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Cossyphodini (Coleoptera: Tenebrionidae: Pimeliinae) in , Namibia and adjacent regions: New species and records, key...

Article in Zootaxa · October 2013 DOI: 10.11646/zootaxa.3721.4.3 · Source: PubMed

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The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. Zootaxa 3721 (4): 351–364 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3721.4.3 http://zoobank.org/urn:lsid:zoobank.org:pub:0A63253B-03A6-4332-903E-1A17D366C2D3 Cossyphodini (Coleoptera: Tenebrionidae: Pimeliinae) in South Africa, Namibia and adjacent regions: New species and records, key to genera, and Old World species catalogue

WOLFGANG SCHAWALLER1,2 1Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart, . E-mail: [email protected] 2Contribution to Tenebrionidae, no. 117.—For no. 116 see: Zoology in the Midle East 60, 2014

Abstract

Cossyphodini Wasmann, 1899 (Tenebrionidae: Pimeliinae) from South Africa, Namibia and adjacent regions are revised, and newly collected material is presented. All species are figured, including as much as possible type specimens of African taxa for comparison. New species are described: Cossyphodes andreaei sp. n. and Cossyphodes caecus sp. n. (both South Africa), Cossyphodes naukluftensis sp. n. (Namibia). New synonyms include: Cossyphodes freudei Basilewsky, 1957 (Cossyphodes uhligi Ferrer, 2004 syn. n.), Cossyphodes wasmanni Reichensperger, 1915 (Cossyphodes brincki Andreae, 1961 syn. n., Cossyphodes reichenspergeri Andreae, 1961 syn. n.). Lectotypes are designated for Cossyphodes arnoldi Brauns, 1925 and Cossyphodes wasmanni Reichensperger, 1915. The aedeagus is of lesser value for species separation. Keys to the four genera in South Africa and adjacent regions, as well as species of Cossyphodes in South Africa are pre- sented. A catalogue of all Cossyphodini from the Old World including full references is added.

Key words: Tenebrionidae, Pimeliinae, Cossyphodini, South Africa, Namibia, taxonomy, new species, new synonyms, key to genera, Old World catalogue, myrmecophyly

Introduction

Ant nest beetles of the tenebrionid tribe Cossyphodini Wasmann, 1899 are rarely collected with only few scattered systematic papers on Old World taxa. Basilewsky (1950a, 1950b, 1952, 1957, 1960) described new species of Cossyphodinus Wasmann, 1899 and Cossyphodes Westwood, 1851 from Central Africa, Angola and Namibia. The South African species were compiled by Andreae (1961), unfortunately some of the given names are not valid (see remarks in catalogue below). Ferrer (1990, 1996, 2006) and Ferrer & Collingwood (1993) treated records from Kenya, Zambia, and Mozambique including new species of Cossyphodes and Cossyphodinus. Robiche (2005) added a species from Benin. Schawaller (2006) first recorded the tribe from Socotra, and Schawaller et al. (2011) from Arabia and Iran. For the adjacent Indian fauna only two taxa are recorded (Waterhouse 1882, Wasmann 1899), probably both are synonymous. The structure of the aedeagus was found to be simple and quite uniform thoughout the tribe. Thus, taxa were described without considering this character. I also resisted from the risky dissections of single or rare minute type specimens, and present herein figures of the aedeagus of only two newly described species (Figs. 8, 16). Previously, the cossyphodines were classified either as their own family or as the subfamily Cossyphodinae Wasmann, 1899 of Tenebrionidae (for example Steiner 1980, Schawaller 2006). Matthews et al. (2010) published plausible arguments to consider them downranked as a tribe Cossyphodini Wasmann, 1899 of the tenebrionid subfamily Pimeliinae Latreille, 1802. The unusual features of this group are mostly adaptations to myrmecophilous habits. In the course of identification work of numerous newly collected specimens of Cossyphodini from South Africa and Namibia in the Ditsong National Museum of Natural History (the former Transvaal Museum) in Pretoria, it became necessary to re-examine and photograph African type material of as much as possible, summarise all references and catalogue Old World taxa.

Accepted by A.D. Smith: 19 Sept. 2013; published: 16 Oct. 2013 351 Depositories

BMNH The Natural History Museum, London, United Kingdom HNHM Hungarian Natural History Museum, Budapest, Hungary MNB Museum für Naturkunde, Berlin, Germany MNHN Muséum national d'Histoire naturelle, Paris, MRAC Museé Royal de l'Afrique Centrale, Tervuren, Belgium MZLU Museum of Zoology, University Lund, Sweden NMP National Museum, Department Entomology, Prague, SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany TMSA Ditsong National Museum of Natutral History, Pretoria, South Africa ZFMK Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany ZSM Zoologische Staatssammlung, München, Germany

Key to genera of Cossyphodini in South Africa, Namibia and adjacent regions

1 Number of tarsomeres 5,5,4; antenna 9-segmented with 1-segmented club (so far only Northern South Africa and Namibia)...... Cossyphodinus – Number of tarsomeres 5,4,4; antenna 11-segmented with 2-segmented club...... 2 2 Body spindle-like; keels and lateral margin of pronotum distinctly dentate...... Cossyphodites – Body broad and nearly flat; keels and lateral margin of pronotum smooth ...... 3 3 Pronotum without any keels; lateral margin of pronotum rounded, pronotum widest near base (only one species) ...... Paramellops – Pronotum at least with one, but often with several longitudinal keels; lateral margin of pronotum parallel at least in posterior half, pronotum widest in the middle (several species) ...... Cossyphodes

Species of Cossyphodini in South Africa, Namibia and adjacent regions

Cossyphodinus basilewskyi Ferrer, 1993 (in Ferrer & Collingwood 1993) (Fig. 1)

Examined type material. Zambia, Lusaka, Kafue, Kafue River, 1200 m, 22.XI.–2.XII.1987, leg. R. Mourglia, holotype ZSM. Remarks. See under Cossyphodinus robichei below. Type locality. "Kafue". Distribution. Zambia.

Cossyphodinus joannae Basilewski, 1960 (Fig. 6)

Examined type material. None. New material. Tanzania, Mt. Meru, eastern slopes, Momella, 5000 ft. (1525 m), 19.–22.III.1981, leg. H. J. Bremer, 1 ex. HNHM.—Tanzania, Mt. Kitumbeine, Olchoromyokie, 02°55'S/36°14'E, 2111 m, 9.–11.XI.2011, leg. R. Smith & H. Takano, 1 ex. BMNH. Remarks. The new records are closely situated to the type locality at Mt. Longido, all in the northern border region of Tanzania next to Kenya. Type locality. "Longido, Masai Distr." Distribution. N Tanzania.

352 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER FIGURES 1–6. Dorsal view of African Cossyphodini. 1. Cossyphodinus basilewskyi Ferrer, 1993, Zambia, holotype ZSM. 2. Cossyphodinus beccarii (Gestro, 1872), Ethiopia, non-type HNHM. 3. Cossyphodinus bremeri Ferrer, 1993, Kenya, paratype TMSA. 4. Cossyphodinus bukavuensis Basilewsky, 1950, Congo, paratype ZFMK. 5. Cossyphodinus goergeni Robiche, 2005, Benin, holotype MNHN. 6. Cossyphodinus joannae Basilewski, 1960, Tanzania/Mt. Meru, non-type HNHM. Scale: 1 mm.

Cossyphodinus robichei Ferrer, 2006

Examined type material. None. Remarks. The drawing of the dorsal view by Ferrer (2006) is quite similar to the drawing of Cossyphodinus basilewskyi from Zambia by Ferrer (in Ferrer & Collingwood 1993), probably both are conspecific. Type locality. "Nampula". Distribution. Northern Mozambique.

COSSYPHODINI (COLEOPTERA: TENEBRIONIDAE) IN AFRICA Zootaxa 3721 (4) © 2013 Magnolia Press · 353 Cossyphodites woodroofei (Péringuey, 1885) (Fig. 26)

Cossyphodes woodroofei Péringuey, 1885

Examined type material. None. New material. South Africa, Willowmore, 25.III. & 1.IV.1902, leg. H. Brauns, 2 ex. ZFMK.—South Africa, Willowmore, 30.VII.1900, leg. H. Brauns, 1 ex. TMSA.—South Africa, Willowmore, without date, leg. H. Brauns, 6 ex. HNHM (det. Basilewsky).—South Africa, Willowmore, without date, leg. H. Brauns, 2 ex. BMNH. Type locality. “Somerset East”. Distribution. South Africa (Eastern Cape).

Paramellops bewicki (Wollaston, 1861) (Fig. 25)

Cossyphodes bewicki Wollaston, 1861 (not Wasmann 1899) Cossyphodes voigti Reichensperger, 1926 syn.

Examined type material. South Africa, Cape of Good Hope, holotype of bewicki BMNH.—South Africa, Western Cape, Muizenberg (misspelled as Muigenberg), 14.VI.1925, 2 syntypes of voigti ZFMK. New material. South Africa, Western Cape, Matjiesfontein, Witteberge, 33°16'S/20°30'E, 1600 m, 25.X.1978, leg. S. Endrödy-Younga, 15 ex. TMSA, 4 ex. SMNS.—South Africa, Western Cape, 15 km N Bredasdorp, 34°24'S/ 20°02'E, 20.IX.1985, leg. S. Endrödy-Younga, 1 ex. TMSA. Type localities. "Cape of Good Hope, Atlantic side" (bewicki); "Cape, Muizenberg" (voigti) (misspelled as Muigenberg by Basilewsky 1952). Distribution. South Africa (Western Cape).

Cossyphodes andreaei sp. n. (Figs. 7–8)

Type material. Holotype (♂): South Africa, "Transvaal", Waterberg, Lapalala Wilderness, 23°49'S/20°17'E, 16.VIII.1975, leg. S. Endrödy-Younga, TMSA.—Paratypes: Same data as holotype, 1 ex. TMSA, 1 ex. SMNS.— South Africa, "Transvaal", Pretoria, Waterkloof, 25°43'S/28°11'E, 9.IX.1978, leg. S. Endrödy-Younga, 2 ex. TMSA.—South Africa, "Transvaal", Pretoria, VII.1953, leg. A. v. Peez, 2 ex. ZSM. Description. Body dark brown without colour pattern, eyes darker. Body length 2.5–2.6 mm. Head semicircular, clypeus not separated, clypeal lines indistinct, straight; anterior and lateral margins somewhat bent upwards, frons without impression or tubercles; head surface with regular microgranulation; eyes sickle-shaped, composed by two rows of large, dark ocelli, each row with about five ocelli; antennae 11-segmented with two large apical antennomeres forming club, as usual for the genus. Pronotum 2.2 times broader than median length, posterior corner rectangular, acute, anterior corner rounded; all margins unbordered; surface with same microgranulation and microsetation as on head, disc on each side with three fine longitudinal keels, internal keel sometimes shortened anteriorly, additionally with traces of a medial keel, for direction, distances of these keels, see Fig. 7; prosternal process elongate triangular with protruding acute posterior angle. Elytra widest shortly before base, 1.2 times longer than broad; elytra with four primary keels and without any secondary keels, internal main keel present only in anterior quarter of elytra, for direction, distances of these keels, see Fig. 7; surface with the same microgranulation and microsetation as on head and pronotum. Legs without peculiarities, tarsal formula 5-4- 4 as usual for the genus. Aedeagus with trianguar acute apicale (Fig. 8). Diagnosis. C. andreaei n.sp. can be recognised by lacking secondary elytral keels in combination with a rudimentary inner main keel present only in the anterior quarter of elytra (Fig. 7), and by eyes with two rows of ocelli. C. machadoi Basilewsky, 1952, from Angola is similar, but the inner elytral keel is complete and the eyes narrower with only one row of ocelli.

354 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER Etymology. Named in honour of Hans Andreae jun. (Cape Town), who recognised this species already as new and named it "pretoriensis", but designated no type material and gave no description for a valid name according to the nomenclatorical rules (see remarks in catalogue below).

FIGURES 7–13. Dorsal view of African Cossyphodini. 7–8. Cossyphodes andreaei sp. n., RSA/Waterberg, paratype SMNS and aedeagus. 9. Cossyphodes arnoldi Brauns, 1925, Zimbabwe, lectotype TMSA. 10. Cossyphodes naukluftensis sp. n., Namibia/Naukluft, holotype TMSA. 11. Cossyphodes angolanus Basilewsky, 1952, holotype MRUC. 12. Cossyphodes wasmanni Reichensperger, 1915 (Cossyphodes brincki Andreae, 1961 syn. n., RSA/Hluhluwe, holotype MZLU). 13. Cossyphodes wasmanni Reichensperger, 1915, Zimbabwe, lectotype TMSA. Scale: 1 mm, for aedeagus: 0.5 mm.

COSSYPHODINI (COLEOPTERA: TENEBRIONIDAE) IN AFRICA Zootaxa 3721 (4) © 2013 Magnolia Press · 355 Cossyphodes angolanus Basilewsky, 1952 (Fig. 11)

Examined type material. Angola, Serra de Moco, Luimbale, 12°30'S/15°10'E, 2000 m, IX.1949, leg. A. de Barros Machado, holotype MRUC. Type locality. "Luimbale". Distribution. Angola.

Cossyphodes arnoldi Brauns, 1925 (in Wasmann & Brauns 1925) (Fig. 9)

Examined type material. Zimbabwe (labelled as S Rhodesia), Bembesi, 2.I.1913, leg. G. Arnold, 2 syntypes TMSA, 1 specimen designated herewith as lectotype. Lectotype designation. The description was based on an unspecified number of syntypes. Therefore, a lectotype was designated in order to fix a single name-bearing type. Remarks. See also under Cossyphodes naukluftensis sp. n. from Namibia below. Type locality. “Bembesi”. Distribution. Zimbabwe.

Cossyphodes braunsi Andreae, 1961 (Fig. 14)

Cossyphodes bewicki Wasmann, 1899 (not Wollaston, 1861)

Examined type material. None. New material. South Africa, Eastern Cape, Port Elizabeth, Algoa Bay, leg. H. Brauns, 2 ex. HNHM (det. Basilewsky as C. bewicki), 1 ex. BMNH (det. as C. bewicki).—South Africa (labelled as Kapland), Port Elizabeth, leg. H. Brauns, 2 ex. ZFMK (coll. Reichensperger). Type locality. “Port Elisabeth”. Distribution. South Africa (Eastern Cape).

Cossyphodes caecus sp. n. (Figs. 15–16)

Type material. Holotype (♂): South Africa, Northern Cape, Richtersveld NP, Doringpoort, 28°34'S/16°56'E, 7.IX.1976, leg. S. Endrödy-Younga, TMSA.—Paratypes: Same data as holotype, 1 ex. TMSA, 1 ex. SMNS.— Western Cape, Elands Bay, XI.1948, leg. C. Koch, 3 ex. TMSA, 1 ex. SMNS.—South Africa, Mpumalanga (labelled as N Transvaal), Waterberg, Haakdoringboom Farm, 24°11'S/27°50'E, 13.II.1976, leg. A. Strydom, 1 ex. TMSA. Description. Body dark brown without colour pattern. Body length 2.6–2.8 mm. Head semicircular, clypeus not separated, clypeal lines indistinct, straight; anterior and lateral margins somewhat bent upwards, frons with two pairs of weak keels, internal keels reduced to two tubercles each; head surface with regular microgranulation; eyes completely absent, a short longitudinal keel instead of ocelli; antennae 11-segmented with two large apical antennomeres forming club. Pronotum 1.7 times broader than median length, posterior corner rectangular, acute, anterior corner rounded; all margins unbordered; surface with same microgranulation and microsetation as on head, disc on each side with three distinct, complete longitudinal keels, additionally with a distinct medial keel, for direction, distances of these keels, see Fig. 15; prosternal process elongate triangular with protruding acute posterior angle. Elytra widest shortly before base, 1.4 times longer than broad; elytra with four primary keels and distinct secondary keels, internal main keel present only on anterior third of elytra, for direction, distances of these keels, see Fig. 15; surface with the same microgranulation and microsetation as on head and pronotum. Legs without peculiarities, tarsal formula 5-4-4 as usual for the genus. Aedeagus with trianguar acute apicale (Fig. 16).

356 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER Diagnosis. Cossyphodes caecus sp. n. can be recognised by the lack of eyes without any ocelli present, by two pairs of weak keels on the head, by distinct complete keels on the pronotum, and by distinct main keels as well as present secondary keels on the elytra. Cossyphodes braunsi from Eastern Cape (Fig. 14) also lacks ocelli, but the head is without any keels or tubercles, the pronotum with only weak external keels, and the elytra with extinct inner keels. Etymology. The name refers to the lack of eyes, caecus in Latin means blind.

Cossyphodes freudei Basilewsky, 1957 (Figs. 18–19)

Cossyphodes uhligi Ferrer, 2004 syn. n.

Examined type material. Namibia (labelled as S. W. Afr.), Otjiwarongo Distr., Abachaus (Farm), 4.I.1956, leg. G. Hohbohm, holotype and 4 paratypes of freudei ZSM.—Namibia, Kavango, Buffalo Camp, 18°09'S/21°42'E, 28.II.1992, leg. M. Uhlig, holotype of uhligi MNB. New material. Namibia (labelled as Hereroland), Otjikonjo Farm, 21°09'S/17°55'E, 24.VI.1978, leg. S. Endrödy-Younga, 1 ex. TMSA, 1 ex. SMNS.—Namibia, Kaokoveld, Ohopoho, 18°04'S/13°55'E, 400 m, 6.II.1975, leg. S. Endrödy-Younga, 10 ex. TMSA, 4 ex. SMNS.—Namibia, Kaokoveld, 38–44 km NW Ohopoho, 17°47'S/ 13°44'E, 13.II.1975, leg. S. Endrödy-Younga, 4 ex. TMSA.—Namibia, Kaokoveld, Sesfontein, 19°08'S/13°36'E, 3.II.1975, leg. S. Endrödy-Younga, 1 ex. TMSA.—C Namibia, Ganab, NE Hillgap, 23°08'S/15°35'E, 7.VII.1978, leg. S. Endrödy-Younga, 8 ex. TMSA, 3 ex. SMNS.—C Namibia, Ganab Water, 23°06'S/15°32'E, 28.V.1975, leg. S. Endrödy-Younga, 1 ex. TMSA.—C Namibia, Arechadamaberg, 23°00'S/15°37'E, 28.V.1975, leg. S. Endrödy- Younga, 1 ex. TMSA.—C Namibia, Us Pass (= Us-hoogte Pass), 23°03'S/15°40'E, 2.III.1975, leg. S. Endrödy- Younga, 1 ex. TMSA. Synonymy. The strength of the pronotal and elytral keels is slightly variable, but not distinctly different between the types of C. freudei and C. uhligi from the same area in northern Namibia, all other external characters agree. Thus, Cossyphodes uhligi is considered as a junior synonym. Remarks. Most of the above listed specimens from Namibia have the pronotum with a weak medial keel, in some specimens this keel is reduced and only present in the anterior part of the pronotum. In the examined type specimens this medial keel is completely missing. All other external characters coincide. Additionally, some specimens have the head without any keels (as the types), other specimens have the head with a weak external keel (but not with a pair of distinct keels). Thus, the presence or absence of a weak keel on the head, as well as of a pronotal medial keels are considered just as infraspecific variations. "C. matabele", mentioned but not described by Andreae (1961), also fits into this variation. Quite similar is also C. machadoi Basilewsky, 1952 from adjacent Angola, but in this species the eyes are reduced and the head is with a pair of distinct keels. Type localities. "Abachaus, Otjiwarongo" (freudei), "Kavango" (uhligi). Distribution. Namibia.

Cossyphodes machadoi Basilewsky, 1952 (Fig. 16)

Examined type material. Angola, Boa Entrada, Gabela, 10°51'S/14°19'E, 1100 m, XI.1949, leg. A. de Barros Machado, holotype MRUC. Type locality. "Gabela". Distribution. Angola.

COSSYPHODINI (COLEOPTERA: TENEBRIONIDAE) IN AFRICA Zootaxa 3721 (4) © 2013 Magnolia Press · 357 FIGURES 14–20. Dorsal view of African Cossyphodini. 14. Cossyphodes braunsi Andreae, 1961, RSA/Port Elizabeth, non- type HNHM. 15–16. Cossyphodes caecus sp. n., RSA/Richtersveld, paratype SMNS and aedeagus. 17. Cossyphodes kundelunguensis Basilewsky, 1950, Congo, paratype ZSM. 18. Cossyphodes freudei Basilewsky, 1957, Namibia, holotype ZSM. 19. Cossyphodes freudei Basilewsky, 1957 (Cossyphodes uhligi Ferrer, 2004 syn. n., Namibia, holotype MNB). 16. Cossyphodes machadoi Basilewsky, 1952, holotype MRUC. Scale: 1 mm.

358 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER Cossyphodes mourgliai Ferrer, 1993 (in Ferrer & Collingwood 1993) (Fig. 22)

Examined type material. Zambia, Lusaka, Kafue, Kafue River, 1200 m, 22.XI.–2.XII.1987, leg. R. Mourglia, holotype ZSM (labelled as Cossyphodites). Type locality. "Kafue". Distribution. Zambia.

Cossyphodes naukluftensis sp. n. (Fig. 10)

Type material. Holotype (sex unknown): Namibia (labelled as S. W. Afr.), Naukluft Park, 24°16'S/16°15'E, 26.X.1974, leg. S. Endrödy-Younga, TMSA. Description. Body reddish brown without colour pattern. Body length 2.3 mm. Head semicircular, clypeus not separated, clypeal lines indistinct, straight; anterior and lateral margins somewhat bent upwards, frons with a pair of distinct, complete external keels and a pair of a weak internal keels, anterior of internal keel with tubercle; head surface with regular microgranulation; eyes narrow sickle-shaped, composed by two rows of dark ocelli, each row with about five ocelli; antennae 11-segmented with two large apical antennomeres forming club. Pronotum 1.5 times broader than median length, posterior corner rectangular, acute, anterior corner rounded; all margins unbordered; surface with same microgranulation and microsetation as on head, disc on each side with three complete longitudinal keels, additionally with distinct medial keel, this medial keel shortly interrupted in anterior part, for direction, distances of these keels, see Fig. 10; prosternal process elongate triangular. Elytra widest shortly before base, 1.4 times longer than broad; elytra with four primary and secondary keels, internal main keel only present near base of elytra, all main keels convergent from base towards tip, for direction, distances of these keels, see Fig. 10; surface with the same microgranulation and microsetation as on head and pronotum. Legs without peculiarities, tarsal formula 5-4-4. Aedeagus unknown, sex not examined (see introduction). Diagnosis. C. naukluftensis sp. n. is similar to C. arnoldi Brauns, 1925, known from Zimbabwe (Fig. 9). Both share the narrow body shape, the head with a pair of distinct keels and with a tubercle before the internal keel, and the distinct pronotal keels with interrupted medial keel. Both species can be separated by a completely different pattern of main elytral keels (Figs. 9–10). In C. naukluftensis sp. n. the elytral keels are bent outwards near base and convergent from base towards tip, in C. arnoldi the main keels are parallel in anterior half of elytra and convergent only in posterior half; in C. naukluftensis sp. n. the internal main keel is present only near base, in C. arnoldi this keel is complete. Etymology. Named after the Naukluft Mountains, where the holotype was collected.

Cossyphodes raffrayi Gestro, 1874 (Fig. 21)

Examined type material. None. New material. Ethiopia (labelled as Abyssin.), 3 ex. ZFMK (coll. Reichensperger).—Yemen (labelled as W Aden Prot.), Dhala, 4800 ft. (1460 m), 14.IX.1937, leg. H. Scott & B. M. Britton, 2 ex. BMNH (det. Basilewsky).—Tanzania (labelled as Africa or.), Katona, Shirati, 1909, 5 ex. HNHM (det. Basilewsky).—Tanzania, Arusha, 1350 m, 8.III.1960, leg. Szunyoghy, 2 ex. HNHM (det. Basilewsky). Remarks. Obviously, this species reaches also the Arabian Peninsula in Yemen. Unfortunately, this record was unknown when treating the Arabian fauna (Schawaller et al. 2011). Type locality. "Abyssinie". Distribution. Ethiopia (Reichensperger 1913); Yemen, Tanzania (new records).

COSSYPHODINI (COLEOPTERA: TENEBRIONIDAE) IN AFRICA Zootaxa 3721 (4) © 2013 Magnolia Press · 359 FIGURES 21–26. Dorsal view of African Cossyphodini. 21. Cossyphodes raffrayi Gestro, 1874, Tanzania, non-type HNHM. 22. Cossyphodes mourgliai Ferrer, 1993, Zambia, holotype ZSM. 23. Cossyphodes uviranus Basilewsky, 1951, Congo, non- type ZFMK. 24. Cossyphodes vandami Andreae, 1961, RSA/Pretoria, paratype MZLU. 25. Paramellops bewicki (Wollaston, 1861), RSA/Matjiesfontein, non-type TMSA. 26. Cossyphodites woodroofei (Péringuey, 1885), South Africa, non-type HNHM. Scale: 1 mm.

360 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER Cossyphodes vandami Andreae, 1961 (Fig. 24)

Examined type material. South Africa, Gauteng (labelled as Transvaal), 16 miles NE Pretoria, X.1954, leg. G. Rudebeck, 3 paratypes MZLU. New material. South Africa, Mpumalanga (labelled as E Transvaal), Waterval River Pass, 24°54'S/30°21'E, 6.XI.1980, leg. S. Endrödy-Younga, 6 ex. TMSA, 1 ex. SMNS.—South Africa, Limpopo, Krüger NP, camp 1 km E Pafurires, 22°25'S/31°13'E, 31.I.1994, leg. S. Endrödy-Younga, 1 ex. TMSA, 1 ex. SMNS. Remarks. The examined paratypes of C. vandami have the elytra somewhat more narrowed towards tip as specimens of C. wasmanni, and also somewhat higher elytral keels, but all other external characters agree. At present, these differences are considered as specific. Type locality. “Pretoria, Zoohill”. Distribution. South Africa (Gauteng, Mpumalanga, Limpopo).

Cossyphodes wasmanni Reichensperger, 1915 (Figs. 12–13)

Cossyphodes wasmanni Brauns in litteris Cossyphodes brincki Andreae, 1961 syn. n. Cossyphodes reichenspergeri Andreae, 1961 syn. n.

Examined type material. Zimbabwe (labelled as S Rhodesia), Bulaweyo, 20.VII.1913, leg. G. Arnold, 1 syntype TMSA, labelled as type of wasmanni by Brauns, and labelled probably not by the author also as paratype of reichenspergeri TMSA, designated herewith as lectotype of wasmanni Reichensperger.—Zimbabwe (labelled as S Rhodesia), Bulaweyo, no date, leg. G. Arnold, 2 syntypes ZFMK (coll. Reichensperger).—South Africa, KwaZulu/ Natal, Hluhluwe Game Reserve, 17.IV.1951, leg. B. Brinck & G. Rudebeck, holotype of brincki MZLU. New material. Zambia (labelled as N Rhodesia, Namwala, III.1913, leg. H. C. Dollman, 4 ex. BMNH (det. Basilewsky).—Zambia (labelled as NW Rhodesia, Chinenga, 25.X.1913, leg. H. C. Dollman, 1 ex. BMNH.— Botswana, Okawango, Maxwee Lagoon, 28.XI.1975, leg. Russell-Smith, 1 ex. TMSA.—Botswana, near Francistown, Selkirk Mine, 21°32'S/27°77'E, 1.–28.III.2008, leg. D. H. Jacobs, 1 ex. SMNS.—Namibia (labelled as Hereroland), Dai Papel Farm, 20°55'S/17°39'E, 24.VI.1978, leg. S. Endrödy-Younga, 1 ex. TMSA, 1 ex. SMNS.—South Africa, Northern Cape, Tswalu NR, 27°16'S/22°23'E, 12.II.2005, leg. R. Müller, 1 ex. TMSA.— South Africa, Limpopo, N Soutpansberg, 22°54'S/29°41'E, leg. S. Endrödy-Younga, 4 ex. TMSA, 2 ex. SMNS.— South Africa, Gauteng, Cullinan, Windy Brow, 25°41'S/28°30'E, 8.I.2002, leg TMSA staff, 1 ex. TMSA.—South Africa, KwaZulu-Natal (labelled as Zululand), 1905, leg. Traegårdt, 2 ex. ZFMK.—South Africa, KwaZulu-Natal, Pietermaritzburg, no further data, leg. Schwarze, 1 ex. NMP. Synonymy. Reichensperger (1915) described C. wasmanni based on a series of syntypes, which he received from H. Brauns already with the in litteris name C. wasmanni, but did not fix a holotype. Therefore, a lectotype was designated in order to fix a single name-bearing type. Later, Andreae (1961) described C. reichenspergeri on the basis of the same series and separated both in his key by only minute differences of the proportions of head and pronotum. These differences are just infraspecific variations, consequently C. reichenspergeri is considered a junior synonym of C. wasmanni. The holotype of C. brincki shows no differences to the lectotype of C. wasmanni, thus C. brincki is considered as a further synonym. Reichensperger (1915) mentioned C. wasmanni also from Umfolozis (= Hluhluwe), the type locality of C. brincki. These syntopical records are a further argument for synonymy. Remarks. See under C. vandami. Type localities. “Bulaweyo” (wasmanni, reichenspergeri), “Hluhluwe Game Reserve” (brincki). Distribution. Zimbabwe (type locality), South Africa (KwaZulu-Natal) (Reichensperger 1915); South Africa (Limpopo, Northern Cape, Western Cape, Gauteng), Zambia, Botswana, Namibia (new records).

COSSYPHODINI (COLEOPTERA: TENEBRIONIDAE) IN AFRICA Zootaxa 3721 (4) © 2013 Magnolia Press · 361 Cossyphodes wollastoni Westwood, 1851

Examined type material. None. New material. Capverdes, S. Jorge, 28.III.1981, leg. T. van Harten, 1 ex. SMNS.—Capverdes, no further data, coll. Reichensperger, 1 ex. ZFMK.—St. Helena Island, Peak Dale, XII.2005–I.2006, leg. N. P. Ashmole et al., 3 ex. BMNH (det. Barclay & Mendel). Type locality. "Insula Maderae". Distribution. Madeira, Canary Islands, Capverdes, St. Helena Island.

Key to the species of Cossyphodes in South Africa

1 Eyes completely absent, without ocelli...... 2 – Eyes present, sickle-shaped, composed by two rows of dark ocelli ...... 3 2 Head without keels or tubercles, pronotum and elytra with extinct inner keels and with feeble external keels...... C. braunsi – Head with weak keels and tubercles, pronotum with distinct keels and additionally with distinct medial keel, elytra with dis- tinct keels and secondary keels ...... C. caecus sp. n. 3 Elytra without secondary keels and extinct inner main keel present only in the anterior quarter of elytra, pronotum with only traces of a medial keel...... C. andreaei sp. n. – Elytra with distinct secondary keels, pronotum with distinct medial keel ...... 4 4 Elytra more narrowed towards tip, dorsal keels high (Fig. 24)...... C. vandami – Elytra broader towards tip, dorsal keels low (Figs. 12–13) ...... C. wasmanni

Catalogue of Cossyphodini of the Old World

Remarks. In the revision of the South African Cossyphodini Andreae (1961) described some new species according to the rules of zoological nomenclature, but also mentioned and named additional “new species” without any descriptions or designations of type specimens (“Cossyphodes fulvus” from Ethiopia, “Cossyphodes godfreyi” from South Africa/Kingwilliamstown, “Cossyphodes matabele” from Zimbabwe/Bulawayo, “Cossyphodes natalensis” from South Africa/Pietermaritzburg, “Cossyphodes pretoriensis” from South Africa/Pretoria, and “Cossyphodes stevensoni” from Zimbabwe/Bulaweyo). These names are not valid and are not included in this catalogue. Mimocossyphus minor Pic, 1923 from Egypt is a doubtful taxon and also not included; most probably it does not belong to Cossyphodini at all.

Cossyphodes Westwood, 1851 Cossyphodes andreaei sp. n. (Fig. 7) South Africa Cossyphodes angolanus Basilewsky, 1952 (Fig. 11) Angola Cossyphodes arnoldi Brauns, 1925 (Fig. 9) Zimbabwe Cossyphodes asiricus Schawaller, 2011 Saudi Arabia (Asir Mts.) Cossyphodes braunsi Andreae, 1961 (Fig. 14) South Africa Cossyphodes bewicki Wasmann, 1899 (not Wollaston, 1861) Cossyphodes caecus sp. n. (Fig. 15) South Africa Cossyphodes freudei Basilewsky, 1957 (Figs. 18–19) Namibia Cossyphodes uhligi Ferrer, 2004 syn. n. Cossyphodes kundelunguensis Basilewsky, 1950 (Fig. 17) Congo Cossyphodes machadoi Basilewsky, 1952 (Fig. 16) Angola Cossyphodes mourgliai Ferrer, 1993 (Fig. 22) Zambia Cossyphodes naukluftensis sp. n. (Fig. 10) Namibia Cossyphodes raffrayi Gestro, 1874 (Fig. 21) Ethiopia, Yemen, Tanzania Cossyphodes uviranus Basilewsky, 1951 (Fig. 23) Congo Cossyphodes vandami Andreae, 1961 (Fig. 24) South Africa

362 · Zootaxa 3721 (4) © 2013 Magnolia Press SCHAWALLER Cossyphodes wasmanni Reichensperger, 1915 (Figs. 12–13) Zimbabwe, Botswana, Namibia, Cossyphodes brincki Andreae, 1961 syn. n. South Africa Cossyphodes reichenspergeri Andreae, 1961 syn. n. Cossyphodes wollastoni Westwood, 1851 Madeira, Canary Islands,Capverdes, St. Helena Island

Cossyphodinus Wasmann, 1899 Cossyphodinus basilewskyi Ferrer, 1993 (Fig. 1) Zambia Cossyphodinus beccarii (Gestro, 1872) (Fig. 2) Ethiopia Cossyphodes beccarii Gestro, 1872 Cossyphodinus bremeri Ferrer, 1993 (Fig. 3) Kenya Cossyphodinus bukavuensis Basilewsky, 1950 (Fig. 4) Congo Cossyphodinus decimcarinatus (Ferrer, 1990) Kenya Cossyphodes decimcarinatus Ferrer, 1990 Cossyphodinus goergeni Robiche, 2005 (Fig. 5) Benin Cossyphodinus indicus Wasmann, 1899 India Cossyphodinus joannae Basilewski, 1960 (Fig. 6) Tanzania Cossyphodinus katanganus Basilewsky, 1950 Congo Cossyphodinus leleupi Basilewsky, 1950 Congo Cossyphodinus makuguensis Basilewsky, 1952 Congo Cossyphodinus robichei Ferrer, 2006 Mozambique

Cossyphodites Brauns, 1901 Cossyphodites woodroofei (Péringuey, 1885) (Fig. 26) South Africa Cossyphodes woodroofei Péringuey, 1885

Paramellon Waterhouse, 1882 Paramellon sociale Waterhouse, 1882 NW India, Iran, Arab Emirates Paramellon socotraense Schawaller, 2006 Socotra Island

Paramellops Andreae, 1961 Paramellops bewicki (Wollaston, 1861) (Fig. 25) South Africa Cossyphodes bewicki Wollaston, 1861 (not Wasmann, 1899) Cossyphodes voigti Reichensperger, 1926 syn.

Acknowledgements

For the trustful loan of specimens from the collections under their care, or for help with loans I thank Dirk Ahrens (Bonn), Michael Balke (München), Max Barclay (London), Roy Danielsson (Lund), Jiři Hájek (Prague), Bernd Jäger and Manfred Uhlig (Berlin), Antoine Mantilleri (Paris), Ottó Merkl (Budapest), Marc de Meyer (Tervuren), Luboš Purchart (Brno) and Ruth Müller (Pretoria). The photographs were taken by Johannes Reibnitz (Stuttgart) with a Leica DFC320 digital camera on a Leica MZ16 APO microscope and subsequently processed by him with Auto-Montage (Syncroscopy) software. Thanks are due also to two referees for their corrections and suggestions.

References

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