TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Zootaxa 3746 (4): 597–599 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Correspondence ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3746.4.7 http://zoobank.org/urn:lsid:zoobank.org:pub:6045D5B0-90A3-4610-A130-5FFAC69A1EF4

The genus Orestias (Teleostei: Cyprinodontidae): nomenclatural errors in the assignation of species names

FRANCO CRUZ-JOFRÉ1, MOISÉS A. VALLADARES1, IRMA VILA2 & MARCO A. MÉNDEZ1,3,4 1Laboratorio de Genética y Evolución, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Casilla 653, Ñuñoa, Santiago, Chile 2Laboratorio de Limnología, Departamento de Ciencias Ecológicas, Facultad de Ciencias, Universidad de Chile, Las Palmeras 3425, Ñuñoa, Chile 3Instituto de Ecología y Biodiversidad (IEB), Universidad de Chile, Las Palmeras 3425, Ñuñoa, Chile 4Corresponding author. E-mail: [email protected]

Orestias Valenciennes 1839 is a fish genus of the family Cyprinodontidae Gill, 1865, endemic to the Altiplano in the range. The genus includes 45 species (44 valid and one species inquirenda) according to Eschmeyer & Fong (2013); more than half inhabit the basin (23 species). The rest of the species have been described from different Altiplano aquatic systems, from Lake Lacsha in the center of to the Ascotán salt pan in northern Chile (9°S to 22°S) (Lauzanne 1982; Vila 2006; Vila et al. 2011). Due to the high species diversity, the genus Orestias has been used as a study model for speciation processes. Sympatric speciation has been proposed for Lake Titicaca (Baker et al. 2001), and allopatric speciation for the rest of its geographic distribution (Parker & Kornfield 1995). The genus Orestias was originally proposed by Valenciennes (1839). This author indicated “Puisqu’ils habitent les points les plus élevés des Andes, et qu’il faut les nommer comme nouveaux, je propose, pour désigner ce genre, le nom d’Orestie (d’Ορεστιας, nymphe des montagnes)”. Along with the description of the genus, Valenciennes (1839) assigned specific binominal names to species that he mentioned by their common names, but provided no descriptions. These names are nomina nuda and are excluded from the nomenclature by Article 12.3 of the International Code of Zoological Nomenclature (ICZN 1999). Later, Valenciennes (1846) repeated the etymological origin of the name Orestias and made a detailed (in ICZN terms) description of the following species (those mentioned in 1839 without descriptions or valid indications are indicated with *):

Orestias cuvieri,* dedicated to Georges Cuvier (1769–1832); type species of the genus by the subsequent designation of Eigenmann (1910); common name: umanto. Orestias albus;* common name: carache blanco. Orestias luteus;* common name: carache amarillo. Orestias humboldti,* dedicated to Alexander von Humboldt (1769–1859) (synonym of O. cuvieri, according to Parenti 1984); common name: peje reye. Orestias pentlandii,* dedicated to Joseph Barclay Pentland (1797–1873); common name: boga or boguilla. Orestias jussiei, dedicated to Joseph de Jussieu (1704–1779); common: ispis. Orestias agassizii, O. agassii or O. agassisii, dedicated to Jean Louis Rodolphe Agassiz (1807–1873) (see comments below); common name: puru. Orestias mulleri, possibly dedicated to Johann Friedrich “Fritz” Theodor Müller, (1822–1897). Orestias owenii, dedicated to Richard Owen (1804–1892) (synonym of O. jussiei, according to Parenti 1984).

Valenciennes did not mention the grammatical gender of the word orestias; however, he clearly specified its Greek origin and meaning. The word ?ρεστιάς (orestias) is a feminine noun1, which in Greek mythology corresponds to the singular of ?ρεστιάδες (orestiades), which refers to feminine divinities (nymphs) who live in the mountains2. According to the ICZN (Art. 31.2), all names of species, if they correspond to a Latin or Latinized adjective, must concur in gender with the generic name with which they combine. It is thus surprising that Valenciennes (1846) assigned adjectives with masculine suffixes as specific names to species of the genus Orestias (e.g. albus and luteus, Latin adjectives which mean white and yellow, respectively).

1. Morwood, J. & Taylor J. (2002) Pocket Oxford Classical Greek Dictionary, p 233. 2. Smith, W. (1867) Dictionary of Greek and Roman Biography and Mythology, Vol. II, 1216.

Accepted by A. Dubois: 25 Nov. 2013; published: 13 Dec. 2013 597 TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Another nomenclatural inconsistency in the report of Valenciennes (1846) occurred with the species of Orestias dedicated to Jean Louis Agassiz, which was spelled three different ways in the paper. In the Table of Contents (Table XV) and in the table of figures of volume XVIII it was called O. agassizii, in the central text in which the species was described it was called O. agassii (p. 238), and finally it was called O. agassisii in the Figure (p. 536). The result has been that some authors have used O. agassii (Tchernovin 1944; Parenti 1984), while others used O. agassizii (Garman 1876, 1895; Eigenman & Allen 1942). The International Commission of Zoological Nomenclature anticipated this kind of situation, and stipulates that when two or more names are assigned to the same species and published in the same paper, the nomenclatural precedence is fixed by the First Reviser (Art. 24.2 ICZN 1999). In the case of the species dedicated to Agassiz, O. agassii was recognized by Tchernavin (1944) as the correct name; however this author only reviewed two spellings (O. agassii and O. agassisii) from Valenciennes (1846), instead the third form (O. agassizii) was reviewed from Garman (1876), this proceeding overriding to Tchernavin as First Reviser, since according to the Code if a name is spelled in more than one way in the original work, the first author to have cited them together and to have selected one spelling as correct is the First Reviser (Art. 24.2.3 , ICZN 1999). In our opinion the First Reviser must be Parenti (1984), who recognized O. agassii as the correct name for the species, after review the three spellings from Valenciennes (1846). By this nomenclatural act O. agassii acquired validity and is the name, which must be used in the future. Since the descriptions of Valenciennes (1846), 38 species of Orestias have been validly described. Most of them were identified with specific names allusive to the localities where they were found, e.g. O. uruni Tchernavin, 1944 (for the Uruni Bay), O. parinacotensis Arratia, 1982 (for the Parinacota Wetland), O. laucaensis (for the Lauca River), O. ascotanensis Parenti, 1984 (for the Ascotán salt pan), O. chungarensis Vila & Pinto, 1986 of Lake Chungará and O. piacotensis Vila, 2006, described for Piacota Lagoon. Other species have been dedicated to persons, such as O. tomcooni Parenti, 1984 (dedicated to Tom Coon) and the most recently described species O. gloriae Vila, Scott, Méndez, Valenzuela, Iturra & Poulin, 2011(dedicated to Gloria Arratia). In all these cases the combinations with the generic name Orestias have been correctly designated (Art. 31.1, ICZN 1999) and do not require changes. However, some authors have used masculine Latin or Latinized adjectives as specific names, which is inconsistent with the grammatical gender of Orestias. Since Orestias is a feminine noun, both for grammatical reasons and in order to comply with Article 31.2 (ICZN 1999), these specific names should concur grammatically with the name of the genus with which they are associated (Cazzaniga 2011). Also, as indicated by the Code, if the gender ending of the species group is incorrect it must be changed to the correct form (Art. 34.2). Thus we propose that the Latin gender of some specific names of Orestias must be changed (Table 1). This modification corresponds to an obligatory correction and does not modify the authorship of the original descriptions of the species (Art. 50.3.2, ICZN 1999).

TABLE 1. Specific names which require modification according to the ICZN.

Original name Meaning Corrected Name1 Author Orestias albus White Orestias alba Valenciennes, 1846 Orestias luteus yellowish Orestias lutea Valenciennes, 1846 Orestias frontosus having several faces Orestias frontosa Cope, 1876 Orestias olivaceus olive-colored Orestias olivacea Garman, 1895 Orestias minutus Small Orestias minuta Tchernavin, 1944 Orestias minimus very small, the smallest Orestias minima Tchernavin, 1944 Orestias gymnotus that is naked Orestias gymnota Parenti, 1984 Orestias mundus clean, nitid Orestias munda Parenti, 1984 Orestias robustus robust, strong Orestias robusta Parenti, 1984

1Adjetives of the first type in Latin have the first and second declinations (-us, masculine; -a, feminine; -um, neuter)

It is probable that several authors (Cope 1876; Garman 1895; Tchernavin 1944; Parenti 1984) simply maintained the adjectives in masculine form following Valenciennes (1846), who incorrectly assigned the masculine gender to the species O. albus and O. luteus. The modifications proposed here do not alter the stability defined by the Code, since they conserve the names of the species (correcting the gender) with their corresponding authors and dates. These changes serve to order and define nomenclature criteria in a genus used as a model in speciation and evolutionary studies, and that is currently under systematic studies (Esquer-Garrigos et al. 2013; Vila et al. 2013). In this contribution we make a necessary correction to comply with the Code, correcting a nomenclatural error with at least 170 years of history.

598 · Zootaxa 3746 (4) © 2013 Magnolia Press CRUZ-JOFRÉ ET AL. TERMS OF USE This pdf is provided by Magnolia Press for private/research use. Commercial sale or deposition in a public library or website is prohibited.

Acknowledgements

We thank Néstor Cazzaniga, Gonzalo Collado and Alain Dubois for useful suggestions and comments on the manuscript. This study was financed by grants FONDECYT 1110243, 1110188, Programa de Cooperación Internacional CONICYT: grant ECOS-CONICYT C10B02 and grant REDES130016.

Literature cited

Baker, P.A., Seltzer, G.O., Fritz, S.C., Dunbar, R.B., Grove, M.J., Tapia, P.M., Cross, S.L., Rowe, H.D. & Broda, J.P. (2001) The History of South American Tropical Precipitation for the Past 25,000 Years. Science, 291, 640–643. http://dx.doi.org/10.1126/science.291.5504.640 Cazzaniga, N. (2011) El género de los géneros: una guía para formar y coordinar nombres científicos en zoología. Editorial de la Universidad Nacional del Sur. Bahía Blanca, Argentina, 446 pp. Cope, E.D. (1876) Note on the ichthyology of Lake Titicaca. Journal of the Academy of Natural Sciences of Philadelphia, 8, 185–187. Eigenmann, C.H. (1910) Catalogue of the fresh-water fishes of tropical and south temperate America. Reports of the Princeton University expeditions to Patagonia 1896-1899. Zoology, 3 (4), 375–511. Eigenman, C.H. & Allen, W.R. (1942) Fishes of western South America. I. The intercordilleran and Amazonian lowlands of Peru. II. The High Pampas of Peru, and Northern Chile with a revision ofthe Peruvian Gymnotidae, and of the genus Orestias. University of Kentucky, Kentucky, USA, 494 pp. Eschmeyer, W. & Fong, J. (2013) Species of fishes by family/subfamily. Online catalogue of fishes. Electronic version. Available from: http://researcharchive.calacademy.org/research/ichthyology/catalog/fishcatmain.asp (accessed 16 April 2013) Esquer-Garrigos, Y., Hugueny, B., Koerner, K., Ibañez, C., Bonillo, C., Pruvost, P., Causse, R., Cruaud, C. & Gaubert, P. (2013) Non-invasive ancient DNA protocol for fluid-preserved specimens and phylogenetic systematics of the genus Orestias (Teleostei: Cyprinodontidae). Zootaxa, 3640 (3), 373–394. http://dx.doi.org/10.11646/zootaxa.3640.3.3 Garman, S.W. & Agassiz, A. (1876) Exploration of Lake Titicaca. I. Fishes and Reptiles. Bulletin of the Museum of Comparative Zoology, 3, 273–278. Garman, S.W. (1895) The cyprinodonts. Memoirs of the Museum of Comparative Zoology at Harvard College, Cambridge, USA, pp. 150–152. ICZN [International Commission on Zoological Nomenclature] (1999) International Code of Zoological Nomenclature. Fourth Edition. International Trust for Zoological Nomenclature, London, 106 pp. Lauzanne, L. (1982) Les Orestias (Pisces, Cyprinodontidae) du Petit lac Titicaca. Revue d’Hydrobiologie Tropicale, 15, 39–70. Parenti, L. (1984) A taxonomic revision of the Andean killifish genus Orestias (, Cyprinodontidae). Bulletin of the American Museum of Natural History, 178, 107–214. Parker, A. & Kornfield, I. (1995) Molecular perspective on evolution and zoogeography of cyprinodontid killifishes (Teleostei; Atherinomorpha). Copeia, 1995 (1), 8–21. http://dx.doi.org/10.2307/1446795 Tchernavin, V.V. (1944) A revision of the subfamily Orestiinae. Proceedings of the Zoological Society of London, 114, 140–233. http://dx.doi.org/10.1111/j.1096-3642.1944.tb00218.x Valenciennes, A. (1839) Poissons d'Amerique. Rapport sur quelques poissons d'Amerique rapportes par M. Pentland du lac Titicaca et des autres points eleves des Andes. L'Institut, 276, 118. Valenciennes, A. (1846) Chapitre XII Des Orestias. In: Cuvier, G. & Valenciennes, A. (Eds.), Histoire Naturelle des Poissons (Volume XVIII, Livre XVIII). Paris, Francia, pp. 221–244. Vila, I. & Pinto, M. (1986) A new species of Killifish (Pisces, Cyprinodontidae) from the Chilean Altiplano. Revue d’Hydrobiologie Tropicale, 19, 233–239. Vila, I. (2006) A new species of Killifish in the Genus Orestias (Teleostei; Cyprinodontidae) from the Southern High Andes, Chile. Copeia, 2006 (3), 471–477. http://dx.doi.org/10.1643/0045-8511(2006)2006[472:ansoki]2.0.co;2 Vila, I., Scott, S., Méndez, M.A., Valenzuela, F., Iturra, P & Poulin, E. (2011) Orestias gloriae, a new species of cyprinodontid fish from saltpan spring of the southern high Andes (Teleostei: Cyprinodontidae). Ichthyological Exploration of Freshwaters, 22, 345–353. Vila, I., Morales, P., Scott, S., Poulin, E., Véliz, D., Harrod, C. & Méndez, M. (2013) Phylogenetic and phylogeographic analysis of the genus Orestias (Teleostei; Cyprinodontidae) in the Southern Chilean Altiplano: the relevance of ancient and recent divergence processes in speciation. Journal of Fish Biology, 82, 927–943. http://dx.doi.org/10.1111/jfb.12031

NOMENCLATURAL ERRORS IN GENUS ORESTIAS Zootaxa 3746 (4) © 2013 Magnolia Press · 599