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The Generic and Infrageneric Taxa Described by Ethelwynn Trewavas

The Generic and Infrageneric Taxa Described by Ethelwynn Trewavas

Environmental Biology o f 41: 55-61,1994. © 1994 Kluwer Academic Publishers. Printed in the Netherlands.

The generic and infrageneric taxa described by Ethelwynn Trewavas

P. Humphry Greenwood J.L.B. Smith Institute o f , Private Bag 1015, Grahamstown, 6140 South Africa

Either as the sole author, or in collaboration (especially with C. Tate Regan), Ethelwynn Trewavas was re­ sponsible for describing and naming at least 462 genera, subgenera, species and subspecies. Doubtless, in compiling this index (which, intentionally does not include nomina nova she established) I have overlooked Trewavas taxa; for those oversights I can only apologise.

In many respects, as she herself often remarked, aspects of systematics, and read extensively in the Ethelwynn entered the field of taxonomic ichthyol­ literature of evolutionary biology. Those wide- ogy at a fortunate time. It was a time when large ranging interests were very obvious throughout my collections of fishes were arriving in the British Mu­ long association with her, particularly so during the seum (Natural History), many from the still poorly time when, in the early 1960’s, Donn Rosen and I sampled Great Lakes of Africa, others from the were roughing-out our ideas of a phylogenetic clas­ then little explored deeper oceanic waters. Above sification for teleosts (Greenwood et al. 1966) and all, she had the great advantage of having Tate Re­ often had long and beneficial discussions with her. gan as her guide and mentor. He too was fortunate Her few publications in the field of higher-level in having Ethelwynn as his student and assistant, are surprising, too, when one recalls that and both were lucky to be working at a period when her major doctoral research (on the hyoid and la­ financially and philosophically the Museums of the rynx of Anura) was in large part stimulated by No­ world were able to concentrate their activities ble’s work on the phylogeny of the group, a study mainly on research. The times of penury and expen­ which had paid scant attention to that important as­ sive theme park-like exhibitions were yet to come. pect of the ’ anatomy. Then, too, her early Ethelwynn was not only a tireless worker, but a association with Tate Regan was at a time when he gifted taxonomist, one not merely of the ‘stamp-col­ was preparing his major synthetic work on the clas­ lecting’ variety, but a true biologist who recognised sification of teleosts (later published in the 14th edi­ the corpses she handled as once living entities and tion of the Encyclopaedia Britannica, 1929), one, to­ essential elements of the biotopes from whence gether with his earlier papers, long accepted as the they were collected. I have not been able to carry definitive work in that field. out a detailed analysis to determine the current sta­ Perhaps her reticence in this sphere of systematic tus of all the taxa in whose recognition and descrip­ ichthyology stemmed from her high regard for Tate tion she was involved; however, for the African Ci- Regan’s work coupled with the fascination that chlidae (and non-) my estimate would be African cichlids, especially the tilapiines, held for that more than 80% are still regarded as valid - and her, and her strong views on the pragmatic impor­ that is despite the activities of the many taxonomists tance to fishery biologists of sound alpha-level tax­ now studying the freshwater fishes of the continent. onomy. Interestingly, and somewhat surprisingly, little of Be that as it may, all those who were to be in­ Ethelwynn’s work was concerned with higher-level volved later either in taxonomy or working systematics, especially the phyletic inter- and intra­ with those fishes in applied research, are deeply in­ relationships of the groups she studied. This is sur­ debted to Ethelwynn, as are the many people who prising because she certainly was interested in those benefited from her tutelage and knowledge. 56

The list of taxa which follows has been arranged, neatus; mocoensis firstly in higher taxonomic categories, and then 1938 (17) Barbus wohlerti within those groups, by families. Within a family the 1941 (23) Barbus arabicus members of each taxonomic rank (i.e. , sub­ 1941 (23) brittoni; tibanica genus, species or subspecies) are listed in chrono­ 1955 (50) Typhlogarra widdowstoni logical (i.e. annual) order of their publication. The 1974 (94) Labeo camerunensis year of publication is followed, in parentheses, by the number allocated to that particular publication Chauliodontidae in Noakes (1994). The type species (abbreviated as Sp. nov. 1939 (4) Chauliodus danae t.s.) and its author is given for each genus and sub­ Astronesthidae genus. Gen. nov. 1929 (4) Diplolynchus (t.s. D. lucifer The names of the taxa are as originally published; no attempt has been made to indicate the present Regan & Trewavas, 1929) status of a taxon or changes in its nomenclature (i.e. Heterophotus (t.s. H. ophistoma Re­ synonymy, allocation to other genera, etc.). Esch- gan & Trewavas, 1929) meyer (1990) should be consulted for details about Rhadinesthes (t.s. Astronesthes deci- the nomenclatural status of all generic names in the mus Zugmayer, 1911) list and for the current familial placement of the Neonesthes (t.s. N. macrolynchnus genera. Regan & Trewavas, 1929) Spp. nov. Astronesthes caulophorus; cyclopho- OSTARIOPHY SI tus; filifer; leucopogon; longiceps; Series Anotophysi luetkeni; neopogon; oculatus Kneriidae Borostomias macristius; macro- Sp. nov. 1936 (15) Kneria polli phthalmus; panamaensis; schmidti Series Otophysi Diplolynchus bifllis; lucifer; monone- Clariidae ma Sp. nov. 1936 (15) Clarias cavernicola Heterophotus ophistoma Mochokidae Neonesthes macrolynchnus Sp. nov. 1974 (94) disneyi Melanostomiidae Gen. nov. 1930 (5) Chirostomias (t.s. C. pliopte- Gen. nov. 1943 (28) Irvineia (t.s. I. voltae Tre- rus Regan & Trewavas, 1930) wavas, 1943) Haplostomias (t.s. H. tentaculatus Spp. nov. 1943 (28) Irvineia voltae Regan & Trewavas, 1930) 1964 (70) Irvineia orientalis Thysanactis (t.s. T. dentex Regan & Amphiliidae Trewavas, 1930) Sp. nov. 1936 (15) Amphilius lentiginosus Trigonolampa (t.s. Trig, miriceps Re­ Characidae gan & Trewavas, 1930) Sub-gen.nov. 1949 (39) Oligosarcus (Paroligo- Sub-gen. nov. Bathophilus (Notopodichthys), sarcus), t.s. O. pintoi Amaral-Campos, 1945 t.s. B. brevis Regan & Trewavas, Sp. nov. 1936 (15) Micralestes argyrotaenia 1930 Cyprinidae Bathophilus (Trichochirus), (t.s. B. Gen. nov. 1955 (50) Typhlogarra (t.s. T. wid- paweeni Regan & Trewavas, 1930) dowstoni Trewavas, 1955) Eustomias (Achirostomias), t.s. E. li- Spp. nov. 1973 (88) Acanthobrama telavivensis pochirus Regan & Trewavas, 1930 1936 (15) Barbus breviceps; dossoli- 57

Eustomias (Dinematochirus), t.s. E. Pachystomias atlanticus fissibarbis Pappenheim, 1914 Photonectes achirus; caerulescens; di­ Eustomias (Haploclonus), t.s. E. em- nema; fimbria; leucospilus; mono- barbatus Regan & Trewavas, 1930 dactylus; ovibarba; parvimanus; Eustomias (Nominostomias), t.s. E. phyllopogon bibulbosus Parr, 1927 Idiacanthidae Eustomias (Spilostomias), t.s. E. Idiacanthus panamensis braueri Zugmayer, 1911 Malacosteidae Eustomias (Triclonostomias), t.s. E. Spp. nov. Aristostomias lumifer; polydactylus; drechseli Regan & Trewavas, 1930 uncodentatus; xenostoma Eustomias (Urostomias), t.s. E. ma- Malacosteus danae crurus Regan & Trewavas, 1930 LOPffllFORMES Photonectes (Melanonectes), t.s. P. For current family placements, see Eschmeyer dinema Regan & Trewavas, 1930 (1990). Photonectes (Microchirichthys), t.s. Gen. nov. 1932 (8) Amacrodon (t.s. Thaumat- f! parvimanus Regan & Trewavas, ichthys binghami Parr, 1927) 1930 Anomalophryne (t.s. Haplophryne Spp. nov. Bathophilus brevis; chironema; hudsonius Beebe 1929) longipes; melas; paweeni; proxi- Caranactis (t.s. Caranactis pumilus mus; schizochirus Regan & Trewavas, 1932) Echiostoma guentheri Centrocetus (t.s. Centrocetus spinulo- Eustomias acinosus; barbura; bimar- sus Regan & Trewavas, 1932) ginatus; bituberatus, botrypogon, Centrophryne (t.s. Centrophryne spi- brevifilis; dactylobolus; dendriti- nulosa Regan & Trewavas, 1932) cus; drechseli; embarbatus; frondo- Chirophryne (t.s. Chirophryne xeno- sus; furcifer; globulifer; leptobolus; lophus Regan & Trewavas, 1932) lipochirus; macronema; macrurus; Cryptolychnus (t.s. Cryptolychnus melanobolus; melanonema; mela- micractis Regan & Trewavas, nostigma; monoclonus; monodac- 1932) tylus; parri; patulus; pyrifer; ramu- Ctenochirichthys (t.s. Ctenochirich- losus; schmidti; silvescens; simplex; thys longimanus Regan & Trewa­ tenisoni; triramus; trituberatus; xe- vas, 1932) nobolus; variabilis Nannoceratias (t.s. Nannoceratias Haplostomias bituberatus; tentacula- denticulatus Regan & Trewavas, tus 1932) Lamprotoxus paucifilis, phanobran- Teleotrema (t.s. Teleotrema microph- chus thalmus Regan & Trewavas, 1932) Leptostomias analis, gracilis; haplo- Trematorhynchus (t.s. Rhynchocera- caulus; longibarba; leptobolus; ra- tias leucorhinus Regan, 1925) mosus Tyrranophryne (t.s. Tyrranophryne Melanostomias albibarba; biseriatus; pugnax Regan & Trewavas, 1932) heteropogon; macrophotus; mar- Xenoceratias (t.s. Xenoceratias longi- garidfer; melanocaulus; melanopo- rostris Regan & Trewavas) gon; spilorhynchus Sub. gen. nov. Dolopichthys (Leptacanthich- thys), t.s. D. gracilispinis Regan, Trematorhynchus exiguus; obliqui- 1925 dens Dolopichthys (Microlophichthys), Terranophryne pugnax t.s. D. microlophus Regan, 1925 Xenoceratias brevirostris; hetero- Dolopichthys (Pentherichthys), t.s. rhynchus; laevis; longirostris; mi- D. atratus Regan & Trewavas, 1932 cracanthus . nov. Aceratias edentulata ATHERINIFORMES Caranactis pumilus Atherinidae Caulophryne ramulosa Spp. nov. 1940 (22) Craterocephalus lacustris Caulophryne acinosa CYPRINODONTIFORMES Centrocetus spinulosus Poeciliidae Centrophryne spinulosa Sp. nov. 1948 (38) Mollienisia elegans Chaenophryne atriconus; columnif- Aplocheilidae er; crenata; fimbriata; haplactis; Sp. nov. 1949 (38) Rivulus roloffi macractis; melanodactylus; melan- Cyprinodontidae orhabdus; parviconus; pterolo- Sp. nov. 1949 (38) Procatopus lacustris phus; ramifera PERCIFORMES Chirophryne xenolophus Teraponidae Cryptolychnus micractis; paucidens Sp. nov. 1946 (32) Therapon adamsoni Cryptosaras normani; pennifer; val- Sciaenidae divae Gen. nov. 1962 (64) Ctenochirichthys longimanus Miracorvina (t.s. Sciaena angolensis Dolopichthys atratus; brevifilis; carls- Norman, 1935) bergi; cirrifer; claviger; cristatus; Pentheroscion (t.s. Sciaena mbizi diadematus; digitatus; exiguus; fla- Poll, 1950) gellifer; frondosus; heteronema; 1964 (68) Dendrophysa (t.s. Umbrina implumis; jubatus; macronema; russellii Cuv., 1830) mirus; multifilis; nigrifilis; penna- 1966 (73) Chrysochir (t.s. Otolithus tus; plumatus; pollicifer; ptilotus; aureus Richardson, 1846) pullatus; simplex; schmidti; thysa- 1977 (100) Afroscion (t.s. Argyroso- nophorus; venustus mus thorpei M.M. Smith, 1977) Edriolychnus macracanthus; radians; Austronibea (t.s. Austronibea oedo- roulei genys Trewavas, 1977) Gigantactis exodon; ovifer; sexfilis Boesemania (t.s. Corvina microlepis Himantolophus danae Bleeker, 1858) Lasiognathus beebi Paranibea (t.s. Nibea semiluctuosa Linophryne argyresca; corymbifera; Cuv., 1830) eupogon; racemifera Sonorlux (t.s. Sonorlux fluminis Tre­ Lophodolus dinema wavas, 1977) Mancalias bifilis; xenistius Sub-gen. nov. 1962 (64) Melanocetus cirrifer Pseudotolithus (Fonticulus), t.s. Cor­ Nannoceratias denticulatus vina nigrita Cuv., 1830 Rhynoceratias altirostris Teleotrema microphthalmus 59

Pseudotolithus (Hostia), t.s. Corvina 1973 (89) Pterochromis (t.s. Pelmato- moorii Gunther, 1865 chromis congicus Bouleng., 1897) Spp. nov. 1977 (100) 1989 (118) Buccochromis (t.s. Para­ Austronibea oedogenys tilapia nototaenia Bouleng., 1902) Panna heterolepis Cheilochromis (t.s. Sonorlux fluminus euchilus Trewavas, 1935) 1987 (116) Atrobucca geniae (t.s. Haplochromis Cichlidae quadrimaculatus Regan, 1922) Gen. nov. 1935 (14) Ctenopharynx (t.s. Hemichromis in­ Aristochromis (t.s. Aristochromis termedins Gunther, 1893) christyi Trewavas, 1935) Dimidichromis (t.s. Haplochromis Christyella (t.s. Christyella nyasana strigatus Regan, 1922) Trewavas, 1935) Eclectochromis (t.s. Haplochromis Cyathochromis (t.s. Cyathochromis ornatus Regan, 1922) obliquidens Trewavas, 1935) Exochromis (t.s. Cyrtocara anagenys Diplotaxodon (t.s. Diplotaxodon ar- Oliver, 1984. genteus Trewavas, 1935) Fossorochromis (t.s. Tilapia rostrata Genyochromis (t.s. Genyochromis Bouleng., 1899) mento Trewavas, 1935 Hemitaeniochromis (t.s. Haplochro­ Labidochromis (t.s. Labidochromis mis urotaenia Regan, 1922) vellicans Trewavas, 1935) Maravicochromis (t.s. Haplochromis Lichnochromis (t.s. Lichnochromis ericotaenia Regan, 1922) acuticeps Trewavas, 1935) Naevochromis (t.s. Haplochromis (t.s. Melanochromis chrysogaster Trewavas, 1935) melanopterus Trewavas, 1935) (t.s. Hemichromis li- Petrotilapia (t.s. Petrotilapia triden- vingstonii Gunther, 1893) tiger Trewavas, 1935) Nyassachromis (t.s. Haplochromis Trematocranus (t.s. Trematocranus breviceps Regan, 1922) microstoma Trewavas, 1935) Placidochromis (t.s. Haplochromis 1961 (63) Chetia (t.s. Chetia flaviven- longimanus Trewavas, 1935) tris Trewavas, 1961) Platygnathochromis (t.s. Haplochro­ 1962 (65) Barombia (t.s. Barombia mis melanotus Regan, 1922) maclareni Trewavas, 1962) (t.s. Chromis kirkii (t.s. Paratilapia mariae Gunther, 1893) Holly, 1930) (t.s. Haplochromis 1972 (87) Konia (t.s. Tilapia eisen- ahli Trewavas, 1935) trauti Trewavas, 1962) (t.s. Haplochromis (t.s. Myaka myaka Trewavas, woodi Regan, 1922) 1972) Taeniochromis (t.s. Haplochromis N.B. Although Green and Corbet were co-authors holotaenia Regan, 1922) of the paper in which these taxa were described, Taeniolethrinops (t.s. Haplochromis Trewavas was the sole author of the taxonomic sec­ praeorbitalis Regan, 1922) tion. Tramitichromis (t.s. Tilapia brevis Bouleng., 1908) 60

Tyrannochromis (t.s. Haplochromis chus; taeniolatus; tetraspilus; triae- macrostoma Regan, 1922) nodon; virgatus Sub-gen. nov. 1983 (110) (Valli- Labidochromis vellicans cola), t.s. Tilapia amphimelas Hil- Lethrinops intermedia gen., 1905. Lichnochromis acuticeps 1984 (111) (Trophe- Melanochromis brevis; labrosus; mel­ ops), t.s. Pseudotropheus trophe- anopterus; perspicax; vermivorous ops Regan, 1921 Petrotilapia tridentiger Spp. nov. 1928 (2) Haplochromis gowersi; max- Pseudotropheus elegans; fuscus; lu- illaris; melanopterus; michaeli; ob- cerna; microstoma tusidens brevis 1928 (3) Haplochromis diplotaenia; Trematocranus auditor; brevirostris; eutaenia; plagiodon; tridens microstoma 1931 (7) Lethrinops alta; christy i; cyr- 1938 (19) Haplochromis bullatus tonotus; furcicauda; furcifer; lati- 1941 (24) Tilapia karongae; lidole; ceps; lituris; longimanus; lunaris; placida macracanthus; microstoma; oculta; 1943 (27) Gobiochromis irvinei parvidens; trilineata; variabilis 1952 (43) Lamprologus christyi; 1933 (12) Haplochromis annectidens; pleuromaculatus; sexfasciatus beadlei; dolorosus; elegans; engys- 1953 (45) Limnochromis christyi toma; labiatus; Umax; rudolfianus; 1960 (60) Petrochromis famula taurinus; vellifer; vicarius 1962 (65) Barombia maclareni Pelmatochromis exul Pelmatochromis loennbergi Tilapia inducta; leucosticta; vulcani Tilapia eisentrauti; steinbachi 1935 (14) Aristochromis christyi 1964 (69) Serranochromis janus;spei; macrochir; rostrata stappersi Christyella nyasana 1966 (79) Tilapia mortimeri; ruvumae Cyathochromis obliquidens 1969 (81) Haplochromis albolabris Diplotaxodon argenteus 1972 (87) Konia Genyochromis mento Myakamyaka Haplochromis balteatus; chrysogas- Stomatepia pindu ter; cynaneus; decorus; epichoria- See note under 1972 (87), page 59 regarding correct lis; euchilus; fenestratus; festivus; authorship of these names. formosus; gracilis; heterodon; het­ 1973 (89) angolensis erotaenia; incola; insignis; labifer; 1974 (94) Chromidotilapia finleyi labiodon; labridens; lobochilus; 1975 (95) Tilapia baloni longimanus; marginatus; mola; 1975 (96) Nanochromis gabonicus mollis; microcephalus; nigritaenia- 1976 (98) Hemichromis fugax tus; nitidus; obtusus; oculatus; or- 1984 (112) Aulonocara maylandi thognathus; ovatus; pardalis; phe- Sub-spp. nov. 1935 (14) Haplochromis margi­ nochilus; pholidophorus; pictus; natus vuae pleurospilus; pleurostigma; polyo- Pseudotropheus tropheops gracilior don; protostoma; purpurans; semi- 1936 (15) Haplochromis philander palatus; serenus; speciosus; spiloti- dispersus 61

1952 (43) Lamprologus savoryi eloti- gatus References cited Lamprologus savoryi pulcher 1966 (78) Tilapia shirana chilwae Eschmeyer, W.N. 1990. Catalog of the genera of recent fishes. California Academy of Sciences, San Francisco, 697 pp. 1983 (10) Oreochromis macrochir Greenwood, P.H., D.E. Rosen, S.H. Weitzman & G.S. Myers. mweruensis 1966. Phyletic studies of teleostean fishes, with a provisional Oreochromis niloticus baringoensis classification of living forms. Bull. Amer. Mus. Nat. Hist. 131: Oreochromis niloticus filoa 339-456. Noakes, D.L.G. 1994. Lifetime list of publications by Ethelwynn Oreochromis niloticus sugutae Trewavas. Env. Biol. . 41: 51-54 (this volume).

Acknowledgement

I am indebted to Huibre Tomlinson for her efforts in preparing this manuscript at very short notice.