae

were recorded from New Caledonian birds, including

the endemic species Myophthiria neocaledonica Maa,

and the widespread species Olfersia aenescens

Thomson, Ornithoctona plicata, Ornithoica exilis

(Walker), and Ornithophila metallica (Schiner) (Maa

& Marshall 1981, Maa 1989).

MATERIALS AND METHODS

Material based on this study will be deposited in

the Museum National d'Histoire Naturelle, Paris,

France (MNHP) and Canadian National Collection of

Insects, Ottawa, Canada (cnc). Specimens of O.

pusilla were borrowed from the United States

National Museum of Natural History, Washington,

USA (usnM).

Terms follow those of McAlpine (1981), except

male terminalia where terms of Cumming et al.

(1995) are followed. Male and female abdomens were

removed, macerated in hot 85% lactic acid, and ex-

amined in glycerin.

TAXONOMY

Ornithoica pusilla group

Species of the O. pusilla group are distinguished

from the remaining species of Ornithoica on the basis

of female tergite 6 medially divided and by clusters of

more than 30 pairs of large anchor-like spines near

the apex of the abdomen (Maa 1966).

Key to species of O. pusilla group

1. Apex of female abdomen with dense cluster of

anchor-like spines, base of spines in contact with

neighbouring spines; female tergite 6 subtrian-

gular; male laterite 6 bearing one long, stout and

one:shontisetal mann, O. pusilla (Schiner)

- Apex of female abdomen with anchor-like spines

not densely clustered, base of spines not in con-

tact with neighbouring spines; female tergite 6

rectangular; male laterite 6 bearing five long,

SOO GALS 4 0.0.0 0.0 00.000.505 O. caledonica sp.n.

129

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 1-6. Female and male abdomens and male terminalia of Ornithoica caledonica sp. n. — 1, female, dorsal view; 2, female,

ventral view; 3, male terminalia, lateral view; 4, female laterite 7; 5, male, dorsal view; 6, male, ventral view.

Abbreviations: hypd, hypandrium; Itg 7, laterite 7; pgt, postgonite; ph, phallus; phapod, phallapodeme; pregen pl, pregenital

plate; pregen tb, pregenital tubercle; span pl, supra-anal plate.

130

Ornithoica caledonica sp. n.

(figs. 1-6)

Type examined. — Holotype 9 : ‘NEW CALEDONIA /

Parc Rivière Bleue / 1992-94; ex. Kagu / coll. F.

Beugnet, / Y. Létocart & R. Gadet’; ‘HOLOTYPE /

Ornithoica | caledonica | Sinclair’ (MNHP).

Paratypes: Same data as holotype (1d, 39, cnc;

26,39, MNHP).

Recognition. — Females distinguished from O.

pusilla by a rectangular tergite 6, narrowly divided

and lacking long stout setae, crescent-shaped supra-

anal plate, pregenital tubercle bearing one long seta

and one long spine-like seta, subtriangular laterite 7,

and apex of abdomen with anchor-like spines less

densely clustered. Males are distinguished by 5 long,

strong setae on laterite 6 and by the slender ventral

arm of the phallapodeme.

Etymology. — Named in reference to the type lo-

cality.

Description

Female. — Wing length 3.25-3.5 mm

Thorax. Presutural scutum (prescutum) with long,

pale setae, about as dense as scutellum; postsutural

scutum (scutum) largely bare, with transverse row of

pale setae along posterior margin. Scutellum with 2

long and 3 shorter pairs of dark, subapical setae.

Anterior margin of prosternum truncate.

Wing. Microtrichia covering only extreme tip of

cell r,,,; cells r,. and m,,, mostly covered by mi-

crotrichia except for basal fifth; cell cuA, with patch-

es along vein.

Abdomen. Richly spinose; syntergite 1+2 with

short, fine setae; dorsal setae on tergites 3-6 shorter

than length of tergites; lateral setae on tergites 3-5

more pronounced; lateral setae on tergite 5 twice

length of tergite; tergite 5 longer than 3 or 4; tergite 6

narrowly divided medially into pair of broad rectan-

gular sclerites, lacking long stout setae. Laterite 2 with

marginal setae longer than tergites 3 and 4 combined;

ventrolateral margin of laterite 2 with short, spine-

like setae; laterite 7 subtriangular, with 3 strong setae

and 4 setulae. Dorsolateral membranous region above

level of spiracles lacking setae. Sternite 1 with numer-

ous marginal setae and 1 median spine-like seta; disc

of venter with 7-8 rows of slender setae, basal ring

pale; setae lacking anterior to pregenital tubercle; pre-

genital tubercle paired, bearing 1 long strong seta and

1 long spine-like seta. Anchor-like spines clustered in

2 widely separate groups; spines equal in size; base of

spines not in contact with neighbouring spines.

Supra-anal plate with anterior and posterior sclerites

SINCLAIR: Hippoboscidae from New Caledonia

clearly separated, both crescent-shaped; anterior scle-

rite thinly sclerotized; posterior sclerite with 2 pairs of

setulae, outer pair longer than median; pregenital

plate somewhat ‘“L’-shaped.

Male. — Similar to female. Tergites 3-5 broader

than female; tergite 6 widely divided into pair of sub-

rectangular sclerites; laterite 6 slender, bearing 5 long,

strong setae; spiracle 7 lying anterior to 6; ventrolat-

eral setae mounted on slightly raised sockets.

Terminalia: Phallapodeme (basal apodeme) subtrian-

gular, ventral edge constricted forming long slender

process; postgonite or gonostylus (tip of penis valve)

strongly arched and tapering, margin with row of se-

tulae; phallus (aedeagus) slender, tapering to point,

encased in membranous sheath.

Remarks. — Prior to this study, the O. pusilla group

was monotypic. Ornithoica pusilla is widespread

among the atolls and small islands of the central

Pacific, and found on a variety of hosts (Maa 1966,

Maa 1989).

Ornithoica caledonica was collected on the kagu, an

endemic bird of uncertain relationships, assigned to

the Rhynochetidae of the order Gruiformes (rails,

cranes, buttonquail and allies) (Lindsey 1993). The

kagu is restricted to La Grand Terre or the main is-

land of New Caledonia.

Ornithoctona plicata (von Olfers)

Material examined. — NEW CALEDONIA: Parc

Rivière Bleue, ix.1993-xii.1994, ex. Notou, F.

Beugnet, Y. Létocart & R. Gadet (2d ,cnc).

Remarks. — Specimens of Ornithoctona plicata were

collected on the notou, an endemic pigeon (Ducula

goliath). This hippoboscid is widespread on islands in

the Pacific and Indian Oceans and has been recorded

from about 50 genera of land and aquatic birds (Maa

& Marshall 1981). In New Caledonia, O. plicata was

previously recorded from Grande Terre and the

Loyalty Is. (Maa 1989). In New Guinea, O. plicata

has been collected on Ducula zoeae and in Vanuatu

(New Hebrides) it has been collected on D. pacifica

(Maa 1969, Maa & Marshall 1981).

ACKNOWLEDGEMENTS

Sincere thanks to F. Beugnet (Ecole Vétérinaire de

Lyon, Marcy L'Etoile) for encouraging me to exam-

ine these specimens. All specimens were collected as

part of a project supported by the crrap agency of

New Caledonia. F. C. Thompson (usnm) is thanked

for loan of specimens of O. pusilla.

131

TIJDSCHRIET VOOR ENTOMOLOGIE, VOLUME 140, 1997

REFERENCES

Cumming, J. M., B. J. Sinclair, & D. M. Wood, 1995.

Homology and phylogenetic implications of male geni-

talia in Diptera - Eremoneura. — Entomologica scandi-

navica 26: 120-151.

Delacour, J. A., 1966. Guide des oiseaux de la Nouvelle-

Caledonie. — Delachaux & Niestlé. 172 pp.

Lindsey, T., 1993. Flutter of hope for a national emblem. —

GEO Australia. 15(3): 50-58.

Maa, T. C., 1966. Studies in Hippoboscidae (Diptera). The

genus Ornithoica Rondani (Diptera: Hippoboscidae). —

Pacific Insects Monographs 10: 10-124.

Maa, T. C., 1969. Synopsis of the genera Ornithophila amd

Ornithoctona with remarks on their habitat diversification

(Diptera: Hippoboscidae). — Pacific Insects Monographs

20: 1-23.

132

Maa, T. C., 1989. 114 [Chapter] Family Hippoboscidae.

Pp. 785-789 in Evenhuis, N.L. (Ed.) Catalog of the

Diptera of the Australasian and Oceanic Regions. —

Bishop Museum Press and E.J. Brill. 1155 pp.

Maa, T.C. & A. G. Marshall, 1981. Diptera Pupipara of the

New Hebrides: Taxonomy, zoogeography, host and ecol-

ogy. — Quarterly Journal of Taiwan Museum 34(3,4):

213-232.

McAlpine, J. F., 1981. Morphology and terminology -

Adults [Chapter] 2. — In: McAlpine, J.F. et al. Manual of

Nearctic Diptera. Vol. 1. — Agriculture Canada Mono-

graph 27: vi + 1-674.

Received:20 March 1996

Accepted: 4 February 1997

J. van TOL

National Museum of Natural History, Leiden, The Netherlands

THE GENUS PROCORDULIA MARTIN IN WESTERN

MALESIA (ODONATA, CORDULIIDAE)

Descriptions and records of Malesian Odonata, 4 *)

Tol, J. van, 1997. The genus Procordulia Martin in western Malesia (Odonata, Corduliidae).

Descriptions and records of Malesian Odonata, 4. — Tijdschrift voor Entomologie 140: 133-

146, figs. 1-27. [rss 0040-7496]. Published 31 October 1997.

The species of the genus Procordulia occurring in Malaysia, the Philippines and Indonesia, excl.

New Guinea, are discussed and a key to the species is provided. P. papandayanensis is described

from Java, and P. lompobatang and P. rantemario from SW Sulawesi. These new species all be-

long to the P. sambawana group of species.

J. van Tol, National Museum of Natural History, P. O. Box 9517, 2300 RA Leiden, The

Netherlands.

Key words. — Procordulia; Malesia; Java; Sulawesi; new species.

*) Part 3: Tijdschrift voor Entomologie 138: 131-141.

All species of Procordulia Martin of the western

part of Malesia are assigned to an apparently mono-

phyletic group, the so-called ‘sambawana group. P.

sambawana Foerster was formerly considered a wide-

spread, although rather variable species (e.g. Lieftinck

1930) in which possibly several geographical races

could be distinguished. Later, Lieftinck (1977: 165)

provisionally split off the Javan specimens for which

the name P. karnyi Fraser was available. Although this

observation was cryptically published without com-

ment, this opinion was followed in the recent check-

list by Davies & Tobin (1985), but not by Tsuda

(1991). Lieftinck also mentioned the existence of two

undescribed species on Sulawesi (Celebes) (Lieftinck

1977: 167). All further studies on Corduliidae by

Lieftinck remained, however, unfinished and unpub-

lished.

In the present paper descriptions or diagnoses of

eight species from the Lesser Sunda Islands, Java,

Sumatra, Borneo, Sulawesi (Celebes) and Mindanao,

are provided. Three species are described as new to

science. This paper focuses on the species of western

Malesia, although also new species from New Guinea

are available in various collections. Also, a first at-

tempt is made to understand the relationships of the

Malesian Corduliidae at a higher level. A more de-

tailed analysis of the species groups now united in

Procordulia, and a general discussion of the Indo-

Australian genera of Corduliidae, has to await the de-

scriptions of new Papuan representatives of

Procordulia and Hemicordulia. Finally, the biogeogra-

phy of the western Malesian species is briefly dis-

cussed.

Abbreviations for museums and institutions follow

the ‘codens’ in Arnett & Samuelson (1986).

SYSTEMATIC PART

Procordulia Martin, 1907 [1906]

Type species. — Cordulia affinis Selys, 1871 [select-

ed Ris 1910: 437].

The closely related genera Hemicordulia Selys and

Procordulia can be distinguished (Martin 1907) by (a)

the presence of auriculae in Procordulia (absent in

Hemicordulia), (b) the presence of a cross-vein in the

anal triangle of Procordulia (absent in Hemicordulia),

and (c) a distinct anal angle in the hind wing of

Procordulia (rounded in Hemicordulia).

The distinguishing characters of both genera are,

however, not so clear (e.g. Watson et al 1991: 218)

anymore, especially since the discovery of several in-

termediate forms in the Pacific. The status of both

genera is further evaluated in the discussion section of

this paper.

133

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

134

Key to the males of western Malesian species of

Procordulia

1. Base of male superior appendage in lateral view

distinctly separated from the main stem (figs. 3-

4). Usually 6 Ax in hind wing ............. 2:

— Base of male superior appendage in lateral view

smoothly continuing into the main stem.

Characteristically 5 Ax in hind wing

2. Males with two Cux in hind wing, the distal Cux

forming an infratriangle (fig. 24). Superior ap-

pendage with sharp lateral tubercle. Lesser Sunda

Islands, Mindanao and ? Sulawesi

— Males with one Cux in hind wing. Superior ap-

pendage with or without lateral tubercle. Java and

S Ula Wes IEA e U. SIME ER AEN QE a 4

3. Superior appendage very long and slender, c. 2.8

mm (fig. 3). Hind wing 35-37 mm. Lesser Sunda

ISA 0 IS MEME ART P. sambawana (Foerster)

— Superior appendage more robustly built with tip

club-shaped. Hind wing 40 mm. Mindanao ...

BER OUR AEEA edet P. moroensis Lieftinck

4. Lateral tubercle on superior appendage approxi-

mately in the middle and sharp (fig. 4). Hind

wing characteristically with 5 Ax and 7 Px.

Sulawesussr ran Asa el POPE) Aon 5

— Superior appendage without lateral tubercle (fig.

12). Hind wing characteristically with 6 Ax and 8

BSE Matt ANRT ER Et Er P. karnyi Fraser

5. Superior appendage in lateral view slender (fig.

4), with lateral tubercle hardly visible in dorsal

Vo (ir) CE RE P. lompobatang sp. n.

— Superior appendage in lateral view relatively ro-

bust (fig. 5), with lateral tubercle conspicuous in

dorsal view (fig. 18) ...... P. rantemario sp. n.

6. Superior appendage with a subterminal, bluntly

shapedhtuberclei(fie. 14)MJava Reno 0 enn

NRW LL, P. papandayanensis sp. n.

— Superior appendage straight, with a lateral carina,

but without a tubercle (figs. 15-16)

7. Abdomen fusiform, relatively short. Hind wing

32-33 mm. N Borneo ... P. fusiformis Lieftinck

— Abdomen only somewhat constricted at segment

3. Hind wing 34-35 mm. Java

POONA. ita RU à RIEN P. artemis Lieftinck

VAN Tor: Procordulia of western Malesia

Procordulia artemis Lieftinck

(figs. 15, 19, 28)

Procordulia artemis Lieftinck, 1930: 159-162, figs. 23-25

[original description, habitat]. Lectotype: Kawah

Kamodjan. 19.iv.1930. Lft. in RMNH [examined]. —

Lieftinck 1933: 399-429 [biology, descriptions];

Lieftinck 1934: 431-432 [material, habitat, habits, ovipo-

sition]; Lieftinck 1971: 75 [lectotype designation]; Lief-

tinck 1977: 164 [compared with P. fusiformis].

Diagnosis. — Small and slender species. Male with

superior appendage slender, in lateral view in the

middle hardly wider that at base; base not conspicu-

ously heavier, tip not club-shaped (figs. 15, 19).

Arculus in forewing approximately midway between

Ax1 and Ax2. One Cux. Female with cylindrical ab-

domen without swollen basal segments. Appendages

c. 1.8 mm. Genital valve short, projecting caudad

from sternite 8, c. 1/4 of the length of segment 9,

shorter than wide, margin with V-shaped excision

with a depth of c. 1/4 of length of valve.

Apparently closely allied to P. fusiformis, which,

however, has a conspicuous spindle-shaped abdomen.

Superior appendages of both species very slender,

without lateral tubercle; those of artemis c. 7-8 times

as long as width in middle, and with acute tip in dor-

sal view (fig. 19), those of fusiformis c. 6 times as long

as width in the middle, with a sharp tip (fig. 22).

Ecology. — Lieftinck (1933) provides an interesting

account of the ecology of this species. Larvae were

found in swamps and crater lakes.

Remark. — Locality data of the lectotype as given by

Lieftinck (1971) are a free interpretation of the data

actually available with the specimen. This method

was also followed for many species.

Material examined. — Long series from Java

(RMNH).

Distribution. — Java, South Sumatra.

Procordulia fusiformis Lieftinck

(figs. 16, 22, 28)

Procordulia fusiformis Lieftinck, 1977: 162-164, fig. 3 [orig-

inal description, distribution].

Diagnosis. — Small size and spindle-shaped ab-

domen distinguish this species from other Procordulia

Figs. 1-10. Procordulia species. 1-3, P. sambawana, 4, P. lompobatang, 5-10, P. cf. lompobatang. — 1, anal appendages of male

holotype, dorsal view; 2, idem, right lateral view; 3, superior appendage male, left lateral view; 4, superior appendage male,

left lateral view; 5-7, female Bulluballija, abdomen left lateral view, dorsal view, and valvula vulvae in ventral view; 8-10, fe-

male Loka, abdomen left lateral view, dorsal view and valvula vulvae in ventral view.

135

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

species (Lieftinck 1977).

Male with superior appendage (figs. 16, 22) slen-

der, but not as straight as in P. artemis; base some-

what heavier, top straight, not club-shaped, without

lateral tubercle. Arculus in forewing about midway

between Axl and Ax2. One Cux in hind wing.

Apparently most closely related to P. artemis (see un-

der that species).

Material examined. — N. Borneo. Mt. Kinabalu.

Mesilau Camp. 4.iv.1964. 5000 ft. Coll. S. Kueh,

1d paratype (in RMNH, ex BMNH).

Distribution. — Northern

? Sarawak).

(Sabah,

Borneo

Procordulia karnyi Fraser

(He 11213, 21526328)

Procordulia karnyi Fraser, 1926: 472-473 [original descrip-

tion, d 9 Java, Mt Tengger]. — Lieftinck 1930: 162-164

[type discussed]; Lieftinck 1971: 95 [lectotype designat-

ed: Mts. Tengger, 1200 m, 8 Dec 1920, H. Docters van

Leeuwen]; Davies & Tobin 1985: 65 [specific status];

Tsuda 1991: 225 [as synonym of P. sambawanal.

Procordulia sambawana |pro parte]. — Lieftinck 1953: figs.

58-59 (and ? 60) [d anal appendages, genitalia, # last ab-

dominal segments].

Description

Male adult. — A relatively large and robust

Procordulia. Head densely setose, darkbrown at up-

perparts. Labrum pale yellowish white, mandibles

and labrum pale brown; clypeus olive-brown with

pale coloured transversal depression against labrum.

Frons brownish with bilobed upper parts with blue-

green metallic shine, coarsely punctate. Depressions

between frons and vertex deep; vertex high, truncat-

ed, punctulate, metallic green; occiput dark brown,

slightly convex behind, shining. Eyes brown in dried

specimens.

Synthorax rather robust, brilliant metallic green,

areas against sutures more brownish without metallic

shine, synthorax covered with long, slender, creamish

white setae; ventral side of thorax and spaces between

wings light brown. Legs long and slender;

trochanters, fore femur and posterior side of middle

and hind femur brown, legs otherwise dark brown.

Wings almost hyaline with a very slight yellowish tint

covering the whole surface, colouring somewhat more

conspicuous at wing tip and wing base; veins dark

brown. Neuration rather similar to that in P. sam-

bawana, pterostigma oblique, in fore and hind wing

2.0 mm wide; membranula brownish grey, paler part

at wing base intermediate between that of P. sam-

bawana and of papandayanensis (fig. 26). Nodal index

in forewing 6.8 | 9.6 or 6.9 | 8.7, in hind wing 8.5 |

6.8 or 8.6 | 6.7. Forewing with one cross-vein in tri-

angle, subtriangle three-celled; one Cux. Hind wing

without cross-veins in triangle; one Cux, no subtrian-

136

gle. Both wings without cross-vein in supratriangle,

no supplementary bridge cross-vein at subnodus.

Arculus oblique, in hind wing situated midway be-

tween Axl and Ax2. Anal angle of hind wing with

distinct edge, anal triangle with a short vein in its api-

cal third.

Abdomen slender, slightly constricted in the mid-

dle of segment 3, then gradually enlarged towards the

end of segment 6, where it is broadest. Basal segments

slightly swollen, much higher than wide, greatest

width 3 mm. Segment 3-5 rounded above, then sub-

triangular in diameter, from about the middle of seg-

ment 6 to the middle of segment 9 with an indistinct

median longitudinal crest. Venter flattened. Segment

10 with crest on dorsum. Auriculae small, knob-like,

brown. Coloration dark bronzed-black, first three

segments with distinct metallic green or coppery-red

shine; middle segments with slight coppery-red reflex

and segment 9-10 almost black. Dorsum of segment

1-2 covered with long setae. Sides of segment 1-2 and

basal half of 3 very shining. Ventral sides of tergites

dull brownish, their terminal fifth darkened.

Accessory genitalia on segment 2 not very prominent,

dark brown. Genital lobe subtriangular in general

outline, with acute tip; posterior hamuli somewhat

shorter than the lobe, thick at base, then constricted

and considerably narrowed, strongly curved inwards,

then outwards, straight in the median plane.

Anal appendages with superiors 2.75 mm long, in

dorsal view (fig. 21) relatively slender at base, distal

part club-shaped; base rather heavy, with a sudden

transition between the base and the main stem; on

stem no or hardly any trace of a lateral tubercle (figs.

ED):

Female. — Similar to the male, but head more

brownish, vertex brown without metallic shine.

Wings generally with more conspicuous yellowish

shine than male; nodal index forewing 5.9 | 9.6, hind

wing 8.6 | 7.7. Hind wing with one Cux. Abdomen

more robust than male, with purple shine. Basal seg-

ments slightly swollen; appendages c. 2.3 mm; genital

valve projecting caudad from sternite 8, c. 1/3 the

length of segment 9, somewhat shorter than wide

with V-shaped excision c. 2/5 the length of the valve.

Differential diagnosis. — Males of Procordulia of

Java can be distinguished by the shape of the superior

appendage (figs. 11-12, 14-15). For distinguishing

characters with P. papandayanensis, see under that

species.

Ecology. — Possibly a species inhabiting running

waters. The data of Lieftinck (1933, sub nom. P. sum-

bawana) may or may not include material of P.

karnyi. In the light of the records of adults taken in

1961 (see below), I presume that at least the ultimate

larva from E Java (Mt. Lawoe, little fall near Serangan

above Madioen, ca. 1300 m, 29 Sep 1927, leg. A.

van Tor: Procordulia of western Malesia

Figs. 11-16. Procordulia species. 11-12, 14-16, superior appendage of male in lateral view, 13, last abdominal segments of fe-

male. — 11, P. karnyi, holotype, right lateral view; 12-13, P. karnyi, Java, Baturraden; 14, P. papandayanensis, Java, Mt.

Papandajan; 15, P. artemis, Java; 16, P. fusiformis.

Vorstman) can be attributed to P. karnyi.

Other material examined. — Java: E Java, Tengger

Mis; 38) Dec 1920, He Di via. 1d. [lectotypel

(RMNH); E Java, Tengger Mts, 6 Dec 1920, H.

Docters van Leeuwen, 12 (BMNH) [not seen]; W

Java, Preanger, Sitoe Lembang, 1600 m, Dec 1936,

F. C. Drescher, 16 (BMNH); S Java, Mt. Tangkuban

Prahu, 1300-1400 m, 16 (BMNH, ex coll. Lieftinck);

Batoerraden, Mt. Slamat, ca. 2500 ft, F. C. Drescher,

PrmDec 19275 19 Apr 192952, Apr 1930, 33 TS

(BMNH, ex coll. Lieftinck); Java occ., H. Fruhstorfer,

38 19 (Museum Berlin) [not seen]; W Java, Situ

Lembang, 1600 m, Dec 1936, F. C. Drescher, 5$

(RMNH); W Java, slopes of Mt Gedeh and Pangrango,

series of both sexes (RMNH); Mt. Patuha,

Patuhawattee, 1750 m, 3 Jun 1935, J. van Marle, 19

(RMNH); E Java, Mt. Ardjuno, Djunggo, 1600 m, Jan

1936, M. E. Walsh, 16 (RMNH); E. Java, Mt. Lawu,

2000 m, Tjemorosewu, 24 Feb 1961, ‘Vole en forêt

vierge très près du sol, vol rapide, capture difficile,

137

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 17-22. Superior anal appendages of male Procordulia. — 17, P. lompobatang; 18, P. sambawana; 19, P. artemis, 20, P. pa-

pandayanensis, 21, P. karnyı, 22, P. fusiformis. Scale bar 1 mm.

semble ici assex commune’, P. Jauffret, 18 19

(RMNH, ex coll. A. Heymer). — Sumatra. W Sumatra,

Lebong Tandai, 1920-23, C. J. Brooks, 16 (headless)

(BMNH); Central W Sumatra, Mt. Dempo, Gunung

Agung Estate, 1500-2000 m, 11 Sep 1941, W. C.

Verboom, 14 (bred from larva, emerged Bogor 2 Jan

1942) (RMNH).

Distribution. — Java, Sumatra.

138

Procordulia lompobatang sp. n.

(figs. 4-10, 17, 23, 28)

Procordulia irregularis Martin, 1907 [1906]: 16-17 [original

description] [partim, male Celebes only].

? Procordulia sumbawana. — Martin 1907 [1906]: 17 [par-

tim].

Procordulia sambawana. — Lieftinck 1953: figs. 63-64 [d

anal appendages].

Procordulia celebensis Lieftinck, ms name.

Type material. — Holotype d: ‘H. Fruhstorf.

Celebes mer.’ [white, round, handwritten], “Neonyx

nitens [white, handwritten in unknown hand],

‘Procordulia sumbawana (Frst) / det vdWleele]’

[white with black margin, handwritten], ‘det MA

Lieftinck 75 / Procordulia celebensis Lieft. / holotype’

[white, partly in print], ‘RMNH Leiden / Procordulia

lompobatang Van Tol / det. J. van Tol, 1988’ [white,

partly in print], ‘Holotype’ [red, in print] in RMNH. —

Paratypes: Lansbg Celebes ?, 1d in 1RSN [this speci-

men is also a paralectotype of Procordulia irregularis

Martin]; Celebes mer, 16 (Fruhstorfer) in RMNH; S

Celebes, Lompa Battau, 3000’, Mar 1896, 46

(Fruhstorfer) in MNHN; SW Celebes, Mt Lompo-

batang, Parasalawaki, 1600-1650 m, 3, 4 and 8 Jul

1936, 3d (L. J. Toxopeus) in RMNH; S Celebes,

Kanrapia, 3 km E of Buluballija, 2000 m, 30 Oct

1965, 28 (R. Straatman) in RMNH; SW Celebes,

Lompobatang complex, 1100 m, Malino resthouse, 2

Jun 1982, 14 (M. A. Lieftinck) in RMNH.

Other material. — Celebes mer, 1 2 (Fruhstorfer) in

rmnh; S Celebes, Lompa Battau, 3000’, Mar 1896,

12 (Fruhstorfer) in MNHN; S Celebes, Loka, Mt

Lompobatang, 1200 m, May 1949, 19 (C. Franssen)

in RMNH; S Celebes, 82 km ESE of Makassar,

Buluballija, 1700 m, 12 Oct 1965, R. Straatman,

12(R. Straatman) in RMNH.

I have also examined in sMFD Frankfurt 116 and

22, with various labels (nos. O-37065 to O-37075,

37104 and 37100, ex collection Ris), but probably all

belonging to the series collected by H. Fruhstorfer on

the Lompobatang in March 1896. I have not exam-

ined these specimens since 1986, and thus did not in-

clude them in the type series. I have not included any

females in the paratype series, since at least two types

of females can be distinguished (see below). It is not

clear whether one or more species are involved.

Description

Male adult, holotype. — A relatively large and ro-

bust Procordulia. Head densely setose, dark brown at

upper parts. Labium pale yellowish white, mandibles

and labrum yellowish, labrum with subtriangular,

olive-brown basal marking; clypeus olive-brown.

Frons yellowish with the bilobed upperparts with

blue-green metallic shine, coarsely punctate.

Depressions between frons and vertex deep; vertex

high, truncate, punctulate, metallic green; occiput

dark brown, slightly convex behind, very shining.

Eyes brown in dried specimens.

Synthorax rather robust, brilliant metallic green,

covered with long, slender, creamish white setae; ven-

tral side of thorax and spaces between wings light

brown. Legs long and slender, trochanters and anteri-

or side of fore and middle legsbrown; legs otherwise

darkbrown. Wings almost hyaline with a very slight

van Tor: Procordulia of western Malesia

yellowish tint covering the whole surface, although

somewhat more conspicuous towards the wing base.

Neuration very similar to that in P. sambawana; neu-

ration dark brown; pterostigma oblique, in fore and

hind wing 1.85 mm wide; membranula brownish

grey, somewhat lighter at extreme base. Nodal index

of holotype fore wing 6.8 | 8.6, hind wing 7.5 | 5.7.

Fore wing with one cross-vein in triangle, subtriangle

three-celled; one Cux. Hind wing without cross-veins

in triangle; one Cux (no subtriangle). Arculus

oblique, in hind wing in all specimens situated mid-

way between Axl and Ax2. Anal angle (fig. 23) in

hind wing with distinct edge, triangle with a short

vein in its apical third. Membranula dark-grey, only

somewhat paler at its very base.

Abdomen slender, slightly constricted in the mid-

dle of segment 3, then gradually enlarged towards the

end of segment 6, where it is broadest. Basal segments

slightly swollen, much higher than wide, greatest

width 3.25 mm. Segment 3-5 rounded above, then

subtriangular in diameter, from about the middle of

segment 6 to the middle of segment 9 with an indis-

tinct median longitudinal crest. Venter flattened.

Segment 10 with crest on dorsum. Auriculae small,

knob-like, brown. Coloration dark bronzed-black,

first three segments with distinct metallic green shine;

middle segments with slight coppery-red reflex and

segment 9-10 almost black. Dorsum of segment 1-2

covered with long setae; sides of segment 1-2 and

basal half of 3 very shining. Ventral sides of tergites

dull brownish, their terminal fifth darkened.

Accessory genitalia on segment 2 not very prominent,

dark brown. Genital lobe subtriangular in general

outline, with acute tip; posterior hamuli of about

equal length as the lobe, thick at base, then constrict-

ed and considerably narrowed, strongly curved in-

wards, then outwards, straight in the median plane.

Anal appendages with superiors 2.9 mm long, in

dorsal view (fig. 17) relatively slender at base, distal

part more knob-like; a small tubercle just visible

about halfway in lateral view (fig. 4), the distal part

beyond the tubercle somewhat widened, especially at

ventral side.

Female. — Generally as male. Head as male, but

vertex with metallic shine; occiput castaneous.

Synthorax as male; hind wings in teneral specimens

with yellow tint, adult female wings brownish yellow,

especially against the wings. Neuration very similar to

P. sambawana, including two Cux in the hind wing

(only one female with one Cux in left hind wing);

frontal side of pterostigma 2.5 mm (Loka) or 2.0 mm

(Buluballija); arculus oblique, in hind wing midway

between Axl and Ax2. Abdomen stoutly built, hard-

ly constricted. Genital valve and appendages with

much variation between the specimens. Loka speci-

men (figs. 8-10) with genital valve protruded from

189

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 23-26. Base of hind wing of male Procordulia. — 23, P. lompobatang, 24, P. sambawana; 25, P. papandayanensis 26, P.

karnyi.

140

abdomen with angle of 60°, length c. 1/2 of segment

9, V-shaped excision c. 1/3 of valve; anal appendages

long and slender, 2.7 mm. Buluballija specimen (figs.

5-7) with much larger valve, 2/3 the length of seg-

ment 9, V-shaped excision less deep; anal appendages

straight and shorter, 1.8 mm long.

Ecology. — A species of mountainous areas.

Distribution. — SW Sulawesi (Celebes) (fig. 28).

Procordulia moroensis Lieftinck

fig. 28

Procordulia moroensis Lieftinck, 1977: 164-167, figs. 4-5

[original description, distribution Mindanao]. Holotype

3, Mindanao I., Upper Eden, 1000 m, 2.xi.1965 (in

USNM) [not examined].

Diagnosis. — Adequately described by Lieftinck

(1977).

Remarks. — In the original description, Lieftinck

(1977) notes that ‘this stout species approaches the

two Procordulia occurring in Celebes, both of the P.

sambawana group, …. In the characters used here it

stands apart from the Sulawesi species by the 6 Ax

veins in the hind wing. The superior anal appendages

are indeed very similar to the Sulawesi taxa.

Remarkably, it seems to be especially close to P.

rantemario, which was unknown to Lieftinck. In lat-

eral view, however, the superiors are quite different,

in P. moroensis more or less club-shaped, and P.

rantemario smoothly widening nearly from the base.

P. rantemario is also c. ten percent smaller than P. mo-

roensis.

Procordulia papandayanensis sp. n.

figs. 14, 20, 25, 28

? Procordulia sumbawana; Lieftinck 1933: 429 [larvae] [at

least part of the material was collected on sites where the

present species has been found].

Type material. — Holotype d: ‘W Java, 2250 m /

Mt. Papandajan / pondok, 9.vii.1934 / M. A.

Lieftinck’, with on reverse side ‘P. sambawana | dt.

Lieftinck’ [white label, both texts in Lieftinck’s

hand], ‘Museum Leiden / ex verz. M. A. Lieftinck’

[printed], ‘Holotype’ [red, printed], ‘RMNH Leiden

holotype / Procordulia papandayanensis Van Tol / det

J. van Tol, 1996’ [partly printed, partly handwritten]

UvT 6787]. — Paratypes: All specimens from W Java.

Gunung Gedeh, c. 2000-2600 m, 116; Gunung

Papandajan, 2300-2600 m, 74; Gunung Tangkuban

Prahu, 1400 m, 16 (all RMNH); W Java, Mt.

Papandajan, 5000 ft, 23 Nov 1929, 18 (C.

Longfield) in BMNH (det. P. sumbawana by C.

Longfield).

van Tor: Procordulia of western Malesia

Description

Male. — A relatively large and robust species of

Procordulia, stouter than P. sambawana, although

with some variation. Head densely setose, darkbrown

at upperparts, paler otherwise. Labium and man-

dibles pale yellowish white, labrum more brownish.

Clypeus olive-brown with a rectangular, paler depres-

sion along labrum. Frons light to olive-brown, or

dark yellowish, with the bilobed upperparts with

blue-green metallic shine, coarsely punctate.

Depression between frons and vertex deep; vertex

high, truncated, punctulate, with only inconspicuous

metallic green shine, less than in other species; oc-

ciput chestnut brown, slightly convex posteriorly,

very shining, densely setose, rear side with a dense

fringe of long setae. Eyes brown in dried specimens.

Synthorax robust, metallic green lees brilliant than

related species, covered with long, slender, creamish

white setae; ventral side of thorax and spaces between

wings light brown. Legs long and slender, trochanters

brown, legs otherwise dark brown. Wings almost hya-

line with a yellowish tint covering the whole surface,

especially near the wing base. Neuration dark brown;

pterostigma oblique, in fore and hind wing c. 2.0 mm

wide; membranula off-white, darker posteriorly.

Nodal index fore wing 6.8 | 8.6, in hind wing 7.5 |

5.7 in nearly all specimens. Fore wing with one cross-

vein in triangle, supratriangle without cross-veins,

subtriangle three-celled, one Cux. Hind wing with-

out cross-veins in triangle and supra-triangle; one

Cux. Arculus oblique, in hind wing in all specimens

situated midway between Axl and Ax2. Anal angle

(hind wing) with distinct edge; anal triangle with a

short vein in its apical third. Supplementary bridge

cross-vein at subnodus.

Abdomen more robust than in P. sambawana,

slightly constricted in the middle of segment 3, then

gradually somewhat enlarged toward the end of seg-

ment 6, where it is broadest. Basal segments slightly

swollen, much higher than wide, greatest width 3.2

mm. Segment 3-5 rounded above, then subtriangular

in diameter, from about the middle of segment 6 to

the middle of segment 9 with an indistinct median

longitudinal crest. Venter flattened. Segment 10 with

crest on dorsum. Auriculae small, knob-like, brown.

Coloration dark bronzed-black, first three segments

with metallic shine greenish or coppery-red; middle

segments with slight coppery-red reflex and segment

9-10 almost black. Dorsum of segment 1-2 sparsely

covered with long setae. Sides of segment 1-2 and

basal half of 3 very shining. Ventral sides of tergites

dull brownish, their terminal fifth darkened.

Accessory genitalia on segment 2 not very promi-

nent, dark brown. Genital lobe subtriangular in gen-

eral outline, with acute tip; posterior hamuli some-

what shorter than the lobe, thick at base, then

141

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

constricted and considerably narrowed, strongly

curved inwards, then outwards, straight in the medi-

an plane.

Anal appendages with superiors 2.5 mm long in

dorsal view; in dorsal view (fig. 20) relatively slender

at base, slightly widening from base to apex, the tips

somewhat curved outwards; in lateral (fig. 14) view

transition of base to stem smooth without distinct an-

gle, at c. 1/4 from the top a bluntly shaped ventral tu-

bercle; top club-shaped.

Female. — Unknown.

Differential diagnosis. — The males of Procordulia

species of Java can be distinguished by the shape of

the superior appendage. In P. papandayanensis the top

is more or less club-shaped with a sub-terminal ven-

tral blunt tubercle, which is absent in the other

species. In P. karnyi the appendage is rather robust,

while the transition of the base to the stem is charac-

teristically distinct as in P. sambawana. The ap-

pendage of P. artemis is much more slender and tubi-

form.

Distribution. — Java, where is seems to be confined

to higher altitudes.

Procordulia rantemario sp. n.

(figs. 5,10, 18, 28)

Type material. - Holotype d: Indonesia. SW

Sulawesi. 30 km NE of Enrekang. SW slope of Gn

Rantemario. Sg. Gowa Sarumpa'pa at Gowa camp

(Pos 2). Fast flowing stream, rapids, ponded sites.

Primary rain forest. Shade; w=3-5 m, d=10-30 cm.

Altitude 1800 m. 3°24'15"S 120°00'30"E. 6-9 Nov

1993. Leg. J. van Tol [JvI no 1730] (in RMNH). —

Paratypes 16 39, same data [JvT 1731-1734] (16

19 in MBBJ, other specimens in RMNH).

Description

A relatively small and slender representative of the

P. sambawana group.

Male. — [JvT 1730]. Head and thorax densely se-

tose. Head: Labium yellowish white, mandibles and

labrum yellowish, labrum with subcircular brown

base, rest orange-yellow; frons olive-brown with

bilobed dorsal parts brownish black with blue-green

metallic shine, coarsely punctate; depression between

frons and vertex deep; vertex brown with hardly de-

tectable metallic shine, high, truncate, punculate; oc-

ciput brown, slightly convex, finely punctulate. Eyes

in dried specimens dark green below and brown

above.

Synthorax slender, greenish with blue metallic

shine; ventral side of thorax pale brown. Legs long

and slender; trochanter of fore and middle leg and an-

terior side of tranchanter of hind leg middle brown;

142

anterior side of femora middle brown (in hind leg re-

duced to longitudinal stripe); remaining parts of legs

brownish black. Wings hyaline with very slight yel-

lowish tint; anal triangle and cubital space of hind

wing with dark yellow tint; neuration similar to P.

sambawana, nervature dark brown, pterostigma

oblique, in fore and hind wing anteriorly approxi-

mately 1.9 mm wide; membranula isabel coloured at

base, more brownish near apex; nodal index fore wing

6.8 | 9.5, hind wing 7.5 | 5.6; triangle of fore wing

with one cross-vein, subtriangle three-celled, one Cux;

triangle of hind wing without cross-vein, one Cux (no

subtriangle); Arculus oblique, in fore wing closer to

Ax2 than to Ax1; anal angle in hind wing with distinct

edge, anal triangle with short vein in its apical third.

Abdomen slender, slightly constricted at the mid-

dle of segment 3, then gradually enlarged towards the

end of segment 6, where it is broadest. Segments 2-9

with conspicuous green metallic shine; basal segments

slightly swollen, higher than wide; from segment 4-9

with lateral carina, making segments triangular in di-

ameter; from middle of segment 4 to middle of seg-

ment 10 with (especially rostrally rather inconspicu-

ous) dorsal carina, base of segment 10 with

conspicuous crest; auriculae small, knob-like, brown;

coloration segments 1-9 bronzed black with green

metallic shine on all segments, but segment 10 rather

dull and hardly metallic; dorsum of segment 1-2 with

long setae; dorsal sites of tergites with brownish yel-

low markings against lateral carina, more or less cen-

tral on segment 4, covering anterior three-quarters of

segments 5-7, basal half of segment 8 and basal quar-

ter of segment 9; ventrum most light brown.

Accessory genitalia on segment 2 not very prominent,

dark brown; genital lobe elongate subtriangular, more

straight than in P. lompobatang, tip acute; posterior

hamuli in ventral view of about the same length as the

lobe, solid at base, then constricted and considerably

narrowed, curved inwards, the tip curved outwards as

well as towards the body. Anal appendages with supe-

rior approximately 2.5 mm long, in dorsal view (fig.

18) much more robust than in sambawana or lompo-

batang, basal half more or less straight up to a con-

spicuous extero-lateral tooth; the distal half curved

inwards, only somewhat widening with distal one-

third approximately parallel; in lateral view (fig. 5)

with conspicuous edge at base (as in lompobatang),

side margin running distally in lateral tooth, knob-

like distal part conspicuously widening, starting prox-

imally to lateral tubercle; inferior appendage rather

short.

Measurements: Hind wing 36 mm; abdomen incl.

appendages 36 mm.

Variation: Male paratype has wing formula

forewing 5.8 | 8.5, hindwing 7.5 | 5.7. Other charac-

ters similar as in holotype.

Female. — Generally as the male, but much more

robust. Head as male, but labium rather pale, yellow-

ish white; wings hyaline without conspicuous yellow

tint; neuration similar to male, also one Cux in hind

wing (compare P. lompobatang); frontal side of ptero-

stigma ca. 2.0 mm; Arculus oblique, in hind wing

halfway between Axl and Ax2. Abdomen rather

stout, only somewhat constricted at posterior half of

segment 3 and base of segment 4. Genital valve rather

short, covering approximately two-fifths of entral side

of segment 9, emarginate with emargination approxi-

mately one-third of length of valve; anal appendages

straight, ca. 2 mm.

Etymology. — Rantemario, name of the mountain

of the type locality; a noun in apposition.

Distribution. — Only known from the type locality,

Gn. Rantemario, one of the tops in the Latimojong

mountains.

Procordulia sambawana (Foerster)

figs. 1-3, 18, 24, 28

Somatochlora sambawana Foerster, 1899: 64-65 [holotype

d, Insel Sumbawa / zwischen Floris / u. Sumba / d.

Pagenstecher don. 1898] [ummz, examined].

Procordulia sambawana (Foerster). — Lieftinck 1936: 148-

149 [material Lombok and Sumba only]; Lieftinck 1953:

191-193, figs. 61-62 [partim, only records Lesser Sunda

Islands] [references, material Sumbawa, geographical

variation]; Lieftinck 1954: 122 [partim, only specimens

Lesser Sunda Islands] [synonymy, range, ecology].

Procordulia sumbawana (Foerster) [incorrect emendation];

Martin 1907: 17 [partim, only records Lesser Sunda

Islands]; Lieftinck 1930: 162-164 [partim, only records

Lesser Sunda Islands] [comparison with P. artemis].

Other material examined. — Lesser Sunda Islands,

W Flores, Rana Mese, 1300 m, 5-8 Apr 1958, A. M.

R. Wegner, 96 (RMNH); Soemba, Fruhstorfer, 16

(IRSN); Insel Sumbawa bei Flores, 1898, Dr. A.

Pagenstecher, 19 (ummz) [with Cux 2 | 1]; Lombok,

Sambalun, 4000’, Apr 1896, H. Fruhstorfer,

19 (RMNH). — [Doubtful record]: Celebes, Bua

Kraeng, 5000 ft, Feb 96, Fruhstorfer, 16 (BMNH)

[with Lieftinck’s label ‘P. spec. nov. Celebes’].

Lieftinck examined: Lombok, Sapit 2000’, Apr

1896, H. Fruhstorfer, 24 19 (zmus).

Description

Male. — A relatively large and robust Procordulia,

although with some variation. Head densely setose,

dark brown at upperparts, paler otherwise. Labium

and mandibles pale yellowish white, labrum more

brownish. Clypeus olive-brown with a rectangular,

paler depression along labrum. Frons light brown or

dark yellowish with the bilobed upperparts with blue-

green metallic shine, coarsely punctate, Depressions

between frons and vertex deep; vertex high, trunctat-

van Tot: Procordulia of western Malesia

ed, punctulate and hardly shining, metallic green; oc-

ciput chestnut-brown, slightly convex posteriorly,

very shining, rear side with a dense fringe of long se-

tae. Eyes brown in dried specimens.

Synthorax relatively slender, brilliant metallic

green, covered with long, slender, creamish white se-

tae; ventral side of thorax and spaces between wings

light brown. Legs long and slender; trochanters and

anterior side of fore and middle legs brown; legs oth-

erwise dark brown. Wings almost hyaline with a very

slight yellowish tint covering the whole surface.

Neuration with nervature darkbrown; pterostigma

oblique, in fore and hind wing c. 2.0 mm wide; mem-

branula brownish grey, off-white in basal 1/6th.

Nodal index in fore wing 6.9 | 9.6, in hind wing 9.6 |

5.8 or 8.6 | 6.8. Fore wing with one cross-vein in tri-

angle, supratriangle without cross-veins, subtriangle

three-celled, one Cux. Hind wing without cross-veins

in triangle and supra-triangle; characteristically with

two Cux. Arculus oblique, in hind wing in all speci-

mens situated in the middle between Ax1 and Ax2.

Anal angle (hind wing) with distinct edge; triangle

with a short vein in its apical third. Supplementary

bridge cross-vein at subnodus.

Abdomen slender, slightly constricted in the mid-

dle of segment 3, then gradually somewhat enlarged

towards the end of segment 6, where it is broadest.

Basal segments slightly swollen, much higher than

wide, greatest width 2.75 mm. Segment 3-5 rounded

above, then subtriangular in diameter, from about the

middle of segment 6 to the middle of segment 9 with

an indistinct median longitudinal crest. Venter flat-

tened. Segment 10 with crest on dorsum. Auriculae

small, knob-like, brown. Coloration dark bronzed-

black, first three segments with distinct metallic green

shine; middle segments with slight coppery-red reflex

and segment 9-10 almost black. Dorsum of segment

1-2 sparsely covered with long setae. Sides of segment

1-2 and basal half of 3 very shining. Ventral sides of

tergites dull brownish, their terminal fifth darkened.

Accessory genitalia on segment 2 not very prominent,

dark brown. Genital lobe subtriangular in general

outline, with acute tip; posterior hamulus shorter

than the lobe, thick at base, then constricted and con-

siderably narrowed, strongly curved inwards, then

outwards, straight in the median plane.

Anal appendages with superiors 3.5 mm long, in

dorsal view relatively slender at base, widening at c.

1/5 from the base and distal 4/5th more or less

straight; a sharp, tooth-like tubercle at about midway

in lateral view.

Distribution. — Lombok, ? Sumba, Sumbawa,

Flores, ? Sulawesi.

Differential diagnosis. — Can be distinguished from

other species in this group by the virtually straight su-

143

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

perior anal appendage with a sharp lateral tubercle ap-

proximately midway, and the presence of two Cux in

the hindwing.

Remarks. — The present data are insufficient to de-

cide whether P. sambawana is indigenous to Sulawesi.

It is not unlikely that the Sulawesi male mentioned

above was mislabelled. Mislabelling of Indonesian

material collected by Fruhstorfer is, unfortunately,

common (see Van Tol 1987: 163). On the other

hand, the collections of apparently genuine Sulawesi

females of Procordulia are heterogeneous. All speci-

mens have two Cux in the hind wing, a character oc-

curring otherwise only in the males of Procordulia

sambawana. Lieftinck considered the Sulawesi speci-

men distinct from P. sambawana, but I can not dis-

tinguish this specimen from material from the Lesser

Sunda Islands.

Discussion

Affinities of genera

Several efforts have been made in the past (e. g.

Martin 1907, Lieftinck 1953, 1977) to understand

the systematics of the genus Procordulia Martin and

related genera, or the Corduliidae in general

(Needham 1908, Williamson 1908). Nevertheless,

the relationships of the corduliine genera, and those

of the species included in Procordulia plus

Hemicordulia, remain poorly understood (e.g.

Watson 1981: 1148).

It is also unclear which character or characters are

diagnostic for each genus. In the Pacific islands, the

easternmost part of their ranges, both genera cannot

be kept apart based on the characters used so far.

Lieftinck (1977) also notes that females are difficult

to assign to any of both genera. Besides, the species of

the Australian region, particularly several included in

Procordulia so far, differ from their Malesian con-

generics in many ways. Especially the position of the

New Zealand Procordulia grayi (Selys, 1871), origi-

nally described in the genus Cordulia, subgenus

Epitheca and later included in Somatochlora by

Martin (1907), has been questioned already several

times (see Rowe 1987). The Procordulia species of

Malesia, on the other hand, are quite uniform and

presumably form a monophyletic group. Their pat-

tern of speciation may contribute to our understand-

ing of the biogeographical history of this region.

Although a more detailed systematic and biogeo-

graphical analysis has to wait for the future when also

the species of New Guinea, Australia and the islands

in the Pacific have been included, a few notes may

summarize our present state of understanding.

Most indo-australian Corduliinae are united in

Procordulia and Hemicordulia. A few oriental species

with obvious Palaearctic relationships are attributed

144

to Cordulia Leach, while several aberrant Malesian

species are arranged in monospecific genera as

Antipodochlora Fraser, Guadalca Kimmins, Hetero-

naias Needham & Gyger and Pentathemis Karsch (see

Watson 1969). The relationships of these genera are

unknown, since they have been characterized by aut-

apomorphies only. As noted above, the difference be-

tween the two genera that mainly concern us here is

less distinct in the eastern parts of their ranges.

Hemicordulia mumfordi Needham from the

Marquesas Islands is remarkable in this respect, since

it has a cross-vein in the anal triangle. Based on a pre-

liminary cladistic analysis, it appears that this cross-

vein, also being more oblique, is not homologous to

that in Procordulia. For this analysis the South-

American genus Rialla was used as outgroup. The

choice of a remotely related outgroup in phylogenet-

ic analysis is certainly not without problems, and the

results of this cladogram have to be used with care.

The genus Hemicordulia is aberrant within the

Corduliidae for two characters, viz. it lacks an angu-

lated base of the male hind wing, as well as auricles

(oreillets). The characters should be interpreted as

autapomorphies. Hemicordulia is not restricted to

Australia and Southeast Asia. H. virens (Rambur) oc-

curs on Mauritius, H. similis (Rambur) on Mada-

gascar and the Seychelles, while H. asiatica is wide-

spread and known from Uganda, South India,

Ceylon and Assam (Pinhey 1962). Fraser (1949) at-

tributed the occurrence of Hemicordulia on the

Mascarene islands and Madagascar to the strong mi-

gratory tendencies of some species. He also states that

‘there is good evidence to show that they [A. virens

and A. similis] are lineal descendents of H. asiatica.

Both remarks would be in support of an Indo-

Australian origin of the genus, with westward migra-

tion in a later phase. The preliminary analysis seems

to indicate that Procordulia grayi and P. jacksoniensis

are not closely allied to Procordulia + Hemicordulia (as

defined by their type species).

Biogeographical notes

The distribution of the western Malesian species of

Procordulia is illustrated in fig. 27. The genus is

known from the southern tip of Sumatra, Java,

Lombok, Sumba, Sumbawa and Flores, the south-

western peninsula of Sulawesi, the northern part of

Borneo and the island of Mindanao. This distribu-

tion pattern may be compared with the comments on

the biogeographical relationships between the

Philippines and Sulawesi as summarized by Vane-

Wright (1990) and of the relationship of microconti-

nents in this region by Michaux (1991, 1994, 1996).

Relationship between Sulawesi and the Philippines

have been proposed earlier by Wallace and by Croizat

(1958), who suggested a relation between Sulawesi

Procordulia

artemis

fusiformis

karnyi

lompobatang

moroensis

papandayanensis

rantemario

sambawana

Sp oane Lo

Fig. 27. Distribution of western Malesian Procordulia species.

and the Philippines via Halmahera. Vane-Wright

demonstrates the faunal regions of the Philippines,

and postulates that the Philippines, rather than

Sulawesi, will provide the key to our understanding of

the evolution of Southeast Asia. His conclusion is

that the Philippines have a derived Sunda Shelf fau-

na. Due to recent geological uplift the land area of the

islands is considerably larger now than before. These

conditions may have induced exchanges between fau-

nal elements from the Philippines to Sulawesi and

from Sulawesi to the Moluccas.

Studies of McCabe & Cole (1989), on the other

hand, indicate that the Banda, Sulu and Celebes Sea

are composed of old oceanic crust surrounded by

younger, actively spreading oceanic crust. Michaux’

conclusion of these data is, that western Sulawesi,

eastern Borneo and parts of Mindano formed a single

tectonic unit and share a common history. This mi-

crocontinent would be of Gondwanic origin, and

have a collision age of (possibly) late Cretaceous.

Thirdly, also more recent climatic changes may

have influenced the possibilities for dispersal of

mountainous species. It is largely unknown what

van Tor: Procordulia of western Malesia

kind of vegetation dominated during the Pleistocene

glaciations, but lower sea levels may have opened cor-

ridors for species now confined to higher altitudes.

The relative close relationships of the species of the

P. sambawana group, as based on morphological

characters, suggest a Pleistocene rather than a

Cretaceous time of splitting of the lineages.

ACKNOWLEDGEMENTS

This paper started on annotations by the late Dr.

M. A. Lieftinck. I have carefully re-examined and

measured all specimens myself, and all conclusions in

this paper are mine. I gratefully acknowledge the kind

help and hospitality of Stephen J. Brooks (BMNH),

and Mark O'Brien (UMMz).

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Received: 3 January 1997

Accepted: February 1997

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Tijdschrift voor Entomologie

Volume 140, no. I

Articles

129

133

P. N. Bellinger & W. N. Ellis

Generic names of Collembola: Supplement 1984-1996.

M. V. Kozlov

The new aurora species-group of the genus Nemophora Hoffmannsegg, with two

new species from Taiwan (Lepidoptera, Adelidae)

N. Nieser, H. Zettel & P. P. Chen

Ten new species of Rhagovelia (Heteroptera: Veliidae) from Sulawesi (Indonesia).

J. T. Polhemus

Seven new species of Hydrotrephes China (Helotrephidae: Heteroptera) from

Sulawesi.

R. Puplesis, A. Diskus & E. J. van Nieukerken

Stigmella divina sp. n., a remarkable species from Turkmenistan and Turkey

(Lepidoptera, Nepticulidae).

O. Rohfritsch

Morphological and behavioural adaptations of the gall midge Lasioptera arundinis

(Schiner) (Diptera, Cecidomyiidae) to collect and transport conidia of its fungal

symbiont.

L. M. Roth

The cockroach genera Pseudothyrsocera Shelford, Haplosymploce Hanitsch, and

Episymploce Bey-Bienko (Blattaria: Blattellidae, Blattellinae).

R. Schouten

Revision of the genus Metaeuchromius Bleszynski (Lepidoptera: Pyralidae:

Crambinae).

B. J. Sinclair

A new species of Ornithoica Rondani from New Caledonia (Diptera: Hippoboscidae).

J. van Tol

The genus Procordulia Martin in western Malesia (Odonata, Corduliidae).

Book review

128

B. A. Bengtsson, 1997. Scythrididae. — Microlepidoptera of Europe (P. Huemer,

O. Karsholt & L. Lyneborg eds.) vol. 2. [E. J. van Nieukerken]

Published 31 October 1997 ISSN 0040-7496

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A journal of systematic and evolutionary

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‚Netherlands Journal of Entomology

Published by the Netherlands Entomological Society

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À journal of systematic and evolutionary entomology since 1858

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APR 2 7 1998

Paur L. TH. BEUK

A HARVARD

REVISION OF THE RADHA GROUP OF THE GENUS INIVERSITY

PLATYLOMIA STAL, 1870 (HOMOPTERA, CICADIDAE)

Beuk, P. L. Th., 1998. Revision of the radha group of the genus Platylomia Stal, 1870

(Homoptera, Cicadidae). — Tijdschrift voor Entomologie 140 [1997]: 147-176, figs. 1-51, tab.

1. [rssN 0040-7496]. Published 26 March 1998.

The name radha group is proposed for a monophyletic group of six species of the genus

Platylomia Stal, 1870. The species are found in India, Bhutan, Nepal, Indo-China, Peninsular

Thailand, China, and Taiwan. Five species are redescribed [P. bivocalis (Matsumura, 1907),

P. bocki (Distant, 1882), P. ficulnea (Distant, 1892), P. pendleburyi Moulton, 1923, and P.

radha (Distant, 1881)] and one species (P. malickyi) is described as new. P. operculata Distant,

1913, and P. similis (Distant, 1888) are brought into synonymy with P. radha. Lectotypes are

designated for Dundubia bocki, Cosmopsaltria ficulnea, P. operculata, D. radha, and D. similis.

A key to the males and distribution maps of the species are presented. The features character-

ising the P. radha group and the relations between the species in the group are briefly dis-

cussed.

P. L. Th. Beuk, Institute for Systematics and Population Biology (Zoological Museum),

University of Amsterdam, Plantage Middenlaan 64, NL-1018 DH Amsterdam, Netherlands.

E-mail: [email protected]

Key words. — Platylomia, radha group; phylogeny; taxonomy; new species; Southeast Asia.

The genus Platylomia was erected by Stal (1870) as

a subgenus of Cosmopsaltria Stal, 1866, to accommo-

date the single species Cicada flavida Guérin-Méne-

ville, 1834, that was distinguished from the species of

Cosmopsaltria sensu stricto by a broader thorax and

more undulating lateral margins of the thorax. Distant

(1905) raised Platylomia to generic level, redefined it

and gave the following diagnosis: head as broad as or

broader than anterior margin of mesonotum; head as

long as or just shorter than distance between eyes;

pronotum as long as distance between anterior margin

of mesonotum and ‘base’ of cruciform elevation; later-

al margin of pronotal collar always with a tooth that

usually is pointed; abdomen much longer than length

of head, pronotum and mesonotum; timbal organs

completely covered; rostrum just reaching between to

beyond posterior coxae; opercula curved to the lateral

part of the abdomen, elongate with a proximal con-

striction and a rounded or attenuate apex; and tegmi-

na and wings hyaline but often infuscate on veins.

Based on these diagnostic characters, a large variety

of species has been gathered in the genus Platylomia,

notwithstanding the fact that not all species fully fit

the diagnosis, and that a lot of variation can be ob-

served in the structure of the genitalia (compare e.g.

Hayashi 1978: fig. 24 [P. saturata (Walker, 1858)]

and fig. 4 below).

The genus Platylomia is currently placed in the

subtribe Dundubiaria of the tribe Dundubiini to-

gether with the genera Orientopsaltria Kato, 1944,

Dundubia Amyot & Serville, 1843, Macrosemia Kato,

1925, Meimuna Distant, 1905, Haphsa Distant,

1905, Ayesha Distant, 1905, and Khimbya Distant,

1905 (Duffels & Van der Laan 1985). The genus

contains about 35 species (Metcalf 1963; Duffels &

Van der Laan 1985; Beuk 1996) from the Southeast

Asian mainland, the Greater Sunda Islands, Sulawesi,

and the Philippines. The genus has not yet been sub-

ject of a phylogenetic study and it is not clear whether

it represents a monophyletic unit or not. However, it

appears that at least some monophyletic groups with-

in Platylomia can be recognised. In the present paper

the species of one such group are revised and the

‘Platylomia radha group’ is introduced to accommo-

date them. The group at present comprises six species

(including one new) distributed with certainty in

India, Bhutan, Nepal, Indo-China (Burma,

Thailand, Laos, Cambodia, Vietnam), Peninsular

Thailand, China, and Taiwan. Records from Japan

could not be confirmed.

This paper is part of a study into the phylogeny

and biogeography of Platylomia and the other genera

of the Dundubiaria. The results from the phylogenet-

ic reconstruction below are too preliminary to draw

147

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

any conclusions about the biogeography of this

group.

MATERIAL AND METHODS

References in the literature to species treated below

were checked whenever possible. Older references

were traced using Metcalf (1963) and Duffels & Van

der Laan (1985). References that could not be

checked are marked with an asterisk (*).

The following abbreviations were used for collec-

tions mentioned in the descriptions:

BMNH Natural History Museum, London (former

British Museum Natural History)

BPBM Bernice P. Bishop Museum, Honolulu

CASC California Academy of Sciences, San Fran-

cisco, California

FOYP Forest Office of Yunnan Province, China

ISNB Institute Royale des Sciences Naturelles de

Belgique, Bruxelles

IZUI Institut für Zoologie der Universität Inns-

bruck, Innsbruck

MCZ Museum of Comparative Zoology, Har-

vard University, Cambridge, Massachusetts

MNKM Muzium Negara Malaysia, Kuala Lumpur

MNHN Museum National d’Histoire Naturelle,

Paris

MSNG Museo Civico di Storia Naturale ‘G.

Doria’, Genova

NHMS Naturhistoriska Riksmuseet, Stockholm

NSMT National Science Museum (Natural His-

tory), Tokyo

NWAS Museum of Entomology, North-western

Agricultural University, Yangling, Shaanxi

OPU Osaka Prefecture University, Osaka

SEMK Snow Entomological Museum, Lawrence,

Kansas

SUU Saitama University, Urawa

TARI Taiwan Agricultural Research Institute,

Taichung

USNM Unites States National Museum, Smithson-

ian Institution, Washington D.C.

UZMK Universitets Zoologiske Museum, Koben-

havn

ZEMK Zoologisch Forschungsinstitut und Muse-

um Alexander Koenig, Bonn

ZMAN Instituut voor Systematiek en Populatie-

biologie (Zoölogisch Museum), Amster-

dam

The following geographical sources have been con-

sulted: ‘Andrees allgemeiner Handatlas’ (Ambrosius

1921), ‘Atlas van Tropisch Nederland’ (Anonymous

1938), ‘The Times Atlas of the World’ (Anonymous

1994), ‘Nelles Road Atlas Southeast Asia excluding

Indonesia’ (Anonymous 1992), ‘Thailand, Indochina

148

& Burma Handbook’ (Eliot et al. 1994), GEonet

Names Server of the U.S. Defence Mapping Agency

(www URL: http://www.nima.mil/gns/html/index.html).

The locations of several collection sites of L. Fea were

determined with Gestro (1904).

Arrows in illustrations point to certain important

features mentioned in the descriptions or other parts

of the text.

PAUP 3.1.1 was used to perform the cladistic analy-

sis to study the relationships between the species of

the P. radha group.

Descriptions were made from numerous specimens

from collections. It should be noted that many species

when alive or freshly collected will be green rather

than ochraceous or brownish.

Measurement were made using a sliding calliper.

Most specimens measured were selected at random or

all available specimens were measured but sometimes

specimens were selected to include extremes of both

ends of the range of variation.

PHYLOGENY OF THE P. radha GROUP

A preliminary phylogenetic analysis was carried out

to investigate the relationships within the P. radha

group. Three outgroups were used for this analysis:

two distant relatives, viz. Meimuna mongolica

(Distant, 1881) and Platylomia tonkiniana (Jacobi,

1905), and one closer relative, viz. Platylomia flavida

(Guérin Méneville, 1834). The characters used are

discussed below and the matrix is given in table 1.

1. — Shape of tergite 3: 0, longer laterally than medi-

ally; 1, virtually equally long laterally and medially.

In the P. radha group and P. flavida tergite 3 is vir-

tually equally long laterally and medially, and the an-

terior margin of tergite 3 is virtually straight. In all

other species of the Dundubiaria, except those of the

Dundubia terpsichore group (see Bloem & Duffels

1976: fig. 1) and Platylomia larus (Walker, 1858)

from Sri Lanka, tergite 3 is considerably longer later-

ally than medially and the anterior margin of tergite 3

is distinctly concave.

2. — Posterolateral and posterior spots on pronotal

collar: 0, present; 1, absent.

The basic pattern in the Dundubiini consists of

several pairs of markings on the pronotal collar, usu-

ally an anterolateral pair, a posterolateral pair and a

posterior pair. Relative positions and sizes can vary

and often all three pairs are present though one may

be less distinct. In a number of groups one or more

spots are reduced or even absent, for example, in most |

species of Dundubia all spots are absent (see also

Overmeer & Duffels 1976, Beuk 1996). The pos- |

terolateral and posterior spots are absent only in some |

Table 1. Character state matrix for the species of the P.

radha group and outgroups used in the cladistic analysis.

3 4 5

M. mongolica 0

P. tonkiniana 0

JP flavida 1

P. bivocalis 1

P. pendleburyi 1

P. malickyi 1

P. radha 1

P. bocki 1

P. ficulnea 1

rie mi SiS

jee eee HE oo

ler se RO

FSG

groups en several not-closely related species (e.g., P.

amicta (Distant, 1889) and relatives, P. radha group

and Platylomia virescens Distant, 1905).

3. — Shape of timbal covering: 0, shorter than broad;

1, as long as or longer than broad.

In the Dundubiini the timbal coverings are usually

shorter than broad at the base. The timbal coverings

are as long as or longer than broad at base in the

species of the P. radha group and in several other

species (e.g., in species of Aola Distant, 1905).

However, the character of this feature is not identical

in the P. radha group and the other species. In the

species of the P. radha group the timbal coverings are

elongated and the timbal organs are almost complete-

ly covered. In the other species the timbal coverings

are narrowed medially and the timbal organs are thus

partly exposed.

4. — Central fasciae pronotal disc; 0, wholly or partly

developed and black; 1, present as dot on posterior

margin of disc or weakly developed and brown.

Similar to the markings on the pronotal collar the

basic pattern in the Dundubiini consists of an elabo-

rate pattern of dark markings on the pronotal disc.

The pattern of markings usually consists of darkening

of the lateral margins, dark fasciae on the oblique fis-

sures and dark central fasciae. In some groups these

markings are strongly reduced so that at most the

central fasciae are distinct but even these can be whol-

ly or partly absent, e.g., in most species of Dundubia

(see also Overmeer & Duffels 1976, Beuk 1996).

In some species of the Dundubiini the transverse

part of the suture between the pronotal disc and

pronotal collar is darkened while the markings on the

disc are otherwise mostly reduced. It is possible that

the dark dot on the posterior margin of the disc is a

remainder of this transverse darkening and not a re-

mainder of the central fasciae. In that case the results

of the analysis carried out here do not change because

then character state 1 would read ‘absent or only

weakly developed and brown.’

Beuk: The Platylomia radha group

5. — Shape uncus lobes: 0, gradually narrowing from

bases onwards (fig. 37); 1, rather abruptly narrowing

near bases (figs. 4, 16).

In the Dundubiaria the shape of the uncus lobes

ranges from two more or less flat lobes to a rather

complex three-dimensional structure. Irrespective of

this shape there usually is a gradual narrowing from

near the basal part of the uncus towards the distal

margin of the uncus lobes even though the lobes may

broaden again distally. In only few species there is a

more abrupt narrowing near the basal part of the un-

cus.

The analysis (exhaustive search) resulted in a single

most parsimonious tree (tree length 5; CI = 1.0, RI =

1.0) which is given in fig. 1. The tree given shows the

two distant relatives in a basal polytomy.

There are some conclusions that can be drawn

from this preliminary analysis. The elongation of the

timbal coverings (character 3 state 1) is a synapomor-

phy for the species of P. radha group. Within the P.

radha group P. radha, P. bocki and P. ficulnea are

grouped together on the basis of the more abrupt nar-

rowing of the uncus lobes near their bases (character

5 state 1). These species are indeed very similar as is

also clear from the descriptions below. Most charac-

ters that can be used to separate these species (mark-

ings on tegmina, shape of opercula) do not have

much phylogenetic significance (see also Beuk 1996).

The clade with P. radha is placed in polytomy with

P. pendleburyi and P. malickyi because they share the

(almost) complete reduction of the central fasciae

M. mongolica

P. tonkiniana

P. flavida

P. bivocalis

P. pendleburyi

P. malickyi

P. radha

P. bocki

P. ficulnea

Fig. 1. Cladogram showing relationships between species of

P. radha group with M. mongolica, P. tonkiniana and P.

flavida as outgroup. Synapomorphies (0??1) are indicated;

see table 1 for the matrix and text for characters.

149

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

(character 4 state 1). Whether this is a good synapo-

morphy should become clear after further analysis of

other groups in the Dundubiaria since this character

state can also be found in most species of Dundubia

(see also Beuk 1996).

Only tentative conclusions can be drawn about the

phylogenetic position of the P. radha group. Within

Platylomia it appears that P. flavida is a close relative

since it shares a synapomorphy with the P. radha

group: the shape of tergite 3 (character 1 state 1). P.

flavida may constitute the sistergroup of the P. radha

group. From the analysis it would appear that the ab-

sence of the posterolateral and posterior markings on

the pronotal collar (character 2 state 1) would also be

a synapomorphy for the clade P. flavida + P. radha

group but as remarked above this state also occurs in

Dundubia and it might indicate a relationship be-

tween this clade and Dundubia.

TAXONOMY

Platylomia Stal, 1870

Cosmopsaltria subg. Platylomia Stal, 1870. Type-species (by

monotypy): Cicada flavida Guérin-Méneville, 1834

Platylomia; Distant (1905) [raised to generic level]

The Platymia radha group

Diagnosis

Large cicadas: d: 40.0-56.5 mm, 9: 38.0-48.0

mm. Body generally ochraceous brown, head and

thorax with some darker markings. Opercula uni-

colorous brownish, only darkened in lateroproximal

corner, male opercula elongate. Dorsal part of ab-

domen usually darker than head and thorax. Timbal

coverings of male elongate, at least as long as broad at

base. Pygofer rather narrow; basal part of uncus short,

narrower than maximum width uncus, and little glo-

bose.

Head. — Postclypeus brown to castaneous, slightly

darker dorsally and usually paler on anteromedial

spot, little swollen. Anteclypeus brownish, some-

times darkened on lateral surfaces. Vertex brownish,

usually darker on vertex lobes and with dark brown

to black markings in area of ocelli and often

along posterior margin. Rostrum brownish, darkened

at tip only; reaching between to distinctly be-

yond hind coxae but never further than halfway ster-

nite 1.

Thorax. — Pronotum slightly to distinctly broader

than head, brown to castaneous but sometimes with

greenish tinge. Pronotal disc with dark brown to

black pattern ranging from small split medial spot at

transverse part of pronotal suture to more elaborate

150

pattern including central fasciae and transverse bands

just posterior of anterior margin and on transverse

part of pronotal suture. Pronotal collar broad, medi-

an length 0.2-0.3 times the median length of prono-

tal disc; posterior margin black; lateral part of collar

with darker brown to blackish markings that may be

connected with transverse band on pronotal suture;

anterolateral corner with lateral tooth. Mesonotum

ochraceous brown to brown, sometimes with green-

ish tinge on paler parts, occasionally posterior half

partly castaneous; disc with pattern of narrow fasciae

consisting at least of paramedian fasciae, lateral fasci-

ae usually absent. Cruciform elevation concolorous

with mesonotal disc or slightly paler. Area enclosed

by arms of cruciform elevation and parts of mesono-

tal disc occasionally with waxy coating. Katepimeral

lobe usually longer than broad at base.

Tegmina and wings. — Tegmina hyaline to brown-

ish hyaline, always with indistinct brownish reticula-

tion along distal margin and sometimes in apical cells;

tegmina either without markings, or with markings

on second, third, fifth, and seventh apical cells and at

apices of longitudinal veins of apical cells, or interme-

diate; basal cell partly fumose. Wings hyaline to pale

brownish hyaline, infuscate along very narrow strip

posterior of first anal vein.

Legs. — All legs ochraceous to brownish with vary-

ing dark pattern on fore legs, mid tibiae and tarsi, and

hind tibiae and tarsi. Fore femur posteroventrally

with middle spine shorter than proximal spine; prox-

imal spine slender, middle spine more pointed than

proximal spine, often broadened at base and thus

more triangular; distal spine short but distinct, either

pointed or blunt, sometimes also broadened at base.

Hind tibiae with 3-5 anterodorsal spines and 2-6 an-

teroventral spines, occasionally with single anterior

spine.

Male operculum. — Operculum rather variable in

length and shape, reaching from anterior margin of

fifth to just beyond posterior margin of seventh ab-

dominal segment; ochraceous to brown but some-

times with greenish tinge and sometimes partly cov-

ered with waxy coating, some parts near latero-

proximal corner dark brown to black. Medial margin

weakly convex for short distance proximal of con-

striction, distal of constriction convex for at least

some distance. Apex lateral of midline, rounded or

angularly rounded to gully-shaped. Lateral margin

convex for short distance proximal of constriction,

concave at level of timbal covering, distal of constric-

tion convex but often straight or sometimes even con-

cave on distal third.

Male abdomen. — Abdomen 1.2-1.5 times as long

as head and thorax together. Dorsal part of tergites

brownish to dark castaneous but usually paler lateral-

ly, tergites 3-7 either with or without darker spots

near lateral margin; sternites and ventral parts of ter-

gites paler than dorsal parts but sometimes less so on

segments 6-7. Tergites usually with a pattern of waxy

coating and/or whitish dusting. Posterior margins of

tergites (5) 6-7 with spinules. Sternite 7 with very

shallow to rather deep posteromedial emargination.

Timbal covering brownish to castaneous but some-

times with greenish tinge, about 1.0-1.3 times as long

as wide.

Male genitalia. — Yellowish brown to dark brown.

Basal pygofer lobes narrow and ridge-like to broad

and rounded. Basal part of uncus little globose, short,

and rather narrow. Uncus lobes short and gradually

narrowing distally or first narrowing and then some-

what broadening distally; distal margin ranging from

smooth and sinuous to adorned with lobule or trian-

gular projections; laterodistal corner produced to

varying extend; lateral margin distally sometimes

adorned with dorsal (inner) flange or with one or

more small triangular projections.

Female operculum. — Operculum greenish to

brownish and often darkened lateroproximally;

reaching little beyond anterior margin of segment 3,

only little curved around abdomen laterally.

Female abdomen. — Abdomen 0.9-1.1 times as

long as head and thorax together, castaneous brown

Beuk: The Platylomia radha group

to largely blackish dorsally but paler laterally and ven-

trally, tergites 3-6 (7) either with or without darker

spots near lateral margin. Tergites usually with pat-

tern of waxy coating and/or whitish dusting. Poster-

ior margins of tergites (3) 4-8 with short dark spines

but only laterally on tergites (3) 4-5, on tergite 6 ei-

ther only laterally or along whole length, on tergite 7

along whole length and on tergite 8 only medially.

Sternite 7 with angular to angularly rounded postero-

medial emargination, sometimes in median lobe.

Female genitalia. — Pygofer brownish to dark casta-

neous or blackish; distal part of ovipositor sheath dark

castaneous to blackish. Dorsal margin of pygofer in

lateral view weakly concave and with slight bump at

base of caudodorsal beak; ventral margin somewhat

stronger convex. Ovipositor sheath short, at most

reaching little beyond apex of caudodorsal beak.

Key to the males of the Platylomia radha group

1. Tegmina with distinct markings on basal veins of

second, third, fifth, and seventh apical cells, and

at apices of longitudinal veins of apical cells

(when in doubt both alternatives can be fol-

— Tegmina with indistinct markings only on basal

| (6)

85 90 95 100 105 110 115 120 sE

204 |

IR +

5h DI

+ N

107 dt

da N |

5 QE ax |

+ à + 15

7 AN :

TS af din al “à A

rx) |

8 QW’ à Ch Uro

Je IN De]

x SE a wen an Be =

75 80 85 90 95 100 105 110 115

Fig. 2. Localities of 2. radha.

151

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

veins of second and third apical cells or without

markıneser et arte Ra 5

Tegmina distinctly brownish infuscate along

veins. Tergites 5 and 6 completely covered with

whitish dust obscuring ground-colour of tergites.

hen ah ede CEE sente din. Sellia P. bivocalis

Tegmina at most slightly infuscate along veins.

Tergites 5 and 6 at least partly without whitish

dust nl ee 3

Opercula attenuate at apex and long, reaching to

anterior margin of sixth abdominal segment or

further (HE 27) ee P. ficulnea

Opercula rounded at apex and short, at most

reaching to anterior margin of sixth abdominal

Seprment (tess 20345749) Re, 4

Distal part of opercula diverging from abdomen,

so well visible in dorsal view. Uncus lobes gradu-

ally narrowing towards distal margin (fig. 45).

Larger species: body length more than 50 mm....

Sr ee ee P. pendleburyi

Distal part of opercula close to abdomen, if diver-

gent than not broadly rounded at apex. Uncus

lobes distally slightly broader than at about

halfway from bases (figs. 16, 30). Smaller species:

bodyjlenethylessithant> Om nnn 6

Apices of opercula variable in shape, distal third

of opercula gradually narrowing or attenuate

(figs. 11-13); operculum usually reaching further

than halfway sixth abdominal segment, if shorter

than narrowed towards apex and medial concave

casal P. radha

Apices of opercula rounded and at most little nar-

rowed, opercula reaching no further than anteri-

or margin of sixth abdominal segment (figs. 20,

34) dora AMER ERN La Lino CAE 6

Transverse part of pronotal suture with narrow

black band may not be connected to markings on

lateral part of pronotal collar. Distal part of oper-

culum broad, narrowing little towards apex (fig.

20). Distal margins of uncus lobes convex near

mediodistal corner and concave near laterodistal

corner; laterodistal corner angular to slightly

pointed (figs. 16, 19). Margins of uncus lobes at

most with one or two small wart-like outgrowths

ORE ENE cet Net en P. bocki

Transverse part of pronotal suture with broad .

black band that is connected to markings on lat-

eral part of pronotal collar. Distal part of opercu-

lum narrowing somewhat towards apex (fig. 34).

Distal margin of uncus lobes with medial lobule

that carries two small distal projections at its dis-

tal margin (figs. 30, 33). Lateral margin near lat-

erodistal corner with several short, pointed pro-

Jéctionsi(figs 9021053) PERSA P. malickyi

Platylomia radha (Distant, 1881)

(figs. 2, 4-15)

Dundubia radha Distant, 1881: 634. — Lectotype é (here

designated) of Dundubia radha Distant: ‘Type [printed

on round label with red margin], ‘Masuri / Hills’ [hand-

written], ‘radha / Dist.’ [Distant’s handwriting], ‘Distant

Coll. / 1911-383.’ [printed].

Dundubia radha. — Distant 1882: 160; Atkinson 1884: 225;

Atkinson 1886: 163; Distant 1888a: 292; Distant 1888b:

455.

Dundubia similis Distant, 1888a: 292. — Lectotype d (here

designated) of Dundubia similis Distant: ‘Type’ [printed

on round label with red margin], ‘Sikkim’ [handwritten],

‘similis / Dist.’ [Distant’s handwriting], ‘Distant Coll. /

1911-383.’ [printed]. Syn. n.

Cosmopsaltria similis. — Distant 1889: 45; Distant 1890: pl.

V figs. 10-10b; Distant 1892b: xii.

Cosmopsaltria radha. — Distant 1889: 46, pl. IV figs. 9-9b;

Distant 1890: 54; Distant 1892b: xii; Noualhier 1896:

254; Distant 1897: 17; Noualhier & Martin 1904: 179.

Platylomia similis. — Distant 1906a: 102; Distant 1906b: 61;

Distant 1912: 49; Paiva 1919: 372; Moulton 1923: 98

[partim, unspotted tegmina], 102-103 [partim; Sikkim,

Assam], 167 [partim; Sikkim, Assam]; Kato 1932: 166;

Metcalf 1963: 625.

Platylomia radha. — Distant 1906a: 105, fig. 31; Distant

1906b: 61; Distant 1912: 49; Distant 1913: 560; Distant

1917: 101; Kato 1932: 166; Mathur 1953: 140; Metcalf

1963: 623; Hayashi 1978: 177, figs. 20-21; Duffels &

Van der Laan 1985: 122; Lei & Li 1994: 92, 94.

Platylomia operculata Distant, 1913: 559. — Lectotype d

(here designated) of Platylomia operculata Distant: "Type

/ H.T. [printed in round label with red margin], ‘Indo-

China. / (R. Vitalis.) [printed], ‘Platylomia / operculata /

type Dist.’ [Distant’s handwriting], “1913-222.” [print-

ed]. Syn. n.

Platylomia operculata. — Distant 1917: 101; Metcalf 1963:

623.

Not: Moulton 1923: 98 [partim; faint infuscations on

tegmina], 100, 102-103 [partim; Thailand], 167 [partim;

Thailand] (undescribed species of Platylomia).

P. radha is the most widespread species of the P.

radha group and displays the largest amount of varia-

tion in body size, shape of the opercula, and shape of

the distal part of the uncus lobes. Specimens with at-

tenuate opercula resemble P. feae but can readily be

recognised by the absence of distinct markings on the

tegmina. Specimens with more rounded opercula

mostly resemble P. bocki.

P. radha appears to be very closely related to P.

bocki. The main differences can be found in the oper-

cula and the male genitalia. The opercula of P. radha

are short or long but always narrowed near the apex

or even attenuate, whereas those of P. bocki are short

and broad almost to the apex. The male genitalia dif-

fer in the shape of the pygofer and the uncus. The

basal pygofer lobes are smaller in P. radha and reach

not as far posteriorly between the lateral margins of

the pygofer as in P. bocki (compare figs. 4 and 16).

The ventral (outer) surface of the uncus lobes has a

Beuk: The Platylomia radha group

a Fig. 3.

Bee Localities of P. bivocalis, P.

| °N : : x

| bocki, P. ficulnea, P. malickyi

1 30 and P. pendleburyi.

125

(

+ 120

+15

110

bivocalis

bocki 5

ficulnea

malickyi

pendleburyi

90 95 100 105

ridge running roughly from the mediodistal corners

to the medial part of the basal part of the uncus. In P.

radha this ridge is much lower and smoother than in

P. bocki. The uncus lobes of P. radha have a distinct

flange of variable shape on the lateral margin, where-

as in P. bocki the lateral margins are smooth or (occa-

sionally) provided with one or two small wart-like

outgrowths. P. bocki is generally smaller than P.

radha, the largest specimens of P. bocki being about

the same size as the smallest specimens of P. radha.

Description

Body brownish to castaneous, often with some

parts of head and thorax lighter, mesonotum with

median and paramedian fasciae partly developed,

tegmina without markings but basal veins of second

and third apical cells occasionally almost impercepti-

bly infuscated. Opercula very variable in shape, rang-

ing from relatively short and almost rounded at apex

to long and almost gully-shaped at apex.

Head. — Postclypeus brown to light castaneous,

sometimes paler on area anterior of frontoclypeal su-

ture and ventrally along lateral margin and at clypeal

suture; little swollen, in dorsal view about as long as

distance between frontoclypeal suture and anterior

margin of pronotum. Anteclypeus brownish. Vertex

brownish but somewhat darker on lateral part and on

supra-antennal plates; posterior margin with pair of

black spots close to eyes; part posterior of eyes with

black marking, either rounded or extending to and

along posterior margin of eye; area of ocelli with dark-

er brown to black marking that encloses anterior ocel-

lus and reaches laterally as far as lateral ocelli. Genae

ochraceous, lori darkened along suture with clypeus.

Frontoclypeal suture semicircular but median part al-

most straight, much broader than distance between

lateral margins of lateral ocelli. Rostrum brownish,

darkened at tip only; reaching halfway between hind

coxae to posterior margin of hind coxae.

Thorax. — Pronotum slightly to distinctly broader

than head. Pronotal disc brownish, anterior margin

and often medial area paler; disc immediately posteri-

or of anterior margin with dark brown to black trans-

verse band that is usually broadly interrupted medial-

ly; central fasciae usually not developed except for

two brownish triangular markings with their base at

pronotal suture; transverse part of pronotal suture

with dark brown to black band that is broadest medi-

ally and continues laterally to level of markings on lat-

eral part of pronotal collar. Pronotal collar usually

paler than pronotal disc and with black posterior

margin that broadens slightly in posterolateral cor-

ners; anterolateral part of collar with darker marking

between lateral margin and pronotal suture, marking

darkest at suture and often connected with band on

transverse part of pronotal suture; anterolateral cor-

153

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

9 10

Figs. 4-10. Platylomia radha (Distant, 1881), male. — 4, pygofer and uncus in ventral view, Jingkawksakan; 5, pygofer and un-

cus in right lateroventral view, Jingkawksakan; 6, pygofer in dorsal view, Doi Inthanon; 7, uncus in right lateroventral view,

Doi Inthanon; 8, uncus in right lateroventral view, Tonkin; 9, uncus in right lateroventral view, Ceylan; 10, uncus in right

lateroventral view, lectotype P. radha.

154

ner with small but distinct lateral tooth; posterolater-

al corner rounded but sometimes part of margin only

weakly convex. Mesonotum ochraceous brown to

brown, sometimes even castaneous; median fascia

variable, from almost indistinguishable to blackish;

paramedian fasciae on anterior half of disc black, nar-

row near anterior margin of disc then broadening and

slightly curved medially, on posterior half usually pre-

sent as distinct black or indistinct brownish markings

of varying size in front of cruciform elevation; anteri-

or margin of disc with dark brown to blackish trian-

gular markings immediately lateral of mesonotal fis-

sures; lateral fasciae absent. Cruciform elevation

ochraceous brown to dark brown, concolorous with

or paler than mesonotal disc. Katepimeral lobe (fig.

15) shorter than broad at base, apex rounded. Surface

of katepimeral lobe concave along dorsal margin, dor-

sal margin thus curved outwards; surface covered

with short, waxy hairs. Apex of katepimeral lobe

reaching over base of operculum but sometimes only

just.

Tegmina and wings. — Tegmina almost entirely

hyaline with indistinct brownish reticulation along

BEUK: The Platylomia radha group

Figs. 11-15.

Platylomia radha (Distant,

1881), male. — 11, ab-

domen and right opercu-

lum in lateroventral view,

lectotype P. similis 12,

right operculum in lat-

eroventral view, lectotype

P. operculata, 13, right op-

erculum in lateroventral

view, lectotype P. radha;

14, right timbal covering,

lectotype P. radha; 15, right

katepimeral lobe,

Jingkawksakan

distal margin and sometimes in apical cells; basal

veins of second and third apical cells rarely with al-

most indistinguishable infuscation; basal cell yellow-

ish fumose on anterior half. Veins of tegmen brown-

ish, darker on most cross veins and parts where veins

meet or split. Wings hyaline. Veins of wings brown-

ish, medial vein and veins close to wing margin dark-

er.

Legs. — Legs ochraceous to brownish. Fore femora

ochraceous, posteroventrally with darkened longitu-

dinal stripe; tibiae slightly darker than femora except

on dorsal surface; tarsi brownish. Mid femora ochra-

ceous; tibiae brownish, slightly darker at base and on

apical third, especially ventrally; tarsi brown. Hind

femora ochraceous, slightly darkened anteriorly; tibi-

ae often little darker than femora, dark brown at

joints with femora; tarsi brownish. Fore femur with

gap between middle and distal posteroventral spines

shallow and broad. Hind tibia with two dorsal spines

and usually four but occasionally only three an-

teroventral spines; spines brownish.

Male operculum (figs. 11-13). — Operculum rather

variable in length and shape, reaching about halfway

155

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

fifth abdominal segment to just beyond posterior

margin of seventh abdominal segment, 2.2-3.0 times

as long as maximum width distal of constriction;

ochraceous to brown and frequently some parts cov-

ered with waxy coating, area of lateroproximal corner

and usually also part of lateral margin dark brown to

black but with ochraceous spot on margin just distal

of lateroproximal corner. Medial margin distal of

constriction convex to apex but near apex often less

convex or almost straight, margin on distal third fre-

quently appearing concave because surface becomes

weakly concave along margin. Apex of operculum lat-

eral of midline and variable in shape: either apex

rounded to angularly rounded with operculum grad-

ually narrowing distally (fig. 11), or angular with sur-

face concave along one margin only (usually lateral

margin), or gully-shaped with surface concave along

both medial and lateral margin (fig. 13) and then of-

ten divergent from abdomen. Lateral margin distal of

constriction convex but often less convex or almost

straight near apex, margin on distal third frequently

appearing concave because surface becomes weakly

concave along margin. Constriction at about 0.3 of

length of operculum, lateral concavity longer and

deeper than medial concavity and at deepest point of-

ten almost angular, broadest part of operculum distal

of constriction 1.3-1.5 times as wide as minimum

width in constriction. Distance between opercula at

constrictions about 0.8-1.1 times as wide as mini-

mum width in constriction. Opercula at point of

closest approximation separated for a distance of 0.5-

0.7 times maximum width between opercula at con-

strictions. Operculum close to abdomen but distal

half well removed from abdomen when apex gully-

shaped; surface only weakly convex in longitudinal

direction or concave along margins on distal third,

surface more strongly convex in transverse direction,

especially on distal third.

Male abdomen. — Abdomen about 1.3-1.5 times as

long as head and thorax together. Dorsal part of ter-

gites brownish to castaneous, often somewhat dark-

ened along posterior margins; sternites and ventral

parts of tergites paler than dorsal parts but less so to-

wards genitalia. Tergite 1 laterally and tergite 2 para-

medially usually with white waxy coating, tergite 3

with dense whitish dust closer to lateral margin, ter-

gite 4 usually with smaller area of whitish dusting at

posterior margin, some dusting present on tergites 5-

7 but without distinct pattern, tergite 8 completely

covered with whitish dust. Posterior margin of tergite

7 with many short dark spinules, of tergite 6 and oc-

casionally also of tergite 5 with few spinules on later-

al part. Sternite 7 with shallow to very shallow pos-

teromedial emargination. Timbal covering (fig. 14)

brownish to castaneous, about 1.0-1.1 times as long

as wide; medial margin virtually straight, convergent

156

with lateral margin; mediodistal corner, distal margin

and laterodistal corner rounded; lateral margin al-

most straight.

Male genitalia (figs. 4-10). — Brown; dorsal part of

pygofer somewhat darker than ventral part; uncus

lobes usually darker along lateral and distal margin.

Basal pygofer lobes broad and rounded (figs. 4-5);

hairs on anteroventral margin of pygofer erect, not

very long and continuing on pygofer surface enclosed

by anteroventral margin, hairs shorter on apices of

basal lobes, towards medial part of anteroventral mar-

gin and on surface enclosed by anteroventral margin;

pygofer surface anterior of medial part of anteroven-

tral margin and lateral of anteroventral margin with

scattered hairs, dorsolateral surface covered with

many short hairs. Dorsal part of pygofer as in fig. 6.

Basal part of uncus small, little globose and narrower

than maximum width uncus (figs. 4-5); near bases of

uncus lobes usually with some thick erect hairs and

several thinner hairs more medially, occasionally only

very few hairs present. Uncus lobes (figs. 4-5, 7-10)

short; medial margin weakly concave and curved dor-

sad (inwards) along whole length; mediodistal corner

rounded and also slightly curved dorsad (inwards);

distal margin weakly concave; laterodistal corner

(figs. 7-10) produced and pointed to varying extend,

tip directed anterodorsad (inwards) to laterad; lateral

margin strongly curved medially close to base, distal

part of uncus lobe then gradually broadening towards

distal margin or narrow for some distance and broad-

ening more abruptly near distal margin; lateral mar-

gin near laterodistal corner with flange of irregular

shape. Ventral (outer) surface of uncus lobes with

longitudinal grooves near anterior margin and each

uncus lobe with indistinct ridge from mediodistal

corner almost to basal part of uncus where ridges

nearly meet. Bases of uncus lobes laterally usually

with numerous strong erect hairs both on dorsal (in-

ner) and ventral (outer) surfaces, those on dorsal sur-

face shorter except for occasional very long hairs.

Female operculum. — Operculum brownish, dark-

er basally and along medial margin, lateral margin

castaneous. Lateral margin distal of lateroproximal

lobe weakly convex to weakly concave; laterodistal

corner angularly rounded; distal margin weakly con-

vex but often concave for very short distance near lat-

erodistal corner; mediodistal corner broadly rounded;

medial margin short and convex; lateral and medial

margin of each operculum virtually parallel.

Female abdomen. — Abdomen about as long as

head and thorax together. Dorsal part of tergites cas-

taneous brown but tergites 1-3 little paler, posterior

margins of tergites darkened, tergites without darker

spots near lateral margins; sternites and ventral parts

of tergites paler than dorsal parts. Tergite 2 and ante-

rior margin of tergite 3 with white dusting laterally |

but dusting often rubbed off, dusting on other ter-

gites without distinct pattern. Posterior margins of

tergites 4-5 with some short dark spines laterally, of

tergite 6 with spines along whole length except medi-

ally, of tergite 7 with spines along whole length, of

tergite 8 with scattered spines medially. Sternite 7

with angular to angularly rounded posteromedial

emargination.

Female genitalia. — Dorsal part of pygofer almost

castaneous, remainder brownish; distal part of

ovipositor sheath castaneous. Dorsal margin of py-

gofer shorter than three preceding segments, in later-

al view weakly concave; ventral margin weakly con-

vex. Ovipositor sheath reaching almost as far as or

short distance beyond apex of caudodorsal beak; anal

valve not reaching as far as caudodorsal beak.

Measurements in mm (2: n = 6). — Body length:

6: 44.5-56.5 (51.5 + 3.1) (n= 12), 2: 38.5-44.5

(40.9 + 1.4); head width: d : 12.6-15.3 (14.2 + 0.7)

(NM) 2:7133-15.07(14:07=7 0:6); maximum

pronotum width: d: 12.8-16.8 (15.3 + 0.8) (n =

11), 2: 14.7-16.3 (15.2 + 0.5); tegmen length: ó:

44.5-62.5 (56.6 + 3.0) (n = 12), 2: 53.3-61.5 (57.0

BED):

Variation

P. radha shows considerable variation in several

characters but there seems to be hardly any geograph-

ic pattern. The characters showing variation are body

colour, body length, length of operculum, shape of

apex of operculum, and shape of laterodistal corner of

uncus lobe. The general pattern is that smaller speci-

mens with shorter opercula and opercula with more

rounded apices (fig. 11) tend to come from the north-

western part of the range of P. radha. However, larg-

er specimens with longer, attenuate opercula (as in

fig. 13) have also been found in that part of the range

(see Hayashi 1978: fig. 20). Smaller specimens are

also found in the central part of the range and spec-

imens with more angularly rounded opercula (com-

pare fig. 12) are found in the eastern part of the range.

Variations in shape of laterodistal corner of uncus

lobe (figs. 7-10) and variations in colour show no ge-

ographic pattern at all. Differences in colour are

probably due to different circumstances in killing and

preserving specimens.

Distribution (fig. 2)

P. radha has a wide distribution and is recorded

from southern parts of the Himalaya (Bhutan; India:

Assam, Sikkim; Nepal) through Indo-China (Burma,

Cambodia, Laos, Thailand, Vietnam) to southern

China in the east (Hainan, Sichuan, Yunnan). A

number of records in the literature are doubtful. The

record from Madras (Distant 1881, 1890, 1906a; At-

kinson 1884) could not be confirmed and may have

BEUK: The Platylomia radha group

been caused by wrong location of the type locality in

the Madras Presidency. Since the nearest locality is

more than 1,500 km away the accuracy of this old

record is questioned. Moulton’s (1923) record of P.

similis from Khao Luang, just south of the Kra

Isthmus on the Malay Peninsula refers to a yet unde-

scribed species of Platylomia.

Synonymy

Examination of the lectotypes of P. radha and P.

similis has shown that they differ in several aspects.

The lectotype of P. similis is smaller and paler than

the lectotype of P. radha, its opercula are shorter and

not attenuate, its timbal covering is more triangular,

and the shape of the distal margin and laterodistal

corner of the uncus lobes is different. The lectotypes

of P. radha and P. operculata are more resemblant.

The lectotype of P. operculata is a little larger and

darker than lectotype of P. radha, its opercula are a

little longer and the apices of the opercula are nar-

rowed and not attenuate. The lectotype of P. opercu-

lata is damaged by museum beetle and its genitalia are

partly destroyed. The remains of the genitalia are

identical to those of the lectotype of P. radha.

Examination of further collection material under

P. radha, P. similis and P. operculata has shown that

several characters show variation. The characters giv-

en by Distant (1888a, 1913) to distinguish P. similis

and P. operculata from P. radha (colour, shape of op-

erculum) are two of these characters (see discussion of

variation above). The three species in fact represent

different forms of the same species. Since there are no

characters that seem to justify a separation of P. sim-

ilisand P. operculata from P. radha, P. similis and P.

operculata are herewith synonymised with P. radha.

Remarks

In the descriptions of Dundubia radha, D. similis

and P. operculata neither holotypes were designated

nor were the numbers of specimens upon which the

descriptions were based mentioned. No lectotype

designations were published subsequently. For this

reason the presence of labels on specimens indicating

type status cannot be considered valid type designa-

tions. For each of the three species it applies that spec-

imens belonging to the type series should be consid-

ered syntypes and lectotypes have to be designated

(ICZN Recommendation 73F).

The type material of P. radha was stated to origi-

nate from ‘Madras Presidency; Masuri [Mussoorie]

Hills’ in India (Distant 1881: 684) but it is not clear

whether Distant referred to two localities or to one lo-

cality (wrongly locating the Mussoorie Hills in the

Madras Presidency). Since later publications (Distant

1889; 1906a) only refer to a specimen from the

Madras Presidency in Distant’s collection, the latter

157

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

may have been the case (see also under Distribution).

No specimen from the Madras Presidency could be

found so the specimen from Mussoorie is designated

as the lectotype.

P. similis was described from ‘Sikkim’ on the basis

of material in Distant’s collection. There are several

males from Sikkim at the BMNH but only one male is

from Distant’s collection and corresponds with the

size mentioned for P. similis in the original descrip-

tion. This male is designated as the lectotype.

P. operculata was described from ‘Indo-China’.

Only one male in the BMNH that may have belonged

to the type series is available. This specimen is desig-

nated as the lectotype.

Material examined. — cHINA: Cangyuan [Mengdong],

Yunnan, 1278 m, 12.v.1980, 2 dd (nwas); Cheli

[Yunjinghong], Yunnan, 620 m, 4.v.1957, 1 d (1zas);

Jianfeng Ling, Hainan, 13.iv.1982, Gu Maobing, 2 d, 1 9

(1zas); Jingdong [Jinping], Yunnan, 1170 m, 21.v.1956, 2

3 (1zas); Jinggu [Weiyuan], Yunnan, 1000 m, 16.v.1956,

1 d, 1 @ (izas); Jinghong [Yunjinghong], Yunnan,

3.iv.1991, 1 2 (Nwas); Longlin, Guangxi, 14.vi.1981, 1 d

(Nwas); Mangshi, Yunnan, 900 m, 15.v.1955, 4 d (1zas);

Menghai [Xiangshan], Yunnan, 27.iv.1980, 1 d (Nwas);

Mengla, Yunnan, 9.v.1991, Wang Yenglun, 1 d (Nwas);

Menglun, Yiwubanna, Yunnan, 650 m, 1.v.1964, 2 d

(1zas); Mengyang, Yunnan, 1800 m, 5.vi.1991, 1 d

(Nwas); Pu’er [Ning’er], Yunnan, 13.v.1956, 1 d (1zas);

Ruili [Mengmao], Yunnan, 1350 m, 12.vi.1956, 1 2 (1zas);

Shuangjiang [Mengmeng], Yunnan, 888 m, 31.v.1980, 1 d

(nwas); Simao, Yunnan, 1200 m, 11.v.1957, 1 d (1zas);

Taiping Shan, Hainan, 5.v.1986, Y. Okuma, 1 d (suv);

Xiaomengyang, Yunnan, 850 m, 4.v.1957, 3 d (1zas);

Yaoqu, Yunnan, 1600 m, 4.v.1991, 1 d (Nwas); Yingjiang

[Pingyuan], Yunnan, 22.v.1983, 1 d (Nwas); Yuanjiang

[Lijiang], Yunnan, 550 m, 3.v.1980, 1 d (Nwas); Yunnan,

7.v.1957, 1 & (suu); Zhenyuan, Yunnan, 4.v.1956, 1 d

(IZAS); VIETNAM: Chapa [Cha Pai], Tonkin, Indo-China, v,

R.V. de Salvaza, 1 2 (BMNH); 6 km S. of Dalat [Da Lat],

1400-1500 m, 9.vi-7.vii.1961, N.R. Spencer, 1 5, 3 9

(BPBM); Mt Tam Dao, N. Vietnam, 1300 m, 10.vii-

2.viii.1990, S. Nagai, 1 d, 2 2 (suu); Tam Dao, N.

Vietnam, 27.vii-2.vili.1992, N. Ohbayashi, 1 2 (suu); Tam

Dao, 100 km N.W. from Hanoi, N. Vietnam, 16-

27.v.1991, M. Takakuwa, 1 & (suv); Tonkin, 1 d (MNHN);

Tonkin, vi.1917, R.V. de Salvaza, 2 6, 1 2 (BMNH); CAM-

BODIA: Cambodge, 1886, A. Pavie, 1 d (MNHN); LAOS:

Luang Prabang [Louangphrabang], vi.1916, R.V. de

Salvaza, 1 d (BMNH); Muong You [Muang You], Luang

Prabang [Louangphrabang], 25.v.1919, R.V. de Salvaza, 1

9 (BMNH); Pakkading, Borikhana Province, 22.iii.1965, na-

tive collector, 1 d (BPBM); Paklay [Muang Pak-Lay],

viil.1917, R.V. de Salvaza, 1 d (BMNH); Phon Saly

[Phongsalil, N. Laos, 2.v.1994, Y. Miyake, 2 d (suv);

Xieng Khouang [Xiangkhoang], 2.iv.1919, R.V. de Salvaza,

1 6 (BMNH); same data, 13.iv.1919, 1 d (BMNH); same

data, 15.iv.1919, 7 d (BMNH); same data, 17.iv.1919, 1 6,

1 2 (BMNH); same data, 22.iv.1919, 1 2 (BMNH); same

data, 5.v.1919, 1 9 (BMNH); THAILAND: Bang Khun Klang,

Doi Inthanon, 18°32’N 98°32’E, 1200 m, 10-17.iv.1989,

Chantaramongkol & Malicky, light trap, 1 d (tzur); same

158

data, 17-24.iv.1989, 1 8, 1 2 (izui); same data, 27.iv-

1.v.1989, 6 d (zut); Bangkok à Xieng Mai [Chiang Mail,

1887, Pavie, 1 d (MNHN); Changmai [Chiang Mai] Zoo,

18°49N 98°57E, 400 m, 24.iv-1.v.1989, Chantara-

mongkol & Malicky, light trap, 1 d (zur); Chiang Dao,

Chiang Mai, 1 d (suu); Doi Chang, N. Siam, 4500 ft,

14.iv.1921, E.J. Godfrey, 5 d (BMNH); Doi Chiang Khian,

Chiang Mai, ca. 1250 m, 29.v.1983, H. Kuroko, S.

Moriuti, Y. Arita & Y. Yoshiyasu, 1 2 (opu); Doi Inthanon,

1750 m, 25.vi.1987, M.G. Allen, 1 d (BMNH); Doi

Inthanon, 2000 m, 13.iv.1988, T.W. Harman, 2 d (zman);

Doi Nya Chang, Khun Thal, N. Siam, 15.iv.1941, A.M.

Hemmingsen, 1 d (uzmk); Doi Suthep, Chiang Mai,

1.v.1984, T. Endo, light trap, 1 d (suv); Doi Suthep, N.

Thailand, 10.iv.1983, T. Shiromura, 1 d (NsMT); Fang,

Chiang Mai, ca. 450 m, 15.v.1983, Kuroko, Moriuti, Arita

& Yoshiyasu, 1 d (opu); Fang, Chiengmai [Chiang Mai]

Province, 14.iv.1958, T.C. Maa, 1 d (BPBM); Fang,

Chiangmai [Chiang Mai], 500 m, 12-19.iv.1958, T.C.

Maa, 2 d (BPBM); same data, at light, 12.iv.1958, 1

d (BPBM); same data, at light, 15.iv.1958, 2 4 (BPBM); same

data, at light, 19.iv.1958, 3 d (BPBM); Mount Angka [Doi

Inthanon], Asiatic Primate Expedition, 1 é (mcz); Phu

Khieo, Chaiyapumh [Chaiyaphum] District, 800 m, 2-

4.v.1986, M.G. Allen, 3 d (BMNH); Wiang Pa Pao, Chiang

Mai, N. Thailand, 30.11.1990, 1 &, 1 £ (suu); BURMA:

Ghecu, Carin, 1300-1400 m, iv.1888, L. Fea, 2 d (MsNG);

Jingkawksakan [Tingkawk Sakan], 17.v.1944, L.C. Kuitert,

2 d (semk); Shingbwiyang, 1.v.1944, L.C. Kuitert, 2 d

(SEMK); same data, 13.v.1944, 1 d (sEMK); same data, J.D.

Hitchcock, 1 d (seMK); North Chin Hills, Watoon, 1 d

(BMNH); southern Shan States, Bingham, 1 d (BMNH);

Teinzo, v.1886, L. Fea, 2 d (msn); Tingkawk [Tingkawk

Sakan], 25.v.1944, L.C. Kuitert, 3 d, 1 2 (SEMK); same

data, 29.v.1944, 1 2 (seMK); Upper Burma, 1 d (BMNH);

INDIA: Assam, 1 2 (BMNH); Masuri [Mussoorie] Hills, &

lectotype Dundubia radha Distant (BMNH); Naga Hills,

Doherty, 1 4 (BMNH); Sikkim, d lectotype Dundubia sim-

ilis Distant (BMNH); Sikkim, 3 4, 6 © (BMNH); Sikkim, 1 &

(NHMs); Sikkim, 1 2 (1sN8); Tumlong, Sikkim, Gallasch, 1

3 (ZFMK); BHUTAN: Bhoutan Anglais, 1900, R. Oberthur, 1

® (mnun); Bhoutan, Maria Basti, 1898, Durel, R.

Oberthur, 1 d (MNHN); EXACT LOCATION UNKNOWN: no lo-

cality given, 4 d, 1 ® (BMNH); no locality given, 2 d

(MSNG); no locality given, 1 4 (usnm); Ceylan [Sri Lanka?],

Oberthur, 101-96, 1 & (MNHN); Indo-China, R. Vitalis, 1

d lectotype Platylomia operculata Distant (BMNH).

Females probably belonging here. — Laos: Vien Poukha

[Viangphoukha], Haut Mékong, 3.v.1918, R.V. de Salvaza,

2 2 (BMNH); Thailand: Doi Pui, Chiang Mai, ca. 1300 m,

30.v.1983, Kuroko, Moriuti, Arita & Yoshiyasu, 1 9 (opu);

Phuping Palace [Phuphing Palace], Chang Mai, 30.iv.1984,

T. Endo, 1 2 (suv).

Platylomia bocki (Distant, 1882)

(figs. 3, 16-22)

Dundubia bocki Distant, 1882: 159. — Lectotype d (here

designated) of Dundubia bocki Distant: “Type / H.T.

[printed in round label with red margin], ‘Indo-China. /

(R. Vitalis.) [printed], ‘Platylomia / operculata / type

Dist.’ [Distant’s handwriting], 1913-222. [printed].

Cosmopsaltria bocki. — Distant 1889: 45, pl. IV figs. 11-11b;

Distant 1892b: xii.

Platylomia bocki. — Distant 1906b: 60; Distant 1912: 48;

Moulton 1923: 98; Kato 1932: 166; Metcalf 1963: 615.

Although P. bocki was described in a paper on

Sumatran cicadas and the type locality is stated to be

Sumatra (Distant 1882), Moulton (1923) already

pointed out that this specimen could have come from

either Sumatra or Siam (Thailand). Considering that

the other specimens of P. bocki are all from continen-

tal Southeast Asia and the fact that no other species of

the P. radha group was ever found on Sumatra it

seems most likely that the specimen was collected in

Thailand. P. bocki is a typical species of the P. radha

group and very closely related to P. radha. The main

differences are discussed under P. radha.

Description

Body brownish with some parts on head and tho-

rax lighter, mesonotum with median and paramedian

fasciae partly developed, the tegmina without mark-

ings except for the often infuscated basal veins of sec-

ond and third apical cells. Opercula relatively short

and broad at apex, apical margin either broadly

rounded or even partly straight.

Head. — Very much resembling P. radha. Vertex as

in P. radha but marking on area of ocelli always dis-

tinctly black; part posterior of eyes without black

marking.

Thorax. — Pronotum as in P. radha, but general

pattern of paler and darker brown parts more distinct;

central fasciae usually not developed. Lateral part of

pronotal collar with darker brown marking between

lateral margin and pronotal suture, marking darkest

at suture and often connected with transverse band

on pronotal suture, anterolateral corner not dark-

ened; anterolateral corner occasionally bidentate.

Mesonotum ochraceous brown to brown; median fas-

cia narrow, broadest on posterior half of mesonotum,

usually black and distinct but sometimes brown and

rather indistinct; paramedian fasciae on anterior half

of disc as in P. radha, on posterior half of disc present

as black markings of varying size anterior of cruci-

form elevation; mesonotal fissures ochraceous brown;

narrow black triangular markings immediately lateral

of mesonotal fissures at anterior margin of disc always

distinct and small; lateral fasciae absent; posterior

margin with black spots lateral of anterior arms of

cruciform elevation that occasionally merge with

markings anterior of cruciform elevation. Cruciform

BEUK: The Platylomia radha group

elevation ochraceous to brown, paler than mesonotal

disc. Posterior margin and median depressions of cru-

ciform elevation often with dense whitish coating.

Katepimeral lobe (fig. 22) mostly as in P. radha, apex

reaching over base of operculum.

Tegmina and wings. — Tegmina and wings mainly

as in P. radha. Infuscations on basal veins of second

and third apical cells of tegmina usually quite distinct

though not very dark, usually more distinct on basal

vein of third apical cell. Apices of longitudinal veins

of apical cells occasionally with almost indistinguish-

able spots.

Legs. — Fore legs ochraceous to brownish; femora

ochraceous, anterodorsally, posterodorsally and pos-

teroventrally with darkened longitudinal stripes that

usually are connected by dark apical ring; tibiae

brownish, dorsal surface with ochraceous stripe from

base almost to apex; tarsi brownish. Mid legs brown-

ish; tibiae with dorsal surface and apical quarter dark-

ened; tarsi dark brown. Hind legs ochraceous; tibiae

slightly to distinctly darkened dorsally at base and on

apical fourth; tarsi ochraceous to brown. Fore femur

with gap between middle and posteroventral distal

spines rounded and not very narrow. Hind tibia

sometimes with only two anteroventral spines.

Male operculum (fig. 20). — Not very long, reach-

ing from about anterior margin of fifth to anterior

margin of sixth abdominal segment, 2.1-2.2 times as

long as maximum width distal of constriction; ochra-

ceous to brown, sometimes with greenish tinge, area

around lateroproximal corner and usually also part of

the lateral margin dark brown to black but latero-

proximal corner itself ochraceous. Medial margin dis-

tal of constriction convex but occasionally for some

distance in distal half almost straight. Distal margin

convex, sometimes straightening medially, usually no

distinct apex present. Lateral margin distal of con-

striction convex to apex but occasionally almost

straight about halfway. Constriction at about 0.4 of

length of operculum, lateral concavity longer and

deeper than medial concavity and at deepest point al-

most angular, broadest part of operculum distal of

constriction 1.5-1.7 times as wide as minimum width

in constriction. Distance between opercula at con-

strictions about 1.2-1.4 times as wide as minimum

width in constriction. Opercula at point of closest ap-

proximation separated for a distance of 0.3-0.4 times

maximum width between opercula at constrictions.

Operculum close to abdomen, surface of operculum

both proximal and distal of constriction convex thus

creating impression of transverse fold at level of con-

striction.

Male abdomen. — Abdomen as in P. radha but

slightly paler, about 1.2-1.3 times as long as head and

thorax together. Dorsal part of tergites brownish to

castaneous, tergites often somewhat darkened along

159

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 16-22. Platylomia bocki (Distant, 1882), male. — 16, pygofer and uncus in ventral view, lectotype; 17, pygofer and un-

cus in right lateroventral view, lectotype; 18, pygofer in dorsal view, Vieng Vai; 19, uncus in anterior view, lectotype; 20, ab-

domen and right operculum in lateroventral view, Khao Yai; 21, right timbal covering, Khao Yai; 22, right katepimeral lobe,

Khao Yai.

160

posterior margins; sternites and ventral parts of ter-

gites paler than dorsal parts but less so towards geni-

talia. Tergite 2 with small paramedian areas of

whitish dusting on posterior margin, tergites 3-4 with

small lateral areas of whitish dusting on anterior mar-

gin, tergite 8 with thin whitish dusting, dusting on

other tergites without distinct pattern. Posterior mar-

gin of tergite 7 with many short dark spinules, of ter-

gite 6 and occasionally also of tergite 5 with few spin-

ules on lateral part. Sternite 7 with shallow postero-

medial emargination. Timbal covering (fig. 21) as in

P. radha, ochraceous to ochraceous brown, about 1.1-

1.3 times as long as wide; lateral margin almost

straight to weakly convex.

Male genitalia (figs. 16-19). — Brown, dorsal part

of pygofer somewhat darker than ventral part; uncus

lobes usually darker than remainder of uncus. Basal

pygofer lobes (figs. 16-17) almost as in P. radha but

reaching further between lateral margins of pygofer;

hairs on anteroventral margin of pygofer mostly short

and erect but interspersed with longer hairs, hairs de-

creasing in length towards basal lobes and continuing

on pygofer surface enclosed by anteroventral margin;

pygofer surface anterior and lateral of anteroventral

margin with scattered hairs. Dorsal part of pygofer as

in fig. 18. Basal part of uncus small, little globose and

narrow (figs. 16-17), near bases of uncus lobes with

occasional short erect hairs. Uncus lobes (figs. 16-17,

19) with medial margin weakly concave; mediodistal

corner rounded; distal margin convex near mediodis-

tal corner, concave near laterodistal corner; laterodis-

tal corner angular to slightly pointed; lateral margin

strongly curved medially close to base and uncus lobe

then broadening slightly towards distal margin, later-

al margin usually smooth near laterodistal corner but

occasionally with one or two small wart-like out-

growths. Each uncus lobe on ventral (outer) surface

with curved broad ridge from mediodistal corner al-

most to basal part of uncus where ridges nearly meet

(fig. 19). Surface of uncus lobes covered with short

erect hairs along margins, especially medial margin;

ventral (outer) surface with very few hairs; dorsal (in-

ner) surface with strong erect hairs at bases of uncus

lobes. Uncus with some strong erect hairs laterally

where uncus lobes and basal part of uncus are joined.

Female operculum. — Operculum brownish, dark-

er on lateral margin close to lateroproximal corner.

Lateral margin weakly concave distal of lateroproxi-

mal lobe; laterodistal corner angularly rounded; distal

margin weakly concave on lateral half, weakly convex

on medial half; mediodistal corner broadly rounded;

medial margin short and weakly convex; lateral and

medial margin of each operculum convergent from

base.

Female abdomen. — Abdomen little shorter than

head and thorax together. Dorsal part of tergites cas-

Beuk: The Platylomia radha group

taneous; sternites and ventral parts of tergites little

paler. Tergites without darker spots near lateral mar-

gin. Tergites with thin whitish dusting but without

distinct pattern. Posterior margins of tergites 3-5 with

some short and slender dark spines laterally but spines

continuing more medially on tergite 5, posterior mar-

gins of tergites 6-7 with more numerous and longer

spines along whole length, of tergite 8 with scattered

long spines medially. Sternite 7 with angularly round-

ed posteromedial emargination.

Female genitalia. — Dorsal part of pygofer almost

castaneous, ventral part more brownish; distal part of

ovipositor sheath dark castaneous brown. Dorsal

margin of pygofer shorter than three preceding seg-

ments, in lateral view weakly concave; ventral margin

weakly convex. Ovipositor sheath not reaching as far

as apex of caudodorsal beak; anal valve not reaching

as far as ovipositor sheath.

Measurements in mm (d: n= 5; 2:n= 1).— Body

length: d: 43.5-47.0 (44.7 + 1.0), 2: 39.5; head

width: 4: 13.0-13.9 (13.6 + 0.3), 2: 14.0; maxi-

mum pronotum width: d: 13.8-15.0 (14.5 + 0.4),

Orml5(0;tegmen length: 8:525-55.5.64.1 #121),

2: 54.0.

Distribution (fig. 3)

P. bocki is distributed throughout Indo-China

(Laos, Thailand, Vietnam) and southern China.

Remarks

Moulton (1923: 101) already pointed out that the

‘Type d and only known specimen’ was labelled as

cited above. Since both the locality label and the iden-

tification label are Distant’s, this specimen is desig-

nated as lectotype.

Material examined. — cHına: Cheli [Yunjinghong],

Yunnan, 550 m, 30.iv.1957, 1 & (1zas); Emei [Emei Shan],

Sichuan, 500 m, 22.vii.1974, 1 d (Nwas); Jinghong [Yun-

jinghong], Yunnan, 3.iv.1991, 1 d (Nwas); Mengla, Yun-

nan, 30.iv.1984, 1 dé (Nwas); Menglun, Yunnan,

10.v.1984, 1 d (Nwas); Yaoqu, Yunnan, 1600 m, 4.v.1991,

1 3 (Nwas); Yiwubanna, Yunnan, 650 m, 1.v.1964, 4 &

(1zas); Yunnan, 5.v.1957, 1 d (suu); VIETNAM: Cochin

Chine, 1878, Pierre, 1 d, 1 9 (mnun); Laos: Haut

Mékong, Vieng Vai, 23.v.1918, R.V. de Salvaza, 1 3

(BMNH); Luang Namtha [Louang Namtha], N. Laos,

4.v.1994, Y. Miyake, 4 & (suu); Luang Prabang, Pan Houei

Ket [Ban Houay Ket], 17.111.1920, R.V. de Salvaza, 1 3

(BMNH); THAILAND: Khao Yai NP, 800 m, 14.iv.1988, M.G.

Allen, 3 & (BMNH); Khao Yai NP, 850 m, 19.iv.1988, T.W.

Harman, 1 d (zman); Khao Yai, Nakorn Nayok [Nakhon

Nayok], ca. 800 m, 14.vi.1983, Kuroko, Moriuti, Arita &

Yoshiyasu, 1 d (opu); Phuping Palace [Phuphing Palace],

Changmai [Chang Mai] Prov., North Thailand, v.1983, Y.

Komiya, 1 2 (suu); same data, 15.vi.1983, 1 d (oru);

Saraburi [Sara Buri], 15.iv.1973, Y. Yoshiyasu, 1 d (suv);

LOCALITY UNCERTAIN: Sumatra or Siam, Bock, d lectotype

Dundubia bocki Distant (BMNH).

161

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 23-29. Platylomia ficulnea (Distant, 1892), male, paralectotype Karen Hills. — 23, pygofer and uncus in ventral view; 24,

pygofer and uncus in right lateroventral view; 25, pygofer in dorsal view; 26, uncus in anterior view; 27, abdomen and right

operculum in lateroventral view; 28, right timbal covering; 29, right katepimeral lobe.

162

Platylomia ficulnea (Distant, 1892)

(figs. 3, 23-29)

Cosmopsaltria ficulnea Distant, 1892a: 102. — Lectotype d

(here designated) of Cosmopsaltria ficulnea Distant:

‘Carin Ghecù / 1300-1400 m / L. Fea II-III.88° [printed]

and ‘Cosmopsaltria / ficulnea / type Dist.’ [Distant’s

handwriting].

Cosmopsaltria ficulnea. — Distant 1892b: 154, xii, pl. XV

figs. 21-21b; Fea 1897: 608.

Platylomia ficulnea. — Distant 1906a: 106; Distant 1906b:

61; Distant 1912: 49; Kato 1932: 166; Liu 1940: 115;

Metcalf 1963: 617.

P. ficulnea is closely related to P. radha and P. bocki

but can easily be distinguished from both species by

the presence of the markings on the tegmina. P. ficul-

nea and P. radha are the only species of the P. radha

group with attenuate apices of the opercula. The

structure of the genitalia of both species is also very

similar but in P. ficulnea the basal pygofer lobes are

narrower and the uncus lobes do not have a flange on

the lateral margin but occasionally they have a small

pointed protuberance.

Description

Body brownish to castaneous with some parts of

head and thorax lighter; mesonotum with median

and paramedian fasciae partly developed; tegmina

with infuscations at basal veins of second, third, fifth,

and seventh apical cells, and at apices of longitudinal

veins of apical cells. Opercula broad and attenuated at

apex.

Head. — Postclypeus and anteclypeus brown; post-

clypeus little swollen, in dorsal view about as long as

distance between frontoclypeal suture and anterior

margin of pronotum. Vertex brown but lateral parts,

central part of supra-antennal plates, part posterior of

eyes and area enclosed by ocelli darker; posterior mar-

gin with pair of dark brown spots close to eyes. Genae

and lori brown. Frontoclypeal suture as in P. radha.

Rostrum as in P. radha; reaching posterior margin of

hind coxae.

Thorax. — Pronotum distinctly broader than head.

Pronotal disc brown but sometimes with greenish

tinge, anterior margin and medial area paler; disc im-

mediately posterior of anterior margin between para-

median lobes with indistinct to dark brown transverse

band that is broadly interrupted medially; central fas-

ciae not developed except for two brownish triangular

markings with their bases at pronotal suture; trans-

verse part of pronotal suture with black band that

does not continue laterally to level of markings on lat-

eral part of pronotal collar. Anterolateral part of

pronotal collar concolorous with pronotal disc, re-

mainder of paler than disc; posterior margin black

and sometimes broadening slightly in posterior cor-

ners; anterolateral corner angular or with small but

distinct lateral tooth; posterolateral corner rounded.

Beuk: The Platylomia radha group

Mesonotum brown, somewhat darker on posterior

half of disc and sometimes with greenish tinge on

paler parts; median fascia brown, concolorous with

posterior half of disc so indistinguishable there; para-

median fasciae on anterior half of disc castaneous and

running immediately medial of mesonotal fissures,

narrow near anterior margin of disc then broadening

and slightly curved medially; mesonotal fissures

slightly paler than remainder of mesonotal disc; ante-

rior margin of disc with castaneous triangular mark-

ings immediately lateral of mesonotal fissures; lateral

fasciae absent. Cruciform elevation concolorous with

anterior half of mesonotal disc but anterior arms

sometimes paler. Katepimeral lobe (fig. 29) shorter

than broad at base; apex rounded. Surface of

katepimeral lobe concave in dorsoventral direction

and convex in longitudinal direction; surface covered

with short, almost waxy hairs, margins with slightly

longer, fine hairs. Apex of katepimeral lobe just reach-

ing over base of operculum.

Tegmina and wings. — Tegmina almost entirely

hyaline with indistinct brownish reticulation along

distal margin and in apical cells; basal veins of second,

third, fifth, and seventh apical cells with distinct

brown infuscation, infuscation on base of third apical

cell often continuing on anterior basal vein of fourth

apical cell; apices of longitudinal veins of apical cells

with rounded dark brown spots; basal cell almost

completely yellowish fumose except for very narrow

hyaline strip along posterior margin. Veins of tegmen

ochraceous to brownish but sometimes with greenish

tinge, darker along cordial fold, on most cross veins,

and on distal of forks in veins. Wings hyaline except

for very narrow strip posterior of first anal vein. Veins

of wings brownish but sometimes with greenish tinge,

medial vein and veins close to wing margin darker.

Legs. — Fore legs brownish; femora posteroventral-

ly with darkened longitudinal stripe; tibiae slightly

darkened distally; tarsi slightly darkened, especially

distally. Mid legs brownish; tibiae brownish, slightly

darkened at base and darkening slightly on distal half

towards apex; tarsi little darker than femora. Hind

legs brownish; tibiae little darker than femora but less

so on band at about one fifth from base; tarsi little

darker than femora. Fore femur with gap between

middle and distal posteroventral spines shallow and

not very broad. Hind tibia with three dorsal spines

and four anteroventral spines; spines brownish.

Male operculum (fig. 27). — Operculum broad and

attenuate at apex, reaching between anterior and pos-

terior margin of sixth abdominal segment, about

(2.5) 2.9 times as long as maximum width distal of

constriction; brown but sometimes slightly darker on

distal part, lateral margin in lateroproximal corner

dark brown to black but lateroproximal corner itself

paler. Medial margin distal of constriction smoothly

163

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

convex to apex, on distal third appearing concave be-

cause of shape of apex. Apex of operculum gully-

shaped, lateral of midline, and divergent from ab-

domen. Lateral margin distal of constriction first con-

vex but concave on distal third. Constriction at

0.3-0.4 of length of operculum, medial concavity

very shallow, lateral concavity longer and only slight-

ly deeper than medial concavity, broadest part of op-

erculum distal of constriction 1.1-1.3 times as wide as

minimum width in constriction. Distance between

opercula at constrictions 0.5-0.7 times as wide as

minimum width in constriction. Opercula at point of

closest approximation separated for a distance of 0.5-

0.6 times maximum width between opercula at con-

strictions. Operculum close to abdomen on proximal

part, distal half slightly divergent from abdomen; sur-

face convex in longitudinal direction near medial and

lateral margins only, surface more strongly convex in

transverse direction, especially on distal third.

Male abdomen. — Abdomen 1.4-1.5 times as long

as head and thorax together. Dorsal part of tergites

brown to castaneous with anterior margins of tergites

1-3 and posterior margins of tergites 1-2 ochraceous,

posterior margins of tergites 4-6 dark brown, lateral

parts always paler; sternites and ventral parts of ter-

gites ochraceous to brownish. Dorsal part of tergites

3-4 (5) with dense white pilosity in anterolateral cor-

ners, tergite 4 with paramedian, transverse oblong

spots of white dusting close to anterior margin. Pos-

terior margin of tergite 7 with many short dark spin-

ules, of tergite 6 with longer spinules along whole

length but few medially, of tergite 5 with few spinules

on lateral part. Sternite 7 with posteromedial emar-

gination considerably deeper than in P. radha. Tim-

bal covering (fig. 28) brownish to castaneous, about

as long as wide; medial margin weakly convex; distal

margin rounded; lateral margin weakly convex to

straight.

Male genitalia (figs. 23-26). — Ochraceous brown;

dorsal part of pygofer somewhat darker. Basal pygofer

lobes (fig. 23) as in P. radha but not as broad and

reaching further between lateral margins of pygofer;

hairs on anteroventral margin of pygofer and on py-

gofer surface enclosed by anteroventral margin as in

P. radha but slightly shorter and interspersed with oc-

casional long hairs; hairs on outer surface of pygofer

as in P. radha. Dorsal part of pygofer as in fig. 25.

Basal part of uncus short, little globose and narrower

than maximum width uncus (fig. 23), near bases of

uncus lobes usually with erect hairs but these hairs

shorter and thinner than in P. radha. Uncus lobes

(figs. 23-24, 26) short; medial margin, mediodistal

corner and distal margin in P. radha, without pointed

protuberances; laterodistal corner somewhat pro-

duced and bluntly pointed, tip directed to anterior;

lateral margin strongly curved medially at base, uncus

164

lobe narrowest about halfway to distal margin, lateral

margin weakly convex from narrowest point in uncus

lobe to laterodistal corner; lateral margin sometimes

with small pointed protuberance near laterodistal cor-

ner. Ventral (outer) surface of uncus lobes virtually

smooth. Bases of uncus lobes with erect hairs laterally

but only very close to base of uncus and with few

erect thick hairs on lateral margin; dorsal (inner) sur-

face with scattered erect thick hairs; ventral (outer)

surface with short fine hairs on distal half only.

Female. — Unknown.

Measurements in mm (d: n = 4). — Body length:

6): 47.0255.5)(52.0) 3.0): head width: 6135152

(14.5 + 0.6); maximum pronotum width: d': 14.7-

16.5 (15.7 + 0.6); tegmen length: d: 54.5-62.0

(593255)

Distribution (fig. 3)

This species is so far only recorded from Burma

and India (Assam) but the record from Assam

(Distant 1906a) could not be confirmed.

Remarks

Cosmopsaltria ficulnea was described from an un-

known number of specimens from ‘Carin Ghecù’ and

‘Karen Hills’. The type series of P. ficulnea consists of

four males, two males at both the MSNG and the

BMNH. One male in each of these collections bears a

type label but neither can be considered to be a holo-

type as this was not mentioned with the decription

nor was a lectotype designation ever published. The

specimen at MSNG with Distant’s type label is desig-

nated as lectotype. The other three males are labelled

as paralectotypes.

Material examined. — BURMA: Ghecü, Carin, 1300-

paralectotype Cosmopsaltria ficulnea Distant (MSNG);

same data, 1 d paralectotype (BMNH); Karen Hills,

Doherty, 1 d paralectotype (BMNH).

Platylomia malickyi sp. n.

(figs. 3, 30-36)

Type material. — Holotype d: ‘Thailand 24.4.-

1.5.1989 / Changmai Zoo Lichtfalle / 400m,

98°57’E, 18°49’N / Chantaramongkol & Malicky

(1zu1). — Paratypes: VIETNAM: 6 km S. of Dalat [Da

Lat], 1400-1500 m, 9.vi-7.vii.1961, N.R. Spencer, 1

d (BPBM); Datanca, Dalat [Da Lat], S. Vietnam,

22.v.1992, 1 d (suu); Laos: Ban Van Eue, Vientiane

Province, 15-31.v.1965, 1 2 (BPBM); same data,

15.v.1966, native collector, 1 d (BPBM); Lakhon

[Muang Lakhonpheng], 1878, J. Harmand, 1 d

(MNHN); Muong Om [Muang Hom], Luang Prabang

[Louangphrabang], 17.v.1920, R.V. de Salvaza, 1 d

(BMNH); Sen Kam, Haut Mékong, 29.v.1918, R.V.

de Salvaza, 1 d (BMNH); Vieng Vai, Haut Mékong,

23.v.1918, R.V. de Salvaza, 1 6 (BMNH); THAILAND:

Amphoe Muang Chiang-mai [Chiang Mail,

24.iv.1973, Y. Yoshiyasu, 3 d (suv); same data,

25.iv.1973, 1 d (suu); Bang Khun Klang, Doi

Inthanon, 18°32’N 98°32’E, 1200 m, 27.iv-

1.v.1989, Chantaramongkol & Malicky, light trap,

1 d (zur); Chiang Dao, Chiang Mai, 1 6, 1 9

(suu); Chiang Mai, 23.iv.1976, S. Saito, 1 d (suu);

Changmai [Chiang Mai] Zoo, 18°49’N 98°57’E, 400

m, 17-24.iv.1989, Chantaramongkol & Malicky,

light trap, 4 d (1zur); same data, 2 d (ZMAN); same

data, 24.iv-1.v.1989, 4 d (zut); Chiengmai [Chiang

Mail, 22.vi.1952, D. & E. Thurman, 1 d (usnm);

Doi Suthep, Chiang Mai, N. Thailand, 1.v.1984, T.

Endo, 1 d (suv); Doi Suthep, East slope,

15.vii.1962, E.S. Ross & D. Cavagnaro, 1 d (casc);

Mae Sa, Chiang Mai, 2.v.1984, T. Endo, 1 d, 1 9

(suu); Phu Khieo Wildlife Sanctuary, Khon San,

Chaiyaphum Province [Changwat Chaiyaphum],

N.E. Thailand, 16°30’N 101°46’E, 800 m, evergreen

rain forest, at light, 13-15.v.1988, M.J.D. Brendell,

1 d (BMNH); Wiang Pa Pao, Chiang Rai Prov., 2-

10.v.1990, 1 d (suu); BURMA: Maymyo, 29.iv.1901,

1 2 (BMNH); Maymyo, v.1910, H.L. Andrewes, 1 9

(BMNH).

Other material examined. — cHINA: Dale, Yunnan,

1650 m, 31.vi.1991[?], 1 d, 1 2 (1245); Jingdong

Uinping], Yunnan, 1200 m, 29.v.1956, 1 d (Nwas);

Lincang, Yunnan, 1110 m, 23.vi.1980, 1 d (roy);

Menghaineshan, Yunnan, 1200-1300 m, 24.iv.1957,

1 & (izas); Nefu, Yunnan, 1350 m, 2.vi.1980, 1 &

(roy); Shuangjiang [Mengmeng], Yunnan, vi.1977,

1 & (Foyp); Simao, Yunnan, 1350 m, 11.v.1957, 1 3

(1ZAs); Xishuangbanna, Yunnan, 750 m, 1 d (1zas);

Yunnan, vi.1953, 1 d (Nwas); THAILAND: Phu Rua,

Loei, c. 800 m, 15-19.viii.1987, S. Moriuti, T. Saito,

Y. Arita & Y. Yoshiyasu, 1 9 (OPU); EXACT LOCATION

UNKNOWN: S. Annam, 1918, C.B. Kloss, 1 d

(MNKM).

P. malickyi and P. bocki are the smallest species of

the P. radha group but P. malickyi can easily be dis-

tinguished from P. bocki by the presence of at least

some distinct markings on the tegmina and the more

elaborate light-dark pattern on the pronotum and

mesonotal disc. P. malickyi displays a large amount of

variation in the markings on the tegmina but some

distinct markings are always present. The pronotum

and mesonotal disc also show considerable variation

in the degree of darkening; in some cases the black

posterior margin of the pronotal collar and the black

band across the transverse part of the pronotal suture

completely enclose the pale area on the posterior part

of the pronotal collar.

BEUK: The Platylomia radha group

Description.

Body ochraceous brown to castaneous brown with

darker pattern on head and thorax, ventral parts paler

than dorsal parts; opercula short and rounded at apex;

legs predominantly brownish. Tegmina usually with

markings at basal veins of second, third, fifth, and

seventh apical cells, and at apices of longitudinal veins

of apical cells.

Head. — Postelypeus brown to castaneous but paler

along clypeal suture and lateral margins, dorsal part

usually somewhat darker than ventral; postclypeus lit-

tle swollen, in dorsal view about as long as distance

between frontoclypeal suture and anterior margin of

pronotum. Anteclypeus concolorous with ventral

part of postclypeus or somewhat darkened on lateral

surfaces. Vertex brownish to castaneous but paler on

supra-antennal plates and area posterior of supra-an-

tennal plates, pale parts sometimes extending as far as

posterior margin; area around ocelli with black mark-

ing that just or just not reaches frontoclypeal suture

anteriorly, marking posteriorly usually not reaching

posterior margin of vertex but if reaching margin

then enclosing brown spot medially at margin and ex-

tending laterally towards eyes along margin; posterior

margin with pair of black spots close to eyes. Genae

ochraceous brown to brown but darker just below an-

tennae. Supra-antennal plates with one or more

curved grooves, posterior groove joins lateral part of

frontoclypeal suture. Frontoclypeal suture semicircu-

lar to distinctly trapezoid, medial part much broader

than distance between lateral margins of lateral ocelli.

Rostrum ochraceous brown to brown, darkened at tip

only, reaching almost to distinctly beyond posterior

margin of hind coxae but never further than halfway

sternite 1.

Thorax. — Pronotum slightly to distinctly broader

than head. Pronotal disc mainly brownish to casta-

neous, anterior margin and area enclosed by central

fasciae paler, central fasciae darker and sometimes al-

most blackish, broadening posteriorly and forming

distinct dark brown to black triangular spots that usu-

ally meet at posterior margin of disc; disc immediate-

ly posterior of anterior margin with dark brown to

black transverse band that is interrupted medially and

narrows laterally, band at level of interruption con-

nected with central fasciae; transverse part of pronotal

suture with dark brown to black band that is con-

nected with central fasciae; medial part of band large-

ly on disc, lateral part largely on collar. Pronotal col-

lar ochraceous brown to brown with black posterior

margin that broadens in posterior corners, lateral part

collar occasionally entirely dark except anteriorly; col-

lar lateral of disc darker, sometimes with exception of

area around lateral tooth; dark area on lateral part of

collar always connected with dark band on transverse

part of pronotal suture and occasionally also connect-

165

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 30-36. Platylomia malickyi sp. n., male, paratype Muong Om. — 30, pygofer and uncus in ventral view; 31, pygofer and

uncus in right lateroventral view; 32, pygofer in dorsal view; 33, uncus in right anterolateral view; 34, abdomen and right op-

erculum in lateroventral view; 35, right timbal covering; 36, right katepimeral lobe.

166

ed with black posterior margin of collar; anterolateral

corner with small lateral tooth, ranging from blunt

and angular to pointed; posterolateral corner rounded

but sometimes part of margin only weakly convex.

Mesonotum ochraceous brown to castaneous; medi-

an fascia usually distinct, dark brown to black and

broadened about halfway between anterior margin

and cruciform elevation; paramedian fasciae on ante-

rior half of disc dark brown to black and running im-

mediately medial of mesonotal fissures, fasciae nar-

row near anterior margin of disc then broadening and

curved medially till they meet or almost meet with

median fascia where latter is broadened; paramedian

fasciae on posterior half of disc present as dark brown

to black markings of varying size in front of cruciform

elevation, only rarely extending anteriorly and meet-

ing with paramedian fasciae on anterior half of disc;

mesonotal fissures ochraceous to ochraceous brown,

much paler than any other part of mesonotal disc; an-

terior margin of disc with narrow dark brown to black

triangular spots immediately lateral of mesonotal fis-

sures; lateral fasciae on anterior half of disc present as

black markings at anterior margin of disc and these

markings sometimes merging with triangular spots

lateral of mesonotal fissures; lateral fasciae on posteri-

or half of disc ranging from indistinct dark brown

spots lateral of cruciform elevation to distinct black

fasciae reaching halfway to anterior margin, lateral

fasciae posteriorly often fused with remnants of para-

median fasciae in front of cruciform elevation.

Cruciform elevation somewhat darker than mesono-

tal fissures, posterior margin at least laterally dark

brown to blackish, sometimes also medially.

Katepimeral lobe (fig. 36) rather variable of shape,

shorter or longer than broad at base; apex rounded to

slightly angular. Surface of katepimeral lobe almost

flat to weakly concave along dorsal margin, covered

with short hairs only. Apex of katepimeral lobe only

just to distinctly reaching over base of operculum.

Tegmina and wings. — Tegmina almost entirely

hyaline with indistinct brownish reticulation along

distal margin and sometimes in apical cells; basal

veins of second and third apical cells along whole

length with dark brown markings that sometimes ex-

tend for short distance along longitudinal veins back

to base of tegmen and may cause the markings to

merge, fifth and seventh apical cells usually with

smaller dark brown spots but these spots sometimes

absent or indistinct; apices of longitudinal veins of

apical cells with rounded dark brown spots but these

spots sometimes absent or indistinct on posterior part

of tegmina; basal cell yellowish to pale brown fumose.

Veins of tegmen brownish, darker on most cross veins

and parts where veins meet or split. Wings hyaline.

Veins of wings brownish, medial vein, and veins close

to wing margin darker.

Beuk: The Platylomia radha group

Legs. — Fore legs ochraceous to brownish; femora

posteroventrally with darkened longitudinal stripe,

apically with dark brown ring that may be interrupt-

ed dorsally, anterior and anteroventral surface often

somewhat darkened; tibiae brown, dorsal surface with

paler stripe on basal quarter to half; tarsi brown. Mid

legs ochraceous brown; femora sometimes darkened

anteriorly and posteriorly at apex; tibiae darkened

dorsally at base and on apical quarter; tarsi brown.

Hind legs ochraceous; femora sometimes with very

indistinct dorsal stripes; tibiae darkened dorsally at

base and with darker ring just before apex; tarsi

ochraceous brown. Fore femur with gap between

middle and distal posteroventral spines shallow to

deep, narrow.

Male operculum (fig. 34). — Not very long, reach-

ing just beyond anterior margin of fifth to just be-

yond anterior margin of sixth abdominal segment,

2.1-2.3 times as long as maximum width distal of

constriction; ochraceous to brown, lateral margin

darkened close to lateroproximal corner of opercu-

lum. Medial margin distal of constriction convex but

occasionally less convex for some distance in distal

third. Apex of operculum rounded, only rarely with a

tendency to be slightly angular, tip from just medial

to just lateral of midline. Lateral margin distal of con-

striction strongly convex to apex. Constriction at 0.4-

0.5 of length of operculum, lateral concavity longer

and deeper than medial concavity, broadest part of

operculum distal of constriction 1.4-1.6 times as wide

as minimum width in constriction. Distance between

opercula at constrictions about 1.0-1.1 times as wide

as minimum width in constriction. Opercula at point

of closest approximation separated for a distance of

0.4-0.5 times maximum width between opercula at

constrictions. Surface of operculum convex except at

level of constriction thus creating impression of trans-

verse fold at level of constriction.

Male abdomen. — Abdomen about 1.2-1.3 times as

long as head and thorax together. Dorsal part of ter-

gites unicolorous brownish to castaneous; sternites

and ventral parts of tergites somewhat paler. Lateral

parts of tergites 2-4 with thin whitish dusting, dust-

ing on other tergites without distinet pattern. Only

posterior margin of tergite 7 with short dark spinules.

Sternite 7 with very shallow posteromedial emargina-

tion. Timbal covering (fig. 35) brownish, paler than

dorsal part of tergites, about 1.0-1.2 times as long as

wide; medial margin virtually straight; mediodistal

corner, distal margin and laterodistal corner rounded;

lateral margin almost straight, convergent with medi-

al margin.

Male genitalia (figs. 30-33). — Pygofer brown to

castaneous on dorsal side, remainder paler; uncus

brown to castaneous, often paler on basal part and ad-

joining part of uncus lobes. Basal pygofer lobes (fig.

167

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

30) narrower than and not as broadly rounded as in

P. radha; hairs on anteroventral margin of pygofer

erect and not very long, decreasing in length only lit-

tle towards basal lobes and continuing on pygofer sur-

face enclosed by anteroventral margin, pygofer sur-

face anterior of medial part of anteroventral margin

with scattered hairs. Dorsal part of pygofer as in fig.

32. Basal part of uncus short, little globose and nar-

rower than maximum width uncus (figs. 30-31), near

bases of uncus lobes with occasional short erect hairs.

Uncus lobes (figs. 30-31, 33) broad; medial margin

weakly concave to straight; mediodistal corner round-

ed, occasionally with short blunt spine on dorsal (in-

ner) side; distal margin with narrowing lobule on lat-

eral half, apical margin of lobule curved dorsad (in-

wards) and two-tipped, each tip broad and very short,

lateral margin of lobule often with short blunt spine

that is curved dorsad (inwards); laterodistal corner

angular, usually with two short but pointed protuber-

ances and often a third one more proximal on lateral

margin; lateral margin shallowly concave. Ventral

(outer) surface of uncus lobes with short grooves run-

ning toward lateral margins of distal lobules. Ventral

(outer) surface of uncus lobes with many short, thin

erect hairs on proximal half and several thicker erect

hairs close to basal part of uncus; dorsal (inner) sur-

face of uncus lobes with long, thick erect hairs near

lateral margin on proximal half, otherwise with many

short, thin erect hairs, especially close to margins.

Female operculum. — Operculum pale brownish,

lateral margin dark brown to black except at latero-

proximal lobe. Lateral margin evenly convex distal of

lateroproximal lobe; laterodistal corner angularly

rounded; distal margin weakly concave to evenly con-

vex; mediodistal corner broadly rounded; medial

margins short and convex and strongly divergent.

Female abdomen. — Abdomen 0.9-1.0 times as

long as head and thorax together. Dorsal part of ter-

gites castaneous brown but on tergites 1-2 paler, pos-

terior margins darker, tergites 3-6 with indistinct

darker spots near posterolateral margin; sternites and

ventral parts of tergites paler but less so on segments

6-7. Tergite 2, lateral parts of tergites 3-4 and medial

parts of tergites 5-7 with thin whitish dusting.

Posterior margins of tergites 3-6 with short dark

spines laterally, of tergite 7 with more numerous and

longer spines along whole margin, of tergite 8 with

scattered spines except laterally, medial spines more

slender than paramedial spines. Sternite 7 with angu-

lar posteromedial emargination.

Female genitalia. — Lateral and ventral part of py-

gofer brownish, dorsal part more castaneous but paler

along mid-line; distal part of ovipositor sheath casta-

neous. Dorsal margin of pygofer shorter than three

preceding segments, in lateral view weakly concave;

ventral margin weakly convex. Ovipositor sheath

168

reaching as far as apex of caudodorsal beak; anal valve

not reaching as far as caudodorsal beak.

Measurements in mm (d:n= 5; 9: n= 3). — Body

length: d: 40.0-43.0 (41.7+ 0.8), 2: 38.0-39.5

(38.8 + 0.6); head width: d : 12.3-12.8 (12.6 + 0.1),

9: 13.1-13.6 (13.5 + 0.3); maximum pronotum

width: d: 12.9-13.9 (13.3 + 0.3), 2: 13.9-14.9

(14.5 + 0.4); tegmen length: d : 46.5-50.5 (48.7 +

LD), Se LOSES G28 22 1D),

Variation

The markings on the tegmen show some variation.

The spots on the apices of the longitudinal veins of

the apical cells are very variable in size and can be pre-

sent at all apical cells, or gradually weakening from

the first to the seventh apical cell and almost be ab-

sent there, or very indistinct or absent altogether.

Likewise can the markings on the basal veins of sec-

ond, third, fifth, and seventh apical cells be indistinct

and they can even be absent on the basal veins of the

fifth and seventh apical cell. The tendency for reduc-

tion of wing markings is especially notable in the fe-

males from Burma. These specimens are also the

specimens with the most extensive darkening on

pronotum and mesonotum.

Distribution (fig. 3)

P. malickyi is known from Indo-China (Burma,

Laos, Thailand, and Vietnam) and southern China

(Yunnan).

Etymology

The species is named after Dr Hans Malicky who

works on Trichoptera and who is one of the collectors

of the series that constitutes half of the type material.

Platylomia bivocalis (Matsumura, 1907)

(figs. 3, 37-44)

Cosmopsaltria bivocalis Matsumura, 1907: 97. — Myers

1929: 137, 222; Kato 1932: 96.

Platylomia bivocalis. — Distant 1912: 49; *Matsumura 1913:

72, pl. IX fig. 2; Schumacher 1915a: 79; Schumacher

1915b: 111; Matsumura 1917: 198, 211; Sonan 1921:

[165]; Kato 1925: 20, 44; Kato 1927: 28; *Matsumura

1930: [15], pl. III fig. 2; Kato 1931: 51 [17], 64 [29];

Matsumura 1931: [1237, text figure]; Esaki 1932: [1704,

fig. 3365]; Schmidt 1932: 126; Kato 1932: 206, 217,

328, pl. XXI fig. 4, pl. XXXI fig. N; *Kato 1933a: pl.17

fig. 4, pl. 18 fig. 5; Kato 1933b: 11; Kato 1938a: 308;

Kato 1938b: 18; Kato 1956: 96, 118, 122, 137; Metcalf

1963: 615; Ishida 1968: 3; Duffels & Van der Laan

1985: 120.

P. bivocalis is the only species of the P. radha group

that does not occur on the mainland of the Southeast

Asia or the Thai-Malay Peninsula. It can easily be dis-

tinguished from the other species of the group by the

brownish hyaline tegmina, the darker brown infusca-

tion along the veins in at least the distal part of the

tegmina, the extensive darkening on tergites, and the

pattern of white dusting on tergites 3, 5, 6, and 8.

Description

Head and thorax brownish, abdomen more casta-

neous; pronotum with well developed central fasciae;

mesonotum with well developed median and parame-

dian fasciae and with spot-like lateral fasciae; tegmina

with markings on basal veins of second, third, fifth

and seventh apical cells and on apices of longitudinal

veins, but often all veins in distal part of wing some-

what infuscated. Opercula broad and not very long.

Head. — Postclypeus and anteclypeus ochraceous

brown to brown, dorsal and anterior parts of post-

clypeus slightly darker except for anteromedial spot

and small area at frontoclypeal suture; postclypeus lit-

tle swollen, in dorsal view about as long as distance be-

tween frontoclypeal suture and anterior margin of

pronotum. Vertex brownish, darker on vertex lobes ex-

cept along eyes and darker on anterior part of supra-

antennal plates; posterior margin with pairs of black

spots close to eyes and posterior of eyes; ocelli enclosed

in brownish to black marking, anterior margin of

marking shallowly concave and generally just not

reaching frontoclypeal suture, lateral margins of mark-

ing either weakly convex or straight, and convergent

posteriorly, posterior margin deeply and narrowly con-

cave medially and (almost) reaching anterior margin of

pronotum. Genae ochraceous, lori darkened along su-

ture with clypeus. Frontoclypeal suture trapezoid, me-

dian part only little broader than distance between lat-

eral margins of lateral ocelli. Rostrum brownish, dark-

ened at extreme tip only; reaching about halfway

between to posterior margin hind coxae.

Thorax. — Pronotum distinctly broader than head.

Pronotal disc brownish but often with greenish tinge;

disc immediately posterior of anterior margin often

with dark brown to black, medially narrowly inter-

rupted transverse band; central fasciae distinct and

meeting on pronotal suture, sometimes continuing

on anterior margin, fasciae broader near anterior and

posterior margin of disc, narrow in between and often

narrowly interrupted; anterior oblique fissure nar-

rowly darkened; transverse part of pronotal suture

with narrow to very narrow dark brown to black band

that is often interrupted just lateral of central fasciae,

band not continuing laterally across pronotal collar.

Pronotal collar concolorous with or slightly paler

than pronotal disc, posterior margin black; lateral

part of collar with darker brown marking at pronotal

suture at lateral lobe of disc; lateral margin at same

level slightly darkened; anterolateral corner with

blunt lateral tooth. Anterior margins of lateral and

paramedian lobes, fissures between these lobes as well

Beuk: The Platylomia radha group

as pronotal suture often with extensive waxy coating.

Mesonotum concolorous with pronotum; median

fascia black and rather broad, narrowest at anterior

and posterior margins of disc; paramedian fasciae on

anterior half of disc black, narrow to very narrow and

more brownish near anterior margin of disc, slightly

curved towards central fascia medially, fasciae on pos-

terior half usually present as black elongated markings

in front of and sometimes extending on to anterior

arms of cruciform elevation; anterior margin of disc

with small blackish triangular markings immediately

lateral of mesonotal fissures; lateral fasciae present

only as black spots in posterolateral corners of

mesonotal disc. Cruciform elevation concolorous

with mesonotal disc, posterior margin narrowly

black, anterior arms sometimes darkened. Mesonotal

disc often with extensive waxy coating on anterior

and lateral margins and in depressions of cruciform

elevation. Katepimeral lobe (fig. 44) almost as in P.

radha. Surface of katepimeral lobe concave along dor-

sal margin and dorsal margin thus curved outwards;

surface covered with short, waxy hairs and margin

with slightly longer hairs. Apex of katepimeral lobe

just or just not reaching over base of operculum.

Tegmina and wings. — Tegmina brownish hyaline

but especially along veins in distal and posterior part

more brownish infuscated and with indistinct brown-

ish reticulation along distal margin and sometimes in

apical cells; basal veins of second, third, fifth, and sev-

enth apical cells usually distinctly infuscated; apices of

longitudinal veins of apical cells with indistinct dark

brown spots; basal cell yellowish brown fumose ex-

cept along posterior margin. Veins of tegmen brown-

ish, darker in distal and posterior part. Wings brown-

ish hyaline but paler than tegmina. Veins of wings

dark brown, medial and cubital veins paler.

Legs. — Fore legs brownish; femora posteroventral-

ly with darkened longitudinal stripe; tibiae posterior-

ly and ventrally slightly darkened, dorsally at base

dark brown. Mid legs brownish, only tibiae slightly

darkened dorsally at base and on apical quarter. Hind

legs brownish, tibiae dorsally darkened at base and on

apical quarter and tarsi. Fore femur with gap between

middle and distal posteroventral spines shallow and

broad. Hind tibiae with 3-5 anterodorsal spines and

4-6 anteroventral spines, occasionally with single an-

terior spine.

Male operculum (fig. 42). — Operculum reaching

about halfway fifth abdominal segment, 2.1-2.3 times

as long as maximum width distal of constriction;

brownish but distal part often somewhat darker, area

of lateroproximal corner and usually also part of later-

al margin into constriction dark brown to black but

margin just distal of lateroproximal corner ochraceous

for short distance; some parts covered with waxy coat-

ing, usually along margins. Medial margin distal of

169

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 37-44. Platylomia bivocalis (Matsumura, 1907), male, Pingtung Hsien. — 37, pygofer and uncus in ventral view; 38, py-

gofer and uncus in right lateroventral view; 39, pygofer in dorsal view; 40, uncus in right anterolateral view; 41, uncus in an-

terior view; 42, abdomen and right operculum in lateroventral view; 43, right timbal covering; 44, right katepimeral lobe.

170

constriction convex to apex but sometimes less convex

to almost straight close to apex. Apex of operculum

rounded, tip lateral of midline. Lateral margin distal of

constriction first convex but on distal third almost

straight or even weakly concave. Constriction at 0.3-

0.4 of length of operculum, lateral concavity longer

and deeper than medial concavity, broadest part of op-

erculum distal of constriction 1.4-1.6 times as wide as

minimum width in constriction. Distance between

opercula at constrictions about 1.0-1.3 times as wide

as minimum width in constriction. Opercula at point

of closest approximation separated for a distance of

0.7-0.9 times maximum width between opercula at

constrictions. Operculum close to abdomen; surface

of part distal of constriction convex.

Male abdomen. — Abdomen about 1.4 times as

long as head and thorax together. Dorsal part of ter-

gites dark castaneous but tergite 1, medial part of ter-

gite 2, sometimes anteromedial part of tergite 3 and

lateral parts of tergites 2-6 brownish; sternites and

ventral parts of tergites pale castaneous brown.

Tergite 1, anterior margin of tergite 3 except medial-

ly, tergites 5, 6 and 8 covered with white waxy coat-

ing. Posterior margin of tergite 7 with many dark

spinules except at extreme lateral part. Sternite 7 with

very shallow posteromedial emargination. Timbal

covering (fig. 43) brownish, about 1.1-1.2 times as

long as wide; medial margin straight to weakly con-

vex; mediodistal corner, distal margin and laterodistal

corner rounded; lateral margin almost straight, con-

vergent with medial margin.

Male genitalia (figs. 37-41). — Yellowish brown;

dorsal part of pygofer dark brown except on desclero-

tised part; uncus lobes darker at bases and along later-

al and dorsal margins. Basal pygofer lobes (fig. 37)

close to lateral margins of pygofer, narrow and ridge-

like, rounded in lateral view (fig. 38); anteroventral

margin of pygofer and pygofer surface enclosed by it

mostly with short and erect hairs but basal lobes and

parts of anteroventral margin leading to them with

long erect hairs; pygofer surface anterior of medial

part of anteroventral margin with scattered hairs; out-

er surface of pygofer on posterior half with numerous

short hairs interspersed with scattered hairs that in-

crease in length towards posterior margin. Dorsal part

of pygofer as in fig. 39. Basal part of uncus rather

broad, somewhat globose and with small posterome-

dial knob that may or may not be distinct (figs. 37-

38); surface near bases of uncus lobes with occasional

long and thick erect hairs. Uncus lobes (figs. 37-38,

40-41) short and gradually narrowing towards distal

margin; medial margins curved dorsad (inwards),

weakly concave near aedeagal opening and weakly

convex near mediodistal corners or more or less paral-

lel; mediodistal corner angularly rounded to rounded

and often slightly curved dorsad (inwards); distal

Beuk: The Platylomia radha group

margin slightly curved dorsad (inwards), weakly con-

cave medially and close to laterodistal corner with

short, pointed triangular projection; laterodistal cor-

ner produced into small, pointed projection; lateral

margin evenly concave on distal half. Ventral (outer)

surface of uncus lobes with short longitudinal grooves

at distal margin, especially near mediodistal corner,

and often with transverse grooves about halfway to

distal margin. Ventral (outer) surface of uncus cov-

ered with short erect hairs, these hairs more sparse on

distal part; dorsal (inner) surface with more numer-

ous and thicker hairs; bases of uncus lobes with long

and thick erect hairs both on ventral and dorsal sur-

face, those on dorsal (inner) surface somewhat short-

er than those on ventral (outer) surface.

Female operculum. — Operculum greenish to

brownish, narrowly dark at meracanthus and along

lateral margin but anterior part of lateroproximal lobe

brownish. Lateral margin weakly convex distal of lat-

eroproximal lobe but sometimes weakly concave near

laterodistal corner; laterodistal corner angular to an-

gularly rounded; distal margin straight or concave for

short distance; mediodistal corner broadly rounded;

medial margin straight, medial margins strongly di-

vergent from bases of opercula.

Female abdomen. — Abdomen 1.0-1.1 times as

long as head and thorax together. Dorsal part of ter-

gites largely blackish, more brownish near lateral mar-

gins, tergites 1-2 brownish medially but in paler spec-

imens also on tergites 3-5 and on anterior margins of

tergites 2-3; tergites 3-7 with small oblong spots lat-

erally; sternites and ventral parts of tergites brownish.

Tergites 1-3 with whitish dusting laterally, tergites 5-

6 and 8 completely covered with white waxy coating.

Posterior margins of tergites 3-6 with some short dark

spines laterally, of tergite 7 with spines along whole

length but fewer medially, of tergite 8 with scattered

spines medially. Sternite 7 with angular to angularly

rounded posteromedial emargination.

Female genitalia. — Dorsal part of pygofer blackish

but in paler specimens more brownish medially, espe-

cially at base of caudodorsal beak, remainder of py-

gofer brownish; distal part of ovipositor sheath black-

ish. Dorsal margin of pygofer just longer than two

preceding segments, in lateral view weakly concave;

ventral margin weakly convex. Ovipositor sheath

reaching little though distinctly beyond apex of cau-

dodorsal beak; anal valve distinctly not reaching as far

as caudodorsal beak.

Measurements in mm (d: n= 6; 9: n = 5). — Body

length: &: 47.5-51.5 (50.2 = 1.3), 2: 39.5-44.0

(42.0 + 1.2); head width: d : 13.3-14.5 (14.1 + 0.4),

9: 13.5-14.0 (13.8 + 0.1); maximum pronotum

width: 7 14.4-16.0. (55 =2705), 2: 14.5-15.7

(15.1 + 0.4); tegmen length: d: 50.5-57.5 (54.9 +

I VESOO-555 (G37 22 143)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 45-51. Platylomia pendleburyi Moulton, 1923, male. — 45, pygofer and uncus in ventral view, holotype; 46, pygofer and |

uncus in right lateroventral view, holotype; 47, pygofer in dorsal view, Khao Luang; 48, uncus in anterior view, Khao Luang; |

49, abdomen and right operculum in lateroventral view, Khao Luang; 50, right timbal covering, Khao Luang; 51, right |

katepimeral lobe, Khao Luang.

172

Biological notes

Matsumura (1907) described the song of P. bivo-

calis as consisting of two calls, ‘sha sha’ and ‘ho ho’,

that are called at the same time. He also stated that

the adults often sit under Pandanus bushes.

Distribution (fig. 3)

Taiwan. The record from Japan is a misinterpreta-

tion by Metcalf (1963) of data provided by Esaki

(1932) and Kato (e.g. 1932, 1933b).

Remarks

The type material of P. bivocalis was not examined.

The original description (Matsumura 1907) and sub-

sequently published illustrations (Matsumura 1931;

Esaki 1932; Kato 1932) leave no doubt that it is iden-

tical with the species described and illustrated here.

Material examined. — Tarwan: Changlo [Ta-Kung-lu],

Pintung Hsien [Ping-tung Hsien], 16.vi.1976, H.

Makihara, 1 d, 1 2 (suu); Kankan [Kang-kang], vi.1909,

H. Sauter, 1 d, 1 9 (BMNH); same data, 1 d (USNM);

Kankau [Kang-k’ou] (Koshun), iv.1912, H. Sauter, 1 à

(USNM); O-Luan-Bi [O-luan-pi], 8.vii.1973, M. Hayashi, 1

d (suu); same data, 9.vii.1973, 2 d, 2 2 (suu); Ouluanpi

[O-luan-pil, Pingtung Hsien [P’ing-tung Hsien],

26.v.1982, K.C. Chou & C.C. Pan, 1 d (rari); Ouluanpi

[O-luan-pil Coast Forest, Pingtung Hsien [P’ing-tung

Hsien], 27.vi.1981, T. Lin & C.C. Pan, 3 6, 6 $ (rar);

same data, 2 d, 2 2 (zman); S. Formosa, xi.1893, J. De La

Touche, 2 2 (BMNH).

Platylomia pendleburyi Moulton, 1923

(figs. 3, 45-51)

Platylomia pendleburyi Moulton, 1923: 69, 98, 100, 103,

167, pl. V figs. 22-22b. — Holotype d : ‘Type’ [printed on

round label with red margin], “Peninsular Siam, / Nakon

Sri Tamarat / Khao Luang / 3300 FT. / April 1* 1922. /

H.M. Pendlebury’ [printed except for altitude and date],

‘Platylomia pendleburyi / Type d Moulton / 1922’

[Moulton’s handwriting]( BMNH) [examined].

Platylomia pendleburyi — Metcalf 1963: 623.

P. pendleburyi is the only species of the P. radha

group confirmed from Peninsular Thailand and is as

yet restricted to that area. P. pendleburyi, P. radha,

and P. ficulnea are the largest species of the P. radha

group. P. pendleburyi can easily be distinguished from

P. radha by the presence of distinct markings on the

tegmina, and from P. ficulnea by the rounded and

shorter opercula.

Description

Body brownish, medial part of tergites more casta-

neous; pronotum with small (sometimes split) medi-

an spot only; mesonotum with four black markings

on posterior margin; tegmina with markings on basal

veins of second, third, fifth, and seventh apical cells,

Beuk: The Platylomia radha group

and at apices of longitudinal veins. Opercula broad

and not very long, divergent from abdomen.

Head. — Postclypeus and anteclypeus brownish,

dorsal part of postclypeus slightly darker except for

small area at frontoclypeal suture; postclypeus little

swollen, in dorsal view about as long as distance be-

tween frontoclypeal suture and anterior margin of

pronotum. Vertex brownish but darker on vertex

lobes, on lateral part of supra-antennal plates and pos-

terior of eyes; area between ocelli blackish. Genae and

lori brownish but sometimes with greenish tinge, up-

per part of genae slightly darker. Frontoclypeal suture

and rostrum as in P. bivocalis.

Thorax. — Pronotum slightly to distinctly broader

than head. Pronotal disc brownish but sometimes

with greenish tinge, only marking consisting of small

(sometimes split) medial spot at pronotal suture.

Pronotal collar slightly paler than pronotal disc,

sometimes with greenish tinge, posterior margin

black; lateral part of collar with indistinct brown

marking close to lateral margin of collar; anterolater-

al corner with pointed lateral tooth at level of broad-

est point of pronotal disc; posterolateral corner

rounded. Mesonotum slightly darker than pronotal

disc; paramedian fasciae on anterior half of disc

brownish to blackish and narrow, especially at anteri-

or margin of mesonotum, on posterior half of disc

present as two black markings in front of anterior

arms of cruciform elevation; anterior margin of disc

either with or without triangular markings just lateral

of mesonotal fissures; lateral fasciae very indistinct or

present only as black spots near posterior margin of

mesonotal disc. Medial part of cruciform elevation

concolorous with pronotal collar, lateral part slightly

darker. Katepimeral lobe (fig. 51) shorter than broad

at base; apex angularly rounded. Surface of katepi-

meral lobe almost flat, only weakly concave along

dorsal margin; surface covered with short, waxy hairs

and margin with slightly longer hairs. Apex of katepi-

meral lobe just reaching over base of operculum.

Tegmina and wings. — Tegmina faintly yellowish

hyaline with a brownish reticulate pattern between

veins in distal and posterior part of the tegmina; basal

veins of second and third apical cells with narrow

brownish clouding along whole length, fifth and sev-

enth apical cells and apices of longitudinal veins of

apical cells with rounded brown spots; basal cell yel-

lowish brown fumose except along posterior margin.

Veins of tegmen brownish. Wings very faintly yellow-

ish hyaline, very narrowly clouded posterior of first

anal vein. Veins of wings brown, darker on basal part

of medial vein and on veins close to wing margin.

Legs. — Fore femora brownish, slightly darker on

posteroventral longitudinal stripe and on posterior

surface near apex; tibiae dark brown but paler on dor-

sal surface of proximal three quarters; tarsi very dark

173

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

brown. Mid legs brownish; tibiae with blackish basal

ring, dorsal surface dark brown on apical three quar-

ters and ventral surface dark brown on apical two

thirds; tarsi very dark brown. Hind legs brownish;

femora ventrally darker at apex; tibiae with blackish

basal ring. Fore femur with gap between middle and

distal posteroventral spines deeper and narrower than

in P. bivocalis.

Male operculum (fig. 49). — Operculum reaching

about halfway fifth abdominal segment, 2.6-2.9 times

as long as maximum width distal of constriction;

brownish but sometimes with greenish tinge, distal

part sometimes darker, area of lateroproximal corner

and lateral margin proximal of constriction dark

brown to black and enclosing brownish spot on lat-

eroproximal lobe. Medial margin distal of constric-

tion weakly convex to apex, medial margins almost

parallel for some distance. Apex of operculum obtuse-

ly rounded. Lateral margin distal of constriction first

convex but partly straight on distal third. Constric-

tion at about 0.4 of length of operculum, medial con-

cavity very shallow; broadest part of operculum distal

of constriction 1.3-1.4 times as wide as minimum

width in constriction. Distance between opercula at

narrowest part of opercula about 1.5-1.9 times as

wide as minimum width of opercula. Opercula at

maximum width of distal part of opercula separated

for a distance of about 0.9-1.0 times distance between

opercula at narrowest part of opercula. Operculum

close to abdomen proximal of constriction only and

somewhat divergent from abdomen from level of

constriction onwards; surface of part distal of con-

striction convex and in addition curved towards ab-

domen laterally.

Male abdomen. — Abdomen about 1.3-1.4 times as

long as head and thorax together. Dorsal part of ter-

gites dark castaneous, medially and laterally more

brownish; tergites 2-7 laterally with darker brown

spots near posterior margin; sternites and ventral

parts of tergites brownish, slightly darker on posterior

segments. Tergite 8 with thin whitish dusting, dust-

ing on other tergites without distinct pattern. Poster-

ior margin of tergites 5-6 sometimes with few small

dark spinules laterally, of tergite 7 with dark spines

along whole length. Sternite 7 with very shallow

rounded posteromedial emargination. Timbal cover-

ing (fig. 50) brownish but sometimes with greenish

tinge, 1.1-1.2 times as long as wide; medial margin

straight; distal margin and laterodistal corner round-

ed; lateral margin weakly convex, convergent with

medial margin.

Male genitalia (figs. 45-48). — Brownish; dorsal

part of pygofer somewhat darker; uncus more casta-

neous at basal part and bases of uncus lobes. Basal py-

gofer lobes (fig. 45) and hairs on pygofer as in P. bivo-

calis, but basal lobes narrower. Dorsal part of pygofer

174

as in fig. 47. Basal part of uncus short, narrower and

less globose than in P. bivocalis (fig. 45-46), near

bases of uncus lobes with occasional long and thick

erect hairs. Uncus lobes (figs. 45-46, 48) short and

gradually narrowing towards distal margin; medial

margins curved dorsad (inwards), straight to concave;

mediodistal corner smoothly rounded; distal margin

straight but close to laterodistal corner with short,

blunt triangular projection; laterodistal corner round-

ed but continuing into projection on distal margin;

lateral margin weakly concave for most of its length

but convex at base and near laterodistal corner. Ven-

tral (outer) surface of uncus lobes almost void of

grooves near mediodistal and laterodistal corners.

Hairs on surface of uncus lobes as in P. bivocalis but

less dense.

Female operculum. — Operculum brownish, dark-

ened in lateroproximal corner and along lateral mar-

gin. Lateral margin distal of lateroproximal lobe first

weakly convex but weakly concave near laterodistal

corner; laterodistal corner angularly rounded; distal

margin weakly convex; mediodistal corner rounded;

medial margins short, straight, and divergent.

Female abdomen. — Abdomen about as long as

head and thorax together. Dorsal part of tergites cas-

taneous, laterally slightly paler, tergites 3-6 with small

oblong spots laterally that almost form line; sternites

and ventral parts of tergites brownish. Tergites with

thin whitish dusting but without distinct pattern.

Posterior margins of tergites 3-4 with few short dark

spines laterally, of tergite 5 with short dark spines ex-

cept medially, of tergites 6-7 with spines along whole

length but fewer medially on tergite 6, of tergite 8

with few short spines medially. Sternite 7 with angu-

lar posteromedial emargination in median lobe.

Female genitalia. — Dorsal part of pygofer dark cas-

taneous but more brownish medially and just anteri-

or of caudodorsal beak, remainder of pygofer brown;

caudodorsal beak and distal part of ovipositor sheath

dark castaneous. Dorsal margin of pygofer about as

long as four preceding segments, in lateral view weak-

ly concave; ventral margin somewhat more convex.

Ovipositor sheath reaching distinctly beyond apex of

caudodorsal beak; anal valve reaching about halfway

to apex of caudodorsal beak.

Measurements in mm (d:n = 3; 9: n= 1). — Body

length: tdi: 5100-5515 63.2 21.6), 2: 48:0; head

width: 6: 16.0-16.1 (16.1 + 0.1), 2: 16.2; maximum |

pronotum width: d: 16.2-17.6 (16.9 # 0.5), 2: 17.0; |

tegmen length: à : 63.0-64.5 (64.2 + 0.9), 2: 66.5.

Distribution (fig. 3) |

P. pendleburyi is only known from'the type locality |

on Peninsular Thailand. The records from Banguey |

Island and Malay ‘Archipelago’ by Metcalf (1963) are |

misinterpretations of comments by Moulton (1923). |

Remarks

The collection date on the holotype labels was mis-

read as April 14th 1922 and cited as such by Moulton

(1923: 100). No other material was mentioned ex-

plicitly in the description so that the other two speci-

mens cannot be considered as paratypes, even though

the female now examined was labelled as paratype

and a female is mentioned in the description.

Material examined. — THAILAND: Khao Luang, Nakon Sri

Tamarat [Nakhon Si Thammarat}, 3500-4000 ft,

13.11.1922, H.M. Pendlebury, 1 9 (BMNH); same data, 1 d

(MNKM); same data, 3300 ft, 1.iv.1922, H.M. Pendlebury, 1

3 holotype Platylomia pendleburyi Moulton (BMNH); same

data, 2000 ft, 2.iv.1922, H.M. Pendlebury, 1 d (BMNH).

ÄCKNOWLEDGEMENTS

This study would not have been possible without

the help of curators in several institutes: Mr M. D.

Webb and Mrs J. Margerison-Knight (BMNH), Dr G.

A. Samuelson and Ms K. Kami (BrBM), Dr P. H.

Arnaud (casc), Dr Lei Zhongren and Dr Zhang

Yalin (Foyp and Nwas), Dr P. Grootaert and Mr R.

Detry (isnB), Dr W. D. Schedl (1zu1), Dr P. D. Per-

kins (mcz), Dr H. J. Zainal Abidin Jamaluddin

(MNKM), Dr M. Boulard and Dr T. Bourgoin

(MNHN), Dr V. Raineri (msn), Mr P. Lindskog

(nHms), Dr. M. Tomokuni (NsMT), Mr. J. van Tol

(RMNH), Dr V. P. Gapud and Dr R. W. Brooks

(SEMK), Dr. M. Hayashi (suu and opw), Dr S. J. Fang

(rari), Dr R. C. Froeschner and Dr M. Epstein

(usnm), Dr N. Moller Andersen (UZMK) and Dr H.

Lampe (zeMK). Dr W. Schedl (1zu1) kindly allowed

me to retain some specimens for our own collection.

I am indebted to Dr. J. P. Duffels (Institute of

Systematics and Population Biology, University of

Amsterdam) for his critical reading of the manuscript.

Mr G. Verlaan prepared specimens after the genitalia

were dissected. The investigations were supported by

the Life Sciences Foundation (SLW), which is sub-

sidised by the Netherlands Organisation for Scientific

Research.

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Received: 30 November 1996

Accepted: 15 May 1997

KJELL ARNE JOHANSON

Museum of Zoology, University of Bergen, Norway.

HELICOPSYCHE MALICKYI SP N. FROM BORNEO

WIRICHORMER:MEELICORLSTEHIDNE)

Johanson, K. A., 1998. Helicopsyche malickyi sp. n. from Borneo (Trichoptera, Helicopsychi-

dae). — Tijdschrift voor Entomologie 140 [1997]: 177-180, figs. 1-10. [1ssn 0040-7496].

Published 26 March 1998.

Helicopsyche malickyi sp. n. (Trichoptera: Helicopsychidae) is described from Sarawak,

Malaysia. This is the first description of a species of Helicopsychidae from Borneo, and the

third species from the Malesian Archipelago.

Correspondence: Kjell Arne Johanson, Museum of Zoology, University of Bergen, Muséplass

3, N-5007 Bergen, Norway. E-mail: [email protected].

Keywords. — Helicopsyche malickyi sp.n.; Helicopsychidae; Trichoptera; Malaysia; Borneo.

With the description of Helicopsyche malickyi sp.n.,

the Helicopsychidae now includes 173 extant de-

scribed species (Johanson 1995, 1997a, Malicky

1994, Malicky and Chantaramongkol 1993) and

forms a moderately large group of caddis flies. The

highest species numbers are found in Antillean, Indo-

Chinean and Ceylonese subregions (Johanson

1997b). The family is mainly confined to tropical and

subtropical areas and the larvae seems to prefer stream

habitats. The Helicopsychidae species are recognized

by having larvae which build dextrally coiled cases

made from sand grains glued together by silk. The

adults can be distinguished from species in other fam-

ilies by the presence of an abdominal VIth sternal

process in most species, and the absence of nigma in

both fore and hind wing fork 2.

The Indo-Malayan Helicopsychidae fauna com-

prises six species: Cochliopsyche angusta (Ulmer,

1951); Helicopsyche boniata Malicky & Chantara-

mongkol, 1992; H. lata Ulmer, 1951; A. puttula

Malicky & Chantaramongkol, 1992; H. anaksaku

Malicky, 1995 and H. anaktangga Malicky, 1995.

However, only C. angusta and H. lata are previously

described from the Malesian Archipelago (Ulmer

1951, 1955, 1957) (fig. 1) and species records indi-

cated absence of Helicopsychidae from Sarawak

(Kimmins 1955). The southeast Asian Helicopsy-

chidae fauna is considered richer than indicated by

known species number and Huisman (1991) added

to the knowledge two undescribed Helicopsyche spe-

cies from Borneo (Sabah) (fig. 1), which remain yet to

be described. The present record from Sarawak thus

adds valuable information to the understanding of

the biogeographic history of Helicopsychidae in

southeast Asia.

Helicopsyche malickyi sp. n.

(figs 2-10)

Type material. — Holotype 6, Malaysia, Sarawak,

Ist Div., Semongoh Forest Res., 1°25’N, 110°17’E,

15-19. XI. 1976, Malaise trap over stream [P. S.

Cranston]. Natural History Museum, B. M. 1977-

19. In Canada Balsam.

Description

With the two-jointed maxillary palp H. malickyi

sp. n. is easily distinguished from the two other Heli-

copsychidae from the Malesian Archipelago: C. an-

gustaand H. lata which have four and three joints, re-

spectively. The spherical interantennal warts, distally

reduced Cu, in the fore wing and the pointed abdom-

inal VIth sternal process in H. malickyi sp.n. indicate

a close relation to other Oriental and Palearctic Heli-

copsychidae. H. malickyi sp.n. is similar to the north-

ern Thailand Helicopsyche namtok Malicky, 1993, but

is easily separated by the IXth segment of the genitalia

which is pointed dorsally and ventrally; by the prima-

ry branch of the gonocoxite which is larger and paral-

lel-sided; by the bifurcated secondary branch of gono-

coxite; and Xth tergum which is slightly shorter. Both

scape and maxillary palp are short. In most

177

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 1.

The distribution of Helicopsy-

chidae in southeast Asia, New

Guinea and northern Australia,

with Indonesia, Malesian is-

lands and Philippines empha-

sized in grey. The Helicopsyche

species of the Malesian islands

are indicated by separate sym-

bols. Other species are mapped

as Stars.

Helicopsychidae the vertex is rounded in lateral view

and is only slightly visible in lateral view. H. malickyi

sp.n. is separated from other Helicopsyche by the

slightly dorsally produced vertex; Xth tergum of the

genitalia which is slender in lateral view, gently

curved ventrally and with two pairs apical setae; and

the posterior margin of the gonocoxal primary branch

which is highly undulated, and with two pointed pos-

tero-ventral processes.

Description

Head (fig. 2). — With short scape, about twice the

length of the pedicel. Maxillary palp short, the basal

joint about as long as scape and distal joint. Eye large

in lateral view, covering large parts of head. Cephalic

warts rounded in dorsal view and separated by the

postero-dorsad produced vertex. Postantennal warts

situated between scapes, spherical. Pronotum with

two pairs setal warts. Mesonotum and metascutellum

with one pair setal warts. Spur formula 1,2,4.

Wings (fig. 3). Fore wing length 3.0 mm, apical-

ly pointed. Fork 1 originates near median part of Dc,

broad. Fork 2 about as long as fork 1. Crossvein R-M

about 1.5X longer than basal part of R. Fork 3 pre-

sent, with stalk about half as long as M,., and half M,

length. Forks 4 and 5 absent. Crossvein between Cu,

and Cu, short. A,, meets posterior wing margin at

about one third the wing length. Hind wing length

2.4 mm, with 13 hamuli. All forks absent. M,., about

half the R,., length. Cu, and Cu, run parallel towards

wing margin. A, and A, diverge towards wing margin.

Abdomen. — With sternites II-V well reticulated.

VI th sternal process (figs 4 and 5) long, situated pos-

teriorly and with rounded apex.

Genitalia (figs 6-10). — IXth segment trianguloid in

lateral view, with dorsal and ventral parts slightly

pointed (fig. 6); anterior marginal and short ventral

178

Cochliophylax angusta (Ulmer, 1951)

Helicopsyche lata Ulmer, 1951

unidentified Helicopsyche

Helicopsyche malickyi sp.n.

Helicopsychidae from outside

the Malesian Archipelago

longitudinal apodeme present. The dorsal apodeme

continues into a longitudinal central apodeme run-

ning into Xth tergum. Anterior part of IXth segment

deeply concave in dorsal and ventral view (figs 7 and

8). Posterior part of IXth sternite extends into a short

bi-lobed process (fig. 8). Superior appendage situated

laterally, long, club shaped and covered by long setae.

Gonocoxite, in lateral view (fig. 6), large, with large

dorsal primary branch and ventral, bifurcated, poste-

riorly pointed ventral branch. Primary branch is par-

allel-sided, with rounded apex, smooth anterior and

undulated posterior margins; undulations are associ-

ated with short setae. Ventral branch divides into two

slightly dorsally curved and pointed processes; in ven-

tral view broad and covered by setae. Xth tergum, in

lateral view (fig. 6), slender, parallel-sided and slight-

ly curved ventrad; in dorsal view with convex lateral

margins, apex divided by a deep, trianguloid central

cleft and two minor lateral clefts, two pairs short setae

present near apex. Phallus, in lateral view (fig. 9),

slender, with posterior part smoothly curved ventrad,

in dorsal view (fig. 10), with basal half slender and

distal part strongly expanded laterally, about 3x

broader than the median part.

Etymology

Malickyi, named after Dr. Hans Malicky in recog-

nition of his contribution to the understanding of the

Oriental Trichoptera fauna.

BIOGEOGRAPHICAL NOTES

By not listing any taxa of Helicopsychidae, some |

authors (Kimmins 1955, Malicky 1995) indicate that

the number of Helicopsychidae species in the Male- |

sian Archipelago is low. Thus, when considering the |

Helicopsychidae, the Malesian Archipelago may con- |

JOHANSON: Helicopsyche malickyi sp. n

Cu, Cu; M344

Figs 2-10. Helicopsyche malickyi sp.n. — 2, Head, lateral view; 3, Right wings; 4, VIth sternal process, lateral view; 5, VIth ster-

nal process, ventral view; 6, Genitalia, lateral view; 7, Male genitalia, dorsal view; 8, Genitalia, ventral view; 9, Phallus, later-

al view; 10, Phallus, dorsal view.

179

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

stitute an actual low-diversity area. However, as is the

case in other areas (Johanson 1993, Johanson 1997b,

Schmid 1993), the family appears to be disjunctively

distributed and is expected to be diverse in certain lo-

calities. This may be the case for the Helicopsychidae

of the Malesian Archipelago as well and is supported

by the findings of two more species (Huisman 1991).

Thus, it might be expected that more species will ap-

pear from that area in the future.

ACKNOWLEDGEMENT

Dr. Hans Malicky gave valuable information on

Helicopsyche namtok Malicky, 1993.

REFERENCES

Huisman, J., 1991. A study of Trichoptera in Sabah and

Sarawak. — Proceedings of the 6th International Sym-

posium on Trichoptera. Adam Mickiewicz University

Press, Poznan. Serie Zoologie: 275- 278.

Johanson, K. A., 1993. New Helicopsyche von Siebold, 1856

(Trichoptera: Helicopsychidae) from East Africa. —

Annales de limnologie 29 (2): 139-155.

Johanson, K. A., 1995. A catalog of the Helicopsychidae

(Insecta: Trichoptera) of the World. — Bulletin Zoölo-

gisch Museum, Universiteit van Amsterdam 14 (7): 101-

123.

Johanson, K. A., 1997a. Description of Helicopsyche giboni

180

sp.n. from Madagascar (Trichoptera: Helicopsychidae). —

Braueria 24: 5-6.

Johanson, K. A., 1997b. Zoogeography and diversity of the

snail case caddisflies (Trichoptera: Helicopsychidae). —

Pp. 205-212 in Holzenthal, R.W. and O.S.Flint Jr. (eds).

Proceedings of the 8th International Symposium on

Trichoptera. Ohio Biological Survey, Columbia.

Kimmins, D. E., 1955. Results of the Oxford University

Expedition to Sarawak, 1932. — Sarawak Museum

Journal 6 (5): 374-442.

Malicky, H., 1994. Zwei neue Helicopsyche (Helicopsy-

chidae) aus Perak, Malaysia. — Braueria 22: 4.

Malicky, H., 1995. A preliminary list of Trichoptera from

Bali. — Braueria 22: 4.

Malicky, H. & P. Chantaramongkol, 1993. Neue Tricho-

pteren aus Thailand. Teil 2: Rhyacophilidae, Philopota-

midae, Polycentropodidae, Ecnomidae, Psychomyidae,

Xiphocentronidae, Helicopsychidae, Odontoceridae. —

Linzer biologische Beiträge 25: 1137-1187.

Ulmer, G., 1951. Köcherfliegen (Trichopteren) von den

Sunda Inseln (Teil I). — Archiv für Hydrobiologie,

Supplement 19: 1-528.

Ulmer, G., 1955. Köcherfliegen (Trichopteren) von den

Sunda Inseln (Teil I). — Archiv für Hydrobiologie,

Supplement 21: 408-608.

Ulmer, G., 1957. Köcherfliegen (Trichopteren) von den

Sunda Inseln (Teil II). — Archiv für Hydrobiologie,

Supplement 23: 109-470.

Received: 23 May 1997

Accepted: 20 October 1997

Q. LI & M. XUE

Shandong Agricultural University, China

A NEW SPECIES OF THE GENUS AMMOPHILA KIRBY

(HYMENOPTERA, SPHECIDAE) FROM CHINA

Li, Q. & M. Xue, 1998. A new species of the genus Ammophila Kirby (Hymenoptera,

Sphecidae) from China. — Tijdschrift voor Entomologie 140 [1997]: 181-183, figs. 1-9. [Issn

0040-7496]. Published 26 March 1998.

Ammophila heteroclypeola sp. n. is described from Xingjiang, China. It is closely related to A.

clypeola Yang et Li.

Correspondence: Li Qiang, Department of Plant Protection, Shandong Agricultural Uni-

versity, Taian, Shandong, 271018, P. R. China.

Key words. — Hymenoptera; Sphecidae; Ammophila; new species; China.

Ammophila is the largest genus in the subfamily

Sphecinae. It includes 204 medium to large size

predatory solitary wasps. Kohl (1906) revised the

Palaearctic species, and his work is still a valuable ref-

erence for the species identifications. Roth (1928)

keyed the species of North Africa. Beaumont (1963)

keyed the species of the nasuta group. Tsuneki (1967,

1971a, 1971b, 1971c, 1976) provided informative

references and keys for the identification of east Asian

forms. Bohart & Menke (1976) revised the genera of

Sphecidae of the world, and listed 187 species of

Ammophila. Wu & Zhou (1996) keyed the Chinese

species.

In the long course of a study on the fauna of Sphe-

cinae from China, we published some descriptive pa-

pers (Li et al. 1994; Li & Yang 1989, 1990; Yang &

Li 1989). While investigating the second part of the

Sphecinae material deposited in the insect collections

of the Zhejiang Agricultural University (rczau) and

the Institute of Zoology, Academia Sinica (ICIZAS),

one new species of Ammophila was discovered. We

follow Bohart and Menke (1976) in the usage of mor-

phological terms. The abbreviations used in the text

are as follows.

Pop Postocellar distance.

oop ocellocular distance.

IODP interocular distance at anterior ocellus.

IODc interocular distance at clypeus.

Ammophila heteroclypeola sp. n.

(figs. 1-9)

Type material. — Holotype female, China,

Xingjiang, Wujiagu, 15.vi.1982, C. Hong, ICZAU.

Paratypes (2): 1 female, China, Xingjiang, Yanzhei,

950-1170 m, 6.vii.1958, C. Li, ıcızas; 1 male,

China, Xingjiang, Wusu, 420-460 m, 25.vi.1957, C.

Hong, ICIZAS.

Description

Female. — Body length 14.8-15.1 mm. Black; ante-

rior half of tegulae, abdominal petiole, tergites I-IV

and sternites II-IV wholly, tergite V and sternite V

partly yellowish red; tibia and tarsi of legs, mandibles

and antennae brown to dark brown; apical black por-

tion of abdomen without steel blue luster; wings pale

yellowish brown, veins brown to dark brown. Upper

clypeus and lower frons areas, pronotal lobe, prepec-

tus, mesopleuron and posterior sides of propodeum

covered with dense appressed silver hair; gena, pro-

thorax except collar above, mesosternum, metapleu-

ron, metasternum, propodeum except propodeal en-

closure, coxae and femora of legs with sparse

appressed silver hair; head and thorax with long,

erect, white hair.

Clypeus densely punctate, disc slightly bulging, an-

terior margin with two lateral teeth (fig. 1). Frons

densely punctate, without supra-antennal projection.

Vertex sparsely punctate. POD : OOD : IODP : IODC =

181

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

C S

Figs. 1-9. Ammophila heteroclypeola sp. n. — 1-5, Female; 6-9, Male. 1, Head, frontal view; 2, Antenna; 3, Prothorax, lateral

view; 4, Tarsi I-II of fore leg; 5, Forewing; 6, Head, frontal view; 7, Gonostyle, lateral view; 8, Volsella, ventral view; 9, Penis

valves, lateral view. — Scale line (mm) for figs. 1, 3: 1.16; 2: 1.00; 4: 0.80; 5, 6: 1.34; 7, 8,9: 0.32.

22 : 30 : 96 : 72. Relative length of antennal pedicel :

flacellomere GAVIN xe aI SrA 305287

Is 117%

Pronotal collar (fig. 3) length : width = 34 : 90,

shiny, sparsely punctate, with median furrow.

Scutum shiny, sparsely punctate, sides of anterior half

and medial region of posterior half irregularly, trans-

versely or obliquely rugose-striate. Scutellum shiny,

longitudinally rugose-striate and punctate. Metano-

tum shiny, densely punctate. Mesopleuron with epis-

ternal sulcus, other sculptures covered by dense ap-

pressed silver hair. Metapleuron coarsely, irregularly

rugose-striate. Propodeal enclosure with a median ca-

rina, sides of it irregularly obliquely rugose-striate;

lateral area of propodeum coarsely, irregularly rugose-

striate.

Submarginal cell III of forewing with petiole (fig.

5). Fore legs with foretarsal rakes (fig. 4); hind leg,

relative length of 1st tarsomere : 2nd : 3rd = 115 : 65

>23:

Length of abdominal petiole : tergite I : II = 145 :

125 : 108.

182

Male. — Body length 15 mm. Similar to female.

Tergites I-III and sternites I-III wholly, Tergite IV

and sternite IV partly yellowish red; mandibles, an-

tennae, wing tegulae, legs, abdomen except yellowish

red area dark reddish brown; veins yellowish brown

to brown. Anterior margin of clypeus without tooth

(fig. 6). POD : OOD : IODP : IODc = 21 : 30 : 96 : 41.

Relative length of antennal pedicel : flagellomere I: II

: III: VII: IX = 14 : 46 : 28 : 28 : 17 : 15. Pronotal

collar length : width = 40 : 86, Scutum coarsely,

densely and obliquely rugose-striate. Fore leg without

foretarsal rake. Hind leg, relative length of 1st tar-

somere : 2nd : 3rd = 110 : 59 : 58. Length of abdom-

inal petiole : tergite I : Il = 164: 144 : 102. Genitalia:

gonostyle (fig. 7), volsella (fig. 8) and penis valves

(fig. 9).

Etymology

The name is derived from Greek betero (=different)

and clypeola (also derived from Greek), the name of

its closest relative.

Remarks

This species is closely related to A. clypeola Yang et

Li, 1990. It can be easily distinguished from the latter

by the clypeal disc being only slightly bulging, the

sculpture and punctures of the scutum, metapleuron

and propodeum, the coloration of the body, ap-

pressed silver hair on clypeus, frons, mesopleuron,

metapleuron and lateral area of propodeum, and by

the male genitalia.

ACKNOWLEDGMENTS

We are grateful to Professor Jinhua He (Zhejiang

Agricultural University, Hanzhou) and Professor

Yan-Ru Wu (Institute of Zoology, Academia Sinica,

Beijing) for providing us with specimens deposited in

the insect collections under their care.

REFERENCES

Beaumont, J. de, 1963. Les Ammophiles palearctiques du

groupe de nasuta (Hym. Sphecid.). — Revue Suisse de

Zoologie 70: 1-24.

Bohart, R. M. & A. S. Menke, 1976. Specid Wasps of the

World, a Generic Revision. — University of California

Press, Berkeley, 695 pp.

Kohl, F. F., 1906. Die Hymenopterengruppe der

Sphecinen. III. Monographie der Gattung Ammophila

W. Kirby. Abteilung A. Die Ammophilinen der paläark-

tischen Region. — Annalen des Naturhistorischen

Museums in Wien 21: 228-382.

Li, Q., J. Li, & C. Yang, 1994. A classification study on

Ammophila Kirby from Shandong with description of a

new species (Hymenoptera: Sphecidae). — Journal of

Shandong Agricultural University 25(3): 289-292.

Li & XUE: New Ammophila from China

Li, Q. & C. Yang, 1989. Three new species of Sphecinae in

China (Hymenoptera: Sphecidae). — Journal of

Shandong Agricultural University 20(4): 33-37.

Li, Q. & C. Yang, 1990. Four new species of Ammophila

(Hymenoptera: Sphecidae) from Nei Mongol. —

Entomotaxonomia 12: 259-266.

Roth, P., 1928. Les Ammophila de l'Afrique de Nord. —

Annales de la Société Entomologique de France 97: 153-

240.

Tsuneki, K., 1967. Studies on the Formosan Sphecidae

(III), the subfamily Sphecinae with special reference to

the genus Ammophila in Eastern Asia (Hymenoptera). —

Etizenia 26: 1-24.

Tsuneki, K., 197la. Ergebnisse der zoologischen

Forschungen von Dr. Z. Kaszab in der Mongolei.

Sphecidae I-II. — Acta Zoologica Academiae Scientiarum

Hungaricae 17: 139-217.

Tsuneki, K., 1971b. Studies on the Formosan Sphecidae

(IX), a supplement to the subfamily Sphecinae (Hym.). —

Etizenia 53: 1-7.

Tsuneki, K., 1971c. Spheciden aus der Inneren Mongolei

und dem Nordlichen China. — Etizenia 58: 1-38.

Tsuneki, K., 1976. A fourth contribution to the knowledge

of Sphecidae (Hymenoptera) of Manchuria, with remarks

on some species of the adjacent regions. — Kontyu, Tokyo

44 (3): 288-310.

Wu, Y. & Q. Zhou, 1996. Economic insect fauna of China,

Fasc. 52, Hymenoptera: Sphecidae. — Science Press,

Beijing, 197 pp.

Yang, C. & Q. Li, 1989. Four new species of the genus

Ammophila Kirby from Shanxi Province (Hymenoptera:

Shecidae). — Entomotaxonomia 11 (1-2): 105-110.

Received: 30 March 1997

Accepted: 16 June 1997

183

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

184

INSLUrGIENEZE ENG

Nankai University, Tianjin

IDENTMINOFSOMEZYXGUSESRECIESDESCRIBED

FROM TAIWAN BY B. POPPIUS

(HETEROPTERA: MIRIDAE)

Lu, N.& L. Y. Zheng, 1998. Identity of some ‘Lygus species described from Taiwan by B.

Poppius (Heteroptera: Miridae). — Tijdschrift voor Entomologie 140 [1997]: 185-189, figs. 1-

16. [1ssN 0040-7496]. Published 26 March 1998.

Type specimens of seven ‘Lygus species, described by B. Poppius in 1912-1915 from Taiwan

(=Formosa), are studied. The following new combinations are established: Apolygus eous

(Poppius), Apolygus fuhoshoensis (Poppius), Apolygus kosempoensis (Poppius), Lygocoris

(Neolygus) bipuncticollis (Poppius), Lygocoris (Neolygus) v-nigrum (Poppius). Lectotypes for

Lygus disciger Poppius, Lygus kosempoensis Poppius, and Lygus sauteri Poppius are designated,

and some additional descriptive data are provided.

Correspondence: Le-Yi Zheng, Department of Biology, Nankai University, Tianjin 300071,

P. R. China. E-mail: [email protected].

Key words. — Poppius; Taiwan; Miridae; ‘Lygus; lectotype designations; new combinations.

B. Poppius (1912, 1915) described a number of

Miridae from the Indo-Pacific region under the all-

embracing generic name ‘Lygus (Carvalho 1959,

Schuh 1995). The type specimens of a part of these

species, preserved in the Hungarian National Natural

History Museum, have been examined and briefly

noted by Carvalho (1980). Recently, we examined

some type specimens of Poppius' ‘Lygus species, col-

lected from Taiwan (=Formosa), which are deposited

in the ‘Deutsche Entomologische Institut, Ebers-

walde (per) and the Zoological Museum of the

University of Helsinki (zuur). The results are here

reported, some descriptive data additional to the orig-

inal descriptions are given, lectotypes are designated,

and new combinations established. Measurements are

given in millimetres. The terminology for vesical

structures follows Yasunaga (1991).

Apolygus kosempoensis (Poppius, 1915) comb. n.

(figs. 1-5)

Lygus kosempoensis Poppius, 1915: 26. — Lectotype 4 (pre-

sent designation) [glued on card, antennal segments 3

and 4, right middle leg, left fore and left hind legs miss-

ing]: ‘Kosempo Formosa H. Sauter 1911’ (print); ‘7. VIT

(print); ‘Lygus kosempoensis n. sp. ‘(hand-written);

‘Mus. Zool. H: fors Spec. typ. No. (print), 10181 Lygus

kosempoensis Popp. (hand-written) [zmun]. [examined]

Other material examined. — Paralectotypes 24 (present

designation): 1d [glued on card, all legs missing (except for

all coxae)], ‘Kosempo Formosa H. Sauter 1911’ (print); 7.

VII (print); ‘Mus. Zool. H: fors Spec. typ. No. (print),

10182 Lygus kosempoensis Popp. (hand-written); 16

[glued on card, antennal segment 4, fore legs (except coxae),

left middle leg (except coxa) missing], ‘Kosempo Formosa

H. Sauter 1911’ (print); ‘7. VIP (print); ‘Mus. Zool. H: fors

Spec. typ. No. (print), 10183 Lygus kosempoensis Popp.

(hand-written)’ [zmuH].

Additional description. — Vertex width : head

width = 0.30 in male. Cuneus about 1.3 times as long

as its basal width. Body length 3.60-3.65, width 1.80-

1.85. Head length 0.25-0.28, width 1.02-1.05.

Antennal segments lengths 0.41-0.50: 1.30-1.46:

0.69-0.74: 0.55-0.58. Pronotum length 0.78-0.80,

width 1.51-1.57. Corium length 1.76-1.87; cuneus

length 0.63-0.66; claval commissure length 0.69-

OWS

Male genitalia (lectotype dissected) (figs. 1-5). —

Parameres as in figs. 2-5. Vesical spicule absent; ven-

tral sclerite S-shaped; wing-shaped sclerite triangular,

outer margin with big spines; lateral sclerite slender,

slightly twisted; sublateral sclerite developed, broad;

seminal duct swollen greatly, secondary gonopore

broad.

Note. — A female specimen [zmun], similarly la-

belled as the type series, but without a ‘type’ label,

185

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 1-5. Male genitalia of Apolygus kosempoensis. — 1, vesica; 2-3, left paramere; 4-5, right paramere.

was also examined. The following notes refer to this

female: Vertex width : head width = 0.42 in female.

Body length 3.83, width 1.80. Head length 0.22,

width 0.88. Pronotum length 0.96, width 1.68.

Corium length 1.84; cuneus length 0.63; claval com-

missure length 0.83.

Remarks. — On the basis of the body size and form,

colour pattern, texture of the body surface and the

structure of male genitalia, this species is here placed

in the genus Apo/ygus China.

Apolygus eous (Poppius, 1915) comb. n.

Lygus eous Poppius, 1915: 27. — Holotype d [glued on card,

left antennal segment 4, right antennal segment 3 and 4,

right hind leg (except coxa) missing]: ‘Fuhosho Formosa

H. Sauter’ (printed); ‘7. IX’ (printed), ‘Poppius det.’

(printed); ‘Holotypus’ (printed on reddish brown card);

‘Lygus eous n. sp.’ (hand-written) [pE1].[examined]

Additional description. — Vertex width : head

width = 0.30 in male. Cuneus about 1.7 times as long

as its basal width in male. Body length 3.71, width

1.75. Head length 0.34, width 0.91. Antennal seg-

ments lengths 0.51: 1.32: 0.80: ? (lost). Pronotum

length 0.86, width 1.49. Corium length 1.89; cuneus

length 0.59; claval commissure length 0.77.

Remarks. — On the basis of the external appearance

and the exposed parameres, this species is here recom-

bined with the genus Apolygus China.

186

Apolygus fuhoshoensis (Poppius, 1915) comb. n.

Lygus fuhoshoensis Poppius, 1915: 31.

Material examined. — Holotype d [glued on card, anten-

nae (except left segment 1), right middle and right hind legs

(except coxae) missing, left middle tibia broken near apex,

apical part missing]: ‘Fuhosho Formosa H. Sauter’ (print);

‘7. IX.’ (print); ‘Poppius det.’ (print); ‘Typus’ (print, on pale

brown card); ‘Lygus fuhoshoensis n. sp.’ (hand-written);

‘Holotypus’ (print on reddish brown card) [Der].

Additional description. — Vertex width : head

width = 0.29 in male. Cuneus about 1.2 times as long

as its basal width in male. Body length 3.92, width

1.71. Head length 0.23, width 0.85. Antennae lost.

Pronotum length 0.86, width 1.49. Corium length

1.89; cuneus length 0.59; claval commissure length

0.77.

Remarks. — Judging by the general habitus and the

exposed parameres, this species is here placed in the

genus Apolygus China.

Lygocoris (Neolygus) bipuncticollis (Poppius,

1915) comb. n. (fig. 6)

Lygus bipuncticollis Poppius, 1915: 25. Holotype ¢ [glued

on card, left antennal segment 3 broken (apical part miss-

ing), right antennal segment 4 and right hind leg (except

left coxa) missing]: ‘Kankau (Koshun) Formosa H. Sauter

VII.1912’ (print); ‘Poppius det.’ (print); ‘Holotypus

(print on reddish brown card); ‘Lygus bipuncticollis n.

sp. (hand-written) [DEI].[examined] |

Additional description. — Vertex width : head

width = 0.34 in male. Cuneus about 1.6 times as long

as its basal width in male. Body length 3.92, width

2.39. Head length 0.36, width 1.09. Antennal seg-

ments lengths 0.71: 1.75: 0.85: ? (lost). Pronotum

length 0.94, width 1.62. Corium length 1.90; cuneus

length 0.63; claval commissure length 0.69.

Remarks. — The exposed right paramere (fig. 6)

clearly proved to be of the typical Neolygus form; the

body form and colour pattern also show that it is a

species of the subgenus Neolygus Knight, genus

Lygocoris Reuter.

Lygocoris ( Neolygus) v-nigrum (Poppius, 1915)

comb. n.

Lygus v-nigrum Poppius, 1915: 31. — Holotype d : [glued on

card, left antenna, right antennal segments 3 and 4, fore

and middle legs (except coxae), left hind tibia and tarsus

missing] — ‘Taihorinsho Formosa H. Sauter, 1909’

(print); 7. IX.’ (print); ‘Poppius det.’ (print); ‘typus

(print on pale brown card); ‘Lygus fuhoshoensis n. sp.’

(hand-written); ‘Holotypus’ (print on reddish brown

card) [per] [examined].

Additional description. — Vertex width : head

width = 0.31 in male. Cuneus about 1.7 times as long

as its basal width in male. Body length 4.14, width

1.76. Head length 0.36, width 0.88. Antennal seg-

ments lengths 0.46: 1.42: ?(broken): ? (lost). Prono-

tum length 0.81, width 1.44. Corium length 1.94;

cuneus length 0.72; claval commissure length 0.81.

Remarks. — The body form, colour pattern and the

Lu & ZHENG: Poppius' Lygus’ types

exposed right paramere indicate that it isa member of

the subgenus Neolygus Knight of the genus Lygocoris

Reuter.

Lygocoris (Neolygus) disciger (Poppius, 1915)

Lygus disciger Poppius, 1915: 34.

Lygocoris (Neolygus) disciger; Schwartz & Kerzhner, 1997:

252. — Lectotype (present designation): [left corium

and left membrane, both antennae missing] — ‘Fuhosho

Formosa H. Sauter’ (print); ‘7. IX.’ (print); ‘Lygus de-

sciger n. sp. (hand-written); ‘Mus. Zool. H: fors Spec.

type. No (print), 10176 Lygus disciger Popp. (hand-writ-

ten) [zmuH].[examined]

Additional description. — Vertex width : head

width = 0.37 in female. Cuneus about 1.6 times as

long as its basal width in female.

Appendix. — The following descriptive notes are

based on specimens collected from China: Guang-

dong Province (16) and Yunnan Province (24,29),

which we identified as Lygocoris (Neolygus) disciger af-

ter comparing with the lectotype.

Descriptive notes. — Vertex width : head width =

0.30 in male.

Male genitalia (figs. 7-11). — Parameres as in figs.

8-11. Vesical spicule slender, bending near apex; with

a sclerite structure at centre; with 4 membranous

lobes; seminal duct cylindric, secondary gonopore rel-

ative small. Body length 3.78-4.00, width 1.49-1.62.

Head length 0.25-0.28, width 0.88-0.93. Antennal

segments lengths 0.55-0.58: 1.52-1.66: 0.76-0.92:

Figs. 6-11. Male genitalia of Lygocoris (Neolygus) spp. — 6, Right paramere of L. bipuncticollis; 7-11, L. disciger. — 7, vesica; 8-

9, left paramere; 10-11, right paramere. Scale a: 0.2 mm to fig. 6; b: 0.2 mm to 7-11.

187

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 12-16. Male genitalia of Sabactus sauteri. — 12, vesica; 13-14, left paramere; 15-16, right paramere.

0.50-0.54. Pronotum length 0.63-0.68, width 1.32-

1.40. Corium length 1.76-2.05; cuneus length 0.55-

0.58; claval commissure length 0.81-0.90.

Remarks. — Schwartz & Kerzhner (1997) place this

species in the subgenus Neolygus based on the female

genitalia, complete basal carina of vertex and tibial

spine color; the structure of the vesica, described in

this study, also suggests a Neolygus-form.

Sabactus sauteri (Poppius, 1912) comb. n.

(figs. 12-16)

Lygus sauteri Poppius, 1912: 303.

Sabactus sauteri; Schwartz & Kerzhner, 1997:256, figs. 35-

38. — Lectotype 9 (present designation) [left antenna,

right antennal segment 4 missing]: ‘Chip Chip Formosa

Sauter II 07-09’ (print); ‘Lygus sauteri n. sp. B. Poppius

det.’ (print); ‘Mus. Zool. H: fors Spec. typ. No (print),

9932 (hand-written) Lygus sauteri Popp. (print) [ZMuH].

[examined]

Other material examined. — Two other specimens be-

longing to the type-series, labelled with ‘Lygus sauteri’

(Mus. Zool. H: fors Spec. typ. No 10191: male; Mus. Zool.

H: fors Spec. typ. No 10190: female) which were borrowed

along with the lectotype (Mus. Zool. H: fors Spec. typ. No

9932) obviously belong to the subfamily Deraeocorinae

(Deraeocoris sp.?) and are thus misidentified paralectotypes.

Additional description. — Vertex width : head

width = 0.38 in female. Cuneus about 1.6 times as

long as its basal width.

Appendix. — The following descriptive notes are

base upon examinations of specimens collected from

China: Sichuan Province (19), Guangxi Province

188

(16, 29), Hainan Province (12), Yunnan Province

(16,29) and Taiwan Province (464, 549 [Bishop

Museum]) which we identified as Sabactus sauteri, af-

ter comparing with the lectotype.

Descriptive notes. — Vertex width : head width =

0.30 in male. Body length 3.92-4.52, width 1.85-

2.12. Head length 0.23-0.34, width 0.85-0.88.

Antennal segments lengths 0.44-0.48: 1.33-1.41:

0.71-0.87: 0.62-0.69. Pronotum length 0.88-0.99,

width 1.53-1.70. Corium length 1.70-1.87; cuneus

length 0.58-0.78; claval commissure length 0.61-

0.78.

Male genitalia. — Basal part of sensory lobe of left

paramere strongly protruded; apex of hypophysis

truncate. Hypophysis of right paramere short and

broad, apex truncate. Vesical spicule absent; with 4

membranous lobes, the lateral margin of the largest

one with several large sclerotized spines, others

minutely spinulose on surface; seminal duct short,

swollen at middle, secondary gonopore broad.

Remarks. — We follow here the generic placement

suggested by Schwartz & Kerzhner (1997). However,

the posterior wall of bursa copulatrix of this species

lacks the characteristic bifid inter-ramal lobes found

in the type species Sabactus institutus Distant (Kelton

1955):

ACKNOWLEDGEMENTS

We are grateful to Dr. R. Gaedicke of the Deut-

sches Entomologisches Institut, Eberswalde, Dr. L. |

Huldén of Zoological Museum, University of Helsin-

ki, and Mr. K. Arakaki of Bishop Museum, Honolu- |

lu for the kind loan of valuable specimens. We also

are very thankful to the Foundation of the Education-

al Committee of China for its grants for this project.

REFERENCES

Carvalho, J., 1959. A catalog of the Miridae of the world.

Part IV. — Arqivos do Museo nacional, Rio de Janeiro 48:

1-384.

Carvalho, J., 1980. Analecta Miridologica, IV: observations

on type specimens in the National Museum of Natural

History, Budapest, Hungary (Hemiptera, Miridae). — Re-

vista Brasileira de Biologia 40: 649-658.

Kelton, L. A., 1955. Genera and subgenera of the Lygus

complex (Hemiptera: Miridae). — Canadian Entomolo-

gist 87: 277-301.

Poppius, B., 1912. H. Sauter's Formosa-Ausbeute: Miridae

(Hem.). — Entomologische Mitteilungen 1: 302-394.

Poppius, B., 1915. H. Sauter's Formosa-Ausbeute: Nabidae,

Lu & ZHENG: Poppius' ‘Lygus’ types

Anthocoridae, Termatophylidae, Miridae, Isometopidae

und Ceratocombidae (Hemiptera). — Archiv für Naturge-

schichte 80A (8): 1-80.

Schuh, R., 1995. Plant bugs of the world (Insecta: Hetero-

ptera: Miridae) - systematic catalog, distributions, host

list, and bibliography. — The New York Entomological

Society, New York, 1329 pp.

Schwartz, M. D. & I. M. Kerzhner, 1997. Type specimens

and identity of some Chinese species of the ‘Lygus com-

plex’ (Heteroptera: Miridae). — Zoosystematica Rossica

5(2): 249-256.

Yasunaga, T., 1991. A revision of the plant bug, genus Lygo-

coris Reuter from Japan, Part I (Heteroptera, Miridae,

Lygus-complex). — Japanese Journal of Entomology 59:

435-448.

Received: 6 May 1996

Accepted: 20 August 1997

189

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

190

WoLFRAM MEY

Museum für Naturkunde der Humboldt-Universität Berlin

THE GENUS HYDROPSYCHE PICTET, 1834 ON

ISLANDS IN THE WEST PACIFIC REGION AND

DESCRIPTION OF NEW SPECIES (TRICHOPTERA:

HYDROPSYCHIDAE)

Mey, W., 1998. The genus Hydropsyche Pictet, 1834 on islands in the West Pacific Region and

description of new species (Trichoptera: Hydropsychidae). — Tijdschrift voor Entomologie 140

[1997]: 191-205, figs. 1-40. [rss 0040-7496]. Published 26 March 1998.

Species of the genus Hydropsyche are largely distributed over the islands of the West Pacific.

They are grouped into 4 species groups on the basis of male genitalic characters. The hitherto

known species of the 4 groups are listed, and the distributional area of the groups is demon-

strated. Among the 57 cited species 16 are newly described: H. germanorum sp. n., H. staphylo-

stirpis sp. n., H. taiwanensis sp. n., H. vialigni sp. n., H. initiana sp. n., H. malickyi sp. n., H.

ambonensis sp. n., H. seramensis sp. n., H. palawanensis sp. n., H. gemellata sp. n., H. salki sp. n.,

H. sirimauna sp. n., H. suppleta sp. n., H. naumanni sp. n., H. sulana sp. n., H. bacanensis sp. n.

Correspondence: Dr. W. Mey, Museum für Naturkunde, Humboldt-Universität Berlin,

Invalidenstr. 43, D-10115 Berlin, Germany.

Key words. — Taxonomy; new species; biogeography; SE Asia; West Pacific Island; Sundaland;

Hydropsyche; species groups; Hydropsychidae; Trichoptera.

The species of Hydropsyche Pictet, 1834 show an

interesting rate of endemism and speciation in the is-

lands between Southeast Asia (= SE Asia) and

Australia. All of the larger islands have a number of

Hydropsyche endemics. By contrast, there are no

species with a wide distribution over the entire region

or with a range through several islands or island

groups. Previous records of wide distribution areas for

some species proved to be based on the misidentifica-

tion of some specimens, e.g. H. hobbyi Mosely, 1951

described from Borneo and New Guinea in Mosely

(1951), corrected by Kimmins (1962), and A. bryan-

ti Banks, 1939 recorded from Sumatra, Java and

Sumba in Ulmer (1951), corrected in Mey (1998).

The high degree of endemism of Hydropsyche ob-

served in the West Pacific Region suggests the exis-

tence of further, still unknown species on the islands

which have not yet been sampled for caddisflies. It is

to be expected that Halmahera, Seram, Obi, Sula,

Buru, the Lesser Sunda Islands and especially New

Guinea will have two to four additional Hydropsyche

species each. Furthermore, the relatively better inves-

tigated Greater Sunda Islands, Sulawesi, the

Philippines and Taiwan have diverse caddisfly faunas,

which are by no means adequately surveyed.

Unknown Hydropsyche species are still to be found in

these larger islands and archipelagos.

With this background it is clear, that we have not

yet reached the advanced level of faunistic research,

which is the prerequisite for any serious discussion of

the origin, dispersal or diversity pattern of the genus

in the West Pacific Region. In addition, the phyloge-

netic relationships within Hydropsyche have been

studied only partially (e.g. Malicky 1977, Mey 1996,

Nimmo 1987, Ross 1944). Even the concept of

Hydropsyche and its probable sistergroup is a matter of

much debate (Bueno-Soria 1984, Flint & Bueno-

Soria 1987, McFarlane 1976, Nielsen 1981, Ross &

Unzicker 1977, Schefter 1996, Schefter et al. 1986,

Schmid 1979, Schuster 1984, Tanıda 1986, Tian &

Li 1987). Without a detailed phylogenetic analysis

and without adequate knowledge of the existing

species and their geographic distributions, it is not

possible to use the genus as an example for recon-

structing the evolutionary history of the region and its

biota. However, the data on Hydropsyche so far accu-

mulated are quite sufficient to recognise that the

group is an extremely promising candidate for eluci-

dating dispersal or vicariance patterns and tracing

them further into the past.

191

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

The purpose of this paper is to contribute to the

improvement of the faunistic database of the genus. It

presents the first records of Hydropsyche species from

the Moluccas and from Palawan. In addition, a list of

all described taxa and their distribution has been

compiled. It is thought that this will be the starting

point for an enhanced faunistic research in the region,

which should subsequently lead to a biogeographic

analysis of the genus.

METHODS

Material examined in this study was obtained from

several sources. Most of the caddisfly specimens were

collected with a mercury-vapour light source by sev-

eral entomologists. A portable generator was used as

the power unit. The majority of specimens is pre-

served in 75% ethanol. Examination and drawing of

male genitalia was done after the entire abdomen had

been cleared in boiling kom, washed and replaced in

ethanol. Illustrations were prepared with the aid of a

Wild drawing equipment on a Wild M8 stereo-mi-

croscope.

The shape of the genitalic structures in lateral, ven-

tral and dorsal view varies between species. They pro-

vide clear characters for species separation and recog-

nition, which are best appreciated by comparing the

figures, rather than by long descriptions or written

keys. In contrast to the genitalia, the external charac-

ters (e.g. coloration, mouth parts, wing venation) of

the species are extremely uniform and can not be used

in the identification of species.

DELIMITATION OF THE STUDY AREA

This study concerns the West Pacific region, which

covers most of the islands between continental SE

Asia and North Australia. Within this vast area the

following countries are situated: Indonesia, West

Malaysia (Sarawak, Sabah), Brunei, Philippines,

Taiwan, Papua New Guinea, Solomon Islands.

Hainan is excluded from the present study.

THE SPECIES-GROUPS OF HyDROPSYCHE IN THE

WEST PACIFIC

The Hydropsyche species of SE Asia and Australia

have not attracted much interest, probably because

they are not frequently reported in the literature nor

are there many people working with them. Even in

discussions of the phylogeny of the genus on a world-

wide scale, the South East Asian and West Pacific

species have been disregarded. However, the region

has at least 4 distinct species groups, which have no

further representatives elsewhere (A. pluvialis group,

H. formosana group, H. hamifera group, H. buergersi

192

group). Members of the H. pluvialis group were not

recorded from the Sunda Islands up to now. In addi-

tion, the regions bears a number of isolated species,

which cannot be assigned to any of the defined

species groups (e. g. H. vasoumittra Schmid, 1961). A

fifth group has a subgeneric name of its own:

Ceratopsyche Ross & Unzicker, 1977. I do not use this

name for the moment, because it implies also a sub-

generic level for the other groups. As a consequence,

three new genus group names would have to be pro-

posed now or later. The taxonomic treatment of the

groups, however, is not the subject of the present ar-

ticle. New names should be proposed in accordance

with the results of a sound phylogenetic study. Before

such an analysis is published I prefer to group related

species into species-groups. Although the groups are

informal entities I follow common nomenclatorial

practice and use group-names based on the first de-

scribed species in each group. The definitions of the

groups are based entirely on characters of the male

genitalic apparatus.

1. Hydropsyche newae group

Ceratopsyche Ross & Unzicker, 1977: 305.

morosa group, Schefter & Wiggins 1986: 2.

silfvenii group, Malicky 1977: 2.

penicillata group, Schmid 1965: 137.

Diagnosis

Phallotheca sinuate, with the genital opening on

the dorsal side between paired phallotremal sclerites;

apex of phallotheca extending beyond phallotremal

sclerites, partially membranous and with spicules or

with membranous lobes; dorsolateral membrane sim-

ple or complex; 10th segment with two apical ap-

pendages (= appendices digitiformis).

Distribution

Holarctic and Oriental biogeographic region (fig.

List of species

H. annulata (Ulmer, 1905) Java

H. germanorum sp. n. Sumatra

H. orbiculata Ulmer, 1911 Taiwan

H. staphylostirpis sp. n. Lombok

H. taiwanensis sp. n. Taiwan

H. vialigni sp. n. Sumatra

2. Hydropsyche hamifera group

bryanti-celebes-annulata group, auctorum

javanica group, Mey 1990: 414

Diagnosis

Basis of phallotheca simply bent; phallotremal scle- |

Asia and

Australia

2000 km

——

Mey: Hydropsyche West-Pacific region

Asia and

Australia

Fig. 1. Distributional area of the Hydropsyche newae group. —

The map shows the distributional limits in Asia. The entire

area of the group includes Northern and Middle Europe and

North America.

rites on dorsal side of phallotheca fused, forming a

ring with a shorter or longer elongation on the ventral

side, forming a phallotremal tongue, which gives the

apex of the phallotheca a trifurcate appearance; dor-

solateral membrane with 1 - 3 appendages; 10th seg-

ment with apical appendages and a pair of small

processes on a flat dorsal side.

Distribution

Sunda Islands, Philippines, Wallacea biogeograph-

ic region, New Guinea (fig. 2).

List of species

H. ambonensis sp. n. Ambon

H. bacanensis sp. n. Bacan

H. bifurcata Mey, 1990 Luzon

H. brueckmanni Mey, 1995 Luzon

H. buenafei Mey, 1998 Negros

H. calawiti Mey, 1995 Mindoro

H. cebuensis Mey, 1998 Cebu, Negros

H. celebensis Ulmer, 1951 Sulawesi

H. davisi Mey, 1998 Negros, Mindanao

H. declinans Mey, 1990 Luzon

H. excelsa Mey, 1990 Luzon

H. fascelina Mey, 1998 Mindanao

H. faurai Naväs, 1925 Luzon

H. forcipata Ulmer, 1930 Mindanao

H. gemellata sp. n. Sulawesi

Fig. 2. Distributional area of the

group.

H. gerostizai Mey, 1998

H. hamifera Ulmer, 1905

H. hobbyi Mosely, 1951

H. initiana sp. n.

H. isolata Banks, 1931

H. javanica Ulmer, 1905

H. luzonica Mey, 1990

H. malickyi sp. n.

H. mindanensis Mey, 1998

H. mindorensis Mey, 1995

H. moselyi Kimmins, 1962

H. muelleri Mey, 1998

H. naumanni sp. n.

H. negrosensis Mey, 1998

H. palawanensis sp. n.

H. rizali Banks, 1937

H. salki sp. n.

H. saranganica Ulmer, 1951

H. schintlmeisteri Mey, 1990

H. secundaria Mey, 1998

H. seramensis sp. n.

H. sirimauna sp. n.

H. sulana sp. n.

H. suppleta sp. n.

H. unitaria Mey, 1990

H. villica Mey, 1990

H. sp. n. (Mey, in press)

193

Hydropsyche hamifera

Negros

Sulawesi

Borneo

Sumatra

Borneo

Java

Luzon

Sumatra

Mindanao

Mindoro

New Guinea

Mindanao

Sulawesi

Negros

Palawan

Mindanao

Sulawesi, Moluccas

Java

Mindanao

Seram

Ambon, Seram

Sula

Ambon, Seram

Luzon

Talaud.

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Asia and

Australia

2000 km

Asia and

Australia

2000 km

Fig. 3. Distributional area of the Hydropsyche formosana

group.

3. Hydropsyche formosana group

Mexipsyche Ross & Unzicker, 1977: 305-306, sensu Tian &

Li 1987: 125-126.

Diagnosis

Basis of phallotheca simply bent; phallotremal scle-

rites vestigial or lost; bifid apex of phallotheca mem-

branous on dorsal side, sclerotized ventrally and with

one or two keels; lateral sides of phallotheca with a

pair of proximally produced appendages; endophallus

nearly as long as phallotheca; segment 9 narrowed;

segment 10 with apical appendages.

Distribution

Oriental biogeographic region (fig. 3).

List of species

H. bryanti Banks, 1939

Java, Sumatra

H. banksi Kimmins, 1955 Borneo

H. doctersi Ulmer, 1951 Java

H. formosana Ulmer, 1911 Taiwan, Malaysia

H. irroratella Ulmer, 1951 Java

H. sp. n. (Mey, in press) Lombok

H. sp. n. (Mey, in press) Sumatra

H. sp. n. (Mey, in press) Borneo.

194

Fig. 4. Distribution of the Hydropsyche buergersi group.

4. Hydropsyche buergersi group

Diagnosis

Basis of phallotheca simply bent; phallotremal scle-

rites small, situated at the tip of phallotheca between

a pair of sclerotized, large valves; dorsolateral mem-

brane lacking; phallicata with a membranous area on

the ventral side, just below the phallotremal sclerites;

segment 9 of usual size; segment 10 with short apical

appendages.

Distribution

Philippines, New Guinea, Solomon Islands (fig. 4).

List of species

H. buergersi Ulmer, 1915

H. flynni Korboot, 1964

H. nasuta Ulmer, 1930

H. tapena Kimmins, 1957

New Guinea

Mindanao

Guadalcanal.

The buergersi group is established here to sum-

marise 3 species from New Guinea and one Philip- |

pine species. They do not belong to any of the afore- |

mentioned groups. Interestingly, they show affinities |

to some species of continental South East Asia, e.g.

H. adonis (Malicky, 1996), H. harpagofalcata Mey, |

1995 and H. napaea Mey, 1996. The diagnosis of the

group is tentative, however. The type of H. buergersi

Ulmer, 1915 was deposited in the Zoological Mu- |

seum Berlin, but could not be found again. Neboiss

(1986: 115) reproduced the original drawings. A re-

description of H. buergersi Ulmer, 1915 together with

collecting efforts to obtain new material of related

species are necessary to provide more arguments for

the monophyly and validity of the group.

5. Species incertae sedis

H. closi Navas, 1927

H. testacea Naväs, 1933

Mindanao

New Guinea.

DESCRIPTION OF NEW SPECIES

The descriptions are based exclusively on male gen-

italic characters. The female genitalia are very ho-

mogenous and can hardly be used for separating the

species. In addition, most species occur sympatrically

with other species of the genus, thus rendering the as-

sociation of sexes doubtful.

Homologies of the phallic apparatus in Aydro-

psyche have been interpreted and named differently by

various authors (e.g. Nielsen 1981, Ross & Unzicker

1977, Schmid 1979). The terminology used in this

article follows Schefter & Wiggins (1986: 4-5).

Most of the type material is deposited in the Mu-

seum für Naturkunde der Humboldt Universität Ber-

lin (MNHB). Paratypes of some species are in coll. Ma-

licky (Lunz am See). No explicit indication of

paratype depository means storage in the MNHB.

newae group

Hydropsyche germanorum sp. n.

(figs. 5-6)

Type material. — Holotyped (pinned), Indonesia,

Sumatra, Sumatera Utara, Medan, Tiga Dolok,

13.11.1995, leg. E. Diehl, in MNHB. — Paratypes: 5d,

same data as for holotype; 21d, same locality,

22.11.1995, leg. VE Diehl; 258, same locality,

20.1.1995, leg. E. Diehl; 58, Indonesia, Sumatra,

Sumatera Aceh, Leuser N.P., Ketambe, 22.-

24.1.1995, leg. A. Kallies; 38, 2 2, Sumatera Aceh,

Calang, Kuala Don, Febr. 1996, leg. A. Kallies; 2d,

Indonesia, Sumatra, Sumatera Barat, 12 km E

Padang, Mt.Talang, 1600m, 18.ix.1991, leg. A.

Schintlmeister.

Description. — External characters: Length of

forewing 6.5-7.3 mm. Head and thorax brown, with

golden brown pilosity. Eyes black. Antennae yellow,

with indistinct darker lines on the articulations of fla-

gellomeres, tip of antennae brown, extending to apex

Mey: Hydropsyche West-Pacific region

of forewing. Maxillary palpi brown, with last joint

longer than the two preceding together. Labial palpi

light brown. Legs yellowish. Forewings yellowish

brown, with numerous pale spots dispersed over the

wing, somewhat darkened towards outer wing mar-

in.

Male genitalia (fig. 5-6): Segment 10 with short

fingerlike appendages. Apical segment of inferior ap-

pendage (= harpago) broad, slightly bent and with an

oblique tip. Phallotheca sinuate, with short apical

part. Phallotremal sclerites large, in vertical position

on the dorsal side of phallotheca. Dorsolateral mem-

brane without appendages, but more proximal a small

membraneous window with a small spine. Apical

membrane of phallotheca with indistinct spicules, lat-

eral membrane with a small lobe, directed distally.

Etymology. — The species is named in honour of

the many German entomologists, who collected ma-

terial of this species on several places in Sumatra.

Remarks. — The species is related to H. annulata,

but differs by the vestigial spicules of the apical mem-

brane, the lacking dorsolateral appendages and by the

broad apical segments of the inferior appendages.

Both species occur sympatrically in Padang, Sumatera

Barat.

Hydropsyche staphylostirpis sp. n.

(figs. 7-9)

Type material. — Holotyped (pinned), Indonesia,

Lombok, Narmada, 17.11.1927, leg. Dr.[B.] Rensch,

in MNHB. — Paratypes: 16, 19, same locality,

14.11.1927.

Description. — External characters: Length of

forewing 7 mm, wings golden brown, reticulate pat-

tern weakly developed. Other characters as in H. ger-

manorum sp. n.

Male genitalia (figs 7-9): Segment 9 shortened and

slender in lateral view. Segment 10 with nearly

straight apical appendages. Inferior appendages with

a broad basal segment (= coxopodit) and an elongate

second segment (= harpago). Phallotheca sinuate,

ending with the phallotremal sclerites. Dorsal mem-

braneous area without appendages, but extending

ventrally to form a ventral membraneous part be-

neath the phallotremal sclerites and equipped with

two curved spines. Phallotremal sclerites excavated

apicolaterally.

Remarks. — The new species is very remarkable by

the reduced apical part of the phallotheca and the pair

of spines beneath the phallotremal sclerites. It is a

quite isolated species in the newae group, but shows

affinities to À. annulata.

195

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 5-11. Male genitalia of Hydropsyche spp. — 5-6, H. germanorum sp. n., 5, lateral, 6, ventral; 7-9, H. staphylostirpis sp. n.,

7, lateral, 8, ventral, 9, tip of phallotheca, caudal; 10-11, H. vialigni sp. n., 10, lateral, 11, ventral.

196

Hydropsyche vialigni sp. n.

(figs. 10-11)

Type material. — Holotyped (pinned), Indonesia,

Sumatera Utara, Medan, Tiga Dolok (Holzweg 2),

22.11.1995, leg. E. Diehl, in MNHB. — Paratypes: 24,

same data as holotype.

Description. — External characters: Length of

forewing 9-9.5 mm. Coloration and wing patterns as

in A. germanorum sp. n.

Male genitalia (figs. 10-11): Segment 10 with

curved apical appendages. Apical segment of inferior

appendages rounded apically, not dilated. Phallo-

theca sinuate, with short apical part behind phallo-

tremal sclerites. Phallotremal sclerites large, on dorsal

side of phallotheca. Dorsolateral membrane without

appendages, but more proximally a small membrane-

ous spot with a short spine. Apical membrane of phal-

lotheca with 2 protruding spicules, lateral membrane

with small lobes, reaching tip of phallotheca.

Remarks. — The new species is closely related to H.

germanorum sp. n., but can be distinguished easily by

the larger size of the species and by the unmodified

second joint of the inferior appendages.

Hydropsyche taiwanensis sp. n.

(figs. 12-13)

Type material. — Holotyped (in alcohol), Taiwan,

Fushan Ilan, 25.xi.1995, leg. H. J. Wu, in MNHB. —

Paratypes: 18, 29, same data as holotype; 1d,

Taiwan, Fushan Botanical Garden, 18.iv.1996, leg.

S. H. Yen.

Description. — External characters: Length of

forewings 8 mm. Other characters as in H. germano-

rum Sp. n.

Male genitalia (figs. 12-13): Segment 9 and 10 not

divided dorsally by a depression. Apical appendages

of segment 10 broad, short and medially bent.

Inferior appendages with a short first segment (= cox-

opodit), second segment (= harpago) elongate, with a

rounded tip. Phallotheca sinuate. Dorsolateral ap-

pendages with a short spine. Apical part of phallothe-

ca with long, membraneous appendages, ending with

a short spine. 2 small spikes in the apical membrane.

Phallotremal sclerites large and broadly fused.

Remarks. — H. taiwanensis sp. n. is very similar to

H. orbiculata. The species can be separated by the

form and structure of the phallic apparatus.

Mey: Hydropsyche West-Pacific region

hamifera group

Hydropsyche initiana sp. n.

(figs. 14-16)

Type material. — Holotyped (pinned), Indonesia,

Sumatera Utara, Sipirok, 25.11.1995, leg. E. Diehl, in

MNHB. — Paratypes: 1d (pinned), same data as holo-

type; 10d, Indonesia, Sumatera Utara, Medan, Tiga

Dolok (Holzweg 2), 20.1.1995 (5d), 13.11.1995

(259), 22.u. 1995130) leg. Diehl:

Description. — External characters: Length of

forewing 7-8 mm. Coloration and wing pattern as in

H. germanorum sp. n.

Male genitalia (figs. 14-16): Segment 10 flat, with-

out a carina. Segment 9 with apical appendages and

with a pair of small dorsal processes fused at the base

to form a Y-like structure. Apical segment (= harpa-

g0) of inferior appendages slender and pointed. Phal-

lotheca with a bifid membraneous apical part, armed

with a spine. Ventral tongue of phallotremal sclerites

not reaching the tip of phallotheca. Dorsolateral

membrane with broad appendages, ending with a

long spur each.

Remarks. — H. initiana sp. n. is related with A. ja-

vanica and H. malickyi sp. n. The shape of the 10th

segment and the structure of the phallic apparatus

clearly separates the species.

Hydropsyche malickyi sp. n.

(figs. 17-19)

Type material. — Holotyped (pinned), Indonesia,

Sumatera Utara, Medan, Tiga Dolok (Holzweg 2),

22.11.1995, leg. E. Diehl, in MNHB. — Paratypes: 64,

same data as holotype; 2d, same locality, 13.11.1995;

34, same locality, 20.1.1995, all leg. E. Diehl; 14,

Sumatera Aceh, Tibbing Raja, 15.10.1996, leg. A.

Kallies.

Description. — External characters: Length of

forewing 8-8.5 m. Coloration and wing pattern as in

H. germanorum sp. n.

Male genitalia (figs. 17-19): Segment 10 with slen-

der apical appendages and a pair of small dorsal

processes. Apical segment of inferior appendages slen-

der and ending with a seta. Base of phallotheca broad,

apical portion membraneous dorsally and with two

spines. Ventral tongue of phallotremal sclerites en-

larged and hook-like apically, not reaching tip of

phallotheca. Dorsolateral membrane with broad ap-

pendages and a short spine.

Etymology. — The new species is named in honour

of Hans Malicky, who discovered and described a

great deal of the caddisfly fauna of Sumatra.

197

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 12-19. Male genitalia of Hydropsyche spp. — 12-13, H. taiwanensis sp. n., 12, lateral, 13, ventral; 14-16, H. initiana sp.

n., 14, lateral, 15, ventral, 16, tip of phallotheca, dorsal; 17-19, H. malickyi sp. n., 17, lateral, 18, ventral, 19, tip of phal-

lotheca, dorsal.

198

Mey: Hydropsyche West-Pacific region

Figs. 20-25. Male genitalia of Hydropsyche spp. — 20-21, H. ambonensis sp. n., 20, lateral, 21, ventral; 22-23, H. seramensis sp.

n., 22, lateral, 23, ventral; 24-25, H. palawanensis sp. n., 24, lateral, 25, ventral.

199

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 26-32. Male genitalia of Hydropsyche spp. — 26-27, H. gemellata sp. n., 26, lateral, 27, ventral; 28-30, A. salki sp. n., 28, |

lateral, 29, dorsal, 30, ventral; 31-32, H. sirimauna sp. n., 31, lateral, 32, ventral.

200

Remarks. — The species is related to H. javanica.

The main differences are exhibited by the long phal-

lotheca and the shape of the phallotremal tongue.

Hydropsyche ambonensis sp. n.

(figs. 20-21)

Type material. — Holotyped (in alcohol), Indone-

sia, Ambon, Gunung Sirimau, 500m, 23.vii.1995,

leg. S. Naumann, in MNHB. — Paratypes: 16, same

data as holotype; 34, Indonesia, Ambon, Katalai,

470m, 26.vii.1995, leg. S. Naumann.

Description. — External characters: As in A. ger-

manorum sp. n., wing length 8 mm.

Male genitalia (figs. 20-21): Short apical ap-

pendages and short dorsal processes on segment 10.

Harpago of inferior appendages much smaller than

coxopodit, without peculiarities. Tip of phallotheca

trifid. Tongue of phallotremal sclerites strongly

curved and sclerotized. Dorsolateral membrane with

two pairs of appendages, the smaller with minute

spines apically, the longer ending with a large, bifur-

cate spur.

Remarks. — The new species is a close relative of H.

seramensis sp. n. They differ in the form of the 10th

segment and in the structure of the phallic apparatus.

Hydropsyche seramensis sp. n.

(figs. 22-23)

Type material. — Holotyped (in alcohol), Indone-

sia, Seram, Piljana, Tule, 26.2.1995, leg. P. Salk, in

MNHB. — Paratypes: 1d, 12 , same data as holotype.

Description. — External characters as in H. germa-

norum sp. n. Length of forewing 7.5 mm (d)-8 mm

(DR

Male genitalia (fig. 22-23): Apical appendages of

segment 10 straight, dorsal processes small. Second

segment of inferior appendages short and simple.

Phallotremal tongue longer than lateral tips of phal-

lotheca, angulate at its base and with a small lateral

teeth. Dorsolateral membrane with 2 pairs of ap-

pendages, the smaller with minute spines and the

longer with an apical spur.

Remarks. — H. seramensis sp. n. is related to the pre-

ceding species. The form of the phallotremal tongue

is unique within the group.

Hydropsyche palawanensis sp. n.

(figs.24-25)

Type material. - Holotyped (in alcohol), Philippi-

nes, Palawan, Puerto Princesa, Irawan river,

22.11.1996, leg. J. Petersen, in MNHB. — Paratypes:

Mey: Hydropsyche West-Pacific region

84,99, same data as holotype ( 24 in coll. Malic-

ky); 2d, 19 (pinned), same locality, 24.- 29.xi.1965,

leg. D. Davis, in National Museum of Natural Histo-

ry, Washington (NMNH); 1d (pinned), Philippines,

Palawan, Litso, Amoyan Ck., 57 km N Puerto

Princesa, 10.XII.1965, in NMNH.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 7-8 mm.

Male genitalia (figs. 24-25): Apical segment of in-

ferior appendages slender and bent. Phallotremal

tongue very long and slender, protruding the lateral

sides of phallothecal tip with half of its length.

Dorsolateral membrane with 2 pairs of appendages,

the smaller with minute spines and the longer with a

bent and serrate spur.

Remarks. — The new species is a close relative of H.

mindorensis and H. cebuensis. The main differences

can be found in the structures of the phallotheca.

Hydropsyche gemellata sp. n.

(figs. 26-27)

Type material. - Holotyped (in alcohol), Indo-

nesia, Sulawesi Selatau, Puncal Palopo, Tanah Tora-

ja, 1300m, 19.viii.1995, leg. S. Naumann, in MNHB.

— Paratypes: 26 , same data as holotype.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 9 mm.

Male genitalia (figs. 26-27): Apical appendages of

segment 10 bent downwards. Second segment of in-

ferior appendages simple, with an acute tip.

Phallotremal tongue longer than the lateral sides of

phallothecal tip, which are characteristically curved

upwards. Dorsolateral membrane with 3 pairs of ap-

pendages, the longer with a long spur apically, the

two shorter ones with minute spines.

Remarks. — The nearest relative is obviously A.

hamifera from the same island. In comparison with

the figures of the type of H. hamifera in Neboiss

(1996: 7) the species differs in the structure of the

phallotheca.

Hydropsyche salki sp. n.

(figs. 28-30)

Type material. - Holotyped (in alcohol), Indo-

nesia, Seram, Piljana, 25.2.1994, leg. P. Salk, in

MNHB. — Paratypes: 22 , same data as holotype; 3d,

Indonesia, Sulawesi Selatan, Puncak, 1000m,

22.11.1994, leg. P. Salk; 473, 32 ‚Indonesia, Bacan

Island, Mt. Sibela, 5-8.11.1996, leg. V. Siniaev (26 in

coll. Malicky); 35, 32, Indonesia, Halmahera, Mt.

Talagaramu, 15 km SE Baru, 600m, 22-31.1.1996,

leg. V. Siniaev.

201

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 33-38. Male genitalia of Hydropsyche spp. — 33-34, H. suppleta sp. n., 33, lateral, 34, ventral; 35-36, H. naumanni sp.

n., 35, lateral, 36, ventral; 37-38, H. sulana sp. n., 37, lateral, 38, ventral.

202

Mey: Hydropsyche West-Pacific region

Figs. 39-40. Male genitalia of Hydropsyche bacanensis sp. n. — 39, lateral, 40, ventral.

Description. — External characters as in A. ger-

manorum sp. n. Length of forewing 8-10 mm.

Male genitalia (figs. 28-30): Segment 9 with a dis-

tinct carina. Dorsal processes of segment 10 very

small or lacking. Apical segment of inferior ap-

pendages sinuate. Phallotremal tongue broad, with a

bifid tip. Dorsolateral membrane with one pair of

long appendages, apically armed with minute spines.

Etymology. — The species is dedicated to Peter

Salk, who collected this species during his voyage in

Indonesia and donated the material to the MNHB.

Remarks. — The species is related to H. sirimauna

sp. n. The dorsolateral appendages of the phallotheca

and the second joint of the inferior appendages are

differently shaped.

Hydropsyche sirimauna sp. n.

(figs. 31-32)

Type material. — Holotype d (in alcohol), Indone-

sia, Ambon, Gunung Sirimau, 500m, 23.vii.1995,

leg. S. Naumann, in MNHB. — Paratypes: 5d, Indo-

nesia, Seram, Piljana, 600m, 26.11.1994, leg. P. Salk.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 7.5 mm.

Male genitalia (figs. 31-31): Dorsal processes on

segment 10 lacking. Apical segment of inferior ap-

pendages sinuate in lateral view. Phallotheca slightly

bent upwards. Phallotremal tongue strongly sclero-

tized, as long as the phallotheca. Dorsolateral mem-

brane with a pair of very long appendages, ending

with a sclerotized and serrate tip.

Remarks. — H. sirimauna sp. n. is related with the

preceding species. Both occur sympatrically on

Seram.

Hydropsyche suppleta sp. n.

(figs. 33-34)

Type material. — Holotype d (in alcohol),

Indonesia, Ambon, Katalai, 470m, 26.vii.1995, leg.

S. Naumann, in MNHB. — Paratypes: 1 d, same data as

holotype; 26 , Indonesia, Ambon, 200m, 24.11.1994,

leg. P. Salk; 16, Indonesia, Seram, Piljana, Tule,

600m, 26.2.1994, leg. P. Salk.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 8 mm.

Male genitalia (figs. 33-34): Segment 10 with both

dorsal and apical appendages. Phallotheca slightly

sinuate. Freely protruding apicolateral sides of phal-

203

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

lotheca reduced. Phallotremal sclerites large, as long

as its tongue. Dorsolateral membrane with two short

lobes, the lateral one with a long and distinctively

bent spine.

Remarks. — The new species is a close relative of A.

naumanni sp. n. from Sulawesi. They differ in the

structure of the dorsolateral appendages.

Hydropsyche naumanni sp. n.

(figs. 35-36)

Type material. — Holotype d (in alcohol), Indo-

nesia, Sulawesi Tengah, Taripa, 700m, 26.8.1995,

leg. S. Naumann, in MNHB. — Paratypes: 2d, same

data as holotype.

Description. — External characters as in A. ger-

manorum sp. n. Length of forewing 8 mm.

Male genitalia (figs. 35-36): Segment 10 with both

dorsal and apical appendages. Phallotheca straight.

Freely protruding apicolateral sides of phallotheca re-

duced, directed dorsally. Phallotremal sclerites large.

Dorsolateral membrane with two pairs of ap-

pendages, the longer with a bent spur, the smaller

with minute spines on the tips.

Etymology. — The species is named in honour of

Stefan Naumann, specialist on Emperor Moths

(Lepidoptera, Saturniidae), who collected the species

on Sulawesi and donated the material to the MNHB.

Remarks. — The species is closely related to H. sup-

pleta sp. n. from Seram and Ambon. Both species can

be separated easily by the different structure of the

phallic apparatus.

Hydropsyche sulana sp. n.

(figs. 37-38)

Type material. — Holotyped (in alcohol), Indo-

nesia, Moluccas, Sula Islands, Sanana, 1 km W Wai-

bau, 250m, 3.viii.1995, leg. S. Naumann, in MNHB. —

Paratypes: 56, 22, same data as holotype.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 8-9 mm.

Male genitalia (figs. 37-38): Segment 10 with dor-

sal and apical appendages. Second segment of inferior

appendages short and slender. Phallotheca slightly

sinuate, apicolateral parts enlarged and bent outward-

ly. Phallotremal tongue reaching just the tip of the

phallotheca. Dorsolateral membrane with two pairs

of appendages, the longer with a terminal spur, the

shorter covered with minute spines.

Remarks. — The new species is related to the next

species, H. bacanensis sp. n.

Hydropsyche bacanensis sp. n.

(figs. 39-40)

Type material. — Holotyped (in alcohol), Indone-

sia, Moluccas, Bacan, Mt. Sibela, 2.- 13.11.1996, leg.

V. Siniaev, in MNHB. — Paratypes: 2d, same data as

holotype.

Description. — External characters as in H. ger-

manorum sp. n. Length of forewing 8.2 mm.

Male genitalia (figs. 39-40): Segment 9 with a well

developed carina. Second segment of inferior ap-

pendages with broad base and slender apical half.

Phallotheca with elongated apical parts, broad and

serrate dorsally. Phallotremal tongue not reaching tip

of phallotheca. Dorsolateral membrane with two

pairs of appendages, the slightly longer with a termi-

nal spur and the shorter with minute spines.

Remarks. — H. bacanensis sp. n. is a close relative to

the preceding species, H. sulana sp. n. The morpho-

logical differences in the phallic apparatus are minor,

but constantly expressed.

ACKNOWLEDGEMENTS

I would like to thank those, who have collected

Hydropsyche material in the West Pacific Region and

donated it to me and to the MNHB: Dr. Eddi Diel,

Axel Kallies, Stefan Naumann, Jan Petersen, Peter

Salk, Alexander Schintlmeister, Sheng-Horn Yen.

The work was supported by a grant from the

Deutsche Forschungsgemeinschaft (Me 1085/5-1).

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205

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

206

JAN PATOCKA

Institut für Waldükologie der SAW, Zvolen

DIE PUPPEN EINIGER MITTELEUROPÄISCHEN

UND KANARISCHEN SCYTHRIDIDAE

(LEPIDOPTERA, GELECHIOIDEA)

Patocka, J., 1998. Die Puppen einiger mitteleuropäischen und kanarischen Scythrididae (Lep-

idoptera, Gelechioidea). — Tijdschrift voor Entomologie 140 [1997]: 207-220, figs. 1-120.

[issN 0040-7496]. Published 26 March 1998.

The pupae of the family Scythrididae and the Central European genus Scythris Hübner are

characterized. A key is provided to 16 species from Central Europe and 2 from the Canary is-

lands; these species are briefly described and some biological data are given.

Institut für Waldökologie der SAW, Stürova 2, 96053 Zvolen, Slovakia.

Key words. — Lepidoptera; Scythrididae; Pupae; Taxonomy; Key to species.

Die Puppen der Familie Scythrididae sind bisher

sehr wenig bekannt. So erwähnt z. B. Mosher (1916)

die Puppen zweier nordamerikanischer Arten und

Bengtsson (1984) einer mittel- bzw. nordeuropäi-

schen Art. In der vorliegenden Arbeit werden insge-

samt 18 Arten behandelt: 16 mitteleuropäischen und

2 von den Kanarische Inseln. Die Puppenexuvien der

letzteren erhielt der Autor von Dr Klimesch ohne

eingehendere Angaben. In Mitteleuropa (als Mit-

teleuropa wird hier Deutschland, die Schweiz, Öster-

reich die Tschechische Republik, Polen, die Slowakei

und Ungarn begriffen) sind von dieser Familie bisher

59 Arten bekannt, welche erst in den zwei letzten De-

zennien eingehender untersucht, taxonomisch auf-

geklärt und verläßlich bearbeitet wurden. Die euro-

päischen Arten werden neuerdings von Bengtsson

(1997) bearbeitet. Die Beziehungen der Familie Scy-

thrididae zu anderen Familien der Oberfamilie Gele-

chioidea, hinsichtlich ihrer Puppenmorphologie,

werden in Patocka (1997, im druck) besprochen.

Die Bionomie und die Nahrungspflanzen der Rau-

pen sind nur lückenhaft bekannt. Manche Arten leben

an Moosen (Bryophyta), die Mehrheit aber an Kräu-

tern aus den Familien Chenopodiaceae, Caryophyl-

laceae, Polygonaceae, Crassulaceae, Fabaceae, Cista-

ceae, Lamiaceae, Ericaceae, Onagraceae, Asteraceae

u.a. Die Raupen leben einzeln oder zu mehreren in

einem Gespinst bzw. Gespinstnest unter oder an ihren

INahrungspflanzen und halten sich oft in einer Ge-

spinströhre auf. Dort findet gewöhnlich auch die Ver-

Puppung in einem dichten, weichen Gespinst statt.

Viele Arten bevorzugen offene und warmtrockene,

andere eher frische bis feuchte Standorte und mehrere

sind Gebirgsbewohner.

Die hier erwähnten Arten gehören alle in die Gat-

tung Scythris Hübner, 1825. Bengtsson (1997) unter-

scheidet noch weitere Gattungen, von denen für Mit-

teleuropa noch Parascythris Hannemann, 1960 und

Enolmis Duponchel, 1845 in Betracht kommen. Von

diesen Gattungen konnte der Autor keine Vertreter

untersuchen.

MATERIAL UND METHODE

Das untersuchte Material stammt aus der Samm-

lung des Verfassers zu der viele Fachkollegen Materi-

al beigetragen haben und ferner aus folgenden Mu-

seen: Museum für Naturkunde Berlin, Zoologische

Staatssammlung München und Naturhistorisches

Museum Wien.

Die Puppen bzw. ihre Exuvien wurden aus ihren

Kokons herausgenommen und mit Hilfe eines Stere-

omikroskopes untersucht, gemessen, beschrieben und

gezeichnet. Das hier benützte System folgt im we-

sentlichen Bengtsson (1984, 1997), Huemer & Tar-

mann (1993) und Karsholt & Razowski (1996). Die

Nomenklatur richtet sich außerdem nach einer Liste

der paläarktischen Scythrididae (aus dem Jahr 1987)

welche B. A. Bengtsson dem Autor liebenswürdiger-

weise zur Verfügung stellte. Die puppenmorphologi-

sche Terminologie (vgl. Abb. 1, 5, 6, 32) folgt

Mosher (1916).

207

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 1-15. — 1-8, Scythris potentillella; 9-15, S. limbella. — 1, 9 Habitusbild in Ventralansicht; 2, 10 Labrum und Umgebung;

3, 11 Palpus maxillaris und Umgebung; 4, 13 abdominales Spiraculum und Umgebung; 5, Vertex und Pronotum (linke

Hälfte); 6, 12, Metanotum und Abdomenbasis (linke Hälfte); 7, 14, Abdomenende in Lateralansicht; 8, 15, Abdomenende

in Dorsalansicht.

A = Antennae, A5 - A10 = 5.-10. Abdominalsegment, An = Analnaht, Af = Analfeld, Cl = Clypeus, F = Frons, Fcs = Fronto-

clypealsutur, G = Genae, Gn = Genitalnaht, Hb = Hinterbeine, Lb = Labium, Lbr = Labrum, Mb = Mittelbeine, Md =

Mandibulae, Msn = Mesonotum, Mtn = Metanotum O = Oculi, Pel = Postclypeus, Pm = Palpi maxillares, Pn = Pronotum,

Pr = Proboscis, Vb = Vorderbeine, Vfl = Vorderflügel, Vx = Vertex .

CHARAKTERISTIK DER PUPPEN DER SCYTHRIDIDAE

Mumienpuppen (pupae obtectae), klein bis mit-

telklein, etwa 4-9 mm lang, mittelschlank (Abb. 16)

bis mäßig gedrungen (Abb. 1), in Ventralansicht

spindel- oder keilförmig, im Frontalteil relativ breit

und stumpf abgerundet. Färbung braun (rot-, ocker-

oder gelbbraun). Skulptur oft mittelfein bis mittel-

grob. Meist findet man unregelmäßige Runzelung

oder Netz- bzw. Zellskulptur. Manchmal ist die

Skulptur dunkler als ihre Umgebung. Am Abdomen

gibt es zuweilen auch Punktgrübchen (Abb. 50). Bor-

sten - auch am Abdomen - oft klein bis mittelklein

(Abb. 15, 88), seltener größer (Abb.21) oder nicht

unterscheidbar (Abb. 27, 58). Nicht selten kommen

auch Sekundarborsten (Abb. 112, 115) vor.

Frontoclypealsutur deutlich (Abb. 1). Labrum mit

208

steileren oder schrägeren Seiten, kaudal abgestumpft

oder abgerundet und kaudal manchmal gewölbt

(Abb. 96) häufiger aber tief manchmal spitz aus-

geschnitten (Abb.2, 30, 43, 96). Labium mit Palpi

labiales gut entwickelt (Abb. 1, 2). Manchmal ist die

Basis des Labiums begrenzt undeutlich (Abb. 43, 48)

oder sein Basalteil von den Palpi labiales abgetrennt

(Abb. 59, 60). Palpi maxillares meist vorhanden und

relativ klein (Abb. 18, 24, 31), selten nicht unter-

scheidbar (Abb. 59, 61). Vorderschenkel (die sich |

sonst zwischen Proboscis und Vorderbeinen befind-

en) dagegen immer verdeckt (Abb. 1). Antennae

nicht selten kürzer als die Vorderflügel (Abb. 1, 59),

aneinander angrenzend (Abb. 42, 47, 77) oder von- |

einander isoliert (Abb. |, 53). Proboscis oft lang (Abb. |

28, 53), manchmal aber auch relativ kurz (Abb. 42). !

Vorderbeine kürzer (Abb. 1), seltener länger als die |

PATOCKA: Puppen der Scythrididae

Abb. 16-29. — 16-21, Scythris knochella; 22-27, S. paullella; 28-29, S. clavella. — 16, 22, 28, Habitusbild inVentralansicht; 17,

23, Labrum und Umgebung; 18, 24, Palpi maxillares und Umgebung; 19, 25, Metanotum und Abdomenbasis (linke Hälfte);

20,26, Abdomenende in Lateralansicht; 21, 27, Abdomenende in Dorsalansicht; 29, Vertex und Pronotum (linke Hälfte).

Mittelbeine (Abb. 28, 107). Hinterbeine verdeckt

(Abb. 28, 47) oder sichtbar, klein (Abb. 1) bzw. gros-

ser (Abb. 22) und selten frontal keilartig verjüngt

(Abb. 77). Vorderflügel voneinander isoliert (Abb. 9,

28, 77) oder länger bzw. kürzer aneinander angren-

zend (Abb. 1, 47, 65). Abdomen der Exuvie in Ven-

tralansicht zum Kaudalrand des 4. (Abb. 28, 107) bis

8. oder Basis des 9. Segmentes (Abb. 59) verdeckt.

Pronotum meist länger und breiter als Vertex

(Abb. 5). Thorakales Spiraculum spaltförmig (Abb.

5). Metanotum gewöhnlich breit und mitteltief aus-

geschnitten und seine Frontallappen mittelbreit und

‚abgerundet (Abb. 6, 67). Hinterflügel meist nur bis

\1/5-34 des 2., selten zur Basis des 3. Abdominalseg-

‚mentes sichtbar (Abb. 32, 56, 112). Abdominale

Spiracula manchmal etwas erhaben (Abb. 39, 113).

Kremaster nicht entwickelt. Abdomenende spitzer

(Abb. 65), oder stumpfer abgerundet (Abb. 115),

abgestumpft (Abb. 58) oder etwas konkav (Abb. 21),

zuweilen mit Höckern oder Lappen an den Seiten

\(Abb. 28, 52, 76), selten stumpf zugespitzt (Abb. 82).

Gattung Scythris Hübner, 1825

Mit der Charakteristik der Familie.

Bestimmungstabelle der Arten

1. Enden der Hinterbeine sichtbar (Abb. 1, 22, 77)

MESE RL RR RE 2. 2

— Enden der Hinterbeine verdeckt (Abb. 16, 28,

ADI Ie AA TIA EIER. OI o 7

2. Enden der Hinterbeine lang und frontal keilartig

verjüngt. Antennae berühren sich miteinander

(GINS) AMAA LY RRL RIM RBB SEN S. fuscoaenea

— Enden der Hinterbeine mittelkurz bis kurz und

frontal nicht keilartig. Antennae voneinander

ISOLER CAD DIE N35) ANNE 3

3. Antennae wesentlich kürzer als die Proboscis. En-

den der Hinterbeine klein und kurz (Abb. 1, 9)...

— Antennae nur etwas kürzer oder gleichlang,

zuweilen auch länger als die Proboscis. Enden der

Hinterbeine mittelklein und länger als breit

(Abb 255 NO) en A PER NE 5)

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 30-46. — 30-34, Scythris clavella, 35-41, S. palustris; 42-46, S. inspersella. — 30, 36, 43, Labrum und Umgebung; 31, 38,

44, Palpus maxillaris und Umgebung; 32, 38, 45, Metanotum und Abdomenbasis (linke Häfte); 33, 40, Abdomenende in

Lateralansicht; 34, 41, Abdomenende in Dorsalansicht; 35, 42, Habitusbild in Ventralansicht; 39, Abdominales Spiraculum

und Umgebung; 46 Pronotum (linke Hälfte). As = abdominales Spiraculum, Hfl = Hinterflügel, Mtn = Metanotum.

Grenze Proboscis/Antennae und Proboscis/Vor-

derflügel etwa gleichlang. Vorderflügel grenzen

aneinander an. Puppe in Ventralansicht vor dem

Abdomenende plôtzlich verjüngt (Abb. 1)

ER E A SC Ae i S. potentillella

Grenze Proboscis/Antennae viel länger als Pro-

boscis/Vorderflügel. Vorderflügel voneinander

isoliert. Abdomenende verjüngt sich allmählich

(ATS SS) een: S. limbella

Antennae etwas langer als die Proboscis (Abb.

116). Grenze Palpi maxillares/Mittelbeine relativ

ne (VANE, MILE) PRE S. fallacella

Antennae etwas kürzer als die Proboscis (Abb. 22,

35). Grenze Palpi maxillares/Mittelbeine kurz

(Abb: DA 7). ERRO 6

Clypeus mit deutlichen Borsten (Abb. 35, 36).

Abdomen vor seinem Ende plötzlich verjüngt mit

etwas konkaven Seiten (Abb. 35) … …. S. palustris

Borsten am Clypeus nicht unterscheidbar (Abb.

23). Abdomenende allmählich verjüngt und seine

Seiten nicht konkav (Abb. 22) … … … S. paullella

210

Proboscis, Antennae und Vorderfliigel beinahe

gleichlang (Abb. 28, 95, 107). Borsten am Ab-

domenende hakenartig (Abb. 100) und meist re-

lativ lang und schlank (Abb. 34, 115) ............... 8

Proboscis, Antennae und Vorderflügel nicht

gleichlang (Abb. 16, 42, 47, 59). Wenn nur die |

Antennae etwas kürzer als die Proboscis und Vor-

derflügel (und oft auch sonst), sind die Borsten am |

Abdomenende kurz, nicht hakenartig (Abb. 71, |

83, 88) oder nicht unterscheidbar (Abb. 58).....10 |

Abdomen der Exuvie ventral bis zum 4. Segment |

verdeckt (Abb. 28, 107). Labrum kaudal deutlich |

ausseschnitten\(Abb 130,109) er. cose 9) |

Abdomen der Exuvie ventral bis zum kaudalteil

des 6.Segmentes verdeckt (Abb. 95). Labrum am |

Ende abgerundet, nicht ausgeschnitten (Abb. 96) |

(Kanarische Inseln) … … S. boseanella |

Abdomenende etwas zwei- oder dreilappig (Abb.

28,33,34). Bei der weiblichen Puppe die Vorder-

beine länger als die Mittelbeine und Antennae |

mit Proboscis gleichlang (Abb. 28) ...... S. clavella

PATOCKA: Puppen der Scythrididae

Abb. 47-60. — 47-52, Scythris noricella; 53-58, S. siccella; 54, 60, S. braschiella. — 47, 53, 59, Habitusbild in Ventralansicht;

48, 54, 60, Labrum und Umgebung; 49, 55, Palpus maxillaris und Umgebung; 50, 56, Metanotum und Abdomenbasis (linke

Hälfte); 51, 57, Abdomenende in Lateralansicht; 52, 58, Abdomenende in Dorsalansicht.

Abdomenende abgerundet (Abb. 107, 108, 114,

115). Bei der weiblichen Puppe die Vorderbeine

kiirzer als die Mittelbeine und Antennae als die

ProDOscisi(AbbAl0 See S. seliniella

. Mittelbeine kürzer als die Vorderbeine (Abb. 42,

47). Abdomen dorsal mit Grübchenskulptur

(ABBASSA Ue ee ERT, es 11

Mittelbeine länger als die Vorderbeine (Abb. 53,

65). Abdomen dorsal mit Runzel- bzw. Netz-

skulptur (Abb. 86, 92), selten mit kaum angedeu-

teten Grübchen (Abb. 56)... 12

. Antennae grenzen aneinander etwa so lang wie

die Vorderflügel an. Proboscis relativ lang (Abb.

BON RS EE lot S. inspersella

Antennae grenzen aneinander viel länger als die

Vorderflügel an. Proboscis relativ kurz (Abb. 47)

SE CR NLA DIANO TOLLER S. noricella

. Antennae kürzer als die Mittelbeine (Abb. 59).

Palpi maxillares nicht unterscheidbar (Abb. 61)...

oe S. braschiella

Antennae länger als die Mittelbeine (Abb. 65,

11538

71). Palpi maxillares unterscheidbar (Abb. 68,

74)

Grenze Antennae/Proboscis kürzer als Probos-

is Mordertlüge {ADD ES 3 MOD) PRES 14

Grenze Antennae/Proboscis länger als Probos-

eis/Morderflügeli(ADDR83 139) KATE En 15

. Labrum ziemlich tief und spitz ausgeschnitten

(Abb. 102). Palpi maxillares relativ groß (Abb.

103). Grenze Mittelbeine/Proboscis viel kürzer

als die Antennae/Proboscis (Abb. 101) (Ka-

nantscheilnseln) ee nee S. klimeschi

Labrum seicht und abgerundet ausgeschnitten

oder fast gerade(Abb. 54). Palpi maxillares klein

(Abb. 55). Grenze Mittelbeine/Proboscis wenig

kürzer als Antennae/Proboscis (Abb. 53).............

Di EEN EN S. siccella

. Grenze Antennae/Proboscis kaum oder bis etwa

2X länger als Proboscis/Vorderflügel (Abb. 16,

SEN NEE AIR 16

Grenze Antennae/Proboscis mehr als 4X länger

als Proboscis/Vorderflügel (Abb. 65, 83)......... 117

211

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 61-76. — 61-64, Scythris braschiella; 65-70, S. crassiuscula; 71-76, S. dissimilella. — 61, 68, 73, Palpus maxillaris, bzw.

Oculus und Umgebung; 62, 67, 74, Metanotum und Abdomenbasis (linke Hälfte); 63, 69, 75, Abdomenende in Lateralan-

sicht; 64, 70, 76, Abdomenende in Dorsalansicht; 65, 71, Habitusbild in Ventralansicht; 66, 72, Labrum und Umgebung.

16. Vorderfliigel grenzen aneinander an. Abdomen-

ende spitzer abgerundet (Abb. 89). Labrum seicht

und abgerundet ausgeschnitten (Abb. 90)

PRA A REA AAA A LEA Ea LES S. bengtssoni

Vorderflügel voneinander isoliert. Abdomenende

eher stumpf (Abb. 16). Labrum spitz ausge-

schnitten (Abb RIVER S. knochella

Abdomenende in Ventralansicht mit je einem

Seitenlappen. Proboscis vor dem Endteil etwas

verschmälert (Abb. ZW): S. dissimilella

Abdomenende in Ventralansicht ohne Seitenlap-

pen. Proboscis vor dem Endteil nicht verschmä-

lei Ab b), (E535, (9333) OR EE 18

Vorderflügel grenzen kurz aneinander an. Abdo-

menende eher spitz (Abb. 65). Labrum tief und

spitz augeschnitten| (Abb REI O

SONI ARE NO RARE SE A A e ARE S. crassiuscula

Vorderflügel voneinander isoliert. Abdomenende

eher stumpf (Abb. 83). Labrum seicht ausge-

ScANIttenNI(AbbASA) ILA S. hungaricella

18.

212

Scythris potentillella (Zeller, 1847)

(Abb. 1-8)

Untersuchtes Material. — | Männchen, Deutschland.

Beschreibung. — Puppe 5,4X2 mm, relativ gedrun- |

gen, Abdomen zwischen dem 8.-9. Segment plötzlich

verjüngt. Färbung dunkel goldbraun, Exuvie orange-

braun, schwach glänzend. Skulptur überall deutlich, |

unregelmäßig runzelig-zellartig. Borsten steif, auch |

am Clypeus deutlich, am Abdomenende hakenartig.

Labrum kaudal tief ausgeschnitten. Labium basal |

deutlich begrenzt und wenig erweitert. Palpi maxilla- |

res ziemlich groß und viereckig. Grenze Palpi maxil- |

lares/Oculi kaum länger als Antennae/Palpi maxilla- |

res und Vorderbeine/Genae kurz, nicht länger als |

Palpi maxillares/Mittelbeine. Grenze Mittelbeine

/Proboscis fast halb so lang wie Proboscis/Antennae. |

Vorderflügel grenzen kurz, aber deutlich aneinander

an. Hinterbeine unterscheidbar, recht klein. 7. Abdo- |

minalsegment der Exuvie ventral fast verdeckt und

seine grenze mit dem 8. Segment geschwungen. Pro-

notum viel länger als Vertex, Mesonotum frontal

längsgerippt. Thorakales Spiraculum kurz spaltför-

mig. Metanotum recht breit abgerundet und tief aus-

geschnitten. Abdominale Spiracula breit elliptisch,

wenig erhaben, durch konzentrische Ringelung ge-

säumt. Analfeld mit je einem Eindruck.

Perianalhäkchen vorhanden. Abdomenende abge-

rundet, in Lateralansicht im Dorsalumriß abge-

schrägt und in Dorsalansicht mit zwei Querreihen

von Borsten.

Lebensweise. — Raupe an Acetosella vulgaris (Koch)

Fourr. (= Rumex acetosella) in einer Gespinströhre

entlang des Stengels. Diese Art bevorzugt sandige

Standorte, auch im Gebirge.

Scythris limbella (Fabricius, 1775)

(Abb. 9-15)

Untersuchtes Material. — 1 Weibchen, Deutschland.

Beschreibung. — Puppe 6,3X2,0 mm, mäßig

gedrungen. Abdomen am 8.-9. Segment nicht auffal-

lend verjüngt. Färbung rotbraun, Exuvie hell rotgelb-

braun, etwas glänzend. Skulptur stark und unregel-

mäßig runzelig-zellartig. Borsten auch am Clypeus

deutlich. Morphologisch S. potentillella ähnlich, aber

die Grenze Antennae/Proboscis viel länger als Mittel-

beine/Proboscis oder Proboscis/Vorderflügel. Vor-

derflügel voneinander isoliert. Enden der Hinterbei-

ne sichtbar und winzig. Labrum stark ausgeschnitten.

Grenze Palpi maxillares/Mittelbeine relativ kürzer als

bei S. potentillella. Abdomen der Exuvie ventral bis

zum Kaudalrand des 7. Segmentes verdeckt. Aus-

schnitt des Metanotums relativ schmaler. Abdomi-

nale Spiracula dunkel, kreisförmig und kaum erha-

ben. Abdomenende in Lateralansicht regelmäßig

abgerundet, in Dorsalansicht mit zwei Querreihen

steifer Borsten.

Lebensweise. — Raupe gruppenweise in einem

Gespinst an Blättern und Blüten von Chenopodium

und Atriplex spp. Ruderale und öde Standorte.

Scythris knochella (Fabricius, 1794)

(Abb. 16-21)

Untersuchtes Material. — 1 Männchen, 1 Weibchen,

Deutschland.

Beschreibung. — Puppe 5,1-5,4X 1,4-1,6 mm, rot-

braun, Exuvie heller. Skulptur deutlich, unregel-

mäßig runzelig, Borsten ebenfalls deutlich, aber am

Kopf nicht unterscheidbar. Labrum kaudal stark aus-

|geschnitten und Labium an der Basis deutlicher er-

weitert als bei den vorgehenden Arten. Palpi maxil-

lares eher dreieckig, ihre Grenze mit den Mittelbeinen

kurz. Grenze Genae/Vorderbeine ebenfalls kurz. Vor-

PATOCKA: Puppen der Scythrididae

derbeine und die Grenze Mittelbeine/Proboscis auch

nicht lang. Grenze Proboscis /Antennae etwa 2X

länger als Proboscis/Vorderflügel. Hinterbeine nicht

sichtbar. Vorderflügel voneinander isoliert. Abdomen

der Exuvie ventral bis etwa zu % des 7. Segmentes

verdeckt. Auschnitt des Metanotums mit eher schrä-

gen Seiten. Abdominale Spiracula kreisrund und we-

nig erhaben. Abdomenende stumpf oder etwas aus-

geschnitten und mit starken Borsten versehen.

Lebensweise. — Noch nicht ausreichend bekannt.

Als Nahrungspflanze der Raupe wird Cerastium, aber

auch Thymus spp. angegeben. Trockene offene Stan-

dorte werden bevorzugt.

Scythris paullella (Herrich-Schäffer, 1855)

(Abb. 22-27)

Untersuchtes Material. — | Männchen, 3 Weibchen,

Deutschland.

Beschreibung. — Puppe 4,2-5,3X1,3-1,6 mm, hell-

braun, Exuvie bleich bräunlichgelb. Skulptur un-

regelmäßig runzelig bis zellartig. Borsten am Kopf

kaum unterscheidbar und auch am Abdomen klein.

Labrum ähnlich wie bei den vorherigen Arten, kaudal

ausgeschnitten. Labium relativ klein, an der Basis

mäßig verbreitet. Palpi maxillares klein, fast dreieck-

ig. Ihre Grenze mit den Mittelbeinen und auch Vor-

derbeine/Genae sehr kurz. Proboscis lang und

berührt die relativ großen Enden der Hinterbeine.

Seine Grenze mit den Mittelbeinen und Vorderflü-

geln kurz, mit den Antennae sehr lang. Vorderflügel

voneinander isoliert. Abdomen der Exuvie ventral

fast bis zum Kaudalrand des 7. Segmentes verdeckt.

Pronotum relativ kurz, aber deutlich länger als Ver-

tex. Thorakales Spiraculum spaltförmig und schmal.

Metanotum etwa bis zu % seiner Länge breit ausge-

schnitten. Abdominale Spiracula klein und deutlich

erhaben. Abdomenende stumpf, in Lateralansicht

sein Ventralumriß abgeschrägt. Die männliche Pup-

pe mit einer Erhabenheit ventral am 8. Abdominal-

segment, diese fehlt bei der weiblichen.

Lebensweise. — Raupe an Moosen (Polytrichum

spp.) in feinem Gespinstnest. Lokal an felsigen,

buschigen und ziemlich sonnigen Standorten ebenso

an alkalischen, wie auch sauren Substrat.

Scythris clavella (Zeller, 1855)

(Abb. 28-34)

Untersuchtes Material. — 1 Weibchen, Böhmen (Bo-

hemia).

Beschreibung. — Puppe 6,5X1,9 mm, rotbraun,

Exuvie heller. Skulptur relativ grob, scharf dunkel,

netzartig. Borsten deutlich, auch am Clypeus unter-

215

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 77-90. — 77-82, S. fuscoaenea; 83-88, S. hungaricella; 89-90, S. bengtssoni. — 77, 83, 89, Habitusbild in Ventralansicht;

78, 84, 90, Labrum und Umgebung; 79, 85, Palpus maxillaris und Umgebung; 80, 86, Metanotum und Abdomenbasis (linke

Hälfte); 81, 87, Abdomenende in Lateralansicht; 82, 88, Abdomenende in Dorsalansicht.

scheidbar, am Pronotum deutlicher, am Abdomen

mittelklein. Hakchen am Abdomenende und eben-

falls die Perianalhäkchen lang und schlank hakenför-

mig. Auch zahlreiche überzählige (sekundäre) Bor-

sten vorhanden. Clypeus rauh skulptiert und etwas

höckerig. Labrum abgerundet trapezförmig mit

schrägen Seiten und am Kaudalende mittelseicht aus-

geschnitten. Labium lang und an der Basis mäßig er-

weitert. Palpi maxillares mittelgroß, trapezförmig, an

die Mittelbeine ziemlich lang angrenzend. Auch die

Grenze Genae/Vorderbeine länger als bei S. paullella.

Mittelbeine etwas kürzer als die Vorderbeine. Pro-

boscis grenzt an die Antennae lang an. Vorderflügel,

Antennae und Proboscis etwa gleichlang. Vorder-

flügel voneinander isoliert. Abdomen der Exuvie in

Ventralansicht nur bis zum Kaudalrand des 4. Seg-

mentes verdeckt. Am 5. und 6. Segment sind Spuren

der Abdominalbeine unterscheidbar. Pronotum rela-

tiv breit mit geschwungenem Kaudalrand. Thorakale

Spiracula schmal und unauffällig. Metanotum mittel-

breit und etwa zu Va seiner Länge ausgeschnitten.

Hinterflügel bis zur Basis des 3. Abdominalsegmentes

214

sichtbar. Abdominale Spiracula etwas erhaben. Ab-

domenende - auch in Lateralansicht - etwas eckig

bzw. lappig und im Mittelteil konkav, in Dorsalan-

sicht mit einem kleinen Kaudalhöcker. Analfeld und

seine Umgebung ziemlich rauh.

Lebensweise. — Nahrungspflanze nach Bengtsson

(1997) ist Helianthemum nummularium (L.) Miller.

Die hier beschriebene Puppe wurde von eingetragene

Teucrium chamaedris L. erzogen (persönliche Mit-

teilung von Dr. Vävra, Prag). Die Imago bevorzugt |

warmtrockene Standorte.

Scythris seliniella (Zeller, 1839)

(Abb. 107-115)

Untersuchtes Material. — 1 Männchen, 1 Weibchen,

Polen.

Beschreibung. — Puppe 5,6-6,2X1,6-1,8 mm rot,

bei dem Weibchen eher ockerbraun, Exuvie heller. |

Skulptur relativ grob, scharf und dunkel runzelig-net-

zartig. Borsten deutlich (auch am Clypeus unterschei-

dbar), fein, aus dunkleren Pinnacula entspringend.

Auch zahlreiche sekundäre Borsten vorhanden. Am

10.Abdominalsegment sind die End- und Perianal-

borsten recht zahlreich, lang und schlank, hakenför-

mig. Morphologisch S. clavella ähnlich. Abdome-

nende bei beiden Geschlechtern abgerundet und

nicht lappig. Bei dem Männchen Antennae, Pro-

boscis und Vorderflügel gleichlang und Mittelbeine

kürzer als die Vorderbeine. Bei dem Weibchen An-

tennae etwas kürzer als die Proboscis und Vorder-

flügel; Vorderbeine kürzer als die Mittelbeine.

Lebensweise. — Raupe lebt nach der Mitteilung von

T. Baran (Torun) an Moosen (Musci spp.), aber auch

an Artemisia campestris L. (T. Baran in Vorbereitung).

Auch diese Art bevorzugt warmtrockene und sonnige

Standorte.

Scythris palustris (Zeller, 1855)

(Abb. 35-41)

Untersuchtes Material. — 1 Männchen, 1 Weibchen,

Deutschland.

Beschreibung. — Puppe 4,2-4,6X1,2-1,4 mm,

ockerbraun, Exuvie hell bräunlich ockergelb und

mäßig glänzend. Skulptur scharf und unregelmäßig

runzelig. Borsten klein und steif, auch am Clypeus

deutlich. Morphologisch S. paullella ähnlich. Ab-

domenende am 8. und 9. Segment auffallender ver-

jüngt und dort in Ventralansicht sein Umriß etwas

konkav. Palpi maxillares relativ etwas großer und ihre

Grenze mit den Mittelbeinen länger. Grenze Ocu-

li/Vorderbeine kürzer als bei S. paullella. 7. Abdomi-

nalsegment der Exuvie ventral praktisch bis zum Kau-

dalrand verdeckt. Abdomenende mehr abgerundet

und in Lateralansicht sein Dorsalumriß abgeschrägt.

Lebensweise. — Raupe an Moos (Rhytidiadelphus

sp.) in Gespinströhren. An feuchten und offenen

Standorten.

Scythris inspersella (Hübner, 1817)

(Abb. 42-46)

Untersuchtes Material. — 1 Männchen, Norwegen (Ab-

domenende beschädigt).

Beschreibung. — Puppe 5,5X1,8 mm, rotbraun,

Exuvie hell rötlich braun. Skulptur fein: Am Kopf

und Thorax feine quere und wirre Runzeln, am 1.-8.

Abdominalsegment winzige Punktgrübchen. Borsten

sehr klein. Labrum trapezförmig und kaudal nicht

ausgeschnitten. Postclypeus stärker quergerunzelt.

Labium mittelgroß, seine Basis erweitert und un-

scharf begrenzt. Palpi maxillares klein, viereckig,

seine Grenze mit den Mittelbeinen ziemlich lang.

Grenze Genae/Vorderbeine kurz. Vorderbeine deut-

PATOCKA: Puppen der Scythrididae

lich länger als die Mittelbeine, Antennae länger als die

Proboscis, aber kürzer als die Vorderflügel. Antennae

und Vorderflügel grenzen etwa gleichlang aneinander

an. Hinterbeine nicht sichtbar. 7. Abdominalsegment

der Exuvie ventral bis auf einen schmalen Kaudal-

streifen verdeckt. Pronotum mittelschmal, sein Kau-

dalrand geschwungen. Thorakales Spiraculum spalt-

förmig, dunkel. Metanotum nur etwa zu 1/3 seiner

Länge, breit und abgerundet ausgeschnitten. Sicht-

barer Teil der Hinterflügel überragt wenig die Höhe

der Spiracula am 2. Abdominalsegment. Abdominale

Spiracula mäßig erhaben. Abdomen zum Ende all-

mählich verjüngt.

Lebensweise. — Raupe an Chamerion angustifolium

(L.) Holub, oft gruppenweise, in versponnenen Trieb-

spitzen. Verpuppungskokon weißlich. An frischen

Waldlichtungen, um Waldwege usw.

Scythris noricella (Zeller, 1843)

(Abb. 47-52)

Untersuchtes Material. — 5 Männchen, 5 Weibchen,

Böhmen (Bohemia).

Beschreibung. — Puppe 7-8,5X2,3-2,6 mm, rot-

braun (Exuvie heller), mäßig glänzend. Skulptur mit-

telfein bis mittelgrob, am Kopf und Thorax un-

regelmäßig gerunzelt, am 1.-8. Abdominalsegment

scharfe und dichte Punktgrübchen. Borsten winzig,

am 8. und 9. Abdominalsegment etwas großer. La-

brum trapezförmig, seine Basis, wie auch Postclypeus

stark gefurcht. Labium mittelklein, basal etwas er-

weitert und unscharf begrenzt. Palpi maxillares mit-

telklein, fast dreieckig, ihre Grenze mit den Mittel-

beinen sehr kurz. Grenze Genae/Vorderbeine relativ

länger. Vorderbeine deutlich länger als die Mittel-

beine. Proboscis relativ kurz. Antennae grenzen an-

einander deutlich länger als die Vorderflügel und als

Antennae an die Proboscis an. 7. Abdominalsegment

der Exuvie ventral nur etwa zur Mitte seiner Länge

verdeckt. Pronotum mittelkurz, thorakales Spiracu-

lum auffällig und groß. Metanotum kaum zu Vs sein-

er Länge, breit und abgerundet ausgeschnitten. Hin-

terflügel etwa zu % des 2. Abdominalsegmentes

sichtbar. Abdominale Spiracula deutlich erhaben. 8-

10. Abdominalsegment unscharf voneinander abge-

grenzt. Abdomenende abgestumpft, nur in Lateral-

ansicht abgerundet. Analfeld mit Eindrücken.

Borsten am Abdomenende kaum unterscheidbar.

Lebensweise. — Raupe an Chamerion angustifolium

(L.) Holub, oft gruppenweise, in versponnenen

Triebspitzen. Verpuppungskokon weißlich. Mehr in

Gebirge, an ähnlichen Standorten wie S. inspersella.

215

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 91-106. — 91-94, S. bengtssont; 95-100, S. boseanella; 101-106, S. klimeschi. — 91, 97, 103, Palpus maxillaris und Umge-

bung; 92, 98, 104, Metanotum und Abdomenbasis (linke Hälfte), 93, 99, 105, Abdomenende in Lateralansicht; 94, 100,

106, Abdomenende in Dorsalansicht; 95, 101, Habitusbild in Ventralansicht; 96, 102, Labrum und Umgebung.

Scythris siccella (Zeller, 1839)

(Abb. 53-58)

Untersuchtes Material. — 1 Weibchen, Deutschland.

Beschreibung. — Puppe 4,3X1,2 mm, relativ

schlank. Färbung rötlich ockerbraun, Exuvie heller.

Skulptur fein gerunzelt, Abdomen dorsal papillos

skulptiert. Borsten winzig, am Kopf und Abdome-

nende nicht unterscheidbar. Labrum auffallend breit,

kaudal ein wenig ausgeschnitten. Mandibulae deut-

lich. Labium groß, an der Basis scharf begrenzt und

wenig verbreitet. Palpi maxillares klein und dreieckig,

unscharf begrenzt. Vorderbeine grenzen lang an die

Basis der Antennae an. Oculi schwach abgegrenzt.

Vorderbeine kürzer als die Mittelbeine, diese kürzer

als die Antennae, alle aber viel kürzer als die Pro-

boscis, diese gleichlang wie die Vorderflügel. Ab-

domen der Exuvie ventral bis zum Kaudalrand des 7.

Segmentes verdeckt. Thorakale Spiracula wenig auf-

fällig, spaltförmig. Metanotum etwa zu Va seiner

Länge stumpf ausgeschnitten, Ausschnitt mit schrä-

216

gen Seiten. Hinterflügel nur bis zur Basis des |. Abdo-

minalsegmentes sichtbar. Abdominale Spiracula fast

kreisrund und wenig erhaben. Abdomenende stumpf, |

in Lateralansicht stumpf abgerundet. Analnaht lang,

Analfeld etwas uneben.

Lebensweise. — Raupe in einer Gespinströhre an der

Basis der niedrig wachsenden Kräuter (wahrscheinlich

polyphag) an offenen und sonnigen Stellen.

Scythris braschiella (Hofmann, 1897)

(Abb. 59-64)

Untersuchtes Material:

Deutschland.

3 Männchen, 3 Weibchen,

Beschreibung. — Puppe 4,2-4,6X 1,0-1,4 mm, mit-

telschlank, dorsoventral abgeflacht. Färbung braun,

Exuvie bleich gelbbraun. Skulptur aus feinen Run- |

zeln. Borsten klein, am Kopf und Abdomenende |

nicht unterscheidbar. Labrum breit und abgerundet,

kaudal spitz ausgeschnitten. Labium klein, Basis der |

Palpi labiales verdeckt. Palpi maxillares nicht unter-

scheidbar, auch Oculi kaum abgegrenzt. Vorder-,

Mittelbeine und Antennae kurz. Proboscis so lang

wie die Vorderflügel und sehr lang an diese angren-

zend. Abdomen der Exuvie ventral bis zur Basis des 9.

Segmentes verdeckt. Metanotum zu 2 seiner Länge,

breit und abgerundet ausgeschnitten. Hinterflügel

überragen nur kurz den Kaudalrand des |. Abdomi-

nalsegmentes. Abdomenende stumpf, in Lateralan-

sicht spitzer abgerundet. Analfeld quergerunzelt.

Lebensweise. — Raupe lebt an Armeria maritima

Miller (Willd.). An sandigen Standorten.

Scythris crassiuscula (Herrich-Schäffer, 1855)

(Abb. 65-70)

Untersuchtes Material. — 1 Weibchen, Mähren (Moravia).

Beschreibung. — Puppe 4,1 X 1,4 mm, eher gedrun-

gen. Färbung rotbraun, Exuvie rötlich braungelb,

glänzend. Skulptur mittelgrob, runzelig zellartig, ihre

Elemente etwas dunkler als die Umgebung. Borsten

deutlich, am Kopf nicht unterscheidbar, am Abdo-

menende fein hakenartig; auch die Perianalhäkchen

deutlich. Labrum mit konvergierenden Seiten, am

Ende tief und spitz ausgeschnitten. Labium relativ

lang, Basis erweitert. Palpi maxillares mittelklein,

viereckig, ihre Grenze mit den Mittelbeinen deutlich.

Grenze Genae/Vorderbeine mittelkurz. Vorderbeine

kürzer als die Mittelbeine, Proboscis länger als die

Antennae. Vorderflügel grenzen kurz aneinander,

Proboscis lang an die Antennae an. Abdomen der Fx-

uvie ventral bis zu % des 6. Segmentes verdeckt. Me-

tanotum zu V seiner Länge, mittelbreit und abgerun-

det ausgeschnitten. Abdominale Spiracula klein und

wenig erhaben. Abdomenende relativ spitz.

Lebensweise. — Raupe an Helianthemum spp.

Trockenwarme, offene Standorte; mit Vorliebe am

Kalksubstrat.

Scythris dissimilella (Herrich-Schäffer, 1855)

(Abb. 71-76)

Untersuchtes Material. — 3 Weibchen, Böhmen (Bo-

hemia).

Beschreibung. — Puppe eher schlank, 6,1-6,3 X 1,6-

1,8 mm. Farbung orangebraun, Exuvie heller. Skulp-

tur mittelfein, scharf und unregelmäßig runzelig.

Borsten winzig und auch am Abdomenende klein, am

Kopf nicht unterscheidbar. Labrum mit relativ steilen

Seiten, kaudal mitteltief und spitz ausgeschnitten.

| Mandibulae relativ deutlich. Labium mittelklein,

frontal mäßig erweitert, an der Basis unscharf begren-

zt. Palpi maxillares klein und dreieckig, die Mittel-

beine fast nur punktartig berührend. Auch ihre Gren-

PATOCKA: Puppen der Scythrididae

ze mit Oculi relativ kurz, ähnlich wie die Grenze

Genae/Vorderbeine. Vorderbeine kürzer als die Mit-

telbeine. Proboscis deutlich länger als die Antennae

und fast so lang wie die Vorderflügel, diese ganz kurz

aneinander angrenzend. Grenze Proboscis/Mittel-

beine und Proboscis/Vorderflügel annähernd gleich-

lang. Proboscis am Ende etwas erweitert zwischen

den Enden der Antennae eher verschmälert. Ab-

domen der Exuvie ventral etwa zu 2/3 des 7. Seg-

mentes verdeckt. Metanotum frontal bis % seiner

länge ausgeschnitten, Ausschnitt abgerundet mit

schrägen Seiten. Hinterflügel bis kaudal der Höhe der

Spiracula am 2. Abdominalsegment sichtbar. Ab-

dominale Spiracula klein und wenig erhaben. Ab-

domenende in Ventral- und Dorsalansicht mit ecki-

gen, kaudal gerichteten Seitenausläufern, in

Lateralansicht ziemlich spitz.

Lebensweise. — Raupe an Helianthemum spp.; an

ahnlichen Standorten wie die vorherige.

Scythris fuscoaenea (Haworth, 1828)

(Abb. 77-82)

Untersuchtes Material. — 1 Männchen, 1 Weibchen,

Böhmen (Bohemia).

Beschreibung. — Puppe 5,9-6,3X1,4-1,6 mm,

honigbraun, Exuvie heller. Skulptur mittelfein vor-

wiegend länglich, aber auch quer und unregelmäßig

gerunzelt. Borsten klein. Labrum mittelgroß, nahe

der Basis mäßig eingeschnürt. Palpi maxillares klein

und viereckig, ihre Grenze mit den Mittelbeinen kurz

und die mit den Antennae am längsten. Die spitze

Proboscis länger als die Mittelbeine. Antennae gren-

zen mittelkurz aneinander an und sind etwa so lang

wie die Vorderflügel. Enden der Hinterbeine groß

und frontal keilförmig verjüngt. Abdomen der Exuvie

ventral fast bis zum Kaudalrand des 7. Segmentes

verdeckt. Pronotum relativ lang und hinten

geschwungen begrenzt. Thorakales Spiraculum ziem-

lich auffällig, ihre Umgebung gerunzelt. Metanotum

etwa zu Vs seiner länge, ziemlich breit und abgerundet

ausgeschnitten. Hinterflügel über 2 des 2. Abdomi-

nalsegmentes sichtbar. Abdominale Spiracula er-

haben. Abdomenende stumpf zugespitzt mit deut-

lichen feinen Häkchen, in Lateral- ziemlich ähnlich

wie in Dorsalansicht.

Lebensweise. — Raupe an Helianthemum spp. An

trockenwarmen, offenen Standorten; mit Vorliebe

am Kalksubstrat.

Scythris hungaricella Rebel, 1917

(Abb. 83-88)

Untersuchtes Material. — 1 Männchen, 1 Weibchen,

Slowakei.

217

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Abb. 107-120. — 107-115, Scythris seliniella; 116-120, S. fallacella. — 107, 108, 116, Habitusbild in Ventralansicht (107, 116

Männchen, 108, Weibchen); 109, 117, Labrum und Umgebung; 110, 118, Palpus maxillaris und Umgebung; 111, Vertex

und Pronotum (linke Hälfte); 112, Metanotum und Abdomenbasis (linke Hälfte); 113, abdominales Spiraculum und Umge-

bung; 114, 119, Abdomenende in Lateralansicht; 115, 120, Abdomenende in Dorsalansicht.

Beschreibung. — Puppe 6,0-7,2X2,0-2,2 mm, mit-

telschlank, honigbraun, Exuvie heller. Skulptur mit-

telfein, unregelmäßig runzelig-zellartig. Borsten, auch

am Abdomenende, winzig. Labrum trapezförmig,

kaudal kaum ausgeschnitten. Labium mittelgroß,

seine Basis stark erweitert und unscharf begrenzt. Pal-

pi maxillares klein, denen bei der vorherigen Art ähn-

lich. Grenze Genae/Vorderbeine mittelkurz. Mittel-

beine etwas länger als die Vorderbeine, Proboscis

länger als die Antennae, ist am Ende zugespitzt und

so lang wie die Vorderflügel. Abdomen der Exuvie

ventral bis zum Kaudalrand des 7. Segmentes ver-

deckt. Pronotum mittellang, viel länger als Vertex.

Thorakales Spiraculum kurz und spaltförmig, seine

Umgebung gerunzelt. Metanotum etwa zu Va seiner

Länge ausgeschnitten, Auschnitt abgerundet mit

schrägen Seiten. Abdominale Spiracula deutlich, aber

kaum erhaben. Abdomenende, auch in Lateralan-

sicht, relativ breit abgerundet.

Lebensweise. — Raupe in einem Gespinstnest an

der Basis der Polster von Thymus spp. An trocken-

218

warmen und sonnigen Standorten. Substrat: junge,

effusive Gesteine oder Kalkstein.

Scythris bengtssoni Patocka & Liska, 1989

(Abb. 89-94)

Untersuchtes Material. — | Männchen, Slowakei.

Beschreibung. — Puppe 5,5X1,7 mm, mäßig |

gedrungen, rotbraun, Exuvie hell rötlich braun.

Skulptur mittelfein, vorwiegend querrunzelig-zellar-

tig, am Clypeus, Frons und Vertex stärker. Borsten |

klein, aber deutlich, auch am Clypeus unterscheid- |

bar. Labrum trapezförmig, seine Seiten etwas

geschwungen, kaudal ganz schwach konkav. Labium

mittelgroß, an der Basis mäßig erweitert und unscharf |

begrenzt. Palpi maxillares mittelklein, ihre Grenze

mit den Mittelbeinen relativ lang. Vorderbeine mäßig

kürzer als die Mittelbeine, die spitze Proboscis auch |

kürzer als die Vorderflügel. Antennae wesentlich

kürzer als die Proboscis. Vorderflügel grenzen kurz

aneinander an. Abdomen der Exuvie ventral zu mehr

als 2/3 des 7. Segmentes verdeckt. Pronotum viel

länger als Vertex, kaudal wenig geschwungen begren-

zt. Thorakales Spiraculum kurz und dunkel. Metan-

otum etwa zu ¥2 seiner Länge, breit und abgerundet

ausgeschnitten. Hinterflügel etwa zur Höhe der

Spiracula des 2. Abdominalsegmentes sichtbar. Ab-

dominale Spiracula deutlich erhaben und dunkel.

Abdomenende ziemlich zugespitzt, kaudal abges-

tumpft und abgerundet (auch in Lateralansicht).

Lebensweise. — Raupe durch nächtliches Streifen

gewonnen, fraß bis zur Verpuppung 7hymus spp.

(persönliche Mitteilung von Ing. Turcani). An

warmtrockenen, steppenartigen Standorten.Substrat:

junge effusive Gesteine oder Kalkstein.

Scythris fallacella (Schläger, 1847)

(Abb. 116-120)

Untersuchtes Material. — 2 Männchen, 1 Weibchen,

Deutschland.

Beschreibung. — Puppe 4,0-5,0X 1,4-1,7 mm. Fär-

bung rötlich braun, Exuvie heller, mäßig glänzend.

Skulptur fein, unregelmäßig runzelig-netzartig. 10.

Abdominalsegment lateral mit Mikrobörstchen an

Papillen. Borsten, auch am Clypeus, deutlich, am Ab-

domenende steif. Labrum abgerundet mit steilen

Seiten, kaudal spitz und tief ausgeschnitten. Labium

mittelklein, an der Basis wenig erweitert und überall

scharf begrenzt. Palpi maxillares mittelklein und

viereckig, grenzen an die Mittelbeine relativ lang an.

Grenze Genae/Vorderbeine ebenfalls relativ deutlich.

Vorder- und die etwas längeren Hinterbeine ziemlich

lang. Antennae und Proboscis fast so lang wie die

Vorderflügel, die mittelkleinen Enden der Hinter-

beine berührend. Antennae und Vorderflügel

voneinander isoliert, Antennae grenzen an die Pro-

boscis lang an. Abdomen der Exuvie ventral bis zum

Kaudalrand des 7. Segmentes verdeckt. Pronotum

breit, sein Kaudalrand geschwungen. Thorakales

Spiraculum schmal und spaltförmig. Metanotum

etwa zu ¥2 seiner Länge ausgeschnitten, dieser Auss-

chnitt abgerundet mit schrägen Seiten. Hinterflügel

nur bis zur Basis des 2. Abdominalsegmentes sicht-

bar. Abdominale Spiracula deutlich erhaben. Ab-

domenende stumpf und kaudal etwas ausgeschnitten,

in Lateralansicht sein Dorsalumriß geschwungen.

Lebensweise. — Raupe an Helianthemum spp. Diese

Art bevorzugt offene felsige und sonnige Gebirgsstan-

dorte am Kalksubstrat, manchmal aber auch in tiefer-

en Lagen.

PATOCKA: Puppen der Scythrididae

Kanarische Arten

Scythris boseanella Klimesch, 1986

(Abb. 95-100)

Untersuchtes Material: 3 Männchen, 3 Weibchen. Ka-

narische Inseln.

Beschreibung. — Puppe 7,5-8,5X2,3-2,6 mm. Sie

istam Kaudalende relativ spitz abgerundet, rotbraun,

Exuvie heller gefärbt. Skulptur mittelgrob runzelig-

zellartig und dunkel. Borsten klein, am Kopf kaum

unterscheidbar. Labrum kaudal gewölbt und

abgerundet, hinten nicht ausgeschnitten. Labium

mittelgroß, basal scharf begrenzt und nicht erweitert.

Palpi maxillares klein und eckig, ihre Grenze mit den

Mittelbeinen kurz. Grenze Genae/Vorderbeine meist

punktartig. Mittelbeine länger als die Vorderbeine,

beide kurz. Proboscis so lang wie die Vorderflügel

und die Antennae nur wenig kürzer. Abdomen der

Exuvie ventral fast zum Ende des 6. Segmentes

verdeckt. Pronotum relativ breit, sein Kaudalrand

geschwungen. Thorakales Spiraculum kurz spaltför-

mig. Metanotum zu V seiner länge und abgerundet

ausgeschnitten. Hinterflügel zur Basis des 3. Abdom-

inalsegmentes sichtbar. Abdominale Spiracula kreis-

rund und wenig erhaben. Abdomenende mit

Häkchen bewachsen.

Scythris klimeschi Passerin d’ Entreves, 1986

(Abb. 101-106)

Untersuchtes Material: 3 Männchen, 4 Weibchen. Ka-

narische Inseln.

Beschreibung. — Puppe 4,2-4,6X1,3-1,4 mm. Ab-

domenende mit etwas konkaven Seiten und relativ

spitz abgerundet. Färbung honigbraun, Exuvie heller.

Skulptur relativ fein, eher chagriniert als gerunzelt.

Borsten recht fein, am Kopf kaum unterscheidbar.

Labrum mit steilen Seiten, abgerundet, kaudal tief

und spitz ausgeschnitten. Labium eher klein, Frontal-

teil scharf abgegrenzt und nur wenig erweitert. Palpi

maxillares relativ groß, kurz an die Mittelbeine an-

grenzend. Grenze Genae/Vorderbeine sehr kurz. Mit-

telbeine etwas, Antennae viel länger als die Vorder-

beine, aber kürzer als die Proboscis, welche mit den

Vorderflügeln gleichlang ist. Abdomen der Exuvie

ventral bis zum Kaudalrand des 7. Segmentes

verdeckt. Pronotum relativ kurz, thorakales Spiracu-

lum mittellang, spaltförmig. Metanotum etwa zu Va

seiner Länge ausgeschnitten. Dieser Ausschnitt

stumpf abgerundet mit steilen Seiten. Frontallappen

des Metanotums ebenfalls breit und stumpf abgerun-

det. Hinterflügel bis zur Basis des 3. Abdominalseg-

mentes sichtbar. Abdominale Spiracula klein und et-

219

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

was erhaben. Am Abdomenende sehr kleine Bor-

sten.

DANKSAGUNG

Der Verfasser dankt vielen Fachkollegen, insbeson-

dere L. Aarvik (Äs), T. Baran (Torun), B. A. Bengts-

son (Löttorp), Dr. Gregor (Brno), Dr. Klimesch

(Linz a. d. D.), Ing. Liska, Ing. Skyva (beide Prag),

Ing. Turcani (B. Stiavnica) und Dr. Vävra (Prag) für

die Überlassung der Puppen zum Studium, der Liter-

atur und für Hinweise über die Nahrungspflanzen

mancher Scythris-Arten. Dem Museum für

Naturkunde Berlin (Dr. W. Mey), der Zoologischen

Staatssammlung München (Dr. W. Dierl) und dem

Naturhistorischen Museum Wien (Dr. M. Lödl)

dankt er für die Ermöglichung des Studiums des dort

befindlichen Puppenmateriales, ferner der Alexander

von Humboldt-Stiftung (Bonn) und Hanskarl-Goet-

tling-Stiftung (Freising) für die finanzielle Hilfe. Die

Untersuchungen wurden im Rahmen des Projektes

VEGA Nr. 2029 durchgeführt und finanziell unter-

stützt. Dr. E. J. van Nieukerken dankt der Autor für

die Durchsicht der Arbeit und werte Anmerkungen

und ausbesserungen im Text. Dr. Biesenbaum dankt

er außerdem für die deutschsprachige Korrektur der

ganzen Arbeit.

220

LITERATUR

Bengtsson, B. A., 1984. The Scythrididae (Lepidoptera) of

Northern Europe. — Fauna Entomologica Scandinavica

13: 1-137.

Bengtsson, B. A., 1997. Scythrididae. — In P. Huemer, O.

Karsholt & L. Lyneborg (eds.): Microlepidoptera of Eu-

rope 2. Apollo Books, Stenstrup: |- 301.

Huemer, R. P. & G. Tarmann, 1993. Die Schmetterlinge

Österreichs. Systematisches Verzeichnis mit Verbre-

itungsangaben für die einzelnen Bundesländer. — Tiroler

Landesmuseum Ferdinandeum, Innsbruck, 224 pp.

Karsholt, O. & Razowski J., 1996. The Lepidoptera of Eu-

rope. A distributional checklist. — Apollo Books, Sten-

strup, 380 pp.

Mosher, E., 1916. A classification of the Lepidoptera based

on the characters of the pupa. — Bulletin of the Illinois

State Laboratory of Natural History 12: 1-159.

Patocka, J., 1987. Die Puppen der mitteleuropäischen

Schmetterlinge (Insecta: Lepidoptera): Überfamilien

Yponomeutoidea, Gelechioidea, Schreckensteinoidea,

Epermenoidea und Copromorphoidea. Linzer biologis-

che Beiträge 29 (im Druck).

Received: 27 February 1997

Accepted: 25 November 1997

M. TERZO & P. RASMONT

Laboratoire de Zoologie, Université de Mons-Hainaut, Belgique

CERATINA ZWAKHALSI ET C. VERHOEFFI,

DEUX NOUVELLES ESPECES DE LA REGION

OUEST-PALEARCTIQUE

(HYMENOPTERA, APOIDEA, XYLOCOPINAE)

Terzo, M. & P. Rasmont, 1998. Ceratina zwakhalsi et C. verhoeffi, deux nouvelles espèces de la

region ouest-paléarctique (Hymenoptera, Apoidea, Xylocopinae). — Tijdschrift voor

Entomologie 140 [1997]: 221-236, figs. 1-58, tables 1-6. [Issn 0040-7496]. Published 26

March 1998.

Two new species of small carpenter bees are described from the West Palearctic region:

Ceratina zwakhalsi from South-East Turkey (compared with C. acuta Friese, 1896, C. callosa

(Fabricius, 1794) and C. chalybea Chevrier, 1872) and Ceratina verhoeffi from Morocco (com-

pared with C. saundersi Daly, 1983 and C. maghrebensis Daly, 1983). Distribution maps are

given for all the treated species. Ceratina saundersi is recorded for the first time from Portugal,

the Lampedusa island (South of Sicily), the Italian peninsula and Lybia. The occurence of C.

saundersi in Sardinia and Spain is confirmed.

Correspondence: Université de Mons-Hainaut, Laboratoire de Zoologie, av. Maistriau 19, B-

7000 Mons, Belgium. E-mail: [email protected]

Key words. — Hymenoptera; Apoidea; Xylocopinae; Ceratina; systematics; new species; West-

Palearctic; distribution maps.

A l'occasion de notre révision de la collection de M.

Gijs van der Zanden (Eindhoven, Pays-Bas), deux

nouvelles espèces de cératines de la région ouest-

paléarctique ont été découvertes: C. (Euceratina) ver-

hoeffi sp. n. et C. (Euceratina) zwakhalsi sp. n. Elles

viennent s'ajouter à la liste des espèces de cératines

ouest-paléarctiques connues notamment depuis les

travaux de Friese (1896, 1901) et Daly (1983).

C. zwakhalsi est comparée ici à C. acuta Friese,

1896, C. callosa (Fabricius, 1794) et C. chalybea

Chevrier, 1872 tandis que C. verhoeffi est comparée à

C. saundersi Daly, 1983 et C. maghrebensis Daly,

1983. Les mesures utiles pour caractériser ces espèces

sont décrites en détail par Daly (1973). Elles sont

reprises aux figures 1, 2 et 3 et sont exprimées en cen-

tièmes de millimètres. Elles ont été mesurées notam-

ment sur les holotypes mâles des nouvelles espèces. La

longueur de l'aile est mesurée depuis l'apex de la

plaque humérale jusqu'à l'apex de l'aile. La nomen-

clature des organes est inspirée de Bitsch & Leclercq

(1993). Les individus, autres que les types, ayant

servis à la réalisation des figures et des tableaux provi-

ennent des localités suivantes: C. acuta G: Grèce,

Macedonia, péninsule de Sithonia; C. acuta 9: Bul-

garie, Blageovgrad, Sandanski; C. callosa 3: Espagne,

Teruel, Pozondon; C. callosa 2: Espagne, Valencia,

Loriguilla ; C. chalybea 3: France, Haute Loire, St-

Didier d'Allier; C. chalybea 2: France, Alpes de Hau-

tes Provence, Vachères; C. maghrebensis 3: Maroc,

Marrakech, Marrakech; C. maghrebensis 2: Maroc,

Aguerd el Had; C. saundersi 3 et 9: Espagne, Ali-

cante, Moraira; C. verhoeff? 2: Maroc, Marrakech,

Ouirgane; C. zwakhalsi 9: Turquie, Hakkari,

Varagoz.

Les localisations des données citées sont renseig-

nées en degrés-minutes Greenwich.

Le matériel typique, soit les deux mâles de C.

zwakhalsi et de C. verhoeffi, provenant de la collection

privée de Monsieur Ing. Gijs van der Zanden, appar-

tient maintenant 4 la collection du Nationaal Natuur-

historisch Museum Leiden, Pays-Bas (RMNH, Dr C.

van Achterberg). Le reste du matériel de ces deux es-

pèces et celui des espèces qui leur sont comparées

proviennent essentiellement des musées et collections

suivants: Oberösterich Landesmuseum Linz (OOLL,

Mag. F. Gusenleitner), Ecole Nationale Supérieure

22]

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

C. callosa

C. chalybea

C. acuta

C. zwakhalsi

Coloration générale

Labre

Clypeus

Carène hypostomienne

Lobes pronotaux

Mesonotum

Fémur postérieur

Tibia postérieur

Sternite 6

Carène du tergite 6

Tergite 7

Genitalia

d -9: bleu métallique, y

compris les sternites

d : noir, parfois avec une pe-

tite marque centrale ivoire

2: noir

2: noir, avec parfois une pe-

tite marque centrale ivoire

réduite; ponctation fine et

présente sur tout le clypeus;

environs 8 ponctuations de

front derrière la suture

clypeale postérieure

d -9: étroite et très légère-

ment convexe (fig. 44)

d - 2: ivoire bordés ou non

de noir

d -2: partie dorsale éparse-

ment et irrégulièrement

ponctuée, laissant de petites

plages imponctuées ou des

plages à plus grande densité

de ponctuations; avec, entre

les notauli et la ligne médi-

ane, 2 à 4 ponctuations con-

tigües ou très distantes

d : pilosité ventrale de la

moitié basale en forme de

brosse courte de longueur

constante (fig. 4)

4 : pilosité ventrale plus

longue à l'apex que la plus

grande largeur du tibia (fig.

4); épine basitibiale située au

quart de la longueur du tibia

d : marge postérieure avec

deux larges lobes submé-

dians tronqués à l'apex et

dont la base interne porte un

petit lobe secondaire dirigé

vers l'avant (fig. 10)

2: presque droite, étroite,

avec une légère convexité à

la base

d : arrondi à l'apex, avec des

marges latérales droites ou

concaves (fig. 7)

voir figures 13 et 16

d -®: bleu métallique, ster-

nites noirs

d: ivoire

2: noir

2: comme callosa mais avec

une marque ivoire plus

grande

d - 2: étroite antérieure-

ment, large et convexe

postérieurement (fig. 45)

d -2: noirs ou ivoire bordés

de noir

3 -2: comme callosa

d : pilosité ventrale de la

moitié basale en forme de

brosse courte avec des soies

deux fois plus longues au

tier apical (fig. 5)

d: comme callosa (fig. 5)

d : marge postérieure avec

deux larges lobes submé

dians tronqués à l'apex et

dont la base interne porte un

petit lobe secondaire tron-

que à l'apex et dirigé vers

l'arrière (fig. 11)

bien marquée, suivie d'une

concavité jusqu'à l'apex

acuminé

3: largement arrondi à

l'apex, avec des marges

latérales jamais concaves

(fig. 8)

voir figures 14 et 17

d -?: bleu métallique, y

compris les sternites

: noir avec une petite mar-

3 tit

que centrale ivoire

Q: noir

Q: noir, avec parfois une pe-

tite marque centrale ivoire;

ponctuations de largeur

égale a celles du reste de la

face et bien délimitées; envi-

rons six ponctuations de

front derriére la suture

clypeale postérieure (fig. 25)

d -2: comme zwakhalsi

d -®: entièrement ivoire

3 -®: partie dorsale densé-

ment et régulièrement ponc-

tuée; avec, entre les deux no-

tauli 8 à 10 ponctuations

espacées par 1 fois le di-

amètre d'une ponctuation

au plus (fig. 27)

d : pilosité ventrale de la

moitié basale en forme de

brosse longue (fig. 19)

d: comme callosa (fig. 19)

zwakhalsi (fig. 21)

2: comme callosa

d: pointu à émoussé à

l'apex; en forme de triangle

dont les marges latérales

sont droites ou légèrement

concaves (fig. 20)

voir figures 22 et 23

d -2: bleu métallique, y

compris les sternites

d: ivoire

2: noir

2: noir, avec une marque

ivoire centrale de petite

taille, rarement absente; cer-

taines ponctuations deux à

trois fois plus grandes que

celles du reste de la face, mal

délimitées et éparses; envi-

rons cinq ponctuations de

front derrière la suture

clypeale postérieure (fig. 24)

d -®: très étroite et droite

sur toute sa longueur (fig.

46)

d -®: entièrement ivoire

d -2: partie dorsale éparse-

ment et régulièrement ponc-

tuée; avec, entre les deux no-

tauli 6 à 8 ponctuations

espacées par plus de deux

fois le diamètre d'une ponc-

tuation (fig. 26)

d : pilosité ventrale de la

moitié basale en forme de

brosse courte dont la

longueur croît progressive-

ment depuis la base du fé-

mur jusqu'à sa mi-longueur

(fig. 6)

3: comme callosa (fig. 6)

d: marge postérieure avec

deux petits lobes subme-

dians arrondis à l'apex et

dont la base interne porte un

petit lobe secondaire dirigé

vers l'arrière (fig. 12)

2: comme chalybea mais

avec une concavite centrale

et une convexite basale

moins marquées

d : fortement pointu à

l'apex, formant un angle

aigu (fig. 9)

voir figures 15 et 18

222

TERZO & RASMONT: Nouvelles Ceratina

Fig 1-3. Têtes schématisées, vues de face, avec les mesures pour caractériser les espèces. — 1, Clypeus: longueur (A) et largeur

(B); distances séparant la base du clypeus de l'insertion antennaire (C) et de l'ocelle médiane (D). Carène frontale: longueur

(CF); 2, Antennes: distances séparant les insertions des antennes (A); l'insertion antennaire du bord de l'oeil (B), de l'ocelle

médiane (C) et de l'ocelle latérale la plus proche (D); diamètre de l'insertion antennaire (E). Espace malaire: longueur (A) et

largeur (B) (non représentées); 3, Ocelles: distances séparant les ocelles latérales (A); une ocelle latérale du bord de l'oeil le plus

proche (B) et du bord postérieur de la tête (C); l'apex de l'oeil du bord postérieur de la tête (D); diamètre de l'ocelle médiane

(E). Yeux: longueur de l'oeil (A'); distances interoculaires au niveau du sommet de yeux (B'), des insertions antennaires (C')

et de la base des yeux (D').

Agronomique de Montpellier (Ensam, Prof. EF.

Leclant), Faculté des Sciences Agronomique de

Gembloux (rsacx, Prof. C. Gaspar), Instituut voor

Taxonomische Zoölogie Amsterdam (zma, M W.

Hogenes), Museum für Naturkunde der Humboldt-

Universität Berlin (mNHUB, Dr F. Koch), Muséum

National d'Histoire Naturelle de Paris (MNHNP, Dr J.

Weulersse), Muséum d'Histoire Naturelle de Stras-

bourg (MHNS, Dr J. Matter), Natural History Mu-

seum London (BMNH, M T. Huddleston), Università

di Catania (u.c., Dr V. Nobile), Université de Mons-

Hainaut (umH, Prof. P. Rasmont), coll. Prof. J. Le-

clercq (Gembloux), coll. Prof. L. Plateau (Vandoeu-

vre-les-Nancy), coll. Dr M. Comba (Cecchina di

Albano Laziale), coll. Dr G. Delvare (Montpellier),

coll. Dr J. Hamon (Gaillard), coll. Ir. R. Fonfria (Ey-

galières), coll. Ing. G. van der Zanden (Eindhoven)

(maintenant dans le RMNH), coll. Pater A. W. Ebmer

(Puchenau, Autriche), coll. A. Foucart (Montpellier),

coll. J.-M. Maldès (Montpellier), coll. A. Pauly

(Gembloux), coll. M. Schwarz (Ausfelden, Autriche),

coll. R. Wahis (Gembloux). La somme du matériel

revu par les auteurs s'élève à 1486 spécimens.

Ces données ainsi que les données bibliographiques

de Balles (1925, 1927, 1933), Daly (1983), Pagliano

(1994) et Westrich (1990) ont été gérées à l'aide du

logiciel Microbanque Faune-Flore (Rasmont et al.

1993). Le montant total des spécimens s'élève ainsi à

1935 spécimens. Les cartes ont été établies à l'aide du

logiciel Carto Fauna-Flora 1.2 (Barbier & Rasmont

1995, 1996). La carte orographique de la Turquie a

été saisie au départ de cartes au 1/2.000.000 à l'aide

même logiciel.

La nomenclature des types de distribution est in-

spirée de de Lattin (1967).

Tableau 2. Ceratina spp. Biométrie des mâles. Pour l'explication des mesures A-E, voir fig. 1-3, (unité 0.01 mm).

C. callosa C. chalybea

LEE 700

Nombre de hamuli 7 7

Yeux: A, B, C, D

Clypeus: À * B, C, D

Antennes: À, B, C, D, E

132, 158, 134, 119

64 * 98, 42, 100

42, 29, 45, 56, 16

139, 173, 140, 131

77 * 99, 45, 102

46, 30, 47, 54, 18

C. acuta C. zwakhalsi

vi 460 re 490 URI

6-7 6

128, 143, 120, 95 120, 140, 113, 97

99 atopy ills too) 61 * 78, 35, 85

39, 27, 42, 50, 15 32, 24, 39, 49, 15

Carène frontale 69 64 43 56

| Ocelles: A, B, C, D, E ayy Sil, IS), Si, 16 34, 54, 19, 55, 18 35, 40, 16, 45, 15 29, 43, 14, 41, 15

Espace malaire: A*B 4* 45 6*58 1 * 48 4*35

Nombre d'articles 6 6 6 6

au palpe maxillaire

223

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

POSSO,

puerta,

Fig. 4-18. Ceratina spp., détails du mâle. — 4, 7, 10, 13, 16, C. callosa Gs 5, 8, IA Il chalybea 3; 6,9, 12, 15, 18: C. i

zwakhalsi 3; 4-6, patte postérieure; 7-9, tergite 7 en vue ventrale; 10-12, sternite 6; 13-15, profil droit du genitalia; 16-18, |

genitalia en vue dorsale (moitié gauche) et en vue ventrale (moitié droite). Echelle: 1 mm.

224

TERZO & RASMONT: Nouvelles Ceratina

Tableau 3. Ceratina spp. Biometrie des femelles. Pour l'explication des mesures A-E, voir fig. 1-3, (unité 0.01 mm).

Longueur de l'aile

Nombre de hamuli

Yeux: A, B, C, D

Clypeus: A * B, C,

Antennes: A, B, C, D, E

Carène frontale

Ocelles: A, B, C, D, E

Espace malaire: A*B

Nombre d'articles

C. callosa

600

156, 182, 157, 139

73 * 112, 40, 107

50, 36, 55, 62, 17

37, 54, 24, 62, 17

4*59

C. chalybea

150, 178, 154, 147

77 * 115, 44, 108

49, 36, 54, 61, 17

35, 57, 27, 66, 18

6*63

C. acuta

450

RSS tel

42, 31, 46, 53, 15

31, 48, 20, 51, 15

3 * 49

C. zwakhalsi

440

121, 138, 116, 100

56 * 84, 32, 88

35, 27, 47, 53, 14

29, 44, 15, 49, 13

4*43

au palpe maxillaire

Ceratina (Euceratina) zwakhalsi sp. n.

Locus typicus. — Turquie, Hakkäri, Suvarihalil

Gesidi (37°30'N 43°23'E), 2.500 m.

Etiquettes de l'holotype. — “Turkey Hakkäri Suvari

Halil Pass 27-VI 1985 2500 m C. J. Zwakhals [im-

primée]; ‘Museum Leiden Ceratina mandibularis

Friesed det. G. v. d. Zanden 1986’ [en partie im-

primée et manuscrite]; “Terzo M. det. 1995 Ceratina

zwakhalsi HOLOTYPE’ [mécanographiée sur papier

rouge]. Collection RMNH.

Paratypes (49, 316) — Prov. Agri: Agri (39°55'N

43°03'E), 1d 28.v.1980 leg. M. Schwarz (Coll. Schwarz).

Prov. Bitlis: Nemrut Dagi (38°40'N 42°12'E), 1d

16.viii.1991 leg. K. Warncke (o611); Tatvan (38°30'N

42°16'E), 18 16.viii.1991 leg. K. Warncke (oorr). Prov.

Elazig: Elazig (38°41'N 39°14'E), 15 7.vi.1980 leg. M.

Schwarz, (coll. Schwarz). Prov. Erzincan: Refahiye (39°54'N

38°46'E), 15 23.viii.1991 leg. K. Warncke (oor). Prov.

Hakkari: Hakkari, Suvarihalil Gecidi (37°30'N 43°23'E), 24

2.vi.1980 leg. K. Warncke (oor); Ikiyaka Daglari (37°20'N

44°15'E), 29 94 10.vi.1981 leg. K. Warncke (oot); Tanin

Daglari (37°29'N 43°00'E) 19, 2d 2.vi.1980 leg. K.

Warncke (oom), 38 25.vi.1985 leg. M. Schwarz (coll.

Schwarz); Varagoz (37°15'N 44°08'E), 15 15.vi.1984, 19

17.vi.1984 leg. K. Warncke (ooLL); Yuksekova (37°35'N

44°17'E), 1d 28.vi.1985 leg. M. Schwarz (coll. Schwarz).

Prov. Icel: Sertavul (?), 26 9.vi.1978 leg. M. Schwarz (coll.

Schwarz). Prov. Kahraman Maras: Goksun (38°03'N

36°30'E), 16 10.vii.1990 leg. P. A. Ebmer (coll. Ebmer).

Prov. Kars: Karakurt (40°09'N 42°25'E), 16 1.vi.1988, 14

| 2.vi.1988 leg. K. Warncke (oor); Karakurt (40°10'N

| 42°36'E), 16 23.v.1975 leg. K. Warncke (oot); Karakurt

| (40°10'N 42°29'E), 16 23.v.1975 leg. K. Warncke (ooLL);

| Karakurt (40°08'N 41°39'E), 15 27.v.1980 leg. M. Schwarz,

| (coll. Schwarz). Prov. Sivas: Gurun (38°44'N 37°15'E), 16

| 7.vii.1984 leg. P. A. Ebmer (col. Ebmer); Gurun (38°44'N

| 37°15'E), 1d 3.vi.1978 leg. M. Schwarz (coll. Schwarz). Prov.

| Van: Baskale (38°03'N 44°01'E),

| 12.vii.1984 leg. P. A. Ebmer (coll. Ebmer); Baskale (38°26'N

| 44°26'E), 16 30.v.1980 leg. M. Schwarz (coll. Schwarz); Ercis

Id 9.vi.1984, 16

(39°01'N 43°13'E), 14 3.viii.1983 leg. K. Warncke (oo1t).

Description (Holotype d )

Voir tableaux 1 à 3; pattes postérieures comme à la

figure 6; tergite 7 comme à la figure 9; sternite 6

comme à la figure 12; genitalia comme aux figures 15

et 18. Le sternite 6, le tergite 7 et le genitalia ont été

disséqués par les auteurs et collés sur une étiquette de

plastique transparent.

Espèces voisines et diagnose différentielle

Les trois espèces dont les mâles peuvent être con-

fondus avec C. zwakhalsi sont C. callosa (Fabricius,

1794), C. chalybea Chevrier, 1872 et C. acuta Friese,

1896.

Ces mâles sont facilement reconnaissables par la

pilosité des pattes postérieures et la forme du dernier

segment abdominal. Seul le mâle de C. acuta possède

une pilosité ventrale de la moitié basale du fémur qui

soit uniformement longue (fig. 19). Dans le cas de C.

callosa, cette pilosité est uniformement courte (fig. 4).

C. chalybea diffère de C. callosa par la présence de

quelques soies longues à l'apex de cette pilosité fémo-

rale (fig. 5). C zwakhalsi posséde lui une pilosité fé-

morale courte à la base mais qui s'allonge progressive-

ment vers l'apex (fig. 6). La forme du dernier tergite

permet également de reconnaître facilement ces qua-

tre espèces. L'apex de ce tergite est en effet largement

arrondi chez C. chalybea (fig. 8), moins arrondi et aux

marges latérales concaves chez C. callosa (fig. 7), poin-

tu, formant un angle presque droit avec des côtés

droits chez C. acuta (fig. 20) et trés pointu avec des

marges latérales concaves chez C. zwakhalsi (fig. 9).

On sépare assez aisément les femelles de C. callosa et

C. chalybea des femelles de C. acuta et C. zwakhalsi par

la forme arrondie de la carene hypostomienne chez les

deux premières espèces et droite chez les deux

dernières. C. chalybea se distingue alors de C. callosa

par la présence d'un élargissement de la partie pos-

térieure de la carène hypostomienne. Ces caractères

sont également valables pour les mâles (fig. 44 à 46).

225

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 19-23. Ceratina acuta 3 . — 19, patte postérieure; 20, tergite 7 en vue ventrale; 21, sternites 6; 22, genitalia en vue dor-

sale (moitié gauche) et en vue ventrale (moitié droite); 23: profil droit du genitalia. Echelle: 1 mm.

On distingue beaucoup plus difficilement la femelle

de C. acuta de celle de C. zwakhalsi. Ces deux espèces

possèdent une tache clypeale réduite ou absente. Il

semble toutefois que cette tache soit plus souvent

présente chez C. zwakhalsi que chez C. acuta. On peut

cependant les distinguer par la ponctuation de la face

et du thorax. Dans le cas de C. acuta, ces ponctuations

sont plus petites et plus nombreuses. On peut compter

ainsi au moins 6 ponctuations de front sur l'aire supr-

aclypeale, le long de la suture supraclypeale (fig. 25).

On en compte rarement plus de cing chez C.

zwakhalsi (fig. 24). On peut également compter de 8 à

10 ponctuations sur une droite fictive reliant la base

des notauli du mesoscutum de C. acuta (fig. 27). On

en compte rarement plus de 6 à 8 chez C. zwakhalsi

(fig. 26). Elles sont en outre, chez cette dernière, plus

larges et plus distantes les unes des autres, laissant sou-

vent apparaître de larges zones imponctuées.

Clé

Les seules clés des cératines incluant la région est-

méditerranéenne sont celles de Friese (1896, 1901).

Dans le cas des femelles, selon que l'on soit en pré-

sence d'un spécimen présentant ou pas une marque

blanche sur le clypeus, la clé de Friese nous conduit

respectivement à C. callosa ou à C. acuta. Ce pro-

blème ne se pose pas dans le cas des mâles où l'on

abouti uniquement à C. acuta. La clé de Friese devrait

donc être modifiée comme suit:

226

Femelles

11. Grande taille, 12 à 14 mm de long ...C. chalcites

Latreille

— Peutetulle 69 mm dellone e lla

1 la. Carene hypostomienne courbe..................... 11b

— Carène hypostomienne droiteC. zwakhalsi sp. n.

11b.Carène hypostomienne étroite sur toute sa

longueur Sr re C. callosa Fabricius

— Caréne hypostomienne étroite à la base et élargie

POSÉÉTIEUTEMENT ee C. chalybea Germar

20. Lobes pronotaux noirs, 6 à 7 mm de long......21

— Lobes pronotaux blancs, 6 à 8 mm de long ..20a

20a. 8 à 10 ponctuations entre les bases des notauli

du mesoscutun; au moins 6 ponctuations de

front derrière la suture clypeale postérieure ........

MAR Ano, ANI BAR me C. acuta Friese

— 6à8 ponctuations entre les bases des notauli du

mesoscutun; rarement plus de 5 ponctuations de

front derrière la suture clypeale postérieure ........

eter Bo Me EE ET A ATEN C. zwakhalsi sp. n.

Mäles

9. Clypeus, labre et mandibules entièrement

blanes,leorps;plus verdatre ee

EE RE C. mandibularis Friese

— Mandibules entièrement noires, corps bleu ....9a

9a. Fémur postérieur doté d'une pilosité ventrale

uniformément longue; terite 7 triangulaire à

l'apex, au sommet émoussé et aux marges droites

ou légèrement concaves; labre noir ou doté

d'une petite marque blanche centrale.................

MR i AA e o dar C. acuta Friese

— Fémur postérieur doté d'une pilosité ventrale

courte à la base et s'allongeant vers l'apex; tergite

7 fortement pointu à l'apex, aux marges latérales

concaves; labre presque entièrement blanc … …

Ee PONE oes TR EE C. zwakhalsi sp. n.

Aires de distribution

L' aire de distribution de C. callosa apparaît claire-

ment comme de type atlanto-méditerranéenne (fig.

53). Elle couvre en effet tout le maghreb et s'étend

vers le nord jusqu'à la péninsule ibérique, le sud de la

France, la péninsule italique, la Corse, la Sardaigne et

la Sicile.

L'aire de distribution de C. chalybea est de type

euro-méditerranéenne (fig. 54). On trouve cette es-

pèce depuis le Maghreb jusqu'en Allemagne et en

Autriche vers le nord et jusque dans la province de

Van en Turquie vers l'est.

L'aire de distribution de C. acuta (fig. 53) est de

type est-méditerranéenne étendue vers le nord et vers

l'ouest. On la trouve ainsi principalement en Turquie

et en Grèce. Elle est également présente dans les pays

de l'est comme la Bulgarie, la Roumanie, l'ex-

AT A

c'e ee

TERZO & RASMONT: Nouvelles Ceratina

Yougoslavie et la Tchequie. Un spécimen a également

été capturé dans le sud-est de l'Allemagne. Il apparaît

ici évident que la limite nord-ouest de distribution de

cette espèce n'atteint pas les frontières italienne, pro-

tégée par les alpes, et suisse, trop à l'ouest. Plus au

sud, elle n'a apparement pas franchi la Mer Adria-

tique vers l'Italie, ni atteint la Sicile. Il est donc plus

qu'improbable que C. acuta puisse être présente en

Espagne comme Dusmet (1923), et plus récement

Pérez-Inigo Mora & Gayubo (1989) entre autres, le

signalent. Il se pourrait que ces auteurs aient confon-

du C. acuta avec C. cyanea (Kirby) dont certains

spécimens ont les lobes pronotaux blancs en pénin-

sule ibériques. Vers l'est, on trouve C. acuta jusqu'à la

frontière iranienne. Un spécimen a été trouvé en

Israël et un autre dans les montagnes du Bolshoy.

L'aire de distribution de C. zwakhalsi. (fig. 55)

semble se restreindre à l'est de la Turquie et serait de

type syrienne. Il est cependant possible que cette es-

pèce soit présente ou même originaire d'Iran ou

d'Asie centrale. Tous les individus de cette espèce, à

l'exception d'un seul, on été trouvés à une altitude

supérieure à 2000 mètres. Il s'agirait là de la seule es-

pèce de cératine ouest-paléarctique décrite dont la

distribution soit exclusivement montagnarde.

Fig. 24-27, Ceratina spp . — 24, 26, C. zwakhalsi 2; 25, 27, C. acuta Q. 24, 25, tête vue de face; 26, 27, mesoscutum en vue

dorsale, partie antérieure orientée vers le haut. Echelle: 0,5 mm.

227

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

28 / 29

Fig. 28-43. Ceratina spp., détails du mâle. — 28, 31, 32, 35, 38, 41, C. maghrebensis 3; 29, 33, 36, 39, 42, C. saundersi 3;

30, 34, 37, 40, 43, C. verhoeffi 8 . 28-30, patte postérieure; 31, sternite 4; 32-34, tergite 7 en vue ventrale; 35-37, sternite 6;

38-40, profil droit du genitalia; 41-43, genitalia en vue dorsale (moitié gauche) et en vue ventrale (moitié droite). Echelle: 1

mm.

228

Tableau 4. Ceratina spp. Caractères cuticulaires et pelage.

C. maghrebensis

Coloration générale

Labre

Clypeus

Carène hypostomienne

Lobes pronotaux

Mesonotum

Patte antérieure

Patte médiane

Fémur postérieur

Tibia postérieur

Sternites 2 à 5

Sternite 6

Carène du tergite 6

| Tergite 7

| Genitalia

d -9: métallique à reflets verdatres,

sternites noirätres

d -Q: noir

d : ivoire, étroitement bordé de noir;

ponctuation moins abondante que

verhoeffi sur tout le clypeus

2: avec une tache centrale oblongue

allant du sommet du clypeus

jusqu'au premier quart de la

longueur du clypeus

d -2: étroite et légèrement convexe

sur toute sa longueur (fig. 47)

d -®: entièrement ivoire

d -?: partie dorsale très densément

et irrégulièrement ponctuée; 5 à 6

ponctuations entre les notauli et la

ligne médiane, espacées par une dis-

tance égale ou plus petite que le di-

amètre d'une ponctuation; ponctua-

tions contigües au centre du

mesonotum, directement en arrière

de la ligne médiane

d -?: avec une petite marque ba-

sitibiale ivoire

d -?: avec une petite marque ba-

sitibiale ivoire

d : pilosité de la moitié basale courte

et éparse (fig. 28)

d -?: avec une marque basitibiale

ivoire n'atteignant pas le quart de la

longueur du tibia

d : pilosité ventrale plus courte que

la plus grande largeur du tibia (fig.

28)

6: avec deux larges lobes subme-

} : 8

dians imponctués sur la marge

o o &

posterieure des sternites; marge

postérieure du sternite 5 échancrée

(fig. 31)

d: marge postérieure avec deux

larges lobes submédians peu élevés et

légèrement recouverts à leur base in-

terne par un lobe médian large et

tronqué à l'apex (fig. 35)

9: présente uniquement à l'apex du

tergite où elle développe une petite

convexité qui disparait juste avant

l'apex acuminé du tergite (fig. 51)

d : apex arrondi et très proéminent

(fig. 32)

d : voir figures 38 et 41

TERZO & RASMONT: Nouvelles Ceratina

C.verhoeffi |

C. saundersi

d -2: bleu métallique, sternites

noiràtres

d -®: noir

d: ivoire, très largement bordé de

noir; ponctuation éparse sur tout le

clypeus

2: avec une tache centrale oblongue

allant du sommet du clypeus

jusqu'au premier tiers de la longueur

du clypeus

d -2: comme maghrebensis (fig. 48)

d -2: idem

d -2: partie dorsale densément et

régulièrement ponctuée; 4 à 5 ponc-

tuations entre les notauli et la ligne

médiane, espacées par une distance

plus grande que le diamètre d'une

ponctuation

d -2: comme maghrebensis

d : pilosité de la moitié basale en

forme de brosse très courte, de

longueur constante mais plus dense

que chez verhoeff? (fig. 29)

d -2: marque basitibiale comme

maghrebensis

6: pilosité ventrale comme

maghrebensis (fig. 29)

d : sans lobes submedians imponc-

tués; marge postérieure du sternite 5

plus échancrée que chez maghrebensis

d : marge postérieure avec deux

larges lobes submédians peu élevés et

dont les marges internes sont ra-

menées ventralement vers l'extérieur

(fig. 36)

2 longue et droite sur toute sa

longueur (fig. 52)

d : apex largement arrondi et peu

proéminent (fig. 33)

d: voir figures 39 et 42

d -2: vert métallique à reflets dorés,

sternites noirätres

d: noir, avec une petite marque

ivoire centrale

2: noir

d : ivoire, étroitement bordé de noir;

ponctuation éparse mais un peu re-

groupée sur la bande centrale

9: avec une grande tache centrale

trapézoidale allant du sommet du

clypeus jusqu'au premier quart de la

longueur du clypeus et deux fois plus

large à la base qu'à l'apex

d -9: étroite et moins convexe que

maghrebensis (fig. 49)

d -2: idem

d -2: partie dorsale densément et ir-

régulièrement ponctuée; 4 à 5 ponc-

tuations entre les notauli et la ligne

médiane, souvent espacées par une

distance plus grande que le diamètre

d'une ponctuation; ponctuations

contigües sur les deux tiers

postérieurs du mésonotum, bien en

arrière de la ligne médiane

3 -?: avec une marque basitibiale

ivoire étendue sur plus des deux tiers

de la longueur du tibia

d -®: avec une petite marque ivoire

basitibiale étendue sur près du quart

basal du tibia

d : pilosité de la moitié basale en

forme de brosse très courte et de

longueur constante (fig. 30)

d -?: avec une marque basitibiale

ivoire étendue sur près des deux tiers

de la longueur du tibia

d : pilosité ventrale comme

maghrebensis(fig. 30)

3: sans lobes submedians imponc-

tués; marge postérieure du sternite 5

non échancrée

3: fort semblable à saundersi (fig.

37)

2: comme saundersi

d : apex très largement arrondi et

peu proéminent (fig. 34)

8: voir figures 40 et 43

229

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 44-49.

Ceratina spp., profil gauche

de la tête. La carène hypo-

stomienne est indiquée par

une flèche. — 44, C. callosa 3;

45, C. chalybea 3; 46, C.

zwakhali 08; 47, €

maghrebensis 8; 48, C. saun-

dersi 3; 49, C. verhoeff S.

Tableau 5. Ceratina spp. Mesures et ratio des mâles. Pour l'explication des mesures A-E, voir fig. 1-3, (unité 0.01 mm).

C. maghrebensis

Longueur de l'aile

Nombre de hamuli

Yeux: À, B, C, D

Clypeus: À * B, C, D

Antennes: À, B, C, D, E

Carène frontale

Ocelles: A, B, C, D, E

Espace malaire: A*B

Nombre d'articles au palpe maxillaire

230

C. saundersi

420

109, 125, 104, 89

ITN

33, 22, 40, 47, 14

24, 33, 9, 36, 16

55)

400

111, 114, 97, 83

52 * 64, 32, 80

32, 22, 41, 50, 13

2593 By) By US)

DSR)

C.verhoeffi

460

129, 125, 107, 94

61 * 78, 32, 82

34, 22, 40, 50, 15

56 |

27, 29, 15, 31, 16 |

2 * 42

TERZO & RASMONT: Nouvelles Ceratina

Tableau 6. Ceratina spp. Mesures et ratio des femelles. Pour l'explication des mesures A-E, voir fig. 1-3, (unité 0.01 mm).

Longueur de l'aile

C. maghrebensis

390

Nombre de hamuli 6

Yeux: A, B, C, D 111,126, 110, 94

Clypeus: À * B, C, D SIS) 25 V1

Antennes: A, B, C, D, E 33, 25, 42, 49, 14

Caréne frontale 56

Ocelles: A, B, C, D, E 27, 34, 15, 38, 15

Espace malaire: A*B 2* 41

Nombre d'articles au palpe maxillaire 5

C. saundersi C.verhoeffi

430 440

5 5

127, 128, 116, 101 130, 130, 115, 101

57 * 84, 31, 86 62 * 82, 28, 85

38, 25, 45, 54, 14 39, 23; 46, 55, 15

57 62

27, 35, 18, 41, 16 29 351515; 37.16

2*45 32043

5 5

Derivatio nominis

Cette espèce est dédiée au Dr C. J. Zwakhals

(Arkel, Pays-Bas) qui a collecté l'holotype.

Ceratina (Euceratina) verhoeffi sp. n.

Locus typicus. — Maroc, Marrakech (31°49'N

8°00'W).

Etiquettes de l'holotype. — “Maroc Marrakech 18-

20-V-1950 PM F Verhoeff [imprimée]; ‘Museum

Leiden Ceratina saundersi Daly 3 det. G. v. d.

Zanden 1988’ [en partie manuscrite et imprimée];

‘Terzo M. det. 1994 Ceratina verhoeffi HOLOTYPE

[mécanographiée]. Collection RMNH.

Paratypes. — Prov. Agadir: Taroudant (30°29'N 8°52'W),

12 4d 24.vi.1974 leg. K. Guichard & G. Else (BMNH);

Prov. Marrakech: Imouzzer (31°19'N 7°23'W), 2¢

21.vi.1974 leg. K. Guichard & G. Else (BMNH); Ouirgane

| (B1°11'N 8°05'W), 32 8d 2.vii.1974 leg. K. Guichard &

G. Else (BMNH). Prov. Ouarzazate: Ait Saoun (30°45'N

6°38'W), 1d 14.vii.1975 leg. J. Gusenleitner (BMNH); Tizi

n'Bachkoum (30°39'N 7°17'W), 78 1.vii.1987 leg. M.

Schwarz (OOLL).

Description (Holotype d)

Voir tableaux 4 a 6; pilosité des pattes postérieures

comme a la figure 30; tergite 7 comme 4 la figure 34;

sternite 6 comme à la figure 37; genitalia comme aux

figures 40 et 43. Le tergite 7 et le genitalia ont été dis-

séqués par les auteurs et collés sur une étiquette de

plastique transparent.

| Espèces voisines et diagnose différentielle

Les deux espèces sympatriques qui peuvent être

confondues avec C. verhoeff? sont C. saundersi Daly,

1983 et C. maghrebensis Daly, 1983. Ces espèces sont

comparées à C. verhoeffi dans les tableaux 4 à 6.

Ces trois espèces forment un groupe d'espèces ho-

mogène caractérisé notamment par des palpes maxil-

laires à 5 articles au lieu de six (caractère également

présent chez C. albosticta Cockerell, 1931), une

hanche doté d'une large dent sur la marge ventrale,

donnant à l'apex de la hanche un aspect tronqué bien

différent de la forme conique habituelle (fig. 50) et

par une taille plus petite que celle des autres espèces

du même sous-genre à l'exception de C. dentiventris

Gerstaecker, 1869 et de C. dallatorreana Friese, 1896.

Les auteurs proposent de nommer "groupe de C.

maghrebensis' l'ensemble de ces trois espèces.

On distingue C. verhoeffi, male et femelle, facile-

ment grâce aux taches basitibiales ivoires des pattes

antérieures et postérieures qui s'étendent au delà de la

moitié de la longueur des tibias, ce qui n'est jamais le

cas chez les deux autres espèces. C. verhoeffi arbore

également une coloration vert métallique bien

prononcée contrairement à la coloration bleu de C.

saundersi et à la coloration légèrement verdâtre de C.

maghrebensis. Le mâle de C. maghrebensis se distingue

alors aisément de celui de C. saundersi par la présence

sur les sternites deux à cing de zones transversales im-

ponctuées et élevées en formes de lamelles (fig. 31).

Les femelles se distinguent par la forme de la carène

du sixième tergite abdominal. Celle-ci est pratique-

ment droite chez C. saundersi (fig. 52), tout comme

celle de C. verhoeffi, et courte avec une extrémité net-

tement convexe chez C. maghrebensis (fig. 51).

Clé

On doit à Daly (1983) d'avoir publié la dernière clé

en date pour les cératines du Maghreb. La découverte

de C. verhoeffi dans la région de Marrakech modifie sa

clé comme suit:

231

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Mäles

13 Sternites 2 à 5 non modifiés, entièrement ponc-

LES pene LR rain 13a

- Sternites 2 à 5 dotés chacun d'une zone bilatérale

imponctuée, élevée, en forme de croissant et

dont les marges apicales portent de longues soies

DA NL ae A RS EURE UE PILE RON Ceratina maghrebensis

13a Taches basitibiales blanches des pattes

postérieures et antérieures étendues dorsalement

sur plus des deux tiers de la longueur des tibias;

clypeus entièrement blanc ...... Ceratina verhoeffi

- Taches basitibiales blanches des pattes

postérieures et antérieures n'atteignant pas le

quart de la longueur des tibias; clypeus avec de

larges marges latéro-basales noires......................

N Ceratina saundersi

Femelles

22 Carène médiane du tergite 6, vue de profil, plus

longue et étroite; tache clypeale distincte, large;

corps aux reflets métalliques bleu-vert .......... 22a

- Carène du tergite 6 plus courte, la marge avec

une convexité subapicale distincte; tache

clypeale mal définie, petite ou absente; corps aux

reflets/metalliquesiplusiverdatres nenne

RP De PAL EVE LEG OR Ceratina maghrebensis

22a Taches basitibiales blanches des pattes

postérieures et antérieures étendues dorsalement

sur plus des deux tiers de la longueur des tibias..

LR AOL SERVI RIO SON Ceratina verhoeffi

- Taches basitibiales blanches des pattes

postérieures et antérieures n'atteignant pas le

quare.de laloneueundessubiase nn ne

ÉD ER A EEE TEN TER Be Ceratina saundersi

Aires de distribution

La distribution de C. maghrebensis est de type mau-

ritanienne (fig. 56). On trouve cette espece dans le

centre du Maroc, le nord de l'Algerie et le nord de la

Tunisie. Elle semble ainsi se limiter aux montagnes de

l'Atlas marocaines et algériennes et aux régions

252

Fig. 50-52.

50, profil gauche de la hanche

et du trochanter postérieurs

52 gauches de C. maghrebensis 8 .

51-52, Profil gauche du ter-

gite 6. —51, C. maghrebensis

Echelle: 1 mm.

côtières méditerranéennes algériennes et tunisien-

nes.

La distribution de C. saundersi est de type atlanto-

méditerranéenne (fig. 57). La figure 57 confirme la

présence de cette espèce en Espagne depuis Daly

(1983) et révèle une distribution largement étendue

vers le nord jusqu'alors insoupçonnée:

Prov. Alicante: Benidorm 38°33'N 0°08'W (coll.

J. Leclercq); Llosa de Camacho 38°46'N 0°00'E, 16

21.vii.1989, leg. J. Selfa (ooLL); Moraira 38°40'N

7°57'E, 26 4-10.ix.1989, 16 4-13.ix.1989, 16 26-

27.1x.1989, 12 12-18.ix.1989, 16 6-11.III.1990,

leg. R. Wahis (FSAGx). — Prov. Almeria: Almeria

36°53'N 2°21'W, 1d 1.viii.1984, leg. R. Hensen

(coll. G. van der Zanden). — Prov. Cadiz: 30Km S-

Cadiz 36°13'N 6°16'W, 19 14.viii.1992, leg. K.

Warncke (OOLL). — Prov. Granada: Albunol 36°47'N

3°11'W, 1d 15.iv.1979, leg. W. Schacht (ooLL);

Salobrena 36°46'N 3°35'W, 1d 8.v.1983, leg. W.

Perraudin (ooLL). — Prov. Jaen: Las Correderas

38°22'N 3°30'W, 14 13.vi.1961, 16 14.vi.1961,

leg. J. van der Vecht (RMNH). — Prov. Malaga: Malaga

36°46'N 4°26'W, 16 viii.1987 (rsacx); Ronda

36745 NS 07 We OS vil GONNIE PANIERS

Verhoeff (RMNH); idem, 19 11.vi.1986, leg. W. Per-

raudin (ooLL). — Prov. Sevilla: Sevilla 37°23'N

5°57'W, 14, leg. J. Dusmet (MNHNP). — Prov. Tole-

do: Toledo 39°53'N 4°01'W, 18 25-27.v11.1967,

leg. P.M.F. Verhoeff (RMNH).

Un individu mâle capturé par Friese (MNHUB) et

renseigné “Pyrenaén” montre que C. saundersi atteint |

le nord de sa distribution dans les Pyrénées. Détermi- _

né par Friese comme étant un mâle de Ceratina dalla- |

torreana Friese, 1896, il est très probable qu'il s'agisse

la du spécimen ayant servi à Friese pour décrire le |

mâle de cette espèce.

La carte de distribution de C. saundersi montre |

pour la première fois sa présence au Portugal: Prov. |

Algarve: Alcoutim 37°25'N 7°40'W, 2d 9.viii.1698, |

16 20.viii.1968, leg. K. Warncke (oorr); Olhao |

| 37°04'N 7°49'W, 19 22.viii.1968, leg. K. Warncke

Moorr);:mlaviran 37413 INT ANT 16 29-

| 31.viii. 1984, leg. P. van Ooijen (zMa).

B Sa présence en Sardaigne (Italie) est confirmée

depuis Pagliano, 1994 par les récoltes suivantes:

Caoliari 3904 NY 905E, LOTO vi 1959 Id

13.vii.1959, leg. H. Wolf (muns); idem, 29 16

15.vii.1959, 1d 20.vii.1959, leg. H. Wolf (ooır);

Porto Santoru 39°37 N 9°39'E, 1d 12.vi.1936, leg.

H.G. Amsel (MNHUB); Pula 39°00'N 8°55'E, 1d

20.iv.1960, leg. E. Priesner (OOLL).

L'est de sa distribution atteint l'île de Lampedusa

au sud de la Sicile: Lampedusa 35°31'N 12°37'E, 16

18.v.1987, leg. Arnone (oorr). Elle atteint également

le centre de l'Italie: Prov. Lazio, Terracina 41°19'N

13°11'E, 19 ix.1974, leg. T. Osten (oor). L'île de

Lampedusa et la péninsule italique sont ainsi citées

pour la première fois dans la distribution de C. saun-

dersi.

C. verhoeffi présente une distribution de type mau-

ritanienne stricte (fig. 58). Cette espèce présente

cependant une distribution restreinte, voire endé-

mique, à la vaste plaine de la province de Marrakech,

avec des excursions au sud jusqu'à Taroudant (prov.

Agadir) et à l'est jusqu'à Ait Saoun (prov.

Ouarzazate).

Derivatio nominis

Cette espèce est dédiée au Dr P. M. F. Verhoeff

| (Pays-Bas) qui a collecté l'holotype.

REMERCIEMENTS

Nous remercions tout particulièrement Monsieur

Gijs van der Zanden pour sa collaboration et le prét

de matériel qu'il nous a accordé. Nous remercions

également toutes les personnes qui ont prété ou permi

l'accés au matériel de cette étude et qui sont cités dans

l'introduction. Que tous les récolteurs du matériel

étudiés, cités ci-après, soient également remerciés:

M. Alicata, M. Arnone, J. van Asperen den Boer,

Y. Barbier, M. Blank, K. Bleyl, C. de Jong, Z. Bou-

cek, P. Brandl, L. Cederholm, W. Degroot, R. Des-

mier De Chenon, P.van Doesburg, G. Else, W.

| Gross, K. Guichard, J. & M. Halada, C. van Heijnin-

gen, R. Hensen, C. Jeekel, H. Knorr, M. Kraus, M. la

Greca, R. Leys, W. Linsenmaier, M. Mei, H. Meuf-

fels, N. Mohr, A. Moussa, S. J. van. Ooststroom, T.

Osten, Z. Padr, W. Perraudin, K. Polacek, P. Pronk,

M. Quedenfeldt, W. Schacht, M. Scheiden, J. Selfa,

E. A. M. Speijer, J. Steffen, J. Timmer, S. Tomarchio,

V.S. van der Goot, J. van der Vecht, F. Vinkestijn, J.

T. Wiebes, H. Wolf et C. J. Zwakhals.

TERZO & RASMONT: Nouvelles Ceratina

Nous remercions enfin Mr D. Flagothier pour

avoir réalisé la carte orographique de la Turquie.

REFERENCES

Balles, L. von, 1925. Beiträge zur Kenntnis der Hymenopte-

renfauna Badens. I. Beitrag zur Kenntnis der badischen

Bienen. — Mitteilungen des Badischen Landesvereins für

Naturkunde und Naturschutz in Freiburg im Bresgau,

Neue Folge 1 (23/24): 437-461.

Balles, L. von, 1927. Beiträge zur Kenntnis der Hymenopte-

renfauna Badens. IV. Vierter Beitrage zur Kenntnis der

badischen Bienen (Arten aus dem Gebiete des Oberrheins

und des Bodensees). — Archiv für Insektenkunde des

Oberrheingebietes 2: 199-210.

Balles, L. von, 1933. Beiträge zur Kenntnis der Hymenopte-

renfauna Badens. V. Fünfter Beitrage zur Kenntnis der

badischen Bienen. — Beiträge zur Naturwissenschaftli-

chen Erforschung Badens 12: 189-195.

Barbier, Y. & P. Rasmont, 1995. Carto Fauna-Flora, car-

tographie des données biologiques, logiciel MS-Dos ver-

sion 1.0. — Université de Mons-Hainaut, 93+36+2 pp.

Barbier, Y. & P. Rasmont, 1996. Carto Fauna-Flora, car-

tographie des données biologiques, logiciel MS-Dos ver-

sion 1.2. — Université de Mons-Hainaut, 62 pp.

Bitsch, J. & J. Leclercq, 1993. Hyménoptères Sphecidae

d'Europe Occidentale, Volume 1, Généralités - Crabro-

ninae. — Faune de France 79: 1-330.

Daly, H. V., 1973. Bees of the genus Ceratina in America

North of Mexico (Hymenoptera: Apoidea). — University

of California Publications in Entomology 74: 1-113.

Daly, H. V., 1983. Taxonomy and ecology of Ceratinini of

North Africa and the Iberian Peninsula (Hymenoptera:

Apoidea). — Systematic Entomology, 8: 29-62.

De Lattin, G., 1967. Grundriss der Zoogeographie. —

Gustav Fischer Verlag, Jena, 602 pp.

Dusmet y Alonso, J., 1923. Los Apidos des Espana. VI. Géne-

ros Xylocopa Latr. y Ceratina Latr. - Memorias de la Real

Sociedad Espafiola de Historia Natural 11 (7): 269-304.

Friese, H., 1896. Monographie der Bienengattung Ceratina

(Latr.) (Palearktische Formen). — Termeszetrajzi Füzetek

19: 34-65.

Friese, H., 1901. Die Bienen Europa's (Apidae europaeae).

Theil VI. Solitäre Apiden. — C. Lampe, Innsbruck, 284 pp.

Pagliano, G., 1994. Catalogo degli imenotteri italiani. IV.

(Apoidea: Colletidae, Andrenidae, Megachilidae, Antho-

phoridae, Apidae). — Memorie della Società Entomolo-

gica Italiana, 72: 331-467.

Pérez-Inigo Mora, C. & S. F. Gayubo, 1989. Notas sobre

los apoideos de la Sierra de Bejar. I. Anthophoridae

(Hym. Apoidea). — Studia Oecologica, 6: 313-331.

Rasmont, P., Y. Barbier & A. Empain, 1993. Microbanque

Faune-Flore, logiciel de banques de données biogéograp-

hiques. logiciel Ms-pos version 3.0. — Université de

Mons-Hainaut, jardin Botanique National de Belgique,

xv+200+20+3+34+14 pp.

Westrich, P., 1990. Die Wildbienen Baden-Wiirttembergs.

Allgemeiner Teil, pp.1-431; Spezieller Teil, pp.432-972.

— Verlag Eugen Ulmer, Stuttgart.

Received: 30 June 1995

Revised version received: 20 November 1996

Accepted: 30 April 1997

233

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

@ Ceratina callosa È

W Ceratina acuta

1 Ha

ip an: + n/a

ì £

EN %

Ceratina chalybea È

Fig. 54. Carte de distribution de Ceratina chalybea.

234

TERZO & RASMONT: Nouvelles Ceratina

Ceratina zwakhalsi

«© À“ Gen |

CRP 1.2 rt :

SS i 4 : :

Fig. 55. Carte de distribution de C. zwakhalsi. En grisé: altitudes supérieures à 2000 m.

Ceratina maghrebensis

CEF (12) — 1996

su 1 4

Fig. 56. Carte de distribution de Ceratina maghrebensis.

235

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

PSS

Ceratina saundersi

Fig. 57. Carte de distribution de Ceratina saundersi.

Ceratina verhoeffi

Fig. 58. Carte de distribution de Ceratina verhoeffi.

236

oo OO FF (FFT

ToMoHIDE YASUNAGA' & YuxIinoBu NAKATANT

‘Hokkaido University of Education, Sapporo, Japan

"University of Osaka Prefecture, Sakai, Japan

BEIERERSTERNTPATBEARETIGREENTIVES OF

EUROPEAN DERAEOCORIS OLIVACEUS (FABRICIUS)

(HETEROPTERA: MIRIDAE)*

Yasunaga, T. & Y. Nakatani, 1998. The eastern Palearctic relatives of European Deraeocoris oli-

vaceus (Fabricius) (Heteroptera: Miridae). — Tijdschrift voor Entomologie 140 [1997]: 237-

247, figs. 1-25. [1ssN 0040-7496]. Published 26 March 1998.

The eastern Palearctic deraeocorine plant bugs which have been incorrectly considered conspe-

cific with the European Deraeocoris olivaceus (Fabricius) are revised. Deraeocoris brachialis Stäl,

which has been regarded as a synonym of olivaceus, is reinstated as valid, diagnosed and re-

described. Two new species, D. gibbantennatus from the Primorskij Kraj, Russia and D. ery-

thromelas from Shikoku and Kyushu, south-western Japan, are diagnosed and described. The

European 'true' D. olivaceus is diagnosed, and a key to the species is provided. The zoogeogra-

phy of the four allied species is discussed.

Correspondence: Tomohide Yasunaga, Biological Laboratory, Hokkaido University of Educa-

tion, Ainosato 5-3-1, Sapporo, 002 Japan. E-mail: [email protected]

Key words. — Heteroptera; Miridae; revision; Deraeocoris olivaceus, D. brachialis new species;

key; zoogeography.

*Contribution from the Russia/Japan Cooperative East-Asian Entomological Program, No. 49.

Deraeocoris olivaceus (Fabricius, 1777) is a conspic-

uous deraeocorine plant bug species, originally de-

scribed from Germany, which is well known in Eu-

rope, because of its large size and predaceous habit.

Stäl (1858) described its relative, D. brachialis, from

Irkutsk in southern Siberia, but this species was sub-

sequently synonymised with olivaceus (Kerzhner

1988a). Although Kerzhner (1988b) referred to some

different colour patterns in the specimens from the

Russian Far East, he considered this as intraspecific

variation. His treatment has been followed by subse-

quent authors (Miyamoto & Yasunaga 1989, Yasuna-

ga 1990, Josifov 1992, Yasunaga & al. 1993, Lee &

al. 1994, Vinokurov & Kanyukova 1995a, 1995b).

Yasunaga (1994) illustrated the male genitalia of

specimens with different coloration from Japan and

the Russian Primorskij Kraj, and mentioned that,

what had been called o/ivaceus in this region, includes

at least three different species and is in need of revi-

sion.

Recently, through the courtesy of Prof. M. Josifov

of Bulgarian Academy of Sciences, we obtained sever-

al specimens of ‘true’ European olivaceus, and, upon

examination, the eastern Palearctic specimens were

found to differ significantly in some structures from

the European ones. Therefore, the populations occur-

ring in eastern Eurasia (southern part of Siberia in-

cluding Irkutsk, the Russian Far East, north-eastern

China, Mongolia and Japanese Honshu) cannot be

regarded as conspecific with those of Europe (= true

olivaceus), but should be called brachialis. On the

other hand, specimens from Shikoku and Kyushu,

south-western Japan, correspond to neither olivaceus

nor brachialis in having a bright red and densely pu-

bescent dorsum, impunctate and setose scutellum,

noticeably pubescent hemelytra and different struc-

ture of the male genitalia; they are considered to be-

long to an undescribed species. Further, one of the

authors, Yasunaga, collected a pair of specimens with

entirely blackish bodies from southern Primorskij

Kraj, Russia. Such remarkable blackish specimens

have previously been regarded as a variant of olivaceus

(= brachialis) only (Kerzhner 1988b; Kerzhner pers.

comm.), although distinct differences are found in

the female antennal segmentl and male genitalia as

described below; therefore, these specimens cannot be

237

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Figs. 1-8. Parameres of Deraeocoris spp. — 1-2, D. olivaceus from Bulgaria; 3-8, D. brachialis, 3-4, S. Primor’je, Russia; 5-6,

Mt. Sasagamine, Niigata Pref., Honshu, Japan; 7-8, Mts. Yatsugatake, Nagano Pref., Honshu, Japan. — 1, 3, 5 & 7, left para-

mere; 2, 4, 6 & 8, right paramere. Scales: 0.2 mm.

considered conspecific with olivaceus or brachialis.

In the present paper, the true Deraeocoris olivaceus

(Fabricius), which is restricted to Europe and adja-

cent regions, is diagnosed, and the specific name, D.

brachialis Stäl, is reinstated as valid for the eastern

Palearctic specimens. Two new species, D. gibbanten-

natus from southern Primorskij Kraj, Russia and D.

erythromelas from Kyushu and Shikoku, Japan, are

described. A key is provided to distinguish D. oli-

vaceus and its allies, and the zoogeography is also dis-

cussed.

MATERIAL AND METHODS

In general dried specimens were used. Depositories

of material examined are abbreviated in the text as

follows: Entomological Laboratory, Faculty of Agri-

culture, Kyushu University, Fukuoka, Japan (ELKU);

238

Biological Laboratory, Hokkaido University of Edu-

cation, Sapporo, Japan (HUES); Dr. Miyamoto’s per-

sonal collection, Fukuoka, Japan (Mc); Museum of

Nature and Human Activities, Hyogo, Japan

(MNHA); National Institute of Agro-Environmental

Science, Tsukuba, Japan (NIAES); Zoological Insti-

tute, National Science Museum, Tokyo, Japan

(NsMT); Entomological Laboratory, University of Os-

aka Prefecture, Sakai, Japan (vor); Zoological Insti-

tute, Russian Academy of Sciences, St. Petersburg

(ZMAS).

All measurements in the text are given in millime-

tres. In the synonymic listings only original references |

and selected references subsequent to Carvalho’s cat-

alogue (Carvalho 1957) are cited for each taxon (de-

tailed synonymic listing provided by Schuh 1995).

YASUNAGI & Nakatani: Eastern Palaearctic Deraeocoris

| Figs. 9-12. Vesicae of Deraeocoris spp. — 9, D. olivaceus from Bulgaria; 10-12, D. brachialis, 10, S. Primor je, Russia; 11,

| Marunuma, Gunma Pref., Honshu, Japan; 12, Mts. Yatsugatake, Nagano Pref., Honshu, Japan. Scales: 0.5 mm.

Terms for descriptions of male genitalia

Male genitalia of the plant bugs form, needless to

say, very important taxonomic characters, and it is

necessary to describe and/or illustrate them properly.

Because Deraeocoris olivaceus and allied species pos-

sess five homologous sclerites on the vesica, we use

‚here alphabetical terms to avoid misunderstanding

(see characters A-E in figs. 9-12, 20, 24).

Sclerite A is the most conspicuous, horn-like ap-

pendage, and may be interpreted as the ‘spiculum’.

| The vesicae of olivaceus and allies are principally di-

| vided into three membranous lobes, and the first lobe

| is accompanied by two flat sclerites (B & C), which

are sometimes fused together basally. ‘Sclerite D’ is

| situated on another lobe, tapered and pointed apical-

ly. The fifth sclerite (E) is a partly or entirely sclero-

| tized extension of the third lobe. These five sclerites

form important characters for the classification of D.

olivaceus and allies.

TAXONOMY

Deraeocoris olivaceus (Fabricius)

(figs. 1-2, 9, 25)

Cimex olivaceus Fabricius, 1777: 300.

Deraeocoris olivaceus — Reuter 1888: 648; Kiritschenko

1951: 159, fig. 166; Carvalho 1957: 71; Southwood &

Leston 1959: 206; Leston & Woodroffe 1961: 188; Wag-

ner & Weber 1964: 40; Wagner 1971: 36; Günther &

Schuster 1990: 369; Melber & al. 1991: 92; Gorczyca

1992: 91; Schuh 1995: 615.

Deraeocoris brachialis — Kiritschenko 1951: 160.

Diagnosis

Recognised by the smallest body among the rela-

tives (up to 10.7 mm in length; redescriptions below

indicating the dimension ranges from 8.5-10.5 mm),

narrow head, wide vertex (0.45-0.50 times as wide as

head including eyes), generally short antennal seg-

ments, relatively incrassate antennal segment II (api-

239)

TiJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

cal part more than 1.5 times as thick as base), bicol-

orous antennal segment III (basal part widely pale

and apical part infuscate), and short legs. Very vari-

able species in coloration. In Europe, its closest rela-

tive is D. trifasciatus (Linnaeus), from which olivaceus

is easily separated by the significantly smaller size and

different coloration.

Detailed descriptions were provided by several Eu-

ropean authors (Reuter 1896; Wagner 1952, 1971;

Wagner & Weber 1964, etc.).

Description

Male genitalia. — Parameres as in figs. 1-2; protu-

berance of left paramere not strongly curved (fig. 1).

Vesica (fig. 9) comparatively small in general shape;

sclerite A rather slender and straight; sclerite C short,

accompanied with second triangular process; sclerite

D elongate.

Dimensions. — 4/9 : Body length (from anterior

apex of tylus to posterior tip of membrane) 9.90-

10.68/ 10.02-10.20; head width including eyes 1.40-

1.48/ 1.50-1.53; vertex width 0.65-0.68/ 0.75; length

of antennal segment I 0.90-0.98/ 0.97-1.00, II 2.97-

3.18/ 3.25-3.28, III 1.00-1.05/ 1.05-1.12, IV 0.63-

0.72/ 0.70-0.73; rostral length 3.70-3.83/ ?; mesal

pronotal length including collar 1.92-2.22/ 2.15-

2.23; basal pronotal width 3.27-3.58/ 3.47-3.68;

maximum width across hemelytra 4.15-4.63/ 4.50-

4.73; length of hind femur 3.00-3.73/ 3.12-3.33, tib-

ia 4.00-4.25/ 4.00-4.20, tarsus 1.07-1.18/ 1.12-1.18;

length of hind tarsomere I 0.48-0.50/ 0.50-0.51, II

0.36-0.38/ 0.37-0.40, III 0.45-0.48/ 0.48-0.50.

Distribution

Europe: central part to the Mediterranean area,

Britain, and the European part of Russia, west of the

240

Ural range, and North Africa: Algeria. This species

appears to be restricted to the western Palearctic Re-

gion.

Biology

D. olivaceus has been reported to be associated with

Rosaceae, such as Crataegus, Malus, Pirus and Prunus

(Wagner 1952, 1971, Wagner & Weber 1964), and

in spite of being principally predaceous, it has been

observed to feed on young hawthorn berries (Cratae-

gus) (Southwood & Leston 1959). In Bulgaria, Dr.

Josifov collected two males from Quercus cerris L.(Fa-

gaceae), which we could not confirm as the host

plant.

Material examined. — BuLcaria: 34, 29, Kaloti-

na, 20.vi.1993, M. Josifov (Hues); 2d, Plana u.,

Kokaljane, 1,000 m, on Quercus cerris, 23.vi.1968,

M. Josifov (HUES).

Deraeocoris brachialis Stäl sp. rev.

(figs. 3-8, 10-13, 25)

Deraeocoris brachialis Stäl 1858: 185; Carvalho 1957: 61;

Kulik 1965a: 170; Kulik 1965b: 50; Kerzhner 1978: 37.

Deraeocoris olivaceus — Kerzhner 1988a: 67; Kerzhner

1988b: 794; Miyamoto & Yasunaga 1989: 158; Yasuna-

ga 1990: 665; Josifov 1992: 117; Yasunaga & al. 1993:

146, fig. 7b; Lee & al. 1994: 7; Yasunaga 1994: 41; Vi-

nokurov & Kanyukova 1995a: 13; Vinokurov &

Kanyukova 1995b: 85.

Diagnosis |

Recognised by the enlarged body (more than 11

mm in most specimens), wide head, narrow vertex,

long antennal segments, less incrassate antennal seg- |

ment II, unicolorous dark antennal segment III (only |

extreme base rarely narrowly pale), and long legs.

Very variable species in coloration.

This eastern Palearctic species is a close relative of

olivaceus, from which it can be distinguished by the

characters as diagnosed above and different structure

of the male genitalia. Reuter (1896) provided a de-

tailed redescription, in which he showed that

brachialis is significantly larger than olivaceus. But his

key is currently not applicable because the variable

coloration was used as the key character.

Redescription

Body elongate oval (6 )/ oval and tumid (9); dor-

sal surface usually brownish, shining, widely punc-

tate. Head brown with dark symmetrical marking, or

entirely darkened, shining, oblique and rather wide;

vertex 0.37-0.45 (d)/ 0.39-0.46 (9) times as wide as

head including eyes, with sparse, silky, very short pu-

bescence; tylus, jugum, buccula and gena with dense,

silvery, suberect pubescence. Antenna dark chestnut

brown; segment I and basal part of II sometimes

widely brown or reddish brown; segment II less in-

crassate toward apex, its apical part less than 1.5 times

as wide as base; segments III and IV filiform, extreme

base of segment III rarely pale; length of segments I-

IV (6/9): 1.05-1.18/ 0.99-1.25, 3.40-3.78/ 3.23-

4.00, 1.22-1.50/ 1.30-1.55, 0.76-0.90/ 0.77-0.88.

Rostrum shiny dark chestnut brown, sometimes part-

ly pale, reaching or slightly exceeding middle coxa.

Pronotum very variable in coloration, pale brown

(usually with darkened calli) to widely darkened,

shining, uniformly with dense, fuscous punctures and

laterally with silky, erect pubescence, posterior mar-

gin usually impunctate; collar sombre yellowish

brown to fuscous, shagreened or pruinose, bearing

silky, erect pubescence; mesoscutum shiny fuscous,

with shagreened lateral parts, bearing sericeous pu-

bescence; scutellum shiny dark brown, with pale lat-

eral and posterior margins forming v-shape, shallowly

but distinctly punctate, bearing a few, silky, very

short setae. Hemelytra brown, or pale, reddish, pur-

plish or dark brown, shining, almost glabrous, with

dense, dark punctures; apical part of corium widely

infuscate; ventral surface of embolium densely pubes-

cent; cuneus yellow, brown, orange or red, with al-

| Ways infuscate apical 1/3-1/2 parts; membrane som-

bre pale brown to brown, narrowly pale near apex of

cuneus. Legs uniform with brownish, suberect setae;

femora widely chestnut brown or dark brown, or pale

brown with dark patches; tibiae reddish brown to fus-

cous, each with two yellow bands; tarsi dark brown,

with dense, blush-like setae inward; length of hind fe-

mur, tibia and tarsus (64/9): 3.52-4.05/ 3.25-4.33,

4.75-5.33/ 4.42-5.63, 1.25-1.38/ 1.15-1.50; that of

hind tarsomeres I-III (4/9): 0.50-0.60/ 0.54-0.70,

0.43-0.48/ 0.37-0.51, 0.48-0.60/ 0.44-0.60. Ab-

YASUNAGI & Nakatani: Eastern Palaearctic Deraeocoris

domen pale brown, reddish brown or dark chestnut

brown.

Male genitalia. — Somewhat variable in shape.

Parameres as in figs. 3-8; protuberance of left para-

mere distinctly curved. Vesical sclerite A is strong,

broad and apically hooked; sclerite C developed; scle-

rite D rather shortened (figs. 10-12).

Dimensions. — 8/2: Body length 10.42-12.90/

10.02-12.20; head width including eyes 1.57-1.80/

1.57-1.75; vertex width 0.62-0.70/ 0.75-0.83; rostral

length 3.65-4.25/ 3.62-4.28; mesal pronotal length

including collar 2.25-2.48/ 2.00-2.73; basal pronotal

width 3.72-4.05/ 3.42-4.45; maximum width across

hemelytra 4.67-5.30/ 4.47-5.50.

Distribution

Russia (possibly east of Altai and south of Stanovoj

ranges: South Siberia including Irkutsk and Tuva,

continental Far East and South Sakhalin), Japan

(North and Central Honshu), Mongolia, North-east

China, North- and South-Korea.

Biology

Kerzhner (1978) recognised Salix sp. (Salicaceae)

as its host plant, and Yasunaga (1994) also recorded

willow (Salix spp.) and alder (Alnus spp., Betulaceae).

Predation on larvae of chrysomelid beetles (e.g., Go-

nioctena japonica Chujo et Kimoto, 1960 on alder

and Chrysomela vigintipunctata (Scopoli, 1763) on

willow) has been frequently observed, and cannibal-

ism was also reported (Yasunaga 1990).

Ussurijsk Reserve, 15.vii.1990, T. Saigusa (HuEs); 1d, Us-

surijsk Reserve, W. border, 12-13.vii.1993, light trap, T.

Yasunaga (HUES); 24d, 169, 17 km SW of Krounovka, nr.

Mt. Medvezh’ja, on alder & willow, 5-9.vii.1993, T. Ya-

sunaga (HUES); 2d, Rjazanovka, Khasanskij Dist, 26-

27.vii.1993, light trap, T. Yasunaga (HUES). — CHINA: 1d,

N. Manchuria (= current Heilongjiang Prov. of NE. China),

1940, A. Loukashikin (NIAES). — JAPAN: Honshu: 1d, 19,

Mt. Sasagamine, Niigata Pref., 20-25.vii. 1993, S. Yoshizawa

(nues); 19, Nikko, Tochigi Pref., 19.vii. 1949, J. Aoki (nı-

AES); 19, same locality, 14.vi.1950, M. Yano (Nies); 2d,

same locality, 17-19.vii.1959, T. Maenami (NIAES); 14,

Nebasawa, Oku-nikko, Tochigi Pref., 11.vii.1937, G. Ya-

mamoto (NIAES); 19, Ozegahara, Gunma Pref., 9.vii. 1951,

H. Hasegawa (NIAES); 34, 3 9, same locality, 9-10.vii.1978,

M. Tomokuni (nsMT); 19, Fujimishita, nr. Ozegahara,

Gunma Pref., 22.vii.1987, Y. Notsu (NSMT); 16,

Marunuma, Gunma Pref., 7.vii.1984, M. Hayashi (Hues);

1d, 42, Mt. Kanayama, Yamanashi Pref., 6.vi.1952, M.

Ohno (NIAES); 12, Masutomi, Yamanashi Pref.,

28.vii.1957, S. Kimoto (mc); 19, Daimonzawa, Mt.

Akadake, 2,000-2,500 m alt., Mts. Yatsugatake, Yamanashi

Pref., 30.vii.1987, T. Yasunaga (Hues); 44, Nippara,

Tokyo, 12.vii.1986, M. Tomokuni (NsMT); 16, Komoro,

Nagano Pref., 29.vii.1959, T. Maenami (nIaEs); 19,

Omachi, Nagano Pref., 26.vi.1977, M. Owada (NsMT); 14,

Mikasa Park, Karuizawa, Nagano Pref., 31.vii.1968, H.

241

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Ono (NsMT); 1d, Ueda C., Nagano Pref., vi.1934, S.

Miyamoto (ELKU); 24,29, Todai, Kami-ina, Nagano Pref.,

8-11.vii.1978, M. Owada (NsMT); 19, Sugadaira, Nagano

Pref., 18.viii.1956, K. Baba (Nraes); 16, Shiga Plateau,

Nagano Pref., 6.viii.1974, Y. Hori (NsMT); 19, same locali-

ty, 3-6.viii.1982, M. Sato (NsMT); 36, 49, Mt. Azusayama,

Kawakami Vil., Nagano Pref., 5.vii.1980, Y. Kurosawa

(NsMT); 36, 19, Minoto, Mt. Akadake, 2,000-2,500 m

alt., Mts. Yatsugatake, Nagano Pref., 27.vii.1987, T. Ya-

sunaga (HUES); 16, Inakoba, Mts. Yatsugatake, Nagano

Pref., 18.vii.1939, H. Hasegawa (NIAES); 26, Sanjiro, Ut-

sukushigahara, Nagano Pref., 30.vii.1954, K. Kai (NIAES);

19, Shimashima Valley, Nagano Pref., 3-5.viii.1940, K.

Kurosa (NIAES); 1d, same locality, 7.viii.1955, H. Hasega-

wa (NIAES); 1, Iwana-dome, Shimashima Valley, Nagano

Pref., 14.vii.1982, Y. Abe (murs); 19, Tokugo Pass,

Nagano Pref., 7.viii.1955, H. Hasegawa (NIAES); 39,

Kamikochi, Japan Alps, Nagano Pref., 21.vii.1917, T. Esaki

(ELKU); 1d, 29, same locality, 15.vii.1945, M. Yano; 29,

cality, 12.viii.1952, T. Nakane (Nrars); 2d, 29, same lo-

cality, 3-7.viii.1955, I. Hattori (NIAES); 19, same locality &

collector, 18-19.vii.1956 (NrAES); 28, 3 9, same locality &

collector, 14-18.vii.1960, I. Hattori (NIAES); 29, same lo-

cality, on alder, 21.vii.1957, M. Ohno (Nrars); 19, same lo-

cality, 3.viii.1956, M. Sato (nsMT); 64, 102, Mt. Nyugasa,

Nagano Pref., 29-31.vii.1962, S. Miyamoto (mc); 19, Mt.

Senjodake, Nagano Pref., 3.viii.1958, H. Yamamoto (mc);

1d, Isedaki, Komagatake, Nagano Pref., 1.viii.1962, S.

Miyamoto (Mc); 2d, Nagawa Vil, Nagano Pref,

21.vii.1994, Y. Sawada & T. Ueda (vor); 1d, 19, Kiso-

Ontake, Nagano Pref., 20.vii.1952, H. Hasegawa (NIAES);

29, Hirayu, Gifu Pref., 22-23.vii.1946, K. Ohbayashi

242

> ALOE mm

Figs. 16-20.

Deraeocoris gibbantennatus. —

16, female head in left lateral

view; 17, female antennal seg-

ment I; 18, left paramere; 19,

right paramere; 20, vesica.

19, same locality, 26.vii.1986, S. Miyakawa

(NSMT);

(nsMT); 16, Mt. Wasamata, Nara Pref., 3.viii.1995, T. Hi-

rowatari (vor); 18,29, Kobe C., Hyogo Pref., 20.vi.1940,

K. Kurosa (NIAES).

Deraeocoris gibbantennatus sp. n.

(figs. 14, 16-20, 25)

Deraeocoris olivaceus — Kerzhner 1988b: 794; Yasunaga

1994: 43, fig. 14.

Type Material. — Holotype d, 17 km SW of

Krounovka, near Mt. Medvezh'ja, S. Primorskij Kraj,

Russia, on Alnus sp., 5-7.vii.1993, T. Yasunaga

(zmas). — Paratype: 1 9, same data as for holotype, 8-

9.vii.1993 (HUES).

Diagnosis

This species is easily distinguished from other rela-

tives by the entirely blackish dorsum (fig. 14) and pe-

culiar hump-like projection on the female antennal

segment I (figs. 16-17). The shape of the vesical scle-

rite is also unique to this species.

Description

Body almost entirely shiny black, oblong-oval (9 )/

oval (2); dorsal surface distinctly punctate; ventral —

surface uniformly darkened. Head black, shining,

with silvery, suberect, short pubescence; vertex some-

what pale near inner margin of eye, relatively narrow, :

0.36-0.37 times (&)/ 0.43 times (9) as wide as head

including eyes. Antenna entirely blackish brown,

densely setose; 2 segment I with a dorsal, hump-like

projection at basal 1/4; segment IT rather incrassate

toward apex, apical part 1.6-1.7 times as thick as base;

segments II and IV filiform; length of segments I-IV

(6/2): 1.25/ 1.25, 3.88/ 3.96, 1.50/ 1.48, 0.83/

0.83. Rostrum shiny dark chestnut brown, reaching

middle coxa.

Pronotum black, shining between punctures, with

brown, upright setae laterally, posterior margin some-

what pale; collar shagreened or pruinose, bearing

silky, erect pubescence; mesoscutum black, sha-

greened laterally, with sparse, silky, short pubescence;

scutellum shiny dark chestnut brown, with yellow ex-

treme apex, timid, punctate, bearing very sparse,

silky, short pubescence. Hemelytra unicolorous shiny

black, densely punctate, with silky, short pubescence

along lateral margin; ventral surface of embolium

densely pubescent. Leg dark brown, densely setose;

each tibia with two yellowish bands; tarsi with dense,

blush-like setae inward; length of hind femur, tibia

and tarsus (4/9): 4.00/ 4.03, 5.25/ 5.33, 1.40/ 1.43;

that of hind tarsomeres I-III: 0.55/ 0.60, 0.48/ 0.52,

0.53/ 0.61. Abdomen entirely shiny blackish.

Male genitalia. — Parameres as in figs. 18-19. Vesi-

ca apparently specialised in form (fig. 20); sclerite A

elongate, tapered at apex; sclerite B narrow, widely

fused with a membranous lobe; sclerite C reduced, di-

vided into several smaller plates; sclerite C weak.

Dimensions. — 6/9: Body length 12.90/ 11.94;

head width including eyes 1.78/ 1.74; vertex width

0.65/ 0.75; rostral length 4.00/ 4.28; mesal pronotal

length including collar 2.35/ 2.63; basal pronotal

width 4.00/ 4.35; maximum width across hemelytra

5.08/ 5.75.

Etymology

| From Latin, gibba or gibbus (= hump) in combina-

(tion with antenna, referring to the characteristic

jhump-like projection exhibited on the female anten-

jnal segment I.

Distribution

Russian Far East (S. Primorskij Kraj).

Biology

D. gibbantennatus was found on Alnus sp. (Betu-

\laceae), with which it appears to be associated. It was

{collected together with D. brachialis, but in lower

‚population densities.

YASUNAGI & NAKATANI: Eastern Palaearctic Deraeocoris

Deraeocoris erythromelas sp. n.

(figs. 15, 21-25)

Deraeocoris olivaceus— Yasunaga & al. 1993: 146, fig. 7a; Ya-

sunaga 1994: 42, fig. 15.

Type material. — Holotype d, Momiki, 600-700

m alt., Izumi Vil., Kumamoto Pref., Kyushu, Japan,

at light, 6.vii.1991, K. Yahiro (Hugs). — Paratypes:

Shikoku: 14, Nishikuma, Monobe Vil., Kochi Pref.,

30.vii.1994, M. Takai (Hugs); 19, Teragawa, Hon-

gawa Vil., Kochi Pref., light trap, 18.vii.1993, I. Ya-

mashita (HUES); 16, same locality and collector,

25.vii.1991 (vor); 12, Hongawa Vil., 800-1,400 m

alt., Kochi Pref., on Betula sp., 23.vii.1996, T. Ya-

sunaga (HUES); 19, Mt. Tsurugi, Tokushima Pref.,

light trap, 10.vii.1993, I. Yamashita (nues); 19, Mt.

Odamiyama, Ehime Pref., 12.vii.1994, M. Takai

(HUES). — Kyushu: 14, Mt. Shiratori, Izumi Vil., Ku-

mamoto Pref., 21.vi.1987, R. Noda (Hugs); 19,

same locality, 19.vii.1992, Y. Nakatani (vor); 19,

Mt. Yamaingiri 1,100 m alt., Izumi Vil., Kumamoto

Pref., on Salix sp., 28.vi.1992, T. Yasunaga (fig. 15,

HUES); 1d, same locality, light trap, 16.vii.1994, R.

Matsumoto (Hugs); 12, Mt. Sobo, Oita Pref.,

25.vii.1978, Y. Shono (MNHA); 19, Kamihori,

Miyazaki Pref., 15.vii.1967, S. Tawara (Mc).

Diagnosis

This species is easily distinguished from other rela-

tives by the bright red and partly infuscate dorsum,

silky suberect pubescence on the impunctate scutel-

lum, and uniformly pubescent hemelytra, in addition

to having a quite restricted distribution.

Description

Body elongate oval, large; dorsal surface noticeable

bright red, partly or sometimes widely infuscate, pro-

vided with silky, suberect pubescence; ventral surface

widely darkened. Head dark chestnut brown, shin-

ing, with silky, suberect pubescence; vertex with a

pale spot near inner margin of each eye, comparative-

ly wide, 0.43-0.44 (4 )/ 0.50-0.52 (®) times as wide

as head including eyes; tylus, jugum, buccula and

gena bearing rather dense, silky, long pubescence.

Antenna dark chestnut brown, with dense, erect, long

setae; apical part of segment II more than 1.5 times as

thick as base; segments III and IV somewhat paler, fil-

iform; length of segments I-IV (6/9): 1.12-1.15/

1.22-1.28, 3.52-3.75/ 3.50-4.13, 1.18-1.25/ 1.26-

1.40, 0.75-0.78/ 0.89-0.91. Rostrum shiny dark

chestnut brown, reaching middle coxa.

Pronotum usually bright red, shining, sometimes

tinged with purple or brown, uniformly bearing

brownish, erect pubescence, with dark punctures; cal-

li sometimes infuscate; collar dark grey, shagreened,

with dense, brownish, erect setae; mesoscutum black-

243

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

Fig. 21. Deraeocoris erythromelas, dorsal habitus. Scale: 3 mm.

ish, laterally with greyish pollinosity, bearing sparse,

silky pubescence; scutellum red, shining, triangularly

darkened medially, impunctate or very shallowly

punctate, uniformly clothed with silky, erect pubes-

cence. Hemelytra shiny red, sometimes tinged with

purple or brown, punctate, uniformly clothed with

silky, suberect pubescence; corium and clavus partly

or sometimes widely infuscate; cuneus bright red,

with darkened apical 1/3-1/2 parts; membrane som-

bre greyish brown, with a pale spot near apex of

cuneus. Leg dark chestnut brown; femora and tibiae

with dense, brownish, erect setae; each tibia with

two yellowish white bands; tarsi with dense, blush-

like setae; length of hind femur, tibia and tar-

(sus (6/9): 3.65-3.75/ 3.80-4.25, 4.72-4.93/ 4.97-

15.38, 1.20-1.25/ 1.30-1.38; that of hind tarsomeres

LIII (3/2): 0.48-0.50/ 0.57-0.60, 0.42-0.46/ 0.44-

10.48, 0.49-0.53/ 0.55-0.60. Abdomen shiny dark

brown.

| Male genitalia. — Parameres (figs. 22-23) distinctly

setose; left paramere densely furnished with long sen-

|sory setae (fig. 22). Vesical sclerites A and B long and

slender; sclerite C well developed; sclerite D extended

\basally (fig. 24).

Dimensions. — 6/9: Body length 10.98-12.00/

110.92-12.03; head width including eyes 1.60-1.65/

11.66-1.73; vertex width 0.70-0.73/ 0.85-0.88; rostral

length 3.57-3.93/ 3.82-4.08; mesal pronotal length

including collar 2.35-2.43/ 2.57-2.68; basal pronotal

width 3.85-3.90/ 3.97-4.38; maximum width across

hemelytra 4.81-4.99/ 5.22-5.88.

Etymology

From Greek erythro- (= red) and melas (= black),

referring to the bright red and partly infuscate body

of this new species.

YASUNAGI & NAKATANI: Eastern Palaearctic Deraeocoris

Figs. 22-24.

Male genitalia of Deraeocoris

erythromelas. — 22, left para-

mere; 23, right paramere; 24,

vesica.

Distribution

Japan (central montane areas of Shikoku and

Kyushu).

Biology

D. erythromelas has been found on Betula sp. (Be-

tulaceae) and Salix spp. (Salicaceae). Predation on lar-

vae of the chrysomelid beetle, Chrysomela vigintipunc-

tata (Scopoli, 1763) (Coleoptera) on willow was

observed in Kyushu. A last-instar nymph was found

to have been parasitized by an undetermined nema-

tode (Yasunaga, unpublished data).

Key TO SPECIES OF D. olivaceus AND ALLIES

1. Scutellum shiny and impunctate, with silky,

suberect pubescence; hemelytra distinctly pubes-

CONE) „Ir RIE he ER D. erythromelas

—. Scutellum distinctly punctate, glabrous; hemelytra

lacking noticeable pubescence except on embolium

2. Dorsal surface entirely shiny black; ® antennal seg-

ment I with a peculiar dorsal hump-like projection

SO cece E D. gibbantennatus

— Dorsal surface partly or widely reddish, brownish

or yellowish; 2 antennal segment I lacking such

projection

3. Body larger, usually more than 11 mm in length;

antennal segment I longer, more than 1 mm; an-

tennal segment III unicolorously darkened, or only

extreme base rarely pale; eastern Palearctic species

found on Salicaceae and Betulaceae . .D. brachialis

— Body smaller, up to 10.7 mm in length; antennal

segment less than 1 mm; antennal segment III bi-

colorous, dark with widely pale base; western Pale-

arctic species associated mainly with Rosaceae ...

See Ne een D. olivaceus

245

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

brachialis

ZOOGEOGRAPHY (fig. 25)

The four treated species have in the past been re-

presented as a single species, o/ivaceus, which was con-

sidered to be a widely distributed Palearctic species.

After a careful check of the previous distributional

records, we realized that D. olivaceus, D. brachialis

and D. erythromelas are obviously allopatrically dis-

tributed. D. olivaceus is restricted to the western

Palearctic Region including the Mediterranean area,

D. brachialis is distributed in southern Siberia, the

Russian Far East, Mongolia, the Korean Peninsula

and Honshu, Japan, and D. erythromelas has the

southernmost distribution, being endemic to central

montane areas of Shikoku and Kyushu Islands,

south-western Japan. Only D. gibbantennatus was

found sympatric with brachialis in the Russian Pri-

morskij Territory.

Of the four species, D. olivaceus and D. brachialis

are the most closely related species. However, none of

these has been reported from the region between the

Ural and Altai ranges. The mountains are considered

to form a barrier, dividing the distribution ranges of

the western D. olivaceus and eastern brachialis.

Our observations and previous reports indicate

that these two species have different host preferences;

D. olivaceus is associated with rosaceous host plants

(Southwood & Leston 1959; Wagner 1952, 1971;

Wagner & Weber 1964), while D. brachialis has been

found only on alder (Betulaceae) and willow (Sali-

caceae). As pointed out by Todo & Yasunaga (1996)

and Yasunaga (1996), many deraeocorine species, in

spite of being predaceous, live on restricted plant

species. The different preference for plant species may

be a diagnostic character.

Kerzhner (1988b) regarded specimens, here named

246

Fig. 25.

Distribution of Deraeocoris oli-

vaceus and allies.

erythromelas

as D. gibbantennatus, as conspecific with D.

brachialis, but a peculiar hump-like dorsal projection

on the female antennal segment I is considered to be

an autapomorphy for gibbantennatus (see figs. 16-17).

The shape of sclerites B and C on the male vesica is

also unique to gibbantennatus, which is, therefore,

here regarded to be a good species. |

ACKNOWLEDGMENTS

We wish to express our sincere gratitude to Dr. S.

Miyamoto (Fukuoka, Japan) for his continuous ad-

vice and encouragement, and to Dr. I. M. Kerzhner

(zmas) for his kind information. Yasunaga also

thanks Drs. N. Kurzenko, A. Lelej, Y. Tsuistjakov, V.

Makarkin, A. Egorov, E. Kanyukova and V. Sidrenko

(Institute of Biology and Pedology, Far Eastern

Branch of Russian Academy of Sciences, Vladivostok) |

for their kind assistance during the expedition in the

Russian Far East, and especially Dr. V. Makarkin for!

the Russian translation. Most specimens used in this |

paper were loaned from or donated by the following

individuals, to whom we are much indebted: Dr. M.!

Josifov (Bulgarian Academy of Sciences, Sofia); Dr. S. |

Miyamoto; Dr. M. Tomokuni (NsMT); Dr. T. Mat- |

sumura and Mr. K. Konishi (NIAES); Dr. A. Nakan-

ishi and Dr. Y. Sawada (MNHA); Dr. Y. Abe (Kyoto!

Prefectural University); Mr. S. Gotoh (Tanabe, |

Japan); Dr. T. Hirowatari, Mr. T. Ueda and Mr. Yel

Sawada (vor); Dr. H. Yahiro (Shiga, Japan); Mr. M.

Takai and Mr. I. Yamashita (Nankoku, Japan); Mr.

R. Noda (Fukuoka, Japan); Dr. S. Kamitani (ELKU); |

Prof. T. Saigusa, Mr. T. Nakamura, Mr. K. Masuna- |

ga, Mr. K. Yoshizawa, and Mr. R. Matsumoto

(Biosystematics Laboratory, Kyushu University,

Fukuoka, Japan). |

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*Indirect citation.

Received: 18 December 1996

Accepted: 15 May 1997

247

CLIS LE

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BOOK REVIEW

Stig Andersen, 1996. The Siphonini (Diptera: Tachi-

nidae) of Europe. — Fauna Entomologica Scandi-

navica, Volume 33, E.J. Brill. 148 pp., 6 colour

plates, 275 text figs. Hardback [1sBN 90 04 10731

21. Price HFL 114.50 excl. postage.

Volume 33 of the well-known series ‘Fauna Ento-

mologica Scandinavica’ deals with the European

species of the tribe Siphonini of the family Tachi-

nidae. It is a typical member of the series, considering

the layout, many text figures, few colour plates and

information on distribution of species in Scandinavia.

On the other hand, this volume differs from all the

others in the fact that its scope is restricted to one

tribe and that all European species of this tribe have

been included. Moreover, there are quite a lot of tax-

onomic novelties in this volume, so that it is best to

consider this volume to be a full-grown review of the

European Siphonini.

Whereas the taxonomy of the family Tachinidae is

considered by most dipterologists to be difficult, that

of the tribe Siphonini is even more complicated.

Especially the genus Szphona is quite notorious in this

respect. Until the sixties of this century only four

European species of Siphona were generally accepted.

Mesnil (in Lindner) raised the number of European

species to 13, still leaving many problems unsolved.

Stig Andersen must be praised for continuing Mes-

inil's difficult task, leading to a review of the genus

\Siphona in 1982 containing 17 species. In his present

work, 20 European species of Siphona have been re-

lcognised.

The introductory chapters of the book deal with

the classification and the lifehistory of the Siphonini.

‚The part on classification and phylogeny is quite ex-

tensive, but rather technical for most readers. The

part on the life-history contains some interesting new

(material. Although many host-relations are still un-

discovered, it becomes more and more clear that very

similar species may have quite different hosts.

The main part of the book is dedicated to the tax-

bnomy of the 58 European species, starting with a key

fo genera. Next each genus is characterised, its distri-

pution on a world-scale is given, followed by a key to

he European species. For each species the typical fea-

tures, distribution and biology are discussed. The text

|s illustrated with simple but clearcut black-and-white

figures. Also all male genitalia have been illustrated. It

s my experience that the keys, in general, make a re-

BOOK REVIEW

liable identification possible. Compared with earlier

work, Andersen has introduced several new features,

making it sometimes possible to recognise even aber-

rant specimens. Some previously recognised species

are now considered to be only a variety of a very sim-

ilar taxon (Actia dubitata of crassicornis and Siphona

confusa of variata). Given the many intermediate

forms and the variability of these species, I totally

agree with Andersen's view. On the other hand, six

new species have been described. Some of the new

species can easily be recognised, whereas others are

very similar to related species. It is my experience that

Andersen tends to underestimate the variability in

these cases: for instance Dutch males of Peribaea

hertingi Andersen cannot be recognised with his key.

A major point of criticism concerns the nomencla-

ture used. Scandinavian dipterologists have a tenden-

cy of very strictly interpreting nomenclatural rules,

not seldomly leading to highly impractical results. To

introduce the name resinellae (Schrank, 1824) for the

species previously known as Actia nudibasis Stein,

1924 while Schrank’s types are lost is perhaps not too

big a problem. But to synonymise Siphona mesnili

Andersen, 1982 with Siphona confusa Mesnil, 1961

and S. confuse Andersen, 1982 with S. variata Ander-

sen, 1982 and then introducing Siphona variata for-

ma confusa Andersen’ is very confusing indeed, to say

the least. Finally, renaming the common S. cristata to

S. geniculata (De Geer, 1776) and the very common

S. geniculata to S. urbana (Harris, 1780) is really dis-

astrous. Although formally undoubtedly correct, with

this change of names Siphona geniculata can never be

used without ambiguity. Andersen refers to this only

as ‘some confusion and argues that …the name has

never had any longstanding and unambiguous usage

....Considering that S. geniculata and cristata had

very clearly been separated by Staeger already in 1849

(and repeatedly by Lundbeck in 1927), this defence is

just incorrect. In my opinion, these changes in names

could and must have been prevented.

Andersen's book is a mature review of the Euro-

pean Siphonini. In the vast majority of cases, it will

lead the more experienced student of Tachinidae to

reliable identifications in this complicated tribe.

Therefore, this book will be of great value to all seri-

ous students of the European Tachinidae. However,

for the non-specialists with only little material avail-

able, using this fine book of Stig Andersen may be

more cumbersome. especially in the genus Siphona.

[Theo Zeegers]

TIJDSCHRIFT VOOR ENTOMOLOGIE, VOLUME 140, 1997

BOOK REVIEWS

Ales Lastivka & Zdenek Lastivka, 1997. Nepticuli-

dae Mitteleuropas. Ein illustrierter Begleiter (Lepi-

doptera). — Konvoj, Brno. Hardback, 22X16 cm.

[1sBN 80-85615-61-4]. Price DEM 72.—. Can be or-

dered from Departrment of Zoology and Apicul-

ture, Faculty of Agronomy Muar, Zemedelskä 1,

cz-613 00 Brno.

This small booklet is meant as a diagnostic guide

for 184 species of Nepticulidae (leafmining Lepido-

ptera) occurring in Central Europe in a broad sense.

Apart from traditional Central Europe, it includes

species from Denmark, southern Sweden, The

Netherlands, eastern half of France, northern Italy,

Slovenia and Croatia. The book is therefore a wel-

come addition to the volume in the series Fauna En-

tomologica Scandinavica (vol. 23), which deals only

with the 121 species of Northwest Europe. The book

is written in German, the most important language in

the area covered by the book, although this may ham-

per its usefulness for some countries.

All species are treated on a separate page, with a

short diagnosis and notes on biology, distribution

and habitats. On the same page there are black-and-

white illustrations of a moth, male and female geni-

talia and leafmine, most original, excellently prepared

by the senior author. A few are redrawn from other

sources. There are no keys, but identification can be

reached by comparing the illustrations. This may be

somewhat difficult for those not yet familiar with the

family, especially since similar genitalia are not de-

picted on the same page. Apart from the descriptions,

there is a systematic list of hostplants and moth spe-

cies, a checklist of the species and a distribution cata-

logue.

The book contains descriptions of four new spe-

cies. Although a few more undescribed species are

currently known from this region, this book is fairly

comprehensive for a large part of Europe, excluding

the southern peninsula’s.

Minor points of criticism are the relatively simple

drawings of the female genitalia and the lack of

colour-plates of the adults. The latter, however,

would have increased the price enormously.

In conclusion, a nice booklet, with a well chosen ti-

tle: ‘Ein illustrierter Begleiter’ [an illustrated compan-

ion], to accompany fieldworkers and taxonomists.

[Erik J. van Nieukerken]

250

Vereniging Onderzoek Flora en Fauna, 1997. Jaar-

boek Natuur 1997, De winst- en verliesrekening

van de Nederlandse natuur. — KNNv Uitgeverij,

Utrecht/vorr, Wageningen. [Society for Research

on Flora and Fauna, 1997. Yearbook nature 1997.

Profit and loss account of the Dutch nature.].

Hardback, 17X25 cm. [1sBN 90-5011-108-4].

Price NLG 40.— excl. p&p. Order from KNNV

Publishers, Fax +31-30-236 89 07.

Why reviewing a Dutch book on Dutch nature in

an international entomological journal? There are

very few countries where so many amateur naturalists

are actively collecting data on the occurrence of

plants, animals and insects as in The Netherlands.

Maybe only in Britain there is a higher percentage of

the population active in this field. Usually specialists

in the various groups publish their results in their,

own circles. This book is a nice example of a broad

view on nature: it tries to answer questions like: is the

situation for nature in The Netherlands getting worse

or better? There are treatments for 15 groups: from.

macrofungi, seedplants, to bats, see-organisms and

birds. But there are also insects: Orthoptera (grass-

hoppers and crickets), Odonata (dragonflies), Coleo-

ptera: Carabidae (groundbeetles), Microlepidoptera,

and Rhopalocera (butterflies). In all chapters authors

try to estimate the trends in occurrence. Some species!

are declining, or extinct, others are increasing. The

differences between groups are large, insects belong. |

ing to the groups with severe decline, especially in

dragonflies and butterflies. It is nice to compare the

various distribution maps and tables with numbers of

gridcells in various periods.

Even for those not able to read Dutch, the book

gives an interesting view of nature in an overpopulat-

ed European country: many nature-lovers studying

little nature with few species. For foreign readers it is

a disadvantage that many captions to the figures give

trivial names only. Also an English summary is miss-

ing. Maybe this can be changed in future volumes in

this series. The price is surprisingly low.

[Erik J. van Nieukerken]

Volume 140 1997

Tijdschrift

voor

Entomologie

A journal of systematic and evolutionary

entomology since 1858

Netherlands Journal of Entomology

Published by the Netherlands Entomological Society