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REVUE SUISSE DE ZOOLOGIE

SWISS JOURNAL OF ZOOLOGY

REVUE SUISSE DE ZOOLOGIE

TOME 111—FASCICULE 3

Publication subventionnée par:
ACADEMIE SUISSE DES SCIENCES NATURELLES (SCNAT)
VILLE DE GENEVE
SOCIETE SUISSE DE ZOOLOGIE

VOLKER MAHNERT
Directeur du Muséum d’histoire naturelle de Genéve

CHARLES LIENHARD
Chargé de recherche au Muséum d’histoire naturelle de Genéve

Comité de lecture

Il est constitué en outre du président de la Société suisse de Zoologie, du directeur du
Muséum de Geneve et de représentants des instituts de zoologie des universités
suisses.

Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le
sujet étudié.

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.

Administration

MUSEUM D'HISTOIRE NATURELLE
1211 GENÈVE 6

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

PRIX DE L’ABONNEMENT:

SUISSE Fr. 225.— UNION POSTALE Fr. 230.—

(en francs suisses)

| Les demandes d’abonnement doivent étre adressées
| à la rédaction de la Revue suisse de Zoologie,
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ANNALES

de la

SOCIETE SUISSE DE ZOOLOGIE

et du

MUSEUM D’HISTOIRE NATURELLE
de la Ville de Geneve

tome 111
fascicule 3
2004

E à
kl GENEVE SEPTEMBRE 2004 ISSN 0035 - 418 X

SWISS JOURNAL OF ZOOLOGY

REVUE SUISSE DE ZOOLOGIE

TOME 111—FASCICULE 3

Publication subventionnée par:
ACADÉMIE SUISSE DES SCIENCES NATURELLES (SCNAT)
VILLE DE GENEVE
SOCIETE SUISSE DE ZOOLOGIE

VOLKER MAHNERT
Directeur du Museum d’histoire naturelle de Genève

CHARLES LIENHARD
Charge de recherche au Museum d’histoire naturelle de Genève

Comite de lecture

Il est constitué en outre du president de la Société suisse de Zoologie, du directeur du
Museum de Genève et de représentants des instituts de zoologie des universites
suisses.

Les manuscrits sont soumis à des experts d’institutions suisses ou étrangères selon le
sujet étudié.

La préférence sera donnée aux travaux concernant les domaines suivants: biogéo-
graphie, systématique, évolution, écologie, éthologie, morphologie et anatomie
comparée, physiologie.

Administration

MUSEUM D'HISTOIRE NATURELLE
1211 GENÈVE 6

Internet: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

PRIX DE L’ABONNEMENT:

SUISSE Fr. 225.— UNION POSTALE Fr. 230.—

(en francs suisses)

Les demandes d’abonnement doivent étre adressées
a la rédaction de la Revue suisse de Zoologie,
Museum d’histoire naturelle, C.P. 6434, CH-1211 Genève 6, Suisse

REVUE SUISSE DE ZOOLOGIE 111 (3): 457-523; septembre 2004

Hygronomini e Athetini della Cina con note sinonimiche
(Coleoptera, Staphylinidae)*

Roberto PACE
Via Vittorio Veneto, 13, I-37032 Monteforte d’Alpone (Verona), Italia.
E-mail: pace.ent@tiscali.it

Hygronomini and Athetini from China (Coleoptera, Staphylinidae)
with notes on the synonymy. - The tribe Athetini is represented by a large
number of species in all zoogeographical regions. The taxonomy of this
group is rather difficult and relies heavily on the characters of the male
copulatory organ and of female spermatheca. In the present paper ninety-
five species are recognised. One subgenus of Atheta, Ekkliatheta, and
65 species are described as new. Three new names are proposed: Alevonota
franziana for Tomoglossa franzi Pace, 1991 [nec Alevonota franzi (Palm,
1973)], Emmelostiba xinjiangensis for Alevonota chinensis Pace, 1993 (nec
Emmelostiba chinensis Pace, 1998), Atheta floresensis for Atheta aptera
Israelson, 1985 (nec Atheta aptera Pace, 1984). New synonymies are pro-
posed for nine species and genera: Alevonota pacei Assing, 2002, is placed
in synonymy with Arlantostiba franzi Pace, 1994, Cordalia permutata
Assing, 2002, with Cordalia chinensis Pace, 1993, the genus Euphorbagria
Assing, 1997, with Melagria Casey, 1906, Leptusa acuta Assing, 2002, with
Leptusa anmashanensis Pace, 1996, the genus Chariusa Pace, 2003, with
Leucocraspedum Kraatz, 1859, Liogluta granulipyga Pace, 1998, with
Liogluta langmusiensis Pace, 1998, and the genus Syntemusa Pace, 2002,
with Irmaria Cameron, 1925. The typical series of Atheta namphoensis
Pasnik, 2001, from North Korea includes two genera and two species, the
holotypus d must be attributed to the genus Nepalota with the name of
Nepalota namphoensis (Pasnik). New combinations are proposed for
10 species: 6 for the genus Alevonota, 1 for Emmelostiba, 2 for Irmaria and
1 for Nepalota. Each new species is described and illustrated. All available
distributional data are presented.

Keywords: Coleoptera - Staphylinidae - Aleocharinae - taxonomy - China -
synonymies - Palaearctic Region.

INTRODUZIONE

Con il presente lavoro prosegue la pubblicazione della descrizione delle nuove
Aleocharinae della Cina raccolte dal Dr. Ales Smetana di Ottawa ed altri, dopo quelle
precedenti (Pace, 1993a, 1997, 1998a-d, 1999a-d). Non dovrebbe essere sorprendente

* 186° Contributo alla conoscenza delle Aleocharinae.
Manoscritto accettato il 29.04.2004

458 R. PACE

la constatazione del numero elevato di specie nuove qui descritte, se si tiene presente
che i più alti rilievi montuosi della Cina sono solo in anni recentissimi oggetto di inda-
gini entomologiche sistematiche, particolarmente del Dr. Ale Smetana e d’altri
colleghi: Guillaume de Rougemont di Londra, Marc Tronquet di Molitg Les Bains
(Francia), Michael Schiilke di Berlino e dell’inglese J. Fellowes. Il presente lavoro com-
prende anche specie raccolte da questi colleghi e alcuni esemplari datimi in esame dal
Dr. J. Klimaszewski del Laurentian Forestry Centre di Sainte-Foy (Québec) Canada.

Gli olotipi delle nuove specie sono conservati nel Muséum d’histoire naturelle
di Ginevra (MHNG), nell’ Institut royal des Sciences naturelles de Belgique (IRSN) e
nel Museo di Storia Naturale dell’ Università Humboldt di Berlino (MB).

METODO

La tribù degli Athetini comprende per lo più specie di difficile classificazione,
soprattutto a livello sottogenerico nel genere Arheta Thomson, 1859. L'incertezza che
regna per le specie paleartiche occidentali di questa tribù, si ripete anche in Cina. Con
riferimento soprattutto al genere Atheta, la creazione di «gruppi misti» per le specie
paleartiche occidentali non è che la prova di quest’incertezza. Poiché per il presente
studio era necessario adottare un criterio di scelta ai fini dell’attribuzione sottogenerica
di Atheta e generica per altre specie, ho scelto come carattere prioritario la forma della
spermateca. Da qualche tempo si constata nel genere Atheta, anche da parte di altri
tassonomisti, che nell’ambito del sottogenere Microdota Mulsant & Rey, 1873, la
spermateca ha più o meno la forma della lettera S rovesciata, nell’ambito del sotto-
genere Atheta (Atheta) Thomson, 1859, la parte prossimale della spermateca è sempre
a forma di matassa di spire, da una a molte, nell’ambito del sottogenere Acrotona
Thomson, 1859, la parte prossimale della spermateca descrive una o più spire regolari,
nel sottogenere Philhygra Mulsant & Rey, 1873, la spermateca è assente o è estrema-
mente minuscola, nel sottogenere Datomicra Mulsant & Rey, 1874, essa ha la parte
distale deviata. Assai difficile è rilevare caratteri differenziali sottogenerici
nell’edeago. Nel presente lavoro le attribuzioni sottogeneriche e generiche vanno
considerate piuttosto sicure se è presente la femmina. I casi in cui è noto il solo mas-
chio l’attribuzione sottogenerica deriva da confronti con edeagi di specie note anche
per la femmina. L’armatura genitale interna dell’edeago, infatti, è composta di pezzi la
cui forma varia in lunghezza, larghezza o ispessimento chitinoso secondo le specie. Per
analogia di forma e posizione di questi pezzi è stata qui determinata l’attribuzione
sottogenerica nel caso di presenza di soli maschi. In ogni modo, evidentemente,
quest’attribuzione ha carattere provvisorio. Infine, le specie di Arheta che non presen-
tano una forma di spermateca caratteristica dei sottogeneri indicati sopra e di altri, sono
incluse nel sottogenere Dimetrota Mulsant & Rey, 1873, che nel presente lavoro
corrisponde più o meno ai gruppi misti delle specie paleartiche occidentali di Atheta.

RITROVAMENTO DEL ¢ O DELLA © DI SPECIE NOTE

Pelioptera viatica Pace, 1998
Rinvenuto il d, il cui edeago e il sesto urotergo libero sono illustrati alle
figg. 17-19.

HYGRONOMINI E ATHETINI DELLA CINA 459

Liogluta attenuata Pace, 1998
Rinvenuto il d, il cui edeago è illustrato alle figg. 42-44.

Atheta (Datomicra) parainopinata Pace, 1998
Rinvenuta la 9, la cui spermateca è illustrata in fig. 142.

Atheta (Datomicra) xinlongensis Pace 1998
Rinvenuta la vera 9, la cui spermateca è illustrata in fig. 127. La spermateca
illustrata precedentemente (Pace, 1998: 727) appartiene ad A. regressa Pace, 1998.

Paraloconota gansuensis Pace, 1998
Rinvenuto il d il cui edeago è illustrato alle figg. 188-189.

ELENCO DELLE SPECIE NOTE

Aloconota lanzhouensis Pace, 1998
Aloconota lanzhouensis Pace, 1998b: 424

2 es., China, Sichuan, Gongga Shan, Hailuogou, above Camp 3, 3000 m, 7.VII.1996, leg.
A. Smetana.

DISTRIBUZIONE. Gansu: Xinlong Shan.

Pelioptera (Tropimenelytron) viatica Pace, 1998
Pelioptera (Tropimenelytron) viatica Pace, 1998d: 938

9 es., China, Sichuan, Gongga Shan, Hailuogou, above Camp 3, 3000 m, 7.VII.1998, leg.
A. Smetana; 2 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m,
5.VII.1998, leg. A. Smetana; 1 2, China, Sichuan, 3000 m, 18.VII.1996, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

NOTA. Gusarov (2002), considera Tropimenelytron Pace, 1983, come genere a
se stante, utilizzando i caratteri diagnostici di Sawada. È da precisare che il metodo di
Sawada è fuorviante perché adotta caratteri a livello specifico utilizzandoli come
fossero caratteri generici. In questo modo dei gruppi di specie rischiano di essere
inclusi in generi differenti, rendendo la tassonomia delle Aleocharinae sempre più
incomprensibile. La tradizione vuole che sia la forma della ligula il criterio prioritario
per la separazione dei generi e non una o più setole spostate sui palpi.

Geostiba (Indatheta) rougemonti Pace, 1993

Geostiba (Indatheta) rougemonti Pace, 1993a: 90
1 2, China, N Yunnan, Xue Shan, 3900 m, 25.VI.1996, leg. A. Smetana.
DISTRIBUZIONE. Yunnan e Sichuan.

Hydrosmecta perignota Pace, 1998
Hydrosmecta perignota Pace, 1998b: 417

1 d, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke.
DISTRIBUZIONE. Beijing.

Liogluta lacustris Pace, 1998
Liogluta lacustris Pace, 1998b: 445

3 es., China, Sichuan, Gongga Shan, Hailuogou, lake above Camp 2, 2700 m,
4.VIIL.1998, leg. A. Smetana; 1 2, China, Sichuan, Gongga Shan, Hailuogou, for. above Camp

460 R. PACE

2, 2800 m, 5.VII.1998, leg. A. Smetana; 19 es., China, Sichuan, Gongga Shan, Hailuogou, above
Camp 3, 3000 m, 7.VII.1998, leg. A. Smetana; 4 es., China, N Yunnan, Xue Shan, nr. Zhongdian,
4000-4100 m, 23.VI.1996, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

Liogluta gonggana Pace, 1998
Liogluta gonggana Pace, 1998b: 445

19 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m,
5.V11.1998, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

Liogluta langmusiensis Pace, 1998
Liogluta langmusiensis Pace, 1998b: 450

1 4 e 1 9, China, Sichuan, Gongga Shan, Hailuogou, in front glacier 1, 2850 m,
7.V11.1996, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Langmusi.

Liogluta attenuata Pace, 1998
Liogluta attenuata Pace, 1998b: 450

1 d, China, N Yunnan, Xue Shan, nr. Zhongdian, 4000-4100 m, 23.VI.1996, leg.
A. Smetana.

DISTRIBUZIONE. Gansu: Xilong Shan.

Liogluta dalijiensis Pace, 1998
Liogluta dalijiensis Pace, 1998b: 448

2 es., China, N Yunnan, Xue Shan, nr. Zhongdian, 4000 m, 24-26.VI.1996, leg.
A. Smetana.

DISTRIBUZIONE. Gansu: Dalijia Shan.

Atheta (Acrotona) xishanensis Pace, 1998
Atheta (Acrotona) xishanensis Pace, 1998c: 689

1 9, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke.
DISTRIBUZIONE. Nota di Beijing, Henan, Jangsu e Yongnian.

Atheta (Acrotona) inquinata Cameron, 1939
Atheta (Acrotona) inquinata Cameron, 1939: 407

6 es., China, W Sichuan, 20 km N Sabdé, 3200 m, 11.VII.1998, leg. A. Smetana; 25 es.,
China, Sichuan, Gongga Shan, Hailuogou, in front glacier 1, 2850 m, 7.VII.1996, leg.
A. Smetana.

DISTRIBUZIONE. Specie del Kashmir, nuova per la Cina (Tipi esaminati).

Atheta (Microdota) philamicula Pace, 1998
Atheta (Microdota) philamicula Pace, 1998d: 918

3 es., China, W Sichuan, 20 km N Sabdé, 3200 m, 13.VII.1998, leg. A. Smetana.
DISTRIBUZIONE. Sichuan: Langmusi

HYGRONOMINI E ATHETINI DELLA CINA 461

Atheta (Microdota ) gonggaensis Pace, 1998
Atheta (Microdota ) gonggaensis Pace, 1998d: 916

13 es., China, Sichuan, Gongga Shan, Hailuogou, above Camp 3, 3000 m, 6.VII.1998,
leg. A. Smetana; 1 à, China, SE Sichuan, Jinfo Shan, 27.VI.1996, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

Atheta (Microdota) plactra Cameron, 1939
Atheta (Traumoecia) placita, Cameron, 1939: 319

9 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5.VII.1998,
leg. A. Smetana; 5 es., China, Sichuan, Gongga Shan, Hailuogou, above Camp 3, 3200 m,

7.V11.1998, leg. A. Smetana; 30 es., China, W Sichuan, 20 km N Sabdé, 3200 m, 13.VII.1998,
leg. A. Smetana.

DISTRIBUZIONE. Specie finora nota solo dell’India: Ghum, Tiger Hill. (Tipi
esaminati).

Atheta (Dimetrota) sericoides Pace, 1998

Atheta (Dimetrota) sericoides Pace, 1998c: 695
1de2 22, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke.
DISTRIBUZIONE. Nota finora solo di Beijing (Pechino).

Atheta (Dimetrota) discriminata Pace, 1987

Atheta (Dimetrota) discriminata Pace, 1987: 420
1 2, China, Sichuan, Emei Shan, Leidongping, 2500 m, 18.VII.1998, leg. A. Smetana.
DISTRIBUZIONE. Specie finora nota dell’ India, Kotgarth e Chakrata.

Atheta (Datomicra) peranomala Pace, 1998
Atheta (Datomicra) peranomala Pace, 1998d: 927
7 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5.V11.1998,

leg. A. Smetana; 10 es., China, W Sichuan, 20 km N Sabdé, 3300 m, 14.VII.1998, leg. A.
Smetana.

DISTRIBUZIONE. Yunnan: Ruili e Mengdien.

Atheta (Datomicra) xinlongensis Pace, 1998
Atheta (Datomicra) xinlongensis Pace, 1998c:725

1 d, China, Gansu, Xinlong Shan, cca. 70 km S Lanzhou, 2380 m, 7.VIII.1994, leg.
A. Smetana; 1 d, China, N Yunnan, Xue Shan, nr. Zhongdian, 3900 m, 25.VI.1996, leg.
A. Smetana; 1 d, China, Sichuan, Kangding, 2900 m, 2.VII.1996, leg. A. Smetana; 4 es., China,

Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5.VII.1998, leg. A. Smetana;
17 es., China, W Sichuan, 20 km N Sabdé, 3300 m, 14.VII.1998, leg. A. Smetana.

DISTRIBUZIONE. Gansu: Xinlong Shan.

Atheta (Datomicra) regressa Pace, 1998

Atheta (Datomicra) regressa Pace, 1998c:728
1 2, China, N Yunnan, Xue Shan, Zhongdian, 19-25.VI.1994, leg. D. Kral & J. Farkaé.
DISTRIBUZIONE. Gansu: Xinlong Shan e Sichuan: Gongga Shan.

462 R. PACE

Atheta (Datomicra) parainopinata Pace, 1998
Atheta (Datomicra) parainopinata Pace, 1998c: 725

3 es., China, Sichuan, Gongga Shan, Hailuogou, Lake above camp 2, 2750 m,
4.V11.1998, leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

Atheta (Datomicra) subinopinata Pace, 1998
Atheta (Datomicra) subinopinata Pace, 1998c: 722

20 es., China, W Sichuan, Kangding, 2800-2900 m, 2.VII.1996 e 21.VII.1998, leg.
A. Smetana; 3 es., China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
6.VII.1996, leg A. Smetana; 34 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp

2, 2800 m, 5.VII.1998, leg. A. Smetana; 70 es., China, W Sichuan, 20 Km N Sabdé, 3200 m,
13.VII.1998, leg A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan.

Atheta (Datomicra) graffa Pace, 1998
Atheta (Datomicra) graffa Pace, 1998c: 728

8 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5.VII.1998,
leg. A. Smetana.

DISTRIBUZIONE. Sichuan: Gongga Shan e Gansu: Xinlong Shan.

Atheta (Datomicra) dadopora Thomson, 1867

Atheta dadopora Thomson, 1867: 282

Atheta (Datomicra) dadopora: Lohse 1974: 189; Pace, 1998c: 730
Atheta (Datostiba) poroshirica Sawada, 1977: 243

3 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m, 5.VII.1998,
leg. A. Smetana.

DISTRIBUZIONE. Specie diffusa dall’Europa alla Siberia, all'India settentrionale
e al Giappone. Già nota della Cina (Pace, 1998c).

Atheta (Datomicra) subsericans Cameron, 1939
Atheta (s. str.) subsericans Cameron, 1939: 355

4 es., China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m, 6.VII.1996, leg
A. Smetana; 6 es., China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m,
5.V11.1998, leg. A. Smetana; 1 9, China, N Yunnan, Xue Shan, nr. Zhongdian, 4000-4100 m,
23-24.VI.1996, leg. A. Smetana.

DISTRIBUZIONE. Specie dell’India settentrionale. Già nota per la Cina (Pace,
1998).

Atheta (Oreostiba) shanicola Pace, 1998, figg. 162-163
Atheta (Oreostiba) shanicola Pace, 1998d: 933

1 2, China, N Yunnan, Xue Shan nr. Zhongdian, 4050 m, 24.VI.1996, leg. A. Smetana.
DISTRIBUZIONE. Finora nota solo del Sichuan.

NOTA. Questo esemplare differisce lievemente da quello della serie tipica. In un
primo tempo l’avevo considerato appartenente a nuova specie e non una varietà locale.
Qui si danno le illustrazioni per rendere note le differenze, a mio parere non di livello
specifico.

HYGRONOMINI E ATHETINI DELLA CINA 463

Nepalota pernitida (Pace, 1984), comb. n.

Atheta (Stethusa) pernitida Pace, 1984a: 447
1 9, China, W Sichuan, 20 Km N Sabdé, 3200 m, 15.VII.1998, leg A. Smetana.
DISTRIBUZIONE. Burma: Kalaw.

Nepalota gansuensis Pace, 1998
Nepalota gansuensis Pace, 1998d: 943

32 es., China, Sichuan, Gongga Shan, Hailuogou, in front glacier 1, 2850 m, 7.VII.1996,
leg. A. Smetana; 8 es., China, SE Sichuan, Jinfo Shan, 1750 m, 26.VI.1996, leg. A. Smetana; 9
es., China, Sichuan, Emei Shan, Leidongping, 2500 m, 18.VII.1998, leg. A. Smetana.

DISTRIBUZIONE. Gansu: Xinlong Shan.

Nepalota martensi Pace, 1987

Nepalota martensi Pace, 1987: 412
1 g, China, Yunnan, Kunming, Western Hills, 1800 m, 24.VII.1996, leg. A. Smetana.
DISTRIBUZIONE. Nepal.

Paraloconota gansuensis Pace, 1998
Paraloconota gansuensis Pace, 1998b: 436

17 es., China, Sichuan, Gongga Shan, Hailuogou, head of glacier, 1250 m, 9.VII.1996,
leg. A. Smetana.

DISTRIBUZIONE. Gansu: Dalijia Shan.

Paraloconota yonghaiensis Pace, 1998
Paraloconota yonghaiensis Pace, 1998b: 438

1 2, China, Gansu, Yonghai, cca. 20 km SW Yuzhong, 2700-2800 m, 9.VIII.1994, leg.
A Smetana.

DISTRIBUZIONE. Giä nota del Gansu.

NOTA. Questo esemplare differisce lievemente da quello della serie tipica. In un
primo tempo l’avevo considerato appartenente a nuova specie e non una varietä locale.
Qui si danno le illustrazioni per rendere note le differenze, a mio parere non di livello
specifico.

ELENCO DELLE NUOVE SPECIE

HYGRONOMINI 9. Liogluta sabdensis sp. n.
1. Hygronoma chinensis sp. n. 10. Liogluta infacunda sp. n.
ATHETINI 11. Liogluta kangdingensis sp. n.
2. Gnypeta yingkouensis sp. n. 12. Liogluta caliginis sp. n.
3. Aloconota pulchricarinata sp. n. 13. Liogluta rhomboidalis sp. n.
4. Aloconota umbonis sp. n. 14. Liogluta serpentitheca sp. n.
5.Aloconota jinfoensis sp. n. 15. Liogluta biacusifera sp. n.
6. Pelioptera tronqueti sp. n. 16. Liogluta imitatrix sp. n.
7.Geostibasoma opacisuturale sp. n. 17. Liogluta ignorata sp. n.

8. Amischa alticola sp. n. 18. Liogluta iperintroflexa sp. n.

464

19. Atheta (Philhygra) quadrifalcifera
sp. n.

20. Atheta (Philhygra) perconsanguinea

sp. n.

21. Atheta (Philhygra) longefalcifera sp. n.

22. Atheta (Philhygra) emeicola sp. n.
23. Atheta (Sipalatheta) emeimontis sp. n.
24. Atheta (Acrotona) fellowesi sp. n.
25. Atheta (Acrotona) deuvei sp. n.
26. Atheta (Acrotona) luteihumeri sp. n.
27. Atheta (Microdota) hailuogouensis
sp. n.
28. Atheta (Microdota) miriapex sp. n.
29. Atheta (Microdota) sichuanicola sp. n.
30. Atheta (Microdota) masculifrons sp. n.
31. Atheta (Microdota) gibba sp. n.
32. Atheta (Microdota) neoamicula sp. n.
33. Atheta (Microdota) tardoides sp. n.
34. Atheta (Microdota) zhongdianensis
sp. n.
35. Atheta (Microdota) liaoningensis sp. n.
36. Atheta (Dimetrota) angulispina sp. n.
37. Atheta (Dimetrota) ridenda sp. n.
38. Atheta (Dimetrota) jiudingensis sp. n.
39. Atheta (Datomicra) zhagaensis sp. n.
40. Atheta (Datomicra) consequens sp. n.

41. Atheta (Datomicra) songpanensis sp. n.

DESCRIZIONI

Hygronoma chinensis sp. n.

R. PACE

42. Atheta (Datomicra) falcifera sp. n.

43. Atheta (Datomicra) pertinens sp. n.

44. Atheta (Datomicra) stenotheca sp. n.

45. Atheta (Datomicra) latifalcifera sp. n.

46. Atheta (Datomicra) mesofalcifera
sp. n.

47. Atheta (Datomicra) hamifera sp. n.

48. Atheta (Oreostiba) micromega sp. n.

49. Atheta (Oreostiba) transcripta sp. n.

50. Atheta (Oreostiba) mimomonticola

sp. n.

51. Atheta (Atheta) paralaevicauda sp. n.

52. Atheta (Ekkliatheta) aniiensis sp. n.

53. Nepalota fellowesi sp. n.

54. Nepalota guangdongensis sp. n.

55. Paraloconota puella sp. n.

56. Paraloconota dalijiamontis sp. n.

57. Oroekklina smetanai sp. n.

58. Emmelostiba granulosa sp. n.

59. Emmelostiba bellicosa sp. n.

60. Alevonota pulchricornis sp. n.

61. Alevonota sinensis sp. n.

62. Alevonota foedicornis sp. n.

63. Taxicera sinensis sp. n.

64. Amidobia perconvexa sp. n.

65. Amidobia ming sp. n.

Figg. 1-2

Holotypus 2, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke (IRSN).
DESCRIZIONE. Lungh. 2,7 mm. Corpo debolmente lucido e bruno scuro, con
addome nero; antenne bruno-rossicce con i due antennomeri basali bruni; zampe
rossicce. L’intero corpo è coperto di granulosità fine e fitta, posta su un fondo non

reticolato. Spermateca fig. 2.

COMPARAZIONI. La nuova specie è ben differente dell’europea H. dimidiata
(Gravenhorst, 1806), per il pronoto nettamente trasverso (lungo quanto largo in dimidi-
ata), per avere solo il primo urotergo libero solcato alla base e il primo metatarsomero

molto più lungo dei due seguenti riuniti.

Gnypeta yingkouensis sp. n.

Figg. 3-4

Holotypus 9, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke (IRSN).

DESCRIZIONE. Lungh. 2,2 mm. Corpo debolmente lucido e bruno; antenne
bruno-rossicce; zampe rossicce. Tutto il corpo è coperto di granulosità saliente e fittis-
sima. L’addome presenta una reticolazione evidente. Spermateca fig. 4.

HYGRONOMINI E ATHETINI DELLA CINA 465

COMPARAZIONI. Le specie cinesi del genere Gnypeta Thomson, 1858, finora
sono G. beijingensis Pace, 1998, G. chinensis Pace, 1998, G. immodesta Pace, 1998,
G. lucidula Pace, 1998 e G. yunnanensis Pace, 1998. Per la forma della spermateca la
nuova specie è simile a G. beijingensis, tuttavia 1 penultimi antennomeri sono trasversi
nella nuova specie e lunghi quanto larghi in beijingensis; il pronoto è molto trasverso
nella nuova specie e lo è poco in beijingensis e la parte intermedia della spermateca è
molto più lunga nella nuova specie che in G. beijingensis.

Aloconota pulchricarinata sp. n. Figg. 5-7

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
6. VII.1996, 29°35 N-102°00E, leg A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 3,8 mm. Corpo debolmente lucido e nero; antenne nero-
brune; zampe giallo-brune con 1 tarsi rossicci. La punteggiatura del capo è estrema-
mente superficiale e assente sulla fascia mediana. La granulosità del pronoto è poco
distinta, quella delle elitre è saliente sulla metà interna, verso la sutura, evanescente
sulla metà esterna. La reticolazione del capo e dell’addome è netta, quella del pronoto
è nettissima, a maglie molto trasverse sull’addome. Il capo ha un’impressione discale.
Edeago figg. 6-7.

COMPARAZIONI. Per la forma dell’edeago e per i caratteri sessuali secondari sul
quinto urotergo libero del d, la nuova specie si mostra affine ad A. inaequalis
Cameron, 1939, dell’India settentrionale (holotypus da me esaminato). Tuttavia, il
quinto urotergo libero del & della nuova specie mostra un rilievo mediano posteriore
a Y, mentre in inaequalis lo stesso è a I. L’armatura genitale interna dell’edeago della
nuova specie è robusta, mentre quella di inaequalis è molto evanescente.

Aloconota umbonis sp. n. Figg. 8-10

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, lake above Camp 2, 2750 m,
4.VII.1998, 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 3,9 mm. Corpo debolmente lucido e nero; antenne nere;
zampe giallo-rossicce. La granulosità del capo è distinta, ma assente sulla fascia longi-
tudinale mediana, quella del pronoto è saliente e quella delle elitre è evanescente. La
reticolazione del disco del capo è netta e distinta sul resto del capo, quella del pronoto
e delle elitre è nettissima, quella dell'addome è molto trasversa ed evanescente. Il capo
presenta una debole impressione discale. Il quinto urotergo libero del 4 presenta un
rilievo mediano posteriore molto saliente. Edeago figg. 9-10.

COMPARAZIONI. La nuova specie presenta il quarto antennomero più lungo che
largo e il quinto nettamente trasverso. Per questi caratteri e per altri è ben distinta dalle
restanti specie cinesi note per la sola 9. Le specie note anche per il d non mostrano i
caratteri sessuali secondari degli uroterghi liberi quinto e sesto del d della nuova
specie.

Aloconota jinfoensis sp. n. Figg. 11-13
Holotypus 3, China, SE Sichuan, Jinfo Shan, 1800 m, 29°01N-107°1414E, 27.VI.1998,
1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).
DESCRIZIONE. Lungh. 5,1 mm. Corpo lucido e bruno, con omeri, margine poste-
riore dei due uroterghi liberi basali e pigidio giallo-rossicci; antenne brune con

466 R. PACE

Fico. 1-7

Habitus, spermateca ed edeago in visione laterale e ventrale. 1-2: Hygronoma chinensis sp. n.;
3-4: Gnypeta yingkouensis sp. n.; 5-7: Aloconota pulchricarinata sp. n.

l’antennomero basale bruno-rossiccio; zampe giallo-rossicce. La punteggiatura del
capo è svanita e assente sulla fascia longitudinale mediana. La granulosità del pronoto
è distinta, quella delle elitre è superficiale. La reticolazione del capo è evanescente,
quella del pronoto è molto svanita, quella delle elitre è estremamente superficiale e

HYGRONOMINI E ATHETINI DELLA CINA 467

Fico. 8-13

Habitus ed edeago in visione laterale e ventrale. 8-10: Aloconota umbonis sp. n.; 11-13:
Aloconota jinfoensis sp. n.

quella dell’addome manca. Il disco del capo è impresso. Il primo urotergo libero basale
del d presenta una protuberanza molto saliente e incisa all’indietro, il secondo
urotergo libero ha un tubercolo saliente e il quinto urotergo libero del 4 presenta un
rilievo mediano largo con parte superiore lievemente concava e gradualmente più
saliente all’indietro. Edeago figg. 11-12.

468 R. PACE

COMPARAZIONI. A motivo dei caratteri sessuali secondari del d, la nuova specie
ricorda da vicino A. spectabilis Pace, 1987, del Nepal. La nuova specie se ne distingue
per avere un rilievo a superficie superiore lievemente concava sul quinto urotergo
libero del & , invece di un granulo e per l’armatura genitale interna dell’edeago più
robusta e di forma ben differente da quella di spectabilis.

Pelioptera (Pelioptera) tronqueti sp. n. Figg. 14-16

Holotypus d, Chine, Guanxi, Mass. de Miao’er-Shan, 2000 m, 16.VIII.1994, litière, leg.
M. Tronquet (IRSN).

DESCRIZIONE. Lungh. 2,2 mm. Corpo lucidissimo e bruno, con uroterghi liberi
quarto e quinto neri; antenne nero-pece con antennomero basale giallo-rossiccio;
zampe giallo-rossicce. La granulosità del capo è molto svanita e fine, quella del pro-
noto è distinta e non fitta e quella delle elitre è molto superficiale. La reticolazione del
capo è evanescente, quella del pronoto e dell’addome è distinta, quella delle elitre è
molto svanita e a maglie molto trasverse sull’addome. Edeago figg. 15-16.

COMPARAZIONI. La nuova specie, per la forma dell’edeago, appartiene al grup-
po di specie che fanno capo a P. opaca Kraatz, 1857, dello Sri Lanka, a cui apparten-
gono P. opacoides Pace, 1991 del Nepal e P. zerchei Pace, 1993, di Sumatra. La nuo-
va specie si distingue da tutte queste specie per avere l’edeago molto ristretto all’apice,
in visione ventrale, e un’armatura genitale interna dell’edeago differente.

DERIVATIO NOMINIS. La nuova specie è dedicata al suo raccoglitore, il collega
Marc Tronquet di Molitg Les Bains (Francia), noto studioso di Staphylinidae.

Geostibasoma opacisuturale sp. n. Figg. 20-24

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 13.VII.1998, leg. 1998 China
Expedition J. Farkaè, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypi: 2 dd e 5 9 9, stessa provenienza.

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido con fascia suturale opaca e nero,
antenne comprese; zampe giallo-rossicce. La punteggiatura del capo è svanita, assente
sulla fascia longitudinale mediana. La granulosità del pronoto e delle elitre è distinta e
fine. La reticolazione del capo e del pronoto è distinta, quella delle elitre è netta e
vigorosa su una fascia lungo la sutura e quella dell'addome è a maglie trasverse dis-
tinte. Il capo presenta due carene mediane occipitali. Il pronoto ha un appiattimento
mediano. Edeago figg. 21-22, spermateca fig. 23, sesto urotergo libero del d fig. 24.

COMPARAZIONI. La nuova specie per la forma dell’edeago è sicuramente affine
a G. satyrus Pace, 1998, di Hong Kong. Se ne distingue per gli occhi più corti delle
tempie (più lunghi delle tempie in satyrus) e per la diversa struttura generale
dell’edeago.

Amischa alticola sp. n. Figg. 25-26
Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
6.VII.1996, 29°35N-102°00E, leg A. Smetana (MHNG).
DESCRIZIONE. Lungh. 2,4 mm. Corpo lucido e bruno con uroterghi liberi quarto
e quinto nero-bruni; antenne brune con i tre antennomeri basali di un giallo sporco;
zampe gialle. La granulosità del capo e del pronoto è distinta, quella delle elitre è

HYGRONOMINI E ATHETINI DELLA CINA 469

0,1 mm

À
\

I mm

À Vies
FEN

amie
Loi

IHM N

q *

)

FIGG. 14-19

Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del 4. 14-16: Pelioptera
tronqueti sp. n.; 17-19: Pelioptera viatica Pace, 1998.

superficiale. La reticolazione del capo, del pronoto e delle elitre è superficiale, quella
dell'addome è distinta. Il pronoto presenta un largo solco mediano posteriore.
Spermateca fig. 26.

COMPARAZIONI. La nuova specie presenta il pronoto impresso e la parte prossi-
male della spermateca non avvolta in spire e con bulbo prossimale largo quanto il dis-
tale. Per questi caratteri ed altri è nettamente distinta da A. beijingensis Pace, 1998, A.
rougemonti Pace, 1998 e A. nana Pace, 1998, tutte della Cina.

470 R. PACE

Fico. 20-26

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del 4. 20-24:
Geostibasoma opacisuturale sp. n.; 25-26: Amischa alticola sp. n.

Liogluta sabdensis sp. n. Figg. 27-30

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
13.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypus: 1 ©, China, W Sichuan, 20 km N Sabdé, 3200 m, 15.VII.1998, leg. A.
Smetana.

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e nero, con elitre nero-brune;
antenne nere, zampe rossicce. La punteggiatura del capo e del pronoto è distinta, ma
assente sulla fascia longitudinale mediana del capo, quella delle elitre è fine e molto

HYGRONOMINI E ATHETINI DELLA CINA 471

evanescente. La reticolazione del capo, delle elitre e dell’addome è distinta, quella del
pronoto è netta, sull’addome è a maglie molto trasverse. Edeago figg. 28-29, sper-
mateca fig. 30.

COMPARAZIONI. Per la forma dell’edeago la nuova specie è sicuramente affine a
L. lacustris Pace, 1998, pure della Cina. Se ne distingue per l’edeago meno ampia-
mente arcuato nella regione preapicale, in visione ventrale, per la diversa forma
dell’armatura genitale interna dell’edeago e per l’undicesimo antennomero lungo
quanto i due precedenti riuniti (lungo quanto i tre precedenti riuniti in lacustris).

Liogluta infacunda sp. n. Figg. 31-34

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3300 m, 29°35N-102°23E,
14.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e nero, antenne comprese; zampe
di un giallo sporco, con femori giallo-bruni. La punteggiatura del capo e del pronoto è
svanita e assente per breve tratto sulla fascia longitudinale mediana del capo. La granu-
losità delle elitre è molto evanescente. La reticolazione del capo e dell’addome è
svanita, quella del pronoto è netta e quella delle elitre è distinta, a maglie trasverse
sull’addome. Sesto urotergo libero del 6 fig. 32, edeago figg. 33-34.

COMPARAZIONI. L’edeago della nuova specie è simile a quello di L. langmusien-
sis Pace, 1998 (= L. granulipyga Pace, 1998, syn. n.), pure della Cina. Tuttavia l’apice
dell’edeago della nuova specie è nettamente più stretto di quello di langmusiensis, la
«crista apicalis» dell’edeago è molto più sviluppata nella nuova specie e l’armatura
genitale interna dello stesso edeago è del tutto differente.

Liogluta kangdingensis sp. n. Figg. 35-37

Holotypus d, China, W Sichuan, 15 km W Kanding, rte. 138, 3250 m, 19.VII.1998, leg.
1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 4,0 mm. Corpo lucido e nero, antenne comprese; zampe
brune con tarsi rossicci. La punteggiatura del capo è superficiale. La granulosità del
pronoto è fine e poco evidente, quella delle elitre è distinta. La reticolazione dell’avan-
corpo è superficiale, quella degli uroterghi liberi quarto e quinto è molto trasversa e
molto evanescente, assente sui restanti uroterghi liberi. Edeago figg. 36-37.

COMPARAZIONI. L’edeago della nuova specie ha forma affine a quella dell’ede-
ago di L. gonggana Pace, 1998, tuttavia è più bruscamente flesso al lato ventrale e
l’armatura genitale interna dell’edeago stesso è nettamente più sviluppata e forte di
quella di gonggana. Inoltre la nuova specie presenta occhi assai più corti delle tempie
e non più lunghi, come in gonggana.

Liogluta caliginis sp. n. Figg. 38-41

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
13.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypus: 1 9, stessa provenienza.

DESCRIZIONE. Lungh. 2,8 mm. Corpo lucido e nero, antenne comprese; zampe
di un giallo sporco. La punteggiatura del capo è distinta e assente sulla fascia longitu-
dinale mediana, quella delle elitre è superficiale. La granulosità del pronoto è distinta.

472 R. PACE

1 mm

Ficc. 27-32
Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del d. 27-30:
Liogluta sabdensis sp. n.; 31-32: Liogluta infacunda sp. n.

La reticolazione di tutto il corpo è evidente, a maglie molto trasverse solo sull’addome.
Il disco del capo è appiattito. Edeago figg. 39-40, spermateca fig. 41.

COMPARAZIONI. La parte preapicale dell’edeago della nuova specie è fortemente
ristretto, in visione ventrale. Per questo carattere è possibile che la nuova specie sia
tassonomicamente affine, perché pure esse lo presentano, a L. sinensis Pace, 1998, L.

HYGRONOMINI E ATHETINI DELLA CINA 473

I mm

Figc. 33-38

Edeago in visione laterale e ventrale e habitus. 33-34: Liogluta infacunda sp. n.; 35-37: Liogluta
kangdingensis sp. n.; 38: Liogluta caliginis sp. n.

xiahensis Pace, 1998 e L. lacustris Pace, 1998, tutte della Cina. La parte ristretta
dell’edeago della nuova specie, tuttavia, è più vicina alla «crista apicalis» che
all’estremità apicale dello stesso edeago. L’armatura genitale interna dell’edeago,
inoltre, è del tutto differente da quella delle tre specie a confronto.

474 R. PACE

Fico. 39-44

Edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del d. 39-41: Liogluta
caliginis sp. n.; 42-44: Liogluta attenuata Pace, 1998.

Liogluta rhomboidalis sp. n. Figg. 45-48

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 7.VII.1996, leg A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypi: 3 dd, stessa provenienza, ma anche 2800 m e 9.VII.1996; 3 d d, China, N
Yunnan, Xue Shan nr. Zhongdian, 4000 m, 24-28.VI.1996, leg. A. Smetana; 2 es., China, N

HYGRONOMINI E ATHETINI DELLA CINA 475

FIGG. 45-50

Habitus, edeago in visione laterale e ventrale e spermateca. 45-48: Liogluta rhomboidalis sp. n.;
49-50: Liogluta serpentitheca sp. n.

Yunnan, Xue Shan nr. Zhongdian, 4050 m, 24.VI.1996, leg. A. Smetana; 1 es., China, N Yunnan,
Xue Shan nr. Zhongdian, 4000-4100 m, 23.VI.1996, leg. A. Smetana; 1 es., China, N Yunnan,
Xue Shan nr. Zhongdian, 3800 m, 26.VI.1996, leg. A. Smetana; 1 9, China, W Sichuan, 20’km
N Sabdé, 3200 m, 13.VII.1998, leg. A. Smetana; 1 2, China, W Sichuan, 20 km N Sabdé, 3200
m, 11.VII.1998, leg. A. Smetana.

476 R. PACE

DESCRIZIONE. Lungh. 4,5 mm. Corpo lucido e nero, con elitre nero-brune;
antenne nere; zampe rossicce. La punteggiatura del capo è distinta e poco profonda,
assente sulla fascia longitudinale mediana, quella del pronoto è fine e superficiale.
Assenti sono punteggiatura e granulosità sulle elitre. La reticolazione è distinta su tutto
il corpo, sull’addome a maglie molto trasverse. Edeago figg. 46-47, spermateca fig. 48.

COMPARAZIONI. Per la forma dell’edeago e della spermateca, la nuova specie è
sicuramente affine a L. lacustris Pace, 1998, pure della Cina, da cui differisce per la
parte preapicale ventrale dell’edeago a forma romboidale (forma semiellittica in
lacustris) e per la presenza di una forte spina dell’armatura interna dell’edeago, assente
in lacustris.

Liogluta serpentitheca sp. n. Figg. 49-50

Holotypus 9 , China, Sichuan, Kangding (formely Tatsienlu), 2900 m, 30°03N-102°02E,
2.VII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e nero, con elitre nero-brune;
antenne nere; zampe rossicce con femori bruno-rossicci e tarsi giallo-rossicci. La pun-
teggiatura del capo è superficiale e assente sulla fascia longitudinale mediana, quella
del pronoto è distinta e quella delle elitre è fine e poco distinta. La reticolazione del
capo è netta sul disco e distinta sul resto della sua superficie, quella delle elitre e
dell’addome è netta, a maglie molto trasverse sull’addome.

COMPARAZIONI. Per la forma della spermateca, la nuova specie è sicuramente
affine a L. langmusiensis Pace, 1998, pure della Cina. Se ne distingue per avere il bulbo
distale della spermateca largo quasi il doppio di quello prossimale (poco più largo in
langmusiensis) e per la parte intermedia della stessa spermateca che descrive un arco
molto più ampio di quello di /angmusiensis. Il quarto antennomero della nuova specie,
inoltre, è trasverso, più lungo che largo in Jangmusiensis.

Liogluta biacusifera sp. n. Figg. 51-54

Holotypus d, China, N Yunnan, Xue Shan nr. Zhongdian, 4000 m, 27°49N-99°34E, 24-
26.VI.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypus: 1 2, China, N Yunnan, Xue Shan nr. Zhongdian, 4050 m, 24.VI.1996, leg. A.
Smetana

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e nero, con elitre nero-brune; an-
tenne nere; zampe giallo-brune. La punteggiatura del capo è distinta e assente sulla fas-
cia longitudinale mediana, quella del pronoto è distinta. La punteggiatura o la granu-
losità delle elitre sono molto superficiali. La reticolazione del capo è distinta sul disco
e molto superficiale sul resto della sua superficie, quella del pronoto è evanescente e
quella delle elitre e dell'addome è distinta, a maglie molto trasverse sull’addome.
Edeago figg. 52-53, sesto urotergo libero del d fig. 54.

COMPARAZIONI. Per la forma dell’edeago la nuova specie si mostra affine sia a
L. langmusiensis Pace, 1998, sia a L. gonggana Pace, 1998, entrambe pure della Cina.
Se ne distingue per la presenza di una sporgenza angolare a ciascun lato della regione
intermedia e per l’armatura genitale interna dell’edeago molto più forte e più comp-
lessa di quella delle due specie a confronto.

Liogluta imitatrix sp. n. Figg. 55-56

Holotypus 2, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
13.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

HYGRONOMINI E ATHETINI DELLA CINA 477

FAR)

KE;

pae ua
Be à
D AA a
A
a 8:

Figc. 51-56

Habitus, edeago in visione laterale e ventrale, spermateca e sesto urotergo libero del d. 51-54:
Liogluta biacusifera sp. n.; 55-56: Liogluta imitatrix sp. n.

DESCRIZIONE. Lungh. 4,1 mm. Corpo lucido e nero-bruno; antenne nere; zampe
rossicce. La punteggiatura del capo è fine e superficiale e assente per stretto tratto sulla
fascia longitudinale mediana, quella delle elitre è poco distinta. La granulosità del
pronoto è fine e svanita. La reticolazione del capo è distinta, quella del pronoto e delle

elitre è netta e quella dell’addome è svanita, a maglie molto trasverse. Spermateca
fig. 56.

478 R. PACE

COMPARAZIONI. La spermateca della nuova specie ricorda da vicino quella di /.
lacustris Pace, 1998, pure della Cina. Se ne distingue per avere la parte prossimale net-
tamente più lunga e il bulbo distale quasi sferico (piriforme in lacustris). Gli occhi del-
la nuova specie, inoltre, sono più corti delle tempie, mentre in /acustris sono lunghi
quanto le tempie.

Liogluta ignorata sp. n. Figg. 57-58

Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3200 m,
29°35N-102°00E, 7.VII.1996, leg A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e nero, con elitre nero-brune;
antenne nere con i due antennomeri basali nero-bruni; zampe rossicce. La granulosità
del capo è fine e distinta, assente sulla fascia longitudinale mediana, quella del pronoto
è fine e saliente e quella delle elitre è poco distinta. La reticolazione del capo è nettis-
sima sul disco e superficiale sul resto della sua superficie, quella del pronoto e delle
elitre è netta e quella dell’addome è distinta, a maglie molto trasverse. Il pronoto
presenta un debole solco mediano posteriore. Spermateca fig. 58.

COMPARAZIONI. La spermateca della nuova specie è simile a quella di L. attenu-
ata Pace, 1998, pure della Cina. Se ne distingue per la presenza dell’introflessione
apicale del bulbo distale della stessa spermateca poco profonda (profondissima in
attenuata) e per la parte prossimale dello stesso organo molto lunga e sviluppata (gra-
dualmente attenuata in attenuata). La nuova specie, inoltre, presenta il quarto
antennomero trasverso e non più lungo che largo, come in attenuata.

Liogluta iperintroflexa sp. n. Figg. 59-60

Holotypus 2, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
15.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e nero, con le elitre nero-brune;
antenne nere; zampe rossicce con femori bruno-rossicci. La punteggiatura del capo è
ombelicata, superficiale e assente sulla fascia longitudinale mediana, quella del pro-
noto è distinta e quella delle elitre è poco distinta. La reticolazione del capo è svanita,
quella del pronoto e dell'addome è distinta, a maglie molto trasverse sull’addome, e
quella delle elitre è netta. Spermateca fig. 60.

COMPARAZIONI. Per la profonda introflessione apicale del bulbo distale della
spermateca, la nuova specie si presenta affine a L. xiaheorum Pace, 1998, pure della
Cina. Il bulbo distale della spermateca della nuova specie, tuttavia, è largo tre volte il
bulbo prossimale della stessa spermateca e non due volte e mezzo come in xiaheorum
e la parte prossimale della stessa spermateca della nuova specie è strettamente flessa,
mentre è ampiamente arcuata in xiaheorum.

Atheta (Philhygra) quadrifalcifera sp. n. Figg. 61-65

Holotypus & , China, Sichuan, Gongga Shan, Hailuogou, lake above Camp 2, 2750 m,
29°35N-102°00E, 5-9.VIII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypi: 2 G 4, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3200 m,
7.VI.1996, leg A. Smetana.

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e nero-bruno, con quarto urotergo

libero nero e iridescente; antenne nere con i tre antennomeri basali rossicci; zampe

HYGRONOMINI E ATHETINI DELLA CINA 479

113 SIN
Aal“ II TE
Fi dpt APE
5e rye
= MAL ANS

f
Al

I mm

= 61 62
Figc. 57-63

Habitus, spermateca ed edeago in visione laterale e ventrale. 57-58: Liogluta ignorata sp. n.;
59-60: Liogluta iperintroflexa sp. n.; 61-63: Atheta (Philhygra) quadrifalcifera sp. n.

giallo-rossicce. La punteggiatura del capo è distinta solo ai lati. La granulosità del
pronoto è distinta e quella delle elitre è superficiale. La reticolazione del capo è nettis-
sima sul disco e svanita ai lati, quella del pronoto è pure nettissima, quella delle elitre
è netta e quella dell’addome è distinta, a maglie molto trasverse sull’addome. Il

480 R. PACE

pronoto presenta un debole solco mediano posteriore. Edeago figg. 62-63, sesto uro-
tergo libero del d fig. 64.

COMPARAZIONI. L’edeago della nuova specie è simile a quello di A. dalijiensis
Pace, 1998, ma il suo apice è lineare nella nuova specie e fortemente ricurvo in dali-
jiensis. L’armatura genitale interna dell’edeago della nuova specie, inoltre, presenta
due forti spine ricurve, assenti nell’edeago di dalijiensis. Il pronoto della nuova specie
è più trasverso di quello della specie a confronto.

Atheta (Philhygra) perconsanguinea sp. n. Figg. 65-68

Holotypus d, China, N Yunnan, Xue Shan, nr. Zhongdian, 3900 m, 27°49N-99°34E,
25.V1.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e bruno; antenne brune con l’an-
tennomero basale bruno-rossiccio; zampe giallo-rossicce. La granulosità del capo è
saliente solo ai lati, quella del pronoto è evidente e quella delle elitre è distinta. La reti-
colazione del capo è netta e a maglie molto ampie sul disco e svanita ai lati, quella del
pronoto è nettissima, quella delle elitre è netta e quella dell’addome è distinta, a maglie
molto trasverse.

COMPARAZIONI. L’edeago della nuova specie è simile, in visione laterale, a
quello di A. inexpectata Pace, 1985 del Nepal, ma ha dimensione maggiore e la sua
parte apicale, in visione ventrale, è strettissima e a lati paralleli (parte apicale trian-
golare in inexpectata). L’armatura genitale interna dell’edeago, inoltre, è più forte nella
nuova specie.

Atheta (Philhygra) longefalcifera sp. n. Figg. 69-72

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m,
29°35N-102°00E, 5.VII.1998, leg. 1998 China Expedition J. Farkaé, D. Kral, J. Schneider,
A. Smetana (MHNG).

Paratypus: 1 d, China, W Sichuan, 20 km N Sabdé, 3200 m, 13.VII.1998, leg.
A. Smetana.

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero; antenne nere con l’anten-
nomero basale rossiccio; zampe rossicce. La punteggiatura del capo è molto svanita e
assente sulla fascia longitudinale mediana. La granulosità del pronoto e delle elitre è
fine e distinta. La reticolazione del capo è netta sul disco e svanita ai lati, quella del
pronoto è pure netta, quella delle elitre e dell'addome è distinta, a maglie molto
trasverse sull’addome. Edeago figg. 70-71, sesto urotergo libero del d fig. 72.

COMPARAZIONI. In visione ventrale l’edeago della nuova specie è simile a quello
di A. inexpectata Pace, 1985 del Nepal, ma la forte armatura genitale interna
dell’edeago della nuova specie non si ritrova nella specie a confronto.

Atheta (Philhygra) emeicola sp. n. Figg. 73-74
Holotypus 9, Chine, Mont Emei, 26.IV.1987, leg. Thierry Deuve (IRSN).
DESCRIZIONE. Lungh. 3,0 mm. Corpo lucido e nero-bruno, con elitre giallo-

brune; antenne nero-brune con antennomero basale bruno-rossiccio; zampe di un giallo

sporco, con femori posteriori bruni e medi giallo-bruni. La punteggiatura del capo è

composta di punti grandi e svaniti, assenti sulla fascia longitudinale mediana, quella

HYGRONOMINI E ATHETINI DELLA CINA 481

FIGG. 64-69

Sesto urotergo libero del ¢, habitus ed edeago in visione laterale e ventrale. 64: Atheta
(Philhygra) quadrifalcifera sp. n.; 65-68: Atheta (Philhygra) perconsanguinea sp. n.; 69: Atheta
(Philhygra) longefalcifera sp. n.

delle elitre è evanescente. La granulosità del pronoto è poco distinta. La reticolazione
del capo è netta, quella del pronoto, delle elitre e dell’addome è molto svanita, a maglie
molto trasverse sull’addome. Spermateca fig. 74.

482 R. PACE

COMPARAZIONI. La spermateca della nuova specie è simile a quella di A.
melanocera Thomson, diffusa nell’Europa centrale e settentrionale, in Siberia e Nord-
america, tuttavia la parte prossimale della spermateca è molto attenuata nella nuova
specie (larga in melanocera) e il bulbo distale della stessa spermateca è molto più
sviluppato nella nuova specie che in melanocera.

Atheta (Sipalatheta) emeimontis sp. n. Figg. 75-78

Holotypus d, China, Sichuan, Emei Shan, Leidongping, 2500 m, 29°32N-103°12E,
18.VII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypi: 15 es., stessa provenienza; 1 d e 2 9 ©, Chine, Mont Emei, 26.IV.1987, leg.
Thierry Deuve.

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e bruno-rossiccio, con capo e
uroterghi liberi terzo, quarto e base del quinto, bruni; antenne rossicce con l’undi-
cesimo antennomero bruno-rossiccio; zampe rossicce. La punteggiatura del capo è
fine. La granulosità del pronoto è distinta e quella delle elitre è saliente. La retico-
lazione del capo e del pronoto è molto svanita, quella delle elitre è distinta e quella del-
l’addome è estremamente superficiale, composta di maglie molto trasverse. Edeago
figg. 76-77, spermateca fig. 78.

COMPARAZIONI. La forma della spermateca della nuova specie è simile a quella
di A. algarum Pace, 1999, di Hong Kong , ma la nuova specie presenta elitre più corte
del pronoto che è più largo di esse. L’edeago della nuova specie è più regolarmente
arcuato al lato ventrale, mentre in algarum è sinuato, con un’armatura interna serpen-
tiforme, assente nell’edeago di algarum.

Atheta (Acrotona) fellowesi sp. n. Figg. 79-81

Holotypus d, China, Guangxi Pr., Nong Gang, nr. Long Rui, 26.V.1998, leg. J. Fellowes
(IRSN).

DESCRIZIONE. Lungh. 2,0 mm. Corpo lucido e di un giallo sporco, con elitre e
quarto urotergo libero giallo-bruni; antenne nero-brune con i due antennomeri basali e
la base del terzo gialli; zampe gialle. La punteggiatura del capo è ombelicata e svanita.
La granulosità del pronoto è fine e saliente, quella delle elitre e dell’addome è distinta,
a maglie molto trasverse sull’addome. La reticolazione del capo è estremamente
svanita, quella del pronoto e dell'addome è assente e quella delle elitre è distinta.
Edeago figg. 80-81.

COMPARAZIONI. L’edeago più simile a quello della nuova specie, a motivo della
presenza di forti spine basali interne dell’armatura genitale dello stesso edeago, è
quello di A. quadrispina Pace, 1987, del Nepal, ma manca nell’edeago di questa specie
un accentuato angolo ventrale, evidente invece nell’edeago della nuova specie e la
parte apicale dello stesso edeago, in visione ventrale, è molto più stretta in quadrispina.

DERIVATIO NOMINIS. La nuova specie è dedicata al suo raccoglitore, l’inglese
J. Fellowes.

Atheta (Acrotona) deuvei sp. n. Figg. 82-83

Holotypus 1 2, Chine, Mont Emei, 26.IV.1987, leg. Thierry Deuve (IRSN).
Paratypi: 2 2 2, China, Sichuan, Emei Shan, Leidongping, 2500 m, 18.VII.1998, leg. A.
Smetana.

HYGRONOMINI E ATHETINI DELLA CINA 483

0,1 mm

fob 11)!
ML 32) 1
i

Na
ITS

IRA
ENTER
È

Fico. 70-75

Edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 70-72: Atheta
(Philhygra) longefalcifera sp. n.; 73-74: Atheta (Philhygra) emeicola sp. n.; 75: Atheta (Sipa-
latheta) emeimontis sp. n.

DESCRIZIONE. Lungh. 2,3 mm. Corpo lucido e nero-bruno, con elitre e pigidio
bruni; antenne brune con i quattro antennomeri basali bruno-rossicci; zampe rossicce.
La granulosità dell’avancorpo è fine e distinta. La reticolazione del capo, del pronoto
e dell’addome è superficiale, quella delle elitre è molto evanescente, a maglie molto
trasverse sull’addome. Spermateca fig. 83.

484 R. PACE

0,1 mm

j ie È
o EIN ci È

ae 4
ci ira]
È © i

vi
ge
È

FIGG. 76-83

Edeago in visione laterale e ventrale, spermateca e habitus. 76-78: Atheta (Sipalatheta) emei-
montis sp. n.; 79-81: Atheta (Acrotona) fellowesi sp. n.; 82-83: Atheta (Acrotona) deuvei sp. n.

COMPARAZIONI. La spermateca della nuova specie è simile a quella di A. xisha-
nensis Pace, 1998, pure della Cina, ma la parte prossimale è più prolungata, sicché
descrive una spira e mezzo e non una sola, come in xishanensis. Il pronoto della nuova
specie, inoltre, è lievemente più largo delle elitre, mentre in xishanensis è nettamente

HYGRONOMINI E ATHETINI DELLA CINA 485

più stretto e gli antennomeri quarto e quinto sono trasversi nella nuova specie e lunghi
quanto larghi in xishanensis.

DERIVATIO NOMINIS. La nuova specie è dedicata al suo raccoglitore, Thierry
Deuve.

Atheta (Acrotona) luteihumeri sp. n. Figg. 84-85

Holotypus ©, China, SE Sichuan, Jinfo Shan, 1750 m, 29°01N-107°14E, 26.VI.1998,
leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypus: 1 ©, stessa provenienza.

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e nero-bruno, con uroterghi liberi
quarto e quinto neri e omeri e uroterghi liberi primo e secondo rossicci; antenne nero-
brune con i tre antennomeri basali rossicci; zampe rossicce. La punteggiatura del capo
è fine e distinta. La granulosità del pronoto è distinta e quella delle elitre è svanita. La
reticolazione del capo è molto svanita, quella del pronoto è molto superficiale, quella
delle elitre è fine e distinta e quella dell’addome è assente. Spermateca fig. 85.

COMPARAZIONI. La spermateca della nuova specie è simile a quella di A. irreg-
ularis Pace, 1984, della Birmania, ma la parte prossimale di quella della nuova specie
è nettamente più breve di quella corrispondente a irregularis. A. irregularis, inoltre,
presenta lunghe setole isolate ai lati del corpo e sulle tibie, assenti nella nuova specie.

DERIVATIO NOMINIS. Il nome della nuova specie significa «omeri gialli».

Atheta (Microdota) hailuogouensis sp. n. Figg. 86-89

Holotypus 9, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3050 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farkaë and P. Kabaték. (MHNG).
Paratypus: 1 9, stessa provenienza.

DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e giallo-bruno, con pronoto opaco
e pigidio giallo-rossiccio; antenne brune con i due antennomeri basali e la base del
terzo giallo-rossicci; zampe gialle. La granulosità del capo e del pronoto è distinta,
quella delle elitre è svanita e quella dell’addome è distinta. La reticolazione del capo è
netta sul disco e svanita ai lati, quella del pronoto è vigorosa e quella delle elitre e
dell’addome è distinta, composta di maglie lievemente trasverse sull’addome. Edeago
figg. 87-88, spermateca fig. 89.

COMPARAZIONI. L’edeago della nuova specie presenta alcuni caratteri dell’ede-
ago di A. bitruncata Pace, 1990, del Nepal, come la stretta parte apicale dello stesso
edeago. La nuova specie, tuttavia, presenta elitre lunghe quanto il pronoto e non molto
più lunghe come in bitruncata.

Atheta (Microdota) miriapex sp. n. Figg. 90-92

Holotypus d, China, Sichuan, Emei Shan, Leidongping, 2500 m, 29°35N-103°21E,
18.VII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DescRIZIONE. Lungh. 2,1 mm. Corpo lucido e bruno, con pronoto e pigidio
giallo-bruni; antenne brune con i due antennomeri basali bruno-rossicci; zampe gialle.
La granulosità del capo è fitta e svanita, quella delle elitre è finissima e quella delle
elitre è superficiale. La reticolazione del capo è netta solo nel fondo della depressione
mediana, al di fuori di essa è superficiale. La reticolazione del pronoto e dell’addome

486 R. PACE

FiGG. 84-89

Habitus, spermateca ed edeago in visione laterale e ventrale. 84-85: Arheta (Acrotona) lutei-
humeri sp. n.; 86-89: Atheta (Microdota) hailuogouensis sp. n.

è netta, quella delle elitre è molto svanita, a maglie poco trasverse sull’addome. Edeago
figg. 91-92.

COMPARAZIONI. Finora nessuna specie nota mostra l’apice dell’edeago dilatato
all’apice, in visione ventrale, come quello della nuova specie.

HYGRONOMINI E ATHETINI DELLA CINA 487

Atheta (Microdota) sichuanicola sp. n. Figg. 93-96

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana er al. (MHNG).

DESCRIZIONE. Lungh. 2,7 mm. Capo e pronoto debolmente lucidi, resto del
corpo lucido. Corpo poco convesso e nero-bruno, con elitre brune; antenne nero-brune
con i due antennomeri basali bruni; zampe gialle. La granulosità del capo è distinta,
assente sulla fascia longitudinale mediana che è largamente solcata. La granulosità del
pronoto è finissima, quella delle elitre è poco saliente e quella dell’addome lo è. La reti-
colazione dell’avancorpo è netta, quella dell’addome è distinta, a maglie lievemente
trasverse. Edeago figg. 94-95.

COMPARAZIONI. La nuova specie presenta l’edeago che mostra alcuni caratteri
dell’edeago di A. amiculoides Cameron, 1939, del Kashmir. L’armatura genitale
interna dell’edeago della nuova specie, tuttavia, è forte e complessa, semplice in
amiculoides, e l’edeago di amiculoides non è bruscamente ristretto, in visione ventrale,
come quello della nuova specie.

Atheta (Microdota) masculifrons sp. n. Figg. 97-101

Holotypus d, China, N Yunnan, Xue Shan, nr. Zhongdian, 4050 m, 27°49N-99°34E,
24.VI.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypi: 3 dd e 2 9 9, stessa provenienza.

DESCRIZIONE. Lungh. 3,6 mm. Corpo lucido e nero-bruno, con elitre brune; an-
tenne nero-brune con i tre antennomeri basali rossicci; zampe giallo-rossicce. La gra-
nulosità del capo è saliente solo ai lati, quella del pronoto e delle elitre lo è poco. La
reticolazione del capo è netta, ma ai lati è svanita, quella del pronoto e delle elitre è
evidente e quella dell’addome è distinta, a maglie lievemente trasverse sull’addome.
Edeago figg. 98-99, sesto urotergo libero del d fig. 100, spermateca fig. 101.

COMPARAZIONI. Per la forma della spermateca e dell’edeago, la nuova specie è
avvicinabile ad A. amiculoides Cameron, 1939, del Kashmir (tipi esaminati), ma la
taglia corporea è maggiore (1,8 mm in amiculoides), la parte prossimale della sper-
mateca descrive un ampio arco (breve in amiculoides) e per la più robusta armatura
genitale interna dell’edeago.

Atheta (Microdota) gibba sp. n. Figg. 102-105

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, for. above Camp 2, 2800 m,
29°35N-102°00E, 5.VII.1998, leg. 1998 China Expedition J. Farkaé, D. Kral, J. Schneider,
A. Smetana (MHNG).

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero-bruno, con elitre giallo-
brune; antenne nere con i due antennomeri basali e base del terzo di un giallo sporco;
zampe gialle. La granulosità del capo è distinta, quella del pronoto è appena saliente
sulla fascia longitudinale mediana e distinta ai lati, quella delle elitre è poco saliente.
La reticolazione del capo e dell’addome è netta, a maglie ondulate trasverse sull’ad-
dome, quella del pronoto e delle elitre è distinta. Il capo presenta una depressione
trasversa tra gli occhi. Edeago. 103-104, sesto urotergo libero del d fig. 105.

COMPARAZIONI. L’edeago della nuova specie è così al di fuori dagli schemi
consueti di quest’organo che non è comparabile con specie affini note.

488 R. PACE

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i
J ti
El 4
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\\

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)
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93 Cinna

Fico. 90-95

Habitus ed edeago in visione laterale e ventrale. 90-92: Atheta (Microdota) miriapex sp. n.; 93-
95: Atheta (Microdota) sichuanicola sp. n.

Atheta (Microdota) neoamicula sp. n. Fig. 106-107

Holotypus 2, China, Sichuan, Gongga Shan, Hailuogou, head of glacier 1, 2850 m,
29°35N-102°00E, 9.VII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,6 mm. Avancorpo debolmente lucido, addome lucido.
Corpo nero pece, con elitre bruno-rossicce; antenne brune con i tre antennomeri basali

HYGRONOMINI E ATHETINI DELLA CINA 489

\
ANUS Il In I
RL SS Le N

\
EU N ll

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AO

0,1 mm

1 mm

101

0,1 mm

FIGG. 96-101
Sesto urotergo libero del 4, habitus e spermateca. 96: Arheta (Microdota) sichuanicola sp. n.:
97-101: Atheta (Microdota) masculifrons sp. n.

rossicci; zampe gialle. La granulosità dell’avancorpo è fitta e poco evidente. La reti-
colazione del capo è svanita, quella del pronoto è distinta, quella delle elitre è poco
evidente e quella dell’addome è distinta, composta di maglie appena trasverse sull’ad-
dome. Spermateca fig. 107.

490 R. PACE

107

FIGG. 102-107

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 102-105:
Atheta (Microdota) gibba sp. n.; 106-107: Atheta (Microdota) neoamicula sp. n.

COMPARAZIONI. L’habitus e la forma della spermateca della nuova specie ricor-
dano da vicino A. amicula (Stephens, 1832), largamente diffusa nella regione pale-
artica. La spermateca della nuova specie, tuttavia, non presenta l’introflessione apicale
del bulbo distale, la parte prossimale della stessa spermateca è più lunga nella nuova
specie che in amicula. Il quarto antennomero della nuova specie, inoltre, è più lungo
che largo, mentre è trasverso in amicula.

HYGRONOMINI E ATHETINI DELLA CINA 491

Atheta (Microdota) tardoides sp. n. Figg. 108-109

Holotypus 2, China, Yunnan, Kunming, Western Hills, 1800 m, 25°04N-102°41E,
24.VII.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,3 mm. Corpo lucido e bruno, con omeri giallo-bruni;
antenne brune con i tre antennomeri basali giallo-rossicci; zampe giallo-rossicce. La
granulositä del capo è saliente e assente su una stretta area mediana anteriore, quella
del pronoto è superficiale, quella delle elitre è distinta. La reticolazione del capo, delle
elitre e dell’addome è distinta, quella del pronoto è netta, a maglie trasverse e ondulate
sull’addome. Spermateca fig. 109.

COMPARAZIONI. Per la forma della spermateca e per gli occhi ridotti, la nuova
specie è avvicinabile nettamente ad A. tarda Cameron, 1939, del Kashmir (tipi esa-
minati), ma quest’ultima specie ha elitre più corte del pronoto. La parte intermedia
della spermateca della nuova specie, inoltre, è più prolungata di quella di tarda.

Atheta (Microdota) zhongdianensis sp. n. Figg. 110-111

Holotypus 9, China, N Yunnan, Xue Shan, nr. Zhongdian, 3900 m, 27°49N-99°34E,
25.V1.1996, leg. A. Smetana, J. Farkaè and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,3 mm. Corpo lucido e bruno, con base delle elitre bruno-
rossiccia, uroterghi liberi quarto e quinto bruni e pigidio giallo-rossiccio; antenne
brune, con i due antennomeri basali giallo-rossicci; zampe giallo-rossicce. La granu-
losità del capo è molto superficiale, quella del pronoto è quasi indistinta e quella delle
elitre è poco saliente. La reticolazione del capo è svanita, quella del pronoto e dell’ad-
dome è netta, a maglie lievemente trasverse sull’addome, quella delle elitre è distinta.
Il capo presenta un largo solco mediano e il pronoto mostra un solco mediano inter-
rotto in avanti. Spermateca fig. 111.

COMPARAZIONI. Gli occhi ridotti, il largo solco mediano del capo e la forma della
spermateca avvicinano tassonomicamente la nuova specie ad A. disputanda Pace,
1984, del Nepal. Il capo, tuttavia, è nettamente reticolato in disputanda e superficial-
mente reticolato nella nuova specie e l’introflessione apicale del bulbo distale della
spermateca è a base larghissima nella nuova specie e assai stretta in disputanda.

Atheta (Microdota) liaoningensis sp. n. Figg. 112-113
Holotypus 9, China, Ying Kou Country, Liaoning Prov., 1987, leg. Li Jing Ke (IRSN).
DESCRIZIONE. Lungh. 2,2 mm. Corpo debolmente lucido e bruno, con addome

nero-bruno; antenne brune; zampe gialle. La granulosità del capo è assente, quella del

pronoto è visibile solo ai lati dove è distinta, quella delle elitre è superficiale. La reti-
colazione del capo e delle elitre è netta, quella del pronoto è vigorosa e quella dell’ad-

dome è distinta, composta su esso da maglie trasverse sugli uroterghi liberi primo a

quarto e a maglie poligonali irregolari sull’urotergo libero quinto. Spermateca fig. 113.
COMPARAZIONI. L’enorme bulbo distale della spermateca della nuova specie si

ritrova pure nella spermateca di A. inflatescens Pace, 1998, pure della Cina. La parte

prossimale della spermateca della nuova specie, tuttavia, è poco assottigliata e

descrivente un arco, mentre quella di inflatescens è molto lunga nella parte intermedia

e descrivente più ampie spire nella regione prossimale. È presente, inoltre, l’introfles-

sione apicale del bulbo distale della spermateca della nuova specie, mentre è assente in

inflatescens.

492 R. PACE

I) A rat
USS

AMIR) PA
AXE

FIGG. 108-113

Habitus e spermateca. 108-109: Atheta (Microdota) tardoides sp. n. sp. n.; 110-111: Atheta
(Microdota) zhongdianensis sp. n.; 112-113: Atheta (Microdota) liaoningensis sp. n.

Atheta (Dimetrota) angulispina sp. n. Figg. 114-117

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
13.VII.1998, leg. 1998 China Expedition J. Farkaë, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypus: 1 &, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3200 m,
7.VII.1996, leg A. Smetana er al. (MHNG).

HYGRONOMINI E ATHETINI DELLA CINA 493

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero, con elitre bruno-rossicce;
antenne nero-brune con i due antennomeri basali bruno-rossicci; zampe di un giallo
sporco. La punteggiatura del capo è distinta e assente sulla fascia longitudinale
mediana. La granulosità del pronoto e delle elitre è distinta. La reticolazione del capo
è netta sul disco e svanita ai lati, quella del pronoto è nettissima, quella delle elitre e
dell’addome è distinta, a maglie molto trasverse sull’addome. Edeago figg. 115-116,
sesto urotergo libero del ¢ fig. 117.

COMPARAZIONI. La nuova specie, per la forma dell’edeago, in visione ventrale,
è sicuramente affine ad A. pergranulosa Pace, 1998, pure della Cina, ma l’armatura
genitale interna dell’edeago è del tutto differente. Inoltre la nuova specie presenta
occhi ridotti, mentre pergranulosa li presenta lunghi quanto le tempie e il margine
posteriore del sesto urotergo libero del d è quasi lineare nella nuova specie e con
quattro lobi in pergranulosa.

Atheta (Dimetrota) ridenda sp. n. Figg. 118-119

Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 2, 2800 m,
29°35N-102°00E, 5.VII.1998, leg 1998 China Expedition J. Farkaë, D. Kral, J. Schneider,
A. Smetana (MHNG).

DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e nero-bruno, con omeri bruno-
rossicci; antenne nero-brune con i tre antennomeri basali giallo-rossicci; zampe giallo-
rossicce. La punteggiatura del capo è distinta e assente sulla fascia longitudinale
mediana. La granulosità del pronoto è distinta, quella delle elitre è svanita. La retico-
lazione del capo è netta sul disco e molto superficiale sul resto della sua superficie,
quella del pronoto è nettissima, quella delle elitre è evanescente e quella dell’addome
è distinta, composta di maglie molto trasverse sull’addome. Spermateca fig. 119.

COMPARAZIONI. Gli occhi ridotti e la forma della spermateca ricordano da vicino
A. disputanda Pace, 1984, del Nepal. Il bulbo distale della spermateca della nuova
specie, tuttavia, è poco più largo che lungo, se misurato dall’angolo d’inizio della parte
intermedia della stessa spermateca, mentre in disputanda è quasi due volte più lungo
che largo e l’introflessione apicale del bulbo distale della stessa spermateca ha la base
più larga. La parte prossimale della stessa spermateca, inoltre, è assottigliata nella
nuova specie e robusta in disputanda.

Atheta (Dimetrota) jiudingensis sp. n. Figg. 120-121

Holotypus ®, Chine, Sichuan, Massif du Jiu Ding Shan, 3500-3600 m, 31.VII.1994, leg.
Marc Tronquet (IRSN).
Paratypi: 2 9 2, stessa provenienza.

DESCRIZIONE. Lungh. 3,2 mm. Corpo lucido e nero, antenne comprese; zampe
rossicce con femori bruni. La granulosità del capo e delle elitre è distinta, quella del
pronoto è saliente. La reticolazione del capo e delle elitre è distinta, quella del pronoto
è netta e quella dell’addome è svanita, tranne nel fondo dei solchi trasversi basali degli
uroterghi dove è grossolana e netta. La reticolazione degli uroterghi liberi è molto
trasversa. Spermateca. 121.

COMPARAZIONI. La forma della spermateca della nuova specie rivela la sua
affinità con A. furtiva Cameron, 1939, dell’India e del Nepal (tipi esaminati). La parte
prossimale di quest’organo, infatti, è lievemente arcuata in entrambe le spermateche e

494 R. PACE

Ent
Br
Zn
Per

119

I mm

0,1 mm

Fico. 114-119

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del d e spermateca. 114-117:
Atheta (Dimetrota) angulispina sp. n.; 118-119: Atheta (Dimetrota) ridenda sp. n.

l’introflessione apicale del bulbo distale della stessa spermateca è in posizione asim-
metrica. I caratteri differenziali sono i seguenti: la parte prossimale e intermedia della
spermateca della nuova specie è nettamente più sottile e più prolungata di quella di
furtiva e l’introflessione apicale del bulbo distale della stessa spermateca della nuova

HYGRONOMINI E ATHETINI DELLA CINA 495

specie è poco distinta, mentre lo è in furtiva. Le setole isolate ai lati del corpo e delle
tibie della nuova specie, inoltre, sono nettamente più lunghe di quelle di furtiva.

Atheta (Datomicra) zhagaensis sp. n. Figg. 122-126
Holotypus d, Chine, Sichuan, Songpan, 3050 m, Zhaga Fall, 5.VIII.1994, leg. Marc
Tronquet (IRSN).

Paratypi: 1 d e 1 9, stessa provenienza.

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e bruno, con capo e uroterghi liberi
quarto e quinto nero-bruni ed elitre giallo-brune; antenne giallo-rossicce sfumate di
bruno verso l’estremità distale; zampe gialle. La granulosità del capo è saliente solo
sulle tempie, sul disco è molto evanescente, quella del pronoto e delle elitre è evidente.
La reticolazione del capo è netta sul disco e svanita ai lati, quella del pronoto e delle
elitre è netta e quella dell’addome è molto superficiale, composta di maglie molto
trasverse. Edeago figg 123-124, sesto urotergo libero del d fig. 125, spermateca fig.
126.

COMPARAZIONI. La nuova specie, per la forma della spermateca, appartiene
sicuramente al gruppo di A. inopinata Pace, 1991, del Nepal. Si distingue per i caratteri
dati nella chiave posta dopo la descrizione della seguente specie.

Atheta (Datomicra) consequens sp. n. Figg. 128-129

Holotypus 2, China, N Yunnan, Xue Shan, nr. Zhongdian, 4050 m, 27°49N-99°34E,
24.VI.1996, leg. A. Smetana, J. Farkaë and P. Kabaték (MHNG).

Paratypi: 3 © 2, stessa provenienza.

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e bruno, con pigidio bruno-rossic-
cio; antenne brune con i quattro antennomeri basali rossicci; zampe giallo-rossicce. La
granulosità del capo e delle elitre è distinta, quella del pronoto è saliente. La retico-
lazione del capo e delle elitre è netta, quella del pronoto è nettissima e quella dell’ad-
dome è molto superficiale, composta di maglie molto trasverse. Spermateca fig. 129.

COMPARAZIONI. La nuova specie è distinta da quelle del gruppo di A. inopinata
Pace, 1991, del Nepal, a cui appartiene per la forma della sua spermateca, per i caratteri
dati nella seguente chiave.

CHIAVE DELLE 9 ® DELLE SPECIE DI ATHETA DEL GRUPPO DI INOPINATA PACE

1 Bulbo prossimale della spermateca sferico, bulbiforme o piriforme ........ 2
- Parte prossimale della spermateca senza bulbo, ma descrivente una o
AUEZSIRUOSIEAM IO A. myohyangsani Pasnik
2 Bulbo prossimale della spermateca piriforme. Cina...... A. consequens Sp. n.
- Bulbo prossimale della spermateca sferico o bulbiforme ................. 3
3 Bulbo distale della spermateca lunghissimo e lineare. Nepal. A. inopinata Pace
= Bulbo distale della spermateca poco o fortemente ricurvo................ 4
4 Bulbordistale della’spermatecarpocorieurVyog LIE PTE EEE CE I
= Bulbordistale della spermateca’fortemente ricurvo, ee ae 7
5 Parte apicale del bulbo distale della spermateca assai piü corta della
Parte,basalexEoreage sr: PAIR ER A. photaechonica Pasnik

- Parte apicale del bulbo distale della spermateca più lunga della parte basale . 6

496 R. PACE

6 Parte prossimale del bulbo distale della spermateca, bruscamente ristret-
tayalliindietror Gina. I. cantate. 6408 RT SERRE A. graffa Pace
- Parte prossimale del bulbo distale della spermateca, gradualmente
niStrettataliindietro Cina... 5.42. oye ea N A. regressa Pace
7 Parte apicale del bulbo distale della spermateca lungo, sicché forma un
AMPIOAICO CIN Alas alah ee RE EE A. subinopinata Pace
- Parte apicale del bulbo distale della spermateca corto, sicché forma un
ATCOIStreto Le 0, HD NES GONE DRE 2 PR 8
8 Parte apicale del bulbo distale della spermateca molto sporgente. Cina

ELLE ARE TIER BEER A. zhagaensis sp. n.
- Parte apicale del bulbo distale della spermateca non sporgente
NIEREN NEE. SERIE LR, LE SD ATL AE ENO A. xinlongengensis Pace

Nota. Le illustrazioni dell’olotipo d e della femmina della specie di Pasnik,
Atheta namphoensis Pasnik, 2001, della Corea del Nord suggeriscono che i due sessi
non sono conspecifici. Il & si rivela appartenere al genere Nepalota, la spermateca
della 9 rivela l'appartenenza al sottogenere Datomicra. Una descrizione formale di
questa specie probabilmente nuova, basata solo su una 9, è prematura e si deve atten-
dere la disponibilità di più materiale, in particolare del à.

Nepalota namphoensis ( Pasnik, 2001), comb. n.
Atheta (Dimetrota) namphoensis Pasnik, 2001: 219, partim (solo l’holotypus
3), la 2 Atheta (Dimetrota) sp.

Atheta (Datomicra) songpanensis sp. n. Figg. 130-133

Holotypus d, Chine, Sichuan, Songpan, versant est, 3100-3300 m, 7.VIII.1994, leg.
Marc Tronquet (IRSN).

DESCRIZIONE. Lungh. 4,1 mm. Corpo lucido e bruno, con elitre giallo-brune;
antenne brune con i due antennomeri basali e la base del terzo giallo-bruni; zampe
gialle. La granulositä del capo è distinta solo sulle tempie, quella del pronoto è distinta
e quella delle elitre è superficiale. La reticolazione del capo e del pronoto è evidente,
quella delle elitre è netta e quella dell’addome è molto superficiale fatta eccezione per
la metà posteriore del quinto urotergo libero che presenta reticolazione netta e poco
trasversa, mentre sui restanti uroterghi liberi è molto trasversa. Il pronoto mostra un
appiattimento mediano posteriore. Edeago figg. 131-132, sesto urotergo libero del d
fe 153;

COMPARAZIONI. A motivo degli occhi molto sviluppati e della forma
dell’edeago, la nuova specie probabilmente è affine ad A. semitibialis Pace, 1998, pure
della Cina, da cui si distingue per l’edeago poco profondamente flesso al lato ventrale,
per l’armatura genitale interna dello stesso edeago molto differente e per il margine
posteriore del sesto urotergo libero del 4 quadrilobato (ondulato in semitibialis).

Atheta (Datomicra) falcifera sp. n. Figg. 134-137

Holotypus &, China, Sichuan, Gongga Shan, Hailuogou, above camp 2, 2800 m,
29°35N-102°00E, 5.VII.1998, leg 1998 China Expedition J. Farcaë, D. Kral, J. Schneider,
A. Smetana (MHNG).

HYGRONOMINI E ATHETINI DELLA CINA 497

he)

121

A Fate — ers N

i A \

fasi \
È Et |
È “Pa i

BU x
2 bs

1mm

Ficc. 120-126

Habitus, spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del 4. 120-121:
Atheta (Dimetrota) jiudingensis sp. n.; 122-126: Atheta (Datomicra) zhagaensis sp. n.

Paratypus: 1 d, China, W Sichuan, 20 km N Sabdé, 3200 m, 14.VII.1998, leg.
A. Smetana.

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e nero, con elitre nero-brune e
pigidio bruno; antenne nere con i due antennomeri basali bruno-rossicci; zampe
rossicce. La granulosità del capo è molto superficiale e assente sul disco, quella del

498 R. PACE

129 130

132

Fico. 127-133
Spermateca, edeago in visione laterale e ventrale e sesto urotergo libero del 3. 127: Atheta
(Datomicra) xinlongensis Pace; 128-129: Atheta (Datomicra) consequens sp. n.; 130-133:
Atheta (Datomicra) songpanensis sp. n.

pronoto molto svanita e quella delle elitre è distinta. La reticolazione del capo è net-
tissima sul disco e molto svanita al di fuori di esso, quella del pronoto è netta, quella
delle elitre è distinta e quella dell’addome è evanescente, composta di maglie molto
trasverse. Edeago figg. 135-136, sesto urotergo libero del d fig. 137.

COMPARAZIONI. La presenza di una forte spina falciforme dell’armatura genitale
interna dell’edeago della nuova specie si riscontra anche nell’edeago di A. regressa

HYGRONOMINI E ATHETINI DELLA CINA 499

Pace, 1998, pure della Cina, ma in quest’ultima specie la spina è nettamente meno
larga, meno ricurva e più lunga. L’apice dell’edeago della nuova specie, inoltre, è as-
sai stretto, in visione ventrale, e non ogivale come in regressa e il margine posteriore
del sesto urotergo libero del 3 è finemente ondulato nella nuova specie e quadriloba-
to in regressa.

Atheta (Datomicra) pertinens sp. n. Figg. 138-141

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
14.VII.1998, leg. 1998 China Expedition J. Farcaë, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero-bruno, con elitre brune,
omeri rossicci e uroterghi liberi primo e secondo e pigidio giallo-rossicci; antenne
brune con i due antennomeri basali e la base del terzo giallo-rossicci; zampe giallo-
rossicce. La granulosità del capo è distinta e assente sulla fascia longitudinale mediana,
quella del pronoto è fine e poco distinta e quella delle elitre è evidente. La reticolazione
è netta su tutto il corpo, sull’addome a maglie molto trasverse. Edeago 140-141, sesto
urotergo libero del à fig. 139.

COMPARAZIONI. A motivo della presenza di vari dettagli della struttura del-
l’edeago, la nuova specie probabilmente è affine ad A. xinlongensis Pace, 1998, pure
della Cina. Differisce da essa per avere una più forte armatura genitale interna del-
l’edeago che è nettamente più largo di quello di xinlongensis nella regione preapicale,
in visione ventrale. Il margine posteriore del sesto urotergo libero del d della nuova
specie, inoltre, è ondulato, mentre è quadrilobato in xinlongensis.

Atheta (Datomicra) stenotheca sp. n. Figg. 143-147

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, for. above camp 2, 2800 m,
29°35N-102°00E, 5.VII.1998, leg 1998 China Expedition J. Farcaë, D. Kral, J. Schneider,
A. Smetana (MHNG).

Paratypi: 14 es., stessa provenienza; 45 es., China, W Sichuan, 20 km N Sabdé, 3200 m,
13.VII.1998, leg. A. Smetana.

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero-bruno; antenne brune con i
due antennomeri basali e la base del terzo giallo-rossicci; zampe giallo-rossicce. La
granulosità del capo è fine e distinta e assente sulla fascia longitudinale mediana, quella
del pronoto è distinta e quella delle elitre è svanita. La reticolazione del capo e del
pronoto è netta, quella delle elitre è distinta e quella dell’addome è molto svanita,
composta di maglie molto trasverse. Edeago figg. 144-145, sesto urotergo libero del 4
fig. 146, spermateca fig. 147.

COMPARAZIONI. La spermateca della nuova specie ha forma simile a quella di A.
dalijiamontis Pace, 1998, pure della Cina, il cui d è sconosciuto. La parte apicale del
bulbo distale della spermateca della nuova specie, tuttavia, è nettamente corta, mentre
è lunga e lievemente ricurva in dalijiamontis e le elitre della nuova specie sono molto
più lunghe del pronoto (poco più lunghe in dalijiamontis).

Atheta (Datomicra) latifalcifera sp. n. Figg. 148-151

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3300 m, 29°35N-102°23E,
14.VII.1998, leg. 1998 China Expedition J. Farcaë, D. Kral, J. Schneider, A. Smetana (MHNG).
Paratypus: 1 d, stessa provenienza.

500 R. PACE

Fico. 134-141

Habitus, edeago in visione laterale e ventrale e sesto urotergo libero del d. 134-137: Atheta
(Datomicra) falcifera sp. n.; 138-141: Atheta (Datomicra) pertinens sp. n.

DESCRIZIONE. Lungh. 3,6 mm. Corpo lucido e bruno, con primo urotergo libero
rossiccio; antenne brune con i due antennomeri basali rossicci; zampe rossicce. La
granulosità del capo è fine e assente sulla fascia longitudinale mediana, quella del
pronoto è pure fine ed evidente, quella delle elitre è superficiale. La reticolazione del
capo e del pronoto è nettissima, quella delle elitre è distinta e quella dell’addome è

HYGRONOMINI E ATHETINI DELLA CINA 501

0,1 mm

142

1 mm

0,1 mm

147

FIGG. 142-148

Spermateca, habitus, edeago in visione laterale e ventrale e sesto urotergo libero del ¢. 142:
Atheta (Datomicra) parainopinata Pace, 1998; 143-147: Atheta (Datomicra) stenotheca sp. n.;
148-151: Atheta (Datomicra) latifalcifera sp. n.

svanita, composta di maglie molto trasverse. Edeago figg. 149-150, sesto urotergo
libero del à fig. 151.

COMPARAZIONI. L’edeago della nuova specie è simile a quello di A. graffa Pace,
1998, pure della Cina, per cui sono molto probabili le affinità tassonomiche. Nella
nuova specie, tuttavia, un pezzo dell’armatura genitale interna dell’edeago è forte-

502 R. PACE

Fico. 149-155

Edeago in visione laterale e ventrale, sesto urotergo libero del d e habitus. 149-151: Atheta
(Datomicra) latifalcifera sp. n.; 152-155: Atheta (Datomicra) mesofalcifera sp. n.

mente sviluppata e robusta, mentre il corrispondente in graffa è relativamente esile. Il
margine posteriore del sesto urotergo libero del & della nuova specie è quadrilobato,
mentre in graffa è bilobato e inciso a metà.

HYGRONOMINI E ATHETINI DELLA CINA 503

Atheta (Datomicra) mesofalcifera sp. n. Figg. 152-155

Holotypus ¢, China, N Yunnan, Xue Shan, nr. Zhongdian, 4050 m, 27°49N-99°34E,
24.VI.1996, leg. A. Smetana, J. Farcaë and P. Kabaték (MHNG).

Paratypus: 16, China, N Yunnan, Xue Shan, nr. Zhongdian, 4000-4100 m, 23.VI.1996,
leg. A. Smetana.

DESCRIZIONE. Lungh. 3,0 mm. Corpo lucido e bruno, con elitre bruno-rossicce
e uroterghi liberi terzo a quinto neri; antenne brune con i quattro antennomeri basali
giallo-rossicci; zampe gialle. La granulosità del capo è distinta e assente sulla fascia
longitudinale mediana, quella del pronoto e delle elitre è saliente. La reticolazione
dell’avancorpo è netta, quella dell’addome è distinta, composta di maglie molto
trasverse. Edeago figg. 153-154, sesto urotergo libero del d fig. 155.

COMPARAZIONI. Per la forma dell’edeago la nuova specie è molto probabilmente
strettamente affine ad A. xinlongensis Pace, 1998, pure della Cina. Se ne distingue per
l’armatura genitale interna dell’edeago più forte e per l’apice dell’edeago, in visione
ventrale, largo, quasi tronco e non appuntito e stretto come quello di xinlongensis. Il
margine posteriore del sesto urotergo libero del 4 è ondulato nella nuova specie e
quadrilobato in xinlongensis.

Atheta (Datomicra) hamifera sp. n. Figg. 156-159

Holotypus d, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E, 14.VII.
1998, leg. 1998 China Expedition J. Farcac, D. Kral, J. Schneider, A. Smetana (MHNG).

Paratypi: 1 d, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3200 m,
7.VII.1996, leg. A. Smetana; 1 d, China, Sichuan, Kangding (formerly Tatsienlu), 2900 m,
2.VII.1996, leg. A. Smetana.

DESCRIZIONE. Lungh. 4,4 mm. Corpo lucido e nero, con elitre brune; antenne
nere con i due antennomeri basali rossicci; zampe giallo-rossicce. La granulosità del
capo è superficiale, quella del pronoto è saliente e quella delle elitre è superficiale. La
reticolazione del capo è netta, quella del pronoto è nettissima, quella delle elitre è
distinta e quella dell’addome è molto svanita, composta di maglie molto trasverse.
Edeago figg. 157-158, sesto urotergo libero del S fig. 159.

COMPARAZIONI. A motivo di più di un carattere, l’edeago della nuova specie è
simile a quello di A. regressa Pace, 1998, pure della Cina. La parte apicale di
quest’organo, in visione ventrale, è stretta e molto più lunga del bulbo basale; due sono
le spine dell’armatura genitale interna dell’edeago ed altro. L’edeago della nuova
specie, tuttavia, è nettamente più arcuato al lato ventrale e una delle due spine dell’ar-
matura genitale interna dell’edeago si è trasformata in un lunghissimo e forte pezzo
falciforme.

Atheta (Oreostiba) micromega sp. n. Figg. 160-161

Holotypus ©, China, Sichuan, Moxi, 1600 m, 29°13N-102°10E, 2.VII.1998, leg. 1998
China Expedition J. Farcac, D. Kral, J. Schneider, A. Smetana (MHNG).

DESCRIZIONE. Lungh. 1,9 mm. Corpo lucido e bruno, con pronoto bruno-ros-
siccio e uroterghi liberi primo e secondo e pigidio giallo-rossicci; antenne nero-brune
con antennomero basale rossiccio; zampe giallo-rossicce. La punteggiatura del capo €
fitta e distinta. La granulosita del pronoto è finissima, quella delle elitre è distinta. La
reticolazione del capo, delle elitre e dell’addome è molto svanita, quella del pronoto è

504 R. PACE

Fico. 156-161

Habitus, edeago in visione laterale e ventrale, sesto urotergo libero del 9 e spermateca. 156-159:
Atheta (Datomicra) hamifera sp. n.; 160-161: Atheta (Oreostiba) micromega sp. n.

assente. Le maglie della reticolazione dell’addome sono poco trasverse. Spermateca
fig. 161.

COMPARAZIONI. La forma della spermateca della nuova specie ricorda da vicino
quella di A. sibirica Mäklin, 1880 (vedi Brundin, 1940), del nord della Siberia. Il bulbo

HYGRONOMINI E ATHETINI DELLA CINA 505

Fico. 162-167

Habitus e spermateca. 162-163: Arheta (Oreostiba) shanicola Pace; 164-165: Atheta (Oreostiba)
transcripta sp. n.; 166-167: Atheta (Oreostiba) mimomonticola sp. n.

distale, tuttavia, è nettamente flesso nella nuova specie e la parte prossimale descrive
una sinuosità. Inoltre, sibirica ha taglia corporea di 3,0 mm.
Atheta (Oreostiba) transcripta sp. n. Figg. 164-165

Holotypus 2, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

506 R. PACE

Paratypus: 1 9, China, Sichuan, Kangding (formely Tatsienlu), 2900 m, 2.VII.1996, leg.
A. Smetana.

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e bruno, con elitre bruno-rossicce;
antenne brune con i tre antennomeri basali giallo-rossicci; zampe gialle. La granulosità
del capo è fine, quella del pronoto e delle elitre è distinta. La reticolazione del capo e
del pronoto è netta, quella delle elitre è svanita e quella dell’addome è distinta,
quest’ultima composta di maglie molto trasverse. Spermateca fig. 165.

COMPARAZIONI. La nuova specie per la forma della spermateca è simile ad
A. sibirica Mäklin, 1880 (vedi Brundin, 1940), del nord della Siberia, ma la parte
intermedia dello stesso organo non descrive un ampio arco come in sibirica ed è pre-
sente nella nuova specie un’introflessione apicale del bulbo distale della spermateca,
assente in sibirica.

Atheta (Oreostiba) mimomonticola sp. n. Figg. 166-167

Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3100 m,
29°35N-102°00E, 8.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucidissimo e nero-bruno, con uroterghi
liberi terzo e quarto neri; antenne nere, con i due antennomeri basali e la base del terzo
rossicci; zampe giallo-brune. La punteggiatura del capo è ombelicata e svanita: La
granulosità del pronoto è distinta, quella delle elitre è poco evidente. La reticolazione
del capo è assente, quella del pronoto è superficiale, quella delle elitre è netta e quella
dell’addome è distinta, quest’ultima a maglie molto trasverse. Spermateca fig. 167.

COMPARAZIONI. La forma della spermateca della nuova specie è simile a quella
di A. monticala Thomson, 1852, diffusa nell’Europa centrale e settentrionale, ma
appartenente al sottogenere Bessobia Thomson, 1858, per avere quattro solchi trasversi
basali degli uroterghi liberi. È simile pure alla spermateca di A. yonghicola Pace, 1998,
pure della Cina, ma questa specie presenta occhi più corti delle tempie e la parte prossi-
male della spermateca più lunga, tanto che descrive due sinuosità, invece di una come
nella nuova specie.

Atheta (Atheta) paralaevicauda sp. n. Figg. 168-169

Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

Paratypus: 1 ?, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
7.V11.1996, leg A. Smetana.

DESCRIZIONE. Lungh. 4,7 mm. Corpo lucido e nero; antenne nere con l’anten-
nomero basale nero-bruno; zampe giallo-brune. La punteggiatura del capo è fine e dis-
tinta, quella del pronoto è poco evidente. La granulosità delle elitre è poco distinta. La
reticolazione del capo e delle elitre è svanita, quella del pronoto è molto superficiale e
quella dell'addome è assente. Spermateca fig. 169.

COMPARAZIONI. A motivo della forma della spermateca, la nuova specie è molto
affine ad A. laevicauda Sahlberg, 1876, specie boreoalpina europea. Se ne distingue
per la parte intermedia della spermateca coperta di tubercoli in modo più esteso e la
parte prossimale più lunga. Inoltre, la sutura delle elitre è più lunga del pronoto (lieve-
mente più corta in /aevicauda), il quarto antennomero è più lungo che largo (trasverso
in laevicauda) e il quinto urotergo libero non è reticolato (reticolato in laevicauda).

HYGRONOMINI E ATHETINI DELLA CINA 507

1 mm

RE re Di

Peril ele rés
Siléa) rit

1 mm

171 172

FIGG. 168-173

Habitus, spermateca, edeago in visione laterale e ventrale e labio con palpo labiale. 168-169:
Atheta (Atheta) paralaevicauda sp. n.; 170-173: Atheta (Ekkliatheta subg. n.) aniiensis sp. n.

Ekkliatheta subg. n. di Atheta

TYPUS SUBGENERIS. Atheta (Ekkliatheta) aniiensis sp. n.

DESCRIZIONE. A motivo della presenza del primo tarsomero posteriore più lungo
dell’immediatamente successivo la nuova specie poteva essere ipoteticamente essere
attribuita al genere Gnypeta Thomson, 1858, ma questo genere comprende specie con

508 R. PACE

ligula divisa fin quasi alla sua base che è stretta e non larga come quella del nuovo
sottogenere.

DERIVATIO NOMINIS. Il nome femminile del nuovo sottogenere significa «Atheta
deviata».

Atheta (Ekkliatheta subgen. n.) anüensis sp. n. Figg. 170-173

Holotypus d, China, Zhejiang Prov., Anii County, ca. 500 m, Long Wan Shan N.R.,
14.V.1996, leg. J. Cooter (IRSN).

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero, con elitre giallo-brune;
antenne interamente nere; zampe nero-brune con tarsi di un giallo sporco e ginocchia
bruno-rossicce. La punteggiatura del capo è molto svanita e assente sulla fascia longi-
tudinale mediana, quella del pronoto è fittissima e svanita e quella delle elitre è fitta e
superficiale. La reticolazione del capo, dell’addome e delle elitre è distinta, quella del
pronoto è molto svanita è distinta è distinta, sull’addome a maglie molto trasverse. Il
disco del capo è largamente concavo. Il pronoto presenta un appiattimento mediano.
Edeago figg. 171-172, labio con palpo labiale fig. 173.

COMPARAZIONI. La ligula a base larga isola la nuova specie dalle restanti del
genere appartenenti ad altri sottogeneri che presentano ligula assai ristretta alla base.
Pertanto è qui proposto il nuovo sottogenere di Arheta.

Nepalota fellowesi sp. n. Figg. 174-177

Holotypus d, China, Guandong, Da Wu Ling, 1600 m, 26.IV.1997, leg. J. Fellowes
(IRSN).

Paratypi: 2 2 2, stessa provenienza.

DESCRIZIONE. Lungh. 3,8 mm. Corpo lucido e nero-bruno, con margine
posteriore degli uroterghi liberi rossiccio; antenne brune, con i due antennomeri basali
giallo-rossicci e l'undicesimo rossiccio; zampe giallo-rossicce. La punteggiatura del
capo è fine e svanita e assente sulla fascia longitudinale mediana. La granulosità del
pronoto è superficiale e quella delle elitre è distinta. La reticolazione del capo è super-
ficiale, quella del pronoto e delle elitre è distinta e quella dell’addome è assente.
Edeago figg. 175-176, spermateca fig. 177.

COMPARAZIONI. Per la forma dell’edeago, la nuova specie si differenzia chiara-
mente dalle specie note, poiché esso è ampiamente arcuato al lato ventrale e l'armatura
genitale interna dell’edeago è del tutto differente. La forma della spermateca non si
osserva uguale o simile in altre specie.

DERIVATIO NOMINIS. La nuova specie è dedicata al suo raccoglitore, l’inglese
J. Fellowes.

Nepalota guangdongensis sp. n. Figg. 178-179

Holotypus 9, China, Guandong Pr, Ding Hu Shan, Biosphere R.,6.V.1998, leg.
J. Fellowes (IRSN).

DESCRIZIONE. Lungh. 3,5 mm. Corpo lucido e rossiccio, con capo nero, pronoto
giallo-rossiccio e metà posteriore del quarto urotergo libero bruno-rossiccio e una
macchia bruna ai lati di ciascun urotergo libero al suo angolo posteriore; antenne
rossicce con il primo antennomero basale e l’undicesimo giallo-rossicci; zampe giallo-

HYGRONOMINI E ATHETINI DELLA CINA 509

175

177

0,1 mm

Fico. 174-178

Habitus, edeago in visione laterale e ventrale e spermateca. 174-177: Nepalota fellowesi sp. n.;
178: Nepalota guangdongensis sp. n.

rossicce con ginocchia rossicce. La granulosità del capo è fine e assente sulla fascia
longitudinale mediana, quella del pronoto è superficiale e quella delle elitre è distinta.
La reticolazione del capo è svanita, quella del pronoto e delle elitre è distinta.
Spermateca fig. 179.

510 R. PACE

RAR ba
\1,\98

FIGG. 179-185

Spermateca, habitus ed edeago in visione laterale e ventrale. 179: Nepalota guangdongensis sp.
n.; 180-182: Paraloconota puella sp. n.; 183-184: Paraloconota yonghaiensis Pace figg.; 185:
Paraloconota dalijiamontis sp. n.

COMPARAZIONI. La spermateca della nuova specie presenta una robusta intro-
flessione apicale del bulbo distale e la parte prossimale bisinuata. Per questi caratteri si
mostra affine a N. gansuensis Pace, 1998 e N. chinensis Pace, 1998, entrambe della
Cina. La nuova specie se ne distingue per avere la parte distale della stessa spermateca

HYGRONOMINI E ATHETINI DELLA CINA 511

nettamente più lunga e la parte prossimale corta e non dilatata apicalmente, non
raggiungente la metà della sua parte intermedia (raggiungendola o superandola nelle
due specie a confronto).

Paraloconota puella sp. n. Figg. 180-182

Holotypus d, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farac & P. Kabatek (MHNG).

DESCRIZIONE. Lungh. 2,7 mm. Corpo lucido e nero, comprese le antenne e le
zampe. La punteggiatura del capo è molto superficiale, assente sulla fascia longitu-
dinale mediana, quella del pronoto e delle elitre è indistinta. La granulosità dell’ad-
dome è evanescente. La reticolazione del capo, delle elitre e dell'addome è distinta,
quella del pronoto è superficiale. Il pronoto presenta un debole e largo solco mediano
posteriore. Edeago figg. 181-182.

COMPARAZIONI. Per la forma dell’edeago, la nuova specie si mostra affine a
P. montium Pace, 1998, ma l’apice dell’edeago della nuova specie è stretto e inciso a
metà e non assai largo e intero come in montium.

Paraloconota dalijiamontis sp. n. Figg. 185-187

Holotypus d, China, Gansu, Dalijia Shan, 46 km W Linxia, 2980 m, 10.VII.1994, leg.
A. Smetana (MHNG).

DESCRIZIONE. Lungh. 4,1 mm. Corpo lucido e nero; antenne bruno-rossicce con
l’antennomero basale bruno; zampe bruno-rossicce, con tarsi rossicci. La punteggia-
tura del capo e del pronoto non è ben distinta, quella delle elitre è indistinta. La reti-
colazione del capo e del pronoto è nettissima, quella delle elitre è distinta, ma composta
di maglie alquanto confuse e quella dell'addome è distinta, composta di maglie
trasverse. Il capo presenta una larga depressione discale e il pronoto mostra un largo
solco mediano, assente in avanti. Edeago figg. 186-187.

COMPARAZIONI. La nuova specie per la forma dell’edeago è tassonomicamente
assai vicina a P. gansuensis Pace, 1998, pure della Cina. Se ne distingue per avere
l’apice dell’edeago con concavità ventrale e soprattutto per la robusta e ricurva
armatura genitale interna dell’edeago.

Oroekklina smetanai sp. n. Figg. 190-193

Holotypus 4, China, N Yunnan, Xue Shan, nr. Zhongdian, 4050 m, 27°49N-99°34E,
24.VI.1996, leg. A. Smetana, J. Farkac and P. Kabaték (MHNG).

Paratypus: 1 2, China, N Yunnan, Xue Shan, nr. Zhongdian, 3900 m, 25.VI.1996, leg.
A. Smetana.

DESCRIZIONE. Lungh. 1,8 mm. Corpo lucido e nero-bruno, con addome nero;
antenne nero-brune; zampe bruno-rossicce. La granulosità del capo è superficiale e
assente sulla fascia longitudinale mediana, quella del pronoto e dell’addome è distinta
e quella delle elitre è saliente. La reticolazione del capo è evanescente sul disco e
assente ai lati, quella del pronoto, delle elitre e dell’addome è distinta, a maglie di reti-
colazione trasverse sull’addome. Edeago figg. 191-192, spermateca fig. 193.

COMPARAZIONI. L’edeago della nuova specie è strettamente arcuato al lato
ventrale e l’apice, in visione ventrale, è diviso in due parti larghe e gradualmente
ristrette: Per questi caratteri è nettamente distinta da O. daxuensis Pace, 1999, pure

512 R. PACE

1 mm

ER (o
Li) Reso:

Fico. 186-193

Edeago in visione laterale e ventrale, habitus e spermateca. 186-187: Paraloconota dalijiamontis
sp. n.; 188-189: Paraloconota gansuensis Pace, 1998; 190-193: Oroekklina smetanai sp. n.

della Cina, che presenta edeago ampiamente bisinuato al lato ventrale e l’apice, in
visione ventrale, diviso in due parti strette e arcuate (vedi Assing, 2002b).

DERIVATIO NOMINIS. La nuova specie è dedicata a colui che con la sue ricerche
ha maggiormente contribuito alla conoscenza delle Aleocharinae della Cina, il Dr.
A. Smetana di Ottawa, noto studioso di Staphylinidae.

HYGRONOMINI E ATHETINI DELLA CINA 513

Ficc. 194-199

Habitus ed edeago in visione laterale e ventrale. 194-196: Emmelostiba granulosa sp. n.; 197-
199: Emmelostiba bellicosa sp. n.

Emmelostiba granulosa sp. n. Figg. 194-196

Holotypus é, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

Paratypi: 2 d d, China, Sichuan, Gongga Shan, Hailuogou, head of glacier 1, 2850 m,
9.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték.

514 R. PACE

DESCRIZIONE. Lungh. 2,4 mm. Corpo lucido e nero, con elitre nero-brune;
antenne brune; zampe bruno-rossicce. La granulositä del capo € molto superficiale,
quella del pronoto e delle elitre è fitta ed evanescente e quella dell’addome è saliente.
La reticolazione del capo è netta, quella del pronoto e dell’addome è distinta e fine.
Edeago figg. 195-196.

COMPARAZIONI. Per la forma dell’edeago la nuova specie è avvicinabile unica-
mente a E. pretiosa Pace, 1985, dell’ India, ma quest’ultima specie presenta elitre molto
più corte del pronoto, oltre a un edeago molto più piccolo.

Emmelostiba bellicosa sp. n. Figg. 197-199

Holotypus 6, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3050 m,
29°35N-102°00E, 6.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,9 mm. Corpo lucido e bruno, con omeri bruno-rossicci
e quarto urotergo libero nero e iridescente; antenne brune con i tre antennomeri basali
giallo-rossicci e il quarto rossiccio; zampe giallo-rossicce. La punteggiatura del capo è
molto superficiale e assente sulla fascia longitudinale mediana. La granulosità del
pronoto è fine e distinta, quella delle elitre è evanescente. La reticolazione del capo è
assente, quella del pronoto è molto svanita, quella delle elitre è distinta e quella
dell’addome è molto superficiale. Edeago figg. 197-198.

COMPARAZIONI. L’edeago della nuova specie è dilatato all’apice, in visione
laterale, come in E. franzi Pace, 1982, del Nepal. Questa specie, tuttavia, presenta elitre
molto più corte del pronoto, armatura genitale interna dell’edeago debole e minore
dilatazione apicale, in visione laterale. Nonostante queste notevoli differenze, l’affinitä
tassonomica tra esse va ritenuta piuttosto certa.

Alevonota pulchricornis sp. n. Figg. 200-202

Holotypus d, Sichuan, Moxi, 1600 m, 29°13N-102°10E, 2.VII.1998, leg. A. Smetana
(MHNG).

DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e bruno, con margine posteriore dei
due uroterghi liberi basali rossicci; antenne rossicce con l’antennomero basale bruno e
il secondo e il terzo bruno-rossicci; zampe giallo-rossicce. La punteggiatura del capo €
appena distinta, quella del pronoto é finissima e assai superficiale. La granulosita delle
elitre è evanescente. La reticolazione del capo è superficiale, quella del pronoto è molto
svanita, quella delle elitre e dell’addome è distinta, a maglie poligonali irregolari
sull’addome. Il capo presenta un debole solco mediano.

COMPARAZIONI. La nuova specie è ben distinta da A. sericata Pace, 1998, pure
della Cina, per l’edeago di maggiori dimensioni, con armatura genitale interna netta-
mente differente.

Alevonota sinensis sp. n. Figg. 203-205

Holotypus 3, Chine, Guanxi, Massif de Yuanpao-Shan, 500-1300 m, 11-13.VIII.1994,
leg. Marc Tronquet (IRSN).

DESCRIZIONE. Lungh. 3,7 mm. Corpo lucido e bruno, con capo nero-bruno e i
due uroterghi liberi basali e pigidio giallo-bruni; antenne rossicce con i quattro anten-
nomeri basali nero-bruni; zampe giallo-rossicce. La punteggiatura del capo e del

HYGRONOMINI E ATHETINI DELLA CINA 515

Ficc. 200-205

Habitus ed edeago in visione laterale e ventrale. 200-202: Alevonota pulchricornis sp. n.;
203-205: Alevonota sinensis Sp. n.;

pronoto è rada e quasi indistinta. La granulosità delle elitre è fine e molto evanescente.
La reticolazione del capo, delle elitre e dell'addome è distinta, quella del pronoto è

lievemente evanescente, a maglie poligonali irregolari sull’addome. Edeago figg.
203-204.

516 R. PACE

COMPARAZIONI. L’edeago della nuova specie possiede l’armatura genitale
interna nettamente differente, sia da quella di A. sericata Pace, 1998, pure della Cina,
che da quella della nuova specie sopra descritta.

Alevonota foedicornis sp. n. Figg. 206-207

Holotypus 9, China, Sichuan, Emei Shan, Leidongping, 2500 m, 29°32N-103°21E,
18.VII.1996, leg. A. Smetana, J. Farkac and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 4,0 mm. Corpo lucido e bruno scuro; addome bruno-
rossiccio con base del quarto urotergo libero e base del quinto bruni; antenne giallo-
rossicce con antennomero basale bruno; zampe giallo-rossicce. La granulosità del capo
è debolmente saliente, quella del pronoto è netta e quella delle elitre è distinta. La
reticolazione del capo è ben visibile, quella del pronoto è nettissima e quella delle elitre
e dell'addome è evidente, a maglie lievemente trasverse sull’addome. Il pronoto
presenta uno stretto solco mediano.

COMPARAZIONI. La nuova specie presenta una spermateca simile a quella di A.
subcorticalis (Cameron, 1939), comb. n. (olim Atheta (Traumoecia) subcorticalis
Cameron, 1939: 315, holotypus 2 esaminato, olim Tomoglossa subcorticalis: Pace,
1991: 835), dell’India. Se ne distingue per aver la parte intermedia della spermateca
nettamente più breve della lunghezza del bulbo distale dello stesso organo, mentre in
subcorticalis ha questa porzione nettamente più lunga del bulbo distale corrispondente.
L’introflessione apicale del bulbo distale della spermateca, inoltre, è forte nella nuova
specie ed esile in subcorticalis e la taglia della spermateca della nuova specie è quasi
il doppio maggiore che quella di subcorticalis. Pochi e sottili sono i caratteri differen-
ziali esterni.

Taxicera sinensis sp. n. Figg. 208-209

Holotypus 9, China, Sichuan, Gongga Shan, Hailuogou, head of glacier 1, 2850 m, 29°35N-
102°00E, 9.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

DESCRIZIONE. Lungh. 2,6 mm. Corpo lucido e nero, comprese le antenne e le
zampe con i tarsi. La punteggiatura del capo è distinta e assente sulla fascia longitu-
dinale mediana, quella del pronoto è evidente e quella delle elitre è poco profonda. La
reticolazione dell’avancorpo è netta, quella dell'addome è superficiale, composta di
maglie trasverse sull’addome. Il pronoto presenta una depressione mediana posteriore.
Spermateca fig. 209.

COMPARAZIONI. Sia la spermateca. che molti caratteri esterni differenziano la
nuova specie da 7. smetanai Pace, 1998, pure della Cina. Il bulbo distale della sper-
mateca della nuova specie è più sviluppato, con introflessione apicale appena visibile
(molto profonda in smetanai) e la parte prossimale dello stesso organo è robusta e pre-
senta un’appendice, assente in smetanai. Il quarto antennomero è trasverso nella nuova
specie e più lungo che largo in smetanai e il pronoto mostra due fossette mediane
posteriori, invece di una, come in smetanai.

Amidobia perconvexa sp. n. Figg. 210-211

Holotypus ©, China, Sichuan, Gongga Shan, Hailuogou, above camp 3, 3000 m,
6.VII.1996, leg A. Smetana, J. Farkac and P. Kabaték (MHNG).

HYGRONOMINI E ATHETINI DELLA CINA sf

1 mm

ANNE |
AN J
INN Sale Je

1 mm

0,1 mm
0,1 mm

209 210

Fico. 206-211

Habitus e spermateca. 206-207: Alevonota foedicornis sp. n.; 208-209: Taxicera sinensis sp. n.:
210-211: Amidobia perconvexa sp. n.

DESCRIZIONE. Lungh. 3,1 mm. Avancorpo debolmente lucido, addome lucido.
Corpo nero, con elitre e pigidio nero-bruni; antenne nere con i tre antennomeri basali
rossicci; zampe rossicce. La granulosità del capo è poco distinta, quella del pronoto è
evidente. La punteggiatura delle elitre è distinta. La reticolazione del capo e del

518 R. PACE

pronoto è netta, quella delle elitre è poco distinta e quella dell’addome è distinta,
composta di maglie molto trasverse. Spermateca fig. 210.

COMPARAZIONI. La spermateca distingue nettamente la nuova specie da A.
smetanai Pace 1998, pure della Cina, innanzitutto per la taglia maggiore, di 1/4 più
lunga. L’introflessione apicale del bulbo distale della spermateca della nuova specie ha
base stretta, mentre quella di smetanai è larghissima e la parte intermedia della stessa
spermateca è accentuatamente arcuata nella nuova specie e appena in smetanai. Le
antenne della nuova specie, inoltre, sono bicolori nere con base rossiccia, mentre in
smetanai sono unicolori nere.

Amidobia ming sp. n. Figg. 212-213

Holotypus 9, China, W Sichuan, 20 km N Sabdé, 3200 m, 29°35N-102°23E,
14.VII.1998, leg. A. Smetana (MHNG).

DESCRIZIONE. Lungh. 4,3 mm. Corpo lucido e nero, con elitre nero-brune;
antenne nere; zampe rossicce con femori bruno-rossicci. La punteggiatura del capo è
superficiale, anche sul disco, quella delle elitre è fine e poco evidente. La granulosità
del pronoto è fine e distinta. La reticolazione del capo è evanescente, quella del pro-
noto è distinta, quella delle elitre e dell’addome è netta, a maglie molto trasverse
sull’addome. Spermateca fig. 213.

COMPARAZIONI. La nuova specie è distinta da A. smetanai Pace, 1998, pure della
Cina, per le elitre molto meno lunghe, ma soprattutto per la spermateca che è forte-

0,1 mm

213

FicG. 212-213
Habitus e spermateca. 212-213: Amidobia ming sp. n.

HYGRONOMINI E ATHETINI DELLA CINA 519

mente arcuata nella parte intermedia (appena situata in smetanai) e per l’introflessione
apicale del bulbo distale della stessa spermateca assai meno larga alla base che quella
di smetanai.

NOTE SINONIMICHE, COMBINAZIONI NUOVE E NOMI NUOVI

Oltre alle sinonimie e alle nuove combinazioni segnalate sopra nel corso delle
descrizioni, sono proposte le seguenti modifiche resesi necessarie grazie alla mia
migliore conoscenza attuale della fauna delle Aleocharinae cinesi.

Alevonota ocularis (Pace, 1991), comb. n.
Tomoglossa ocularis Pace, 1991: 835

Alevonota nepalensis (Pace, 1987), comb. n.
Ousipalia nepalensis Pace, 1987: 404

Alevonota basicornis (Pace, 1985), comb. n.

Schistoglossa basicornis Pace, 1985: 105
Tomoglossa basicornis: Pace, 1991: 835

Alevonota venusta (Pace, 1984), comb. n.
Ousipalia venusta Pace, 1984: 318

Alevonota franziana, nom. n., comb. n.
Tomoglossa franzi Pace, 1991: 833
(nec Alevonota franzi (Palm, 1976), Assing, 2002c: 63, olim Sipalia franzi Palm, 1976:
71)
Emmelostiba xinjiangensis Pace, nom. n., comb. n.
Alevonota chinensis Pace, 1993: 110
(nec Emmelostiba chinensis Pace, 1998: 440)
Leucocraspedum Kraatz, 1859: 51
Chariusa Pace, 2003: 166, syn. n.
Irmaria Cameron, 1925: 48
Syntemusa Pace, 2002: 232, syn. n.

Irmaria kinabaluensis (Pace, 2002), comb. n.
Syntemusa kinabaluensis Pace, 2002: 235

Irmaria smetanai (Pace, 2002), comb. n.
Syntemusa smetanai Pace, 2002: 235

Cordalia chinensis Pace, 1993: 82
Cordalia permutata Assing 2002a: 5, syn. n.

Va rilevato che la prima pubblicazione delle spermateca e dell’habitus di Cordalia
vestita (Boheman, 1858) è del 1992 (Pace, 1992: 243). Assing (2002a) non ne fa cenno.

Atlantostiba Pace, 1994: 72, genus bonum
Alevonota Thomson, 1858, sensu Assing, 2002: 63, partim

Imprudentemente Assing ha messo in sinonimia Atlantostiba Pace, 1994, senza un’ac-
curata analisi dei caratteri distintivi tra Alevonota e questo genere, riservandosi di af-
fermare solo «...the former (Atlantostiba) is a junior secondary homonym of the latter

520 R. PACE

(Sipalia franzi Palm)». Non solo, ma non ha pubblicato la forma della spermateca di
Atlantostiba franzi Pace, un fatto eccezionale questo perché Assing pubblica sempre
edeago o spermateca di specie recentemente da me descritte e pure illustrate con la
spermateca ed edeago, oltre che con l’habitus. Pongo qui a confronto i caratteri distin-
tivi tra Alevonota e Atlantostiba. Ogni punto (*) significa presenza.

Atlantostiba Alevonota
Spermateca piegata a Z e
Spermateca piegata a 7 e
Pronoto poco trasverso e
Pronoto ben trasverso °
Protibie setolose .
Protibie spinulose °
Palpi labiali di 3 articoli °
Palpi labiali di 2 articoli °
Edeago con due appendici apicali °
Edeago senza appendici apicali °
Ligula a due lembi accostati e
Ligula a due lembi separati e

Atlantostiba franzi Pace, 1994: 74
Alevonota pacei Assing, 2002c: 63, syn n.

Leptusa anmashanensis Pace, 1996: 30
Leptusa acuta Assing 2002d: 978, syn. n.
L’edeago di L. acuta è pressoché sovrapponibile con esattezza a quello di L. an-
mashanensis Pace.
Melagria Casey, 1906: 230
Euphorbagria Assing, 1997: 64, syn. n.

La spermateca del genere Euphorbagria Assing ha forma tipica delle specie del genere
Melagria Casey, diffuso anche nella regione etiopica.

Atheta (Geostibops) floresensis Pace, nom. n.

Atheta (Geostibops) aptera Israelson, 1985: 13, (nec Atheta (Microdota) aptera Pace,
1984: 320)

RINGRAZIAMENTI

Per avermi affidato in studio il materiale oggetto del presente lavoro rivolgo i
miei più cordiali ringraziamenti ai colleghi Dr. Ales Smetana di Ottawa, Guillaume de
Rougemont di Londra. e Marc Tronquet di Molitg Les Bains (Francia). Per il prestito
di tipi ringrazio il Dr. P. M. Hammond del Museo di Storia Naturale di Londra, il Dr.
Lothar Zerche del D.E.I di Eberswalde e il Dr. Manfred Uhlig del Museo Zoologico
dell’Università Humboldt di Berlino.

HYGRONOMINI E ATHETINI DELLA CINA 521

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in ! te Bu
a dee

REVUE SUISSE DE ZOOLOGIE 111 (3): 525-538; septembre 2004

A preliminary contribution to Platyceps najadum (Eichwald, 1831)
and systematic status of Coluber atayevi Tunijev & Shammakov,
1993 (Reptilia: Squamata: Colubrinae)

Beat SCHATTI
Apartado postal 383, San Miguel de Allende, Gto. 37700, Republica Mexicana.

A preliminary contribution to Platyceps najadum (Eichwald, 1831) and
systematic status of Coluber atayevi Tunijev & Shammakov, 1993. -
External morphological features of Platyceps najadum and the distribution
of this species in Iran and Turkmenistan are analysed. Based on morpho-
logical characters and mtDNA sequence data (12S rDNA), Coluber atayevi
Tunijev & Shammakov, 1993 is ranked as a subspecies of the Slender racer,
P. najadum atayevi comb. n. This taxon seems to have a restricted dis-
tribution in the Kopet Dag.

Keywords: Platyceps najadum ssp. - Coluber atayevi - Iran - Kopet Dag -
morphology - systematics - mtDNA.

INTRODUCTION

Eichwald (1831) described 7yria najadum from the vicinity of Baku in Azer-
bajdzhan. Eichwald’s, or the Slender, racer is distributed from the eastern Adriatic
Region (Dalmatia) to the Caucasus and northeastern Iran. Mertens & Müller (1928)
referred this species to the genus Coluber auct.

Schätti & McCarthy (2001) considered Tyria najadum Eichwald, 1831,
Zamenis dahlii var. collaris Müller, 1878 (see Schätti et al., 2001), and Coluber (s. 1.)
schmidtleri Schätti & McCarthy, 2001 to be closely related to Zamenis rhodorachis
Jan, 1863 and allied taxa (see Schätti, 1993). The najadum group is a distinct evolu-
tionary lineage within the monophyletic racer genus Platyceps Blyth, 1860 (Schätti &
Utiger, 2001).

Tunijev & Shammakov (1993) described Coluber atayevi on the basis of
twenty-three specimens from the vicinity of Saivan in the “Bakharden Region” of
Turkmenistan. They noted “significant morphological divergence” compared with
“other representatives of the najadum-rubriceps complex” and concluded that their
new taxon replaces the Slender racer in the “western and central Kopet-Dag”.
Darevskij & Orlov (1994) questioned the validity of C. atayevi. Schätti & Utiger
(2001) excluded this taxon from their systematic re-arrangement of Old World racer
genera for want of morphological criteria allowing a clear distinction from Platyceps
najadum. For the time being, Atajev’s racer parades under the operational term
Coluber sensu lato.

Manuscript accepted 06.04.2004

526 B. SCHÄTTI

This study investigates external morphological characters of Platyceps najad-
um, with special emphasis on features considered to separate this species from Coluber
(s. 1.) atayevi, and presents molecular (12S rDNA) data.

MATERIAL AND METHODS

The material used for this investigation was obtained from the following insti-
tutions: The Natural History Museum [British Museum (Natural History)], London
(BMNH), California Academy of Sciences, San Francisco (CAS), Field Museum of
Natural History, Chicago (FMNH), Muséum d’histoire naturelle, Genève (MHNG),
Muséum National d’Histoire naturelle, Paris (MNHN), Museo civico di Storia naturale
‘Giacomo Doria’, Genova (MSNG), National Museum, Department of Zoology,
Prague (NMP), Naturhistorisches Museum, Wien (NMW), Forschungsinstitut und
Naturmuseum Senckenberg, Frankfurt on Main (SMF), Zoological Institute, Russian
Academy of Sciences, St. Petersburg (ZISP), Institut für Systematische Zoologie [for-
merly Zoologisches Museum], Museum für Naturkunde, Humboldt-Universität, Berlin
(ZMB), and Zoologische Staatssammlung, München (ZSM). Further acronyms used in
the text are CRS (Caucasian Reserve, Sochi, Krasnodar Region, Russia), MTKD
(Staatliches Museum für Tierkunde, Dresden), and RM (J. Robert Macey, Berkeley:
field tags).

The specimens from Iran and Turkmenistan examined for the purpose of this
study are: BMNH 1920.3.20.3 (Gorgan, Golestan Province, 36°50’N 54°29’E, 9,
Ingoldby); CAS 182948-50 (Turkmenistan: “environs of Saivan Village, Saivan-
Nokhur Plateau, western Kopet-Dag, Bakharden Region”, d, 9 9, paratypes of
Coluber atayevi), CAS 185185-86 and 185188-94 (“elev. 1200-1300 m, 38°30’N
56°47’E, 2 km SE (airline) of Saivan, Ashgabad Region”, 5 4 4, ? ©, paratypes of C.
atayevi), FMNH 141612 (Shahrabad [Shahrabad-e-Khavar], Khorasan, 37°30’N
56°51’E, unsexed [tail completely truncated immediately behind anal scute], Street
Expedition), FMNH 141649 (Chalus, Mazandaran, 36°40’N 51°25’E, juv. d, Street &
Lay 1962); MHNG 1403.92 (Lar [Elburs], Teheran (Central) Province, 35°22’N
49°42’E, 2), MHNG 1403.93 (Polur, Teheran, 35°52’N 52°03’E, 3), MHNG 2626.56
(vic. of Mahniyah, Hamadan/Zanjan border, 35°31’N 49°05’E, 4); MNHN 3582
(“Perse”, 6, Aucher-Eloy); MSNG 30313 (“Isphahan”, dg, Doria); NMP6V 35563
(“50 km NNE Avaj”, Zanjan, approx. 35°34’N 49°13’E [Avaj], juv. 9); NMW 20138
(“Pera, Khoi” [Khoy], West Azarbaijan, ca. 38°33’N 44°58’E, d, Zugmayer); SMF
30095 (“Shahi” [Qa’emshar], Mazandaran, 36°28’N 52°53’E, 9), SMF 67214 (“Sarab
bei Miyandoab”, West Azarbaijan, ca. 36°57’N 46°06’E [Miandowab], 9); ZISP
13626 (Gorgan, juv., Kirichenko 1914 [in bad shape, ventral and subcaudal counts un-
known]), ZISP 18624 (Turkmenistan: Firjuza, 37°56’N 58°04’E, © , Gorbunov 1973);
ZMB 31841 (“Persisch Kurdistan”, 2); ZSM 6.1968 (Gok Shalu, Miandowab, 9),
ZSM 55.1972 (“Tschalus” [Chalus Valley], Mazandaran, ca. 36°38’N 51°38’E, d).

The morphological description of Platyceps najadum is based on data from
roughly 250 specimens. A detailed analysis of this sample will be presented in a forth-
coming study. The dorsal scale formula gives the number of longitudinal rows at the
tenth ventral, at midbody (50% of the total number of ventrals) and in front of the anal
scute. Reductions in the number of dorsal scale rows on the posterior part of the body

PLATYCEPS NAJADUM AND COLUBER ATAYEVI 527

are situated either ‘high’ (paravertebral or vertebral) or ‘low’ (lateral). Their position
along the trunk is expressed in terms of ventrals and as a percentage of the total number
thereof (%ven); these values are based on the mean of the right and left side counts.
The tail / body ratio is the tail length divided by the snout-vent length, i.e., from the tip
of the snout to the posterior border of the anal scute.

For this paper, a partial sequence of the mitochondrial small ribosomal subunit
(12S rRNA) was obtained from tissue samples of the following specimens: C. atayevi
Tunijev & Shammakov CAS 185188 (RM 9545, 99% ethanol, liver), Platyceps kare-
lini (Brandt) MHNG 2443.3 (frozen muscle), P. najadum (Eichwald) MHNG 2626.56
(ibid.), and P. variabilis (Boulenger) MHNG 2456.71 (ibid.). The GenBank numbers
are AY647231-34. The technique of DNA extraction as well as PCR and sequencing
procedures are described in Utiger er al. (2002). The obtained data were added to those
of Hemorrhois hippocrepis (L.) and seven Platyceps spp. (including two P. najadum)
from an existing sequence file consisting of two partially analysed mitochondrial
genes, cytochrome oxidase subunit I (COD and 12S rRNA (Schätti & Utiger, 2001).
For the new samples, only the latter gene region was investigated. The lacking COI
characters were coded as missing.

TABLE 1. Sequence properties and tree reconstruction parameters.

gaps: missing gaps: 5th character state
unweighted MP unweighted MP weighted MP

Lenght of sequence alignment (COI/12S) 1116 (513/603) 1116 (513/603) 1116 (513/603)

Total of variable characters (COI/12S) 308 (150/158) 317 (150/167) 317 (150/167)
- parsimony-informative (COV12S) 191 (108/83) 197 (108/89) 197 (108/89)
Number of most parsimonious trees 1 1 1
Tree length 592 622 258.8
Rescaled consistency index (RC) 0.32 0.316 0.708

All phylogenetic analyses were performed with PAUP* version 4.0b10 for Mac
(Swofford, 1998), and the weighted maximum parsimony (MP) analysis is described
in Utiger et al. (2002) and Utiger & Schätti (2004). Gaps in the 12S rDNA sequence
were treated as fifth character state. After a first run with heuristic search and tree-bi-
section reconnection (TBR) branch swapping, characters were weighted with the
rescaled consistency index (RC, Farris, 1989) and a second heuristic search was per-
formed. Nonparametric bootstrap values (Felsenstein, 1985) with 1000 replicates were
calculated for both character types (see Discussion).

RESULTS

MORPHOLOGY

Platyceps najadum has eight or nine supralabials, the fourth and fifth or fifth
and sixth entering the eye; with the exception of MNHN 3582 from “Perse” (Duméril
et al., 1854), all Iranian specimens examined as well as an individual from
Turkmenistan (ZISP 18624) have eight supralabials. The preocular and anterior sub-
ocular are single; posterior subocular scales are absent. The number, size and arrange-

528 B. SCHÄTTI

ment of the temporals is highly variable; usually, there are two anterior and three
posterior scales; in most cases, the lower anterior temporal is larger than the upper.
Fragmentation of the temporals is noted in the type series of Coluber najadum
albitemporalis Darevskij & Orlov, 1994.

Usually, there are ten (nine to eleven) sublabials, the anterior four or five in
contact with the first chin shield (inframaxillary), which is shorter and slightly broader
than the posterior. Normally, the posterior pair of chin shields is separated anteriorly
by one or two granular scales and two or three rows caudally. However, the size and
number of the scales separating the posterior chin shields is highly variable; in some
specimens, the latter are in contact for almost their whole length as, for instance, in
ZSM 55.1972.

I
jee

are

cK
=

Fic. 1
Ventral head scales of Platyceps n. najadum ZSM 55.1972 (left) and ZSM 48.1918 (right).

Platyceps najadum has 205-236 (4 4 205-225, 9 © 208-236) ventrals and 100-
140 (6 & 100-140, 2 © 103-137) paired subcaudals; the anal scute is paired. There are
207-236 (3 d 207-222, 2 2 219-236) ventrals and 107-133 (6 d 109-132, 2 2 107-
133) subcaudals in specimens from Iran.

Generally, the highest ventral counts are recorded in eastern Anatolia and
adjacent regions (including SMF 67214). The minimum for subcaudals (100) in
specimens with complete tails was registered in ZISP 14092 from Sochi (Krasnodar).
Strauch (1873) and Nikolskij (1916) noted 98 subcaudals in the “Kaukasus” (no. 1741,
“Dr. Höft, 1844”) and 99 for a specimen from the Transcaspian Region. Considerably
lower subcaudal counts reported, for instance, by Latifi (1991) and Darevskij &
Szczerbak (1993) are probably from individuals with incomplete tails.

PLATYCEPS NAJADUM AND COLUBER ATAYEVI 529

Platyceps najadum normally has 19-19-15 or 19-19-13 dorsal scale rows. Both
patterns are found in specimens from the southern border of the Caspian Sea to northern
Khorasan. The reduction pattern in two females (MHNG 1403.92, SMF 30095) is
‘high-low’ or ‘low-high’ at 61-89%ven; ZSM 55.1972, a male with 13 dorsal scale rows
prior to the anal scute, has the reductions ‘low-high-high’ at 61-62% (first and second)
and 85%ven (third); FMNH 141612 (unsexed, see Material and Methods) shows the
sequence ‘low-high’ at 61-64%ven; ZISP 18624 (2) from the Kopet Dag (Firjuza) has
two reductions involving low (first) and high (second) levels at 64-65%ven.

The maximum total length reported for Platyceps najadum is approx. 140 cm
(Cattaneo, 2001). The longest male examined, i.e., MHNG 1358.72 from Zadar
(Dalmatia), attains 920 + 395 mm. The maximum snout-vent length in females is
913 mm in MHNG 2447.68 from Mt. Ararat, Turkey (tail incomplete). The body is
slender and the tail / body length ratio in adults at least 0.32 (2 2) to 0.34 (dd); in
Anatolia some specimens, e.g., CAS 105234 (9) from the vicinity of Anamur, attain
values of 0.50.

Head, body, and tail are generally olive, light brownish, or faint grey. The
supralabials, preocular, and postoculars are white or yellowish. Often, there is a dark
spot below the eye or an oblique stripe running from the lower border of the orbit to
the temporal region. The lateral portion of the neck and forebody shows conspicuous
white-edged ocelli or blotches with dark (black to dark olive) centres; these markings
may be replaced by, or mixed with, narrow bars on the anterior flanks. Size and dis-
tinctiveness of the posterior lateral markings reduce progressively, and the light edges
of the ocelli and blotches are absent. The anterior lateral markings may be confluent
mid-dorsally forming transverse bands. The chin, venter, and underside of the tail are
uniform light (white, cream, or yellowish). Mertens (1940) noted a brick red (“ziegel-
rot”) hue of the posterior subcaudals in SMF 30095.

Populations with two large and conspicuous milky dots on the nape occur in the
vicinity of Lenkoran and Archivan, southern Azerbajdzhan (Coluber najadum
albitemporalis Darevskij & Orlov).

Specimens having one or several cross-bands on the nape and neck are recorded
from Anatolia, Syria, and Iraq to the Caucasus and Turkmenistan (e.g., BMNH
1920.3.20.3, MHNG 1403.92, NMP6V 35563, ZISP 18624, and ZSM 6.1968). In the
eastern part of the distribution range occur individuals with a dorsal colour pattern
composed of fine black spots behind the neck that may extend onto midbody as exem-
plified by NMW 20138, SMF 30095, and ZSM 55.1972. Specimens virtually devoid
of any dorsal markings are known from various areas and seem to be common in
central western Iran (e.g., MHNG 2626.56, MSNG 30313).

DISTRIBUTION

Platyceps najadum lives from Dalmatia across the southern Balkans, the Ionian
and Aegean islands including Lefkada [Levkäs] and Lipsos (new records), Anatolia,
the Levant south to Lebanon and the Syrian lava desert, Iraq, the Caucasus north to the
Krasnodar Region and Dagestan, Azerbajdzhan, northern Iran, and in the Kopet Dag,
Turkmenistan (e.g., Strauch, 1873; Baran, 1976; Bannikov er al., 1977; Darevskij &
Szczerbak, 1993).

530 B. SCHÄTTI

Based on the catalogue entries of the St. Petersburg herpetological collection, Ananjeva
and Orlov (1977) concluded that a juvenile specimen (ZISP 11151) collected in 1910 by K.
A. Satunin at “Derbel in Caucaso” (Nikolskij, 1916: 93) came from Dzhebel (39°38’N 54°14’E)
in the Bolshoj Balkhan area, Turkmenistan. Probably following these authors, Szczerbak &
Golubev (1981) mentioned this individual from the Transcaspian Region. However, ZISP 11151
is from Derbent (42°03’N 48°18’E) in Dagestan (Nikolskij, 1916: 94).

In Iran, Platyceps najadum is reported from large parts of the western mountain
region (e.g., MHNG 2626.56, NMP6V 35563) south to Esfahan Province (Blanford,
1876: MSNG 30313), and the Caspian Region (e.g., SMF 30095, Qa’emshar).
Northeastern records and collecting sites include BMNH 1920.3.20.3 and ZISP 13626
from Gorgan (Baran, 1976; Ananjeva & Orlov, 1977), FMNH 141612 (Shahrabad, N
Khorasan), Emamrud (“Shakhrud”, 36°25’N 54°58’E, ca. 1200-1500 m a.s.l.) in
Semnan Province, “Kent-Parsian” between 36°22’-37°15’N and 55°25’-56°45’E (fide
Adamec, 1981) as well as Darreh Gaz and Quchan (Fig. 2) in northern Khorasan
(Kessler, 1872; Derjugin, 1905; Latifi, 1991). The southern distribution limit is not
well documented (see below).

Apart from an unspecified record from the Transcaspian Region (leg. Matvejev
1904) quoted by Czarevskij (1915), Platyceps najadum is recorded from Mt. Khozly
and Kara Kala settlement to as far east as the Gaudan Valley (Ashkhabad - Quchan
road) separating the central and eastern Kopet Dag (Fig. 2), between ca. 400 m (Kara
Kala) to 27000 m on Mt. Dushak (Varenczov, 1894; Ananjeva & Orlov, 1977;
Rustamov & Shammakov, 1979; Szczerbak & Golubev, 1981; Szczerbak et al., 1986;
Starkov, 1988; Atajev & Shammakov, 1990; Atajev ef al., 1991, 1994). The map
(Fig. 2) does not include three records from Turkmenistan, i.e., a specimen collected
by Bilkevich near “Ashkhabad” (Nikolskij, 1905), probably a vague origin, and two
unlocated places, “Garrygala” and “Imarat” (Atajev et al., 1991, 1994).

Besides Derjugin’s (1905) record from “Kent-Parsian” in the border area of
Semnan and Khorasan Provinces (see above), various specimens mentioned from the
Irano-Turkmenian border region are presumably lost (see Ananjeva & Orlov, 1977).
This is, for instance, the case with an individual obtained at “Ashkhabad” (37°57’N
58°23’E, see above) or Varenczov’s (1894) material (determined by N. M. Kulagin,
fide Nikolskij, 1905) collected near km 20 along the road from Ashkhabad to the
Gaudan pass (37°37’N 58°24’E) and at “Suluklju”. New observations from 1400-1500
m near Suljukli in Germob District (Sczcerbak et al., 1986: “Germab”, see Fig. 2) were
reported by Szczerbak & Golubev (1981).

Tunijev & Shammakov (1993) did not cite, for instance, the occurrence of the
Slender racer near the Arvaz pass at above 1800 m a.s.l. (active at 8°C) and in the
vicinity of Kénekesir (Kojnja-Kasyr, “Kojne-Kesyr”) reported by Atajev &
Shammakov (1990) and Starkov (1988), respectively (Fig. 2). Atajev er al. (1994)
reported the species from “Garrygala” west of Mt. Dushak (see above). Further pu-
blications dealing with the herpetofauna of Turkmenistan such as, for instance,
Bobrinski (1923), Bogdanov (1962), Shammakov (1971), Rustamov & Shammakov
(1982), or Shammakov er al. (1993) do not procure any pertinent information about
this species, and Atajev (1985: map 86) gave but four localities based on previous
records (i.e., Mt. Dushak, Firjuza, Suljukli, and “Kuchanskoje shosse”). The occur-
rence of Platyceps najadum in the vicinity of Firjuza as first reported by Ananjeva &
Orlov (1977) was confirmed by observations in May 1979 (Szczerbak et al., 1986).

PLATYCEPS NAJADUM AND COLUBER ATAYEVI 531

TURKMENISTAN

FIG. 2

Distribution of Platyceps najadum in Turkmenistan (Kopet Dag) and North Khorasan, Iran.
Localities in the Kopet Dag are Firjuza (ZISP 18624, Ananjeva & Orlov, 1977) at 37°56°N
58°04’E as well as Arvaz (38°14’N 57°09’E, pass, approx. 1’840 m a.s.1.), Dagish and Dashtoj
gorge (Babazo District: B), Mt. Dushak (ca. 37°58’N 57°54’E), vicinity of Germob (ca. 38°01’N
57°44’E), Kara Kala (38°34’N 56°44’E, 400 m), Karanki gorge (Babazo District: B, 1°200-
1°400 m), 15 km NE Kénekesir (“Kojne-Kesyr”, ca. 38°17’N 57°00’E), Mt. Khozly (ca.
38°35’N 56°48’E, “about 18 km south of Bami”), Mirza Dag (38°01’N 57°36’E), the vicinity of
Saivan (38°30’N 56°47’E, type locality of Coluber atayevi), Suljukli (Germob District: G), and
km 20 on the Ashkhabad - Quchan road (Varenczov, 1894; Rustamov & Shammakov, 1979;
Szczerbak & Golubev, 1981; Szczerbak et al., 1986; Starkov, 1988; Atajev & Shammakov, 1990;
Atajev et al., 1991; Tunijev & Shammakov, 1993). Babazo and Germob are border districts with
Iran (Sczcerbak, 1986: map). Iranian localities include Shahrabad (37°30’N 56°51’E, FMNH
141612) and two collecting sites from Latifi (1991), i.e., Darreh Gaz (“Darehghaz”, 37°27’N
59°07’E) and Quchan (“Ghoochan”, 37°06’N 58°30’E).

Latifi (1991) mentioned Platyceps najadum from Kerman, Baft (29°12°N
56°36’E) and Khash (28°14’N 61°15’E) in southeastern Iran. At least two of these
indications are possibly based on records (ZISP 9286, 9288) of “Zamenis dahlii Fitz.”
from “Kaskin in Kirmano orient.” (ca. 27°30’N 60°22’E, fide Anderson, 1999) and

532 B. SCHÄTTI

“Kerat in Chascht-Adano” (Nikolskij, 1899). The series (ZISP 9285-88) belongs to
P. rhodorachis (Jan) as noted by Ananjeva & Orlov (1977). The illustration of P.
najadum in Nikolskij (1916: pl. 1.2) or, for instance, Terentjev & Chernov (1936: pl.
IV, same specimen) is not based on ZISP 9286 as indicated.

Latifi (1991) also reported Platyceps najadum from Fars and Khuzestan
(Ahvaz) in southwestern Iran. These references as well as a specimen from
Boyerahmad-va-Kogiluye (Yasuj, approx. 30°40’N 51°36’E) collected by Frynta et al.
(1997) are in need of confirmation and re-identification, respectively. These records
require comparison with P. schmidtleri (Schätti & McCarthy, 2001).

DISCUSSION

Although Tunijev & Shammakov (1993) had, apart from the type series of
Coluber atayevi, no further specimens from Turkmenistan at hand, they assigned all
populations of Platyceps najadum auct. of the “western and central Kopet-Dag, from
the surroundings of the Kara-Kala settlement in the west to the Sulukli Spring in the
east” (see Distribution) to this taxon. According to these authors, C. atayevi differs
from P. najadum (Eichwald) in cranial pholidosis (supralabials, chin shields), head and
body shape including cross-section of the trunk, development of the ventral edges,
lengths and proportions, and in dorsal colour pattern.

As far as head scales are concerned, Tunijev & Shammakov (1993: fig. 3) stated
that the supralabials of Coluber atayevi are “weakly distinguished from the throat [sic]
scales, whereas all upper labials of C. najadum are strongly pronounced”. Furthermore,
the posterior “genial scutes” (chin shields) “contact one another along the mental
groove, rarely having a few isolated granules between the posterior pair” in C. atayevi
but are “widely separated” by “always 2-4 rows of well-developed scales” in Platyceps
najadum. The illustrations in Tunijev & Shammakov (1993) simply show that there is
no difference between the two taxa in the size of the supralabials. The posterior chin
shields are completely separated, for instance, in CAS 185186, 185189, and 185191
(paratypes of C. atayevi), and there is considerable variation in the arrangement, num-
ber, and development of the median granular scales in P. najadum (Fig. 1).

With regard to head shape, Atajev’s racer has been diagnosed as having a “nar-
row, sharp, and flat head with the rostrum beveled downward”. The drawings of
Platyceps najadum in Tunijev & Shammakov (1993: figs 3.a-c) show a bloated head
which does by no means represent the contour in living specimens. Differences in gen-
eral body shape and habitus between the two taxa as insinuated by Tunijev &
Shammakov (1993: fig. 2) cannot be confirmed either.

Tunijev & Shammakov (1993) described the lateral edges of the ventrals (“ab-
dominal ridges”) in Coluber (s. 1.) atayevi as “practically absent” (“in contrast to”
Platyceps najadum) and the cross-section of the body round, whereas it is thought to
be “rectangular [...] in C. najadum and C. rubriceps” (i.e., P. collaris, see Schatti et al.,

1) ZISP 9285-87 from “Kirmano orient.” and “in Sargado” were collected in Sistan-va-
Baluchistan. ZISP 9288 was obtained on 13 April 1898 in the vicinity of Herat (34°34’N
60°33°E), eastern Khorasan (Zarudnyj, 1898; Nikolskij, 1905; Anderson, 1999).

PLATYCEPS NAJADUM AND COLUBER ATAYEVI 533

2001). In fact, these features are variable in P. najadum and, in preserved specimens,
finally depend on factors such as killing and preservative agent and state of the
specimen.

The maximum snout-vent length of 533 mm indicated for Atajev’s racer
(Tunijev & Shammakov, 1993: CRS 421) is exceeded, for instance, in CAS 185185
with almost 60 cm. Differences in the relative tail length (“1/4 of total length” vs. “1/3”
in najadum) are correlated with the number of subcaudals (see below). The tail/body
ratio in Coluber atayevi is 0.32-0.36 (3 3 0.34-0.36, 2 2 0.32-34). Body length for the
examined northeast Iran Platyceps najadum (all females) ranges from ca. 25 to 84 cm.
FMNH 141612 (sex unknown) has a snout-vent length of ca. 64 cm, ZISP 18624 from
Turkmenistan ca. 32,5 + 13 cm. The tail/body length ratio in these specimens is 0.31
(BMNH 1920.3.20.3) to 0.39.

The examined paratypes of Coluber atayevi have a dark spot below the eye and
the nape and neck are ocellated. Usually, there are two to five ocelli (and transverse
bars in CAS 185185) followed by up to five longitudinal rows of small dark dots
extending to midbody or beyond (e.g., CAS 185185-86, 185189, 185191); posteriorly,
they are reduced to small points situated mid-dorsally. Tunijev & Shammakov (1993)
emphasise that “brown colors prevail instead of olive-green ones” as in Platyceps
najadum. However, the coloration of the Slender racer includes olive, brownish, or
grey hues, and there is no character of dorsal colour pattern allowing a distinction of
these taxa.

To sum up, “significant morphological divergence” between Atajev’s racer
(Saivan population) and Platyceps najadum as stated by Tunijev & Shammakov (1993)
does not correspond to the facts. However, qualitative differences vis-a-vis Slender
racers from more western regions exist in the degree of separation of the posterior chin
shields, relative tail length, and peculiarities of the temporal scales as well as low ven-
tral and subcaudal counts that are, strangely enough, not notified by Tunijev &
Shammakov (1993).

In at least nine paratypes of Coluber atayevi, viz., CAS 182948-50, 185185
(right side), 185186, 185190-91, 185193, and 185194 (left), the upper first temporal is
vertically divided. No information regarding the remaining type material (CRS series)
is available (Tunijev & Shammakov, 1993: Tbs 1-2). In any case, this configuration is
uncommon in Platyceps najadum (e.g., MHNG 2626.56).

The Saivan specimens (CAS series) have 203-224 ventrals (4 4 203-209, 2 2
214-224) and 94-104 subcaudals (dd 94-104, £ £ 97-101) compared to 205-225
(dd) and 208-236 (9 ?) ventrals and 100-140 (3 &) and 103-137 (2 2) subcaudals
in Platyceps najadum (see Morphology). Tunijev & Shammakov (1993: Tb. 2) gave
197-214 ventrals and 90-106 subcaudals for the type series of Coluber atayevi (sexes
not separated)2). In P. najadum from Iran examined for the purpose of this study there
are 207-236 (4 d 207-222, 2 2 219-236) ventrals and 107-133 (Sd 109-132, 29
107-133) subcaudals. ZISP 18624 (2) from Turkmenistan has 222 ventrals and 125
subcaudals (not 112 as erroneously stated by Ananjeva & Orlov, 1977).

2) According to Tunijev & Shammakov (1993: Tb. 1), a paratype deposited in the Caucasian
Reserve, Sochi (CRS 424) has only 190 ventrals and 90 subcaudals.

534 B. SCHÄTTI

The sum of the ventral and subcaudal counts for the Saivan series (examined
specimens) ranges from 299-322 (dd 299-313, 2? 314-322). These values are
usually considerably higher in Platyceps najadum. Only in the case of three males from
the Krasnodar Region (ZISP 3695: 316, ZISP 14092: 309) and the Ararat area
(NMP6V 35564.1: 314) they are below 320 (ZSM 55.1972 has 321). Females normal-
ly have values distinctly over 330 and at least 324-325 in the Ararat area, Georgia and
Dagestan (MTKD 14274, NMP6V 35564.2, and ZISP 11151); BMNH 1920.3.20.3
from Gorgan (Golestan) has 329 (222 ventrals, 107 subcaudals). Varenczov’s (1894)
presumably lost specimen from the Gaudan road (see Distribution), probably a female,
had 225 ventrals and 101 subcaudals (tail incomplete?).

The description of Atajev’s racer is based on a comparison of the type series in
the CRS collection (holotype and nine paratypes) collected around Saivan with a
limited number (ten specimens) of Platyceps najadum “from various areas of the
Caucasian Isthmus” and information from literature including questionable data on
P. collaris Müller [as Coluber rubriceps (Venzmer)].

Except for the type series of Coluber atayevi, no morphological data are
reported for populations from the Bakharden region (i.e., Kara Kala, Mt. Khozly). This
is also true for specimens from collecting sites between Kénekesir and Mt. Dushak
along the border region with Iran (Fig. 2). By all means, the subadult female from
Firjuza (ZISP 18624) with 347 ventrals and subcaudals (see above), entire (unfrag-
mented) anterior temporals, a low (first) and high posterior reduction at 64-65 %ven,
and a dorsal colour pattern composed of a collar and distinct bars along the neck
conforms to typical Platyceps najadum auct.

The configuration of the chin shields and temporals (see above) as well as
particularly low ventral and subcaudal counts in the Saivan population are probably
due to a genetic bottleneck?). Molecular data reveal divergence of the Saivan
population vis-à-vis Platyceps najadum auct. (Fig. 3). However, the genetic distance is
smaller than between typcial specimens of the Slender racer examined for the purpose
of this study and MHNG 2626.56 from the northern Zagros range.

The reader may argue that COI and 12S rDNA data were used for the tree
reconstruction (Fig. 3) although a number of COI sequences are missing. However,
conspecific samples with a complete data set are available, giving the tree some
significance. Unweighted and weighted MP analyses of both gene fragments resulted
in the same topology. Bootstrap values with 5000 replicates are virtually identical with
those presented in the phylogeny (Fig. 3). Uncorrected genetic distances within the
Platyceps najadum group are 5.2% (in prep.), 8.6% for P. collaris vs. P. najadum, and
8.5-12.3% for Hemorrhois hippocrepis vs. the remaining species analysed within the
scope of this study.

Based on morphological and molecular data, Coluber atayevi Tunijev &
Shammakov is considered a subspecies of Plaryceps najadum Eichwald with a limited

3) “Increasing climatic aridization in the Holocene can be the cause of disappearance of C.[olu-
ber] atayevi in the foothills and severe restriction of its area in the middle-altitude and high-
altitude parts of the western and central Kopet-Dag and, possibly, to the breakup of the area
into several local refuges. The “primitive” morphological features were preserved in the
absence of contacts with closely related forms” (Tunijev & Shammakov, 1993: 8).

PLATYCEPS NAJADUM AND COLUBER ATAYEVI 535

Platyceps najadum atayevi*

Platyceps najadum ssp.
Platyceps najadum ssp.
Platyceps najadum ssp.*
Platyceps collaris

Platyceps ventromaculatus
Platyceps elegantissimus
Platyceps karelini x

Platyceps cf. rhodorachis
Platyceps rogersi

Platyceps florulentus

Platyceps variabilis*

Hemorrhois hippocrepis
—— 10 changes

FIG. 3

Weighted maximum parsimony (MP) tree of two partial gene sequences, COI and 12S rDNA
(only 12S in taxa with an asterisk) and the Palaearctic horseshoe snake Hemorrhois hippocrepis
(L.) as out-group. Bootstrap values (1000 replicates) from weighted (upper) and unweighted
(lower values) MP analysis.

geographic distribution. Supposedly, P. najadum as understood in this paper is made up
of several distinct taxa including incipient species, and this systematic complex is in
need of a more detailed investigation (in prep.).

ACKNOWLEDGEMENTS

The author is grateful to the following persons for the loan of specimens from
the collections in their respective care: Natalja Ananjeva and Ilja S. Darevskij (St.
Petersburg), E. Nicolas Arnold and Colin McCarthy (London), Lilia Capocaccia and
Roberto Poggi (Genoa), Robert C. Drewes and Jens V. Vindum (San Francisco), Ulrich
Gruber (Munich), Rainer Giinther (Berlin), Ivan Ineich (Paris), Konrad Klemmer and
Günther Köhler (Frankfurt on Main), Jiri Moravec (Prague), Franz Tiedemann
(Vienna), and Harold C. Voris (Chicago). Vincent Ziswiler and Paul Ward provided

536 B. SCHÄTTI

laboratory facilities, and Urs Utiger (all Zurich) performed the molecular analyses. I
express a most special thank you to Ted Papenfuss and Robert Macey (Berkeley) for
the tissue sample of a paratype of Coluber atayevi. Corinne Charvet drew the
illustration of the chin region and the distribution map. Alan Resetar (Chicago) helped
to clarify collecting sites of the Street expedition to Iran.

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REVUE SUISSE DE ZOOLOGIE 111 (3): 539-550; septembre 2004

New species of coelotine spiders (Araneae, Amaurobiidae)
from northern Thailand II

Pakawin DANKITTIPAKUL! & Xin-Ping WANG?

! Terrestrial Arthropod Research Unit, Department of Biology, Faculty of Science,
Chiang Mai University, Chiang Mai 50200, Thailand.

1 The University of Auckland, Private Bag 92019, Auckland, New Zealand.

2 Research Associate, Department of Entomology, California Academy of Sciences,
San Francisco, CA 94118, USA.

2 Coordinator of Statistical Research, Brooks Center for Rehabilitation Studies, Health
Science Center, University of Florida, P.O. Box 100185, Gainesville, FL 32610, USA.

New species of coelotine spiders (Araneae, Amaurobiidae) from
northern Thailand II. - Four new coelotine spiders are described from the
provinces of Chiang Mai and Lamphun in northern Thailand. The species
Draconarius lateralis sp. n., D. paralateralis sp. n. and D. pseudolateralis
sp. n. belong to a species group here called the /ateralis group. Draconarius
elatus sp. n. is unique among coelotines in its peculiar male palp and is
placed in a group of its own, the elatus group. Comments on the distribution
and diversity of coelotines in northern Thailand are given.

Keywords: Araneae - Amaurobiidae - Coelotinae - Draconarius - new
species - taxonomy - zoogeography - Thailand.

INTRODUCTION

This paper is the second contribution to the knowledge of the spider family
Amaurobiidae of Thailand. In our previous publication on the subfamily Coelotinae,
which gives the first record of amaurobiids from Thailand, six new species have been
described (Dankittipakul & Wang, 2003). These include five Draconarius species from
Chiang Mai Province and a Coelotes species from Chiang Rai Province. In the present
paper we describe four additional new species from the provinces of Chiang Mai and
Lamphun. These species do not fit well into any coelotine genus and are placed only
tentatively in Draconarius at this moment. Three of them, D. lateralis sp. n., D. para-
lateralis sp. n. and D. pseudolateralis sp. n., belong to a distinct lineage, here called
the /ateralis group, and can be recognized by laterally expanded spermathecae and
bifurcate conductor apices. The fourth species, D. elatus sp. n., belongs to a group of
its own, here called the elatus group, which differs from all other coelotines by its
small size, short and elevated RTA, short cymbial furrow, reduced tegular sclerite, and
large embolic base. The species groups established here are entirely phenetic and a
phylogenetic analysis, including the species groups assigned by Wang (2002) and
(2003), is required.

Manuscript accepted 25.02.2004

540 P. DANKITTIPAKUL & X.-P. WANG

MATERIAL AND METHODS

All illustrations were made with an Olympus SZX-9 stereomicroscope
equipped with a drawing tube. Body measurements are in millimetres. Measurements
of leg segments were taken from the dorsal side. Epigyna were drawn in natural and
cleared state (immersing in lactic acid for 20-30 minutes). Male palps were drawn in
retrolateral and ventral view. The specimens examined are deposited in the collections
of the Museum d’histoire naturelle, Geneve (MHNG), in the Department of Ento-
mology at the California Academy of Sciences (CAS), and in Pakawin Dankittipakul’s
collection (PDC) which will also be deposited in the MHNG.

Abbreviations used in the text and in the figures: ALE, anterior lateral eyes;
AME, anterior median eyes; C, conductor; CD, copulatory duct; CL, conductor
lamella; DC, dorsal apophysis of conductor (= conductor dorsal apophysis according
to Wang, 2002); E, embolus; LE, lateral eyes; ME, median eyes; MOQ, median ocular
quadrangle; PA, patellar apophysis; PLE, posterior lateral eyes; PME, posterior median
eyes; RDTA, retrolateral dorsal tibial apophysis (= lateral tibial apophysis according to
Wang, 2002); RTA, retrolateral tibial apophysis; SH, spermathecal head.

TAXONOMY
Draconarius Ovtchinnikov, 1999

The lateralis group

Females can be recognized by the absence of epigynal teeth and the presence of
laterally extended spermathecae; males by the absence of a median apophysis and the
presence of a bifurcate conductor apex. The /ateralis group species are similar to D.
penicillatus (Wang et al., 1990) by having laterally extended spermathecae but differ
by the absence of epigynal teeth in females and by the absence of a median apophysis
and the presence of a bifurcate conductor apex in males.

Draconarius lateralis sp. n. Figs 1-4

Type locality: THAILAND, Chiang Mai Province, Chiang Dao District, Doi Chiang Dao
Wildlife Sanctuary, Pha Taeng at the foot of Doi Chiang Dao, 500 m.

Type material: Holotype: Ö , from the type locality, evergreen forest along stream, pitfall
trap, 23.xi.-22.xii.1990 (MHNG). Paratypes: 146 , 19, same data as for holotype (MHNG); 34,
pitfall trap, 25.x.-23.x1.1990 (MHNG); 66, pitfall trap, 22.x11.1990-15.1.1991 (CAS, PDC AM-
0451); 39, pitfall trap, 15.i.-23.11.1991 (MHNG, CAS, PDC AM-0452 ). All specimens leg. P. J.
Schwendinger.

Etymology: The specific name refers to the laterally extended spermathecae.

Diagnosis: Males can be distinguished from those of D. paralateralis sp. n. and
D. pseudolateralis sp. n. by the long patellar apophysis of the palp and the broad
conductor apex bifurcation (Figs 1, 2); females by the anteriorly situated atrium
(Fig. 3).

Description: è (holotype). Total length 9.64. Carapace 4.54 long, 3.21 wide.
Abdomen 4.62 long.

COELOTINE SPIDERS FROM NORTHERN THAILAND II 541

Fics 1-4

Draconarius lateralis sp. n., 4 holotype (1, 2) and 2 paratype (3, 4). Left palp, ventral (1) and
retrolateral (2) view. Epigynum, ventral view (3). Vulva, dorsal view (4). Scale lines 0.5 mm (1,
2), 0.25 mm (3, 4).

Eye sizes and interdistances: AME 0.18, ALE 0.18, PME 0.15, PLE 0.15; AME-
AME 0.10, AME-ALE 0.10, PME-PME 0.18, PME-PLE 0.21, ALE-PLE 0.10; MOQ
0.43 long, anterior width 0.40, posterior width 0.43. Clypeus height 0.17. Promargin of
chelicerae with 5 teeth, retromargin with 5.

542 P. DANKITTIPAKUL & X.-P. WANG

Leg measurements:

I II III IV
Femur 4.33 3.95 3.50 4.16
Patella + Tibia 4.21 4.51 3 4.96
Metatarsus 4.34 SB 3.61 4.35
Tarsus 351 2.92 2.52 21/8)
Total 16.19 SA 13.34 16.20

Palp (Figs 1, 2): Patellar apophysis (PA) fairly long, with pointed apex; retro-
lateral tibial apophysis (RTA) occupying more than half of tibial length; retrolateral
dorsal tibial apophysis (RDTA) short, triangular, close to RTA; cymbial furrow 3/4 of
cymbial length; basal lamella of conductor (CL) large; conductor (C) broad, with broad
apex bifurcation; dorsal apophysis of conductor (DC) small; median apophysis absent;
embolus (E) long, slender, originating posteriorly.

? (paratype). Total length 10.91. Carapace 4.65 long, 3.13 wide. Abdomen 6.01
long.

Eye sizes and interdistances: AME 0.15, ALE 0.25, PME 0.25, PLE 0.20; AME-
AME 0.13, AME-ALE 0.13, PME-PME 0.10, PME-PLE 0.25, ALE-PLE 0.13; MOQ
0.56 long, anterior width 0.46, posterior width 0.58. Clypeus height 0.13. Promargin of
chelicerae with 5 teeth, retromargin with 5.

Leg measurements:

I II III IV
Femur 3.61 3:35 2.80 3.86
Patella + Tibia 4.53 4.01 3.04 4.53
Metatarsus 3.30 2.60 DES 3.76
Tarsus 2.01 1.59 1.39 1.62
Total 13.45 1155 9.74 1387,

Epigynum and vulva (Figs 3, 4): No epigynal teeth present; atrium situated
quite far anteriorly, with widely recurved anterior ridge, internal structure visible
through thin tegument; copulatory ducts (CD) broad, situated anteriorly, strongly scle-
rotized; spermathecal heads (SH) located posteriorly; spermathecae laterally extended
and convoluted.

Distribution and habitat: Known only from the type locality in one of the rare
evergreen low land forest of northern Thailand.

Draconarius paralateralis sp. n. Figs 5-10

Type locality: THAILAND, Chiang Mai Province and District, Doi Suthep-Pui National
Park, Doi Suthep, forest above Pin Pak Pai Waterfall and Tham (cave) Ryssie, 1180 m.

Type material: Holotype: ¢, from the type locality, pitfall trap, 4.x1.-6.xii.1987 (MH-
NG). Paratypes: from the type locality, 49, pitfall trap, 1.-30.1.1987 (MHNG, CAS, PDC AM-
0453); 19, pitfall trap, 18.1.-19.11.1986 (MHNG); 2°, pitfall trap, 30.111.-28.1v.1987 (MHNG,
CAS); 16, pitfall trap, 4.1.-7.11.1988 (CAS); Doi Pui, 1580 m, 24 , leaf litter sample, 26.x.2000
(MHNG TH00/02). All specimens leg. P. J. Schwendinger.

Etymology: The specific name derives from the similarity between D. paralateralis sp.
n. and D. lateralis sp. n.; both possess laterally extended spermathecae.

COELOTINE SPIDERS FROM NORTHERN THAILAND II 543

Fics 5-10

Draconarius paralateralis sp. n., 3 holotype (5, 6), d paratype (7, 8) and © paratype (9, 10).
Left palp, ventral (5) and retrolateral (6) view. Conductor (7). Patellar and tibial apophyses (8).
Epigynum, ventral view (9). Vulva, dorsal view (10). Scale lines 0.5 mm (5-8), 0.25 mm (9, 10).

Diagnosis: Males can be distinguished from those of D. lateralis sp. n. and D.
pseudolateralis sp. n. by their short conductor apex bifurcation (Fig. 7) and by a blunt
patellar apophysis on the palp (Figs 6, 8); females by the presence of a triangular ridge
on the epigynum (Fig. 9), by relatively small copulatory ducts, and by large lateral ex-
tensions of their spermathecae (Fig. 10).

Description: 3 (holotype). Total length 10.05. Carapace 4.72 long, 3.21 wide.
Abdomen 5.12 long.

Eye sizes and interdistances: AME 0.18, ALE 0.23, PME 0.19, PLE 0.20; AME-
AME 0.11, AME-ALE 0.76, PME-PME 0.20, PME-PLE 0.25, ALE-PLE 0.10; MOQ

544 P. DANKITTIPAKUL & X.-P. WANG

0.55 long, anterior width 0.50, posterior width 0.54. Clypeus height 0.24. Promargin of
chelicerae with 5 teeth, retromargin with 5.
Leg measurements:

I II III IV
Femur 3.54 3.35 3.02 4.06
Patella + Tibia 4.56 4.05 3.55 4.75
Metatarsus 3.43 2.90 DT 3.66
Tarsus 1.70 TRS, 1.43 1.51
Total 13.23 11.81 10.71 13.98

Palp (Figs 5-8): Patellar apophysis relatively short, with blunt apex; RTA long;
retrolateral dorsal tibial apophysis slightly separated from RTA; cymbial furrow more
than half of cymbial length; basal lamella of conductor large; conductor broad, with
small apex bifurcation (Fig. 7); dorsal apophysis of conductor small; median apophysis
absent; embolus long, slender, originating posteriorly.

? (paratype). Total length 10.63. Carapace 4.96 long, 3.25 wide. Abdomen 5.13
long.

Eye sizes and interdistances: AME 0.18, ALE 0.20, PME 0.16, PLE 0.20; AME-
AME 0.13, AME-ALE 0.10, PME-PME 0.19, PME-PLE 0.25, ALE-PLE 0.07; MOQ
0.57 long, anterior width 0.50, posterior width 0.52. Clypeus height 0.22. Promargin of
chelicerae with 5 teeth, retromargin with 4.

Leg measurements:

I II III IV
Femur 4.39 4.10 3.64 4.37
Patella + Tibia SD 5102 2410) 5.55
Metatarsus 4.59 4.05 3.82 5.06
Tarsus 4.27 218 1.73 2.05
Total 18.97 15.30 11.89 17.03

Epigynum and vulva (Figs 9, 10): No epigynal teeth present; epigynum with an-
terior triangular ridge; atrium small; copulatory ducts round, anteriorly situated,
strongly sclerotized; spermathecal heads posteriorly located; spermathecae with large
and strongly convoluted lateral extensions.

Distribution and habitat: Known only from the type locality. All specimens ex-
amined were collected in evergreen hill forest between 1000 m and 1600 m.

Draconarius pseudolateralis sp. n. Figs 11-14

Type locality: THAILAND, Chiang Mai Province and District, Doi Suthep-Pui National
Park, Doi Suthep, near Monthathan Waterfall, 650 m.

Type material: Holotype: 4, from the type locality, pitfall trap, 14.x11.1986-10.1.1987
(MHNG). Paratypes: 116, 39, same data as for holotype (MHNG; CAS; PDC AM-0456-9);
1d, 19, 650 m, pitfall trap, 30.xi.-14.xii.1996 (MHNG); 134, 650 m, pitfall trap, 10.1.-
26.11.1997 (MHNG; CAS); 276, 22, 650 m, pitfall trap, 10.i.-6.11.1997 (MHNG); 2¢, 500 m,
leaf litter sample, 15.1.1986 (MHNG). All leg. P. J. Schwendinger. Chiang Mai Province,
Chomthong District, Doi Inthanon National Park, Doi Inthanon: 500 m, deciduous dipterocarp
forest, pitfall trap: 14, 25.i-26.11.2000, leg. P. Dankittipakul & S. Sonthichai (PDC AM-0460):
28, 25.xii.1999-25.1.2000, leg. P. Dankittipakul & S. Sonthichai (PDC AM-0461-2); 750 m,

COELOTINE SPIDERS FROM NORTHERN THAILAND II 545

Fics 11-14

Draconarius pseudolateralis sp. n., d holotype (11, 12) and © paratype (13, 14). Left palp, ven-
tral (11) and retrolateral (12) view. Epigynum, ventral view (13). Vulva, dorsal view (14). Scale
lines 0.5 mm (11, 12), 0.25 mm (13, 14).

deciduous dipterocarp-pine forest, 3 2, leaf litter sample, 15.xii.1999, leg. P. Dankittipakul & S.
Sonthichai (MHNG; CAS); Pha Mon, 1000 m, pine forest, 16, 27.xi.1987, leg. P. J.
Schwendinger (MHNG). Lamphun Province, Mae Tha District, Doi Khun Tan National Park,
Doi Khun Tan, 750 m, deciduous dipterocarp forest, 16, collected 20.x.2002, molted
20.xii.2002, leg. P. Dankittipakul (MHNG).

546 P. DANKITTIPAKUL & X.-P. WANG

Etymology: The specific name derives from the similarity between D. pseudolateralis sp.
n., D. paralateralis sp. n. and D. lateralis sp. n., all of which possess laterally extended sper-
mathecae.

Diagnosis: Males can be distinguished from those of D. lateralis sp. n. and D.
paralateralis sp. n. by a short, blunt patellar apophysis of the palp and by a long, slen-
der conductor apex bifurcation (Figs 11, 12); female by large copulatory ducts situated
close to each other (Fig. 14) and by differences in their atrial shapes (Fig. 14).

Description: & (holotype). Total length 8.45. Carapace 4.83 long, 3.31 wide.
Abdomen 3.64 long.

Eye sizes and interdistances: AME 0.15, ALE 0.18, PME 0.16, PLE 0.18; AME-
AME 0.13, AME-ALE 0.07, PME-PME 0.15, PME-PLE 0.20, ALE-PLE 0.07; MOQ
0.47 long, anterior width 0.42, posterior width 0.46. Clypeus height 0.24. Promargin of
chelicerae with 5 teeth, retromargin with 5.

Leg measurements:

I II III IV
Femur 4.03 5:55 3.25 3.96
Patella + Tibia DI 4.32 SA 5.08
Metatarsus 3.92 3.30 3.10 4.15
Tarsus 2.05 1.84 1.56 1.90
Total 15.13 13.01 11.62 15.09

Palp (Figs 11, 12): Patellar apophysis short, with blunt apex; RTA long; retro-
lateral dorsal tibial apophysis slightly separated from RTA; cymbial furrow more than
half of cymbial length; basal lamella of conductor large; conductor broad, with one
small and one large apex bifurcation; dorsal apophysis of conductor small but larger
than in the previous two species; median apophysis absent; embolus long, slender, pos-
terior in origin.

? (paratype). Total length 9.05. Carapace 4.51 long, 3.10 wide. Abdomen 3.50
long.

Eye sizes and interdistances: AME 0.10, ALE 0.15, PME 0.15, PLE 0.16; AME-
AME 0.13, AME-ALE 0.07, PME-PME 0.15, PME-PLE 0.18, ALE-PLE 0.07; MOQ
0.45 long, anterior width 0.36, posterior width 0.45. Clypeus height 0.23. Promargin of
chelicerae with 5 teeth, retromargin with 5.

Leg measurements:

I II III IV
Femur 3.30 3.05 23 3:99
Patella + Tibia 4.21 3.56 3.03 4.06
Metatarsus 3.05 2.62 225 3.33
Tarsus 1.32 1.30 1.13 1.62
Total 12.08 10,53 9.14 12.56

Epigynum and vulva (Figs 13, 14): No epigynal teeth present; atrium small,
umbrella-shaped, much wider than long (Fig. 13); copulatory ducts round, anteriorly
situated and close to each other; spermathecal heads round, posteriorly located;
spermathecae large and strongly convoluted lateral extensions (Fig. 14).

COELOTINE SPIDERS FROM NORTHERN THAILAND II 547

Distribution and habitat: Known from four localities with seasonally dry and
fairly disturbed habitat in northern Thailand.

The elatus group

Females can be recognized by their long epigynal teeth and broad, round sper-
mathecae; males by the small size, the elevated, short RTA (less than half of tibial
length), the very short cymbial furrow (almost invisible), the reduced tegular sclerite,
and the large embolic base. Both sexes have three promarginal (one of them are tiny
and almost invisible) and three retromarginal cheliceral teeth (tiny, bud-like and not

sharp).

Draconarius elatus sp. n. Figs 15-18

Type locality: THAILAND, Chiang Mai Province and District, Doi Suthep-Pui National
Park, Doi Suthep, Tham Ryssie, 1180 m.

Type material: Holotype: d, from the type locality, evergreen hill forest, pitfall trap,
4.x11.1986-2.1.1987, leg. P. J. Schwendinger (MHNG). Paratypes: from the type locality, 19,
134 , pitfall trap, 30.iii.-28.1v.1987 (MHNG); 34, 19, pitfall trap, 22.iv.-7.vi.1986 (CAS); 14,
pitfall trap, 18.iii.-22.iv.1986 (MHNG); 14, pitfall trap, 30.11.1987 (MHNG); 1 à, pitfall trap,
19.i1.-18.111.1986 (MHNG); 29, pitfall trap, 2-29.viii.1987 (MHNG); 46 , pitfall trap, 30.v.-
2.vii.1987 (MHNG), 126, 19, pitfall trap, 28.iv.-30.v.1987 (MHNG). All leg. P. J.
Schwendinger. Doi Suthep, 960 m (evergreen hill forest): 19, pitfall trap, 30.vii.30.viii.1986
(MHNG); 16, pitfall trap, 2.i.-5.iii.1987 (MHNG); 16, pitfall trap, 5.iii.-4.iv.1987 (MHNG);
24, pitfall trap, 28.iv.-30.v.1987 (MHNG); 26, pitfall trap, 4-28.iv.1987 (MHNG); 19, pitfall
trap, 26.111.-30.vi1.1996 (MHNG). All specimens leg. P. J. Schwendinger. Chomthong District,
Doi Inthanon National Park, Doi Inthanon: 19 , 1780 m (evergreen hill forest), 3.111.1987, leg. P.
J. Schwendinger (MHNG); 34, 1500 m (evergreen hill forest), pitfall trap, 21.1x.-23.x.1999
(CAS); 16, pitfall trap, 20.11.-25.111.2000 (CAS); 16 , leaf litter sample, 25.11.2000 (PDC AM
0156); 26 , pitfall trap, 25.iv.-27.v.2000 (PDC AM 0157-8); 19, leaf litter sample, 15.X.1999,
(CAS); 19, leaf litter sample, 25.iii.2000 (PDC AM 0159); 16, pitfall trap, 25.v.-24.vi.2000
(PDC AM 0160). 14, 510m (dry dipterocarp forest), pitfall trap, 25.vi.-29.vii.2000 (CAS); 24,
29, leaf litter sample, 29.iv.2000 (PDC AM 0161-64). 1 9, 750m (dipterocarp forest with pines),
leaf litter sample, 15.xii.1999 (CAS); 1 9, leaf litter sample, 15.v.2000 (CAS); 14, 1000 m (dry
dipterocarp forest with pines), pitfall trap, 23.xi.-25.x11.1999 (PDC AM 0165); 16, pitfall trap,
26.iii.-29.1v.2000 (PDC AM 0166); 1 4, pitfall trap, 25.v.-24.vi.2000 (PDC AM 017); 19, pitfall
trap, 26.iv.-27.v.2000 (MHNG); 1°, leaf litter sample, 29.vii.2000 (PDC AM 0168). All latter
specimens leg. S. Sonthichai & P. Dankittipakul.

Etymology: The specific name refers to the short, elevated RTA in males of this species.

Diagnosis: Males can be easily recognized by the short (less than half of tibia
length), broadly rounded, strongly elevated RTA, the very short cymbial furrow, the
reduced tegular sclerite, and the broad embolic base (Figs 15, 16); female by their long
epigynal teeth and broad, round spermathecae (Figs 17, 18). D. elatus sp. n. is usually
less than 5 mm long and thus one of the smallest known coelotines.

Description: è (holotype). Total length 3.78. Carapace 1.67 long, 0.74 wide.
Abdomen 1.56 long.

Eye sizes and interdistances: AME 0.09, ALE 0.10, PME 0.08, PLE 0.11; AME-
AME 0.07, AME-ALE 0.06, PME-PME 0.06, PME-PLE 0.13, ALE-PLE 0.03; MOQ
0.27 long, anterior width 0.25, posterior width 0.26. Clypeus height 0.13. Promargin of
chelicerae with 3 teeth (including a tiny tooth), retromargin with 3 small, bud-like
teeth.

548 P. DANKITTIPAKUL & X.-P. WANG

Fics 15-18

Draconarius elatus sp. n., ¢ holotype (15, 16) and © paratype (17, 18). Left palp (partly ex-
panded), ventral (15) and retrolateral (16) view. Epigynum, ventral view (17). Vulva, dorsal view
(18). Scale lines 0.5 mm.

Leg measurements:

I II III IV
Femur 1.62 1.44 1237, 1.79
Patella 0.62 0.62 0.55 0.65
Tibia 1.337] 1.10 1.03 1.62
Metatarsus 1.24 1.03 1.24 1.89
Tarsus 0.96 0.86 0.82 1.10

Total 5.81 5.05 5.01 7.05

COELOTINE SPIDERS FROM NORTHERN THAILAND II 549

Palp (Figs 15, 16): Patellar apophysis small; RTA strongly elevated, broadly
rounded, short, less than half of tibial length; retrolateral dorsal tibial apophysis short,
wide; cymbial furrow very short, almost invisible; conductor with broad base and
slender apex; dorsal apophysis of conductor present; tegular sclerite reduced; median
apophysis spoon-like; embolic base large; embolus relatively short, originating
posteriorly.

2 (paratype). Total 3.55 length. Carapace 1.41 long, 0.69 wide. Abdomen 1.00
long.

Eye sizes and interdistances: AME 0.09, ALE 0.14, PME 0.10, PLE 0.11; AME-
AME 0.06, AME-ALE 0.36, PME-PME 0.10, PME-PLE 0.13, ALE-PLE 0.06; MOQ
0.27 long, anterior width 0.24, posterior width 0.27. Clypeus height 0.12. Promargin of
chelicerae with 3 teeth (including a tiny tooth), retromargin with 3 small, bud-like
teeth.

Leg measurements:

I II III IV
Femur 1.62 1.44 1.34 118722
Patella 0.55 0:59 0.58 0.58
Tibia 1.48 1177, 1.00 1.68
Metatarsus 1.20 1.03 1.34 1.93
Tarsus 193 0.93 0.82 19197
Total 5.98 Su 5.08 7.08

Epigynum and vulva (Figs 17, 18): Epigynal teeth long; atrium short and broad,
situated posteriorly above epigastric furrow (Fig. 17); copulatory duct small; sper-
mathecae broad, round, slightly separated from each other (Fig. 18).

Distribution and habitat: Known from deciduous and evergreen forests in the
Doi Suthep-Pui (960-1180 m) and Doi Inthanon (510-1000 m) National Parks.

ZOOGEOGRAPHY

Draconarius paralateralis sp. n. and D. elatus sp. n. are here reported from
evergreen hill forests at about 1100-1600 m on Doi Suthep-Pui, whereas D. pseudo-
lateralis sp. n. was collected from dry dipterocarp forests at lower altitudes (500-650
m) on the same mountain. Another species, D. anthonyi Dankittipakul & Wang, was
previously reported near the summit of Doi Pui (1500-1680 m), which is only about
1 km away from the summit of Doi Suthep (Dankittipakul & Wang, 2003). The syn-
topic occurrence of four congeneric species (D. paralateralis sp. n., D. pseudolateralis
sp. n., D. elatus sp. n. and D. anthonyi), partially separated by altitude, is remarkable
and emphasizes the high biodiversity of this mountain. Four species are also known
from a nearby mountain (Doi Inthanon), i.e. D. elatus sp. n., D. inthanonensis
Dankittipakul & Wang, 2003, D. siamensis Dankittipakul & Wang, 2003, and D. sub-
ulatus Dankittipakul & Wang, 2003. Although it is not clear whether the species of the
lateralis group are really congeneric with other members of Draconarius, each moun-
tain evidently harbors four amaurobiids species. At present a total of ten coelotine
species are known from Thailand.

550 P. DANKITTIPAKUL & X.-P. WANG

It is also interesting to note that although D. paralateralis sp. n. and D.
pseudolateralis sp. n. are represented by fairly low numbers of mature males and
females, the corresponding numbers of immature specimens collected are rather high.
They were present in several instars at any time, indicating that the spiders live for
more than one year.

ACKNOWLEDGEMENTS

We are grateful to Dr Peter J. Schwendinger for providing material from his
private collection and for the loan of specimens from the MHNG. We thank Dr Charles
Griswold for allowing the depository of some types in the collection of the Department
of Entomology, California Academy of Sciences. Work on this paper was carried out
while the first author was based at Chiang Mai University. He is grateful to Prof.
Saowapa Sonthichai for providing research facilities there.

The Thailand Research Fund TRF/BIOTEC Special Program for Biodiversity
Research and Training Grant (project number BRT_R 145002) supported the first
author at the beginning of this project. The Royal Forest Department gave permission
to collect spiders in protected areas.

REFERENCES

DANKITTIPAKUL, P. & WANG, X.-P. 2003. New species of coelotine spiders (Araneae, Amauro-
biidae) from northern Thailand I. Revue suisse de Zoologie 110(4): 723-737.

OVTCHINNIKOV, S. V. 1999. On the supraspecific systematics of the subfamily Coelotinae
(Araneae, Amaurobiidae) in the former USSR fauna. Tethys Entomological Research |:
63-80.

WANG, X. P. 2002. A generic-level revision of the spider subfamily Coelotinae (Araneae,
Amaurobiidae). Bulletin of the American Museum of Natural History 269: 1-150.

WANG, X. P. 2003. Species revision of the coelotine spider genera Bifidocoelotes, Coronilla,
Draconarius, Femoracoelotes, Leptocoelotes, Longicoelotes, Platocoelotes, Spiri-
coelotes, Tegecoelotes, and Tonsilla (Araneae, Amaurobiidae). Proceedings of the Cali-
fornia Academy of Sciences 54(26): 497-660.

WANG, J. F., YIN, C. M., PENG, X. J. & XIE, L. P. 1990. New species of the spiders of the genus
Coelotes from China (Araneae: Agelenidae) (pp. 172-253). In: Spiders in China: One
Hundred New and Newly Recorded Species of the Families Araneidae and Agelenidae.
Hunan Normal University Press.

REVUE SUISSE DE ZOOLOGIE 111 (3): 551-562; septembre 2004

An enigmatic cave-dwelling ground beetle: Trechus barratxinai
Espanol, 1971 (Coleoptera, Carabidae, Trechinae, Trechini)*

Vicente M. ORTUNO
Departamento de Biologia Animal, Facultad de Biologia, Universidad de Alcala,
E-28871 Alcala de Henares, Madrid, Spain. E-mail: vicente.ortuno@uah.es

An enigmatic cave-dwelling ground beetle: Trechus barratxinai
Espanol, 1971 (Coleoptera, Carabidae, Trechinae, Trechini). - An im-
portant population of Trechus barratxinai Espanol, 1971 was discovered in
the Penyes de Roset, near the type locality (abyss of Barratxina, Xixona-
Alicante, East Spain). This species, originally considered as cavernicolous,
shows epigeal and lapidicolous life habits, as now discovered. A detailed
laboratory study revealed the presence of wings which had been overlooked
in the concise description of the species. The development of wings
confirms the suspicion that, as in other species in this genus, 7. barratxinai
has geophilic, lucifugous and hygrophilous habits but is not a cavernicolous
(troglobiont) species. The species description is completed with details on
the anatomy of the female genitalia. Also, the taxonomic position of T. bar-
ratxinai is questioned and its position within the “7. austriacus-group”, near
T. diecki Putzeys 1870, its geographical neighbour, is proposed. Finally, the
first data on the species biology and on its habitat characteristics are given.

Keywords: Carabidae - Trechus - redescription - systematics - ecology -
caves - Iberian Peninsula.

INTRODUCTION

Trechus Clairville, 1806 is a large genus, including more than 500 species
(Casale et al., 1998), with new ones being discovered every year. Presently it is known
from the Holarctic region and East Africa. From the Iberian Peninsula 49 species are
recorded (Serrano, 2003); of these, 18 have strictly hypogeous life with morphological
adaptations to the subterranean environment, but only 14 of these species are known
exclusively from caves. As for the Spanish-Levante fauna, the following are the known
hypogeous Trechus species, all of them limited to the Alicante province: T. martinezi
Jeannel, 1927, T. alicantinus Espafiol, 1971, T. beltrani Toribio, 1990 and 7. barratxi-
nai Espafiol, 1971. The first three species belong to the same species group, whilst the
position of T. barratxinai is enigmatic, it only has been found once in the abyss of
Barratxina in Xixona (Alicante, Spain). The citation of the species in the Cave of the

* This work has been funded by the Ministerio de Ciencia y Tecnologia within the
“Ramon y Cajal” plan.
Manuscript accepted 04.02.2004

552 V. M. ORTUNO

Farallön, Riöpar (Albacete, SE Spain) (Lencina er al., 1990) probably refers to an
undescribed species of Trechus. Espanol (1971) placed T. barratxinai into the
planipennis-pandellei-group (Trechus pyrenaeus-group and Trechus pandellei-group
sensu Jeannel, 1927). The author asserts that this species is the first cave-dwelling of
the mentioned group, whereas other species of the group are epigeous and more or less
orophilous. Neither its status as cavernicolous species (Jeanne, 1976; Sendra &
Zaragoza, 1982; Zaragoza & Sendra, 1988; Belles, 1987; Zaballos & Jeanne, 1994) nor
the close affinities with T. planipennis Rosenhauer, 1856 from Sierra Nevada (Espanol,
1971) had ever been questioned. Based on new morphological and ecological data the
author is reconsidering the affinities of this species within the genus Trechus.

MATERIAL AND METHODS

Examined specimens of 7. barratxinai: 3 males, 2 females, Penyes de Roset,
850 m.s.m., 30SYH17, Xixona (Alicante), 23-3-2003, V.M. Ortufio & J.A. Zaragoza
leg. 10 males, 7 females, idem, 7-5-2003, V.M. Ortufio & J.A. Zaragoza leg. 8 males,
idem, 15-6-2003, V.M. Ortufio leg. The specimens are deposited in the collection of
V.M. Ortufio (Dept. of Animal Biology, Alcalä University, Spain) and 2 males, 2 fe-
males in the Museum d’histoire naturelle Genève (Switzerland) and 1 male and 1 fe-
male in the collection M. Toribio (Tres Cantos, Spain).

Also, the following Trechus have been examined in this study: 7. martinezi
(three specimens from Cueva de las Maravillas, Alicante; with aborted wings),
T. quadristriatus (Schrank, 1781) (six specimens from Tres Cantos, Madrid, with func-
tional wings), 7. diecki Putzeys, 1870 (eight specimens from caves in Andalucfa) and
T. planipennis (two specimens from Laguna de las Yeguas, Sierra Nevada, Granada).
Other species of Carabidae (accompanying fauna) have also been studied.

All measurements were made using a calibrated ocular grid set in a microscope
and stereomicroscope ocular. The measurements of the wings are summarised in Table
I. Two indices proposed by Den Boer et al. (1980) were used to estimate relative wing
length (RL) and relative wing area (RA): RL = MLW /MLE; RA = MLW x MWW /
MLE x MWE (MLW = maximum wing length: MLE = maximum elytron length;
MWW = maximum wing width; MWE = maximum elytron width).

The aedeagus was extracted from the abdomen and separated from the tergal
apodemal ring; the parameres were separated from the associated membranes in the
surface of the median lobe. After examination, genital preparations were included in
DMHF (dimethyl hydantoin formaldehyde) and put on over acetate sheet.

A routine procedure was followed to prepare the female reproductive
appendages for scanning microscopy. The last abdominal segments of the female were
gently squeezed with forceps and introduced in saturated solution of KOH for 8 hours.
They then were washed in Scheerpeltz’s solution and opened dorsally for checking the
alkaline digestion. Staining was carried out with Chlorazol black E® in watery solution
for 1-3 minutes under visual control. The excess of dye was removed by washing in
KOH and the structures washed again in Scheerpeltz’s solution. The dissected
structures were placed in a watchglass with Scheerpeltz’s solution under a stereo-

AN ENIGMATIC CAVE-DWELLING GROUND BEETLE 553

microscope with a drawing tube for observation and drawing. Selected dissections
were mounted onto a slide and made permanent.

RESULTS AND DISCUSSION

REDESCRIPTION OF TRECHUS BARRATXINAI ESPANOL, 1971 (Figs 1, 2, 3a-b, 4a-b)

Diagnosis. Winged species (brachypterous). Macrophthalmic, depigmented
with microreticulate integument. Pronotum transverse and slightly cordate: hind angles
sharp and sides slightly sinuous in basal margin. Elytra convex, slightly protruding
shoulders, with five well-defined striae (the 6th and 7th not very defined, almost miss-
ing the striae). Both sexes show similar external morphology, except for the first two
segments of anterior tarsus dilated in males. Median lobe of the aedeagus slightly
curved; inner piece of median lobe simple. Female genitalia: genital shield with a line
of thorn-shape setae on gonosubcoxite; spermatheca long and slender lacking sper-
mathecal gland.

Description. Length: 3,9-4,9 mm (Fig. 1). Head (to anterior end of clypeus) a
little wider than long; in dorsal view two deep frontal sulci border ocular areas on both
sides and fronto-clypeal area towards the center. Eyes large but not prominent (ocular
convexity not very conspicuous). Tempora almost as long as the eyes. Antennae fili-
form, densely setulose, in particular from 2nd to 11th antennomere. Mandible promi-
nent, sharp. Labial and maxilary palps as typical in the genus. Cephalic chaetotaxy:
two pairs of supraocular setae (anterior and posterior); two setae at both sides of
clypeus (outer one largest); six setae on labrum, lateral setae more elongate; one seta
in sulcus of each mandible.

Pronotum one third wider than long, with the sides slightly sinuous in basal
half; maximum width slightly anterior the middle half; basal margin rectilinear and
smoothly broken toward the rear angles; basal foveae smooth and deep; hind angles
sharp and slightly diverging. Disk convex, divided lengthway by central sulcus branch-
ing off towards anterior and posterior angles. Lateral channel with regular width.
Pronotal chaetotaxy: one anterior seta on each side (almost in the first third) and one
posterior seta next to hind angle.

Elytra oval-shaped, approximately three times longer than wide; humeral mar-
gin well-defined, basally right and perpendicular to the sagittal plane. Disk convex
with transverse microsculpture (Fig. 2a); five well-defined striae (1st-Sth), the 6th and
7th almost missing. Elytral chaetotaxy: marginal umbilical series typical of Trechus
(humeral area with four equidistant setae; subapical area with four setae, two anterior
and two posterior). Each elytron with one setigerous pore beginning at 2nd stria
(scutellar pore), two discal setae in the 3rd stria (anterior seta in anterior quarter and
posterior setae behind half), similarly subapical seta present at end of 2nd-3rd stria, two
smaller ones near apical margin. Elytra with sparse micropubescence (only visible with
the microscope).

Short wings (brachypterous species) (Fig. 2b), only a little longer than the ely-
tra (Table I).

Legs long and slender, without special characteristics; foretibia with a longitu-
dinal sulcus; first two tarsal segments of male foreleg dilated.

554

V. M. ORTUNO

Fic. 1
Habitus of Trechus barratxinai Espafiol, 1971 (Scale: 1 mm).

AN ENIGMATIC CAVE-DWELLING GROUND BEETLE 555

FIG. 2

Morphological details of Trechus barratxinai Español, 1971: a) elytral microsculpture; b) wing.
(Scale a: 0.05 mm; scale b: 1 mm).

Aedeagus 0.6 mm long, (Figs 3a-b) with the median lobe not very arched; short,
thick and blunt apex; apical lamina subsymmetrical. Distal extreme transforming into
sagittal lamina. Inner sac very simple with a triangular lamina; there is superimposed
a field of spines in the wall of the praeputial tube. Parameres similar, but of different
lengths, both tetrasetulose in distal extreme.

Female genitalia (Figs 4a-b): external genitalia formed by dimerous IX
gonopods (gonocoxites and gonosubcoxites) and IX laterotergites. Each gonocoxite

556 V. M. ORTUNO

FIG. 3

Aedeagus. Trechus barratxinai Espanol, 1971: a) median lobe, left and right paramere in lateral
view; b) median lobe in dorsal view. Trechus diecki Putzeys, 1870 from Covadura (Sorbas,
Almeria): c) median lobe in lateral view. (Scale a,b,c: 0.2 mm).

AN ENIGMATIC CAVE-DWELLING GROUND BEETLE 557

FIG. 4

Female genitalia in ventral view. Trechus barratxinai Español, 1971: a) left genital shield; b)
spermathecal complex. Trechus diecki Putzeys, 1870 from Covadura (Sorbas, Almeria): c) sper-
mathecal complex. (Scale a: 0.1 mm; scale b,c: 0.3 mm).

558 V. M. ORTUNO

unguiform, with two thorn-shaped setae of considerable size on dorsal surface (the
largest located near external edge). Small groove near apex and above ventral surface,
with two fine sensorial setae. Gonosubcoxite subtriangular, a little longer than wide
and oblique with respect to laterotergite, with four large thorn-shaped setae; smaller
setae (1 to 2) found in other areas close to base. Wing-shape, slightly sclerotized IX
laterotergite with one group of setae over basal margin (approximately 20) and one
more internal group (approximately 6). Internal genitalia completely membranous;
tube-shaped vagina-bursa; spermatheca digitiform (long and narrow) with a diverti-
culum in the basal tract; associate gland absent (spermathecal or accessory gland). The
odd oviduct makes contact with spermathecal complex at the same point as where the
spermatheca opens, displaying some longitudinal folds; interior densely covered in
microfringes.

Variability. Important morphological variations have not been observed, and
the same applies to wing development: the relative length and relative area of the wings
is very similar in eight specimens randomly selected (5 males and 3 females) (Table I;
Fig. 5). Remarkable size differences among imagoes have been observed (see above).

TAXONOMIC POSITION AND BIOGEOGRAPHICAL COMMENTS

The overall body shape (tempora, pronotum and basal foveae, elytra) and the
development of the wings (brachypterous species) in 7. barratxinai are discordant with
the features of both the 7. pyrenaeus-group and the 7. pandellei-group. Also, the
aedeagus of 7. barratxinai is very different from that shown by the species in these

TABLE I. Values for six quantitative characters used in the analysis of Trechus wings. MLW =
maximum wing length; MWW = maximum wing width; MLE = maximum elytron length; MWE
= maximum elytron width; RL = relative wing length (in mm); RA = relative wing area (in
mm?).

sex MLW MWW MLE MWE RL RA

Trechus barratxinai 1 female 2.99 1.20 2.81 0.84 1.06 1.52
2 f 2.81 1.02 2.63 0.98 1.06 1.11
3 male 2.58 0.80 2.41 0.89 1.07 0.96
4 2.99 0.93 Dexa 0.98 11117 1.11
5 f 3.03 0.98 2.63 0.98 1.15 1.15
6 m 3.21 1.16 2.67 1.07 1.20 1.30
7 m 2.99 1.02 2.63 1.02 1513 1.112)
8 m 2.67 0.84 2.36 0.93 1.13 1.02
Trechus quadristriatus | m 5.26 2.18 2.67 0.84 1.97 SU
2 f SALE 1.96 2.67 0.80 1.92 4.70
3 f 4.37 1.83 2.41 0.89 1.81 SN
4 m 4.68 1.60 2.54 0.89 1.84 3.31
5 m SAIT 1.91 DID, 0.98 1.90 3.70
6 f 5.13 1.91 2.63 0.93 1.95 4.00

Trechus martinezi

Bram
©
N
Nn
©
Oo
un

NESS
©
m
—
WW
Uo
©
—
OO
©
©
1

AN ENIGMATIC CAVE-DWELLING GROUND BEETLE 559

T. quadristriatus

Relative wing area(mm 5)

T. barratxinai

T. martinezi

"0,0 0,4 0,8 1,2 1,6 2,0 2,4
Relative wing length (mm)

FIG. 5

Wing development in three species of Trechus: T. martinezi Jeannel, 1927 (species with aborted
wings), T. barratxinai Espafiol, 1971 (brachypterous species) and 7. quadristriatus (Schrank,
1781) (species usually with functional wings).

groups, including 7. planipennis, supposedly the most closely related species.
Relationships of 7. varendorffi Deville, 1903 from Corsica and 7. planipennis from
Sierra Nevada are quite close (demonstrated by several morphological characters and
the aedeagus structure due to their common origin in the old Corso-Betican mountain
range -Early Oligocene-), and it would not be surprising if, according to taxonomic and
geographical criteria, 7. barratxinai showed a great resemblance and also a certain eco-
logical relationship with the two species mentioned. But on the contrary, there is a great
morphological difference between 7. barratxinai and the species of the 7. pyrenaeus-
group (see above). Probably 7. barratxinai may in fact be a species of the Trechus aus-
triacus-group, with a great morphological resemblance of 7. barratxinai with T. diecki
(see below); also, both species are geographical neighbours. The 7. austriacus-group
proposed by Jeannel (1927), at the moment and with some upgrades, is formed by three
lineages. A first subgroup is formed by Trechus crucifer La Brülerie, 1875 (different
areas in the eastern Mediterranean), Trechus schmalfussi Baehr, 1983 (Grecia), Trechus
creticus Moravec, 2000 (Crete), Trechus diecki diecki Putzeys, 1870 (Southern Spain),
Trechus diecki maculipennis Bedel, 1897 (Algeria) and, as proposed here, Trechus
barratxinai (Alicante, Spain). A second subgroup, known from the Middle East, is
formed by Trechus saulcyi Jeannel, 1921, Trechus libanensis La Brülerie, 1875 and
Trechus pamphylicus Jeanne, 1996. Finally, if we accept the synonymies proposed by
Casale & Laneyrie (1982), the third subgroup is monospecific and is formed by
Trechus austriacus Dejean, 1831, from central and eastern Europe, but also from

560 V. M. ORTUNO

Middle East. However, this subgroup should be throughly revised to clarify the status
of some taxa synonymised by Casale & Laneyrie (1982) such as: Trechus subacumi-
natus Fleischer, 1898, Trechus labruleriei Jeannel, 1921, Trechus angustus Sahlberg,
1908 and Trechus olympicus La Brulerie, 1876.

Thus, in my opinion, the evidence for a close relationship between 7. barratxi-
nai and T. diecki is rather strong. Many morphological characteristics are common to
both species in the same phyletic line, in particular the genital (male and female) ana-
tomy. The aedeagus is almost identical in both species (Figs 3a,c), as well as the parts
of the inner sac that show great taxonomic and systematic significance. A supple-
mentary argument for a close relationship between T. barratxinai and T. diecki is the
extraordinary resemblance of the female genitalia in both taxa, in particular in the sper-
mathecal complex (Figs 4b,c): both genitalia have a long, digitiform spermatheca with
a diverticulum in the basal tract. On the contrary, T. plannipennis shows a more simple
structure without diverticulum that, in turn, is different and more complicated than
other Trechus genitalia (see, e.g., Salgado & Ortuno, 1998).

From all this evidence I propose that the ancestors of 7. barratxinai took refuge
during the Miocene in eastern Iberia. Subsequent climatic changes during periods of
high dryness decimated this phyletic line. They found shelter in “area-islands” with
high humidity and low temperature, an optimal combination for this hygrophilous
species, the Penyes de Roset being one of these refuges (also, other hygrophilous cara-
bids and Prionoglaris stygia Enderlein, 1909 a troglophilous Psocoptera, have been
located there).

In spite of the taxonomic relationship of 7. barratxinai and T. diecki, it is
possible to differentiate both species by the following characteristics: 7. diecki presents
dark pigmentation, tetramaculate elytra, deeper striae (6th and 7th well-defined) and
convex interstriae, deeper microreticulation integument (especially in the elytra),
shorter tempora and aedeagus with median lobe not very arched; 7. barratxinai shows
unicoloured and depigmented integument, more superficial striae (6th and 7th not well-
defined) and not very convex interstriae, shallow microreticulation, longer tempora
and, finally, slightly more arched median lobe.

ECOLOGICAL FEATURES

The Penyes de Roset is a very special area in the orography from Alicante
province. They are cliff-rocks of calcareous character. Many of the rocky walls
collapsed and formed abyss and cracks of tectonic origin. The substrate is very spongy
and porous in the base of the cliff-rocks forming screes. This place rises on a vast xeric
landscape of the Subcomarca from Canal de Xixona, forming small havens of high
humidity and low temperature. In view of these peculiarities, it deserves mention a
small list of the more common rupicolous plants: Hedera helix L., Pistacia terebinthus
L., Umbilicus horizontalis (Guss.) DC.; Lavatera maritima Gouan, Ruscus aculeatus
L., Verbascum sp., Rosmarinus officinalis L., Thymus sp., and different species of
Asplenium and Sempervivum.

Many geophilic carabids of extreme or limited habitats show stable populations.
In these cases dispersal is not necessary and the imagoes fix their brachypterous
condition (Darlington, 1943). This same phenomenon occurs in 7. barratxinai, that has

AN ENIGMATIC CAVE-DWELLING GROUND BEETLE 561

stabilized the character “reduced wings” (Fig. 5). According to this distribution model
we are dealing with a telospecies or senile species (now in contraction; see Rapoport,
1975: 68). T. barratxinai is very lucifugous hiding in shady areas and accidentally
inside the abyss and cracks. T. barratxinai should not be considered as a cavernicolous
species (without further qualification) and much less as troglobite (for example, see
Jeanne, 1976). In any case, there are reasons to believe that it is a troglophilous species;
this eco-physiological trend is favoured by the “pre-adaptation” within this Trechus
lineage (hygrophilous and lucifugous species), and also by the very cracked substratum
with numerous underground refuges. The troglophilous behaviour of T. barratxinai
also appears in other species of the group (or phyletic line) of Trechus austriacus. For
example, the epigeous species T. austriacus is located in the caves of the Balkan
Peninsula (Jeannel, 1941) and Bulgaria (Heynderycx, 1996); T. saulcyi appears in
caves of Turkey, Syria and Cyprus, occasionally (Decu er al., 2001); T. diecki is
common, but not exclusive, in the caves of the south-eastern of the Iberian Peninsula
(Mateu, 1956; Jeanne, 1967; Tinaut, 1998); Finally, 7. crucifer is known as troglo-
philous species in some areas of the southern and northern Mediterranean (Coiffait,
1954; Heynderycx, 1996).

To conclude, 7. barratxinai is a geophilic, lucifugous and hygrophilous species
that, however, lacks important adaptive features to the subterranean environment: the
wing is much less reduced than in cavernicolous species, for example 7. martinezi
(Fig. 5). On the other hand, some species have large wings and are capable of flight (7.
quadristriatus, which exceptionally might be brachypterous) (Fig. 5); the wings of 7.
barratxinai do undoubtedly not allow flight (Fig. 2b, 5). This species has lapidicolous
life in fresh and humid places of the Penyes de Roset; in this area the temperature is
about 15°C during summer. The very cracked substratum facilitates colonization of the
abyss (specimen of the type series) and the mesocavernous shallow substratum (8
specimens studied in this work). 7. barratxinai coexists with other species of carabids,
e.g. Carabus (Mesocarabus) helluo Dejean, 1826, Orthomus expansus Mateu, 1952,
Percus (Pseudopercus) guiraoi Pérez-Arcas, 1869, Laemostenus (Pristonychus) terri-
cola (Herbst, 1783), Platyderus sp., Cryptophonus tenebrosus (Dejean, 1829), Har-
palus (Harpalus) atratus Latreille, 1804, Harpalus (Harpalus) wagneri Schauberger,
1926, Licinus punctatulus granulatus Dejean, 1826 and Brachinus (Brachynidius)
sclopeta (Fabricius, 1792), some of them previously not recorded from the Spanish-
Levante.

ACKNOWLEDGEMENTS

I wish to express my thanks to Juan A. Zaragoza for his collaboration in the
biospeleological expeditions of 2003 and an anonymous reviewer for helpful
comments on a first draft. I also thank Jaime Potti for his revision of the English.

REFERENCES
BELLES, X. 1987. Fauna cavernicola i intersticial de la peninsula ibérica i les illes balears. CSIC.,
Ed. Moll, Mallorca, 207 pp.

CASALE, A. & LANEYRIE, R. 1982. Trechodinae et Trechinae du Monde. Tableau des sous-
familles, tribus, series phylétiques, genres, et catalogue general des espèces. Memoires
de Biospéologie 9: 1-226.

562 V. M. ORTUNO

CASALE, A., VIGNA-TAGLIANTI, A. & JUBERTHIE, Ch. 1998. Coleoptera Carabidae (pp. 1047-
1081). Jn: JUBERTHIE, C. & DECU, V. (eds). Encyclopaedia Biospéologica. Vol. 2. Société
de Biospeologie, Moulis-Bucarest.

COIFFAIT, H. 1954. Mission Henri Coiffait au Liban (1951). Coléoptères cavernicoles et endoges.
Archives de Zoologie Experimentale et Generale 91(4): 423-436.

DARLINGTON, P.J., Jr. 1943. Carabidae of mountains and islands: data on the evolution of isolat-
ed faunas, and on atrophy of wings. Ecological Monographs 13: 37-61.

DECU, V., JUBERTHIE, Ch., BAKALOWICZ, M. & TABACARU, I. 2001. Turquie (pp. 2007-2026). In:
JUBERTHIE, C. & DECU, V. (eds). Encyclopaedia Biospéologica. Vol. 3. Société de
Biospéologie, Moulis-Bucarest.

DEN Boer, P.J., HUIZEN, T.H.P., DEN BOER-DAANJE, W., AUKEMA, B. & DEN BIEMAN, C.F.M.
1980. Wing polymorphism and dimorphism in ground beetles as stages in an evo-
lutionary process (Coleoptera: Carabidae). Entomologia Generalis 6(2-4): 107-134.

EsPaNOL, F. 1971. Nuevos tréquidos cavernicolas de la fauna española (Col. Caraboidea).
Publicaciones del Instituto de Biologia Aplicada de Barcelona 51: 89-96.

HEYNDERYCX, J. 1996. Les Carabidae troglobies de Bulgarie. Lambillionea 96(2): 292-294.

JEANNE, C. 1967. Carabiques de la Péninsule Ibérique (5° note). Actes de la Société linnéenne de
Bordeaux 104(10), ser. A: 3-22.

JEANNE, C. 1976. Carabiques de la Péninsule Ibérique (2° supplément). Bulletin de la Société
linnéenne de Bordeaux 6(7-10): 27-46.

JEANNEL, R. 1927. Monografie des Trechinae. Morphologie comparée et distribution géo-
graphique d’un groupe de Coléoptères (II livraison). L’Abeille 33: 1-592.

JEANNEL, R. 1941. Faune de France, Coléoptères Carabiques, Première Partie. Lechevalier,
Paris, 571 pp.

LENCINA, J.L., ANDUJAR, A. & RUANO, L. 1990. Algunas citas de interés de la fauna de
coleöpteros de la provincia de Albacete. Revista de Estudios Albacetenses 27: 101-121.

MATEU, J. 1956. Atranus collaris (Menetries) (Col. Carabidae). Género y especie nueva para la
fauna espafiola y observaciones sobre caräbidos ibéricos. Eos 32(1-4): 135-140.

Rapoport, E.H. 1975. Aerografia. Estrategias geogräficas de las especies. Fondo de Cultura
Economica, Mexico, 214 pp.

SALGADO, J.M. & ORTUNO, V.M. 1998. Two new cave-dwelling beetle species (Coleoptera:
Carabidae: Trechinae) of the Cantabrian Karst (Spain). The Colepterists Bulletin 52(4):
351-362.

SENDRA, A. & ZARAGOZA, J.A. 1982. Invertebrados cavernicolas del Pais Valenciano. Lapiaz 10:
14-22.

SERRANO, J. 2003. Catalogo de los Carabidae (Coleoptera) de la Peninsula Ibérica. Monograffas
S.E.A. 9. Sociedad Entomolögica Aragonesa, Zaragoza, 130 pp.

TINAUT, A. 1998. Artrépodos terrestres de las cavidades andaluzas. Zoologica baetica 9: 3-28.

ZABALLOS, J.P. & JEANNE, C. 1994. Nuevo catalogo de los caräbidos (Coleoptera) de la Peninsula
Ibérica. Monografias S.E.A. 1. Sociedad Entomolögica Aragonesa, Zaragoza, 159 pp.

ZARAGOZA, J.A. & SENDRA, A. 1988. Fauna cavernicola de la provincia de Alicante. Ayudas a la
investigaciön 1984-1985. Inst. Est. “Juan Gil-Albert”. Diputaciôn de Alicante 3: 11-35.

REVUE SUISSE DE ZOOLOGIE 111 (3): 563-565; septembre 2004

Une espèce nouvelle de Pseudosinella provenant du Portugal
(Insecta: Collembola). XVII® contribution

Maria Manuela DA GAMA
Instituto do Ambiente e Vida, Departamento de Zoologia da Universidade de Coimbra,
3004-517 Coimbra, Portugal.

A new species of Pseudosinella from Portugal (Insecta: Collembola).
XVIIth contribution. - Pseudosinella angelae sp. n. is described from a
typical Mediterranean landscape dominated by Quercus suber L. in the
surroundings of Lisbon. Morphological comparison with similar species is
made, in order to establish the genealogical position of the new species.

Keywords: Insecta - Collembola - Pseudosinella - new species - Portugal.

INTRODUCTION

Cette étude s’insere dans un projet financé par l’Union Européenne,
‘BIOASSESS — European Biodiversity Assessment Tools’, dont l’objectif principal
consiste dans le développement d’un groupe d’instruments efficaces pour l’évaluation
de la biodiversité, afin de contröler les changements dans les patrons de biodiversité
des organismes vivants. Ces changements sont causés par les modifications dérivées de
politiques communautaires dans l’usage du sol, comme l’agriculture, la sylviculture,
l’urbanisme, etc.

Les matériaux qui concernent la description de cette espèce nouvelle ont été
récoltés dans un bois méditerranéen typique, dont l’espece dominante est le chéne-
liege (Quercus suber). Il s’agit d’une zone ou l’intervention humaine est faible, située
sur le rivage gauche du fleuve Tage à 20 km de Lisbonne.

Les types de la nouvelle espèce sont déposés au Muséum d’histoire naturelle de
Genève (MHNG) et au Musée Zoologique de l’Université de Coimbra (MC).

Pseudosinella angelae sp. n. Figs 1-2

STATION ET TYPES

Portugal, Companhia das Lezirias, environs de Lisbonne, bois “montado” de
chéne-liège (Quercus suber) avec une strate herbacée dominée par Cistus salvifolius
L., Calluna vulgaris (L.) Hull et Ulex australis Clemente.

L’holotype et 2 paratypes, en préparation, IV. 2002, sont déposés au MHNG.
4 paratypes, en préparation, IV. 2002, sont déposés au MC.

DESCRIPTION

Taille: 0,8-1 mm. Le pigment bleu se trouve uniquement dans les 2+2 yeux, de
taille identique, placés l’un à côté de l’autre. Rapport entre la longueur des antennes et

Manuscrit accepté le 28.11.2003

564 M. M. DA GAMA

‚N / 5: No
|

SI a

FIG. 1
Pseudosinella angelae sp. n.: Chétotaxie dorsale de l’abd. II, côté gauche.

FIG. 2
Pseudosinella angelae sp. n.: Griffe 3, face antérieure.

UNE NOUVELLE ESPECE DE PSEUDOSINELLA 565

la diagonale céphalique variant de 1,2 à 1,4. L’organe antennaire III présente la
structure habituelle. Articles antennaires sans écailles. Macrochètes dorsaux:
R011/10/0101+2. Le macrochete céphalique dorsolatéral est accompagné d’une
trichobothrie. Chétotaxie de l’abdomen II: pa B qj (Fig.1). Soie accessoire ‘s’ de
l’abdomen IV absente. Base du labium: M, Mr E L, L;: r est rudimentaire et les
autres soies sont distinctement ciliées. La dent impaire de la griffe est située à environ
68-70% de sa créte interne et la dent proximale postérieure est un peu plus développée
que la dent proximale antérieure (Fig. 2). Empodium lancéolé. Ergot tibiotarsal non
spatulé. Les dents apicale et anteapicale du mucron sont subégales.

DISCUSSION

La formule chétotaxique des macrochètes dorsaux, de l’abdomen II et de la base
du labium de la nouvelle espèce est identique à celle de P. zygophora (Schille, 1908)
et de P. picta Börner, 1903, qui ne possèdent pas non plus de soie accessoire s sur
l’abdomen IV. Néanmoins P. angelae sp. n. se distingue de ces deux espèces par le
nombre des yeux (5+5 chez P. zygophora et 4+4 chez P. picta) et par des petits details
concernant la structure de la griffe, de l’empodium et de l’ergot tibiotarsal.

Ces trois espèces doivent appartenir à la lignée généalogique dérivée de Lepido-
cyrtus pallidus Reuter, 1890 emend. Gisin, 1965 (Gisin, 1967; Gama, 1984): p présent
sur l’abdomen II, r rudimentaire sur la base du labium.

P. obanae Gruia, 1998, provenant de Roumanie, qui fait partie de cette méme
lignée généalogique, semble se distinguer de la nouvelle espèce uniquement par la
chétotaxie du thorax II sans macrochètes. Toutefois la disposition des yeux est
différente, car ils sont placés l’un derrière l’autre chez P. obanae et l’un à côté de
l’autre chez P. angelae sp. n.

BIBLIOGRAPHIE

GAMA, M. M. da 1984. Phylogénie des espèces européennes de Pseudosinella (Collembola:
Entomobryidae). Annales de la Société royale zoologique de Belgique 114: 59-70.

GISIN, H. 1967. Espèces nouvelles et lignées évolutives de Pseudosinella endogés. Memorias e
Estudos do Museu zoolögico da Universidade de Coimbra 301: 1-21.

GRUIA, M. 1998. Sur la faune de Collemboles de l’écosystème exokarstique et karstique de
Movile (Dobrogea du Sud, Mangalia, Romania). Mémoires de Biospéologie 25: 45-52.

N j de II 4
7 i pa Ve: ee Lee |

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REVUE SUISSE DE ZOOLOGIE 111 (3): 567-583; septembre 2004

Une nouvelle espèce d’Hyperolius du Cameroun
(Amphibia, Anura, Hyperoliidae)

Jean-Louis AMIET
48, rue des Souch£res, F-26110 Nyons.

A new species of Hyperolius from Cameroon (Amphibia, Anura, Hyper-
oliidae. - Previously mentioned as “Hyperolius sp. 6”, Hyperolius cameru-
nensis sp. n. is distributed in a small area of south-west Cameroon. It differs
from related species by its phase III colour pattern, with dorsum marked of
more or less numerous large rounded red spots, and ventrum brightly
coloured of orange and white-speckled black. This pattern is exhibited by
females and about a quarter of the reproducing males. In phase II, H. came-
runensis is very similar to H. riggenbachi, but it is smaller and has a broader
head. H. camerunensis is a farmbush species, differing from other
Cameroonian anurans by its horizontal and vertical distribution. A definition
of the species group to which it is related is proposed.

Keywords: Amphibia - Anura - Hyperoliidae - Hyperolius - Cameroon -
systematics - ecology.

INTRODUCTION

La batrachofaune du Cameroun peut étre estimée à environ 200 espèces, dont
plusieurs ne sont pas encore décrites. Parmi ces dernières se trouve l’Hyperolius qui
fera l’objet de cet article. Reconnu comme espèce distincte et probablement nouvelle
il y a déjà une trentaine d’années, il n’a été jusqu’ici mentionné que sous le nom
d’«Hyperolius sp. 6» (Amiet, 1975). Grâce au travail d’ensemble que Schigtz (1999) a
consacré aux rainettes africaines, il est possible de confirmer qu’il s’agit effectivement
d’une espèce nouvelle pour la science, et apparemment endémique du sud-ouest du
Cameroun.

MATERIEL ET METHODES

Les 82 spécimens étudiés ci-après ont tous été captures de nuit par l’auteur. Les
mâles ont, en général, été repérés grâce à leur activité vocale; les femelles ont été
trouvées en compagnie des mâles, deux étant même en amplexus. Il s’agit donc dans
tous les cas d’individus adultes. Un élevage à partir d’une ponte produite par l’un des
couples a permis de connaître la livrée juvénile.

Dans plusieurs stations où l’espèce a été entendue, il n’y a pas eu de prélè-
vement de spécimens. Une vérification de visu a été faite dans les sites où elle pouvait
être syntopique d’Hyperolius riggenbachi (Nieden, 1910), car les cris des deux espèces
se ressemblent beaucoup.

Manuscrit accepté le 08.12.2003

568 J.-L. AMIET

Les données faunistiques de la carte de la Fig. 3 incluent les relevés auditifs.
Pour plus de précisions sur les méthodes de relevé bio-acoustique et de cartographie,
on pourra se reporter à Amiet (1983). Dans le tableau 1, les coordonnées géographiques
sont celles des mailles (carrés de 10 minutes de côté) où les spécimens ont été capturés.

TABLEAU 1. Principales données relatives au matériel d’H. camerunensis étudié. Explications

dans le texte sous «Matériel étudié».

Nombre de Numéros de Localité Coordonnées Altitude Date de
spécimens collection de la maille capture

9 M2, 1 M3 71.332-341 Ebone 9°50'-10° x 4°50'-5° 450-500 30.11.71
2M2,2M3,1F 71.508-509;

71.539 ; Mwandong 9°40'-50' x 4°50'-5° = 1200 29.IV.71

71.604-605
6 M2, 1F 71.719-725 Santchou 9°50'-10° x 5°10'-20' = 700 24.V.71
1 M3 71.758 Fotabong 9°50'-10° x 5°30'-40' 1050-1100 30.V.71
1 M3 71.849 Mwakoumel 9°40'-50' x 4°50'-5° = 1200 26.V.71
4M2 71.789-792 Mwakoumel 9°40'-50' x 4°50'-5° = 1200 30.V.71
3M2 71.1082-1084 Santchou 9°50'-10° x 5°10'-20' = 700 22.IX.71
1M2 72.089 Ebone 9°50'-10° x 4°50'-5° 450-500 29.1.72
4 M2, 4 M3 72.223-230 Manengolé 9°50'-10° x 4°50'-5° = 600 23.11.72
3M3 73.033-035 Mwakoumel 9°40'-50' x 4°50'-5° = 1200 23.1.73
2 M2, 2M3 73.542-545 Bakwat 9°50'-10° x 4°50'-5° = 850 26.X11.73
2M2,1M3,1F 75.089-90;

75.153-154 Mankwat 9°50'-10° x 5°00'-10' = 950 23.11.75
5 M2, 1 M3 75.231-233;

75.245-247 Falaise des Mbos 10°00'-10' x 5°20'-30' = 900 25.V.75
6 M2, 1F 79.068-074 Magba 11°10'-20' x 5°50'-6° =700 23.V1.79
1 M2, IF 79.099-100 Ndikitolé 10°40'-50' x 4°40'-50' = 750 28.V1.79
2M2,1M3,1F 80.078-80;

80.082 Mélong II 9°50'-10° x 5°00'-10' = 850 05.V11.80
1F 80.081 Ekom Nkam 10°00'-10' x 5°00'-10' = 600 07.V11.80
1F 81.032 Falaise de Foréké 10°00'-10' x 5°20'-30' = 1100 20.11.81
1 M3,1F 81.036-37 Falaise de Foréké 10°00'-10' x 5°20'-30' = 850 20.11.81
2 M2 82.004-005 Bodipo 10°40'-50' x 4°20'-30' = 700 28.1.82
3 M2 86.021-023 Makouopnsap 11°00'-10' x 5°10'-20' 600-650 18.111.86
3 M2 86.029-031 Yambeta 10°50'-11° x 4°40'-50' = 700 16.X.86

Les mensurations ont été faites au pied à coulisse, au dixième de mm, pour la
longueur museau-anus (L), la largeur de la tête, en arrière des yeux (T) et la longueur
de la jambe, du milieu du genou au milieu du talon (J). Pour la téte, les rapports
morphometriques ont été établis à partir de dessins à la chambre claire de la téte, vue
de dessus, de 10 individus pris au hasard. Les rapports sont calculés à partir des valeurs
suivantes: — espace entre les narines (IN), — distance entre la narine et la commissure
palpébrale antérieure (NO), — distance entre les deux commissures palpébrales (OE), —
largeur de la téte en arrière des yeux (T).

La description des livrées, particulièrement importante pour les Hyperolius, est
basée sur les notes de terrain, les spécimens préservés et, surtout, 64 diapositives de
nombreux spécimens photographiés sur le vif.

Les vocalisations de trois individus ont été enregistrées sur le terrain, dans des
localités différentes (magnétophone Uher Report 4000), et ont fait l’objet de sona-

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 569

grammes réalisés au Laboratoire de Zoologie de l’Université de Copenhague grâce à
l’obligeance de A. Schigtz.

Les noms des auteurs et les dates de publication des noms d’espèces cités dans
le présent travail n’ont pas été mentionnés dans le texte. On les trouvera ci-après, les
genres et les espèces étant classés par ordre alphabétique: Afrixalus dorsalis (Peters,
1875), A. paradorsalis Perret, 1960, Amnirana albolabris (Hallowell, 1856), Bufo
maculatus Hallowell, 1855, Chiromantis rufescens (Giinther, 1868), Hyperolius acuti-
rostris Peters, 1875, H. adametzi Ahl, 1931, H. balfouri (Werner, 1907), H. bolifambae
Mertens, 1938, H. bopeleti Amiet, 1979, H. cinnamomeoventris Bocage, 1866, H.
concolor (Hallowell, 1844), H. fusciventris Peters, 1876, A. guttulatus Güther, 1858,
H. kuligae Mertens, 1940, A. ocellatus Günther, 1858, H. pardalis Laurent, 1947, A.
phantasticus (Boulenger, 1899), H. platyceps (Boulenger, 1900), H. riggenbachi
(Nieden, 1910), H. tuberculatus Mocquard, 1897, H. viridiflavus (Duméril & Bibron,
1841), Leptopelis aubryi (Duméril, 1856), L. calcaratus (Boulenger, 1906), L. notatus
(Buchholz & Peters, 1875).

Hyperolius camerunensis sp. n.

MATERIEL ETUDIE

Le tableau 1 donne pour chacun des prélèvements, classés par ordre chronologique, les
informations suivantes: — nombre de spécimens (M2: mâles en livrée de phase II, M3: mâles en
livrée de phase III, F: femelles); — numéros de collection de l’auteur; — localité d’origine, telle
qu’elle figure sur les étiquettes de collection (il s’agit en général de petits villages relativement
proches des points de capture effectifs des spécimens; les localités Mwakoumel et Mwandong,
ainsi que Falaise des Mbos et Falaise de Foréké, sont en fait synonymes); — coordonnées géo-
graphiques de la maille de 10 minutes de côté où ont été trouvés les spécimens; — altitude du site
de capture; — date de capture.

Holotype: mâle de Bakwat, 26.XII.1975, déposé au Musée d'histoire naturelle de
Genève sous le numéro MHNG 2643.57 (n° Amiet: 73.544). Principaux caractères mentionnés
ci-après.

Paratypes: tous les autres spécimens du tableau 1. Vingt d’entre eux sont déposés au
Musée de Genève sous les numéros MHNG 2643.58 à 82, vingt autres au Muséum national
d'Histoire naturelle de Paris sous les numéros MNHN 2003.0040 à 2003.0064.

DIAGNOSE

Hyperolius de taille moyenne, à museau court et tégument lisse, proche
d’H. riggenbachi, dont il diffère par sa tête plus large, sa taille plus faible et, surtout,
par sa livrée en phase III: — dos et flancs, ou seulement ces derniers, ornés de taches
rouge vif, la plupart sub-circulaires, entourées d’un liséré clair, — face ventrale orange
dans sa partie centrale, noire avec de gros points blancs sur sa périphérie. Espèce
parasylvicole endémique du sud-ouest du Cameroun.

ETYMOLOGIE

Le nom spécifique fait allusion au fait que cette espèce est propre au territoire
camerounais.

DESCRIPTION
1 — Principaux caractères de l’holotype

Mensurations (en mm) et proportions (en %): L: 24; T: 9,1; avant-bras, du coude
au poignet; 5,8; cuisse; 10,2; J: 12,1; pied, tarse compris: 17,8; T/L: 37,9; J/L: 50,4;

570 J.-L. AMIET

IN/T: 28,1; NO/IN; 72,7; NO/OE: 60,5; bord postérieur du disque gulaire/largeur de la
mandibule au méme niveau: 37,4.

Livrée: livrée diurne de phase III. En alcool: face supérieure de la tête, du corps,
des avant-bras et des jambes, et tranche externe des pieds de teinte gris clair, avec des
macules de forme arrondie ou grossièrement bacilliforme de teinte rouge grenat, plus
foncé sur le pourtour, et entourées d’un mince liséré jaune clair; flancs de méme cou-
leur, les taches étirées longitudinalement et s’étendant jusqu’à la région loréale.
Dessous des régions pectorale et abdominale blanchätre, le tégument opaque; cuisses,
pieds et mains teintés d’orangé très pâle. Pigmentation dorsale et latérale sur le vif: voir
photo d de la planche I.

2 — Habitus, proportions

Espèce de taille moyenne pour le genre (moyenne pour 73 mâles: 2,4 cm, pour
9 femelles: 3 cm), de forme plutôt ramassée et aplatie dorso-ventralement, la tête rela-
tivement large, un peu plus chez le mâle (moyenne T/L: 37,51%) que chez la femelle
(36,08%). Par leur habitus, les mâles ressemblent à ceux d’H. ocellatus (plus petits) ou
d’H. riggenbachi (plus grands et un peu plus élancés) et les femelles à celles d’H.
pardalis.

Si on compare les mâles d’H. camerunensis à ceux d’H. riggenbachi, très
ressemblants en phase II, on constate (Tab. 2 et histogrammes de la Fig. 1) que: — 1°)
H. camerunensis est plus petit qu’H. riggenbachi, avec une moyenne de 23,94 mm
pour le premier et de 27,76 pour le second, — 2°) la tête est proportionnellement plus
large chez camerunensis que chez riggenbachi, avec des rapports T/L de 37,5% et
32,5% respectivement.

Sur la face supérieure du corps et des membres, le tégument est lisse, d’aspect
satiné; sur la face ventrale, il est plus mince et transparent, sauf chez les individus en
phase III (voir ci-après), chez lesquels il est opaque. Il n’y a aucune trace de spinosite.
Les seuls reliefs tégumentaires correspondent à quelques petites excroisssances rictales.

Museau très court, obtus: vu en plan, il apparaît tronqué ou très légèrement
arqué entre les narines; la distance entre la commissure palpébrale antérieure et la
narine (NO) représente entre les deux tiers et les trois quarts du diamètre de l’ceil (OE),
et des 2/3 aux 4/5 de la distance entre les narines (Tab. 2). Narines largement écartées,
séparées par un espace qui atteint presque le tiers de la largeur de la téte en arrière des
yeux (Tab. 2). Canthus très largement arrondi. Tympans non différenciés, comme chez
les autres Hyperolius, cependant perceptible chez certains spécimens fixés en Eclairage
rasant.

Membres sans particularités notables. Doigts (Fig. 2) avec des disques ter-
rninaux bien développés, un peu plus larges que longs aux doigts 2, 3 et 4. Ils sont
réunis par une petite palmure ne dépassant guère, au doigt 3, le premier tubercule sous-
articulaire, mais se prolongeant le long des doigts jusqu’aux disques. Les disques des
orteils (Fig. 2) sont aussi larges (1 et 5) ou un peu plus larges (2, 3 et 4) que longs. Le
développement de la palmure pédieuse est semblable à celui de nombreux autres
Hyperolius: de part et d’autre de l’orteil 4, elle s’étend jusqu’au tubercule sous-articu-
laire distal mais se poursuit en une étroite frange jusqu’au disque.

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 571

TABLEAU 2. Quelques mensurations et rapports morphométriques chez H. camerunensis et, pour
comparaison, chez l’espèce voisine A. riggenbachi. Abréviations: voir «Matériel et méthodes».

L m J DIE "I SINTANENOIN (NO/0E
È N 73 73 73 73 73 10 10 10
S max. 280 108 148 40,88 54,89 32,00 91,20 75,60
S min. 210 77 10200 33,46, 45:65, 28.10.28. 772207 759,60
È moyen 23940 1899 1228 371,51 5123 2896 77,40) 67.23
sl &c.-t. 15 5,8 8,6 kam 222 132 7,08 5,4
<<
2 | x N 30 30 30 30 30 10 10 10
È max. 308 102 14853642 52,38 33337 | 105,88 178126
S min. 240 80 1149 2729,82) 45.125 25,207 77000. 55.81
= Moya 217.6 3037 13620 32.54 49109152929 1280,45 | 6591
= Gero Se 6,4 8,6 STA 712690 3:07 10,66 6,04
È N 9 9 9 9 9
S max 324 119 172 3710453555
5 min. 270 99 130 33,89 46,42
ù moy. 293,4 1059 148,6 36,08 50,58
Sx éc.-t. 16,5 6,9 27 132,56
= a
2 | $ N 6 6 6 6 6
à max. 380 126 200 33,06 52,91
à min. 285 97 1390052758 47.94,
È moy. 336 HID 2 1168:32 33710850
= éc.-t. — 38,4 11,8 235000073 (2510:

3 — Caractères sexuels secondaires

Taille. Les mâles sont plus petits que les femelles, la taille moyenne des
premiers (23,94 mm) représentant environ 80% de celle des secondes (29,34 mm),
mais il y a un léger chevauchement des valeurs de la longueur museau-anus: le plus
grand mâle mesure 28 mm et la plus petite femelle 27 mm.

Glande (ou disque) gulaire. De forme trapézoïdale ou sub-rectangulaire, son
extension transversale est très variable: la largeur de son bord postérieur peut repré-
senter de 37% à 69% de celle de la mandibule, mesurée au même niveau (mensurations
effectuées sur 6 mâles pris au hasard).

Glandes tégumentaires. Dans la région pectorale et sur la face interne de
l’avant-bras, le tégument est, chez les mâles, un peu épaissi et piqueté de pores. Ces
aires glandulaires, qui existent chez divers autres Hyperolius, ne sont pas toujours
faciles à observer.

4 — Livrée

Bien mieux que les critères de forme et de proportions, les caractéristiques de
la livrée permettent de séparer cette espèce des espèces voisines.

H. camerunensis peut se présenter sous des livrées très différentes, qui se
succèdent dans le temps et correspondent donc à des phases, et non à des morphes.

572 J.-L. AMIET

BH. riggenbachi |
OX. camerunensis |

Pourcentages

210-229 230-249 250-269 270-289 290-309
Classes L

Pourcentages

29-30,9 31-329 33-349 35-369 37-38,9 39-409
Classes T/L

Fic. 1

Variation de la taille (L) et du rapport largeur de la tête / longueur museau-anus (T/L) en pour-
centage du nombre d'individus chez 30 mâles d’H. riggenbachi et 73 mâles d’H. camerunensis.

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 973

5 mm

FIG. 2

Face palmaire de la main, à gauche, et face plantaire du pied, à droite, chez un individu
d’H. camerunensis (n° 79.073).

Trois phases peuvent être reconnues, évidemment reliées par des intermédiaires: —
phase 1: livrée des juvéniles néo-métamorphosés; — phase IT: livrée arborée, sur les
sites de reproduction, par une proportion importante des mâles, de l’ordre de 75%; —
phase III: dans les mêmes conditions, elle correspond à la livrée des autres mâles et de
la totalité des femelles. La livrée varie aussi en fonction du cycle nycthéméral (en fait,
avec la lumière et l’hygrométrie), et il y a donc, pour toutes les phases et leurs inter-
mediaires, une «livrée diurne» et une «livrée nocturne». Cette variation affecte essen-
tiellement la face dorsale; elle entraîne une modification des couleurs, en particulier
des contrastes, mais non des dessins (constituant le «patron» ou «pattern»).

Phase I. — Le tégument de la face dorsale, légèrement transparent, est d’un vert
jaunätre passant au turquoise vers l’arrière, assez régulièrement piqueté de chro-
matophores brun-noir. Deux fines lignes blanches latéro-dorsales partent de l’aine,
passent au-dessus de l’ceil, se prolongent sur le canthus et se rejoignent au bout du
museau (PI. I, k). En livrée nocturne, les lignes claires ressortent moins sur le fond et
les chromatophores ont tendance à former des taches nébuleuses plus foncées qui
parsèment la face dorsale. Ventralement, le tégument est transparent, vert teinté de
turquoise, et il n’y a aucune trace de pigment jaune ou rouge.

574 J.-L. AMIET

Phase II. — En livrée diurne, toute la face supérieure est d’un vert pâle, plus ou
moins nuancé de jaune ou de turquoise. Les lignes latéro-dorsales sont relativement
plus larges que dans la phase précédente (de l’ordre de 1,5 à 2 mm) et d’un blanc pur
(pl. I, a).

En livrée nocturne, le contraste entre le fond et les bandes s’atténue. La
pigmentation foncière, plus ou moins envahie de taches sombres, est alors d’un vert
terne, pouvant tirer sur le brun clair. Les bandes latéro-dorsales deviennent d’un brun
rougeätre ferrugineux et sont bordées par des concentrations de chromatophores
brunätres. C’est le cas aussi pour les canthus, ornés de bandes brunes plus ou moins
larges, pouvant persister longtemps après l’éclaircissement de la livrée nocturne (PI. I,
betc).

Sous les membres et la gorge, le tégument est transparent, vert clair teinté de
jaune ou de turquoise. L’aspect du reste de la face ventrale varie suivant le degré d’évo-
lution de la livrée. Dans un premier stade, le tégument de la poitrine et de l’abdomen,
opaque, est d’un blanc pur sauf, éventuellement, une «réserve» transparente médio-
abdominale (PI. I, j, individus de gauche). A un stade un peu plus avancé, la zone trans-
parente disparaît alors que la face ventrale devient jaune citron, la pigmentation
blanche ne subsistant qu’en périphérie, dans la région latéro-ventrale, où elle s’étend
sur la moitié inférieure des flancs. Le disque gulaire est jaunätre. Les doigts et les
orteils sont vert-jaune avec, chez certains individus, un léger lavis rouge.

Phase III. — La livrée, très différente des précédentes, est caractérisée par des
macules de couleur rouge sur le dos et les flancs, ou seulement sur ces derniers. Ce
système chromatique permet de séparer d’emblée H. camerunensis des espèces
voisines.

Livree diurne. Toutes les parties apparentes en position de repos ont une teinte
foncière claire, allant du gris perle au bistre verdätre, en passant par le jaune-beige, le
gris-beige ou le bistre. Sur ce fond ressortent des taches bien circonscrites, arrondies ou,
plus rarement, étirées; de couleur rouge vif, elles sont irrégulièrement bordées de
noiràtre et entourées d’un mince liséré clair. Le nombre, la taille et la répartition de ces
taches montrent une importante variabilité individuelle entre les deux extrêmes
suivants: — taches nombreuses mais relativement petites sur tout le dos ainsi que sur les
avant-bras et les jambes, — ou taches localisées sur les flancs et la région loréale, de
forme plutôt étirée que circulaire, le restant de la face dorsale étant uni (Pl. I, dà g et i).

Livrée nocturne. Le fond va du gris brunätre au brun chocolat, les macules sont
d’un rouge grenat et les lisérés clairs plus apparents (PI. I, h).

Face ventrale. Elle est caractérisée par le contraste entre deux types de pigmen-
tation différents: — une pigmentation hétérogène, constituée de gros points blancs sur
un fond noir ou noiràtre, — une pigmentation unie, orange vif ou rouge corail. La
première s’étend sur le pourtour de l’abdomen, d’où elle remonte sur les flancs, sur la
région gulaire, qu’elle envahit plus ou moins, et sur le dessous des jambes et des
membres antérieurs. La pigmentation orange couvre tout l’abdomen, une plus ou
moins grande partie de la gorge et le dessous des cuisses. La pigmentation rouge est
propre aux pieds et aux mains. Les femelles se distinguent légèrement des mäles par
leur maculation plus contrastée et mieux circonscrite, ainsi que par leurs couleurs
encore plus éclatantes (PI. I, j, individus de droite).

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 575

PLANCHE I. Variation du chromatisme chez H. camerunensis. De aä c, mäles en phase II (a:
livrée diurne, b et c: livrée nocturne); de d ah, mâles en phase III (d: holotype, h: livrée nocturne
très foncée); i: femelle; j: face ventrale de deux mâles en phase II (à gauche) et d’un mâle et une
femelle en phase III (à droite); k: jeune individu en phase I. Tous les individus sont à la méme
échelle, sauf ceux des figures j et k.

576 J.-L. AMIET

Livrées intermédiaires. — Le passage de la phase I (juvéniles obtenus par
élevage à partir d’une ponte) à la phase II (mäles adultes trouvés sur les sites de
reproduction) n’est pas connu. On peut supposer que l’un des deux caractères propres
à la phase II, l’opacification du tégument ventral, se produit tardivement puisque chez
certains individus elle n’est pas encore complète quand ils arrivent sur les sites de
reproduction.

Aucun des 73 mäles capturés ne présente de livrée réellement intermédiaire
entre les phases II et III. Deux individus montrent toutefois quelques petites taches
noirätres irrégulières sur les flancs, amorces du dessin noir et blanc beaucoup plus
étendu de la phase III; chez l’un, les bandes latéro-dorsales sont encore perceptibles,
alors qu’elles ont disparu chez l’autre. Un caractère de transition plus fréquent chez les
mâles en phase II est la présence d’un lavis rouge sur les doigts et les orteils, annonçant
la coloration rouge corail caractéristique de la phase III.

On peut déduire des observations précédentes que le développement de la livrée
de phase III a lieu après la première période de reproduction, mais il subsiste deux
interrogations sur les modalités de ce processus: —1°) la transition est-elle rapide (c’est
le cas chez H. concolor, où le passage à la phase verte unie se fait en quelques heures:
obs. pers.), ou se déroule-t-elle progressivement pendant les quelques mois qui
séparent la première et la seconde périodes de reproduction; — 2°) tous les mâles sont-
ils susceptibles de passer à la phase III, ou bien certains restent-ils «bloqués» en phase
II? Seuls des élevages de longue durée permettraient de répondre à ces questions.

DISTRIBUTION ET ECOLOGIE

La présence d’H. camerunensis a été relevée dans 24 stations qui se répartissent
entre 17 carrés de 10 minutes de côté. Si on compare le second chiffre à celui
qu’atteignent des espèces affines (A. ocellatus: 105, A. bolifambae: 40, H. pardalis:
38) ou se reproduisant dans des biotopes similaires (A. tuberculatus: 80, H platyceps:
59, H. cinnamomeoventris: 53, H. kuligae: 41), on pourrait considérer H. cameru-
nensis comme une espèce plutôt rare.

Il faut toutefois relativiser ce point de vue en tenant compte de son extension
géographique. Comme le montre la carte de la Fig. 3, celle-ci se limite en fait à une
faible portion du territoire camerounais, à tel point qu’H. camerunensis y est, avec A.
adametzi, l’espèce la moins largement répandue du genre (exception faite des avancées
d’espèces «occidentales»). Dans les limites de ce domaine restreint, l’espèce apparaît
plutôt comme commune, les carrés où elle a été trouvée représentant environ 60% des
carrés prospectés.

Son aire de répartition dessine une bande grossièrement en forme de V, d’une
centaine de km de largeur au maximum, recouvrant les piedmonts et les basses pentes
des massifs avancés de la Dorsale camerounaise (monts Koupé, Nlonako et
Manengouba) ainsi que la vallée du fleuve Mbam, affluent de la Sanaga coulant à l’est
des plateaux bamiléké et bamoun. Il est possible qu’elle s’étende un peu plus vers le
nord, aussi bien pour sa branche ouest que pour sa branche est, car la batrachofaune de
ces secteurs est mal connue; en revanche, une plus grande extension vers l’ouest et le
sud, régions densément prospectées, est improbable.

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN SI

FIG. 3

Carte du Cameroun au sud du 8° paralléle montrant la distribution d’H. camerunensis et de deux
especes apparentees, H. riggenbachi et H. pardalis. La trame correspond a des carrés de 10
minutes sexagésimales de côté. La courbe de niveau de 900 m ainsi que le cours du fleuve
Sanaga et de son affluent le Mbam ont été figures.

Cette aire présente la particularité de recouper des territoires climatiques et des
types de végétation très différents. Dans sa partie ouest et sud-ouest, la pluviosité est
très élevée, avec une moyenne annuelle de précipitations atteignant au moins deux
metres, et souvent beaucoup plus. La végétation, luxuriante, a pour climax la forét sem-
pervirente de type atlantique (ou biafréen). Au contraire, la partie sud-est et est corre-
spond à une des régions les moins arrosées du Cameroun méridional, avec des
moyennes annuelles pouvant s’abaisser jusqu’à 1,5 m. Elle est couverte de savanes et
de galeries ou massifs de forêt semidécidue (ou mésophile), qui représente le climax.

L’amplitude d’extension verticale d’H. camerunensis est d’un peu moins de
1000 m: les stations les plus basses, entre Nkongsamba et Yabassi, se situent vers
450 m, les plus élevées, sur les pentes sud du plateau bamiléké («falaise des Mbos»)
atteignent 1200 m. L’espéce est ainsi représentée dans environ les deux-tiers supérieurs
de l’étage planitiaire et dans la frange inférieure de l’étage submontagnard (sensu
Letouzey, 1968). H. camerunensis n’est donc pas un orobionte, mais ce n’est pas non
plus une espèce planitiaire au sens strict puisqu'il manque dans la plaine littorale et le
bassin de Mamfé. Ce type de distribution, illustré par le transect de la fig. 4, est
exceptionnel dans la batrachofaune camerounaise.

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NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 579

H. camerunensis est une espèce «parasylvicole», c’est-à-dire inféodée aux
milieux secondarisés résultant de la destruction ou de la dégradation de la forét, en
général sous l’action de l’homme (cf. Amiet, 1989). En période de reproduction, les
mäles peuvent étre entendus dans des brousses secondaires plutöt denses, constituées
de grandes Graminées, de Marantacées, de Zingibéracées, d’arbustes héliophiles et de
rejets d’arbres forestiers. De tels milieux sont particulièrement fréquents là où les
routes et pistes recoupent des cours d’eau. Dans la partie orientale de son aire, où la
forêt laisse souvent la place à de grandes étendues de savanes, H. camerunensis évite
celles-ci et se cantonne dans les zones dégradées des galeries forestières. A sa limite
supérieure d’extension, dans la «falaise des Mbos», il se mêle à A. riggenbachi,
«descendu» du plateau bamiléké. Dans ce site, la ressemblance des livrées de phase II
et des vocalisations rend la distinction des mâles difficile sur le terrain, mais la
syntopie des deux espèces a été confirmée par la capture de femelles qui, elles, ne
posent pas de problème d’identification.

Bien que les postes de chant des mâles ne soient jamais bien loin de petites
rivières ou de ruisseaux, il est probable que le développement des têtards, comme chez
tous les Hyperolius, se déroule dans des collections d’eau stagnante ou faiblement
renouvelée qui en sont proches.

La distribution d’H. camerunensis ne correspond à aucun des «aréotypes» qui
peuvent être reconnus dans la batrachofaune camerounaise. Sa localisation à l’ouest de
la Sanaga suggère une appartenance à 1’«élément ouest-camerounais» (Amiet, 1975),
ensemble d’une douzaine d’espèces endémiques concentrées dans le sud-ouest du
Cameroun (certaines peuvent déborder légèrement sur le territoire nigérian, dans les
Rumpi Hills). La partie occidentale de son aire de répartition coïncide effectivement
avec le «centre de gravité» de l’élément ouest-camerounais, mais il s’en distingue très
nettement par l’importante extension de son domaine vers le nord-est et l’est, sans
équivalent chez les autres espèces de cet ensemble. Un autre élément de divergence
réside dans son écologie parasylvicole, car toutes les espèces de l’élément ouest-
camerounais sont fondamentalement sylvicoles. De façon plus générale, on relèvera le
caractère paradoxal que présente une distribution aussi limitée, alors que les Anoures
parasylvicoles sont d’ordinaire largement répandus (y compris hors du Cameroun) et
que l’espèce est assez euryécique pour prospérer aussi bien dans des régions hyper-
humides (avant-postes de la Dorsale camerounaise) que dans des régions relativement
sèches (vallée du Mbam).

CYCLE ANNUEL D'ACTIVITÉ VOCALE, PHONOCENOSES ET VOCALISATIONS

Le graphique de la figure 5 montre la distribution dans le temps des relevés
durant lesquels H. camerunensis a été entendu. Les enseignements à en tirer sont
limités car les prospections dans la zone de distribution de l’espèce ont été trop inéga-
lement réparties dans l’année. Un fait s’en dégage toutefois: l’activité vocale peut se
poursuivre tout au long de l’année, avec probablement un maximum bien marqué en
début de saison des pluies, c’est-à-dire de mars à mai suivant les secteurs. Ce cycle
correspond tout à fait au schéma général, semi-opportuniste, déjà décrit pour les
espèces parasylvicoles (Amiet, 1989). Dans les zones climatiques où l’espèce est

580 J.-L. AMIET

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Distribution par mois des relevés au cours desquels a été notée l’activité vocale d’H. cameru-
nensis.

représentée, les petites saisons séches (juillet-aoüt) sont atténuées, voire nulles, et il n’y
a alors qu’une seule saison des pluies: il serait intéressant de savoir s’il existe, comme
dans les régions a climat équatorial typique à quatre saisons, un second maximum, et
quand il se situe.

Le terme de «phonocénose» a été proposé récemment pour désigner «des
ensembles d’espèces qui exercent leur activité vocale... pendant la méme période de
l’année et dans des sites présentant des caractéristiques écologiques similaires»
(Amiet, 2001). Pour la même raison que précédemment, la mise en évidence des
phonocénoses auxquelles participe H. camerunensis est fortement biaisée. Quelques
points peuvent cependant être dégagés de la comparaison des 30 relevés (effectués
dans 24 sites) au cours desquels H. camerunensis a été entendu.

— L'espèce la plus fréquemment associée à H. camerunensis a été une autre
rainette des formations secondaires boisées: Afrixalus paradorsalis (24 relevés sur 30).

— Vient ensuite Leptopelis calcaratus (13/30), espèce forestière qui déborde
souvent sur les formations secondaires quand elles jouxtent la forêt et sont suffi-
samment riches en buissons et petits arbres.

— Parmi les autres espèces préférant le farmbush arboré, ont été aussi relevés
Leptopelis aubryi (9/30), Hyperolius ocellatus (7/30), plus forestier que parasylvicole,
Chiromantis rufescens (5/30) et Leptopelis notatus (6/30), qui recherche surtout les
raphiales et galeries marécageuses.

— Parmi les espèces les plus typiques des formations secondaires, on note encore
la fréquence de Bufo maculatus (10/30), Amnirana albolabris (10/30) et Hyperolius
tuberculatus (7/30).

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 581

— Dans la partie est de l’aire d’H. camerunensis, ainsi que dans la plaine des
Mbos, l’extension des savanes se traduit par la présence d’espèces de farmbush riche
en Graminées (Afrixalus dorsalis: 5/30), voire d’espèces savanicoles (Hyperolius
balfouri: 4/30).

Les appels sont de brefs trilles, rapidement répétés, dont les sonagrammes de la
Fig. 6a et b montrent la structure. Celle-ci, dans ses grandes lignes, est semblable à
celle des appels des autres espèces du méme groupe (voir ci-après), en particulier
H. riggenbachi (Fig. 6c et d). La comparaison des sonagrammes d’individus des deux
espèces (y compris ceux qui n’ont pas été reproduits ici) montre que les trilles
d’H. camerunensis sont émis dans la bande de 4 à 5 kHz, alors que ceux d’H. riggen-
bachi se situent dans celle de 3 à 4 kHz. Effectivement, quand les deux espèces sont
réunies sur un méme site de reproduction, leurs appels, malgré leur ressemblance,
peuvent être distingués par la tonalité plus aiguë et plus claire d’H. camerunensis, s’op-
posant à celle, plus grave et comme «räpeuse», d’H. riggenbachi.

Aux trilles peuvent se substituer des «djuitt», notes brèves et appuyées (Fig. 6e),
qui paraissent émises lorsqu’un mäle pergoit un autre individu dans son champ visuel.
Ces cris brefs s’entendent surtout au crépuscule, quand les mäles prennent possession
de leur poste de chant, source fréquente de rivalité entre individus proches.

AFFINITÉS

Parmi les 24 espèces d’Ayperolius de la faune camerounaise, quatre doivent
être rapprochées d’H. camerunensis: H. ocellatus, H. pardalis, H. riggenbachi et
H. bolifambae. Toutes partagent en effet, de façon exclusive, la combinaison de carac-
tères suivante: — 1°) livrée en phase I verte avec de fines lignes latéro-dorsales claires;
— 2°) absence totale de patron MD, c’est-à-dire de macules dorsales symétriques autres
que les bandes latéro-dorsales (Amiet, 1978); — 3°) livrée en phase III vivement
colorée, aussi bien sur la face ventrale que sur la face dorsale (sauf, pour ce dernier
point, chez H. bolifambae); — 4°) vocalisations constituées de trilles; — 5°) écologie
parasylvicole, jamais savanicole; — 6°) distribution limitée à l’Afrique centrale.

Le second caractère est important, car il sépare radicalement les espèces de ce
groupe de celles, relativement nombreuses dans la batrachofaune camerounaise, qui
arborent un patron de type MD, soit dans une livrée unique (comme A. acutirostris ou
H. bopeleti, par exemple), soit dans une livrée correspondant à une morphe (A. platy-
ceps ou H. adametzi, par exemple) ou à une phase (H. concolor). Le troisième carac-
tere pourrait suggérer un rapprochement avec les nombreuses formes du «groupe viri-
diflavus», sensu Schigtz (1999), souvent pourvues, au moins sur la face dorsale, d’une
livree très colorée, parfois considérée comme «aposématique». Il ne s’agit là que d’une
convergence car ce groupe, comme l’a souligné Schigtz (op. cit.), possede de nom-
breux caractères propres qui le différencient de tous les autres Hyperolius, y compris
les espèces mentionnées ci-dessus.

En revanche, les caractères précédents, sauf le dernier, se retrouvent chez une
espece polytypique à distribution occidentale, atteignant l’ouest du Cameroun: A. fus-
civentris. Comme un phénomène de convergence portant sur l’ensemble de ces carac-
tères est improbable, on peut considérer que H. fusciventris est phylétiquement proche

582 J.-L. AMIET

FIG. 6
Sonagrammes de deux individus d’H. camerunensis (a et b: cri normal, e: cri court) et, pour
comparaison, de deux individus d’H. riggenbachi (c: H. r. hieroglyphicus, d: H. r. riggenbachi).
a: Mwakoumel, 26.V.71, b: Bakwat, 26.XII.73, c: Fongo-Tongo, 28.IV.71, d: Ribao, 22.XII.71,
e: Bakwat, 26.XII.73 (même individu que b). Sonagrammes A. Schigtz et Laboratoire de Zoo-
logie de l’Université de Copenhague.

NOUVELLE ESPECE D’HYPEROLIUS DU CAMEROUN 583

des espèces discutées ici. Des affinités, plus lointaines, paraissent aussi exister avec
deux autres espèces représentées dans la faune camerounaise, H. guttulatus et H. phan-
tasticus.

Si, en s’appuyant sur les caractères morphologiques, on essaye de concevoir les
relations phylétiques à l’intérieur du groupe formé par les espèces affines
d’H. camerunensis, on peut admettre qu’ H. ocellatus et H. bolifambae sont relati-
vement éloignés des trois autres espèces. Le premier s’en sépare, entre autre, par l’évo-
lution des bandes latéro-dorsales, qui fusionnent sur le museau en formant un triangle
caractéristique, et aussi par l’habitus des femelles, plus «longiligne» que chez les
autres espèces. Le second se distingue par son habitus (museau plus pointu) et sa livrée
dorsale en phase III dépourvue de maculation foncée, arborée par tous les mâles
parvenus au stade reproducteur. Parmi les trois espèces restantes, H. camerunensis
paraît plus proche d’H. riggenbachi (similitude des livrées en phase II, de la pigmen-
tation ventrale en phase III et des vocalisations) que d’H. pardalis, avec lequel il
partage toutefois la variabilité de la maculation dorsale en phase III chez les mâles.

REMERCIEMENTS

Plusieurs collègues m’ont apporté leur aide pour le présent travail: A. Schigtz a
réalisé les sonagrammes, J.-L. Perret a relu le manuscrit et V. Mahnert m’a offert toutes
les facilités pour le publier. Qu'ils trouvent ici l’expression de ma gratitude pour leur
amicale collaboration.

RÉFÉRENCES
AMIET, J.-L. 1975. Écologie et distribution des Amphibiens Anoures de la région de
Nkongsamba (Cameroun). Annales de la Faculté des Sciences de Yaoundé 20: 33-107.

AMIET, J.-L. 1978. A propos d’Hyperolius platyceps (Boulenger), H. kuligae Mertens et
H. adametzi Ahl (Amphibiens Anoures). Annales de la Faculté des Sciences de Yaoundé
25: 221-256.

AMIET, J.-L. 1983. Un essai de cartographie des Anoures du Cameroun. Alytes 2(4): 124-146.

AMIET, J.-L. 1989. Quelques aspects de la biologie des Amphibiens Anoures du Cameroun.
Année Biologique 28(2): 73-136.

AMIET, J.-L 2001. Un nouveau Leptopelis de la zone forestière camerounaise (Amphibia, Anura,
Hyperoliidae). Alytes 19(1): 29-44.

LETOUZEY, R. 1968. Etude phytogéographique du Cameroun. Encyclopédie Biologique, LXIX.
P. Lechevalier, Paris, 508 pp.

SCHISTZ, A. 1999. Treefrogs of Africa. Chimaira, Frankfurt am Main, 350 pp.

Haba | epi. 0

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REVUE SUISSE DE ZOOLOGIE 111 (3): 585-598; septembre 2004

Contributi al riordinamento sistematico dei Peritelini paleartici
V. Istituzione di un nuovo genere e descrizione di due nuove specie
(Coleoptera, Curculionidae, Entiminae)

Helio PIEROTTI! & Cesare BELLO2
I strada di Selvana 1, I-31100 Treviso (Italia).
2 via Vespucci 11/a, I-31033 Castelfranco Veneto (Italia).

Contributions towards a taxonomic rearrangement of Palaearctic
Peritelini. V. Erection of a new genus and description of two new species
(Coleoptera, Curculionidae, Entiminae). - The authors diagnose a new
genus of Palaearctic Peritelini: Heteromeiropsis gen.n., for Heteromeira
zariquieyi Solari, 1954 (comb.n.), and describe two new species of this
genus. A dichotomic key as well as illustrations of their genital apparatuses
are also provided.

Keywords: Coleoptera - Curculionidae - Peritelini - Heteromeiropsis
gen. n. - taxonomy.

INTRODUZIONE

Heteromeira è un genere creato da Solari (1954) con specie-tipo Meira varie-
gata A. & F. Solari, 1903, per raggrupparvi “tutte le specie di Meira e Pseudomeira
degli autori, le quali hanno il margine esterno dello scrobo nettamente curvato e che
per ciò non possono essere ascritte a Pseudomeira, Dolichomeira, Meira, perchè in
questi tre generi il margine esterno dello scrobo è quasi retto”.

Lo stesso autore distingueva poi Heteromeira (e Meira) da Pseudomeira e
Dolichomeira per il rostro nelle prime “diviso dal capo, almeno superiormente”. In
realtà, quest’ultimo carattere non è, tuttavia, presente in Heteromeira zariquieyi —
endemica della Spagna nord-orientale (Catalogna) e descritta da Solari nel medesimo
lavoro — mentre già la descrizione originale evidenziava che “la nuova specie ... si dis-
tingue a prima vista da tutte le specie descritte ... per l’appendice squamulata dello
scrobo tanto sviluppata, che raggiunge la fronte e così profonda da rendere fortemente
angoloso fino all’altezza della fronte il suo margine interno, (laddove) nelle altre
Heteromeira manca l’appendice dello scrobo ed il rostro è arrotondato fra la fine dello
scrobo e la fronte (per cui, in definitiva,) zariquieyi non ha affini fra le specie del
genere Heteromeira”.

Se a questo carattere esattamente rilevato da Solari (1954) si aggiungono gli
altri più oltre evidenziati, sembra giustificato proporre, per Heteromeira zariquieyi e
per le due nuove specie affini che vengono qui descritte, l’istituzione di un nuovo
genere.

Manoscritto accettato il 23.04.2004

586 H. PIEROTTI & C. BELLÒ

Sul punto, ci sembrano del resto assolutamente condivisibili le considerazioni
svolte da Dellacasa et al. (2001).
Le collezioni nelle quali sono conservati gli esemplari studiati sono indicate con
le seguenti sigle rispettive:
BEL collezione Bellò - Castelfranco Veneto (Treviso - Italia)
MHNG collezione Muséum d’histoire naturelle - Genève
PIE collezione Pierotti - Treviso (Italia)
SOL collezione Solari (in Museo Civico di Storia Naturale - Milano)
TOS collezione Torres Sala (in Museo Valenciano de Historia Natural - Valencia)

CHIAVE

Chiave per il riconoscimento del nuovo genere nell’ambito dei Peritelini europei.

1 Rostro in visione dorsale delimitato all’apice da un robusto cercine
davanti alla base di tutto il bordo anteriore dello pterigio. Protibie sul
bordo esterno distintamente allargate all’apice
OR She Da (Centricnemus, Peritelus, Simo partim)
- Rostro in visione dorsale delimitato all’apice dal bordo anteriore dello
pterigio, talora con un breve accenno di cercine sul prolungamento del
bordo laterale del clipeo. Protibie sul bordo esterno allargate, diritte o
smussate:all’apice... 7... Ne A A TE 2
2 Rostro normalmente arrotondato ai lati dell’epifront
(Meira, Gymnomorphus, Pseudomeira, Dolichomeira, Heteromeira, Meirella,
Lepretius, Leptomeira)
- Rostro appiattito dorsalmente ai lati dell’epifronte fin presso il margine
OCUIATE Re aie none QI LEE ELLES 3
3 Vertice non più alto del margine oculare. Pterigi molto sviluppati.
Staturamassiore: 9 92729 OE (Simo partim)
- Vertice distintamente più alto del margine oculare. Pterigi non o appena
SalentiStaturaminore #6 :9-52 Mme m. en Heteromeiropsis gen. n.

DESCRIZIONE DEL NUOVO GENERE

Heteromeiropsis gen. n.

Heteromeira Solari, 1954: 40 partim; Pierotti & Bellö, 1998: 97 partim; Alonso Zarazaga &

Lyal, 1999: 171 partim; Alonso-Zarazaga, 2002: 24.

Diagnosi. Specie di dimensioni medie o piccole (3,9-5,2 mm.), caratterizzate -
rispetto alle altre specie attribuite al gen. Heteromeira - da pterigi non particolarmente
salienti, sutura frontale distinta, rostro appiattito dorsalmente ai lati dell’epifronte fin
presso il margine oculare, sutura epicranica indistinta, clipeo incavato, vertice distinta-
mente piü alto del margine oculare e da sacco interno del pene munito di un’armatura
orifiziale, costituita da una placca munita di una spinula, due armature mediane ed
un’armatura genitale, quest’ultima di tipo periteloide “puro” (espressione con la quale
intendiamo fare riferimento alla struttura dell’armatura genitale comune a Peritelus
familiaris Boheman, sphaeroides Germar e ruficornis Brisout, mentre le altre

PERITELINI PALEARTICI V 587

Heteromeira ne presentano una “versione” caratteristica). Le specie appartenenti al
gen. Heteromeira, inoltre, non superano ad occidente la Liguria orientale, la Corsica e
la Sardegna.

Specie tipo. Heteromeira zariquieyi Solari, 1954.

Descrizione. Forma allungata. Vestitura dorsale costituita da squame rotondeg-
gianti, fortemente embricate, terree, unicolori o qua e là più chiare, spesso disposte a
formare una fascia longitudinale mediana sul pronoto e macchie irregolari e mal deli-
mitate sulle elitre, e da setole più o meno allungate.

Capo debolmente ipognato. Rostro breve, più o meno trasverso, in visione
dorsale delimitato all’apice dal bordo anteriore dello pterigio, appiattito dorsalmente ai
lati dell’epifronte fin presso il margine oculare; clipeo alquanto stretto, incavato e su-
tura clipeo-frontale ribordata; pterigi non particolarmente salienti. Vertice distinta-
mente più alto di quest’ultimo, più o meno sullo stesso piano dell’epifronte. Occhi sub-
arrotondati o alquanto allungati (Fig. 1.a). Foramen occipitale subtriangolare (Fig. 1.d).
Antenne con scapo discretamente robusto e funicolo ad articoli non progressivamente
ingrossati; clava col primo articolo spesso più o meno peduncolato, almeno nella 9,
munito di lunghe setole scure come gli articoli 4-7 del funicolo. Mandibole a funzio-
namento orizzontale, robuste, tozze, a parete interna concava (Fig. 2.a); mascelle a
galea modesta e lacinia bene sviluppata (Fig. 2.b); labium scutiforme (Fig. 2.c).

Pronoto per lo più subquadrato o trasverso, raramente più lungo che largo, largo
alla base più o meno quanto al margine anteriore, arrotondato ai lati.

Elitre più o meno allungate, più o meno appiattite sul dorso, talora con la
massima larghezza in corrispondenza degli omeri, più (4) o meno (9) affusolate nel-
la regione apicale. I primi due ventriti rudimentali e fusi a ridosso del metasterno; i
ventriti II-VII visibili e progressivamente poco ristretti, i primi due più sviluppati e
fusi insieme ma ancora distinti, gli altri tre mobili (Fig. 1.c).

Zampe robuste, con protibie incurvate verso l’interno avanti l’apice (Fig. 1.b)
ed onichi bene sviluppati; unghie connate.

Pene allungato (Fig. 3.b), fortemente sclerificato ai lati, discretamente sul lato
ventrale, debolmente sul dorso, con sacco interno munito di un’armatura orifiziale,
costituita da una placca munita di una spinula, due armature mediane spinuliformi e
un’armatura genitale di tipo periteloide “puro” (Figg. 7, 8, 9); temones alquanto lunghi.

Tegmen a manubrium discretamente breve (Fig. 3.a).

Spiculum ventrale con manubrium allungato e margo apicalis della lamella più
o meno profondamente inciso nel mezzo (Fig. 3.c).

Coxiti privi di stili (Fig. 3.d).

Spermateca di forma alquanto variabile, ma con cornus sempre bene sviluppato
(Fig. 6).

Derivatio nominis. Femminile, dalla somiglianza (opsis, in greco) con il genere
Heteromeira Solari, 1954.

Distribuzione geografica. Genere noto finora solo della Spagna orientale e della
Francia sud-occidentale.

Note ecologiche. Le specie assegnate a questo genere si rinvengono in lettiera
di foresta a latifoglie o ai piedi di Quercus, Olea, Pistacia.

588 H. PIEROTTI & C. BELLÒ

FIG. 1

Heteromeiropsis zariquieyi: a, capo in visione dorsale (schematico); b, protibia sinistra d
(schematico); c, ventriti (schematico); d, foramen occipitale.

TABELLA DELLE SPECIE DEL GEN. HETEROMEIROPSIS

1 Elitre anche presso l’apice con setole reclinate. Solco longitudinale
mediano dell’epifronte interrotto presso la base di questa (Spagna
centro-orientale: Valencia) MI: PIE E ee ee 3. H. valentiana sp. n.

- Elitre sul dorso con setole sollevate, presso l’apice con setole semie-
rette. Solco longitudinale mediano dell’epifronte prolungato fin sul
vertice. Propigidio 9 a margine apicale largamente arrotondato o sub-
troncato (Francia meridionale, Spagna settentrionale) .................. 2
2 Elitre piü allungate, con la massima larghezza in corrispondenza degli
omeri, nel & particolarmente affusolate nella regione apicale. Pronoto
con setole reclinate su tutta la superficie (Spagna nord-orientale:
Catalosna)t. Pisces: E 1. H. zariquieyi (Solari, 1954) comb. n.

PERITELINI PALEARTICI V 589

eet

FIG. 2

Heteromeiropsis zariquieyi: a, mandibula destra dal lato interno, in visione ventrale; b, maxilla
sinistra in visione ventrale; c, labium in visione ventrale. Scale: 0,10 mm.

= Elitre meno allungate, a lati subparalleli, non particolarmente affusolate
nella regione apicale. Pronoto sul dorso con un gruppo di setole semie-
rette presso il margine anteriore (Francia sud-occidentale: Pyrénées-
@rientales)k sh Bee az 2. A. loebli sp. n.

DESCRIZIONE DELLE SPECIE

1. Heteromeiropsis zariquieyi (Solari, 1954) comb. n.

Heteromeira zariquieyi Solari, 1954: 58; Pierotti & Bellò, 1994: 296; Pierotti & Bellò, 1998:
107; Alonso-Zarazaga, 2002: 24.

Esemplari esaminati. Gerona: Figueras (BEL, PIE, holotypus in SOL), 10 Km. w
Figueres, m. 75 (BEL, PIE), Camallera (BEL, PIE).

590 H. PIEROTTI & C. BELLÒ

Fic. 3

Heteromeiropsis zariquieyi: a, tegmen (scala: 0,50 mm); b, pene in visione laterale (scala: 0,50
mm); c, spiculum ventrale (scala: 0,50 mm); d, coxiti (scala: 0,15 mm).

Diagnosi. Specie di media statura (4,2-5,2 mm.), caratterizzata, rispetto alle
altre specie del genere, da elitre particolarmente allungate nel d, a lati subparalleli ed
apice marcatamente ogivale, da setole erette o sollevate sulle elitre ed, in particolare,
rispetto ad A. loebli, da antenne con scapo un po’ più slanciato, da pene in visione
dorsale tricuspidato nella regione apicale e da propigidio 9 a margine apicale larga-
mente arrotondato (Figg. 1, 2, 3, 4.a, 5.a, 6).

Descrizione. (Ancorchè dettagliata nel lavoro originale, riteniamo opportuno
riproporla in termini omogenei con quella delle nuove specie qui di seguito descritte).
Forma robusta, marcatamente allungata. Vestitura dorsale costituita da squame terree,
che sulle elitre lasciano visibili le strie, e da setole robuste, erette o semierette sul
vertice e sulle elitre, reclinate altrove.

PERITELINI PALEARTICI V 59]

——— |

FIG. 4

Apice del pene in visione dorsale: a, Heteromeiropsis zariquieyi; b, Heteromeiropsis loebli; c,
Heteromeiropsis valentiana. Scala: 0,15 mm.

Rostro subquadrato (6) o trasverso (¢), a lati debolmente convergenti in
avanti; fronte, epifronte e vertice percorsi da un unico solco longitudinale mediano;
pterigi poco salienti, ma distinti. Occhi subarrotondati o molto debolmente allungati,
convessi, non sporgenti oltre le guance. Vertice stretto, poco più largo della fronte.
Antenne discretamente robuste, soprattutto nella 2, con scapo più (6) o meno (9)
progressivamente ingrossato dalla base verso l’apice; funicolo con i primi due articoli
poco allungati, ornati anche di setole un po’ più larghe, appiattite, il secondo più corto
dei due seguenti presi insieme, gli articoli 4-7 moniliformi nel à , trasversi nella 9;
clava robusta con il primo articolo svasato (4 ) o peduncolato (9).

Pronoto non o molto debolmente trasverso, sul disco con punteggiatura doppia,
discretamente fitta.

Elitre non o molto debolmente arrotondate ai lati, particolarmente allungate nel
3, più brevi e larghe nella 9, con la massima larghezza in corrispondenza degli omeri,
distintamente ogivali nella regione apicale, con strie di punti ravvicinati evidenti,
ciascuno con una microsetola allungata, ed interstrie debolmente convesse.

Zampe robuste con protibie fortemente (4) o debolmente (£) incurvate verso
l’interno avanti l’apice (Fig. 1.b) e con onichi particolarmente sviluppati.

Propigidio a margine apicale troncato nel d, largamente arrotondato nella ©
(Fig. 5.a).

Pene, in visione laterale, a marcata convessità dorsale, all’apice bruscamente
incurvato (Fig. 3.b); in visione dorsale, distintamente tricuspidato nella regione apicale
(Fig. 4.a); armature del sacco interno: Fig. 7.

592 H. PIEROTTI & C. BELLÒ

FIG. 5

Propigidio 9 in visione dorsale: a, Heteromeiropsis zariquieyi; b, Heteromeiropsis loebli; c,
Heteromeiropsis valentiana. Scala: 0,50 mm

Tegmen: Fig. 3.a.

Spiculum ventrale a margo apicalis distintamente inciso nel mezzo (Fig. 3.c).

Coxiti allungati (Fig. 3.d).

Spermateca: Fig. 6.a.

Distribuzione geografica. Catalogna nord-orientale. La specie sembra diffusa,
ancorché probabilmente in modo puntiforme, in tutta la provincia di Gerona.

Località di cattura desunte dalla letteratura. Gerona: Figueras (1. cl.), Olot, Coll
de Cannes.

Note ecologiche. Raccolta in lettiera di Quercus ilex L. a Figueras ed in lettiera
di Olea europaea europaea L. a Camallera.

2. Heteromeiropsis loebli sp. n.

Serie tipica. Holotypus d: France, Pyr.-Or., Le Tech (tamisage), 15.1V.1977, Löbl
(MHNG); paratypi: France, Pyr.-Or., Le Tech (tamisage), 15.1V.1977, Löbl (MHNG) 4 exx.;
Pyrénées-Orientales, Le Tech, m. 560, N 42° 24° 443” E 02° 31’ 822”, 18.5.2003, Bellò (BEL)
14 exx., Pierotti (PIE) 14 exx.

PERITELINI PALEARTICI V 593

ctu

FIG. 6

Spermateca in visione laterale: a, Heteromeiropsis zariquieyi; b, Heteromeiropsis loebli; c,
Heteromeiropsis valentiana. Scala: 0,10 mm.

Diagnosi. Specie di statura media o piccola (3,9-4,9 mm.), caratterizzata da
elitre non particolarmente allungate, ad apice deboimente ogivale, e da setole erette o
semierette sul vertice, sul dorso del pronoto presso il margine anteriore e sulle elitre,
reclinate altrove ed in particolare, rispetto ad H. zariquieyi, da antenne con scapo un
po’ più robusto, da pene in visione dorsale a profilo cordiforme nella regione apicale e
da propigidio 9 a margine apicale subtroncato.

Descrizione. Forma robusta, discretamente allungata. Vestitura dorsale
costituita da squame terree, qua e là più scure e da setole robuste, erette o semierette
sul vertice, sulle elitre e sul dorso del pronoto presso il margine anteriore, qui rivolte
un po’ in avanti verso l’interno, reclinate altrove.

594 H. PIEROTTI & C. BELLÒ

FIG. 7

Heteromeiropsis zariquieyi: armature del sacco interno del pene: a, armature orifiziali; b,
armatura mediana; c, armatura genitale. Scala: 0,10 mm.

Rostro trasverso, a lati subparalleli (4 ) o debolmente convergenti in avanti (9);
fronte, epifronte e vertice percorsi da un unico solco longitudinale mediano; pterigi
pressochè nulli. Occhi subarrotondati o debolmente allungati, discretamente convessi,
non sporgenti oltre le guance. Vertice stretto, poco più largo della fronte. Antenne
robuste, soprattutto nella 9, con scapo ingrossato già nel primo terzo e funicolo con i
primi due articoli poco allungati, ricoperti anche di setole distintamente più allargate
ed appiattite, il secondo subeguale ai due seguenti presi insieme o un po’ più corto, gli
articoli 4-7 moniliformi o talora (nella ©) trasversi; clava slanciata, col primo articolo
brevemente svasato.

PERITELINI PALEARTICI V 595

Fic. 8

Heteromeiropsis loebli: armature del sacco interno del pene: a, armature orifiziali; b, armatura
mediana; c, armatura genitale. Scala: 0,10 mm.

Pronoto più (2) o meno (d) trasverso, sul disco con punteggiatura doppia,
discretamente fitta.

Elitre più (6) o meno (9) allungate, con omeri smussati (4 ) o discretamente
evidenti (9), largamente ogivali nella regione apicale, con strie di punti ravvicinati
evidenti, ciascuno con una esilissima microsetola, ed interstrie debolmente convesse.

Zampe robuste, con protibie un po’ incurvate verso l’interno avanti l’apice.

Propigidio a margine apicale subtroncato anche nella 2 (Fig. 5.b).

Pene, in visione dorsale, debolmente tricuspidato, con le cuspidi laterali arro-
tondate (Fig. 4.b); armature del sacco interno: Fig. 8.

Spiculum ventrale con margo apicalis della lamella debolmente inciso nel
mezzo.

Coxiti discretamente allungati.

Spermateca: Fig. 6.b.

Derivatio nominis. Dedichiamo la specie al suo primo raccoglitore, il dott. Ivan
Löbl di Ginevra.

596 H. PIEROTTI & C. BELLÒ

Fic. 9

Heteromeiropsis valentiana: armature del sacco interno del pene: a, armature orifiziali; b, ar-
matura mediana; c, armatura genitale. Scala: 0,10 mm.

Distribuzione geografica. Francia sud-occidentale (Pyrénées-Orientales).
Note ecologiche. La specie è stata raccolta al vaglio, al margine di foresta di
latifoglie.

3. Heteromeiropsis valentiana sp. n.

Serie tipica. Holotypus d : Valencia, s.ra des Agulles, m. 390, cami de la font del barber,
2.11.2000, leg. Pierotti (MHNG); paratypi: ibidem, 2.11.2000, leg. Bello (BEL) 5 exx., (MHNG)
1 ex., leg. Pierotti (PIE) 5 exx.; Tabernes, Valencia, Hispania, coll. E.Moroder, 27.V.1934 (TOS)
lex.

Diagnosi. Specie di statura media o piccola (3,9-5 mm.), caratterizzata da elitre
non molto allungate, ad apice un po’ arrotondato anche nel d, da setole reclinate sul
dorso delle elitre e da propigidio d a margine apicale largamente lanceolato, papillato
al centro.

PERITELINI PALEARTICI V 597

Descrizione. Forma robusta, un po’ allungata. Vestitura dorsale costituita da
squame scure con modesti riflessi metallici e da setole robuste, semierette sul vertice,
reclinate sul pronoto e sul dorso delle elitre, alquanto sollevate presso l’apice delle
elitre.

Rostro trasverso, a lati debolmente convergenti in avanti; pterigi poco salienti.
Occhi subarrotondati, poco convessi, non sporgenti oltre le guance. Vertice stretto,
poco più largo della fronte, nel mezzo con una fossetta. Antenne con scapo discreta-
mente robusto, debolmente ricurvo, progressivamente ingrossato dalla base verso
l’apice; funicolo con i primi tre articoli poco allungati, ricoperti anche di setole larghe,
terree con riflessi metallici, i tre seguenti submoniliformi ed il settimo un po’ più lungo
che largo; clava alquanto allungata, con il primo articolo un po’ peduncolato.

Pronoto trasverso, alquanto fittamente punteggiato sul disco.

Elitre debolmente arrotondate ai lati, debolmente ogivali nella regione apicale,
con strie di punti evidenti nonostante la vestitura, ciascuno con una microsetola allun-
gata, ed interstrie alquanto convesse.

Zampe robuste.

Propigidio a margine apicale subtroncato nel d, largamente lanceolato,
papillato all’apice, nella 9 (Fig. 5.c).

Pene, in visione dorsale, progressivamente ristretto verso la regione apicale, qui
debolmente e molto brevemente tricuspidato, con le cuspidi laterali molto arrotondate
(Fig. 4.c); armature del sacco interno: Fig. 9.

Spiculum ventrale con margo apicalis della lamella distintamente inciso nel
mezzo.

Coxiti allungati.

Spermateca: Fig. 6.c.

Derivatio nominis. Dalla regione di provenienza.

Distribuzione geografica. Spagna orientale (Valencia).

Note ecologiche. La specie è stata raccolta sotto Pistacia lentiscus L.

RINGRAZIAMENTI

Ringraziamo molto vivamente il Dott. Giulio Cuccodoro del Museum d’histoire
naturelle di Ginevra ed il Dott. Sergio Montagud del Museo Valenciano de Historia
Natural di Valencia, per averci affidato in studio i Peritelini delle collezioni conservate
nei loro Istituti, consentendoci così di individuare le due nuove specie sopra descritte.

BIBLIOGRAFIA

ALONSO-ZARAZAGA, M.A. 2002. Lista preliminar de los Coleoptera Curculionoidea del area
ibero-balear, con descripciön de Melicius gen.nov. y nuevas citas. Boletin de la S.E.A. -
Sociedad Entomologica Aragonesa 31: 9-33.

ALONSO-ZARAZAGA, M.A. & LYAL, C.H.C. 1999. A World Catalogue of Families and Genera of
Curculionoidea (Insecta: Coleoptera) (Excepting Scolytidae and Platypodidae).
Entomopraxis SCP Edition, Barcelona, 315 pp.

DELLACASA, G., BORDAT, P. & DELLACASA, M. 2001. A revisional essay of world genus-group
taxa of Aphodiinae (Coleoptera Aphodiidae). Memorie della Società Entomologica
Italiana 79: 1-482.

598 H. PIEROTTI & C. BELLÒ

PIEROTTI, H. & BELLÒ, C. 1994. Peritelini nouveaux ou intéressants de la faune thyrrénienne
(Coleoptera Curculionidae Polydrusinae). 2° Contribution à la connaissance de la Tribu
Peritelini. L’Entomologiste 50 (5): 289-296.

PIEROTTI, H. & BELLO, C. 1998. Present knowledge of Palaearctic Peritelini (Coleoptera:
Curculionidae: Polydrosinae). /n: COLONNELLI, E., Louw, S. & OSELLA, G. (eds).
Taxonomy, ecology and distribution of Curculionoidea (Coleoptera: Polyphaga). XX
I.C.E. (1996, Firenze, Italy). Atti del Museo regionale di Scienze naturali di Torino:
81-108.

SOLARI, F. 1954. Proposta di un riordinamento delle Tribü degli Otiorhynchini e dei Peritelini e
creazione di tre nuovi generi di questi ultimi (Col.Curculionidae). Memorie della Società
Entomologica Italiana 33: 33-63.

REVUE SUISSE DE ZOOLOGIE 111 (3): 599-609; septembre 2004

First report of Tischeriidae (Lepidoptera) on the Galapagos
Islands, Ecuador, with descriptions of two new endemic species

Bernard LANDRY! & Lazaro ROQUE-ALBELO?

1 Muséum d’histoire naturelle, C. P. 6434, CH-1211, Geneva 6, Switzerland.
E-mail: bernard.landry@mhn.ville-ge.ch

2 Charles Darwin Research Station, A.P. 17-01-3891, Quito, Ecuador; and
Biodiversity and Ecological Processes Research Group, Cardiff School of
Biosciences, Cardiff University, PO Box 915, Cardiff CF10 3TL, United Kingdom.
E-mail: lazaro@fcdarwin.org.ec

First report of Tischeriidae (Lepidoptera) on the Galapagos Islands,
Ecuador, with descriptions of two new endemic species. - The family
Tischeriidae is reported from the Galapagos Islands for the first time. The
two species found on the archipelago (Astrotischeria scalesiaella B. Landry,
sp. n. and A. alcedoensis B. Landry, sp. n.) are described and illustrated.
Both species are presumed to be endemic and their larvae are leaf miners on
Scalesia species, a group of endemic Asteraceae.

Keywords: Micro moths - Astrotischeria - taxonomy - leaf miners -
Scalesia - Asteraceae.

INTRODUCTION

The Tischeriidae (Tischerioidea) are a group of small, primitive moths
belonging to the Monotrysian Heteroneura (Lepidoptera). They may be the sister-
group of the Ditrysia (Nielsen & Kristensen, 1996; Kristensen & Skalski, 1998)
although Davis (1986) presented an alternative hypothesis of relationships. Some 101
species are described so far, mostly from the Holarctic Region, but species are known
also from the Ethiopian, Oriental, and Neotropical Regions (Puplesis & Diskus, 2003).
Eleven species have been described from the Neotropics, but the region remains poorly
known with regard to its leaf-mining fauna. The region with the most diverse tischeriid
fauna is the Nearctic, with 48 described species (Davis, 1983). Larvae mine leaves of
Fagaceae, Rosaceae, Asteraceae, Rhamnaceae, Malvaceae, and Tiliaceae, in decreasing
order of importance, although Anacardiaceae, Apocynaceae, Combretaceae, Ericaceae,
Euphorbiaceae, Hypericaceae, Sterculiaceae, and Theaceae are also used (Puplesis &
Diskus, 2003).

For the purpose of documenting the micro-moth fauna of the Galapagos archi-
pelago, BL conducted fieldwork on most of the larger islands in 1989 and 1992. Since
1997, LRA and others at the Charles Darwin Research Station on Santa Cruz Island,

Manuscript accepted 18.11.2003

600 B. LANDRY & L. ROQUE-ALBELO

did more collecting and reared more immature stages, leading to a wealth of new
information. This paper is part of a series begun in 1992 (Landry & Gielis, 1992) to
describe new species and report on other native and introduced species of micro-moths
in the Galapagos to develop a better understanding of this important group of insects
for biodiversity management purposes.

MATERIAL AND METHODS

Most of the moths studied here were reared from mined leaves of Scalesia
species (Asteraceae). Mined leaves were collected by staff of the Charles Darwin
Research Station (CDRS), Santa Cruz Island, Galapagos, brought to their quarantine
lab, and placed in cages. Emerging moths were collected and mounted. Two other
series of specimens were available for study. The larger one was collected by BL in
1992 with a portable generator and a mercury-vapour lamp placed in front of a white
sheet (see Landry & Gielis, 1992). The other series was collected at light in 1964 and
was borrowed from the California Academy of Sciences, San Francisco, California,
U.S.A. (CAS).

The holotype and some paratypes of Astrotischeria scalesiaella will be
deposited in the Muséum d’histoire naturelle, Geneva, Switzerland (MHNG). The
other paratypes of A. scalesiaella will be deposited in the CAS, CDRS, National
Museum of Natural History, Washington, D.C., U.S.A. (USNM), and The Natural
History Museum, London, England (BMNH). The single known specimen of
Astrotischeria alcedoensis is deposited in the MHNG.

The species described here were determined to be new by comparing specimens
with the types of seven of the Neotropical species, with the original and more recent
descriptions of four species (Bourquin, 1962; Puplesis & DiSkus, 2003), and with the
treatment of the North American species published by Braun (1972). The types of six
Neotropical species described by Meyrick and Walsingham were examined at the
BMNH. The type of Tischeria unicolor Walsingham is located in the Naturhistorisches
Museum Wien, Vienna, Austria (NMW) and was borrowed for examination.

Genitalia were dissected after the abdomen had macerated in a cold 20% KOH
solution overnight. The dissected parts were kept in lactic acid stained with orange G
for description purposes. They were subsequently stained with chlorazol black and
mounted on slides in Euparal. The head of one male specimen was similarly treated.
The right pair of wings of one male specimen was removed, bleached, stained with acid
fuchsin, and mounted in Euparal.

Illustrations of the adult head and genitalia were made with the AutoMontage®
system using a video camera mounted on a Leica MZ APO stereomicroscope or a Zeiss
Axioskop compound microscope. The wing venation was drawn using a camera lucida
mounted on a compound microscope.

The descriptions and nomenclature of parts are based on Puplesis & Diskus
(2003). Generic characters were not repeated. In the legs, wings, thorax, and abdomen
only the lateral or dorsal exposed surfaces are described; the ventral or median surfaces
of the legs are uniformly whitish beige.

TISCHERIIDAE ON THE GALAPAGOS ISLANDS 601

SYSTEMATIC TREATMENT
Astrotischeria Puplesis & Diskus, 2003

The genus was described to include a monophyletic group of species found on-
ly in the New World. Thirty species are known, seven of which were described from
the Neotropical Region (Argentina, Chile, Ecuador, Peru).

Species of Astrotischeria are characterised by the following synapomorphies:
presence of a dorsal arm on the valva; shortened uncus with arms (with some
exceptions); long, distally bifurcated aedeagus; strongly developed anellus thickened
laterally; enlarged vinculum; stronger development of microtrichia on the female
tergite IX; slightly reduced, narrowed ovipositor lobes; greatly elongated mediobasal
spur of the hind tibia, which is as long as the tibia; and use of Malvaceae and Aster-
aceae as hostplants (Puplesis & Diskus, 2003).

Astrotischeria scalesiaella B. Landry sp. n. Figs 1-2, 4-7, 9-12

Holotype è. [1] “ECU., GALAPAGOS/ Santa Cruz, Los/ Gemelos, 27.v.1992/ M[er-
cury] V[apor] L[amp], /eg. B. Landry” (MHNG). Printed in black ink on white paper. [2]
“Tischeria/ det. Davis 1993”. Hand-written on white card stock with black border in black ink
except for black ink printed “det. Davis 19”. [3] “HOLOTYPE/ Astrotischeria/ scalesiaella/ B.
Landry”. Hand-written in black ink on red card stock. Specimen in perfect condition.

Paratypes, Ecuador: 12 6g, 10 22 from the Galapagos Islands. FLOREANA: 1 d
(dissected, slide BL 1175), Cerro Pajas, 26.x11.1998, ex larva en Scalesia pedunculata (L.
Roque). SANTA CRUZ: 1 2, Los Gemelos, 17.1.2002, [no emergence date], Minador de hojas
Scalesia pedunculata (R. Boada); 1 3 (dissected, BL 1415), same data as holotype; 1 6, 1 ©,
Los Gemelos, 2.ix.2001, emergio 10.ix.2001, Minador Scalesia pedunculata (L. Roque); 2 3,
1 2, Los Gemelos, 11.ix.2001, emergio 28.ix.2001, 30.ix.2001 and no emergence date, Criada
en hojas Scalesia pedunculata (R. Boada); 4 4, Los Gemelos, xi.1999, [no emergence date], En
Scalesia pedunculata (T. Poulson); 1 4, 2 2 (one dissected, BL 1417), Finca Vilema, 2 km W
Bella Vista, 1.1v.1992, M[ercury] V[apor] L[amp] (B. Landry); 1 & (dissected, Wings BL 1416;
Head BL 1420), E[stacion] Cfientifica] C[harles] D[arwin], 6.111.1992, U[Itra] V[iolet] L[ight]
(B. Landry); 1 9, Steve Devine farm, 31.x.1999, [no emergence date], minador de hojas Scalesia
pedunculata (C. Causton); 16 (dissected, BL 1418), 3 2, Horneman Farm, 220 m, 5.iv.1964
(D.Q. Cavagnaro). WoLF: 1 @ (dissected, BL 1419), 7.11.2002, Minador Scalesia baurii (L.
Roque, C. Causton).

Diagnosis. This species and the next are easily distinguished from other small
Galapagos micro-moths with lanceolate wings by virtue of the combination of three
characters of the head: 1- the presence of scales on the proboscis, 2- the drooping labial
palpi, and 3- the rough vestiture of the vertex. This is a combination of characters
found in all Tischeriidae. Gelechioidea have scales on the proboscis as well, and some
have lanceolate wings, but their labial palpi are upturned and their vertex is not rough-
scaled. Astrotischeria scalesiaella can be separated from A. alcedoensis by its smaller
size (forewing length up to 3.02 mm in A. scalesiaella and 4.28 mm in A. alcedoensis),
more prominent orange colour with more diffuse brown markings, wider, shorter and
more angled uncus arms, apically thinner and more pointed dorsal lobes of the valva,
and the shorter cucullus. The female of A. alcedoensis is unknown.

Description. MALE (n=13) (Figs 1, 4, 6, 7, 9-12). Vertex vestiture a mixture of
ochre and brown scales, the ochre scales bicoloured with a paler basal half, the brown
scales with white on basal half and apex; frons vestiture (Fig. 4) mostly greyish brown,

602 B. LANDRY & L. ROQUE-ALBELO

TISCHERIIDAE ON THE GALAPAGOS ISLANDS 603

shining, with row of white or pale ochre scales laterally and apically; interocular index
(n=1): 1.76. Antenna 2/3 length of forewing (n=1); with 20 flagellomeres (n=1); scape
and slender pecten brown and white, sometimes with some pale ochre scales; with 6
rows of scales on flagellomere I, with 2 rows on other flagellomeres; scales on flagel-
lomeres with brown apex and white basal half; flagellomere I 3X length of flagellom-
ere II; sensillae chaeticae about 1.5X length of flagellomere on flagellomere II, about
2/3 length of flagellomere on last one. Haustellum 3X length of labial palpus; scales on
both beige, sometimes with brown laterally on labial palpus. Maxillary palpus 3-seg-
mented, scaled only on third segment. Thorax mostly brown with beige-tipped scales,
sometimes with ochre scales at tip of tegulae. Foreleg dark brown. Midleg femur dark
brown laterally with pale ochre scales on dorsal edge and beige apically; tibia dark
brown with beige on dorsal row of long and narrow scales, and apex; first tarsomere
with scales dark brown on apical half and white on basal half; tarsomeres I-V mostly
beige with apical dark-brown spot. Hind coxa dark brown with apical row of longer
beige scales; femur mostly white; tibia dark brown with beige along ventral margin and
on ventral and dorsal rows of long and thin scales, with pale ochre apically; tarsomeres
dark brown. Forewing length: 2.46-3.02 mm (holotype: 2.81 mm). Venation (Fig. 9)
with only Sc, R, Rs1, Rs4, MI, CuA, and 1A + 2A clearly defined; with a faint indica-
tion of Rs3; 1A + 2A not forked at base. Length/ largest width: 0.25. With a patch of
microtrichia in anal sector. Colour (Fig. 1) mostly dark brown with ochre scales along
dorsal margin beyond middle, along cubital fold, in apical sector, and as a diagonal bar
or large spot medially, with darker brown markings as two oblique bands before and
beyond middle, first one more inclined than second; sometimes with more ochre scal-
ing (fig. 2); sometimes mostly dark brown with a few ochre scales mostly in apical sec-
tor; scales paler on their basal half, white on those with dark-brown apical half; fringe
greyish brown. Hindwing venation only with Sc + R and A clearly visible. Sockets of
the fringe scales forming spines especially conspicuous on costal margin. Without
microtrichia. Colour brownish grey, with concolorous fringe. Abdomen greyish brown.

Male genitalia (n=3) (Figs 7, 10-12). Socii short, with short, slender setae. Un-
cus arms separate and connecting to tegumen at their most dorsal and most anterior
points; each arm elongate, larger in middle, directed downward, flattened and narrowly
rounded at apex, with a few short setae on dorsal and ventral edges beyond connection
with tegumen. Tegumen hood-like, rounded dorsally and apically, with short setae
dorsolaterally, with very narrow and strongly melanised anterior arms connecting with
vinculum, with broader and lightly melanised posterior arms connecting with mem-
branous ventral wall of tegumen. Vinculum rather broad, triangular in ventral view,
without saccus. Anellus broad, lightly melanised, rectangular, with about five short

Fics 1-8

Galapagos Astrotischeria spp. 1. Holotype of A. scalesiaella; 2. Female paratype of A. scale-
siaella from Santa Cruz, Finca Vilema; 3. Holotype of A. alcedoensis from Isabela, Volcan
Alcedo, reared from Scalesia affinis; 4. Head of male paratype of A. scalesiaella from type
locality; 5. Leaf of S. pedunculata mined by A. scalesiaella on Los Gemelos, Santa Cruz; 6.
Descaled head of male paratype of A. scalesiaella from Santa Cruz, Charles Darwin Station; 7.
Male genitalia of paratype of A. scalesiaella from Floreana, Cerro Pajas, reared from Scalesia
pedunculata; 8. Male genitalia of holotype of A. alcedoensis.

604 B. LANDRY & L. ROQUE-ALBELO

SEELE EEE 47 re RE
a

Fic. 9

Wings of paratype male of Astrotischeria scalesiaella from Santa Cruz, Charles Darwin Station
(frenulum broken) (scale: 0.5 mm) with details of the bases of the costal and inner margins
(scale: 0.25 mm).

setae ventrally. Valva with dorsal arm curved inward at right angle, narrowing, apically
pointed, and with narrow apodemes projecting directly anteriorly; cucullus slender,
slightly curving upward toward apex, with abundant short setation medially, apically
not narrowing and rounded, about as long (1.06 X) as dorsal arm inclusive of apodeme.
Aedeagus very narrow, slightly longer than vinculum + valva, divided in two divergent
branches from about 2/3; each branch apically flattened and with bunch of slender
spines and spinules; gonopore from about base of divergent branches.

FEMALE (n=10). Antenna 0.62 length of forewing (n=1). Forewing length:
2.58-2.83 mm. Frenulum with two acanthae. Forewing length/ largest width: 0.25.
Female genitalia (n=2) (Figs 16-21). Ovipositor lobes short and round, with rather
stout and slightly curved peg setae, the most medially located ones apically bifurcated;
median ridge between ovipositor lobes with a series of three pairs of short setae.
Tergum IX a more or less triangular plate, slightly convex, with convex lateral margins,
emarginated anterior margin, and straight posterior margin, the latter membranous and
set with bunch of 3-4 setae on each side of middle, including one much longer than
others; with strongly melanised narrow band at anterior and anterolateral margins
connecting lateroanteriorly with bases of posterior apophyses; with long and sparse
setation dorsally. Posterior lobes of sternum IX somewhat conical, slightly larger than
ovipositor lobes, with short and long setae, with long, straight and apically narrow
posterior apophyses extending beyond middle of segment VII; anterior lobes of
sternum IX very narrow with short setae directed posteriorly; with two pairs of apo-

TISCHERIIDAE ON THE GALAPAGOS ISLANDS 605

Fics 10-15

Male genitalia of Astrotischeria spp. 10-12. Paratype of A. scalesiaella, same specimen as fig. 7:
10. Posterior end of aedeagus; 11. Tegumen, uncus, and socii; 12. Dorsal arms of valvae, vin-
culum, anellus. 13-15. Holotype of A. alcedoensis: 13. Posterior end of aedeagus; 14. Posterior
end of tegumen, uncus, and socii; 15. Dorsal arms of valva.

physes in membranous area anterior to oviporus: dorsal pair narrow, almost reaching
as far as posterior apophyses (of sternum IX), apically acute; ventral pair broader,
shorter, reaching slightly beyond anterior margin of sternum VIII. Tergum VIII a
narrow rectangular and flat plate slightly broader posteriorly, with one anterolateral
projection on each side directed ventrally and connected to anterior apophyses of
sternum VIII slightly posterad to their middle. Sternum VIII a rather large concave
plate extended laterally, with broadly rounded apical margin set with narrow spinelike

606 B. LANDRY & L. ROQUE-ALBELO

projections. Ductus bursae with small elliptical plate set at right angle in bend beyond
anterior end of shorter apophyses of sternum IX, also with thin spinelike scobination
anteriorly from bend and until connection with corpus bursae, but more concentrated
near tips of longer apophyses. Corpus bursae small and oblong, with tiny spinules on
anterior half. Apex of tergum VII medially with joining pair of median arcs of closely
set scale sockets with elongate narrow scales. Sternum VII terminating in small wider
than long median lobe set with abundant and rather long setation.

Etymology. The specific epithet refers to the generic name of the host plants.

Biology. Adults were reared from leaf mines found on two species of Scalesia
(Asteraceae): S. baurii Robins. & Greenm. and S. pedunculata Hook. filius.

Distribution. Endemic to the Galapagos; found on Floreana, Santa Cruz, and
Wolf.

Remarks. In the female genitalia, my interpretation of tergum IX, which I
believe is dorsal, between terga VIII and X, differs from that of Davis (1998) and
Puplesis and Diskus (2003), who mention that it is ventral from the lobes of tergum X.
In my view, tergum IX of these authors is actually sternum IX because of its position.
In Davis’ (1998, fig. 6.8 N) schematic drawing of the female genitalia of Tischeria
(now Coptotriche) citrinipennella (Clemens), there is no sclerite between tergum VIII
and the ovipositor lobes of tergum X as in Astrotischeria scalesiaella (see Figs 18, 19).

Astrotischeria alcedoensis B. Landry sp. n. Figs 3, 8, 13-15

Holotype 3. [1] “ECU. GALAPAGOS/ Isabela, V[olcan]. Alcedo, en/ Scalesia villosa
affinis, 19 X/ 1999, emergio 29 X 1999/ L. Roque” (MHNG). Computer made in black on white
paper with black borders on top and bottom. Hostplant species name crossed with pencil and new
name added with pencil on last line with an arrow between the two names. [2] “MHNG/ Prép.
micr./ No 2724 3”. Printed black on white card stock except for ink hand written “MHNG”,
number and male sign. [3] “BL 1423 3”. Hand-written in black ink on green paper; upside
down. [4] “HOLOTYPE/ Astrotischeria/ alcedoensis/ B. Landry”. Hand-written in black ink on
red card stock. The right midleg and hindleg are in a gelatine capsule attached to the pin. The
left hindleg is broken beyond the tibia.

Diagnosis. See Diagnosis of Astrotischeria scalesiaella above to distinguish the
two species from each other and from all other known micro-moths of the Galapagos.

Description. MALE (n=1) (Figs 3, 8, 13-15). Vertex vestiture with erect scales
usually tricolored, pale beige at their base, brown subapically, pale beige at their apex,
with a few scales apically shining greyish brown, with appressed scales between
antennal bases entirely pale beige; frons vestiture with shorter scales coloured as erect
scales of vertex, but scales slightly paler at their base; interocular index not calculated.
Antenna about 3/4 length of forewing; approximately 26 flagellomeres (head not
mounted on slide); scape brown and white, with greenish-beige narrow pecten;
flagellomere I with longitudinal rows of white and brown scales; other flagellomeres
with basal scales pale beige and apical scales brown. Haustellum white and brown.
Labial palpus dark brown laterally with white apex, pale beige medially. Maxillary
palpus scaling probably white (not clearly visible). Thorax mostly with tricolored
scales, dirty white at their base, brown subapically, ochre apically. Foreleg coxa with
scales bicoloured dirty white at their base and brown at their apex (sometimes with
apex a little paler beyond brown area), other parts dark brown except for pale beige

TISCHERIIDAE ON THE GALAPAGOS ISLANDS 607

Fics 16-21

Female genitalia of Astrotischeria scalesiaella. 16-17. Paratype from Wolf, reared from Scalesia
baurii: 16. Segments VIII-X and genitalia; 17. Apex of abdomen. 18-19. Paratype from Santa
Cruz reared from Scalesia pedunculata: 18. Dorsal view of segments VIII-X; 19. Lateral view
of same segments. 20-21. Same specimen as on figs 16-17: 20. Section of ductus bursae; 21.
Corpus bursae.

bases of tarsomeres II-IV and all of tarsomere V. Midleg as in foreleg except scales
more bicoloured as on coxa and tarsomeres more pale beige except for small patch of
brown at apex of tarsomeres II-IV. Hindleg apparently as in midleg, but damaged.
Forewing length: 4.28 mm (holotype). Venation not studied. Colour a mixture of beige,
brown, and ochre scales, the latter not present in the radial sector, the brown scales
usually tricolored with white at base, brown on apical half and beige at apex as a
narrow band; without strongly marked pattern in unique and somewhat damaged
specimen, but possibly with submedian and postmedian oblique bands as in A. scale-

608 B. LANDRY & L. ROQUE-ALBELO

siaella; fringe pale greyish ochre. Hindwing venation not studied; colour pale greyish
beige; fringe pale greyish ochre. Abdomen colour not recorded.

Male genitalia (n=1) (Figs 8, 13-15). Socii and uncus as in A. scalesiaella
except arms of uncus more slender, slightly longer, less curved, and narrower apically.
Tegumen as in A. scalesiaella except slightly broader and with broader posterior arms.
Vinculum narrower, less triangular, than in A. scalesiaella. Anellus as in A. scalesiaella
except for apparent absence of ventral setae. Dorsal arm of valva with distal half
broader and not as narrowly pointed apically as in A. scalesiaella; cucullus as in A.
scalesiaella except more distinctly bent at about 2/3 and longer, 1.44 X as long as
dorsal arm inclusive of apodeme. Aedeagus as in A. scalesiaella except broadly curved
(possibly an artefact) and base of two apical branches at about 4/5 length; gonopore not
observed.

FEMALE. Unknown.

Etymology. The specific epithet refers to the collecting locality, Volcan Alcedo,
on Isabela Island.

Biology. The unique specimen was reared from a leaf of Scalesia affinis Hook.
filius (Asteraceae).

Distribution. Galapagos endemic; found on Isabela, Volcan Alcedo.

Remarks. The poorly melanised spines of the apical end of the aedeagus
(Fig. 13) may be the result of a premature death.

ACKNOWLEDGEMENTS

We thank the authorities of Galapagos National Park and those of the Charles
Darwin Research Station for allowing fieldwork and for logistical support in 1989,
1992, and 2002. We are grateful to the Charles Darwin Foundation and the Galapagos
Conservation Trust of England for providing financial support for BL’s investigations
at the BMNH in 2000 and in the Galapagos in 2002. We owe a great deal of gratitude
to Prof. Stewart B. Peck, Carleton University, Ottawa, for inviting BL to join his 1989
and 1992 Galapagos expeditions. This fieldwork was supported by an operating grant
to Dr Peck from the Natural Sciences and Engineering Research Council of Canada for
field research on arthropod evolution. For their help and pleasant company in the field
we thank Novarino Castillo, Charlotte Causton, Joyce Cook, Moraima Inca, Ricardo
Palma, S. B. Peck, Bradley J. Sinclair, and Eduardo Vilema. We thank curators Don R.
Davis (USNM), Sabine Gaal-Haszler (NMW), and Kevin Tuck (BMNH) for their
permission to examine specimens in their care. We are thankful to Jean-François
Landry, Agriculture and Agri-Food Canada, for his multi-faceted help while BL was in
Ottawa working on Galapagos moths. Finally, we thank Florence Marteau (MHNG)
for producing the plates and Don Davis, J.-F. Landry, and Rimantas Puplesis for their
important comments on the manuscript.

REFERENCES

BOURQUIN, F. 1962. Microlepidopteros nuevos con sus biologias. Revista de la Sociedad Ento-
mologica Argentina 23: 31-46.

BRAUN, A. 1972. Tischeriidae of America North of Mexico (Microlepidoptera). Memoirs of the
American Entomological Society 28: 1 + 148 pp.

TISCHERIIDAE ON THE GALAPAGOS ISLANDS 609

Davis, D. R. 1983. Tischeriidae (p. 3). In: HoDGEs, R.W. et al. (eds). Check list of the
Lepidoptera of America North of Mexico. E. W. Classey Ltd. and the Wedge
Entomological Research Foundation, London, xxiv + 284 pp.

Davis, D. R. 1986. A new family of Monotrysian moths from Austral South America (Lepi-
doptera: Palaephatidae), with a phylogenetic review of the Monotrysia. Smithsonian
Contributions to Zoology 434: iv + 202 pp.

Davis, D. R. 1998. The Monotrysian Heteroneura (pp. 65-90). Jn: KRISTENSEN, N. P. (ed.). Hand-
book of Zoology, Lepidoptera, Moths and Butterflies, Vol. 1: Evolution, Systematics,
and Biogeography. Walter de Gruyter, Berlin & New York, x + 491 pp.

KRISTENSEN, N. P. & SKALSKI, A. W. 1998. Phylogeny and Palaeontology (pp. 7-25). In:
KRISTENSEN, N. P. (ed.). Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol.
1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin & New York, x
+ 491 pp.

LANDRY, B. & GIELIS, C. 1992. A synopsis of the Pterophoridae (Lepidoptera) of the Galapagos
Islands, Ecuador. Zoologische Verhandelingen, Leiden 276: 1-42.

NIELSEN, E. S. & KRISTENSEN, N. P. 1996. The Australian moth family Lophocoronidae and the
basal phylogeny of the Lepidoptera-Glossata. Invertebrate Taxonomy 10: 1199-1302.

PUPLESIS, R. & Diskus, A. 2003. The Nepticuloidea & Tischerioidea (Lepidoptera) - a global
review, with strategic regional revisions. Lutute Publishers, Kaunas, 512 pp.

REVUE SUISSE DE ZOOLOGIE 111 (3): 611-630; septembre 2004

A revision of the Blattodea, Ensifera and Caelifera
described by H. Fruhstorfer

Hannes BAUR! & Armin CORAY?
! Department of Invertebrates, Natural History Museum, Bernastrasse 15,

CH-3005 Bern, Switzerland. E-mail: hannes.baur@nmbe.unibe.ch (correspondence)
2 Natural History Museum, Augustinergasse 2, CH-4001 Basel, Switzerland.

E-mail: armin.coray@balcab.ch

A revision of the Blattodea, Ensifera and Caelifera described by
H. Fruhstorfer. - Twelve taxa described by Fruhstorfer are revised,
belonging to Blattodea: Blattellidae (1), Ensifera: Phaneropteridae (1),
Tettigoniidae (4) and Caelifera: Acrididae (6). Name-bearing types are
deposited in the Swiss Federal Institute of Technology, Zurich (7), the
Zoologische Staatssammlung, Munich (1) and the Natural History
Museums of Berlin (1) and Geneva (4). Syntype series of two taxa are
missing. In seven cases lectotypes are here designated. Four new
synonymies are established: Tettigoniidae: Ephippiger ephippiger eustra-
tius Fruhstorfer, 1921 = E. diurnus Dufour, 1841; Ephippiger vicheti Harz,
1966 = E. persicarius Fruhstorfer, 1921 stat. rev.; Thamnotrizon chabrieri
malachiticus Fruhstorfer, 1921 = Eupholidoptera chabrieri (Charpentier,
1825). Acrididae: Stenobothrus lineatus fervidior Fruhstorfer, 1921 =
Stenobothrus lineatus (Panzer, 1796). The paper provides detailed informa-
tion on name-bearing types, type localities and the current status of each
taxon.

Keywords: Blattellidae - Phaneropteridae - Tettigoniidae - Acrididae - new
synonymy - lectotype designations - taxonomy - distribution.

INTRODUCTION

Hans Fruhstorfer (1866-1922) was a Swiss naturalist who mainly lived in
Germany (Martin, 1922). Coming from a family in modest circumstances he quickly
gained some wealth because of his extraordinary talents as a salesman of insects. His
financial independence allowed him to travel extensively through South America and
Asia where he spent most of his time collecting insects. His main interest was in butter-
flies and due to innumerable publications and his contributions to the monumental
work of Adalbert Seitz on Rhopalocera (butterflies) he became one of the best known
lepidopterologists of his days (Martin, 1922). Apparently, he established an incredible
number of almost 5000 new names (Seitz, 1922)! Only late in his life, Fruhstorfer
developed a keen interest in orthopteroid insects. In a remarkably short time he

Manuscript accepted 18.03.2004

612 H. BAUR & A. CORAY

assembled a voluminous work in which he mainly investigated the distribution and ori-
gin of Swiss grasshoppers, cockroaches, mantids and earwigs (Fruhstorfer, 1920a-
1920f, 1921a-1921c). For a very long time his main work “Die Orthopteren der
Schweiz und der Nachbarländer...” (Fruhstorfer, 1921c) remained a cornerstone for
faunistic research (Thorens & Nadig, 1997), some criticism by Ramme (1923) and
Zeuner (1931) notwithstanding. Fruhstorfer (1920d, 1921c) also described a series of
new taxa in Blattodea, Ensifera and Caelifera. Although the majority of these new
names have merely infrasubspecific status, twelve were formally published in accor-
dance with the International Code of Zoological Nomenclature (1999, ICZN) and must
be treated as available names (Table 1). In the past, however, many of Fruhstorfer’s
taxonomic decisions did not receive the attention they deserved, which, in turn, is
critical with respect to nomenclatural stability. We therefore present in this revision all

TABLE 1. Synopsis of taxa established by Fruhstorfer

Original names Valid names

BLATTODEA

Ectobius neolividus Fruhstorfer Ectobius vittiventris (A. Costa)

ENSIFERA

Barbitistes obtusus alpinus Fruhstorfer Barbitistes obtusus Targioni-Tozzetti

Ephippiger cruciger eustratius Fruhstorfer syn. n. Ephippiger diurnus Dufour

Ephippiger persicarius Fruhstorfer Ephippiger persicarius Fruhstorfer
stat. rev.*

Ephippiger plinianus Fruhstorfer Ephippiger bormansi Brunner v. W.

Thamnotrizon chabrieri malachiticus Fruhstorfer syn. n. Eupholidoptera chabrieri
(Charpentier)

CAELIFERA

Chrysochraon brachypterus chrysoberyllus Fruhstorfer Euthystira brachyptera (Ocskay)

Omocestus haemorrhoidalis fantinus Fruhstorfer Omocestus haemorrhoidalis
(Charpentier)

Podisma alpina formosanta Fruhstorfer Miramella (Nadigella) formosanta
(Fruhstorfer)

Podisma alpina irena Fruhstorfer Miramella (Kisella) irena
(Fruhstorfer)

Podisma frigida strandi Fruhstorfer Bohemanella frigida (Boheman)

Stenobothrus lineatus fervidior Fruhstorfer syn. n. Stenobothrus lineatus (Panzer)

*= Ephippiger vicheti Harz syn. n.

pertinent data for a proper taxonomic treatment of Fruhstorfer’s names. This includes
detailed information on the name-bearing types, the type locality and the current status
of the taxa concerned. The present paper is thus intended as a further step towards a
List of Available Names (ICZN, Art. 79) in orthopteroid insects.

MATERIAL AND METHODS

Fruhstorfer’s insect collection is preserved in several different institutions
(Horn er al., 1990) and probably also in some private collections. Horn et al. (1990)
mention that a collection of Swiss “Orthoptera” had been donated in 1921 to the Swiss

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 613

Federal Institute of Technology, Zurich (ETHZ). A large part of the collection is also
deposited at the Zoologische Staatssammlung München (Schönitzer, pers. comm.).
This is not surprising, since Fruhstorfer moved to Munich shortly before his death
(Martin, 1922). We traced further material in the Museums of Basle, Berlin, Geneva
and Vienna, but specimens are likely to be encountered in many further collections.
Fruhstorfer (1921c; Martin, 1922) was in contact with many colleagues and he
certainly also exchanged some specimens. For instance, syntypes of several species
were found in the Nadig collection in Geneva.

Fruhstorfer’s collection has not been preserved in its original state. The speci-
mens have been removed from their original drawers and have been integrated in
general collections of the respective groups. This happened for instance in Zurich,
Munich and Vienna. Fruhstorfer’s material can nevertheless be recognized because he
properly labelled his specimens with printed or partly handwritten labels where he
usually noted the locality, date and name of the collector (Figs la, 2a, 3b). The only
difficulty is that he most often failed to add individual identification labels (see
Fig. 2b). From a few remnants it is apparent that he had used single bottom labels (Fig.
1b) to denote a particular taxon in his collection. Such labels are now mostly lost,
which renders the identification of particular specimens rather difficult. This is espe-
cially true for the type series. Fruhstorfer based his new taxa solely on syntypes which
he did not label specially. Therefore, we have accepted as syntypes only those speci-
mens which could be unambiguously recognized by their data label, and which, of
course, fit the description well. For instance, we consider as part of the type series of
Chrysochraon brachypterus chrysoberyllus a specimen labelled “Claro 9.VIII.18”
because Fruhstorfer (1921c: 95) cited exactly this locality and date in the original
description. We thereby assume that he did not mix his new subspecies with any other
grasshoppers mentioned from the same place, e. g. Omocestus haemorrhoidalis fanti-
nus or Gomphocerus rufus. Such taxa can be easily separated, even by a beginner.
However, Fruhstorfer evidently confounded the species of certain genera, for instance
of Ectobius and Ephippiger (Ramme, 1923; Nadig, 1968). Here, we accepted only
specimens with unique data labels as type specimens. A female Ephippiger labelled
“Piandolce 31.X.18” (Figs 3, 6) in the Zoologische Staatssammlung München is thus
regarded as part of the type series of E. plinianus because Fruhstorfer cited the
respective data solely for this Ephippiger species.

ABBREVIATIONS AND DEPOSITORIES

ICZN International Code of Zoological Nomenclature, Fourth edition (1999)

CAN Coll. Adolf Nadig, in MHNG

CKH Coll. Kurt Harz, in MHNG

ETHZ Eidgenössische Technische Hochschule, Entomologische Sammlung,
Zürich, Schweiz

MHNG Museum d’histoire naturelle, Genève, Suisse

NHMB Naturhistorisches Museum, Basel, Schweiz

NHMW Naturhistorisches Museum, Wien, Österreich

NMBE Naturhistorisches Museum, Bern, Schweiz

ZMB Zoologisches Museum, Berlin, Deutschland

ZSM Zoologische Staatssammlung, München, Deutschland

614 H. BAUR & A. CORAY

INFRASUBSPECIFIC NAMES

Fruhstorfer (1921c) introduced a series of new names using the term “forma” or
“forma nova” in combination with a binomen. The content of his descriptions un-
ambiguously reveals that most of these names have merely infrasubspecific rank
(ICZN, Art. 45.6.4). Fruhstorfer, apparently, denoted as a new form some outstanding
morphs which differed in color or size from the majority of specimens of a variable
species. In other words, he named some particular specimens instead of some popula-
tion(s). We think, it is thus sufficiently clear that such “forms” can have no status in
zoological nomenclature, since systematics aims at the delimitation of populations, not
individuals (Mayr & Ashlock, 1991; Cracraft, 1997). For completeness, we list the
respective names and summarize the content of their descriptions. We would like to
stress that these names had never been adopted as the valid names of a taxon. They are
not available and cannot enter the synonymy. The following nineteen names belong
here: Barbitistes obtusus f. “picta” (p. 176, color variety); Barbitistes obtusus f. “viri-
descens” (p. 176, color variety); Chorthippus dorsatus f. “nigripennis” (p. 130, color
variety); Chorthippus dorsatus f. “pallilis” (p. 130, color variety); Conocephalus
tuberculatus f. “persicaria” (p. 191, color variety); Gomphocerus rufus f. “alpicola”
(p. 99, size and color variety); Gomphocerus rufus f. “insubrica” (p. 99, color variety);
Gomphocerus rufus f. “murina” (p. 99, color variety); Gomphocerus rufus f. “pyg-
maea” (p. 99, size variety); Gomphocerus sibiricus f. “deminutus” (p. 103, size
variety); Gomphocerus sibiricus f. “viridopicta” (p. 103, color variety); Mecostethus
grossus f. “vinula” (p. 143, color variety); Platycleis brachyptera f. “deminuta”
(p. 221, 222, size variety); Platycleis brachyptera f. “restricta” (p. 220, 222, size and
color variety); Platycleis brachyptera f. “rhaetorum” (p. 220, 222, color variety);
Platycleis saussureana [sic] f. “rugosa” (p. 223, 224, size and color variety);
Stauroderus miniatus f. “smaragdina” (p. 116, female color variety); Stenobothrus
lineatus f. “interposita” (p. 108, color variety); Stenobothrus lineatus f. “myrina”
(p. 108, color variety).

The names of the four Platycleis-formae are in fact borderline cases. From their
first mentioning and the descriptions, these names might be regarded as subspecific. On
the other hand, Fruhstorfer evidently listed them in addition to a trinomen in his
classification of taxa (see pp. 222, 224). In our opinion, this is a clear indication that
he really intended to give them infrasubspecific rank. As we noted above, these names
have never been used as the valid names of a taxon, Otte’s (1997: 289) adoption of
“deminuta” in the rank of a species notwithstanding. A merely formal elevation in rank,
however, does not make it an available name (compare ICZN, Arts. 11.5.2 and 45.5.1).
The same applies in the case of Yin er al. (1996) who listed “Stauroderus miniatus
smaragdina [sic] Fruhstorfer” as a valid name in their catalogue of World grass-
hoppers.

A further two names, Podisma alpina formosanta and P. a. irena, have also been
introduced as “forma nova” by Fruhstorfer (1921c) but are considered as available
names (see under their respective headings).

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 615

CATALOGUE OF TAXA ESTABLISHED BY FRUHSTORFER

For easy access the taxa are arranged alphabetically within families, i.e. first the
original genus then the species. After name, author, year, and page number where the
species was first described, the name-bearing type (bold if designated here) and type
locality are noted. Where possible the current spelling of a locality is given. Material
starts with name-bearing types. Labels are cited in quotation marks, the text is rendered
exactly as spelt, with semicolons separating the different labels. Mostly original labels
are quoted only, beginning with the uppermost. Square brackets indicate additional
information (such as a red label, print or handwriting, etc.). Remaining specimens of a
type series are listed more summarily, their labels are cited only if necessary. Voucher
specimens are detailed for difficult taxa in the Appendix, otherwise that material is
summarized by reference to the respective depositories. Discussion provides further
information on the type material and mentions additional specimens which are doubt-
fully attributed to the type series or otherwise noteworthy. Taxonomic notes are usually
included. Terminology and abbreviations of morphological terms follow Harz (1969,
1975). We apply the Phylogenetic Species Concept which makes no use of the sub-
species category. Following this concept, populations are classified as species when
they are diagnosably distinct from other populations by a unique combination of fixed
character states (Cracraft, 1989; Nixon & Wheeler, 1990). Status finally lists current
usage of a name.

BLATTODEA
BLATTELLIDAE

Ectobius neolividus Fruhstorfer

Ectobius neolividus Ramme & Fruhstorfer [sic], 1921c: 78, 79. Syntypes d & 2 2, Switzerland:
Canton Ticino, Denti della Vecchia and Canton Grisons, Plotta near Soglio (CAN, ZMB
[examined]).

Material. Syntypes, CAN: 1 @ (directly pinned, right hind tarsus lacking) labelled
“Soglio Anf. 8. [sic] 20 Fruhstorfer [print]; Ectob. © neolividus Fruhst. [Fruhstorfer’s hand, blue
ink, label with border]; Ectobius vittiventris (COSTA) K. HARZ det. [print, blue ink]” (see Fig.
2) and 15 (directly pinned, left antenna, left hind leg and left hind tarsus lacking) labelled
“Soglio Anf. 8. [sic] 20 Fruhstorfer [print]; Ectobius 6 vittiventris Costa [hand] Ramme det.
[print]; Ectobius vittiventris (COSTA) K. HARZ det. [print, blue ink]”. ZMB: 1 @ (directly
pinned, with ootheca; left antenna, right antenna partly, left fore and right mid leg beyond coxae
and hind tarsi lacking) labelled “Dti Vecchia 5.IX.19 Fruhstorfer [print]”.

Discussion. Fruhstorfer (1921c: 78) mentioned further material from the
southern part of the Canton Ticino but did not specify the number of specimens he had.
He (1921c: 79) also examined specimens in the MHNG from “Hyères”, “Sizilien” and
“Curzole in Dalmatien” which could not be traced. The above specimens match the
data given for only one species of Ectobius, E. neolividus, and are thus considered as
syntypes. The collecting date on the labels of the two specimens from Soglio is
indicated as beginning (= “Anf.”) of August whereas it was quoted as end of July by
Fruhstorfer (1921c: 78). Because both dates essentially refer to the same time period,
this minor discrepancy does not, in our opinion, disqualify the specimens as syntypes.

616 H. BAUR & A. CORAY

E. neolividus has been introduced with “Ramme & Fruhstorfer” as authors. But
from the contents the contribution of Ramme is not evident. It is also noteworthy in
this context, that Ramme heavily criticized Fruhstorfer who, apparently, had mis-
understood “Meine ausdrücklich als unverbindlich gekennzeichneten Mitteilungen...”
on Ectobius (Ramme, 1923: 107). Hence, we consider Fruhstorfer as the only person
responsible for both the name and the criteria of availability (ICZN, Art. 50.1), as was
done by subsequent authors (Princis, 1971; Harz & Kaltenbach, 1976; among others).

E. neolividus is currently considered to belong to E. vittiventris (see Ramme,
1923; Harz & Kaltenbach, 1976; Baur et al. 2004). This species is rather variable and
might consist of a complex of sibling species (Bohn, pers. comm.). Therefore, we have
refrained from the designation of a lectotype for E. neolividus.

Status. Junior synonym of Ectobius vittiventris (A. Costa, 1847) (Ramme, 1923:
130).

ENSIFERA

PHANEROPTERIDAE

Barbitistes obtusus alpinus Fruhstorfer

Barbitistes obtusus alpina [sic] Fruhstorfer, 1920d: 77, 82, 85. Lectotype d, Switzerland,
Canton Ticino: Monte Generoso, Crocetta, here designated (CAN [examined]).

Material. Lectotype (directly pinned, right antenna partly and left hind tibia partly lack-
ing) labelled “Generoso-Crocetta 29.VII.19 Fruhstorfer [print]”.

Paralectotypes, ZSM: 1 d (directly pinned) labelled “Generoso-Crocetta 8.VIII.19
Fruhstorfer [print]; Barbitistes obtusus Targ.- Toz. det. D. R. Ragge. 1959 [print]”. ZSM: 1 2
(directly pinned) labelled “Generoso-Crocetta 1.IX.19 Fruhstorfer [print]”, “Barbitistes ob-tusus
[sic] Targ. Toz. 9 [hand] det. K. Harz [print]”.

Discussion. Fruhstorfer introduced Barbitistes obtusus alpinus in the “Tessiner
Wanderbilder” (1920d: 77) where he provided a brief definition of the taxon using the
following statement: “Der Barbitistes aber stellte sich beim Vergleich mit einem
Anfang Juli bei Chiasso gefundenen Exemplar als eine interessante, verkleinerte und
verdunkelte alpine Rasse von obtusus heraus, jetzt alpina Fruhst.”. The connotation as
a race (“Rasse”) formally makes it a subspecific name. A more detailed diagnosis of B.
o. alpinus (note that Barbitistes is masculine, hence B. o. alpinus) was given later by
Fruhstorfer (1921c: 179). He (1920d: 77, 82, 85) collected 2 d,1 £ on three occasions
during July to September 1919 near the south-western slope of Crocetta. The place is
situated at about 1360 m altitude in the northern part of the Monte Generoso massif.
The labels fit exactly the data given by Fruhstorfer (1920d, 1921c) for only one taxon
of Barbitistes, B. o. alpinus. The lectotype is the smallest specimen of the type series
and is thus closest to Fruhstorfer’s definition (1920d: 77). He also explicitly referred to
the respective male on the same page. In order to ensure stability in the application of
this name, we have chosen this specimen as the lectotype (ICZN, Art. 74.7.3).

Examination of the type series and much additional material (CAN, ETHZ,
MHNG, NHMB, NMBE, ZSM) confirms the view of Nadig er al. (1991) that B. o.
alpinus cannot be separated from B. obtusus which is widely distributed in the Canton
Ticino (Thorens & Nadig, 1997).

Status. Junior synonym of Barbitistes obtusus Targioni-Tozzetti, 1881 (Nadig et
MOI 238):

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 617

ne 2

mio L'ontchss | = ES done
Fam 21. Lectotype +
= ae
PEFISBIE sou . Fralet «for 1324 :
SES ANS HARZ ser. | Ides. H. Baur 200%
2a-C 3a-e
Fics 1-4

Labels on specimens, original size (our labels were omitted, except for figure 3e): (1) Para-
lectotype 2 of Podisma alpina formosanta Fruhstorfer; Fig. 1b is a bottom label in Fruhstorfer’s
handwriting. (2) Syntype of Ectobius neolividus Fruhstorfer; Fig. 2b is an original identification
label by Fruhstorfer. (3) Lectotype of Ephippiger plinianus Fruhstorfer; Fig. 3b is a data label in
Fruhstorfer’s handwriting. (4) Lectotype of Ephippiger cruciger eustratius Fruhstorfer; Fig. 4d
shows a different handwriting and is probably not an original bottom label.

TETTIGONIIDAE

Ephippiger cruciger eustratius Fruhstorfer
Ephippigera crucigera eustratia Fruhstorfer, 1921c: 231. Lectotype d, Switzerland, Canton

Bern: Moutier, Klus, here designated (ETHZ [examined]).

Material. Lectotype d (directly pinned, left antenna partly, left hind tarsus partly and
right hind leg beyond coxa lacking, titillators damaged at base by pest insects) labelled “Type
[print, yellow label with border]; 20.IX.19. Moutier Klöti [hand, label with border]; Eph.
ephippiger [print] d [hand] Fieb. [print] vitium Harz f. silvicola [hand] det. Nadig [print]”. Two
further labels were placed near the specimen. One label reads “var eustratia Fruhst. [hand, grey
label with double border]”, the other one shows a note by Nadig “wohl eher: Eph. ephippiger F.
var silvicola Az. Febr 1932 (Nadig.) vide Mit. S. E. G. 1932 [hand, label with double border]”
(Fig. 4).

Paralectotype, ETHZ: 1 d (directly pinned) with similar data to lectotype, considered as
part of the type series.

Discussion. Fruhstorfer (1921c: 231) mentioned material from “Solothurner
Jura, Moutier” but did not specify the number of specimens he had. However, his short
description was actually based on material which was collected in 1919 by E. Klöti-
Hauser “an der Strasse die von Moutier durch die Birsschlucht nach Choindez führt”
(Klöti-Hauser, 1922, see also Nadig, 1932). Of the seven males collected by Klöti-
Hauser, only two survived in the ETHZ collection. The lectotype fits the original
description and is designated to ensure stability in the application of this name (ICZN,
Art. 74.7.3). The composition of the entire type series is uncertain and might include

618 H. BAUR & A. CORAY

more than one taxon. The lectotype designation makes it clear to which taxon this
name actually refers. The names of this and the following Ephippiger taxa first
appeared in combination with Ephippigera Serville, 1831, an unjustified emendation
of Ephippiger Berthold, 1827. These names have to be treated as though they were
published in combination with Ephippiger (ICZN, Art. 51.3.1).

Re-examination of the type series and additional material (Appendix) fully
confirms the synonymy of E. c. eustratius with E. diurnus Dufour (see Nadig, 1932,
sub E. vitium).

Status. Junior synonym of Ephippiger diurnus Dufour, 1841 [= E. vitium of
authors, not Serville, 1831]! syn. n.

Ephippiger persicarius Fruhstorfer Fig. 5

Ephippigera persicaria Fruhstorfer, 1921c: 230, 33. Syntypes dd? 2, Switzerland, Canton
Ticino: Monte Carasso, above Monti di Freghiscio and Alpe di Orina over Cugnasco
(?depository [not located]).

Discussion. Fruhstorfer (1921c: 230, 33) mentioned one female from Monte
Carasso but did not specify the number of specimens he had from Alpe di Orina. The
two localities are separated only by a distance of about three kilometres and are
situated on the southern slopes of elevations north of the Piano di Magadino and the
Ticino river. Monte Carasso is the name of a village at the bottom of the Ticino valley
at about 230 m altitude. But according to Fruhstorfer (1921c: 33, 230), the specimen
was found above Monte Carasso and near Monti di Freghiscio at about 1600-1700 m
altitude. There is only one place at this altitude which is devoid of forest and therefore
suitable as a habitat of Ephippiger, that is the south-western slope of the Cima della
Pianca (Swiss grid co-ordinates 717.9/119.3, 1600-1700 m, see Appendix).

Of the two Ephippiger species described by Fruhstorfer from the Canton Ticino,
a few female syntypes of E. plinianus are kept at the ZSM (see below) but no specimen
of E. persicarius could be traced so far. This is in contrast to Nadig’s claim (1960: 28,
footnote) that “Die Type von E. pliniana [sic] ist unauffindbar (Nadig, 1931), jene von
E. persicaria [sic], ein £ steckt ... in der Zoolog. Sammlung des bayerischen Staates
in Miinchen [= ZSM]”. We thus assume that Nadig simply confounded the species
names in his footnote.

The information given by Fruhstorfer (1921c) concerning his Ephippiger
material from the Canton Ticino is rather confusing and sometimes contradictory (see
Zeuner, 1931 for a critical discussion). His descriptions of the new species, E. persi-
carius and E. plinianus, are short and insufficient for their unambiguous recognition,
original material is scarce. These shortcomings led to serious difficulties concerning

! Audinet-Serville (1831) introduced E. vitium as a replacement name for Locusta
ephippiger Fabricius (= Gryllus ephippiger Fiebig, see Coray & Lehmann, 1998: 88), which is
why the name has to be considered as a junior objective synonym of E. ephippiger (Fiebig)
(ICZN, Art. 72.7). Hence, the name virium Serville cannot be retained to denote the Western
European populations of the E. ephippiger species-aggregate, despite claims to the contrary in
some recent works (Coray & Lehmann, 1998; Coray & Thorens, 2001). For the taxon in
question, E. diurnus Dufour has been adopted (Kruseman, 1988; Oudman er al., 1990; Ingrisch
& Köhler, 1998; all sub E. ephippiger diurnus).

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 619

the identity of the taxa involved and resulted, as we will show below, in the unjustified
neglect of E. persicarius by Harz (1966) and subsequent authors.

In an attempt to clarify the identity of E. persicarius, it is important to consider
the distribution of the Ephippiger species in the Canton Ticino. More recent
investigation have shown that only two species occur there, E. vicheti Harz and E. bor-
mansi Brunner von Wattenwyl, of which detailed data on their horizontal and vertical
distribution are available (Nadig, 1968; Thorens & Nadig, 1997; Baur, Rösti, pers.
observation). A careful evaluation of these data in combination with a re-examination
ofthe remaining material of the Fruhstorfer collection gave us new insights with regard
to the identity of the taxa. Thus, it is evident from the type material deposited in the
ZSM that E. plinianus belongs to E. bormansi (see below). Furthermore, we agree with
Nadig (1968: 343) that specimens in the ETHZ and the NHMW identified by
Fruhstorfer as E. perforatus (Rossi) (see Appendix) belong to E. vicheti and not to a
third species. The identity of E. persicarius, finally, is less obvious because of a lack
of syntypes and the poor original description. Fruhstorfer (1921c: 230) emphasised the
violet or peachy coloration of his specimens which he collected late in the year. We
observed that topotypical specimens are indeed darker in October (Fig. 5) than in the
months before (Baur, Rösti, pers. observation). As pointed out by Zeuner (1931: 33, 34,
sub E. vitium) the other characters given by Fruhstorfer lie within the range of variation
of E. vicheti. Moreover, it is most revealing that only one species, E. vicheti, is present
in the region where the two type localities of E. persicarius are situated. This is evident
from the detailed studies by Nadig (1968) and Thorens & Nadig (1997) and from
recent collections made by the senior author at the type localities. Following these
investigations E. bormansi is known to occur only south of the Ticino river, where it is
found on sunny slopes in grass and small shrubs from about 900 up to 1900 m altitude.
E. vicheti is mostly a vicariate of E. bormansi that lives from the bottom of the valleys
up to about 1100 (-1450) m altitude in fern (Pteridium), bushes and lower trees. But in
places such as the elevations north of the Ticino river, where E. bormansi is absent, E.
vicheti is also encountered at higher altitudes up to 1650 m (Nadig, 1968; Baur, Rösti,
pers. observation). In conclusion, these findings leave no doubt that E. persicarius has
to be regarded as a senior synonym of Ephippiger vicheti Harz, 1966 syn. n. (see also
Nadig, 1968: 343, footnote). The identity of populations from the respective type
localities has been demonstrated by several authors (Harz, 1966; Nadig, 1987) and is
confirmed by examination of the holotype of E. vicheti (from Italy, Trentino, Storo) and
much additional material from the respective regions (Appendix).

E. persicarius evidently forms a diagnosably distinct taxon of the allopatric
E. ephippiger species-aggregate (compare the distribution map in Detzel, 1998: 283).
It occurs on the southern side of the Alps from the Piedmont (Italy) in the west (Nadig,
1987) to Styria (Austria) in the east and Istria (Croatia) in the south-east (Harz, 1966).
It is separated from the Western European E. diurnus Dufour (type locality St.-Sever
in France, Dept. Les Landes) by song characteristics (Duijm, 1990), the absence of a
pronounced color polymorphism in adults (Hartley & Bugren, 1986), less strongly
pointed male cerci, shorter titillators and the lack of paired sclerotized plates on the
female subgenital plate (compare Duijm & Oudman, 1983, figures 1-2,4; sub E.
ephippiger vicheti). The latter character state is shared with the eastern European E.

620 H. BAUR & A. CORAY

ephippiger (Fiebig) (type locality Vienna in Austria) but this species has much longer
titillators (see Nadig, 1987: 332, figure 16) and the apical part of the male cerci strong-
ly pointed as in E. diurnus. Alleged character transitions in the Eastern refuge of
E. persicarius (Harz, 1966, sub E. e. vicheti) might be the result of secondary hybrid
zones which are irrelevant with regard to the classification of taxa as phylogenetic
species (Cracraft, 1989).

Our findings are in contrast to those of Hartley & Warne (1984) who syno-
nymised E. persicarius (sub E. e. vicheti) with E. ephippiger. Their view probably led
to the disregard of E. persicarius in a few subsequent works (Heller, 1988; Heller et
al., 1998; Ragge & Reynolds, 1998), although in the most recent studies E. persicarius
(sub E. vicheti) has been considered as a distinct taxon (Nadig, 1987; Otte, 1997;
Thorens & Nadig, 1997; Ingrisch & Köhler, 1998; Coray & Thorens, 2001; Fontana et
al., 2002; among others). We would like to stress that the proposed synonymy of
Hartley & Warne is actually based on too little evidence (see also Nadig, 1987).
Apparently, they (1984: 46) examined just “a few specimens” of E. persicarius from
Lago Maggiore (Northern Italy) and another two specimens from Skopje (Macedonia)
which they regarded as E. ephippiger but no topotypical material at all.

Status. Ephippiger persicarius Fruhstorfer, 1921c stat. rev.

Ephippiger plinianus Fruhstorfer Fig. 6

Ephippigera pliniana Fruhstorfer, 1921c: 233. Lectotype 9, Switzerland, Canton Ticino: Val
Morobbia, Monti Piandolce, here designated (ZSM [examined]).

Material. Lectotype ® (directly pinned, left antenna partly, right fore tarsus partly and
right mid tarsus lacking) labelled “Type [print, yellow label with border]; Piandolce 31.X.18
[hand] Fruhstorfer [print]; Type von [print] Epphippigera [sic] Pliniana Fruh [hand, orange
label]; Fruhstorfer Orth. d. Schwz [sic] (1921c) p. 233. [back side of former label]; Holotypus
Zool. Staatsammlg. München [print, red with black border]” (Figs 3, 6).

Paralectotypes, ZSM: 2 © (directly pinned) labelled “Mti Croce 29.X.18 [hand, blue ink]
Fruhstorfer [print]; ...” from Monti della Croce at Motto d’Arbino over Bellinzona (Fruhstorfer,
1921c: 234).

Discussion. It is likely that only the second label on the lectotype is from
Fruhstorfer himself. The label “Holotypus ...” has certainly been added later. However,
the information content of the data label matches exactly the data given by Fruhstorfer
(1921c: 234) for only one species of Ephippiger, E. plinianus. He mentioned also
specimens from Monte il Gaggio and Monte Boglia but did not specify the number of
specimens he had. The lectotype fits the original description and is designated to ensure
stability in the application of this name (ICZN, Art. 74.7.3). The composition of the
entire type series is uncertain and might include more than one taxon. The lectotype
designation makes it clear to which taxon this name actually refers.

Re-examination of the lectotype and much additional material (Appendix) fully
confirms the view of authors (Nadig, 1960, 1980; Harz, 1969) that E. plinianus is the
same as E. bormansi which is widely distributed in southern parts of the Canton Ticino
(Thorens & Nadig, 1997).

It is unclear whether E. bormansi is really diagnosably distinct from E. ter-
restris (Yersin). We have not examined enough material of the latter to decide this
question, which is why we rely on the detailed investigations by Nadig (1980) and

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 621

Fics 5-6

(5) A female of Ephippiger persicarius Fruhstorfer from Monti della Gana in Canton Ticino, a
place very close to one of the two type localities, Alpe di Orino. The spread of violet to peachy
tinges on pronotum and abdomen increases with age and characterizes many older specimens in
autumn. According to the original description, the missing syntypes may have had a similar
appearance. Photograph D. Rösti. (6) Lectotype female of Ephippiger plinianus Fruhstorfer, a
Junior synonym of E. bormansi Brunner von Wattenwyl. This species also occurs in the Canton
Ticino but is absent from the region where the type localities of E. persicarius are situated. Scale
1 cm. Photograph L. Schäublin.

622 H. BAUR & A. CORAY

Landman er al. (1989). According to these studies the two species are well separated
over most of their ranges but form a broad hybrid zone in the south-western part of the
Alps. Clinal variation has been observed in several important features within each ta-
xon. On the other hand, Nadig (1980) asserted that the hybrid zones are likely to be the
result of secondary contact. Clearly further work is necessary to solve these problems.
For the time being, we keep E. bormansi as a separate taxon, as was done in some re-
cent works (Nadig, 1987; Otte, 1997; Thorens & Nadig, 1997; Ingrisch & Köhler,
1998; Coray & Thorens, 2001).

Status. Junior synonym of Ephippiger bormansi Brunner von Wattenwyl, 1882
(Harz, 1969: 514).

Thamnotrizon chabrieri malachiticus Fruhstorfer
Thamnotrizon chabrieri malachiticus Fruhstorfer, 1921c: 209, 210. Syntypes £ £, Switzerland,

Canton Ticino: Ligornetto-Tremona and Ligornetto-Meride (CAN, ETHZ [examined]).

Material. Syntypes, CAN: 1 ® (directly pinned, right antenna partly lacking) labelled
“Ligornetto-Meride 17.IX.19 Fruhstorfer [print]; chabrieri Ch. [print, not original]”. ETHZ: 1 9
(directly pinned, left antenna partly and left hind tibia partly lacking; left hind leg pinned below
specimen) labelled “Ligornetto-Tremona Aug. 19 Fruhstorfer [print]”.

Discussion. Fruhstorfer first introduced the name malachiticus in the “Tessiner
Wanderbilder” (1920d: 30, 35, 38, 81) but it was not thereby made available. This hap-
pened only later (Fruhstorfer, 1921c: 210) when a definition of the taxon was provided.
Fruhstorfer mentioned several males and females from Ligornetto, Mendrisio and
Meride in the southern parts of the Canton Ticino but did not specify the total number
of specimens he had. However, only the above two female syntypes could be traced.
They fit the original description and are obviously conspecific. Because the deli-
mitation of species in this genus is almost entirely based on the male genitalia, we
prefer to refrain from the designation of a lectotype.

According to Nadig (1985a: 176) Eupholidoptera chabrieri (Charpentier)
occurs in the Insubrian region, i. e. the region south of the Alps between the Lago
Maggiore and the Lago di Como (including the southern parts of the Canton Ticino).
However, Nadig also stressed that the Insubrian specimens differed slightly from topo-
typical material in the shape of titillators which is why he erected an infrasubspecific
name for the Insubrian populations. In case future investigations should substantiate
the existence of a separate taxon, the name 7. c. malachiticus is available.

Status. Junior synonym of Eupholidoptera chabrieri (Charpentier, 1825) syn. n.

CAELIFERA
ACRIDIDAE

Chrysochraon brachypterus chrysoberyllus Fruhstorfer

Chrysochraon brachypterus chrysoberyllus Fruhstorfer, 1921c: 95, 96. Lectotype 9, Switzer-

land, Canton Ticino: Pizzo Claro, here designated (ETHZ [examined]).

Material. Lectotype © (directly pinned, entire) labelled “Claro 9.VIII.18 [hand, blue
ink] Fruhstorfer [print]”.

Paralectotypes, ETHZ: 19 3, 26 © (directly pinned) from “Piandolce”, “Generoso”,
“Generoso-Crocetta”, “Ligornetto-Meride”, “Dti Vecchia”, “Pizzo Leone”, “Mt. Boglia”, “Mt.
Croce” and “Mt. Carasso”.

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 623

Discussion. Fruhstorfer (1921c: 95) mentioned material from many localities in
the Canton Ticino but did not specify the number of specimens he had. The lectotype
fits the original description very nicely and is designated to ensure stability in the
application of this name (ICZN, Art. 74.7.3). The composition of the entire type series
is uncertain and might include more than one taxon. The lectotype designation makes
it clear to which taxon this name actually refers.

Fruhstorfer (1921c: 96) considered chrysoberyllus as a “Südrasse” which he
separated from specimens north of the Alps solely by the emerald coloration of the
female fore wing. However, our analysis of an extensive material (CAN, ETHZ,
MHNG, NHMB, NMBE, ZSM) revealed that it is actually impossible to separate
chrysoberyllus from E. brachyptera (Ocskay) which is widely distributed in
Switzerland (Thorens & Nadig, 1997).

Status. Junior synonym of Euthystira brachyptera (Ocskay, 1826) (Yin et al.,
1996: 286).

Omocestus haemorrhoidalis fantinus Fruhstorfer
Omocestus haemorrhoidalis fantinus Fruhstorfer, 1921c: 110. Lectotype 3, Switzerland,

Canton Ticino: Pizzo Claro, here designated (ETHZ [examined]).

Material. Lectotype d (directly pinned, right antenna lacking) labelled “Claro. 9.VIIL.18
[hand] Fruhstorfer [print]”.

Discussion. Fruhstorfer (1921c: 110) mentioned material from many localities
in the Canton Ticino but did not specify the number of specimens he had. The lecto-
type fits the original description and is designated to ensure stability in the application
of this name (ICZN, Art. 74.7.3). The composition of the entire type series is uncertain
and might include more than one taxon. The lectotype designation makes it clear to
which taxon this name actually refers.

O. h. fantinus is obviously the same as O. haemorrhoidalis (Charpentier) which
is widespread in southern Switzerland (Thorens & Nadig, 1997). Examination of many
specimens (CAN, ETHZ, MHNG, NHMB, NMBE, ZSM) showed that the differences
in size and coloration mentioned by Fruhstorfer (1921c: 110) are rather variable and do
not allow the separation of his taxon.

Status. Junior synonym of Omocestus haemorrhoidalis (Charpentier, 1825)
(Yin et al., 1996: 462).

Podisma alpina formosanta Fruhstorfer

Podisma alpina forma formosanta Fruhstorfer, 1921c: 164f., 167. Lectotype d, Switzerland,
Canton Ticino: Monte Generoso, Camoscé, designated by Nadig (1989: 206) (ETHZ
[examined]).

Material. Lectotype d (directly pinned, entire) labelled ‘“Generoso-Camoscé, 22.IX.19
Fruhstorfer [print]; M. formosanta FRUHST. det. A. Galvagni 19 [print]; M. for. formosanta
(FRUHST.) [print] & [hand] det. NADIG [print]; LECTO-Holo-Typus [print, red label with
black border]”

Paralectotypes, CAN: 4 d, 5 9 (directly pinned) from “Bedretto [one male with iden-
tification label by Fruhstorfer]”, “Pizzo Leone [one female with original bottom label of the
Fruhstorfer coll. (Fig. 1b)]”, Buffalora”, “Tamaro”. ETHZ: 10 6, 25 ® (directly pinned) from

624 H. BAUR & A. CORAY

“Generoso-Camoscé”, “Generoso-Crocetta”, “Val Osogna”, “Bedretto”, “Passo Predelp”,
“Generoso”, “Tamaro”, “Pizzo Leone”, “Buffalora” and “Monte Boglia”. CKH: 1 d, 12
(directly pinned) from “Bedretto”.

Discussion. Fruhstorfer first mentioned “Podisma alpina forma formosanta” in
the “Walliser Wanderbilder” (1920a: 44) but the name was not thereby made available.
This happened only later (1921c: 164) when a description of the taxon was provided.
Fruhstorfer (1921c: 167) mentioned material from several localities in the Canton
Ticino but did not specify the number of specimens he had. Although the name has
been introduced as a “forma” it must be considered subspecific (ICZN, Art. 45.6.4.1).

Mistshenko (1952: 397) first applied the name formosanta to a subspecies of
Miramella alpina (Kollar). Nadig (1986) finally adopted it for a species of which he
(1989) later gave a thorough analysis of the morphology and distribution.

Status. Miramella (Nadigella) formosanta (Fruhstorfer, 1921c) (Nadig, 1989:
206).

Podisma alpina irena Fruhstorfer

Podisma alpina forma irena Fruhstorfer, 1921c: 251. Syntypes dd 9 9, Italy, South Tyrol:

Penegal and St. Vigil (?Depository [not located]).

Discussion. Fruhstorfer (1921c: 251) mentioned material collected by himself
from Penegal and by W. Ramme from St. Vigil but did not specify the number of
specimens he had. From St. Vigil 5 4,5 9 are present in the ZMB and a further 1 à,
1 2 in the ZSM. Unfortunately, the specimens lack any original labels by Fruhstorfer,
which is why the material cannot be recognized as part of the type series with certainty.
Harz (1975) and Nadig (1989) considered the type series to be lost. Furthermore, the
neotype designation of Harz (1975: 290) is invalid because it fails to meet the quali-
fying conditions of the Code (ICZN, Art. 75.3). Instead of a single specimen Harz
selected a pair of neotypes which had not become the property of a scientific institution
immediately upon publication but remained in his private collection. However, we
think that the lack of type material is not problematic here. Fruhstorfer indicated the
origin of his specimens very precisely which made it possible for Galvagni (1986) and
Nadig (1989) to study topotypical material. Following their investigations, only one
species of Miramella is known from these localities. Hence, we cannot see any excep-
tional circumstances (ICZN, Art. 75) that would justify the designation of a neotype.

The taxonomic history of P. a. irena is nevertheless complicated and requires
further discussion. Galvagni (1954) was the first to apply the name for a subspecies of
Miramella Dovnar-Zapolskij, M. carinthiaca irena. With this action he made this name
definitely available (ICZN, Art. 45.6.4.1). Galvagni’s classification was followed by
Harz (1973, 1975) who included M. c. irena in his new subgenus Kisella Harz, 1973.
Nadig (1985b), following the opinion of Galvagni (in litt.), established the currently
accepted status as a separate species, M. irena. Galvagni (1986) refined the concept of
the species and also found that he had partly misidentified M. irena as M. carinthiaca
in his earlier paper (Galvagni, 1954). Finally, Nadig’s (1989) monograph on Central
European Miramella is most useful, because he meticulously sorted out the morpho-
logical and distributional limits of M. irena and related taxa such as M. carinthiaca.
However, the fact that the two species had been confounded earlier by Galvagni (1954)

BLATTODEA, ENSIFERA AND CAELIFERA DESCRIBED BY H. FRUHSTORFER 625

apparently led to the misidentification of the type species of Kisella, Podisma alpina
carinthiaca, by Harz (1973). Moreover, this taxon had notoriously been attributed to a
wrong author and date (Puschnig, 1910 instead of Obenberger, 1926!) since its
establishment. Hence, we address these problems, which are critical concerning the
identity of the taxa involved, in a separate paper (Baur & Coray, 2004, this volume).
Status. Miramella (Kisella) irena (Fruhstorfer, 1921c) (Nadig, 1989: 186).

Podisma frigida strandi Fruhstorfer

Podisma frigida strandi Fruhstorfer, 1921c: 159. Lectotype d, Switzerland, Canton
Graubünden: Muottas-Muragl, here designated (ETHZ [examined]).

Material. Lectotype 6d (directly pinned, entire) labelled “Muottas-Muraigl [sic]
2200-2400 m 10.8.20 Fruhstorfer [print]”.

Paralectotypes, CAN: 1 4 (directly pinned) from “Muottas Muraigl [sic]”. ETHZ: 2 3,
7 © (directly pinned) from “Muottas Muraigl [sic]”, “Sparrhorn”, “Bedretto” and “Belalp”.

Discussion. Fruhstorfer (1921c: 159) mentioned material from the Alpine belt
throughout Switzerland but did not specify the number of specimens he had. The above
mentioned specimens fit exactly the data given by Fruhstorfer and are thus considered
as part of the type series. The lectotype fits the original description and is designated
to ensure stability in the application of this name (ICZN, Art. 74.7.3). The composition
of the entire type series is uncertain and might include more than one taxon. The
lectotype designation makes it clear to which taxon this name actually refers.

The name Bohemanella frigida strandi had been used to denote the Alpine
populations of B. frigida (Boheman) (e.g. Harz, 1975, sub Melanoplus) but is now
treated as a synonym of the latter (Nadig er al. 1991; Ingrisch & Köhler, 1998 and
others). In case further studies would confirm a separate status of these Alpine
populations, the name Podisma prossenii Puschnig, 1910 should also be considered
(see Ebner, 1937: 149).

Status. Junior synonym of Bohemanella frigida (Boheman, 1846) (Nadig et al.,
1992251):

Stenobothrus lineatus fervidior Fruhstorfer

Stenobothrus lineatus fervidior Fruhstorfer, 1921c: 109, 108. Lectotype à , Switzerland, Canton
Ticino: Monte Bisbino, here designated (ETHZ [examined]).

Material. Lectotype 4 (directly pinned, left mid tarsus lacking) labelled “Mte Bisbino
21.VIIL.19 Fruhstorfer [print]”.

Paralectotypes, ETHZ: 16 d, 1 9 (directly pinned) from “Generoso-Crocetta”, “Ligor-
netto”, “Mt. Boglia” and “Mte Bar”. ZSM: 2 d (directly pinned) from “Mte Boglia”.

Discussion. Fruhstorfer (1921c: 108, 109) mentioned material from several
localities in the Canton Ticino and Valais but did not specify the number of specimens
he had. The name was introduced on p. 108 as “forma fervidior Fruhst.” but on the
following page Fruhstorfer denoted “fervidior subsp. nova” as a distinct Southern race
which also comprised the forma obscura Zacher. For this reason fervidior has to be
treated as a subspecific name. The lectotype fits well the characters given by
Fruhstorfer (1921c: 109) and is designated to ensure stability in the application of this
name (ICZN, Art. 74.7.3). The composition of the entire type series is uncertain and

626 H. BAUR & A. CORAY

might include more than one taxon. The lectotype designation makes it clear to which
taxon this name actually refers.

Fruhstorfer (1921c: 109) mainly stressed the intense, reddish coloration on hind
legs and apex of abdomen of S. /. fervidior. Examination of an extensive material
(CAN, ETHZ, MHNG, NHMB, NMBE, ZSM) revealed that this character is actually
highly variable and does not allow the separation of the taxon from the widespread
S. lineatus (Panzer) (Thorens & Nadig, 1997).

Status. Junior synonym of Stenobothrus lineatus (Panzer, 1796) syn. n.

ACKNOWLEDGEMENTS

We thank Sigfrid Ingrisch (Bad Karlshafen, Germany), Elsa Obrecht, (NMBE),
Daniel Rösti (Wasen 1. E., Switzerland), Christian Kropf (NMBE) and an anonymous
reviewer for critical reading of the manuscript and many useful suggestions. We are
grateful to Andrew Wakeham-Dawson (Executive Secretary, ICZN) for his guidance
concerning the selection of lectotypes. We are also indebted to Otto Kraus (Hamburg,
Germany) for his advice concerning the status of some taxa. The senior author would
specially like to thank D. Rösti for his assistance in field work and for taking some of
the photographs; for the other photographs we are grateful to Lisa Schäublin (NMBE).
For the loan of or information on specimens we thank Alfred P. Kaltenbach and Ulrike
Aspöck (NHMW), Horst Bohn (Zoologisches Institut, Ludwig-Maximilians
Universitàt, Munich, Germany), Daniel Burckhardt (NHMB), Charles Lienhard
(MHNG), Andreas Müller (ETHZ), Adolf Nadig (CAN), Michael Ohl (ZMB) and Klaus
Schönitzer (ZSM).

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APPENDIX

Material of Ephippiger examined in this study. The data of many specimens have
already been published by Nadig (1968, 1987) or Thorens & Nadig (1997) and are thus quoted
in short form.

E. bormansi, CAN: ITALY, Como: Val Sanagra, A. Livea 1 ©; Val Sanagra, S A. Erba
1 d; Monte Grona 2 d; CH, Canton Ticino, Monte Boglia 1 6; Lago di Como: Cortafongrat
3 4; Piemonte: V. d’ Arma, near Cle. di Muro 2 d; over Montaldo di Cosola, 1200-1250 m,
9.9.1990, leg. Nadig 2 d. SWITZERLAND, Canton Ticino: Val Morobbia, Melirolo 3 4, 3 9.

ETHZ: SWITZERLAND, Canton Ticino: Monte Boglia, leg. Fruhstorfer 1 4 larva; Val
da Colla, Corticiasca, Monte Bar, leg. Nadig 1 6, 1 9.

NMBE: SWITZERLAND, Canton Ticino: Miglieglia, Monte Lema, 1620 m,
707.9/99.75, 4.10.2000, leg. Baur 5 d, 3 2; Miglieglia, Mater delle Tagliadelle, 1160 m,
708.7/99.1, 4.10.2000, leg. Baur 6 d, 1 2; Monte Generoso, Pianconce, 1400 m, 722.1/86.4,
24.9.1998, leg. Baur 7 d , 1 2; Monte Generoso, Pianconce, 1400 m, 722.1/86.4, 29.9.1999, leg.
Baur 12 d.

E. diurnus, CAN: FRANCE, Alpes: Col de Perty 5 d,5 2; Guillestre 2 d; Provence:
La Sainte-Baumes, Gémenos 2 à ; Alpes Marittimes: Gréolières, Cheiron 3 4,3 9; Durence, N
Embrun 1 d; Var: Ampus-Chäteaudouble 2 6; Vaucluse: Méouge 1 6; Mont Ventoux 3 d;
Hérault: N Lodève 2 d. SWITZERLAND, Canton Vaud: Mont sur Rolle 1 & ; Sepey 1 6; Mont
sur les Truits 1 9; Canton Bern: Moutier 5 6, 1 9;

CKH: FRANCE, Bouches-du-Rhône: Marseille, surroundings 1 6; Mont Majour, 1 à;
Aveyron: Cause du Larzac, La Couvertoirade 1 6; Lozère: Marvyols 2 d, 2 9; Hérault: Mt.
Caroux 2 6; Nefiès 1 d.

ETHZ: FRANCE, Hautes-Provence: Gavarnie, leg. Sauter 3 d, 1 2; Dordogne:
Lascaux, leg. Sauter 3 d, 2 9; Vaucluse: Luberon, leg. Sauter 3 6; Puy de Dome: St. Victor,
leg. Sauter 1 d, 12. SWITZERLAND, Canton Bern: Moutier, leg. Nadig 1 d.

NHMB: FRANCE, Haut-Rhin: Rufach [= Rouffach], 8.1923 1 &,1 9.

NMBE: FRANCE, Les Landes: Lit-et-Mixe, July 1970, 1 2; Lozère: St-Martin-de-
Lansuscle, Château le Cauvel, 780 m, 12.7.1991, leg. Baur 3 d,2 9, ex larva; Var: Aiguines,
259198541007

630 H. BAUR & A. CORAY

ZSM: FRANCE, Isère: Grenoble, leg. Fruhstorfer 2 ¢.

E. ephippiger, CAN: AUSTRIA, Wien: Eichkogel 1 d, 2 2; Wien 2 &;
Niederösterreich Hainburger Berge, Hundsheimerberg 1 dé; Mautern-Rosatz 1 &;
Guntramsdorf-Richardshof 1 9; HUNGARY, Budapest: Harmashatargh. 1 d,1 2; MACEDO-
NIA, Skopie: Vodno 1 d,5 2; Sar Planina: Prevalao-Mt. Bistra 1 6; Vratnika 2 6,7 9;
Krusevo 1 4; SLOVENIA, Gracnicatal: E Rimske Toplice: 1 2; Gorjanci 2 6, 1 9.

CKH: AUSTRIA, Wien: Eichkogel bei Mödling 1 6; CZECH REPUBLIC, Moravia:
Mohelno 1 4, 1 2; HUNGARY: Pillisszanto 1 6; SERBIA: Podgorica [= Titograd] 1 ®;
MACEDONIA, Bitola 2 6, 1 2.

NMBE: AUSTRIA: Wien, leg. Brunner v. W. 1 d.

E. persicarius, CAN: ITALY, Brianza: Eupilio-M. Cornizzolo 6 6, 6 9; N Brescia:
Lumezzane 1 6, 1 9; Prealpi Bresciane: Capovalle, Monte Stino, 1400-1466 m, 31.8.1992, leg.
Nadig 2 6, 2 2; Como: Vestreno, Madonna Bondo 2 6, 1 2; Chiavenna, Menarola 3 6,1 9;
Dorio, Perdonasco 1 d ; Piemonte: S Biella, La Bessa 2 d,1 2; Lombardio: Apennino, Pso. del
Penice, 1100-1150 m, 8.9.1990, leg. Nadig 2 d, 1 9 ; Pallanza: Cresta della Ceresa 2 & ; Trarego:
Cannero 1 6; SWITZERLAND, Canton Ticino: Bré, Aldesago 2 2; Bré-M.Boglia, Castra 4 9;
Gola di Lago 2 d; CH, Ticino, Bré-Val Morobbia 2 3; Locarno, Cardada 2 4,2 9; N Piano di
Magadino, Sassariente-Cima Sassello 2 d, 1 2; Monte Generoso 1 d; Val d’Arbedo, Monte
Fogal2 8,22%

ETHZ: SWITZERLAND, Canton Ticino: Salvatore 1 9 ; Generoso-Crocetta 1 6, 1 9,
S. Stefano-Balerna 2 ?, Capolago-Meride 3 2, Arzo-Meride 1 ©, Ligornetto-Tremona 2 d, 5
9, Ligornetto 1 9, Losone 1 ?, all leg. Fruhstorfer; San Salvatore, leg. Sauter 2 d; 1 2, Monte
Caslano, leg. Sauter 1 d ; Maroggia, leg. Krüger 1 G.

NHMB: ITALY: Valtellina, Tirano 1 4. SWITZERLAND, Canton Ticino: Mendrisio 1
9, larvae 1 9, Monte Generoso, Pianca comune, 1280 m, 9.8.1988, leg. Coray 1 d.

NHMW: ITALY, Trentino: Storo, holotype d, paratypes 1 4,2 © of E. vicheti Harz [tit-
tilators of holotype were missing!]; SWITZERLAND, Canton Ticino: Capolago-Meride 1 9,
Ligornetto-Tremona | © [both © 2 Fruhstorfer leg. and det. as “E. perforatus”].

NMBE: SWITZERLAND, Canton Ticino: Monte-Carasso, Cima della Pianca, 1660 m,
717.9/119.3, 28.8.2001, leg. Baur & Rösti 4 d; Cugnasco, Monti della Gana, 1400 m,
713.5/113.5, 4.10.2002, leg. Baur & Rôsti 9 d, 2 2; Gudo, Monti di Boscaloro, 1100 m,
713.3/116.5, 2.10.2002, leg. Rösti 1 d, 1 9; Gudo, Fontane di Biasca, 1200 m, 717.1/116.8,
3.10.2002, leg. Baur & Rösti 1 d, 1 2; Val Onsernone, Barione, 13.10.1961, leg. Huber 1 9;
Monte Generoso, Pianconce, 1400 m, 722.1/86.4, 24.9.1998, leg. Baur 4 d, 4 9; Monte
Generoso, Pianconce, 1400 m, 722.1/86.4, 29.9.1999, leg. Baur 1 9; Miglieglia, Mater delle
Tagliadelle, 1160 m, 708.7/99.1, 4.10.2000, leg. Baur 1 d; Miglieglia, N Frecc, 850 m,
708.8/98.25, 4.10.2000, leg. Baur 2 3; Meride, SE Cassina 860 m, 717.3/84.65, 28.9.1999, leg.
Baur 3 d,1 9; 1.10.1999, leg. Baur 2 d, 1 2; Meride, SSE San Giorgio, 1020 m, 717.4/85.4,
23.9.1998, leg. Baur 1 5; Lugano, Lurenzin, 1200 m, 716.6/97.8, 13.9.2001, leg. Rôsti 1 à.

ZSM: SWITZERLAND, Canton Ticino: Ligornetto, 28.7.1919, leg. Fruhstorfer 1 9;
Generoso-Camoscé, 22.9.1919, leg. Fruhstorfer 1 9; Capolago-Meride, 7.9.1919, leg.
Fruhstorfer 1 d.

REVUE SUISSE DE ZOOLOGIE 111 (3): 631-642; septembre 2004

The status of some taxa related to Miramella irena (Fruhstorfer) and
the type of Kisella Harz (Caelifera: Acrididae: Melanoplinae)

Hannes BAUR! & Armin CORAY?
1 Department of Invertebrates, Natural History Museum, Bernastrasse 15,

CH-3005 Bern, Switzerland. E-mail: hannes.baur@nmbe.unibe.ch (correspondence)
2 Natural History Museum, Augustinergasse 2, CH-4001 Basel, Switzerland.

E-mail: armin.coray@balcab.ch

The status of some taxa related to Miramella irena (Fruhstorfer) and
the type of Kisella Harz (Caelifera: Acrididae: Melanoplinae). - The
present paper provides taxonomic notes on four taxa of the genus Miramella
Dovnar-Zapolskij, 1933. Two species, M. irena (Fruhstorfer) and M. carin-
thiaca (Obenberger), which have often been confused are keyed out and
illustrated. A neotype is designated for M. carinthiaca to provide a firm
basis for the application of this name. We also show that this name has
wrongly been attributed to Puschnig, 1910 (instead of Obenberger, 1926) by
all subsequent authors. The currently accepted interpretation of the two taxa
furthermore requires the designation of a lectotype for Pezotettix alpinus
collinus Brunner von Wattenwyl in Künstler, 1864, which is thus fixed as a
junior synonym of M. alpina (Kollar, 1833). Finally, the type species of the
subgenus Kisella Harz, 1973 of Miramella, M. carinthiaca (Obenberger)
[= Podisma alpina carinthiaca Obenberger, 1926], had been misidentified
by Harz. We therefore validly fix the taxonomic taxon involved in the
misidentification, M. irena (Fruhstorfer) [= Podisma alpina irena Fruh-
storfer, 1921], as the type species of Kisella.

Keywords: Miramella carinthiaca - Pezotettix alpinus collinus - lectotype
designation - neotype designation - type species fixation - taxonomy.

INTRODUCTION

In the course of a taxonomic revision of the Caelifera described by H.
Fruhstorfer (Baur & Coray, 2004, this volume) we encountered a complex taxonomic
problem which is related to one of the revised taxa, Miramella irena (Fruhstorfer)
[= Podisma alpina irena Fruhstorfer, 1921]. In the past this species had been
confounded by several authors with M. carinthiaca (Obenberger) [= Podisma alpina
carinthiaca Obenberger, 1926a] (see Galvagni, 1986a) which, in turn, led to the
misidentification of M. carinthiaca as the type species of the subgenus Kisella Harz of
Miramella and to a general instability in the application of these names (Galvagni,
1986b; Nadig, 1989). Here, we address the problems associated with those taxa. The
two species, M. irena and M. carinthiaca, are keyed out and illustrated in order to

Manuscript accepted 18.03.2004

632 H. BAUR & A. CORAY

facilitate their recognition. We also designate a neotype for M. carinthiaca and show
that this name had hitherto been attributed to the wrong author and date. Furthermore,
we select a lectotype for Pezotettix alpinus collinus Brunner von Wattenwyl in
Künstler, 1864, a nominal taxon with an ambiguous status (Galvagni, 1986a). Finally,
we validly fix the type species of Kisella Harz in accordance with Art. 70.3 of the
International Code of Zoological Nomenclature (Fourth edition, 1999, ICZN). For
information concerning material and methods and abbreviations of depositories we
refer to Baur & Coray (2004, this volume).

Podisma alpina carinthiaca Obenberger

Podisma alpinum var. carinthiacum [sic!] Obenberger, 1926a: 179, first available use of
Podisma alpina var. alpina subvar. carinthiaca Puschnig, 1910: 27, 28, unavailable
name. Neotype d, Austria, Carinthia: Nockberge, Grundtal-Grundalm, here designated
(CAN [examined]).

Material. Neotype 4 (Fig. 1, directly pinned, right antenna and left mid tarsus lacking,
genital complex exposed) labelled “A: Kärnten: 82:109 Nockgebiet: Grundtal 1750 m 25.8.82
leg. NADIG [print]; 82: 109 No. 2 ([print] 6 [hand]) MIKROPH. Kopf,Elytra Penis v. Seite
[print, yellow label]; M. ...[deleted word] carin-thiaca [sic] (PUSCH.) [print] ¢ [hand] det.
NADIG [print]; TOPO-Typus [print, red label with black border]” (Fig. 2).

Discussion. The name carinthiaca has consistently been attributed to Puschnig
(1910) by subsequent authors (e.g. Ramme, 1941, 1951; Galvagni, 1954, 1986a,
1986b, 1987; Hölzel, 1955; Harz, 1973, 1975; Nadig 1987, 1989; Otte, 1995; Ingrisch
& Köhler, 1998; Heller er al., 1998; among others), which is wrong. Puschnig (1910:
27) introduced “Podisma alpina var. alpina subvar. carinthiaca” as the fourth name in
addition to a trinomen. Art. 45.5.1 of the Code (ICZN) says that such a name is infra-
subspecific and cannot be made available from its original publication by any sub-
sequent action. Following this article, the first author using the same word in a manner
that satisfies the provisions of the Code has thus established a new name with its own
authorship and date. This first author was Obenberger (1926a: 179) who used the name
in a trinomen as “Podisma alpinum var. carinthiacum [sic]. Though the name was
published as “var.” it has subspecific status (ICZN, Art. 45.6.4) and is deemed to have
been published in the form Podisma alpina carinthiaca Obenberger.

Obenberger (1926a, in Czech) published P. a. carinthiaca in a book on Ortho-
ptera and allied insects from Czechoslovakia and adjacent countries of which a shorter
edition (Obenberger, 1926b) appeared in French in the same year. We have no further
information as to the exact date of publication of these works which is why both are to
be considered as being published simultaneously (December 31, 1926). However, in
the French edition Obenberger referred to his Czech book as “...mon travail original
tchèque” (1926b: 117; compare also his remarks under “N. B.” on p. 233 where he
gave the exact quotation and paging of the Czech work). Therefore, we give prece-
dence to P. a. carinthiaca as published in this Czech edition (Obenberger, 19262). It
might be informative for the reader that the French edition was also published as a
separate reprint with its own paging (1-121). Several authors, for instance Mistshenko
(1952) or Otte (1995), quoted from this reprint.

1 Podisma is actually feminine.

ON MIRAMELLA IRENA AND KISELLA (CAELIFERA) 633

A: Kärnten: 82:109
Nockgebiet: Grundtal

1750 »
25.8.82 leg.NADIG

62:109 No.2 (4)
MIIROPH Kopf, Elytra
Penis v, Seite

eg 2

et. NADIG

Fics 1-2

Neotype d of Miramella carinthiaca: (1) habitus in lateral and in dorsal view. Scale 1 cm.
Photographs L. Schäublin. (2) labels, original size.

It is important to address the question which specimen(s) constitute the name-
bearing type of P. a. carinthiaca. Usually, the author who establishes a new name is
also responsible for the name-bearing type but here this is not evident. For instance,
Obenberger failed to specify any data with regard to the origin and composition of his
material. Apparently, he had not the intention to establish a new taxon, because he
(1926a: 179) quoted the name as “Podisma alpinum var. carinthiacum [sic] Puschnig”.
Yet, an unambiguous bibliographic reference to Puschnig (1910) is missing although
Obenberger (1926a: 24) listed Puschnig among those authors whose work he
considered important. On the other hand, we have found a striking resemblance in the
respective descriptions which are quoted verbatim below:

“Die kurzflügelige alpina-Form zeigt im Vergleich zu niederösterreichischen
Exemplaren noch kürzere, das dritte Hinterleibssegment nicht überragende
Elytrenschuppen, welche ähnlich wie bei Pod. Fieberi Scudd. und Schmidti Fieb. stark
seitlich gerückt sind, ihre ovoide und relativ breite Form jedoch beibehalten haben
(subvar. carinthiaca m.)” (Puschnig, 1910: 27, 28).

“Krovky kratké, kratSf neZ u typické formy [= alpina], avSak Siroké a ovälnt, sil-
né na strany posunuté” (Obenberger, 1926a: 179).

634 H. BAUR & A. CORAY

If we allow for certain discrepancies in the wording due to natural constraints
of language (German versus Czech) and context (diagnosis versus key) then both au-
thors describe essentially the same character. In short, they consider the elytra to be
shorter than in alpina, ovoid, broad, and strongly confined laterally. Evidently,
Obenberger did not mention any additional character. Couplet 9” of his key rather
looks like a mere translation and adaptation of the German text. A similar close match
of descriptions is also present in all other cases where Obenberger (1926a) attributed a
taxon to Puschnig, e.g. Chrysochraon brachypterus var. subcoeruleus (p. 141),
Podisma pedestre var. maius [= major in Puschnig, 1910] (p. 178), Caloptenus italicus
ab. bilineatus (p. 181), among others. Coincidence can hardly be the cause for such a
high degree of congruence. Rather it suggests that Obenberger used Puschnig’s
diagnoses as a template for the respective paragraphs of his key. Because we do not
know of any other publication containing the name let alone a description of P. a.
carinthiaca prior to Obenberger (1926a), Obenberger’s mentioning of Puschnig in
combination with a taxon is actually a reference to that particular work of Puschnig
(1910). Following Art. 72.4.4 of the Code, the specimens cited by Puschnig (1910)
from Carinthia are thus to be considered as the name-bearing type of P. a. carinthiaca
Obenberger. This may look bewildering at first glance but it is convenient, because the
type remains the same with regard to previous works where the authorship had
erroneously been attributed to Puschnig (see above).

According to Nadig (1989: 179) no type material of P. a. carinthiaca is pre-
served in the Museums of Klagenfurt and Vienna where Puschnig’s collection is
assumed to be deposited. Nadig’s statement is confirmed by the curators of the respec-
tive Museums, Drs P. Mildner and A. Kaltenbach (pers. comm.), and our own search
in the NHMW. Furthermore, the “neotypes” erected by Harz (1975: 290) and Nadig
(1989: 179) are invalid because their actions were in clear violation of the regulations
of the Code (ICZN, Art. 75.3). For instance, both authors selected a pair of “neotypes”
which they kept in their private collection (note: contrary to his claim, Nadig actually
did not designate any specimens of his collection as “neotypes”, but he labelled a series
of 23 d, 27 © from Grundtal-Grundalm as “TOPO-typus”). Therefore, the name-
bearing type of P. a. carinthiaca is definitely no more in existence.

This situation is rather unfortunate, in particular with regard to the complicated
taxonomic history of P. a. carinthiaca. Ramme (1941) was the first to use the name in
the current combination with Miramella as “Miramella alpina collina f. carinthiaca
Puschnig”. He (1941: 128) apparently misinterpreted the taxon, because the elytra
were said to be intermediate in length with regard to M. alpina alpina with short and
M. a. collina with long elytra. A respective specimen was figured later by him (Ramme,
1951, plate 4, figure 1b). Ramme’s interpretation was thus in clear contradiction to
Puschnig’s (1910) description and this probably caused much of the later confusion.
The name carinthiaca has subsequently been adopted for a species of Miramella by
Galvagni (1954) but thereby was confounded with M. irena (Fruhstorfer) (see also
Hölzel, 1955: 60). This, in turn, led to the misidentification of P. a. carinthiaca as the
type species of the subgenus Kisella Harz of Miramella (see below). Only the studies
by Galvagni (1986a) and Nadig (1989) have shed some light on these problems and
provide a more satisfying delimitation of the species. With regard to the apparent

ON MIRAMELLA IRENA AND KISELLA (CAELIFERA) 635

difficulties in separating these taxa, however, we think it is justified to designate a neo-
type for P. a. carinthiaca. This provides a firm basis for subsequent investigations and
furthermore allows selection of a type locality among several distant places in
Carinthia [Puschnig (1910: 27) mentioned “Grundlalm [sic]-Schiestlnock [sic]”,
“Saualpe-Weite Alpe”, “Metnitztal (Oberhof, Mödringgraben, Pachlergraben, ...)” as
the origin of his material]. We thus propose for a neotype a male from Grundtal-
Grundalm in CAN (see above). This selection ensures that the application of the name
will rest on Obenberger (1926a) as well as on Galvagni’s and Nadig’s major contri-
butions to the taxonomy of Miramella.

As mentioned above, the differences between M. carinthiaca and M. irena are
subtle. Although Nadig (1989) thoroughly studied both species, he failed to provide a
synthesis in form of a short diagnosis. Below, the most important diagnostic characters
are presented in a key which is based on a re-examination of material from many
collections (see Appendix):

- Elytra (Figs 3, 5) squamipterous, always widely separated medially.
Dark markings on pronotum (Figs 3, 5) usually less extensive, median
keel often only slightly darkened. Penis (Fig. 7) in lateral view with
dorsal valves slender and uniformly narrowing; ventral valves elongate,
at most as broad as dorsal valves basally ................. M. carinthiaca
- Elytra (Figs 4, 6) longer, squamipterous to subbrachypterous, usually
touching medially. Dark markings on pronotum (Figs 4, 6) rather more
extensive, median keel and some of the transverse sulci usually strongly
darkened. Penis (Fig. 8) in lateral view with dorsal valves less slender,
abruptly narrowing in apical third; ventral valves stouter, broader than
COrsalivalVesbasdllyi e e Re a. RS Re EEE M. irena

The two species occur in the south-eastern part of the Alps where they are
largely parapatric in distribution. Their ranges have been investigated by Galvagni
(1986a) and Nadig (1989) who found occasional hybrid populations between M.
carinthiaca and M. irena or M. alpina (Kollar). Yet, the status of these taxa is currently
not under debate.

Status. Miramella (Kisella) carinthiaca (Obenberger, 1926a) stat. rev.

Pezotettix alpinus collinus Brunner von Wattenwyl

Pezotettix alpina var. collina [sic®] Brunner von Wattenwyl in Künstler, 1864: 773. Lectotype
3, Slovakia: Pressburg [= Bratislava], here designated (NHMW [examined]).

Material. Lectotype 3 (directly pinned, remounted by H. Baur; genital complex ex-
posed; tip of left ventral valve (Fig. 9) lacking, however, the missing part was restored in the
drawing according to the intact right ventral valve) labelled “Coll. Br.v.W [print] Pressburg
Seyffert leg. [hand]; det. Br. v. W [print] Podisma alpina Koll. [hand]; 4345. [hand, label with
double border]”. The first two labels are not from Brunner’s hand and probably have been added
later. The number “4345” refers to Brunner’s collection index. The respective page shows an en-
try of the year “1864” on the top left. Lines 4334-4359 are headed by a note that reads “... [?,
illegible] Seyffert im August u. Sept. Pressburg. ...”. On line 4345 is finally stated “... collina”.

2 Pezotettix is actually masculine.

H. BAUR & A. CORAY

636

pra

TESO

na (from South Tyrol: Mendelkamm, Felixeralm). Pronotum and wings of d in lateral view: (5)

(6) M. irena (same locality as ?). Scale 1 mm. Drawings A. Coray.

M. carinthiaca (neotype),

ON MIRAMELLA IRENA AND KISELLA (CAELIFERA) 637

Paralectotypes, 1 6, 1 2 (directly pinned) labelled: “Coll. Br.v.W [print] Pressburg
Seyffert leg. [hand]; det. Br. v. W [print] Podisma alpina Koll. [hand]; Miramella alpina (KOL-
LAR) det. A. Galvagni,1961 [print; this label on 2 only]”. Both specimens stood beside lecto-
type under a bottom label “4345. Pressburg [hand, label with double border]” and are considered
as part of the type series.

The true extent of the type series is unknown. We have examined further specimens from
Vienna and surroundings, Mehadia, and some other localities which might belong to the type
series, but lack any positive evidence such as unambiguous determination labels by Brunner or
a respective entry in the above-mentioned collection index. Thus, they can not be considered as
part of the type series with certainty. This material is summarily listed in the Appendix under M.
alpina and M. irena.

Discussion. Brunner first mentioned a brachypterous form of “Pezotettix alpina
[sic]” from “Wienerwald” and “Krain” in his “Orthopterologische Studien” (1861:
222), but without giving a name. Brunner’s description of Pezotettix alpinus collinus
was then published in a paper by Künstler (1864) (see Mistshenko, 1952) on an out-
break of M. alpina and not first in the “Prodromus” (Brunner, 1882), as stated by many
authors (e.g. Harz, 1975; Galvagni, 1986a, Nadig, 1989; among others). Brunner (in
Künstler, 1864: 773) indicated a rather large distribution of his new taxon by noting
“Die letztere [= collinus] tritt zuerst in der Gegend von Wien auf, breitet sich südlich
in die Krain und östlich bis Mehadia aus, (südlich von Laibach [= Ljubljana] ist sie mir
unbekannt, ebenso wenig in Serbien und Siebenbürgen). Sie findet sich nur im
Hügelland (weder in der ungarischen Ebene, noch im Hochgebirge der Krain und
Serbien’s)”. In other words, collinus is said to occur from Vienna to Mehadia (near
Baile Herculane, Romania) in the East and Krain (i. e. the region North of Ljubljana,
Slovenia) in the South, where it is confined to hilly areas. Naturally, this large and
vaguely defined distribution does also include the specimens from Bratislava in
Brunner’s collection.

Clearly, the distribution of collinus as indicated by Brunner includes several
taxa. For instance, we have seen specimens of M. alpina as well as M. irena in
Brunner’s material from Mehadia (Appendix). Galvagni (1986a: 27) also reported both
species from this locality. Furthermore, the distribution of collinus may even comprise
part of the range of M. carinthiaca (see distribution map in Galvagni, 1986a: 35). As a
consequence, M. irena and M. carinthiaca could possibly be regarded as junior syno-
nyms of collinus. In fact, these three taxa have often been interpreted very differently
in the past, for instance by Mistshenko (1952) and Harz (1975). Therefore, a lectotype
of collinus is designated to ensure stability in the application of this name (ICZN, Art.
74.7.3), which thus becomes a junior synonym of M. alpina. With this interpretation,
we follow the seminal and now widely accepted work of Galvagni (1954, 1986a) and
Nadig (1989) who considered collinus as the brachypterous form of M. alpina. The
best character to separate this form from the very similar M. irena are the male
genitalia (see Nadig, 1989). The penis valves of the lectotype (Fig. 9) very nicely fit
Nadig’s figure (1989: 257, figure 37) of aM. alpina 3 from “Wiener Wald”.

Status. Junior synonym of Miramella (Kisella) alpina (Kollar, 1833) (Galvagni,
1986a: 26, 27).

Kisella Harz

Kisella Harz, 1973: 403 (subgenus of Miramella). Type species: Miramella irena (Fruhstorfer)
[= Podisma alpina irena Fruhstorfer, 1921], here designated.

638 H. BAUR & A. CORAY

Fics 7-9

Male genitalia of (7) Miramella carinthiaca (neotype), (8) M. irena (same specimen as in
Fig. 6) and (9) Pezotettix alpinus collinus (lectotype; the tip of the left ventral valve is actually
lacking, however, the missing part was restored in the drawing according to the intact right
ventral valve). Scale 1 mm. Drawings A. Coray.

ON MIRAMELLA IRENA AND KISELLA (CAELIFERA) 639

Discussion. Harz (1973) established the new subgenus Kisella of Miramella
Dovnar-Zapolskij with “K. c. carinthiaca Puschnig” as the type species by original
designation. As we argued above, he wrongly attributed the authorship of the type
species to Puschnig (1910) instead of Obenberger (1926a). According to Art. 67.7 of
the Code (ICZN) Harz is nevertheless to be considered as having validly fixed the type
species as Miramella carinthiaca (Obenberger). However, this species had been
misidentified by Harz, since he (1973: 403) quoted Hermagor, Carinthia (Austria) as
the origin of his material. Galvagni (1986a) and Nadig (1989) demonstrated that
specimens from this locality undoubtedly belong to M. irena (Fruhstorfer). Moreover,
Harz (1973) actually described the true M. carinthiaca as M. carinthiaca puschnigi,
thus establishing a junior synonym of the former (Galvagni, 1986a: 22; Nadig, 1989:
179). A re-examination of the respective specimens fully confirms these views (see
Appendix). Concerning the selection of a type species, the respective species are
obviously very closely related and probably would be equally eligible as type species
of Kisella. But because Harz connected the type species so clearly with some particular
specimens, we feel that the taxonomic species actually involved in the misiden-
tification, M. irena, would best serve stability and universality here. Probably for
similar reasons, Galvagni (1986b: 70) listed the same taxon as the type species of
Kisella. However, Galvagni’s action was invalid with regard to the Code then in force
(see Coray & Lehmann, 1998: 107) and no valid type fixation has hitherto been made.
In accordance with Art. 70.3 of the Code (ICZN) we now fix the taxonomic species
involved in the misidentification, M. irena (Fruhstorfer) [= Podisma alpina irena
Fruhstorfer, 1921], misidentified as M. carinthiaca (Obenberger) [= Podisma alpina
carinthiaca Obenberger, 1926a] in the original designation by Harz (1973, sub “K. c.
carinthiaca Puschnig”), as the type species of Kisella.

The taxonomic status of Kisella is not yet settled. Galvagni (1986b, 1987) has
elevated all previous subgenera to generic level but there is also a tendency to regard
Miramella as a single unit and to ignore subgenera such as Kisella (Heller et al., 1998;
Coray & Thorens, 2001). The split of Miramella into several taxa by Harz (1973) and
Galvagni (1986b) may indeed not withstand a rigorous cladistic analysis. Here, we
simply follow the last comprehensive study on Miramella by Nadig (1989) who treated
Kisella as a subgenus (see also Coray & Lehmann, 1998; Ingrisch & Köhler, 1998).

Status. Subgenus of Miramella Dovnar-Zapolskij, 1933 (Nadig, 1989: 106).

ACKNOWLEDGEMENTS

We are indebted to Otto Kraus (Hamburg, Germany) for his help concerning the
authorship of Podisma alpina carinthiaca. We thank Elsa Obrecht (NMBE), Christian
Kropf (NMBE) and an anonymous reviewer for critical reading of the manuscript and
many useful suggestions. We are also grateful to Lisa Schäublin (NMBE) for taking
photographs. For the loan of or information on specimens we thank J. Constant
(Institut royal des Sciences naturelles de Belgique, Bruxelles), Alfred P. Kaltenbach
and Ulrike Aspöck (NHMW), Charles Lienhard (MHNG), Paul Mildner (Landes-
museum Kärnten, Klagenfurt, Austria), Andreas Müller (ETHZ), Michael Ohl (ZMB),
and Klaus Schönitzer (ZSM).

640 H. BAUR & A. CORAY

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GALVAGNI, A. 1986b. Attuale struttura sistematica del genere Miramella Dovnar-Zapolskij, 1933,
e proposta per una sua scomposizione in piü generi (Insecta: Caelifera: Catantopidae:
Catantopinae). Arti/ dell’Accademia Roveretana degli Agiati, a. 235, serie VI 25(B):
67-84.

GALVAGNI, A. 1987. The genus Miramella Dovnar-Zapolskij, 1933, in the Balkan and the
Carpathian regions (Pp. 208-218). Jn: BACCETTI, B. M. (ed.). Evolutionary biology of
orthopteroid insects. Ellis Horwood, Chichester, 612 pp.

HARZ, K. 1973. Orthopterologische Beiträge XIII. Atalanta 4: 403-407.

HARZ, K. 1975. Die Orthopteren Europas/The Orthoptera of Europe. Vol. II. W. Junk, The Hague,
939 pp.

HELLER, K.-G., KORSUNOVSKAYA, O., RAGGE, D. R., VEDENINA, V., WILLEMSE, F., ZHANTIEV,
R. D. & FRANTSEVICH, L. 1998. Check-list of European Orthoptera. Articulata, Beiheft
7: 1-61.

HÔLZEL, E. 1955. Heuschrecken und Grillen Kärntens. Carinthia II, 19. Sonderheft: 1-112.

INGRISCH, S. & KÖHLER, G. 1998. Die Heuschrecken Mitteleuropas. Die neue Brehm Bücherei,
629. Westarp-Wissenschaften, Magdeburg, 460 pp.

KÜNSTLER, G. A. 1864. Ueber Heuschreckenfrass. Verhandlungen der kaiserlich-königlichen
zoologisch-botanischen Gesellschaft in Wien 14: 769-746 [recte 776].

NADIG, A. 1987. On the taxonomy and geonomy of the genus Miramella in the Alps, the Jura,
the Vosges, the Schwarzwald, and the Pfalzerwald (pp. 373-376). In: BACCETTI, B. M.
(ed.). Evolutionary biology of orthopteroid insects. Ellis Horwood, Chichester, 612 pp.

NADIG, A. 1989. Die in den Alpen, im Jura, in den Vogesen und im Schwarzwald lebenden Arten
und Unterarten von Miramella Dovnar-Zap. (Orthoptera, Catantopidae) auf Grund
populationsanalytischer Untersuchungen. Arti dell’Accademia Roveretana degli Agiati,
a. 238, serie VI 28(B): 101-262 + pls. 6-9 in fine.

OBENBERGER, J. 1926a. Rovnokridly hmyz (Orthoptera a Dermaptera) republiky Ceskosloven-
ské (se 4 tabulemi a 25 obrazy v textu) [In Czech]. Fauna et flora Cechoslovenica. I.
Nakladem Geské Akademie Véd a Umeni, Praha, VII + 234 + 4 pls.

ON MIRAMELLA IRENA AND KISELLA (CAELIFERA) 641

OBENBERGER, J. 1926b. Orthoptères et dermaptères de la République Tschécoslovaque. Bulletin
international de l’académie tcheque des sciences 27: 113-238 + 4 pls.

OTTE, D. 1995. Orthoptera Species File 4: Grasshoppers [Acridomorpha] C. Orthopterists’
Society and the Academy of Natural Sciences of Philadelphia, 518 pp.

PUSCHNIG, R. 1910. Beiträge zur Kenntnis der Orthopterenfauna von Kärnten. Verhandlungen
der kaiserlich-königlichen zoologisch-botanischen Gesellschaft in Wien 60: 1-60.

RAMME, W. 1941. Die Orthopterenfauna von Kärnten. Carinthia II, 131: 121-131.

RAMME, W. 1951. Zur Systematik, Faunistik und Biologie der Orthopteren von Südost-Europa
und Vorderasien. Mitteilungen aus dem Zoologischen Museum in Berlin (1950) 27:
1-431 + 39 pls.

APPENDIX

Material of Miramella species examined in this study. For abbreviations of depositories,
see Baur & Coray (2004, this volume).

Miramella alpina: INSTITUT royal des Sciences naturelles de Belgique, coll. de Selys
Longchamps: “B. W. Pezott. [sic] alpina Var. Vienne [hand]” 1 8,1 9; “Var. alata [hand]; B. W.
Pezott. [sic] alpina Var. Vienne [hand]” 1 ?. The 3 specimens are listed in Burr’s (1913: 21) cat-
alogue on the de Selys Longchamps collection under “Podisma alpinum Var. collinum [sic]”.

NHMW: AUSTRIA, coll. Brunner, leg. Schiesser: Kaltleitgeben [near Vienna], vii.1858,
4 8,3 2, 25.ix.1858, 2 d, 1 2, and a pair in copula; leg. Brunner: Mauer [near Vienna],
15.vii.1862, 1 9, Ober St. Veit [near Vienna] x.1863, 1 4,1 2.

NMBE: ROMANIA: Mehadia [near Baile Herculane], coll. Brunner 1 6, 1 9.

Miramella carinthiaca: CAN: AUSTRIA, Carinthia: Nockgebiet, Grundtal 4 4, 3 9;
Schiestelnock, Windeben 2 3; Innerkrems, Grünleitennock 2 2; Wöllanernock, Kaiserhügel 4
3; Haidnerhöhe, Hirnkopf 4 4,2 9; Saualpe, Klipitztörl, W side 1 d; Styria: Murau, Frauenalm
4 4,2 2; Saualpe, Schwarzkogel 2 8,3 9; Salzburg: Mautendorf, Speiereck 4 à.

CKH: AUSTRIA, Carinthia: Gurkentaleralpen, Haidnerhöhe, viii. 1958, leg. Hölzel,
coll. Ebner, paratypes 1 d,1 © of M. carinthiaca puschnigi Harz, 1973.

LANDESMUSEUM für Kärnten, Klagenfurt, Austria: AUSTRIA, Carinthia: Gurken-
taleralpen, Haidnerhöhe, leg. Hölzel 11 4,9 9.

NHMW: AUSTRIA, Carinthia: Gurkentaleralpen, Haidnerhöhe, viii. 1958, leg. Hölzel,
coll. Ebner, holotype d, paratypes 1 d,1 © of M. carinthiaca puschnigi Harz, 1973.

Miramella irena: CAN: AUSTRIA, Carinthia: Plöckenpass 1 d; ITALY, South Tyrol:
Mendelkamm, Felixeralm 4 d, 3 2; Gampenjoch, N-side 3 4; Ultental, over Gamperalm 2 d;
Province Trento: V. di Non, V. della Forma 1 ¢, 2 9; Madonna di Campiglio: Meledrio 3 d,
3 2; V. Nambrone: Adamello 1 4, 1 2; Adamello, V. di Fumo 2 & ; Passo Croce Domini: South
2 4,2 2; Rifugio Bazzena 1 4, 1 9; Dolomiten: Passo Giau, S-side 2 4, 2 9; Province Brescia:
€. Tombea 3 8,3 ©.

CKH: AUSTRIA, Carinthia: Karnischer Hauptkamm, Hermagor, 2.8.1953, 1 3,1 9 la-
belled by Harz as “neotypes” of M. carinthiaca (see text); same, 1 6; Oberdrauburg, 26.vi-
11.1962, leg. R. Hess 1 d, 1 9; ITALY, South Tyrol: Dolomiten, St. Ulrich, Anf. ix.1962, leg.
Engel 1 4, 1 9; BOSNIA-HERZEGOVINA: Kupres, 2.viii.1966, leg. Eitschberger 1 3;
Sarajevo, Trebevic, 7.viii.1966, leg. Eitschberger 4 d, 4 2. All specimens identified by Harz as
M. carinthiaca.

ETHZ: AUSTRIA, Carinthia: Obir, Eisenkappelhiitte-Hoffmannsalpe, 1400-1700 m,
29.vii.1987, leg. Ingrisch 2 d; ITALY, Trentino: Dosso Sabion, Pinzolo, 1700 m, 31.vii.1947,
leg. Galvagni 1 à ; 1400 m, 30.vii.1947, leg. Galvagni 3 à ; Passo, Mendola, 1400 m, 7.ix.1946,
leg. Galvagni 1 d; Concei-M. Cadria, Vies, 1800-1850 m, 6.ix.1958, leg. Galvagni 2 6, 1 9;
M. Tremalzo, 1700-1900 m, 28.viii.1959, leg. Galvagni 2 9; Paganella: Becco del Corno,
2000 m, 7.ix.1946, leg. Galvagni 1 d.

NHMW: ROMANIA: Mehadia [near Baile Herculane], leg. Brunner, 23.7.1862, 1 ©,
20.7.1864, 3 2; SLOVENIA: Steiner Alpen [North of Ljubljana], 1859, leg. Hoffmann 1 ©.

ZMB, coll. W. Ramme: AUSTRIA, Carinthia: Plöckenpass, 1200-1300 m, viii.1929,
1 3; Mauthen, viii.1929, 1 ¢; ITALY, South Tyrol: St. Vigil, 7.vii.-8.viii.1913, 2 6,3 9;

642 H. BAUR & A. CORAY

vii.1913, 2 8; St. Vigil, Enneberg, vii.-viii.1913, 1 6, 2 ©; Klausen, 17.vi.-3.viii.1910, 1 g;
Krain: Planina, 26.viii.-5.ix.1912, 2 6, 2 2; Seiseralpe: Schlern, 26.viii.1921, 1 6; Monte
Spinale, 14.viii.1921, 1 d ; Val di Genova, 1500 m, 9.viii.1921, 2 8,2 9; KROATIA, Plitvitka:
Jesera, 400-500 m, viii.1929, 2 2.

REVUE SUISSE DE ZOOLOGIE 111 (3): 643-655; septembre 2004

A new species of Platythyrea from Dominican amber and
description of a new extant species from Honduras
(Hymenoptera: Formicidae)

Maria L. DE ANDRADE

Institut für Natur-, Landschafts- und Umweltschutz, Universität Basel,
Biogeographie, Neuhausstrasse 31, CH-4057 Basel, Switzerland.
E-mail: maria.deandrade@unibas.ch

A new species of Platythyrea from Dominican amber and description of
a new extant species from Honduras (Hymenoptera: Formicidae). -
Two new species of Platythyrea are described in this paper. P. pumilio sp. n.
is described from a single worker in Dominican amber. P. lenca sp. n. is
based on three workers from the Recent Honduran fauna. Among the eight
Recent and the three fossil Neotropical Platythyrea species known to date,
the fossil pumilio shares with the Dominican amber scalprum Lattke the
small size, the incrassated fore femora and the fore basitarsi with a seta
opposite to the strigil, and with the extant Brazilian exigua Kempf the
elongate head, the flat eyes and the minutely denticulate mandibles. The
Honduran /enca sp. n. is very similar to the Costa Rican prizo Kugler. P.
lenca and prizo share the elongate body, the narrow frontal lobes and the
serrate mandibles; this latter character is also present in the fossil P. dentata
Lattke.

Keywords: Hymenoptera - Formicidae - Platythyrea - Dominican amber -
fossil - extinction - new species - Honduras.

INTRODUCTION

The genus Platythyrea is represented by eight Recent and three fossil species in
the New World (see the literature review by Bolton, 1995 and Lattke, 2003). All New
World Platythyrea species are very similar to each other (Kempf, 1964; Brown, 1975).
Nonetheless, Kempf (1964) described Platythyrea exigua from Brazil and stressed it
distinctness. P. exigua is characterized mainly by its small size, elongate head and
armed petiole. Brown (1975) described Plarythyrea zodion from Ecuador stating that
this species, together with exigua, are the sole two distinct New World species. P.
zodion Brown is easily differentiated from all the other species by its small size,
unarmed petiole and head longer than broad. Kugler (1977) described another peculiar
Costa Rican Platythyrea: P. prizo which differs from all other American species mainly
by its serrate mandibles, narrower frontal lobes and slender appendages. Kugler (1977)
added that prizo appears to belong to the Old World clypeata group of Brown (1975).
His attribution to the clypeata group is due to the fact that P. prizo shares with the

Manuscript accepted 22.01.2004

644 M. L. DE ANDRADE

members of this group similar frontal lobe structure, mandible shape, dentition, trunk
shape and palpal formula. Wilson (1985a) and Baroni Urbani (1995) record the
presence of Platythyrea in Dominican amber without giving further details. Lattke
(2003) described three new species of Platythyrea, dentata, procera and scalprum
from Dominican amber. According to Lattke (1. c.), the dentate mandibles of dentata
are also found in prizo Kugler, but this character should be plesiomorphic since denta-
ta is smaller, possesses hind coxae dorsally dentate and the anterior clypeal margin
straight instead of convex. Lattke (1. c.) compared P. procera and P. scalprum with P.
exigua Kempf and P. zodion Brown.

MATERIAL AND METHODS

The following specimen of Platythyrea was examined in one amber sample
from the Dominican Republic, deposited at the Museum of Comparative Zoology,
Harvard University, Massachusetts, USA.

The MCZC Platythyrea specimen has been numbered by myself as MCZC-37.
The numbers 1 to 24 were already used for MCZC amber material in de Andrade &
Baroni Urbani, 1999 and the numbers 25-35 in Baroni Urbani & de Andrade, 2003.
The number 36 was used for a Cyphomymrmex gyne (de Andrade, 2003).

MCZC-37 (Fig. 1). A small dark orange sample 1.1 x 0.8 cm containing a
worker of Platythyrea and impurities. The preservation condition of the ant is good.

The Recent Platythyrea examined for this study are deposited in the following
collections, given here with the relative coden as it will be used in the following text:
MCZC. Museum of Comparative Zoology, Harvard University, Massachusetts, USA.
Courtesy Stefan P. Cover. NHMB. Naturhistorisches Museum of Basel, Switzerland.
Courtesy Dr Michel Brancucci and Dr Daniel Burckhardt.

Measurements and indexes used in the text are defined here as:

HL Head length: the maximum measurable distance between the medial margin of vertex
and the antero-medial margin of clypeus with the head in full frontal view.

HW Head width: maximum head width behind the eyes with the head in full frontal view.

EE Eye length: maximum length of the eye.

SL Scape Length: length of scape shaft, excluding the basal condyle.

HBaL Maximum length of hind basitarsus measured on its external face.

HTiL Maximum length of hind tibia measured on its external face.

HFeL Maximum length of hind femur measured on its external face.

FFeL Maximum length of fore femur measured on its external face.

FFeW Maximum width of fore femur measured on its external face.

WL Weber’s Length: diagonal length of mesosoma from the anterior pronotal border

(excluding neck) to the distal edge of the propodeal lamellae.

INL, Total Length: combined head length in full face view (closed mandibles included),

Weber’s length of mesosoma, petiole length (in profile) and length of gaster (in profile).
Cl Cephalic index: (HW/HL) x 100

SI Scape index: (SL/HL) x 100
FFel Fore Femora index: (FFeW/FFeL) (100)

DESCRIPTIONS
Platythyrea pumilio sp. n. Fig. 1

Material. Holotype worker from Dominican amber, labelled: MCZC-37, in the MCZC
(see section Material and Methods).

NEW CENTRAL AMERICAN PLATYTHYREA 645

FIG. 1

Platythyrea pumilio sp. n. Holotype worker from Dominican amber: head in dorsal view (top),
and profile of the whole specimen (bottom). Distance between two scale bars 0.1 mm.

646 M. L. DE ANDRADE

Etymology. From the Latin substantive pumilio = dwarf, pygmy, referring to the
small size of this species. It is used here as a noun in apposition.

Diagnosis. A Platythyrea species, known from the worker only, sharing charac-
ters with the workers of P. scalprum Lattke and P. exigua Kempf, but differing from
both by the declivous propodeal sides lamellaceous and by the much less pronounced
postero-median petiolar convexity, from scalprum only by the head narrower and by
the petiole shorter and higher, and from exigua only by the fore femora broader, by the
lack of metacoxal tooth and by its smaller size.

Description. Worker. Head elongate, slightly less than 2/3 longer than broad,
with subparallel sides. Vertexal margin medially concave. Vertexal angles slightly pro-
truding posteriorly and subround. Frontal lobes far from each other and gently
rounded. Clypeus declivous anteriorly, with truncate anterior border and weakly
differentiated from the frontal lobes posteriorly. Eyes large, about 1/4 of the head
length and placed dorso-laterally on the anterior half of the head. Mandibles sub-
triangular. Masticatory margin of the mandibles with 7-8 minute, irregular denticles
followed by an apical one. Mandibular sulcus superficially impressed. Palpal formula
3,2, the Mandibular palps not surpassing the postero-medial rim of buccal cavity.
Scapes much shorter than the vertexal margin. First funicular joint slightly longer than
broad. Funicular joints 2-10 slightly broader than long. Last funicular joint about as
long as the sum of joints 9-10.

Mesosoma in profile elongate. Pronotum in dorsal view with barely distin-
guishable humeri. Pronotal suture impressed. Posterior third of the propodeal dorsum
gently declivous posteriorly. Sides between basal and declivous propodeal faces sub-
round. Sides of the declivous face with a lamella reaching the propodeal lobes.

Petiole longer than broad. Anterior face of the petiole in dorsal view weakly
convex medially and weakly angulated latero-ventrally. Posterior face of the petiole in
dorsal view poorly convex medially and unarmed laterally. Postpetiole slightly shorter
than first gastric segment.

Fore femora strongly incrassate. Fore basitarsi with a seta opposite to the strigil.
Hind tibiae slightly shorter than hind femora. Hind basitarsi slightly shorter than 1/3 of
the tibiae. Mid and hind tibiae with paired, pectinate spurs. Tarsal claws with preapical
teeth.

Sculpture. Integument opaque. Head, mesosoma, petiole, postpetiole and gaster
densely and finely reticulate-punctate and with sparse, minute, superficial foveae. Legs
minutely punctate, the punctures more superficial than in the other body parts.

Body densely covered by very short, thin hairs. Apex of the gaster with long
hairs.

Colour. Light brown.

Measurements in mm and indices. TL 3.82; HL 0.75; HW 0.46; EL 0.18; SL
0.44; WL 1.24; PeL 0.45; HFeL 0.59; HTiL 0.57; HBaL 0.41; FFeL (left) 0.54; FFeW
(left) 0.25; FFeL (right) 0.59; FFeW (right) 0.25; CI 61.3; SI 58.6; FFel (left) 46.3;
FFel (right) 42.5.

FIG. 2

Platythyrea lenca sp. n. Paratype worker from Honduras (4 km SW Mina EI Mochito, Depto.
Santa Barbara): head in dorsal view (top), and profile of the petiole (bottom).

NEW CENTRAL AMERICAN PLATYTHYREA 647

Uni Basel

648 M. L. DE ANDRADE

Platythyrea lenca sp. n. Figs 2 & 3

Material examined. Holotype worker from Honduras, labelled: 4 km SW Mina El
Mochito, 1040 m, rotten wood Forest ravine, Depto. Santa Barbara, HONDURAS 14 March
1979, WL Brown; 2 paratype workers same data and collection as the holotype, all in the MCZC.

Etymology. This species is named after the Lenca, an Indian tribe from
Honduras. The name is used here as a noun in apposition.

Diagnosis. A Platythyrea species, known from the worker only, similar to P.
prizo, but differing from it by the erect hairs on the petiole, postpetiole and gaster
longer and denser, by the legs longer, by the eyes smaller and by the scapes longer and
narrower.

Description. Worker. Head slightly elongate and less than 1/3 longer than broad,
with weakly convex sides. Vertexal margin weakly concave medially. Vertexal angles
round. Frontal lobes close to each other and rounded. Clypeus swollen medially.
Anterior border of the clypeus bearing a convex semitransparent lamella. Eyes small,
about 1/7 of the head length and placed dorso-laterally close to the midline of the head.
Mandibles subtriangular. Masticatory margin of the mandibles with 9 teeth followed by
an apical one. Mandibular sulcus superficially marked. Palpal formula 4,4, palpi not
surpassing the posteromedial rim of buccal cavity. Antennae long and slender. Scapes
surpassing the vertexal margin by about 0.32 mm when directed posteriorly. First
funicular joint more than half longer than broad and slightly longer than the following
joints 3-10. Second funicular joint about 1/4 longer than the first joint. Funicular joints
3-10 longer than broad. Last funicular joint slightly longer than the first joint.

Mesosoma elongate in profile. Pronotum in dorsal view with barely distinguish-
able humeri. Pronotal suture impressed. Propodeal dorsum gently declivous posterior-
ly. Area between basal and declivous propodeal faces gently concave and each side with
a small obtuse tooth. Declivous propodeal face marginate. Propodeal lobes rounded.

Petiole about 1/3 longer than broad. Anterior face of the petiole in dorsal view
medially weakly concave and latero-ventrally superficially angulate. Posterior face of
the petiole in dorsal view tumuliform medially and unarmed laterally. Postpetiole in
dorsal view broader than long and shorter than the first gastric segment.

Fore femora weakly incrassate. Hind tibiae about 1/8 shorter than hind femora.
Hind basitarsi about 1/7 shorter than the hind tibiae. Mid and hind tibiae with paired,
pectinate spurs. Tarsal claws with preapical teeth.

Sculpture. Integument opaque. Head, mesosoma, petiole, postpetiole and gaster
densely and finely punctate and with small, superficial foveae, the foveae more
impressed on the petiole, fainter and sparser on the gaster. Legs minutely punctate.

Body densely covered by very short, thin hairs. Petiole, postpetiole and gaster
with erect, sparse hairs. Apex of the gaster with long hairs.

Colour. Ferruginous-brown with slightly lighter antennae and legs.

Measurements in mm and indices (3 specimens examined). TL 7.72-8.38; HL
1.68-1.74; HW 1.16-1.22; EL 0.25; SL 1.52-1.60; WL 2.60-2.64; PeL 0.82-0.86; PeW
0.56-0.60; HFeL 1.78-1.84; HTiL 1.54-1.64; HBaL 1.32-1.42; FFeL 1.56-1.64; FFeW
037-.0397€1:69.0-70.1: ST 903-929; FRel 237.

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Platythyrea prizo Kugler Figs 4,5 & 6

Material examined. COSTA RICA: Heredia, OTS, La Selva field Station, Puerto Viejo
de Sarapiqui, Rio Puerto Viejo, 5.1II.1973, 10°26° N-83°59° W, 3 workers (paratypes numbers
73-292, 73-295, 73-299), J. Wagner & J. Kethley [MCZC].

Complement to biometry:

Measurements in mm and indices (3 specimens examined). TL 7.65-8.22; HL
1.64-1.68; HW 1.18-1.22; EL 0.30-0.32; SL 1.42-1.46; WL 2.56-2.72; PeL 0.83-0.86;
PeW 0.58-0.60; HFeL 1.68-1.72; HTiL 1.48-1.56; HBaL 1.24-1.36; FFeL 1.52-1.58;
FFeW 0.39-0.41; CI 71.1-72.6; SI 86.6-86.9; FFel 25.6-26.3.

IDENTIFICATION KEY FOR THE NEW WORLD PLATYTHYREA

The following identification key to the workers of Platythyrea from the New
World, including all known extant and fossil species, is based on Brown’s (1975) key
for this genus modified to include the species described afterwards by Kugler (1977),
Lattke (2003) and myself (present paper). P. dentata Lattke, known on the gyne only,
is not included in the key, although it may probably be separated from the known
worker-based species with a reasonable degree of confidence (see note below).
Longino (2003), in a web document, states that the mandibular sulcus, used by Brown
(1975) to differentiate P. sinuata from P. pilosula, shows a high degree of variation in
most Costa Rican sinuata and, as a consequence of this, treats sinuata as a junior
synonym of pilosula. This proposal is accepted in the key below, even if the source can
not be considered a publication for nomenclatorial purposes.

1 HW (eyesancluded) > 0:90 mm "LE SR 2
- HW (eyes included) <0:90:mm RO oes SENO NE 7
2 Mandibles’withteeth.. "5:20 00 22004 M OO = see eee 3)
- Mandibles' without teeth ."::::::.......:.42.. 0.0 SORBONA 4
3 Erect hairs on petiole, postpetiole and gaster sparser and shorter. EL

> 0.30 mame eis sas ee ee TE TE prizo
- Erect hairs on petiole, postpetiole and gaster denser and longer. EL

= 0.25 MM... na ern ne PE lenca
4 HW (eyes included) > 1.4 mm. EL > 0.45 mm. Fore basitarsi with 3

setae opposite to the/strigil- 3 Re strenua
- HW (eyes included) < 1.4 mm. EL < 0.45 mm. Fore basitarsi with 1 or

2 setae opposite to the strigil .. .. .......-. 2... 2... SNC 5)
5) Profemur strongly swollen (W/L > 0:38) erre 2 eee angusta
- Profemur less tick (W/L. < 0.38). : :... 222... 2.2.02 2.202 NEE 6
6 Petiolar node > 0.85 mm long. Petiole about 1/4 longer than broad. Fore

basitarsi with 2 setae opposite to the strigil ............ pilosula (= sinuata)

- Petiolar node < 0.85 mm long. Petiole about as long as broad or slightly
longer than broad. Fore basitarsi with 1 seta opposite to the strigil . . . punctata

FIG. 4
Platythyrea prizo Kugler. Paratype worker from Costa Rica (Heredia, La Selva field Station,
Puerto Viejo de Sarapiqui, Rio Puerto Viejo): head in dorsal view (top), and profile of the petiole
(bottom).

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FIG. 6

Platythyrea prizo Kugler. Paratype worker from Costa Rica (Heredia, La Selva field Station,
Puerto Viejo de Sarapiqui, Rio Puerto Viejo): left eye showing interommatidial pilosity.

7 Metacoxal dorsum with a tooth. Posterior margin of petiolar node as

seen from above with lateral corners produced caudad as short, blunt

tcethtorangles4Clypeusidorsallyiswollen att prece exigua
- Metacoxal dorsum unarmed. Posterior margin of petiolar node with at

most weakly armed lateral corners. Clypeus dorsally flat ................ 8
8 Scapes reaching the vertexal margin. TL = 5.9 mm. SI>70......... procera
- Scapes much shorter, not reaching the vertexal border. TL < 5.0 mm. SI < 60 . 9
9 Head elongate, about 1/3 longer than broad. CI<63............... pumilio
= Headishortertabout l//Mlonserithantbroad Cli> 8008 ara rer er oe 10
10 Petiolar node elongate (W/L 0.67). Profemur swollen (W/L 0.47). TL

MOINE ER. ae beato sie. ee che scalprum
- Petiolar node shorter (W/L 0.86). Profemur less thick (W/L 0.41). TL

> Sa Sai? Nele at aan zodion

Note. The original description of P. dentata Lattke, known on a sole gyne from Dominican
amber, gives HW 0.87 mm (eyes excluded). According to Lattke’s drawing the HW of this
specimen (eyes included, to render it comparable with the figures given by Brown (1975) should
be about 0.93 mm. By using the key to the workers above, the gyne of P. dentata falls to couplet
2 (mandibles with teeth) where it can be easily differentiated from both other two species, prizo
and /enca, by the hind coxae dorsally dentate and by the anterior clypeal margin straight instead
of convex.

654 M. L. DE ANDRADE

DISCUSSION

P. pumilio, described in this paper, and scalprum Lattke are the smallest Plary-
thyrea known to date from the New World. P. pumilio shares with the Dominican fossil
P. scalprum Lattke the fore basitarsi with a seta opposite the strigil and the swollen fore
femora, but it differs from it mainly by the petiole shorter and higher and by the head
much narrower. P. pumilio shares with the Brazilian P. exigua Kempf the head elongate
and narrow with flat eyes on the anterior half of the head and by the masticatory mar-
gin of the mandibles minutely denticulate. P. pumilio differs from the Brazilian exigua
mainly by its lack of metacoxal teeth, by its fore femora broader, and by its posterior
petiolar face unarmed. P. pumilio shares with the Ecuadorian zodion Brown a similar
petiolar shape, but it differs from it by the head much longer and by the size smaller.
P. pumilio shares with the other Dominican fossil, procera Lattke the propodeal
declivity with a lamella and the lack of metacoxal teeth. P. pumilio differs from procera
mainly by its smaller size.

P. lenca is very similar to prizo but the characters listed under the diagnosis of
lenca allow to distinguish both species. Since the two species are allopatric, one can
not exclude that they may represent geographic variants of the same species. Lenca and
prizo, moreover, are the sole known Recent New World Platythyrea with narrow
frontal carinae and truly denticulate mandibles. The biometric and pilosity differences
between the two, however, supported by our current understanding of Platythyrea vari-
ability, render improbable the geographic variation hypothesis. Lattke (2003) described
from Dominican amber P. dentata, another species with denticulate mandibles. P. den-
tata Lattke differs from prizo Kugler and /enca mainly by its smaller size, by the hind
coxae dorsally dentate and by the anterior clypeal margin straight instead of convex.
Lattke (2003) considered the sharing of the denticulate mandibles of his fossil species
with the Recent prizo as a plesiomorphic character since there are significant
differences between the fossil and the Recent species.

With the description of P. pumilio the number of fossil Plarythyrea from
Dominican amber increases to 4. The sole two known Recent species on the Island of
Hispaniola are P. strenua Wheeler & Mann and P. punctata Smith. The presence of four
fossil and two extant Platythyrea species on the island of Hispaniola is a further
confirmation, if necessary, of the major role played by local extinction as opposed to
invasion from other sources in shaping the contemporary. Hispaniolan fauna.
Interpretation of this phenomenon is a hot spot in biogeographic research. If, on one
hand, all ant species recorded from Dominican amber are extinct, every published
record of Dominican amber ants can be considered as proof in favour of extinction. On
the other hand, a famous, widely cited, paper by Wilson (1985b) emphasizes the role
of invasion and migration over the one of extinction in shaping the contemporary
Hispaniolan fauna. The apparent discrepancy between the known composition of the
amber fauna and Wilson’s conclusion is easily explained by the fact that Wilson’s
(1. c.) analysis is conducted at genus and not at species level. Baroni Urbani (1995)
showed that local extinction plays the dominant role also at generic level and that local
extinction and not invasion of new taxa is, hence, the most important factor that shaped
the present Caribbean fauna. This argument, as far as I know, was supported again only
by de Andrade & Baroni Urbani (1999) and the remarks above are added to reply an

NEW CENTRAL AMERICAN PLATYTHYREA 655

anonymous referee who suggested that there are other cases of references to extinction
in the Dominican amber fauna.

ACKNOWLEDGEMENTS

I would like to express my warmest thanks to Stefan P. Cover for lending this
and many other interesting amber samples, to Dr M. Brancucci and Dr D. Burckhardt
for the allowing my free access to the collection of the Natural History Museum of
Basel, to Daniel Mathys and Marcel Düggelin of the Lqaboratory of Scanning Electron
Microscopy of the Basel University for helping competently with all SEM photo-
graphs, and to Prof. Dr Cesare Baroni Urbani for critically reading this manuscript.

REFERENCES

ANDRADE, M. L. (DE) 2003. First descriptions of two new amber species of Cyphomyrmex from
Mexico and the Dominican Republic. Beiträge zur Entomologie 53: 131-139.

ANDRADE, M. L. (DE) & BARONI URBANI, C. 1999. Diversity and adaptation on the ant genus
Cephalotes, past and present. Stuttgarter Beiträge zur Naturkunde B 271: 1-889.

BARONI URBANI, C. 1995. Invasion and extinction in the West Indian ant fauna revised: the
example of Pheidole (Amber Collection Stuttgart: Hymenoptera, Formicidae. VIII:
Myrmicinae, partim). Stuttgarter Beiträge zur Naturkunde B 222: 1-29.

BARONI URBANI, C. & ANDRADE, M. L. (DE) 2003. The ant genus Proceratium in the extant and
fossil record. Monografie No 36. Museo Regionale di Scienze Naturali-Torino, 492 pp.

BOLTON, B. 1995. A new general catalogue of ants of the world. Harvard University Press,
Cambridge (Mass.) & London, 504 pp.

BROWN, W. W. 1975. Contributions toward a reclassification of the Formicidae. V. Ponerinae,
tribes Platythyreini, Cerapachyini, Cylindromyrmecini, Acanthostichini and Aenicto-
gitini. Search Agriculture 5: 1-115.

KEMPF, W. W. 1964. Uma nova Plarythyrea do Brasil (Hym., Formicidae). Revista Brasileira de
Entomologia 11: 141-144.

KUGLER, C. 1977. A new species of Platythyrea (Hymenoptera, Formicidae) from Costa Rica.
Psyche (Cambridge) 83: 216-221.

LATTKE, J. 2003. The genus Platythyrea Roger, 1863 in Dominican Amber (Hymenoptera:
Formicidae: Ponerinae). Entomotropica 18: 107-111.

LONGINO, J. 2003. Platythyrea pilosula (F. Smith 1858). 1 p.
http://www.evergreen.edu/ants/genera/platythyrea/species/pilosula/pilosula.html.
World Wide Web document.

WILSON, E. O. 1985a. Ants of the Dominican amber (Hymenoptera: Formicidae). 1. Two new
myrmicine genera and an aberrant Pheidole. Psyche (Cambridge) 92: 1-9.

WILSON, E. O. 1985b. Invasion and extinction in the west Indian ant fauna: evidence from the
Dominican amber. Science 229: 265-267.

a i Nur #

Da us

REVUE SUISSE DE ZOOLOGIE 111 (3): 657-669; septembre 2004

First reports of the genus Nigrobaetis Novikova & Kluge
(Ephemeroptera: Baetidae) from Madagascar and La Réunion
with observations on Afrotropical biogeography

Jean-Luc GATTOLLIAT
Museum of Zoology, P.O. Box 448, CH-1000 Lausanne 17, Switzerland.
E-mail: Jean-Luc.Gattolliat@serac.vd.ch

First reports of the genus Nigrobaetis Novikova & Kluge (Ephemero-
ptera: Baetidae) from Madagascar and La Réunion with observations
on Afrotropical biogeography. - Nigrobaetis colonus sp. n. and Nigro-
baetis cryptus sp. n. are described from La Réunion and from Madagascar,
respectively. This is the first report of the genus from either island and is the
first description of a mayfly from La Réunion. These amazing discoveries
are of biological importance for the understanding of colonisation processes
of the islands in the Western part of the Indian Ocean. The first record of
mayflies from La Réunion, a young (<3My) volcanic island, proves the
potential of mayflies to disperse over long distances. Recently, Nigrobaetis
bethunae Lugo-Ortiz & de Moor, 2000 was described from southern Africa
(from the border between Namibia and Angola); new reports from eastern
South Africa are provided, which greatly extend the distribution range of
this species.

Keywords: Baetidae - Nigrobaetis - new species - systematics - biogeo-
graphy - Madagascar - La Réunion - southern Africa.

INTRODUCTION

Nigrobaetis Novikova & Kluge originally was considered to be a subgenus of
Baetis Leach, and included most of the species of the Baetis niger group (Miiller-
Liebenau, 1969; Novikova & Kluge, 1987). It was thereafter raised to generic level and
redefined (Waltz er al., 1994). Despite the apparent similarities between the different
species encompassed in this genus, few relevant features were used to characterize the
genus. Waltz et al. (1994) proposed the following larval characters: body cylindrical,
prostheca normal or reduced, glossae with few to many dorsal setae, femoral villopore
absent, claws without preapical setae. Lugo-Ortiz & de Moor (2000) considered the
presence of a small interantennal carina as a generic character, despite the fact that
Waltz et al. (1984) did not refer to it and that this character is not mentioned in the
original descriptions (Soldän, 1977; Miiller-Liebenau, 1984; Kang er al., 1994). Other

Manuscript accepted 15.01.2004

658 J.-L. GATTOLLIAT

characters that seem constant in the different species, such as the two kinds of peculiar
setae of the distal margin of the labrum, the truncated third segment of the labial palp,
the few stout setae between prostheca and mola on both mandibles and the humped
posture of the prothorax, were not included in the generic diagnosis. The presence or
absence of hindwing pads, as well as the number of gills, are used for specific
diagnoses (Lugo-Ortiz & de Moor, 2000). There are no useful characters for separating
Nigrobaetis from other related genera at the imaginal stage (Waltz et al., 1994).

The genus Nigrobaetis comprises 15 species, mainly from the Palearctic
(6 species) and Oriental realms (6 species) [terrestrial zoogeographical regions as
defined by Zwick (2003)]. The recent description of a species from the Afrotropical
realm greatly extended the known range of the genus (Lugo-Ortiz & de Moor, 2000).
The simultaneous discovery of the genus on Madagascar and La Réunion is even more
surprising. A recent survey of the Malagasy freshwater macroinvertebrates included
sampling of more than 650 localities all over the island (Elouard & Gibon, 2001).
Based on this material, a study of the systematics of the Malagasy Baetidae increased
the known Malagasy Baetidae fauna from eight species belonging in three genera to 54
species in 22 genera (Gattolliat & Sartori, 2003). Despite this intensive research, the
genus Nigrobaetis was not known to occur on Madagascar. It was only during more
recent research on the impact of deforestation on macroinvertebrate community
composition that larvae of Nigrobaetis were collected in Madagascar.

Ephemeroptera were first found on La Réunion when Starmühlner’s expedition
collected larvae from two streams (Starmühlner, 1979). The occurrence of insects with
low powers of dispersal on an isolated volcanic island of recent origin remains a bio-
geographical riddle. The Baetidae are the only mayfly family that has been found on
this island. The identification of the genus and of its phylogenetic affinity is important
for the understanding of the mechanism of dispersion and colonisation of the different
islands of the western part of the Indian Ocean. McCafferty & Mauremootoo (2001)
tentatively associated the larvae with the genera Afrobaetodes Demoulin or Acanthiops
Waltz & McCafferty. The study of the original material collected by Starmühlner’s
expedition, as well as other specimens collected by the ORE (Organisme Réunionais
des Eaux) and Christian Guillermet, now allow the assignment of all mayflies living in
running waters from La Réunion to the genus Nigrobaetis.

Larvae of Nigrobaetis bethunae were collected from the Limpopo and Incomati
basins in and around the Kruger National Park, South Africa. This constitutes the first
report of the species after its description. It greatly enlarges the previously known
distribution range, which was restricted to the Cunene River, ca. 2500 km distant on
the border between Angola and Namibia. The investigation of this new material adds
further information to the original description, especially concerning the larval
colouration.

As the examined material from Madagascar and La Réunion differs from all the
previously described species, two new species are described here. The holotypes and
part of the paratypes are housed in the Museum of Zoology, Lausanne, Switzerland.
Other paratypes are deposited in the Museum National d’Histoire Naturelle, Paris
(France) and in the Natural History Museum of Vienna (Austria).

THE GENUS NIGROBAETIS IN THE AFROTROPICS 659

DESCRIPTIONS

Nigrobaetis colonus sp. n. Figs 1-11

MATERIAL EXAMINED

Holotype: 1 larva. La Réunion; Tributary of Marsouin River, Ravine Mathurine (road to
Takaka) (F/Ré/9b), 8.04.1974. Leg. F. Starmühlner.

Paratypes: 2 larvae same data as for holotype. 2 larvae, La Réunion; tributary of
Marsouin River, Ravine Mathurine (road to Takaka) (F/Ré/9c) 8.04.1974. Leg. F. Starmühlner.
1 larva, La Réunion, Galets River, Amont de la Prise, 7.02.1995. Leg. ORE (Organisme
Reunionais des Eaux). 2 larvae, La Réunion, Marsouin River, Bébour, 10.1996. Leg. ORE.

LARVA

Maximum length: body 5.2 mm; cerci 2.8 mm; median caudal filament 1.8 mm.

Head. Uniformly medium brownish yellow with vermiform marks on vertex
and frons. Antennae close to each other, with a small interantennal carina.

Dorsal surface of labrum (Fig. 1) with about nine simple stout setae in the distal
half and about six small fine setae in the proximal half; ventral surface with four small
pointed setae near lateral margin; distal margin fringed with two kinds of setae: the
seven lateral ones long and strongly feather-like and the median ones shorter and only
slightly feather-like.

Hypopharynx (Fig. 2): lingua covered with minute stout setae, superlingua with
thin setae apically.

Right mandible (Fig. 3) with two very approximated incisors, outer incisor with
three cusps, inner one with four cusps, outer margin straight; very thin setae in proxi-
mal half of inner margin of incisors; prostheca with one apical elongated denticle and
thirteen short and slender denticles; tuft of few stout setae between prostheca and mo-
la; tuft of setae at apex of mola reduced to two or three setae.

Left mandible (Fig. 4): incisors with seven cusps, outer margin straight; pros-
theca with five broad denticles and a comb-shaped structure; tuft of a few stout setae
between prostheca and mola; tuft of setae at apex of mola absent.

Maxillae (Fig. 5) with four broad teeth, one of them clearly separated but not
opposed to others; two rows of setae, the first one formed by abundant small setae end-
ing with three to five setae twice as long as the others, and the second by two long stout
spine-like setae; row of four stout setae at base of galea; one single stout seta perpen-
dicular to margin of galea; palp two-segmented with a few thin setae especially at the
apex of the second segment, segment 1 approximately 0.8 x length of segment 2,
second segment apically clearly turned outward.

Labium (Fig. 6) with glossae slightly shorter than paraglossae; margins of
glossae with stout setae increasingly longer towards apex, dorsally with about ten thin
setae; paraglossae falcate, with long and stout setae; labial palp three-segmented;
segment 1 subrectangular, 0.8 x length of segment 2 and 3 combined; segment 2 with
a dorsal oblique row of five long pointed setae; segment 3 truncated, with thin setae
more abundant apically and few small pointed setae apically.

Thorax. Thorax uniformly medium brownish yellow without any pattern. Pro-
and mesothorax rounded, giving the larva a humped posture in lateral view. Legs
yellow except for brownish yellow apex of femora.

660 J.-L. GATTOLLIAT

Fics 1-6
Larval structures of Nigrobaetis colonus sp. n.: 1: labrum (left : ventral view; right: dorsal view).
2: hypopharynx. 3: right mandible. 4: left mandible. 5: left maxilla. 6: labium.

Forelegs (Fig. 7). Femora dorsally with one row of nine long and pointed setae,
without another row of setae subparallel to dorsal margin, dorsoapical setal patch
formed by 2 stout setae; ventral margin with stout pointed setae; anterior face almost
bare.

THE GENUS NIGROBAETIS IN THE AFROTROPICS 661

Tibiae with only very thin setae dorsally; ventral margin with a few small
pointed setae; tibiopatellar suture and subproximal arc of setae absent.

Tarsi with only very thin setae dorsally; ventral margin with few small pointed
setae; tarsal claws (Fig. 8) hooked and slender, with one row of about fifteen acute teeth
increasing in length toward the apex.

Hindwing pads absent.

Abdomen. Tergal coloration brownish yellow without any pattern. Texture of
terga as in Fig. 21.

Sterna brownish yellow, with scales and scale bases; posterior margin without
spination.

Gills on segments 1 to 7, elliptic with serrated margin in distal half; tracheation
reduced to a central vein (Fig. 10); first gill half the length of other gills, without
tracheation (Fig. 9); gill 7 similar to gills 2 to 6.

Paraproct (Fig. 11) with very few scale bases, margin with about six long
pointed teeth and a few small ones; postero-lateral extension with a few scale bases,
margin with numerous slender teeth.

Cerci and median caudal filament yellow without dark band or annulation,
abundant thin setae on inner margins.

IMAGOS
Unknown.

ETYMOLOGY

The specific epithet “colonus” is a Latin noun, meaning “pioneer that colonises
a new area”.

ECOLOGY

Based on Starmühlner’s notes, larvae of Nigrobaetis colonus sp. n. were
collected in low to medium water currents (0.50 to 1 m/s) in small streams (1 to 2 m
wide and 0.05 to 0.2 cm deep) (StarmiihIner, 1979). The temperature of the water was
18°C (at 12h) and the pH 7.85. The larvae were not abundant anywhere.

Nigrobaetis cryptus sp. n. Figs 12-22

MATERIAL EXAMINED

Holotype: 1 larva, Madagascar, Rianila Basin, tributary of Sahatany River,
Antanambotsira (near Andasibe), 48°25’49” E, 18°52’53” S, 950 m, 21.02.2002. Leg.
R. Oliarinony.

Paratypes: 3 larvae, same locality as for holotype, 22.05.2001. 2 larvae, same locality,
14.09.2001. 2 larvae (1 larva on slide), same locality, 30.11.2001. 6 larvae, Madagascar, Rianila
Basin, tributary of Sahatandra River, Madiofasina (near Andasibe), 48°26’33” E, 18°55’23” S,
870 m, 13.06.2001. Leg. R. Oliarinony. 9 larvae, same locality, 14.05.2002. 5 larvae, same lo-
cality, 11.09.2001. 2 larvae, same locality, 09.10.2001. 3 larvae, same locality, 13.11.2001. 19
larvae (1 larva on slide), same locality, 27.11.2001.

LARVA
Maximum length: body 2.9 mm; cerci and median caudal filament broken.
Head. Uniformly medium brownish yellow without vermiform marks on vertex
and frons. Antennae close to each other, with a small interantennal carina.

662 J.-L. GATTOLLIAT

Fics 7-11

Larval structures of Nigrobaetis colonus sp. n.: 7: foreleg. 8: tarsal claw. 9: first gill. 10: fourth
gill. 11: paraproct.

Dorsal surface of labrum (Fig. 12) with about six simple stout setae in the distal
half and about five small fine setae in the proximal half; ventral surface with four small
pointed setae near the lateral margin; distal margin fringed with two kinds of setae: the
nine lateral ones long and strongly feather-like and the median ones shorter and slightly
feather-like.

Hypopharynx (Fig. 13): lingua with minute stout setae apically, superlingua
with thin setae apically.

Right mandible (Fig. 14) with two very approximated incisors, outer incisor
with three cusps, inner one with four cusps, outer margin expanded and curved; very
thin setae in proximal half of inner margin of incisors; prostheca with about nine
denticles, the median ones longer than the others: tuft of few stout setae between
prostheca and mola; tuft of setae at apex of mola.

THE GENUS NIGROBAETIS IN THE AFROTROPICS 663

Left mandible (Fig. 15): incisors with seven cusps, outer margin expanded and
curved; prostheca with five broad denticles and a comb-shaped structure; tuft of few
stout setae between prostheca and mola; no tuft of setae at apex of mola.

Maxillae similar to Fig. 5.

Labium (Fig. 16) with glossae slightly shorter than paraglossae; margins of
glossae with stout setae, slightly longer apically, dorsally with about seven medium
setae; paraglossae apically flattened, with long and stout setae; labial palp three-seg-
mented; segment 1 subrectangular, 0.9 x length of segment 2 and 3 combined; segment
2 with a dorsal oblique row of three long and pointed setae; segment 3 truncated, with
thin setae more abundant apically and few small pointed setae apically and ventrally.

Thorax. Brown with a broad central yellow spot, distal part of forewing pads
yellow. Pro- and mesothorax rounded, giving the larva a humped posture in lateral
view. Legs yellow.

Forelegs (Fig. 17). Femora dorsally with one row of eleven long and pointed se-
tae, without another row of setae subparallel to dorsal margin, dorsoapical setal patch
formed by two stout setae; ventral margin with stout pointed setae; anterior face bare.

Tibiae without setae dorsally; ventral margin with a few small pointed setae;
tibiopatellar suture and subproximal arc of setae absent.

Tarsı without setae dorsally; ventral margin with seven small pointed setae;
tarsal claws (Fig. 18) hooked and slender, with one row of about eleven acute subequal
teeth.

Hindwing pads absent.

Abdomen. Terga 1-5 uniformly yellow, terga 6 and 7 brown, tergum 8 brown in
proximal third and yellow in distal two thirds, terga 9 and 10 yellow. Terga with scales
and scales bases; posterior margin with triangular spines (Fig. 21).

Sterna 1-5 yellow, 6 and 7 brown, 8 and 9 yellow; with scales and scales bases;
posterior margin without spination.

Gills on segments 1 to 7, all elliptic with serrated margin in distal half;
tracheation generally absent (Fig. 20), only present in some mature larvae (Fig. 19), gill
1 as in Fig. 9.

Paraproct (Fig. 22) with abundant scale bases, margin with about seven long
pointed teeth and a few small ones; postero-lateral extension with abundant scale
bases, margin with about fifteen teeth.

Cerci and median caudal filament broken.

IMAGOS
Unknown.

ETYMOLOGY

The specific epithet “cryptus” is a Greek adjective meaning “hidden, the
existence of which remains unknown”.
ECOLOGY

The larvae were collected in two small streams (2-3 m wide and 0.15 cm deep)
with slow water currents (0.3 m/s) on the Eastern Coast of Madagascar. The water tem-
perature ranged between 16°C and 23.5°C (seasonal variations) and the pH between

664 J.-L. GATTOLLIAT

Fics 12-16

Larval structures of Nigrobaetis cryptus sp. n.: 12: labrum (left: ventral view; right: dorsal view).
13: hypopharynx. 14: right mandible. 15: left mandible. 16: labium.

6.8 and 7.3. The substratum component was mainly cobbles and gravel. Despite
intensive prospecting in this area, larvae were not collected at any other locality. As the
samples were taken every month during a whole year, observation of the larval
development indicates that the species is bivoltine, with one generation in June and one
in December, at least at these two localities.

THE GENUS NIGROBAETIS IN THE AFROTROPICS 665

Fiss 17 - 22

Larval structures of Nigrobaetis cryptus sp. n.: 17: foreleg. 18: tarsal claw. 19: third gill (mature
larva). 20: fourth gill (young larva). 21: posterior margin of fourth tergum. 22: paraproct.

Nigrobaetis bethunae Lugo-Ortiz & de Moor, 2000
MATERIAL EXAMINED

5 larvae, (S2103), South Africa, Incomati River system, Buffelspruit River, near Caro-
lina (closer to Badplaas), 30°27’02” E, 26°00’06” S, 1260 m, 16.05.2003. 10 larvae, (S2104),
South Africa, Limpopo Basin, Crocodile River, Kruger NP- near Malelane, 31°37 04° E,

666 J.-L. GATTOLLIAT

Comoros Islands

Angola

Mauritius
* Ô

La Réunion

South Africa 2?

200 600 1000 km

FIG. 23

Distribution of Nigrobaetis in southern Africa and in the western part of the Indian Ocean. Star:

N. colonus sp. n. Square: N. cryptus sp. n. Circle: Nigrobaetis bethunae (black: type locality;
grey: new localities).

25°23°57” S, 247 m, 17.05.2003. 10 larvae, (S2108), South Africa, Limpopo Basin, Sabie River,
Kruger NP- near Skukuza, 32°00°02” E, 25°09°42” S, 130 m, 18.05.2003. 4 larvae, (S2113),
South Africa, Limpopo Basin, Sabie River, Kruger NP- near Skukuza Camp, 31°26°16” E,
24°59°16” S, 289 m, 19.05.2003. 2 larvae, (S2114), South Africa, Limpopo Basin, Olifants
River, Kruger NP- near Olifants Camp, 31°40°02” E, 24°01°10” S, 189 m, 20.05.2003. 1 larva,
(S2118), South Africa, Limpopo Basin, Letaba River, Kruger NP- near Olifants Camp,
31°46°08” E, 23°59°10” S, 163 m, 21.05.2003. 1 larva, (S2128), South Africa, Limpopo Basin,
Tshirovha River, near Thohoyandou, 30°23°44” E, 22°48°55” S, 704 m, 25.05.2003. 2 larvae,
(S2134), South Africa, Limpopo Basin, Magoebas River, near Tzaneen, 30°02°34” E, 23°49’51”
S, 818 m, 27.05.2003.

All specimens collected by: Helen Barber-James (AM: Albany Museum, Grahamstown,
South Africa), Ferdy de Moor (AM), Pascale Derleth (EPFL: Ecole Polytechnique Fédérale de
Lausanne, Switzerland; MZL: Musée de Zoologie Lausanne, Switzerland), Jean-Marc Elouard
(IRD: Institut de Recherche pour le Développement, Montpellier, France), Jean-Luc Gattolliat
(MZL), Dominique Holveck (IRD), Michael Monaghan (NHM: Natural History Museum,
London, England), Michel Sartori (MZL).

LARVA

See Lugo-Ortiz & de Moor, 2000; additional morphological information given
in the following: General colouration brownish grey with yellow pattern. Head
brownish grey. Thorax brownish grey with a broad yellow stripe at base of forewing
pads; legs yellow with a transverse brown stripe on femora. Most terga brownish grey
with a yellow central symmetric yellow comma, terga 4, 8 and 9 uniformly yellow. In
light specimen: tergum 2 with a yellow central spot. In dark specimen: most terga
completely brownish grey, terga 8 and 9 yellow. Sterna uniformly yellow, except for

THE GENUS NIGROBAETIS IN THE AFROTROPICS 667

laterally brownish sterna in mature larva sterna. Comparison of this material with ma-
terial from the Cunene River indicates that the colour pattern is both locally and geo-
graphically variable.

IMAGOS
Unknown.

DISCUSSION

Larvae of Nigrobaetis colonus sp. n. and N. cryptus sp. n. differ from those of
N. niger (Linnaeus, 1761), N. digitatus (Bengtsson, 1912) and N. rhithralis (Soldän &
Thomas, 1983) by the number of gills (7 vs 6 pairs), the absence of hindwings and the
absence of stout setae on the dorsal margin of tibiae; from those of N. aciniger (Kluge,
1983) and N. harasab (Soldän, 1977) by the number of gills (7 vs 6 pairs); from those
of N. bacillius (Kluge, 1983), N. gracilis (Bogoescu & Tabacaru, 1957) and all the
Taiwanese species except N. facetus (Chang & Yang, 1994) by the absence of
hindwings.

Only N. bethunae, N. facetus and N. minutus (Müller-Liebenau, 1984) cannot
be easily separated from N. colonus sp. n. and N. cryptus sp. n. as they have the same
number of gills and no hindwing pads. Morphology of these five species is quite
similar. The degree of development of the setae between prostheca and mola of the left
mandible (Figs 4, 15), the spination of the right prostheca (Figs 3, 14), the small size
of the first pair of gills, the absence of a dark band on cerci, the colouration of the
abdomen and the size of the mature larva allow N. colonus sp. n. and N. cryptus sp. n.
to be distinguished from the three other species.

N. colonus sp. n. and N. cryptus sp. n. can be separated mainly by the shape of
the incisors of both mandibles (Figs 3, 4, 14, 15), the shape of the right prostheca (Figs
3,14), the colouration of the abdomen and the size of the larvae.

BIOGEOGRAPHY

Among the 23 genera of Baetidae occurring in Madagascar, Nigrobaetis is the
14th that is present in both Madagascar and Africa. Unlike most others, its distribution
is not restricted to the Afrotropical realm but also includes the Palearctic and Oriental
realms. With regard to the specific diversity of the genus in the different parts of its dis-
tribution area, it is reasonable to assume a Palearctic or Oriental origin. The presence
of Nigrobaetis in southern Africa and in Madagascar is probably the result of a step-
by-step colonisation: northern Africa to southern Africa and finally Madagascar. As has
been assumed for other genera (Gattolliat, 2002; Gattolliat & Sartori, 2003), it is like-
ly that the colonisation of Madagascar took place after its break-off from the
Gondwanan mainland (ca. 165 My). This implies an overseas colonization of more
than 300 km (Fig. 23).

La Réunion is a young volcanic island (ca. 3 My) and the presence of mayflies
there implies overseas dispersal. Based on geographical distance, the probable origin
of the mayflies should be either Mauritius (200 km) or Madagascar (800 km); the
African mainland is 2250 km distant. Nigrobaetis is not known from Mauritius despite

668 J.-L. GATTOLLIAT

the island’s longer history (ca. 8 My). The only mayfly larvae recently collected and
described from Mauritius belong to the genus Cloeon (McCafferty & Mauremootoo,
2001), a genus also present on La Réunion (Starmühlner, 1979). Thus a tentative
conclusion is that Nigrobaetis colonised from Madagascar; however, its limited distri-
bution on Madagascar and La Réunion, and the poor knowledge of the aquatic inver-
tebrates of Mauritius, suggest that Nigrobaetis occurs on Mauritius but has not been
collected.

Nigrobaetis bethunae (southern Africa), N. cryptus sp. n. (Madagascar) and
N. colonus sp. n. (La Réunion) are geographically the closest reported members of the
genus. This morphological study confirms the affinities among the three species.
Nigrobaetis bethunae possesses a wide distribution in southern Africa, but is never
abundant, and direct colonisation from Africa seems unlikely because of the more than
2250 km distance.

As noted above, Nigrobaetis colonus sp. n. is morphologically closely related to
species of the Oriental realm [N. facetus (Taiwan) and N. minutus (Malaysia)]. Because
the Baetidae fauna of the Oriental realm remains poorly known, the recorded
distribution range of Nigrobaetis remains incomplete. The presence of the genus in
India or Sri Lanka is not recorded but is possible. However, even if its presence is
confirmed, an Oriental origin for La Réunion seems unlikely because of the distance;
ca. 3500 km between India and La Réunion.

Considering the present abundance and distribution, common genera should be
the most probable candidates for overseas interchanges. In the case of the Comoros
Islands, which are also recent volcanic islands, only the common and widespread
genera Labiobaetis Novikova & Kluge, 1987, Afroptilum Gillies, 1990 and Cloeodes
Traver, 1938 were successful colonists (unpublished data based on the material
collected by Starmühlner’s expedition). Consequently, why the rare Nigrobaetis was
able to colonize La Réunion while other abundant and widespread genera did not
remains an important question.

In conclusion, the occurrence of Nigrobaetis on La Réunion is surprising and
the origin of Nigrobaetis colonus sp. n. remains unclear, even if an Afrotropical origin
is probably the most likely. Is the small size of Nigrobaetis part of the explanation of
this riddle? Was Nigrobaetis once much more abundant in the Afrotropical area than it
is at present time? Is this genus perhaps more widespread than it is yet recorded? The
question remains open. In clarifying the affinities of Nigrobaetis colonus sp. n.,
molecular phylogenetic studies could greatly help to solve this puzzle and determine
the colonizing source. The presence of Nigrobaetis on La Réunion indicates success-
ful overseas colonisation of at least 800 km and constitutes the final proof that the
dispersal capacities of mayflies are greatly underestimated. It means that colonization
over long distances must be taken in account when biogeographical scenarios are
established.

ACKNOWLEDGEMENTS

I want to thank Ranalison Oliarinony (Museum of Zoology, Lausanne,
Switzerland) for providing the specimens from Madagascar together with ecological
information. I am greatly indebted to Dr M. Hubbard (A&M University, Tallahassee,

THE GENUS NIGROBAETIS IN THE AFROTROPICS 669

Florida, USA) for the loan of Starmühlner’s material, Emilie Barthe and Stephanie
Merlin from ORE (Organisme Réunionais des Eaux) for the gift of specimens collected
during water quality surveys, Christian Guillermet (Antibes, France) for information
concerning Baetidae from La Réunion, and to Helen M. Barber-James (Dept. Fresh-
water Invertebrates, Albany Museum, Grahamstown, South Africa) for the organisation
of our fieldtrip in South Africa. I also want to express my appreciation to Dr Michel
Sartori (Museum of Zoology, Lausanne, Switzerland) and Helen M. Barber-James and
Michael Monaghan (Natural History Museum, London, England) for useful comments
on this paper. This paper is contribution n° 51 to the series “Aquatic Biodiversity of
Madagascar”.

REFERENCES

ELOUARD, J.-M. & GIBON, F.-M. 2001. Biodiversité et biotypologie des eaux continentales mal-
gaches. IRD, Montpellier, 447 pp.

GATTOLLIAT, J.-L. 2002. The genus Cloeon (Insecta : Ephemeroptera) in Madagascar. Mitteillung
der Schweizerischen Entomologischen Gesellschaft 74: 195-209.

GATTOLLIAT, J.-L. & SARTORI, M. 2003. An overview of the Baetidae of Madagascar (pp. 135-
144). In: GAINO, E. (ed.). Research Update on Ephemeroptera and Plecoptera. University
of Perugia, Italy, Perugia.

KANG, S.C., CHANG, H.C. & YANG, C.T. 1994. A revision of the genus Baetis in Taiwan
(Ephemeroptera, Baetidae). Journal of Taiwan Museum 47: 9-44.

LuGo-ORTIZ, C.R. & DE Moor, FC. 2000. Nigrobaetis Novikova & Kluge (Ephemeroptera:
Baetidae): first record and new species from southern Africa, with reassignment of one
northern African species. African Entomology 8: 69-73.

MCCAFFERTY, W.P. & MAUREMOOTOO, J.R. 2001. The first mayfly taken from the island of
Mauritius (Ephemeroptera, Baetidae). Ephemera 2: 105-111.

MÜLLER-LIEBENAU, I. 1969. Revision der europäischen Arten der Gattung Baeris Leach, 1815
(Insecta, Ephemeroptera). Gewässer und Abwässer 48/49: 1-214.

MÜLLER-LIEBENAU, I. 1984. New genera and species of the family Baetidae from West-Malaysia
(River Gombak) (Insecta: Ephemeroptera). Spixiana 7: 253-284.

Novikova, E.A. & KLUGE, N.Y. 1987. [Systematics of the genus Baetis (Ephemeroptera,
Baetidae) and a description of a new species from central Asia] (in Russian). Vestnik
Zoologii 4: 8-19.

SOLDAN, T. 1977. Three new species of mayflies (Ephemeroptera) from the mist oasis of Erkwit,
Sudan. Acta Entomologica Bohemoslovaca 74: 289-294.

STARMÜHLNER, F. 1979. Results of the Austrian hydrobiological mission, 1974, to the Seychelles,
Comores and Mascarene Archipelagos: Part 1, preliminary report. Annalen des Natur-
historischen Museums in Wien 82: 621-742.

WALTZ, R.D., MCCAFFERTY, W.P. & THOMAS, A. 1994. Systematics of Alainites n. gen., Diphetor,
Indobaetis, Nigrobaetis n. stat., and Takobia n. stat. (Ephemeroptera, Baetidae). Bulletin
de la Société d'Histoire Naturelle de Toulouse 130: 33-36.

Zwick, P. 2003. Biogeographical pattern (pp. 94-102). Jn: RESH, V.H. & CARDE, R.T. (eds).
Encyclopedia of Insects. Academic Press.

REVUE SUISSE DE ZOOLOGIE 111 (3): 671-672; septembre 2004

First record of Hyssopus pallidus (Askew, 1964) for Switzerland
(Hymenoptera: Eulophidae)

Kathrin TSCHUDI-REIN, Nayuta BRAND, Ute KÜHRT & Silvia DORN
Institute of Plant Sciences, Applied Entomology, Swiss Federal Institute of
Technology (ETH), Clausiusstrasse 25 / NW, CH-8092 Zurich, Switzerland.

First record of Hyssopus pallidus (Askew, 1964) for Switzerland
(Hymenoptera: Eulophidae). - The parasitoids emerged from apples that
were collected in orchards in the cantons of Aargau and Zurich and were
brought into the laboratory. They were found to have developed on both
known hosts, i.e. Cydia pomonella (L., 1758) and Cydia molesta (Busck,
1916) (Lepidoptera: Tortricidae).

Keywords: Parasitoid - Hyssopus pallidus - Cydia pomonella - Cydia
molesta - first record - Switzerland.

Hyssopus pallidus (Askew, 1964) is a gregarious, idiobiont ectoparasitoid of
late larval instars of the codling moth, Cydia pomonella (L., 1758), a widely distributed
major fruit pest, and of the Oriental fruit moth, Cydia (Grapholita) molesta (Busck,
1916), an economically important pest of stone fruit orchards and in a rapidly
expanding area also of apple orchards (reviewed by Natale er al., 2003). In apple
orchards under integrated and organic regimes augmentative release of this parasitoid
could substitute the labor intensive and therefore costly post-harvest fruit removal
(sanitation) often needed to lower codling moth populations to a level that allows the
successful employment of the mating disruption technique (Mattiacci et al., 1999). The
behavioral ecology of this species is being studied at our institute (Gandolfi et al.,
2003; Mattiacci et al., 1999; Mattiacci et al., 2000) and it is thus with great interest that
we take note of its presence here in Switzerland.

The species was first described by Askew (1964) as Elachertus pallidus from
Castagniers (Alpes Maritimes), France, and from Sofia, Bulgaria, and was then trans-
ferred to the genus Hyssopus by LaSalle and Huang (1994). Morphology and develop-
ment of its immature stages were described by Tschudi-Rein and Dorn (2001).

A total of 26 male and 78 female H. pallidus emerged in July and August in the
laboratory from apples collected between June and August 2003 in three apple
orchards in Northern Switzerland: Endingen (395 m a.s.l., N 47° 31° 46.7°’/ E 8° 17°
1.6”) and Schlatt b. Leuggern (436 m a.s.l., N 47° 33’ 40.5”/ E 8° 13° 20.7”) in the
canton of Aargau and Steinmaur (442 m a.s.l., N 47° 29’ 34.1”/ E 8° 26° 44.1”) in the
canton of Zurich. Apples lying on the ground in the field with indications of infestation
by Cydia species had been collected and brought to the laboratory for an ongoing

Manuscript accepted 12.12.2003

672 HYSSOPUS PALLIDUS IN SWITZERLAND

behavioral study of the codling moth. Upon detection of the emerging parasitoids,
more apples were collected and these were kept singly in containers. This allowed the
tracing of emerging parasitoids back to their host larva in the apple. The host larvae
were identified with the help of larval keys of Brown (1987) and of Pfeiffer (no date).
Male and female A. pallidus were collected from all the 3 orchards from which apples
had been collected, and from their two known hosts, Cydia pomonella and Cydia
molesta. This latter host species was found as a parasitized larva in an apple from the
orchard in Schlatt b. Leuggern in the canton of Aargau.

Voucher specimens have been deposited in the Natural History Museum Geneva.

ACKNOWLEDGEMENTS
We wish to thank Andreas Müller for helpful comments.

REFERENCES

ASKEW, R. R. 1964. On the biology and taxonomy of some European species of the genus
Elachertus SPINOLA (Hymenoptera, Eulophidae). Bulletin of Entomological Research
55: 53-58.

BROWN, R. L. 1987. Tortricidae (Tortricoidea) (pp. 419-427). In: STEHR, F.W. (ed.). Immature
Insects. Kendall / Hunt Publishing Co, Dubuque, 427 pp.

GANDOLFI, M., MATTIACCI, L. & DORN, S. 2003. Mechanisms of behavioral alterations of para-
sitoids reared in artificial systems. Journal of Chemical Ecology 29: 1871-1887.
LASALLE, J. & HUANG, D. 1994. Two new Eulophidae (Hymenoptera: Chalcidoidea) of eco-

nomic importance from China. Bulletin of Entomological Research 84: 51-56.

MATTIACCI, L., HÜTTER, E. & Dorn, S. 1999. Host location of Hyssopus pallidus, a larval para-
sitoid of the codling moth, Cydia pomonella. Biological Control 15: 241-251.

MATTIACCI, L., HÜTTER, L., SCHOCH, D., SCASCIGHINI, N. & DORN, S. 2000. Plant-odour mediates
parasitoid host handling and oviposition in an endophytic tritrophic system. Chemo-
ecology 10: 185-192.

NATALE, D., MATTIACCI, L., HERN, A., PASQUALINI, E. & Dorn, S. 2003. Response of female
Cydia molesta (Lepidoptera: Tortricidae) to plant derived volatiles. Bulletin of
Entomological Research 93: 335-342.

PFEIFFER, D. G. (no date). A key to the internal feeders of apple. Retrieved August 7, 2003, from
http://www.ento.vt.edu/Fruitfiles/AppleInternalKey.html (3 pp).

TSCHUDI-REIN, K. & Dorn, S. 2001. Reproduction and immature development of Hyssopus
pallidus (Hymenoptera: Eulophidae), an ectoparasitoid of the codling moth. European
Journal of Entomology 98: 41-45.

REVUE SUISSE DE ZOOLOGIE 111 (3): 673-682; septembre 2004

A new species of Hyphessobrycon (Characiformes, Characidae)
from the Esteros del Iberä wetlands, Argentina

Adriana E. ALMIRON! , Jorge R. CASCIOTTA!, José A. BECHARA?

& Federico J. RUIZ DIAZ?

! Divisiön Zoologfa Vertebrados, Facultad de Ciencias Naturales y Museo,
UNLP, Paseo del Bosque, 1900 La Plata, Argentina.
E-mail: aalmiron@museo.fcnym.unlp.edu.ar

2 Instituto de Ictiologfa del Nordeste, Facultad de Ciencias Veterinarias, UNNE,
Sargento Cabral 2139, 3400 Corrientes, Argentina.

A new species of Hyphessobrycon (Characiformes, Characidae) from
the Esteros del Ibera wetlands, Argentina.- Hyphessobrycon auca sp. n.
is described from the Esteros del Iberä wetlands. Hyphessobrycon auca
differs from the remaining species of the genus by the presence of one
maxillary tooth with 5 cusps; dentary low with teeth decreasing in size
anteroposteriorly; males with hooks in all fins; mature females with hooks
on pelvic-fin rays; one humeral spot vertically elongated, bounded by a light
area; a second lateral spot may be faint or well developed; a wide lateral
band ending in a conspicuous caudal spot.

Keywords: Characiformes - Characidae - Hyphessobrycon - Esteros del
Iberä wetlands - new species - Argentina.

INTRODUCTION

Nine species of the genus Hyphessobrycon were recorded for the Rio de la Plata
basin in Argentina, eight of them inhabit the Esteros del Iberä wetlands system
(Almirön er al., 2003). The Esteros del Iberd wetlands is one of the freshwater
environments which has a high diversity of fishes in Argentina. More than 126 species
have been recorded from this wetland, a number that represents about one third of the
fish fauna known for Argentina (Casciotta et al., 2003).

The aim of this paper is to describe a new species of the genus Hyphessobrycon
from the Esteros del Iberä wetlands.

MATERIAL AND METHODS

The specimens examined in this study were cleared and counterstained (C&S)
following Taylor & Van Dyke (1985). Measurements are straight distances taken with
calliper to the nearest 0.1 mm. Peduncle length is the distance between last branched
anal-fin ray and hypural joint. Vertebral count includes four vertebrae corresponding to
the Weberian apparatus and also the complex centrum as one element. Asterisk
indicates holotype.

Manuscript accepted 15.01.2004

674 A. E. ALMIRON ET AL.

Material is deposited in the Asociaciön Ictiolögica, La Plata, Argentina (AI);
Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Argentina
(MLP); Museo Argentino de Ciencias Naturales Bernardino Rivadavia, Argentina
(MACN-Ict); Muséum d’histoire naturelle, Genève, Switzerland (MHNG).

COMPARATIVE MATERIAL (standard length, SL): Hyphessobrycon anisitsi (Eigenmann,
1907): MHNG 2493.72, 8 ex., 41.4-42.7 mm SL, (2 C&S), Paraguay, Caazapa, Arroyo Moroti
15 km SE from Tavai. MHNG 2493.063, 6 ex., 35.1-46.5 mm SL, Paraguay, Caazapa, Arroyo
Yvyra-pary, 2 km from Tavai. MHNG 2493.86, 2 ex., 29.5-31.0 mm SL, Paraguay, Caaguazu,
small affluent of the rio Giiyrau-gua. MHNG 2054.2-44, 5 ex., 28.0-38.3 mm SL, Paraguay,
Asunciön, stream in Colonia Thompson. MHNG 2493.88, 2 ex., 28.2-29.4 mm SL, Paraguay,
Guaira, stream Ovie, near of the road from Villarica. AI 133 (ex MHNG 2493.063) 2 ex., 23.0-
37.6 mm SL, Paraguay, Caazapa, Arroyo Yvyra-pary, 2 km from Tavai. Hyphessobrycon arianae
Uj & Gery, 1989: MHNG 2412.79 (holotype), 22.7 mm SL, Paraguay, Dept. Caaguazu, rio
Giiyrau-gua, affl. of the rio Monday, 3 km East of Juan Frutos. MHNG 2412.80-81 (paratypes),
12 ex., 17.6-23.6 mm SL, (2 ex. C&S), same locality as holotype. Hyphessobrycon elachys
Weitzman, 1984: MLP 6431, 3 ex., 16.0-20.0 mm SL, Argentina, Corrientes Province, San
Cosme. Hyphessobrycon eques (Steindachner, 1882): MLP 8999, 3 ex., 23.3-27.9 mm SL,
Argentina, Corrientes, rio Santa Lucia. JRC Pers. Collection, 2 ex., 28.7-29.6 mm SL (C&S),
Argentina, Corrientes Province, Bella Vista, Riacho Carrizal. Hyphessobrycon guarani Mahnert
& Géry, 1987: MHNG 2366.99 (holotype), 29.8 mm SL, Paraguay, rio Alto Paranä in Puerto
Bertoni, Dept. Alto Parand. MHNG 2366.100 (paratypes), 7 ex., 23.5-29.4 mm SL, (2 ex. C&S),
same locality as holotype. Hyphessobrycon igneus Miquelarena, Menni, Löpez & Casciotta,
1981: JRC Pers. Collection, 3 ex., 25.9-29 mm SL Argentina, Esteros del Iberä, Laguna
Fernandez. Hyphessobrycon luetkeni (Boulenger, 1887): MLP 8796, 9 ex., 24.4-35.0 mm SL,
Argentina, Formosa Province, creek in the national road Formosa-Clorinda, 37 Km far from
Clorinda. MLP 6451, 13 ex., 14.2-21.6 mm SL, Argentina, Formosa Province. Hyphessobrycon
meridionalis Ringuelet, Miquelarena & Menni, 1978: MLP 8407, 2 ex., 32.2-34.0 mm SL,
Argentina, Corrientes Province, pond in road Bella Vista-San Roque. JRC Pers. collection, 3 ex.,
33.0-40.9 mm SL (C&S), Argentina, Buenos Aires Province, Berisso, Los Talas ponds.
Hyphessobrycon wajat Almiron & Casciotta, 1999: MLP 9321, (holotype), 27.6 mm SL,
Argentina, Corrientes Province, rio Paranä basin, Laguna Brava. MLP 9322, 5 paratypes, 29.2-
31.0 mm SL, Argentina, Corrientes Province, Laguna Iber4. MHNG 2593.96, 5 paratypes, 28.5-
30.0 mm SL, Argentina, Corrientes Province, Laguna Iberä.

RESULTS

Hyphessobrycon auca sp. n. Figs 1-8, Table 1

Holotype. MACN-Ict 8647, 51.2 mm SL, female, Argentina, Corrientes, Esteros del
Iberä, pond in San Juan Poriahü farm (27°41°53”S-57°12’17”W), coll. J. Casciotta, A. Almirön
& F. Ruiz Diaz, March 2003.

Paratypes. AI 129, 6 ex. (1 C&S), 42.0-49.9 mm SL, Argentina, Corrientes, Esteros del
Iberä, pond in San Juan Poriahi farm (27°41’39””S-57°12’56”W), coll. J. Bechara, J. Ortiz & S.
Sanchez, May 2001. MHNG 2644.023, 4 ex., 42.4-52.8 mm SL, Argentina, Corrientes, Esteros
del Iberä, pond in San Juan Poriahü farm (27°41’39”S-57°12’56”W), coll. J. Bechara, J. Ortiz
& S. Sanchez, May 2001. AI 130, 5 ex., 42.6-50.0 mm SL, Argentina, Corrientes, Esteros del
Iberä, pond in San Juan Poriahü farm (27°42°S-57°11’W), coll. J. Casciotta, A. Almiron & F.
Ruiz Diaz, March 2003.

Non type material: AI 131, 4 ex. (C&S), 43.7-48.9 mm SL, Argentina, Corrientes,
Esteros del Iberä, pond in San Juan Poriah farm (27°42’S-57°11°W), coll. J. Casciotta, A.
Almirön & F Ruiz Diaz, March 2003.

DIAGNOSIS
Hyphessobrycon auca is distinguished from its congeners by the following
combination of characters: one maxillary tooth with 5 cusps; dentary low (dentary

NEW SPECIES OF HYPHESSOBRYCON 675

depth 40.7-47.6 in % of dentary length) with teeth decreasing in size anteroposteriorly;
males with hooks in all fins; mature females with hooks on pelvic-fin rays; one humeral
spot vertically elongated, bounded by a light area; a second lateral spot may be faint or
well developed; a wide lateral band ending in a conspicuous caudal spot.

DESCRIPTION

Morphometrics of holotype and paratypes are presented in table 1. Maximum
body depth located immediately anterior to dorsal-fin origin (Fig. 1). Females slightly
convex on snout, concave between eye to posterior tip of supraoccipital spine, and
convex from supraoccipital to base of first dorsal-fin ray. Males with dorsal profile
straight from snout to posterior tip of supraoccipital spine, slightly convex from this
point to base of first dorsal-fin ray. Dorsal profile slanting ventrally from dorsal-fin
origin to caudal peduncle in both sexes. Dorsal and ventral profile of caudal peduncle
straight or scarcely concave. Ventral profile of body arched from tip of lower jaw to
anal-fin origin, posterodorsally slanted along anal-fin base. Vent between bases of
pectoral and pelvic fins transversally rounded, compressed between pelvic and anal fins.

Dorsal-fin origin almost equidistant from tip of snout and base of caudal fin.
Pelvic-fin base before anterior to vertical through dorsal-fin origin. Adipose fin slightly
anterior to base of last branched anal-fin rays. Females with tip of pectoral fin reaching
pelvic-fin origin, males with tip of pectoral fin surpassing pelvic-fin origin. Females
with tip of pelvic fin not reaching anal-fin origin; males with tip of pelvic-fin
surpassing that origin.

TAB. 1: Morphometric data of the holotype and 15 paratypes of Hyphessobrycon auca sp. n.
Minimum, maximum, and average between parenthesis. Standard length expressed in mm.

holotype males females
SE SEZ 42.0-45.7 45.7-52.8
Percents of SL
Predorsal distance SIT, 53.0-54.2 (53.3) 339-3987 (54:3)
Preventral distance 48.4 46.9-48.5 (47.3) 46.8-50.2 (48.6)
Preanal distance 63.1 59.7-63.7 (62.0) 63.1-66.3 (64.8)
Body depth 36.1 31.7-34.7 (33.4) 34.1-38.3 (36.4)
Dorsal-fin base 133 12.8-14.4 (13.8) 13.1-14.6 (13.8)
Anal-fin base 307 30:8-33.320 1324) 27.8-31.9 (30.3)
Pectoral-fin length 225] 23.0-26.2 (24.1) 21.8-25.2 (23.4)
Pelvic-fin length 177.6 18.5-21.2 (19.6) 17.0-19.9 (18.1)
Distance between pectoral
and pelvic-fin origins 2181 20.6-22.6 (21.4) 19.0-23.7 (21.5)
Distance between pelvic
and anal-fin origins 19.3 14.8-18.9 (17.4) 17.6-20.7 (19.0)
Head length 28.9 28.4-30.5 (29.6) 28.3-30.4 (29.4)
Peduncle depth 11.1 117221208 70107) 10.4-12.0 (11.4)
Percents of HL
Snout length 23.6 23.1-25:6 (23.9) 20.7-27.1 (23.6)
Eye diameter 39.9 37.7-40.0 (38.7) 36.7-42.9 (39.3)
Interorbital length SII 30.8-35.3 (32.2) 30.0-32.8 (31.0)
Postorbital length 43.9 41.7-46.8 (44.0) 40.7-46.9 (43.5)

Maxillary length 297] ITS (29:35) 27723228 (601)

676 A. E. ALMIRÖN ET AL.

Fic. 1

Hyphessobrycon auca sp. n., holotype: MACN-Ict 8647, 51.2 mm SL, female, Argentina,
Corrientes, Esteros del Iberä, pond in San Juan Poriahu farm.

Dorsal fin with ii, 9 rays; posterior margin of dorsal fin straight, last unbranched
and first branched dorsal-fin rays longest. Males with hooks on last unbranched and
first eight branched rays. Hooks very small, directed ventrally. One hook by segment,
maximum 12 hooks on 5" ray.

Anal fin with iii-v, 21-25 rays; 21(2), 22(10), 23(9), 24(5), 25(2*). Males with
posterior margin almost straight; females with last unbranched and first six branched
rays produced forming a lobe. Males bearing small hooks on last unbranched and the
first 21 branched rays. Hooks directed outward and curved dorsally; one pair of hooks
on each segment, maximum 14 pairs on each ray. Branched rays with hooks on poste-
rior branch, few hooks, very small, on distal portion of anterior arm.

Caudal-fin with 1 unbranched and 9 branched principal rays in upper lobe; 8
branched and 1 unbranched principal rays in lower lobe. Males (2 ex.) with few small
hooks on distal tips on middle caudal-fin rays. Lower lobe slightly longer.

Pectoral fin with i,11*-12 rays. Posterior margin of pectoral fin slightly rounded.
Males (3 ex.) with small hooks on rays, one specimen with hooks on all branched rays.

Pelvic fin with 1,6-7* rays, posterior margin of fin slightly rounded. Males with
hooks on unbranched and branched rays. One pair of hooks on each segment curved
rostrally, maximum 14 hooks on each ray. Mature females with hooks on pelvic fins,
rather smaller than those of males (Figs 2-3). Head short, less than 1/3 of SL, mouth
terminal and horizontal; snout short, blunt. Third infraorbital not reaching sensory tube
of preopercle. Lower jaw slightly longer than upper jaw. Five specimens (C&S) with
premaxillary ascending process triangular; alveolar process bearing two series of teeth,
each tooth with a central cusp larger. Outer row with 4 (4 ex.), 5 (1 ex.) teeth, with 3
(4 ex.) to 5 (1 ex.) cusps (Fig. 4). Inner series of premaxilla with 5 wide teeth; sym-
physial tooth slender, with 4 to 5 cusps; remaining teeth with 4 to 9 cusps (Fig. 5), cen-
tral cusp longer than the others. Maxilla with long anterodorsal and laminar processes,
the last one surpassing vertical through anterior orbital margin. One wide maxillary

NEW SPECIES OF HYPHESSOBRYCON 677

Fics 2-3

Hyphessobrycon auca sp. n., left pelvic-fin rays in dorsal view showing the bony hooks: 2,
males; 3, females. Scale= 1 mm.

tooth with 5 cusps (Fig. 4). Dentary low (40.7-47.6% dentary depth/dentary length)
with 8 to 12 teeth decreasing in size anteroposteriorly. Four large teeth with 5 to 8
cusps, 1 or 2 smaller one with 3 to 5 cusps, usually 5 cusps, and 3 to 7 much smaller
with 1 to 3 cusps, usually 3 cusps (Fig. 6).

Lateral series with 32-36 scales; 32(1), 33(1), 34(8), 35(6*), and 36(1). Lateral
line with 5 to 22 perforated scales; 5(1), 7(1), 8(2), 9(4*), 10(2), 11(1), 12(4), 13(4),
15(1), 22(1); two specimens with complete lateral line (34). Five to six scales between
dorsal-fin origin and lateral line; 4 or 5 scales between lateral line and ventral-fin
origin. Ten to twelve scales between supraoccipital process and dorsal-fin origin. Six
to eleven rectangular scales placed on anal-fin base. Scales covering basal fifth of
caudal lobes. Pelvic axillary scale without hooks on its posterior area in males.Gill
rakers: 6-10/10-12. Vertebral counts: 33-34.

Coloration upon capture: Upper half of body darker, wide silvery lateral band
ending in a caudal spot, lower half silvery. Base of caudal- fin lobes black and yellow;
most of the lobe red, distal margin of them hyaline; middle caudal rays black sur-
rounded by yellow. Most of anal fin red, distal margin gray or hyaline. Dorsal fin pale
yellow, pectoral fin hyaline and pelvic fin reddish (Fig. 7).

Coloration in alcohol preserved specimens: Background pale brown, dorsal
region of flanks and head darker, one dark midline along body on dorsum. Dark humer-
al spot vertically elongated, bounded by a clear area. Second lateral spot faint. Dark
wide lateral band ending in a caudal spot. Dorsum of head, premaxilla, and dorsal half
of maxilla, dark.

Dorsal fin hyaline with black chromatophores on the unbranched rays, dark
chromatophores on anterior and posterior margins of branched rays. Anal fin hyaline

678 A. E. ALMIRON ET AL.

Fics 4-5

Hyphessobrycon auca sp. n., 4, upper jaw in lateral view; 5, detail of third tooth of inner pre-
maxillary series in medial view. Scale= 1 mm.

—

FIG. 6
Hyphessobrycon auca sp. n., right lower jaw in medial view. Scale= 1 mm.

NEW SPECIES OF HYPHESSOBRYCON 679

FIG. 7
Hyphessobrycon auca sp. n., upon capture, pond in Esteros del Iberä wetlands.

with large black chromatophores on distal margin of anal fin, forming a faint band in
some specimens. Black caudal spot on caudal peduncle extended on middle caudal-fin
rays; remaining caudal-fin rays with dark chromatophores on their margins. Pectoral
and pelvic fins hyaline, few chromatophores along ray surfaces.

Chromatophores completely covering scales surface on upper half of flank,
more dense on posterior margin. Scales on lower half of flanks with scattered chro-
matophores on their surface.

Sexual dimorphism: Males of Hyphessobrycon auca have a preanal distance
shorter (59.7-63.7 vs. 63.1-66.3% SL) and lower body depth (31.7-34.7 vs. 34.1-38.3%
SL), and caudal peduncle depth (100.0-108.2 vs. 107.5-118.4% caudal peduncle
length). Also they bear bony hooks on all fin rays; hooks present on dorsal and caudal
fin are tiny, thus they are difficult to see in alcoholic preserved specimens. Mature
females bear hooks on pelvic-fin rays, they are smaller than those present on pelvic fin
of males (Fig. 3). Also, females have a small lobe on first six branched anal-fin rays,
absent in males.

ETYMOLOGY

The invariable specific epitet auca is a Mapuche word meaning rebel.

DISTRIBUTION

Hyphesobrycon auca sp. n. is known from two isolated ponds placed inside of
the farm San Juan Poriahu in the northwest of Esteros del Iberä wetlands (Fig. 8).

680 A. E. ALMIRÖN ET AL.

57°

PARAGUAY

RIO PARANA

Fic. 8

Geographical distribution of Hyphessobrycon auca sp. n., black circle show the localities in the
Esteros del Iberä.

HABITAT AND ASSOCIATED ICHTHYOFAUNA

Hyphessobrycon auca sp. n. was collected in two small lentic waterbodies
located on a large sand bar (150 km long), that was deposed by the rio Paranä during
the Pliocene. From a topographic standpoint, ponds are placed at the headwaters of the
Ibera basin and make part of a complex of interconnected small lakes dispersed over
the sand bar. As many others in the area, ponds were probably generated by eolic and
pseudokarstic processes. Sampled ponds have a surface of about 1.5 and 12.5 ha, their
form is near circular, with a sandy bottom covered by submerged macrophytes domi-
nated by Egeria naias. In the littoral areas, patches of the water hyacinth Eichhornia
azurea and some unidentified Gramineae were observed.

Ponds and the connected aquifer are fed only by rain water, and they have low
conductivity (11.6-20.8 wS cm) and acidic pH (5.6-6.8). Waters are very soft
(hardness 2-5 mg CaCo3 I"!), low in chloride (0.8 to 1.9 mg I!), as well as in alkalinity
(2-9 mg CaCo3 ll). Nutrient content was also negligible (less than 10 yg I! of
dissolved orthophosphate). Transparency is high, being impossible to estimate with

NEW SPECIES OF HYPHESSOBRYCON 681

Secchi disk due to the low depths in sampling areas (0.6-1.3 m). Dissolved oxygen
concentrations were generally high during sampling (6.7-10.2 mg I, 86.1-110.9%
saturation), and water temperature varied from 13.5 to 20.8°C.

Although different kinds of environments were sampled in the area, including
larger lakes, marshes, and small creeks, the new species was only found in the two de-
scribed ponds. The other fish caught with a purse seine in the same samples belonged
to the following species: Acestrorhynchus pantaneiro, Aphyocharax rathbuni, Chara-
cidium rachovii, Characidium cf. zebra, Cyphocharax cf. spilotus, Hyphessobrycon
eques, Hoplias malabaricus, Metynnis mola, Moenkhausia intermedia, Serrasalmus
spilopleura, Serrapinnus kriegi, Apistogramma borelli, Apistogramma commbrae,
Crenicichla lepidota, Cichlasoma dimerus, Gymnogeophagus balzanii, Laetacara dor-
sigera, and Phalloceros caudimaculatus.

REMARKS

The following species of the genus Hyphessobrycon were recorded from the
Rio de la Plata basin in Argentina: H. anisitsi, H. elachys, H. eques, H. guarani,
H. igneus, H. luetkeni, H. meridionalis, H. reticulatus, and H. wajat. All of them were
registered from the Esteros del Iberä wetlands except A. guarani. The color pattern of
H. eques, H. elachys, H. guarani, H. igneus, and H. reticulatus differenciate these
species from H. auca. Hyphessobrycon eques has the ground colour of body red and
H. elachys is greenish with yellow whereas H. auca is mostly silvery. Hyphessobrycon
guarani bear a black spot on dorsal-fin rays, absent in H. auca; H. igneus has the
caudal fin red whereas in H. auca is black, yellow and red. Finally, H. reticulatus has
a narrow lateral band and a dark line along the base of anal fin whereas H. auca has a
wide lateral band and it lacks a line along the anal-fin base. Hyphessobrycon guarani
has 2 to 5 maxillary teeth, Hyphessobrycon luetkeni has 2 to 3, and H. wajat 3 to 5,
whereas H. auca has only one. Hyphessobrycon meridionalis has 26 to 30 branched
anal-fin rays whereas H. auca has 21 to 25.

Hyphessobrycon auca is similar to H. anisitsi, however the new species differs
from H. anisitsi in having always one maxillary tooth with 5 cusps instead of 3 to
5 cusps. The dentary of H. auca is lower than those of A. anisitsi (dentary depth 40.7-
47.6 vs. 50.0-53.3 in % of dentary length), and the teeth decrease in size antero-poste-
riorly whereas in H. anisitsi the dentary bears 4 large teeth followed by 4 to 8 much
smaller. Males of H. auca bear hooks on dorsal-fin rays (3 ex.) and hooks on caudal
and pectoral-fin rays (2 ex.), hooks on dorsal, caudal, and pectoral-fin rays were not
found in males of A. anisitsi.

Hyphessobrycon auca is the only species of the genus with some males bearing
hooks on all fins. Hooks on pelvic-fin rays in mature females of H. auca, appear to be
a feature shared with H. anisitsi (1 female with pelvic-fin hooks). Within characids,
females bearing hooks on pelvic-fin rays was only reported for the genus Cheirodon
(Malabarba, 1998). As this author noted, hooks are usually visible in large females and
they are smaller than those of males.

682 A. E. ALMIRÖN ET AL.

ACKNOWLEDGEMENTS

We would like to express our gratitude to Marcos Garcia Rams, the owner of
the farm San Juän Poriaht for his hospitality and to C. Tremouilles (UNLP) for help
with figures. This project was financed by the National Geographic Society (Grant
7314-01) and Universidad Nacional del Nordeste (Special Grant for Iberä Project).

REFERENCES

ALMIRON, A., CASCIOTTA, J., BECHARA, J., ROUX, P., SANCHEZ, S. & TOCCALINO, P. 2003. La
ictiofauna de los esteros del Iberä y su importancia en la designaciön de la reserva como
sitio Ramsar (pp. 75-85). In: ALVAREZ, B. (ed). Fauna del Iberä. Editorial Universitaria
de la Universidad Nacional del Nordeste (EUDENE), Corrientes, Argentina, 375 pp.

CASCIOTTA, J. R., ALMIRON, A. E. & BECHARA, J. A. 2003. Los peces de la Laguna Iberd.
Editorial AI Margen, La Plata, Argentina, 203 pp.

MALABARBA, L. R. 1998. Monophyly of the Cheirodontinae, characters and major clades
(Ostariophysi: Characidae) (pp. 193-233). In: MALABARBA, L. R., REIS, R. E., VARI, R.
P., Lucena, Z. M. & LUCENA, C. A. S. (eds). Phylogeny and Classification of neotropical
fishes. EDIPUCRS, Porto Alegre, 603 pp.

TAYLOR, W. R. & VAN DYKE, G. C. 1985. Revised procedures for staining and clearing small
fishes and other vertebrates for bone and cartilage study. Cybium 9: 107-119.

REVUE SUISSE DE ZOOLOGIE 111 (3): 683; septembre 2004

Erratum

NG, H. H. & KOTTELAT, M. 2004. Amblyrhynchichthys micracanthus, a new species of
cyprinid fish from Indochina (Cypriniformes: Cyprinidae). Revue suisse de Zoologie
111 (2): 425-432.

Due to the insufficient printing quality of Fig. 1 published in the above mentioned
paper (p. 427) a new figure is printed here.

Fic. 1
Amblyrhynchichthys micracanthus, CMK 13137, paratype, 68.1 mm SL.

oso ET si wnat:

REVUE SUISSE DE ZOOLOGIE

Tome 111 — Fascicule 3

PACE, Roberto. Hygronomini e Athetini della Cina con note sinonimiche
(Coleopteras Staphylinidae),. . ee... rs ner TE
SCHATTI, Beat. A preliminary contribution to Platyceps najadum (Eichwald,
1831) and systematic status of Coluber atayevi Tunijev &
Shammakov, 1993 (Reptilia: Squamata: Colubrinae) .............
DANKITTIPAKUL, Pakawin & WANG, Xin-Ping. New species of coelotine
spiders (Araneae, Amaurobiidae) from northern Thailand II ........
ORTUNO, Vicente M. An enigmatic cave-dwelling ground beetle: Trechus
barratxinai Espafiol, 1971 (Coleoptera, Carabidae, Trechinae, Tre-
CERTA ee IO
DA GAMA, Maria Manuela. Une espèce nouvelle de Pseudosinella pro-
venant du Portugal (Insecta: Collembola). XVII® contribution ......
AMIET, Jean-Louis. Une nouvelle espèce d’Hyperolius du Cameroun (Am-
phibia@AnurasHyperolüdae)ar se sr EEE
PIEROTTI, Helio & BELLÒ, Cesare. Contributi al riordinamento sistematico
dei Peritelini paleartici. V. Istituzione di un nuovo genere e descrizione
di due nuove specie (Coleoptera, Curculionidae, Entiminae)........
LANDRY, Bernard & ROQUE-ALBELO, Lazaro. First report of Tischeriidae
(Lepidoptera) on the Galapagos Islands, Ecuador, with descriptions of
twomewzendemieispeciesHs pra ie lion an
BAUR, Hannes & Coray, Armin. A revision of the Blattodea, Ensifera and
@aeliterardescenibediby.Hr Brubstorfer. 245 en Sons,
BAUR, Hannes & Coray, Armin. The status of some taxa related to
Miramella irena (Fruhstorfer) and the type of Kisella Harz (Caelifera:
iNerididae,Melanoplinae)ae. ti e RR
DE ANDRADE, Maria L. A new species of Platythyrea from Dominican
amber and description of a new extant species from Honduras
(Eiyymenopteraskormicidae)ea nr ET
GATTOLLIAT, Jean-Luc. First reports of the genus Nigrobaetis Novikova &
Kluge (Ephemeroptera: Baetidae) from Madagascar and La Réunion
with observations on Afrotropical biogeography .................
TSCHUDI-REIN, Kathrin, BRAND, Nayuta, KUHRT, Ute & Dorn, Silvia. First
record of Hyssopus pallidus (Askew, 1964) for Switzerland
(Hymenoptera Eulophidae) ea ar ne set hae AT ee
ALMIRON, Adriana E., CASCIOTTA, Jorge R., BECHARA, José A. & RUIZ DIAZ,
Federico J. A new species of Hyphessobrycon (Characiformes,
Characidae) from the Esteros del Iberä wetlands, Argentina........
ERRATONG (NG SEs HNSAIKOTTEFAT IM 2004 er. st

Pages

457-523

525-538

539-550

551-562

563-565

567-583

585-598

599-609

611-630

631-642

643-655

657-669

671-672

673-682
683

REVUE SUISSE DE ZOOLOGIE

Volume 111 — Number 3

PACE, Roberto. Hygronomini and Athetini from China (Coleoptera, Staphy-
linidae)\withimotes onthe synonymy 300. MR
SCHATTI, Beat. A preliminary contribution to Platyceps najadum (Eichwald,
1831) and systematic status of Coluber atayevi Tunijev &
Shammakov, 1993 (Reptilia: Squamata: Colubrinae) .............
DANKITTIPAKUL, Pakawin & WANG, Xin-Ping. New species of coelotine
spiders (Araneae, Amaurobiidae) from northern Thailand II ........
ORTUNO, Vicente M. An enigmatic cave-dwelling ground beetle: Trechus
barratxinai Espafiol, 1971 (Coleoptera, Carabidae, Trechinae, Tre-
CHIMERA UL CEE
DA GAMA, Maria Manuela. A new species of Pseudosinella from Portugal
(Insecta: Collembola) XVII! Contribution." SII ee
AMIET, Jean-Louis. A new species of Hyperolius from Cameroun (Amphi-
biaeAnura bby peroludae) #222 =... 25 RENE ONE NE
PIEROTTI, Helio & BELLO, Cesare. Contributions towards a taxonomic
rearrangement of Palaearctic Peritelini. V. Erection of a new genus and
description of two new species (Coleoptera, Curculionidae, Entimi-
TAC) Fen ete : Per See ET MAZIONE
LANDRY, Bernard & ROQUE-ALBELO, Lazaro. First report of Tischeriidae
(Lepidoptera) on the Galapagos Islands, Ecuador, with descriptions of
twornewzendemieispeeiesen meer ciate ye TT IE
BAUR, Hannes & Coray, Armin. A revision of the Blattodea, Ensifera and
@aeliferaldesenibed/by/H.Eruhstorferan. ur. rt RR
BAUR, Hannes & CoraAy, Armin. The status of some taxa related to
Miramella irena (Fruhstorfer) and the type of Kisella Harz (Caelifera:
Aendidae@Melanoplnae) DE un. se SELE INR A
DE ANDRADE, Maria L. A new species of Platythyrea from Dominican
amber and description of a new extant species from Honduras
(Fiymenoptera:#FOrRMICIA 26) RME We) A ean oe eee eee
GATTOLLIAT, Jean-Luc. First reports of the genus Nigrobaetis Novikova &
Kluge (Ephemeroptera: Baetidae) from Madagascar and La Réunion
with observations on Afrotropical biogeography .................
TSCHUDI-REIN, Kathrin, BRAND, Nayuta, KUHRT, Ute & Dorn, Silvia. First
record of Hyssopus pallidus (Askew, 1964) for Switzerland
(Eiymenoptera#Eulophidae)r. ar A Soe TERRE eee
ALMIRON, Adriana E., CASCIOTTA, Jorge R., BECHARA, José A. & Ruiz DIAZ,
Federico J. A new species of Hyphessobrycon (Characiformes,
Characidae) from the Esteros del Iberä wetlands, Argentina........
ERRATUM (NG) HEHE 8AKOTTERAT ME 2004) eee

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GASTROPODS OF THE CHANNEL AND ATLANTIC OCEAN:
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NAEINETSIAWUEST& K.NMAREBA, 1992. MR ee en ea nee Fr.
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R. DESQUEYROUX-FAUNDEZ & S.M. STONE, 190 p., 1992...................... SE

ATLAS DE REPARTITION DES AMPHIBIENS
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THE MARINE MOLLUSKS OF THE GALAPAGOS ISLANDS:
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CATALOGUE OF THE SCAPHIDIINAE (COLEOPTERA: STAPHYLINIDAE)

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ATR AAS po 2 OO De ION ee er Fr.

REVISION DER PALAARKTISCHEN ARTEN DER GATTUNG BRACHYGLUTA
THOMSON, 1859 (COLEOPTERA, STAPHYLINIDAE) (1. Teil)
(Instrumenta Biodiversitatis VI), G. SABELLA, CH. BUCKLE, V. BRACHAT
CAEC-BESUCHETAVIA2 8 Sip 2004 RR DE LO Fr.

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and double-spaced, on A4 (210 x 297 mm) or equivalent paper and all pages should be numbered. All margins
must be at least 25 mm wide. Authors must submit three paper copies (print-outs), including tables and figures, in
final fully corrected form, and are expected to retain another copy. Original artwork should only be submitted with
the revised version of the accepted manuscript.

We encourage authors to submit the revised final text on a disk (3,5), or on a CD-R, using MS-WORD or a
similar software. The text should be in roman (standard) type face throughout, except for genus and species names
which should be formatted in italics (bold italics in taxa headings) and authors’ names in the list of references (not
in other parts of the text!), which should be formatted in SMALL CAPITALS. LARGE CAPITALS may be used for
main chapter headings and SMALL CAPITALS for subordinate headings. Footnotes and cross-references to specific
pages should be avoided. Papers should conform to the following general layout:

Title page. A concise but informative full title plus a running title of not more than 40 letters and spaces, full
name(s) and surname(s) of author(s), and full address(es) including e-mail address(es) if possible.

Abstract. The abstract is in English, composed of the title and a short text of up to 200 words. It should
summarise the contents and conclusions of the paper and name all newly described taxa. The abstract is followed
by up to 10 keywords, separated by hyphens, which are suitable for indexing. Some of the terms used in the title
may be omitted from the list of keywords in favour of significant terms not mentioned in the title.

Introduction. A short introduction to the background and the reasons for the work.

Material and methods. Sufficient experimental details must be given to enable other workers to repeat the
work.-The full binominal name should be given for all organisms. The International Code of Zoological Nomen-
clature must be strictly followed. Cite the authors of species on their first mention.

Results. These should be concise and should not include methods or discussion. Text, tables and figures should
not duplicate the same information. New taxa must be distinguished from related taxa. The abbreviations gen. n., sp.
n., syn. n. and comb. n. should be used to distinguish all new taxa, synonymies or combinations. Primary types must
be deposited in a museum or similar institution. In taxonomic papers the species heading should be followed by
synonyms, material examined and distribution, description and comments. All material examined should be listed in
similar, compact and easily intelligible format; the information should be in the same language as the text. Sex
symbols should be used rather than “male” and “female” (text file: $= 4, £= ©).

Discussion. This should not be excessive and should not repeat results nor contain new information, but
should emphasize the significance and relevance of the results reported.

References. The author-date system (name-year system) must be used for the citation of references in the text,
e.g. White & Green (1995) or (White & Green, 1995). For references with three and more authors the form Brown
et al. (1995) or (Brown et al., 1995; White er al., 1996) should be used. In the text authors’ names have to be
written in standard type face. However, in the list of references they should be formatted in SMALL CAPITALS (see
below). The list of references must include all publications cited in the text and only these. References must be
listed in alphabetical order of authors, in the case of several papers by the same author, the name has to be repeated
for each reference. The title of the paper and the name of the journal must be given in full in the following style:
PENARD, E. 1888. Recherches sur le Ceratium macroceros. Thèse, Genève, 43 pp.

PENARD, E. 1889. Etudes sur quelques Héliozoaires d’eau douce. Archives de Biologie 9: 1-61.
MERTENS, R. & WERMUTH, H. 1960. Die Amphibien und Reptilien Europas. Kramer, Frankfurt am Main, XI + 264 pp.
HANDLEY, C. O. Jr 1966. Checklist of the mammals of Panama (pp. 753-795). In: WENZEL, R. L. & Tipton, V. J.

(eds). Ectoparasites of Panama. Field Museum of Natural History, Chicago, XI + 861 pp.

Tables. These should be self-explanatory, not integrated in the text-file, with the title at the top, organised to fit
122 x 180 mm, each table on a separate sheet and numbered consecutively.

Figures. These may be line drawings or half tones, not integrated in the text-file, and all should be numbered
consecutively. Figures should be arranged in plates which can be reduced to 122 x 160 mm. Drawings and lettering
should be prepared to withstand reduction. Magnification should be indicated with scale lines. Authors should refrain
from mixing drawings and half tones. If possible, originals of figures (ink drawings, photographs, slides) should be
submitted (together with the revised version of the accepted manuscript). Original drawings will not be returned
automatically. The Revue suisse de Zoologie declines responsibility for lost or damaged slides or other documents. If
electronically scanned figures are submitted on disk or CD-R (never in MS-Powerpoint format !), this should be
clearly indicated on the print-out of these figures enclosed with the print-out of the text. The following minimum
resolutions should be respected: scanned line drawings: 600 dpi; scanned photographs: 300 dpi; digital photographs:
5 millions of pixels.

Legends to figures. These should be typed in numerical order on a separate sheet.

Proofs. Only page proofs are supplied, and authors may be charged for alterations (other than printer’s errors) if
they are numerous.

Offprints. The authors receive a total of 25 offprints free of charge; more copies may be ordered at current prices
when proofs are returned.

Correspondence. All correspondence should be addressed to

Revue suisse de Zoologie, Muséum d’histoire naturelle, CP 6434, CH-1211 Genève 6, Switzerland.
Phone: +41 22 418 63 33 - Fax: +41 22 418 63 O1. E-mail: volker.mahnert @ mhn.ville-ge.ch
Home page RSZ: http://www.ville-ge.ch/musinfo/mhng/page/rsz.htm

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