A TAXONOMIC REVISION OF THE ANDEAN KILLIFISH GENUS ORESTIAS (CYPRINODONTIFORMES, CYPRINODONTIDAE) LYNNE R. PARENTI BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 178: ARTICLE 2 NEW YORK: 1984 A TAXONOMIC REVISION OF THE ANDEAN KILLIFISH GENUS ORESTIAS (CYPRINODONTIFORMES, CYPRINODONTIDAE) LYNNE R. PARENTI Research Associate, Department ofIchthyology American Museum ofNatural History BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 178, article 2, pages 107-214, figures 1-72, tables 1-12 Issued May 9, 1984 Price: $8.40 a copy Copyright ©) American Museum of Natural History 1984 ISSN 0003-0090 CONTENTS Abstract ............................... 110 Introduction ............................... 110 Acknowledgments ............................... 113 Abbreviations ............................... 114 Note on Materials and Methods ............................... 115 Relationships of Killifishes of the Tribe Orestiini ............................... 116 Phylogenetic Analysis ...................................... 122 Squamation and Neuromast Pattern ........... .................... 123 Hyobranchial Apparatus ............................... 126 Fins ............................... 129 Chromosomes ............................... 130 Sexual Dimorphism and Dichromatism .............. ................. 131 Jaw and Jaw Suspensorium ...................................... 133 Skull ............................... 134 Meristic Characters ............................... 135 Morphometric Characters ............................... 146 Explanation of Synapomorphy Diagrams .............. ................. 150 Key to Orestias Species ............................... 160 Systematic Accounts ...................................... 165 Genus Orestias Valenciennes ............................... 165 Orestias cuvieri Valenciennes ............................... 167 Orestias pentlandii Valenciennes .......................-1-68 Orestias ispi Lauzanne ...................... 169 Orestiasforgeti Lauzanne ...................... 170 Orestias mulleri Valenciennes ...................... 171 Orestias gracilis, New Species ........................... 172 Orestias crawfordi Tchernavin ...................... 173 Orestias tutini Tchemavin ........................... 174 Orestias incae Garman ...................... 174 Orestias luteus Valenciennes ...................... 175 Orestias rotundipinnis, New Species ...................... 176 Orestiasfarfani, New Species ...................... 178 Orestias albus Valenciennes ...................... 179 Orestias olivaceus Garman ...................... 180 Orestias silustani Allen ...................... 181 Orestias agassii Valenciennes ...................... 182 Orestias empyraeus Allen ...................... 185 Orestiasfrontosus Cope ...................... 186 Orestias polonorum Tchemavin ...................... 188 Orestias elegans Garman ...................... 188 Orestias jussiei Valenciennes ...................... 189 Orestias puni Tchernavin ...................... 189 Orestias parinacotensis Arratia ...................... 190 Orestias laucaensis Arratia ...................... 190 Orestias tschudii Castelnau ...................... 191 Orestias gymnotus, New Species .... ...... .. .. ... ... ...... .. .. 192 Orestias hardini, New Species ...... .... ..... .... 193 Orestias ctenolepis, New Species .... ..................................... 194 Orestias ascotanensis, New Species.. ..................................... 196 Orestias richersoni, New Species .... ..................................... 197 Orestias multiporis, New Species .... ..................................... 198 Orestias mundus, New Species ...... ..................................... 199 Orestias ututo, New Species ........ ..................................... 200 Orestias gilsoni Tchemavin......... ..................................... 201 Orestias taquiri Tchernavin......... ..................................... 201 Orestias mooni Tchemavin ......... ..................................... 202 Orestias uruni Tchemavin.......... 203 Orestias minimus Tchernavin.............................. ................... 203 Orestias minutus Tchemavin .............................. ................... 204 Orestias tchernavini Lauzanne ............................. ................... 205 Orestias imarpe, New Species............................ ................... 205 Orestias tomcooni, New Species.......................... ................... 206 Orestias robustus, New Species........................... ................... 207 ?Orestias cuvieri X pentlandii .............................. ................... 208 Orestias rospigliosii Eigenmann and Allen................... ................... 209 Orestias pentlandii var. fuscus Garman ..................... ................... 209 Summary .................................................. ................... 209 Literature Cited ............................................ ................... 211 ABSTRACT The ichthyofauna of the high Andes is assumed is a unique squamation and head pore pattern to be of low taxonomic diversity. Only three fish characterized by a prominent lyre-shaped arrange- genera, the killifish Orestias and the catfishes As- ment of minute neuromasts and a prominent me- troblepus and Trichomycterus include species en- dian dorsal ridge of scales running from the top demic to the Altiplano, the high-altitude plateau of the head to the dorsal fin origin. between the eastern and western slopes ofthe cen- Primary characters used to distinguish Orestias tral Andes. The atherinid Basilichthys is known species are overall body shape and squamation doubtfully from the Altiplano. pattern. Diversity of form is marked. There are In the last revision ofthe genus Orestias, Tcher- large, troutlike midwater predators, such as 0. cu- navin (1 944a) recognized 20 species and five sub- vieri, diminutive inshore species such as 0. mi- species. This small number was taken as evidence nutus and 0. minimus, and very wide-headed and of the low diversity of fish of the Titicaca Basin. wide-bodied species, with large, thick granulated However, in the past two years, five new Orestias scales, such as 0. luteus and 0. albus. species have been described, three from Lago Ti- The diversity among Orestias species in Lago ticaca (Lauzanne, 1981) and two from the Alti- Titicaca, the large, high-altitude lake of the Alti- plano of northern Chile (Arratia, 1982). With the plano of Peru and Bolivia, has led to the appli- benefit of large recent collections, 43 species of cation of the term "species flock" to this group of Orestias are recognized in the present revision, 14 lacustrine killifish. However, the Orestias of Ti- being described as new (0. gracilis, 0. rotundi- ticaca do not form a monophyletic group, and it pinnis, 0. farfani, 0. gymnotus, 0. hardini, 0. is recommended that the term "species flock" not ctenolepis, 0. ascotanensis, 0. richersoni, 0. mul- be used for the killifishes limited to the lake. tiporis, 0. mundus, 0. ututo, 0. imarpe, 0. tom- The known range of Orestias extends from cooni, and 0. robustus). northern Peru to northern Chile. It is expected that Seven synapomorphies described here define the number will rise with additional collecting, Orestias as a monophyletic group: (1) pelvic fins because collections are lacking from many of the and fin girdle are absent; (2) vomer is absent; (3) small, isolated lakes of the Altiplano; also, single middle anal and middle dorsal fin radials are car- specimens of apparently new species are known, tilaginous, rather than ossified; (4) bony anterior yet remain undescribed. and posterior ceratohyal are separated ventrally The hypothesis of a close relationship between by a large gap filled with cartilage; (5) first postclei- Orestias and the Anatolian cyprinodonts is re- thrum is absent; (6) anguloarticular lacks a ventral viewed. extension parallel to the retroarticular; and (7) there INTRODUCTION The killifish genus Orestias (Cyprinodon- fish genera with some species endemic to the tiformes, Cyprinodontidae) is endemic to the Altiplano of the central Andes. high-altitude lakes and tributary streams of More than halfofthe 43 species of Orestias the Peruvian, Bolivian, and Chilean Andes. are endemic to the closed Titicaca Basin, and In the present revision the genus is consid- of those, 23 are known only from Lago Ti- ered to comprise 43 species. The range of ticaca, the large (8100 km2 surface area), high- Orestias (fig. 1) extends from Ancash Prov- altitude (3803 m) lake of the Altiplano, sit- ince, northern Peru to Antofagasta Province, uated between lat. 15°S and 17°S and between northern Chile. The distribution is coinci- long. 68°W and 7 1°W (with the midpoint of dent only in part with the catfish genera As- the lake at approximately 1 5045'S and troblepus and Trichomycterus (see Arratia, 69°25'W) in Peru and Bolivia (Richerson, 1982 and Arratia et al., 1978) and the sil- Widmer, and Kittel, 1977). verside genus Basilichthys,' the only other In the last revision of Orestias, Tchernavin (1 944a) formally recognized 20 species and I The genus Basilichthys is known as an Altiplano en- five subspecies. Recently, three more species demic from just one specimen, 17.5 mm SL, FMNH have been described from Lago Titicaca 94184, collected by W. R. Allen, 1918-1919, from a (Lauzanne, 1981), and two from the Alti- pool below Acora, Peru, a village near Chucuito, near plano of northern Chile (Arratia, 1982). In Bahia de Puno (fig. 22). the present paper, 14