Zootaxa 3608 (6): 440–456 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN1175-5334(online edition) http://dx.doi.org/10.11646/zootaxa.3608.6.2 http://zoobank.org/urn:lsid:zoobank.org:pub:8B60241F-8229-49B8-9B28-0CAC90614E38 Checklist of the Inland Fishes of El Salvador CALEB D. MCMAHAN1, WILFREDO A. MATAMOROS1,2, FRANCISCO S. ÁLVAREZ CALDERÓN3, WENDY YAMILETH HENRÍQUEZ3, H. MICHELLE RECINOS3, PROSANTA CHAKRABARTY1, ENRIQUE BARRAZA4 & NÉSTOR HERRERA4 1LSU Museum of Natural Science (Ichthyology), Department of Biological Sciences, Louisiana State University, 119 Foster Hall, Baton Rouge LA 70803 USA. E-mail: [email protected]. 2Department of Biological Sciences, University of Southern Mississippi, 118 College Dr., Hattiesburg, MS 39406 USA. E-mail: [email protected]. 3Universidad de El Salvador, Facultad de Ciencias Naturales y Matemáticas, Escuela de Biología, Final 25 Avenida Norte, Ciudad Universitaria, San Salvador, El Salvador. E-mail: [email protected], [email protected], [email protected]. 4Ministerio de Medio Ambiente y Recursos Naturales, Col. y Calle Las Mercedes, San Salvador, El Salvador. E-mail: [email protected], [email protected]. Abstract The inland fish fauna of El Salvador and its distribution was originally described in 1925 by Samuel Hildebrand. That work has been the main source of information for freshwater fishes of El Salvador up to today. Based on the combination of an intensive literature review, electronic database searches, re-identification of museum specimens, and fieldwork, we hereby provide an updated checklist of the inland fishes of El Salvador. This checklist provides distributional data at the Salvadoran hydrographical and political (by department) levels. The checklist is systematically arranged at the ordinal and familial level and then alphabetically therein. The freshwater fish fauna of El Salvador includes 101 species divided into 64 genera, 29 families, and 14 orders. According to their supposed tolerance to salinity, 73% of these species are peripheral, 23% secondary, and only 4% are primary freshwater fishes. One species is endemic to the country, Amatitlania coatepeque. The low number of primary freshwater fishes and endemics is comparable to the Central American Pacific slope in particular, as well as northern Central America in general. Key words: Pisces, Central America, hydrographic zones, Middle America Resumen La fauna de peces de las aguas interiores de El Salvador y su distribución fueron originalmente descritas en 1925 por Samuel Hildebrand. Hasta hoy dicho estudio ha sido la fuente principal de información en cuanto a peces de agua dulce de El Salvador. Basados en una combinación de intensa revisión de la literatura, búsqueda en bases de datos electrónicas, verificación de colecciones de museos y trabajo de campo, proveemos una lista actualizada de los peces de aguas interiores de El Salvador. La lista incluye datos de distribución al nivel de las unidades hidrológicas salvadoreñas, así como también de los departamentos políticos. La lista esta ordenada sistemáticamente por orden y familia y después alfabéticamente. La ictiofauna de agua dulce de El Salvador está formada por 101 especies, en 64 géneros, 29 familias y 14 órdenes. Según su tolerancia a la salinidad, 73% de las especies son periféricas, 23 por ciento son secundarias, y solo un cuatro por ciento se encontró como primarios de agua dulce. Una especie es endémica del país, Amatitlania coatepeque. El bajo número de especies primarias de agua dulce así como de especies endémicas es comparable con otras áreas del Pacifico centroamericano en particular, así como al norte de Centroamérica en general. Key words: distribution, Neotropics, rivers, lake, Middle America 440 Accepted by L. Page: 10 Dec. 2012; published: 23 Jan. 2013 Introduction El Salvador is the smallest of the Middle American countries, as well as the only one with coastline solely on the Pacific slope. It is bordered by Guatemala to the north and northwest, and by Honduras to the north and northeast. The Gulf of Fonseca separates El Salvador from Honduras and Nicaragua in the southeast. The Sierra Madre de Chiapas runs along northern El Salvador, and to the south is a mountainous chain of volcanoes, with a central plateau positioned between these two mountain ranges. Lowland tropical savannah largely characterizes the area from the mountains to the Pacific Ocean (Marshall 2007). Politically, the country is divided into 14 departments (Fig. 1) and has a total land area of approximately 23,040 km2 (22,720 km2 land and 320 km2 freshwater). Most of the rivers found in El Salvador are part of the Río Lempa basin, with a few exceptions such as the Río Goascorán at the border with Honduras. However, multiple hydrographic zones are recognized within the country, and these are used herein (Fig. 2). Within the highlands are several crater lakes of volcanic origin (e.g. Lago Ilopango, Lago Coatepeque, Lago Güija), and just north of San Salvador is the Cerrón Grande Dam damming the Río Lempa. The reservoir created by this dam (Lago Suchitlán) is the largest body of freshwater in the country. As typical of the Neotropics, El Salvador has a tropical climate with rainy (May through October) and dry (November through April) seasons. In addition to being the smallest of the Middle American countries, El Salvador is also the most densely populated, with a population of nearly 6,000,000 people. The first review of fishes from El Salvador was by Hildebrand (1925). Samuel Hildebrand and Fred Foster of the US Bureau of Fisheries were sent to survey fishes (both freshwater and marine) of the country, and their checklist (Hildebrand 1925) was a summary of that expedition. Prior to this expedition, only two species records existed for the country. The most recent significant freshwater expeditions are those by the Natural History Museum of Sweden (NRM; 2006), and the Louisiana State University Museum of Natural Science in the United States (LSUMZ; 2011). In addition to native species, in past years multiple species of freshwater fishes have been introduced into the country for aquaculture and food. It should be noted that substantial material of marine fishes collected off the coast of El Salvador exist from recent expeditions by the Smithsonian National Museum of Natural History (USNM) and Smithsonian Tropical Research Institute (STRI). Those specimens are deposited in the USNM. In addition, an expedition to the coastal waters of El Salvador was conducted by the Natural History Museum at the University of Kansas (KU; 2008). Some recent work has also focused on biology of fishes from El Salvador (e.g. Hernández-Rivas & Calderón 1974, Phillips 1981, Phillips 1983, Burns & Ramírez 1990, Álvarez, et al. in review). The purpose of this updated and revised checklist is to provide an overview of the inland fishes of the country, as we presently understand them. Here we refer to inland fishes as freshwater (primary, secondary) and peripheral species. As such, this information is intended to provide baseline data for future systematic and biogeographic studies, as well as illuminate where more sampling efforts are needed. In addition, checklists such as this are crucial to agencies interested in the biodiversity of fishes in El Salvador and their conservation. Materials and Methods Distributional and diversity data for El Salvador inland fishes were obtained by several methods. Initially, a thorough review of scientific literature was conducted for studies on fishes of El Salvador. Species occurrence records were further obtained by querying the inter-institutional online database FishNet2 (www.fishnet2.org). Finally, in order to supplement the data obtained by the above sources, we conducted an expedition to El Salvador from 29 May to 11 June 2011. In the field, fishes were captured by the use of multiple methods including seines, cast nets, and electrofishing. Specimens were preserved and deposited in the LSU Museum of Natural Science. All species presented in this checklist have vouchered specimens in museum collections. Comments are provided for species where annotations are necessary (e.g. substantial range extensions). This checklist is arranged evolutionarily at the ordinal and familial levels following Nelson (2006). Genera and species within families are arranged in alphabetical order. Valid species names, their authorities, and year of publication follow Eschmeyer (2012), with the exception of Rhamdia guatemalensis (sensu Perdices et al. 2002). Each family is classified by salinity tolerance (primary, secondary, peripheral) following Myers (1949). When available, Salvadoran Spanish and English names (following Fischer et al. 1995 and Nelson et al. 2004) are also provided for each species, as well as the origin of each species (native unless otherwise noted as introduced or endemic). Finally, distributional ranges are presented at the political (department) level (Fig. 1) and at the hydrographical level (Fig. 2). All rivers in CHECKLIST OF EL SALVADOR FRESHWATER FISHES Zootaxa 3608 (6) © 2013 Magnolia Press · 441 El Salvador drain to the Pacific Ocean; therefore, slope is not included in the checklist as we assume that it is implicit that all species occurrences are for the Pacific slope. FIGURE 1. Political map of El Salvador. A=Ahuachapán, B=Sonsonate, C=La Libertad, D=San Salvador, E=La Paz, F=San Vicente, G=Usulután, H=San Miguel, I=Morazán, J=La Unión, K=Santa Ana, L=Chalatenango, M=Cuscatlán, N=Cabañas. FIGURE 2. Hydrographic map showing major hydrographic zones in El Salvador. A=Río Lempa, B=Río Paz, C=Cara Sucia San Pedro, D=Grande de Sonsonate, E=Mandinga Comalapa, F=Río Jiboa, G=Estero Jaltepeque, H=Bahía Jiquilisco, I=Río Grande de San Miguel, J=Sirama, K=Río Goascorán, L=Lago Coatepeque. 442 · Zootaxa 3608 (6) © 2013 Magnolia Press MCMAHAN ET AL. Checklist of El Salvador Inland Fishes ORDER LEPISOSTEIFORMES Family Lepisosteidae Secondary Atractosteus tropicus Gill, 1863 (Machorra, Tropical Gar) Hydrographical zones: Río Paz, Cara Sucia San Pedro Department: Ahuachapán ORDER CLUPEIFORMES Family Engraulidae Peripheral Anchoa curta (Jordan & Gilbert, 1882) (Anchoa corta, Short anchovy) Hydrographical zone: Sirama Department: La Unión Anchoa mundeola (Gilbert & Pierson, 1898) (Falsa anchoveta, False Panama anchovy) Hydrographical zone: Río Paz Departments: Ahuachapán, Usulután Anchoa spinifer (Valenciennes, 1848) (Anchoa de fondo, Spicule anchovy) Hydrographical zones: Mandinga-Comalapa, Sirama Departments: La Libertad, La Unión Anchoa starksi (Gilbert & Pierson, 1898) (Ancho de starks, Starks’ anchovy) Hydrographical zone: Sirama Department: La Unión Anchoa walkeri Baldwin & Chang, 1970 (Anchoa de Walker, Walker’s anchovy) Hydrographical zones: Mandinga-Comalapa, Sirama Departments: La Libertad, La Unión Anchovia macrolepidota (Kner, 1863) (Anchoveta escamuda, Bigscale anchovy) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Family Pristigasteridae Peripheral Ilisha fuerthii (Steindachner, 1875) (Sardina hacha, Pacific ilisha) CHECKLIST OF EL SALVADOR FRESHWATER FISHES Zootaxa 3608 (6) © 2013 Magnolia Press · 443 Hydrographical zone: Bahía Jiquilisco Department: Usulután Family Clupeidae Peripheral Neoopisthopterus tropicus (Hildebrand, 1946) (Sardinela pelada, Tropical longfin herring) Hydrographical zone: Bahía Jiquilisco Department: Usulután Odontognathus panamensis (Steindachner, 1876) (Sardina machete panameña, Panama longfin herring) Hydrographical zone: Sirama Department: La Unión ORDER CHARACIFORMES Family Characidae Primary Astyanax aeneus (Günther, 1860) (Plateada, Banded tetra) Hydrographical zones: Río Lempa, Río Paz, Cara Sucia San Pedro, Mandinga-Comalapa, Río Jiboa, Río Grande de San Miguel Departments: Ahuachapán, La Libertad, La Paz, Usulután, San Miguel, Morazán, Santa Ana, Chalatenango Roeboides bouchellei Fowler, 1923 (Alma seca, Crystal tetra) Hydrographical zones: Río Lempa, Río Grande de San Miguel Departments: La Libertad, San Miguel, Santa Ana ORDER SILURIFORMES Family Heptapteridae Primary Rhamdia guatemalensis (Günther, 1864) (Filin/Juilin, Guatemalan chulin) Hydrographical zones: Río Lempa, Río Grande de San Miguel Departments: La Libertad, San Salvador, San Miguel, Morazán, Santa Ana, Chalatenango, Cuscatlán Rhamdia laticauda (Kner, 1858) (Filin, Filespine chulín) Hydrographical zone: Río Lempa Department: San Miguel, Santa Ana Family Ariidae Peripheral Ariopsis guatemalensis (Günther, 1864) (Bagre cuatete, Blue sea catfish) 444 · Zootaxa 3608 (6) © 2013 Magnolia Press MCMAHAN ET AL. Hydrographical zones: Río Lempa, Río Grande de San Miguel Departments: San Vicente, Usulután, San Miguel, Santa Ana, Cuscatlán Ariopsis seemanni (Günther, 1864) (Bagre tete, Tete sea catfish) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Bagre panamensis (Gill, 1863) (Bagre chihuil, Chihuil sea catfish) Hydrographical zones: Río Lempa, Mandinga-Comalapa, Bahía Jiquilisco Departments: La Libertad, Usulután, Santa Ana Cathorops raredonae Marceniuk, Betancur-R., & Acero, 2009 (Bagre curadora, Curator’s sea catfish) Hydrographical zone: Bahía Jiquilisco Department: Usulután Cathorops steindachneri (Gilbert & Starks, 1904) (Congo baboso, Steindachner’s sea catfish) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Notarius kessleri (Steindachner, 1877) (Bagre esculpido, Sculptured sea catfish) Hydrographical zone: Sirama Department: La Unión Notarius troschelii (Gill, 1863) (Bagre chili, Chili sea catfish) Hydrographical zone: Sirama Department: La Unión ORDER BATRACHOIDIFORMES Family Batrachoididae Peripheral Batrachoides waltersi Collette & Russo, 1981 (Sapo peludo, Multipored toadfish) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión ORDER MUGILIFORMES Family Mugilidae Peripheral Agonostomus monticola (Bancroft, 1834) (Tepemechin, Mountain mullet) Hydrographical zones: Río Grande de Sonsonate, Mandiga Comalapa, Río Jiboa Departments: Sonsonate, La Libertad, La Paz CHECKLIST OF EL SALVADOR FRESHWATER FISHES Zootaxa 3608 (6) © 2013 Magnolia Press · 445 Mugil curema Valenciennes, 1836 (Chimbera, White mullet) Hydrographical zones: Mandinga-Comalapa, Bahía Jiquilisco, Sirama Departments: La Libertad, Usulután, La Unión Mugil cephalus Linnaeus, 1758 (Lisa, Flathead grey mullet) Hydrographical zone: Río Paz Department: Ahuachapán Mugil hospes Jordan & Culver, 1895 (Lisa hospe, Hospe mullet) Hydrographical zone: Sirama Department: La Unión ORDER ATHERINIFORMES Family Atherinopsidae Peripheral Atherinella argentea Chernoff, 1986 (Pejerrey argénteo, Moon silverside) Hydrographical zones: Río Lempa, Bahía Jiquilisco, Lago Coatepeque Departments: San Salvador, Usulután, Santa Ana Atherinella guatemalensis (Günther, 1864) (Ejote guatemalteco, Guatemala silverside) Hydrographical zone: Bahía Jiquilisco Department: Usulután Atherinella guija (Hildebrand, 1925) (Ejote, Guija silverside) Hydrographical zones: Río Lempa (including Lago Güija) Departments: San Vicente, Santa Ana Atherinella pachylepis (Günther, 1864) (Pejerrey alón, Thickscale silverside) Hydrographical zones: Sirama Department: La Unión ORDER BELONIFORMES Family Hemiramphidae Peripheral Hyporhamphus unifasciatus (Ranzani, 1841) (Pajarito, Common halfbeak) Hydrographical zone: Sirama Department: La Unión ORDER CYPRINODONTIFORMES 446 · Zootaxa 3608 (6) © 2013 Magnolia Press MCMAHAN ET AL. Family Profundulidae Secondary Profundulus guatemalensis (Günther, 1866) (Escamudo guatemalteco, Guatemalan killifish) Hydrographical zone: Río Paz Department: Ahuachahapán Profundulus kreiseri Matamoros, Schaefer, Hernández, & Chakrabarty, 2012 (Escamudo de Kreiser, Kreiser’s killifish) Hydrographical zone: Río Lempa Department: Santa Ana Family Anablepidae Secondary Anableps dowei Gill, 1861 (Cuatro ojos, Pacific foureyed fish) Hydrographical zones: Río Lempa, Río Paz, Río Grande de San Miguel, Sirama Departments: Ahuachapán, La Libertad, San Vicente, Usulután, San Miguel, La Unión, Santa Ana, Cuscatlán Family Poeciliidae Secondary Heterandria anzuetoi Rosen & Bailey, 1979 (Chimbolo, Anzueto’s killifish) Hydrographical zone: Río Lempa Departments: Cabañas, Cuscatlán Poecilia butleri Jordan, 1889 (Topote del Pacífico, Pacific molly) Hydrographical zone: Río Jiboa Department: La Paz Poecilia gillii (Kner, 1863) (Olomina, Gill’s molly) Hydrographical zones: Río Lempa, Cara Sucia San Pedro Departments: Ahuachapán, La Libertad, San Salvador, La Paz, San Miguel, Morazán, Santa Ana Poecilia marcellinoi Poeser, 1995 (Topote de Ilopango, Ilopango molly) Hydrographical zones: Río Lempa, Grande de Sonsonate, Mandinga-Comalapa Departments: Sonsonate, La Libertad, Santa Ana Poecilia salvatoris Regan, 1907 (Topote de El Salvador, Salvadoran molly) Hydrographical zones: Río Lempa, Río Paz, Grande de Sonsonate Departments: Ahuachapán, Sonsonate, San Salvador, Santa Ana, Chalatenango Poecilia sphenops Valenciennes, 1846 (Topote mexicano, Mexican Molly) Hydrographical zones: Río Lempa, Río Paz, Mandinga-Comalapa, Río Grande de San Miguel Departments: Ahuachahapán, La Libertad, San Salvador, Usulután, San Miguel, Santa Ana, Cuscatlán CHECKLIST OF EL SALVADOR FRESHWATER FISHES Zootaxa 3608 (6) © 2013 Magnolia Press · 447 Poeciliopsis pleurospilus (Günther, 1866) (Guatopote manchote, Largespot livebearer) Hydrographical zones: Río Lempa, Río Paz, Grande de Sonsonate, Mandinga-Comalapa, Río Jiboa, Río Grande de San Miguel, Lago Coatepeque Departments: Ahuachapán, Sonsonate, La Libertad, San Salvador, La Paz, San Miguel, Morazán, Santa Ana, Chalatenango, Cuscatlán Poeciliopsis turrubarensis (Meek, 1912) (Guatopote del Pacífico, Barred livebearer) Hydrographical zones: Río Lempa, Bahía Jiquilisco Departments: San Vicente, Usulután ORDER GASTEROSTEIFORMES Family Syngnathidae Peripheral Pseudophallus starksii (Jordan & Culver, 1895) (Pipeta/Falso Caballo, Yellowbelly pipefish) Hydrographical zones: Mandinga-Comalapa, Sirama Departments: La Libertad, La Unión ORDER SYNBRANCHIFORMES Family Synbranchidae Secondary Synbranchus marmoratus Bloch, 1795 (Falsa anguila/Anguila, Marbled swamp eel) Hydrographical zones: Río Lempa, Río Grande de San Miguel, Sirama Departments: Morazán, La Unión ORDER PERCIFORMES Family Centropomidae Peripheral Centropomus armatus Gill, 1863 (Robalo espina larga, Longspine snook) Hydrographical zones: Cara Sucia San Pedro, Bahía Jiquilisco, Sirama Departments: Ahuacahpán, Usulután, La Unión Centropomus medius Günther, 1864 (Robalo aleta prieta, Blackfin snook) Hydrographical zones: Río Lempa, Sirama Departments: Usulután, La Unión Centropomus nigrescens Günther, 1864 (Robalo negro, Black snook) Hydrographical zones: Río Lempa, Bahía Jiquilisco Department: Usulután 448 · Zootaxa 3608 (6) © 2013 Magnolia Press MCMAHAN ET AL. Centropomus robalito Jordan & Gilbert, 1882 (Robalo aleta amarilla, Yellowfin snook) Hydrographical zones: Río Lempa, Bahía Jiquilisco Department: Usulután Family Carangidae Peripheral Caranx caballus Günther, 1868 (Jurel bonito, Green jack) Hydrographical zone: Bahía Jiquilisco Department: Usulután Caranx caninus Günther, 1867 (Jurel común, Pacific crevalle jack) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Chloroscombrus orqueta Jordan & Gilbert, 1883 (Casabe, Pacific bumper) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Oligoplites altus (Günther, 1868) (Zapatero sierrita, Longjaw leatherjacket) Hydrographical zone: Bahía Jiquilisco Department: Usulután Trachinotus kennedyi Steindachner, 1876 (Pámpano plateado, Blackblotch pompano) Hydrographical zone: Bahía Jiquilisco Department: Usulután Family Lutjanidae Peripheral Lutjanus argentiventris (Peters, 1869) (Pargo amarillo, Yellow snapper) Hydrographical zones: Bahía Jiquilisco, Sirama Departments: Usulután, La Unión Lutjanus colorado Jordan & Gilbert, 1882 (Pargo rojo, Colorado snapper) Hydrographical zones: Mandinga-Comalapa, Sirama Departments: La Libertad, La Unión Lutjanus guttatus (Steindachner, 1869) (Pargo lunarejo, Spotted rose snapper) Hydrographical zone: Bahía Jiquilisco Department: Usulután CHECKLIST OF EL SALVADOR FRESHWATER FISHES Zootaxa 3608 (6) © 2013 Magnolia Press · 449