Chec List Annotated Checklist of the Freshwater Ishes of Continental And

PECIES

S continental and insular Costa Rica: additions and

OF nomenclatural revisions ISTS L Arturo Angulo 1*, Carlos A. Garita-Alvarado 1, William A. Bussing 1, 2 1, 2

Rica, 11501-2060. * Corresponding author. E-mail: [email protected]

insular Costa Rica. This checklist, systematically arranged at the ordinal and familial level, includes nomenclatural revisions,

Louisiana State University Museum of Zoology (LSUMZ). Other institutional abbreviations used are as follows: CAS = California Academy of Sciences; LACM = Los Angeles County Museum; STRI = Smithsonian Tropical Research these pioneer works, knowledge about the freshwater Institute; TU = Tulane University Museum of Natural History; UF = University of Florida; and UMMZ = University of new species, the establishment of new inland and of Michigan Museum of Zoology. country records, and the numerous nomenclatural changes The checklist is arranged by order and family following for previously known species, given major advances in Eschmeyer and Fong (2013). Genera and species within a understanding of the status and relationships of many family are arranged in alphabetical order. After the mention groups, contribute to such development. The purpose of of each most inclusive taxa, the number in parentheses the following checklist is to update the known composition larger than genera the number in brackets represent the number of genera. In those higher than family the number conclusions of recent studies on relationships. preceding genera is the number of families. The family tolerance to salinity is listed according to Diversity and distributional data were obtained description of each species and genera follow Eschmeyer (2013). An asterisk after the authority name indicates that localities in Costa Rica was performed between March the taxon is introduced. When available, after the authority name, the common English name was provided followed by the common Spanish name, both following Bussing Pauly (2013) (see Angulo (2013) for a more complete list were issued by the Costa Rican Ministerio de Ambiente of the common and technical Spanish names of the Costa Energía y Telecomunicaciones (MINAET) and the Sistema following the common names indicates the page location Distributional data are presented at the Costa Rican Museo de Zoología of the Universidad de Costa Rica major River drainage basins (Figure 1), for which there are (UCR) was conducted. Finally, species occurrence records were also obtained by querying the inter-institutional drainages, all listed in north to south order). Following the distributional data, altitudinal ranges in meters supplement the data obtained by the above sources. above sea level are also presented. For range expansions Specimens preserved were deposited at UCR and

987 Angulo et al. | Freshwater !shes of Costa Rica

the most speciose families. By far, the vast majority of museum specimens that are associated with the species of interest. Finally, comments are provided for species supposed tolerance to salinity, are peripheral (63.2%), where annotations are necessary in the remarks section followed by secondary (23.6%); only 13.2% are primary (e.g. nomenclatural notes, range extensions, conservation or obligate freshwater. status, etc.). peripheral species, two (Ophisternon aenigmaticum and Poeciliopsis santaelena) are secondary species and only Bryconamericus gonzalezi is a primary species. Twenty-four species appear to be endemic to the country (Amphilophus bussingi, A. diquis, Brachyrhaphis olomina, B. rhabdophora, Bryconamericus terrabensis, Cryptoheros myrnae, C. sajica, Cynodonichthys fuscolineatus, C. glaucus, C. siegfriedi, species), Poeciliidae (23 species), Gobiidae (22 species), Eleotris tubularis, Gobiesox fulvus, Hyphessobrycon

(TO), Parismina (PA), Matina (MA), Sixaola (SI), Isla del Coco (IC), Nicoya (NI), Tempisque (TE), Bebedero (BE), Barranca (BA), Tárcoles (TA), Pirrís (PI),

988 Angulo et al. | Freshwater !shes of Costa Rica quadripunctatus, Poeciliopsis paucimaculata, P. santaelena, Atractosteus tropicus Priapichthys annectens, Pseudocheirodon Térrabae, Pterobrycon myrnae, Roeboides ilseae, Sicydium adelum Distribution: Atlantic slope: Lago de Nicaragua, Río and S. cocoensis). Finally, in addition to the native species, Carassius auratus, Cyprinus carpio, Parismina. Micropterus salmoides, Oncorhynchus mykiss, Oreochromis niloticus, Poecilia reticulata, Pterygoplichthys pardalis and Xiphophorus variatus), included in eigth genera, three families and two orders, are also found in the country. Peripheral.

Elops 5. Addition to fauna. Peripheral. 2011). Eight specimens have been collected in freshwaters Carcharhinus Peripheral. 1. Carcharhinus leucas Megalops Distribution: Atlantic slope: Lago de Nicaragua, Río 6. Megalops atlanticus sábalo real (61). Distribution: Atlantic slope: Lago de Nicaragua, Río m. Peripheral. Pristis Peripheral. 2. Pristis pectinata Albula Distribution: Atlantic slope: Lago de Nicaragua, San Albula vulpes Remarks: This species has been reported in the Addition to fauna. Caribbean coast of Costa Rica by Distribution: Atlantic slope: Matina. 0-2 m. Remarks: This species has been reported in the Nicaragua, and its major branch, the Colorado River, Costa 2010). Four specimens have been collected in freshwaters Rica. Other freshwater records in Caribbean drainages River, Guatemala (Thorson et al River, Honduras (Matamoros et al Peripheral. Anguilla 3. Pristis pristis Anguilla rostrata Distribution: Atlantic slope: Lago de Nicaragua, Río Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Matina and Sixaola. 0-20 m. Peripheral. Myrophis Secondary. Myrophis punctatus Atractosteus eel, tieso gusano. Addition to fauna.

989 Angulo et al. | Freshwater !shes of Costa Rica

Distribution: Atlantic slope: Matina. 0-3 m. drainage. Remarks: This species has been reported in the Peripheral. Anchoa 15. Anchoa curta 10. Myrophis vafer anchovy, anchoa pelada. Addition to fauna. 2011). One specimen has been collected in freshwaters in the Palo Seco River and the Quebrada Vueltas River, drainage. 16. Anchoa lucida anchovy, anchoa bocona. Addition to fauna. Peripheral. Dorosoma 2011). One specimen has been collected in freshwaters in 11. Dorosoma chavesi Anchovia Distribution: Atlantic slope: Lago de Nicaragua, Río Anchovia macrolepidota m. anchovy, anchoa de escama grande. Addition to fauna. Lile 12. (Schreiner and Miranda Ribeiro, 2011). One specimen has been collected in freshwaters Atlantic piquitinga, sardina piquitinga. Addition to fauna. drainage. Distribution: Atlantic slope: Sixaola. 0-66 m. Remarks: One specimen has been collected in Anchoviella freshwaters in the Quebrada Biei River (UCR 1531-10), Sixaola drainage. This species is known only from Nueva Anchoviella elongata Elongate anchovy, anchoveta alargada. Addition to fauna. and Lake Papary near Natal south to Bahia and Espirito Distribution: Atlantic slope: Tortuguero and Parismina. 0-5 m. north range extension of the known distribution in the Remarks: This species has been reported in the western Atlantic. 2010). Ten specimens have been collected in freshwaters 13. herring, sardina rayada. Addition to fauna. 2011). Forty-eight specimens have been collected in the Tortuguero Lagoon, Tortuguero drainage. Lycengraulis Atlantic Sardinella sabretooth anchovy, anchoa dientona. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. Sardinella brasiliensis Remarks: This species has been reported in the Distribution: Atlantic slope: Tortuguero. 0-3 m. 2010). One specimen has been collected in freshwaters Remarks: This species has been reported in the drainage. 2010). Eight specimens have been collected in freshwaters

990 Angulo et al. | Freshwater !shes of Costa Rica

Primary. 25. Astyanax aeneus Carassius Distribution: Atlantic slope: Lago de Nicaragua, 20. Carassius auratus carpa dorada (23). and Coto. 0-1000 m. Remarks: Originally introduced from Asia, this species 26. Astyanax cocibolca ; heard of specimens reaching other waters. Distribution: Atlantic slope: Lago de Nicaragua, Río Cyprinus m. A. cocibolca from 21. Cyprinus carpio that he previously considered as A. nasutus. This species Taiwan in an attempt to strengthen aquaculture activities Astyanax orthodus freshwaters in an unknown pond name in Alajuela (UCR Distribution: Atlantic slope: Sixaola. 1-60 m.

Bramocharax

Primary. Bramocharax bransfordii Cyphocharax Distribution: Atlantic slope: Lago de Nicaragua, Río 22. Cyphocharax magdalenae Parismina. 5-530 m. Carp headstander, capani (126). Bramocharax to be monophyletic and derived from some Astyanax species. On the other hand, Ornelas-García et al Primary. that this genus is not monophyletic and considered all Bramocharax species as morphotypes of Astyanax. The Hoplias close relationship between Bramocharax and Astyanax proposed by Ornelas-García et al 23. Hoplias microlepis by Mirande (2010), which instead relates Bramocharax perro (123). with Oligosarcus provisionally keep this species in Bramocharax following et al. (2003) and Primary. Mirande (2010); however, a comprehensive taxonomic study is needed to test the monophyly and position of this Lebiasina clade.

Brycon Brycon behreae Remarks: This species was called Piabucina boruca in species in the genus following Eigenmann and 30. Brycon costaricensis Angulo and Gracian-Negrete, et al. (2011). This species Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Matina. 0-600 m. Remarks: Angulo and Gracian-Negrete (2013) Primary. described B. costaricensis from specimens of the Atlantic Astyanax considered as B. guatemalensis.

991 Angulo et al. | Freshwater !shes of Costa Rica

Bryconamericus Remarks: This species is endemic to the Pirrís and 31. Bryconamericus gonzalezoi Román-Valencia 2002. Creek tetra, sardina de quebrada. Addition to fauna. Pterobrycon Pterobrycon myrnae 32. Bryconamericus scleroparius Creek tetra, sardinita (113). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Remarks: This species is endemic to the Pirrís and Roeboides 33. Bryconamericus terrabensis Roeboides bouchellei headstander, sardina (116). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, 2002). Barranca. 0-610 m. Carlana Roeboides bussingi Matamoros, Chakrabarty, Carlana eigenmanni Angulo, Garita-Alvarado and McMahan, 2013. Glass sardinita (100). Distribution: Atlantic slope: Lago de Nicaragua, Río Remarks: Matamoros et al. (2013) described R. bussingi R. sp. Hyphessobrycon Roeboides ilseae 35. Hyphessobrycon panamensis sardina (120). tetra, sardinita (105). m. Remarks: This species is endemic to the Pirrís and 36. Hyphessobrycon savagei Remarks: This species is endemic to the Pirrís and Peripheral.

Hyphessobrycon tortuguerae Cathorops Tortuguero tetra, sardinita de Tortuguero (110). Distribution: Atlantic slope: Lago de Nicaragua, Río Cathorops steindachneri m. fauna. Odontostilbe 0-55 m. Odontostilbe dialeptura Pygmy tetra, sardinita (103). One specimen has been collected in freshwaters in the Remarks: This species was called Compsura dialeptura Odontostilbe following Malabarba drainage. however a new generic assignment is necessary for this species. Cathorops tuyra Pseudocheirodon 0-55 m. Pseudocheirodon Térrabae tetra, sardinita (111). 2011). Two specimens have been collected in freshwaters

992 Angulo et al. | Freshwater !shes of Costa Rica

Pimelodella Notarius 51. Pimelodella chagresi Notarius cookei (Acero and Betancur-R, 2002). Sea Rhamdia coast of Costa Rica by as Arius cookei 52. Rhamdia guatemalensis species in the genus Notarius following Bentancur-R. and Distribution: Atlantic slope: Lago de Nicaragua, Marceniuk et al. (2012). One specimen has been collected Tempisque drainage. Sciades Costa Rican members of this genus were referred to the genus R. guatemalensis with R. quelen. On the other Arius hand, Perdices et al. history of the genus in Central America and concluded in the genus Sciades following Marceniuk et al. (2012). that Central American R. guatemalensis are evolutionarily distinct from R. quelen from South America. In the present Sciades assimilis bagre maya. Addition to fauna. from R. quelen et al. (2002), Weber et al. (2003), Remarks: This species has been reported in the Sharpf (2006) and Matamoros et al. 53. Rhamdia laticauda Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Sciades guatemalensis Remarks: This species was called Rhamdia rogersi Sciades seemanni R. laticauda et al. Rhamdia nicaraguensis 0-32 m. barbudo (152). Distribution: Atlantic slope: Lago de Nicaragua, Río Primary. Costa Rican members of this family were referred to the family and Tárcoles. 20-1160 m. Primary. (2006). Trichomycterus 55. Trichomycterus striatus (Meek and Hidelbrand, 50. Remarks: This species was called Nannorhamdia Primary. lineata following Fonchiiichthys et al. 56. Fonchiiichthys uracanthus

993 Angulo et al. | Freshwater !shes of Costa Rica

Remarks: This species was called Rineloricaria Brachyhypopomus uracantha 61. Brachyhypopomus occidentalis Fonchiiichthys following Isbrücker (2001; 2002), Fisch- Distribution: Atlantic slope: Sixaola. 1-60 m. Remarks: This species was called Hypopomus Hemiancistrus occidentalis Brachyhypopomus Hemiancistrus aspidolepis suckermouth, arrisuaca (160). 01) and Parismina (Molina et al. 2010, as Hypostomus panamensis Peripheral. Remarks: This species was called Hypostomus panamensis Oncorhynchus Hemiancistrus as synonym of H. aspidolepis following 62. Oncorhynchus mykiss Isbrücker (2001; 2002), Fisch-Muller (2003) and Ferraris trout, trucha arcoiris (22). Distribution: Atlantic slope: Parismina (A. Molina, pers. Pterygoplichthys Remarks: This species was introduced in Costa Rica Pterygoplichthys pardalis Distribution: Parismina. 20-110 m. Remarks: One specimen has been collected in Parismina drainage. Molina et al. (2010), as P. sp., and Herrera (2012) recorded this species in freshwaters in the Lagoon, Parismina drainage. This species naturally occurs Gobiesox to several countries around the world. In Costa Rica this 63. Gobiesox fulvus species has been introduced most likely by aquarists chupapiedra (250). (Molina et al. 2010; Herrera 2012). Remarks: This species is endemic to the Isla del Coco Primary. Gobiesox nudus Gymnotus Distribution: Atlantic slope: San Carlos, Sarapiquí, Gymnotus cylindricus cuchillo (130). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, 65. Gobiesox potamius Matina and Sixaola. 1-50 m. 60. Gymnotus maculosus Remarks: This species was thought to be endemic to the Nicoya, Tempisque, Bebedero, Barranca, Tárcoles, Distribution: Atlantic slope: Lago de Nicaragua, Río however, McMahan et al. (2013) report this in El Salvador. Primary. Brachyhypopomus occidentalis Peripheral. Costa species of this family know to be present in Costa Rica, Rican members of this family were referred to the family was referred to the family Rhamphichthyidae by Bussing following Saeed et al.

994 Angulo et al. | Freshwater !shes of Costa Rica and Nelson (2006). Rivulus to two endemic species to Cuba and considered as valid genera Anablepsoides, Atlantirivulus, , Atherinella Melanorivulus and Cynodonichthys, subgenera of Rivulus. In the present account we placed in 66. Atherinella argentea Cynodonichthys all Costa Rican rivulids species following this author. Cynodonichthys fuscolineatus m. Atherinella blackburni Remarks: This species is endemic to the San Carlos and Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the Cynodonichthys glaucus 2010). One specimen has been collected in freshwaters drainage. m. Atherinella chagresi Distribution: Atlantic slope: Tortuguero, Parismina, Cynodonichthys hildebrandi Atherinella hubbsi Cynodonichthys isthmensis Distribution: Atlantic slope: Lago de Nicaragua, Río Distribution: Atlantic slope: Lago de Nicaragua, Parismina, Matina and Sixaola. Atherinella milleri Tárcoles. 0-1500 m. Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Cynodonichthys rubripunctatus Atherinella sardina of C. isthmensis . In the present 2011). Membras Cynodonichthys siegfriedi Membras gilberti Landia silverside, pejerrey landia. Addition to fauna. of C. . In the present 2011). Twenty-nine specimens have been collected in drainage. m. Secondary. Oxyzygonectes Cynodonichthys Costa Rican members of this genus were referred to the genus Rivulus Oxyzygonectes dovii ojos blancos (166). and morphological data of 33 rivuline taxa, restricted

995 Angulo et al. | Freshwater !shes of Costa Rica

Brachyrhaphis parismina 0-15 m. (200). Distribution: Atlantic slope: Lago de Nicaragua, Rio Frio, Matina. 5-50 m. Alfaro Brachyrhaphis rhabdophora Alfaro cultratus (203) Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Remarks: This species is endemic to the Pirrís and

Belonesox Brachyrhaphis roseniOlomina (205) Distribution: Belonesox belizanus Brachyrhaphis terrabensisOlomina Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Distribution: -01), Matina. -01).

Brachyrhaphis Gambusia

Brachyrhaphis cascajalensis (Meek and Hildebrand, Gambusia nicaraguensis Olomina. Addition to fauna. Distribution: Atlantic slope: Sixaola. 5-125 m. Distribution: Atlantic slope: Matina. 0-15 m. Remarks: from Costa Rica, although he mentions that all specimens Phallichthys from the Sixaola drainage lack of a caudal black spot (apparently the unique feature present in B. cascajalensis Phallichthys amates that separate this of B. parismina). In the present account, olomina (213). Distribution: Atlantic slope: Lago de Nicaragua, et alet al (2003) and Lucinda and Reis (2005) and on the basis of Bebedero and the Sixaola drainage; however, further studies are needed Phallichthys quadripunctatus Olomina (216). Distribution: Brachyrhaphis holdridgei Remarks: This species is endemic to the Sixaola drainage Distribution: Atlantic slope: Lago Nicaragua, Rio Frío, Phallichthys tico 35-620 m. Distribution: Atlantic slope: Lago de Nicaragua, San Brachyrhaphis olomina Poecilia Distribution: Atlantic slope: Lago de Nicaragua and Poecilia gillii Barranca and Tárcoles. Remarks: et al Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, B. olomina as synonym of B. rhabdophora B. olomina as valid and different from B. rhabdophora, m. since the two species can be completely separated based on the number of gonopodial subterminal spines in ray Poecilia mexicana This species is endemic molly, olomina (222). to the Lago de Nicaragua, Parismina, Nicoya, Tempisque, Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Bebedero, Barranca and Tárcoles drainages, Costa Rica Matina.

996 Angulo et al. | Freshwater !shes of Costa Rica

Poecilia reticulata Guppy, olomina (23). Remarks: This species naturally occurs in Barbados, Xenophallus though it has been widely introduced to several countries Xenophallus umbratilis around the world. In Costa Rica this species has been (210) Distribution: Atlantic slope: Lago de Nicaragua, Río et al. et al. Remarks: This species was called Neoheterandria umbratilis Poeciliopsis Xenophallus following Poeciliopsis elongata toothcarp, olomina (223). Xiphophorus 105. Xiphophorus variatus Variable , platí (23). Poeciliopsis paucimaculata Remarks: This species naturally occurs in Mexico, (225). though it has been widely introduced to several countries around the world. In Costa Rica this species has been Remarks: Coto drainages Poeciliopsis retropinna Strongylura 100. Poeciliopsis santaelena Addition to fauna. 106. Strongylura marina Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the Remarks: This species is endemic to the Potrero Grande 2010). Seven specimens have been collected in freshwaters River, Peninsula de Santa Elena, Nicoya drainage, Costa

101. Poeciliopsis turrubarensis Strongylura timucu Distribution: Nicoya, Tempisque, Bebedero, Barranca, Distribution: Atlantic slope: Tortuguero and Matina. 0-13 m. Remarks: This species has been reported in the Priapichthys 2010). Five specimens have been collected in freshwaters 102. Priapichthys annectens (230). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, in freshwaters in the Tortuguero Lagoon, Tortuguero drainage. Remarks: This species is endemic to the Lago de Peripheral. Tortuguero, Parismina, Matina, Sixaola, Nicoya, Tempisque, Hemiramphus Bebedero, Barranca and Tárcoles drainages, Costa Rica Hemiramphus brasiliensis 103. Priapichthys panamensis Meek and Hildebrand, Distribution: Atlantic slope: Tortuguero. 0-2 m.

997 Angulo et al. | Freshwater !shes of Costa Rica

Remarks: This species has been reported in the 2010). One specimen has been collected in freshwaters 2010). One specimen has been collected in freshwaters in drainage. Hyporhamphus Secondary. Hyporhamphus roberti Synbranchus Distribution: Atlantic slope: Tortuguero. 0-2 m. Remarks: This species has been reported in the 115. Synbranchus marmoratus swamp eel, anguila de pantano (260). 2010). One specimen has been collected in freshwaters in Distribution: Atlantic slope: Lago de Nicaragua, Peripheral. Ophisternon Microphis 116. Ophisternon aenigmaticum Rosen and Greenwood, 110. Microphis lineatus Addition to fauna. Distribution: Atlantic slope: Tortuguero (Gilbert and Oostethus lineatus O. aenigmaticum may occur in et al. (2006) collected six specimens in Rica. of Oostethus brachyurus the genus Microphis following Ferraris (2003) and Kuiter Peripheral. Centropomus Pseudophallus Centropomus armatus 111. Pseudophallus elcapitanensis (Meek and Remarks: This species has been reported in the 0-20 m. 112. Pseudophallus mindi (Meek and Hidelbrand, drainage. Distribution: Atlantic slope: Sarapiquí, Tortuguero, Centropomus ensiferus Parismina, Matina and Sixaola. 0-35 m. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, 113. Pseudophallus starksi Matina and Sixaola. 0-11 m. Remarks: This species has been reported in the 2010). Four specimens have been collected in freshwaters 0-155 m. Syngnathus

Syngnathus scovelli (Evermann and Kendall, Centropomus medius snook, gualaje aleta manchada (265). Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the 0-250 m.

998 Angulo et al. | Freshwater !shes of Costa Rica

120. Centropomus mexicanusLargescale Distribution: Atlantic slope: Tortuguero, Parismina, 2011). One specimen has been collected in freshwaters Remarks: This species has been reported in the 2010). Four specimens have been collected in freshwaters n=3), Tortuguero drainage. Centropomus unionensis snook, mano de piedra. Addition to fauna. 121. Centropomus nigrescens Nine specimens have been collected in freshwaters in the Sixty specimens have been collected in freshwaters in the Peripheral. Epinephelus Maria River (UCR 1150-03, n=3), Tárcoles drainage, the Epinephelus guttatus cabrilla colorada. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the 2010). One specimen has been collected in freshwaters in the Tortuguero Lagoon (UCR 0261-10), Tortuguero drainage. et al. Micropterus Micropterus salmoides Largemouth black bass, black-bass (23). 122. Centropomus parallelus Remarks: This species was introduced in Costa Rica Carlos, Sarapiquí, Tortuguero, Parismina, Matina and heard of specimens reaching other waters. 123. Centropomus pectinatus Peripheral. Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-12 m. Carangoides

Centropomus robalito Carangoides vinctus Cocinero, platanillo. Addition to fauna. 0-50 m. 2011 as Caranx vinctusCaranx vinctus, 125. Centropomus undecimalis María River, Tárcoles drainage. In the present account, we Distribution: Atlantic slope: Tortuguero, Parismina, Carangoides following Matina and Sixaola. 0-50 m. Castro-Aguirre et al. 126. Centropomus viridis robalo blanco. Addition to fauna. Caranx 130. Caranx caballus

999 Angulo et al. | Freshwater !shes of Costa Rica

Addition to fauna. 0-10 m. Remarks: This species has been reported in the Oligoplites saurus and Caribbean coasts of Costa Rica by ). Three specimens have been collected 131. Caranx caninus jack, jurel. Addition to fauna. Tortuguero drainage. 0-55 m. Lutjanus pargo criollo. Addition to fauna. Distribution: Atlantic slope: Tortuguero and Matina. 0-3 m. Remarks: This species has been reported in the 132. Caranx hippos jurel. Addition to fauna. 2010). Four specimens have been collected in freshwaters Distribution: Atlantic slope: Tortuguero and Matina. in the Tortuguero Lagoon (UCR 0261-03, n=3), Tortuguero 0-5 m. Remarks: This species has been reported in the drainage. n=1), Tortuguero drainage, and the Quebrada Westfalia 0-20 m. 133. Caranx latus Colorado snapper, pargo colorado. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-20 m. Remarks: This species has been reported in the Hemicaranx freshwaters in an unknown river name in the Cocos 135. Hemicaranx amblyrhynchus Bluntnose jack, jurelito chato. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the 2010). One specimen has been collected in freshwaters in the Quebrada Camaronal River, the Pavo River and the species in freshwaters in the Paquita River, Pirrís drainage. Oligoplites 136. Oligoplites altus pargo mulato. Addition to fauna. leatherjacket, cuero. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the 2010). Two specimens have been collected in freshwaters 2011). Six specimens have been collected in freshwaters drainage. Oligoplites palometa snapper, pargo caballera. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, Distribution: Atlantic slope: Sarapiquí and Tortuguero.

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Remarks: This species has been reported in the recorded this species in freshwaters in the Hatillo Nuevo 2010). Ten specimens have been collected in freshwaters River, Pirrís drainage. Diapterus rhombeus mojarra, mojarra de estero. Addition to fauna. Distribution: Atlantic slope: Tortuguero and Parismina. 0-12 m. Remarks: This species has been reported in the the Tortuguero Lagoon, Tortuguero drainage. 2010). Five specimens have been collected in freshwaters Addition to fauna. Remarks: This species has been reported in the Parismina drainage. Eucinostomus Eucinostomus argenteus Silver mojarra, mojarra plateada. Addition to fauna. Distribution: Atlantic slope: Matina. 0-6 m. snapper, pargo negro. Addition to fauna. Remarks: This species has been reported in the 2010). Two specimens have been collected in freshwaters 150. Eucinostomus currani Tárcoles drainage, the Quebrada Aserradero River (UCR 151. Eucinostomus dowii palmito plateado. Addition to fauna. coast of Costa Rica by both as E. argenteus). One specimen has been collected Diapterus as valid and distinct from E. argenteus following Allen and Diapterus auratus et al mojarra gaucha. Addition to fauna. et al Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the 152. Eucinostomus melanopterus Addition to fauna. 2010). Seven specimens have been collected in freshwaters Distribution: Atlantic slope: Tortuguero, Parismina and Matina. 0-10 m. Remarks: This species has been reported in the Diapterus peruvianus mojarra, pargo blanco. Addition to fauna. Remarks: This species has been reported in the Hone Creek River (UCR 1352-11, n=11), Matina drainage. Eugerres

1001 Angulo et al. | Freshwater !shes of Costa Rica

153. Eugerres brevimanus drainage. Haemulopsis leuciscus grunt, vieja chinilla. Addition to fauna. Remarks: This species has been reported in the Eugerres plumieri Sarapiquí and Tortuguero. 0-31 m.

Gerres Pomadasys

155. Gerres cinereus 160. Pomadasys bayanus mojarra, mojarra trompetera. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-12 m. Remarks: This species has been reported in the 2010). Two specimens have been collected in freshwaters 161. Pomadasys branickii Sand grunt, burro. Addition to fauna. Remarks: This species has been reported in the 156. Gerres simillimus mojarra, palmito blanco. Addition to fauna. drainage. coast of Costa Rica by as G. cinereus). Seventeen specimens have been collected in 162. Pomadasys crocro 21, n=1), Pirrís drainage, and an unknown river name in Distribution: Atlantic slope: Sarapiquí, Tortuguero, as G. cinereus, recorded this species in freshwaters in the Nosara River, Nicoya drainage, and the 163. Pomadasys empherus grunt, roncador. Addition to fauna. G. cinereus coast of Costa Rica by ). One Peripheral. specimen has been collected in freshwaters in the Lagarto Conodon Pomadasys macracanthus Conodon nobilis Longspine grunt, vieja espinalarga. Addition to fauna. ronco canario. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-51 m. Remarks: This species has been reported in the 2011). Three specimens have been collected in freshwaters 2010). Seven specimens have been collected in freshwaters Peripheral. drainage. Bairdiella

Haemulopsis 165. Bairdiella ronchus ronco rayado. Addition to fauna. Haemulopsis corvinaeformis Distribution: Atlantic slope: Tortuguero. 0-5 m. Roughneck grunt, corocoro gris. Addition to fauna. Remarks: This species has been reported in the Distribution: Atlantic slope: Tortuguero. 0-5 m. Remarks: This species has been reported in the Caribbean coast of Costa Rica by 2010). Two specimens have been collected in freshwaters

1002 Angulo et al. | Freshwater !shes of Costa Rica three specimens in freshwaters in the Tortuguero Lagoon, Tortuguero drainage. drainage.

Cynoscion Peripheral.

166. Cynoscion albus Kyphosus Kyphosus sectatrix chub, chopa blanca. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. 2011). Four specimens have been collected in freshwaters Remarks: This species has been reported in the 2010). One specimen has been collected in freshwaters Larimus drainage. Shorthead drum, Peripheral. Distribution: Atlantic slope: Tortuguero. 0-3 m. Remarks: This species has been reported in the Agonostomus 2010). One specimen has been collected in freshwaters in Agonostomus monticola drainage. Distribution: Atlantic slope: Lago de Nicaragua, Micropogonias Nicoya, Tempisque, Bebedero, Barranca, Tárcoles, Pirrís, Micropogonias furnieri Distribution: Atlantic slope: Tortuguero and Parismina. Joturus 0-3 m. Remarks: This species has been reported in the Joturus pichardi 2010). Six specimens have been collected in freshwaters Distribution: Atlantic slope: Sarapiquí, Tortuguero, Parismina, Matina and Sixaola. 0-600 m. Mugil Parismina drainage. Mugil cephalus Ophioscion Distribution: Atlantic slope: Tortuguero. 0-3 m. Ophioscion typicus Remarks: This species has been reported in the china corredora. Addition to fauna. Bussing and ). Two specimens have been collected Tortuguero drainage. 2011). Two specimens have been collected in freshwaters Mugil curema lisa blanca. Addition to fauna. Peripheral. Distribution: Atlantic slope: Tortuguero, Parismina, Polydactylus Polydactylus virginicus Remarks: This species has been reported in the Addition to fauna. Bussing Distribution: Atlantic slope: Tortuguero, Parismina and ). Three hundred eighty- Matina. 0-6 m. seven specimens have been collected in freshwaters in the Remarks: This species has been reported in the Matina drainage, an unknown river name in Gandoca (UCR

1003 Angulo et al. | Freshwater !shes of Costa Rica

drainage Amphilophus citrinellus cichlid, mojarra (312). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Matina. 0-12 m. Amphilophus diquis in the Quebrada Camaronal River, the Pavo River and the this species in freshwaters in the Guacimal River, Barranca and Coto drainages Amphilophus longimanus drainage. Distribution: Atlantic slope: Lago de Nicaragua, Río Secondary. slope: Nicoya, Tempisque and Bebedero. 0-100 m. Amatitlania described A. kanna, from Atlantic Amphilophus lyonsi slope of Costa Rica and Panama, and A. siquia, from Honduras, Nicaragua and Costa Rica, from specimens that Archocentrus nigrofasciatus Amphilophus rhytisma cichlid, mojarra (332). Amatitlania kanna . Convict Amphilophus rostratus Distribution: Atlantic slope: Sixaola. 0-65 m. Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Amatitlania siquia Matina. 0-200 m. Distribution: Atlantic slope: Lago de Nicaragua, Río Andinoacara Barranca and Tárcoles. Andinoacara coeruleopunctatus Amphilophus Costa Rican Mojarra (310). members of this genus, with exception of A. citrinellus and A. lyonsi, were referred to the genus Astatheros by Bussing Remarks: This species was called Aequides coeruleopunctatus in the genus Amphilophus all Costa Rican Astatheros species account we this species in the genus Andinoacara (sensu following Musilova et al. ; however a comprensive taxomonic work is needed to test Archocentrus the monophyly and position of this group. Archocentrus centrarchus Amphilophus alfari mojarra rayada (336). Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Tárcoles. 2-1150 m. Archocentrus multispinosus Rainbow cichlid, mojarrita (351). Amphilophus altifrons Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Matina. Amphilophus bussingi Remarks: This species was called Herotilapia multispinosa Remarks: This species is endemic to the Sixaola this species in the genus Archocentrus following

1004 Angulo et al. | Freshwater !shes of Costa Rica

Negro National Wildlife Refuge, Río Frío drainage. Oro Cryptoheros Costa Rican members of this genus were referred to the genus Archocentrus Cryptoheros following Cryptoheros myrnae agencies in an attempt to strengthen aquaculture activities Remarks: This species is endemic to the Sixaola Parachromis drainage Parachromis dovii Cryptoheros sajica Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Remarks: 0-600 m. and Coto drainages Parachromis loisellei Cryptoheros septemfasciatus Distribution: Atlantic slope: Lago de Nicaragua, Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, Matina. 12-600 m.

Hypsophrys Parachromis managuensis Hypsophrys nematopus Distribution: Atlantic slope: Lago de Nicaragua, Distribution: Atlantic slope: Lago de Nicaragua, Río Parismina, Matina and Sixaola. Remarks: This species was called Neetroplus nematopus Tomocichla Costa Rican this members of this genus were referred to the genus Theraps species in the genus Hypsophrys following Chakrabarty provisionally Tomocichla following Burgues (2000), Allgayer (2002), Kullander (2003) and Hypsophrys nicaraguensis however a comprensive taxomonic work is needed to test the monophyly and position of this Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, clade. 5-200 m. Tomocichla sieboldii Oreochromis

Oreochromis niloticus 200. Tomocichla tuba Addition to fauna. Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, and A. Angulo, pers. comm.), San Carlos, Sarapiquí (C. Matina and Sixaola. Garita, A. Angulo and B. Naranjo, pers. comm.), Tortuguero, Remarks: This species was called Theraps underwoodi Parismina (A. Molina, pers. comm.), Matina (A. Angulo, this species as synonym of Tomocichla tuba following slope: Nicoya (A. Angulo, pers. comm.), Bebedero, Tárcoles Burgues (2000), Allgayer (2002), Kullander (2003) and Remarks: Eight specimens have been collected in Vieja Aguacate River (UCR 2015-01, n=2), San Carlos drainage, 201. Vieja maculicaudaBlackbelt cichlid, n=3), Tárcoles drainage. Cabrera et alet al Distribution: Atlantic slope: Tortuguero, Parismina,

1005 Angulo et al. | Freshwater !shes of Costa Rica

Matina and Sixaola. 0-5 m. Remarks: McMahan et al. (2010) sugested the genus Paraneetroplus for this species based on a molecular phylogeny; however, we provisionally keep this species on Vieja until a comprensive morphological diagnosis for the proposed clades are made. Dormitator maculatus Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-12 m. Halichoeres Eleotris 202. Halichoeres aestuaricola Addition to fauna. Eleotris amblyopsis Distribution: Atlantic slope: Tortuguero, Parismina, coast of Costa Rica by ). Two Matina and Sixaola. 0-15 m. specimens have been collected in freshwaters in the Eleotris picta 203. Halichoeres notospilus Nicoya, Tempisque, Bebedero, Barranca, Tárcoles, Pirrís, Addition to fauna. 210. Eleotris pisonis coast of Costa Rica by Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-60 m.

Peripheral. 211. Eleotris tecta Dactyloscopus

Dactyloscopus amnis 212. Eleotris tubularis Cocos sleeper, vieja de Cocos. Addition to fauna. fauna. Remarks: This species has been reported in the saltwaters of the Cocos Island National Park, Costa Rica coast of Costa Rica by ). Thirty- three specimens have been collected in freshwaters in the been collected in freshwaters in an unknown river name in Isla del Coco drainage. This species is endemic to the Isla Pirrís drainage. Peripheral. Erotelis Hypsoblennius 213. Erotelis armiger 205. Hypsoblennius maculipinna spot blenny, cachudito aleta manchada. Addition to fauna. four specimens have been collected in freshwaters in 2011). Two specimens have been collected in freshwaters drainage. drainage. Peripheral. Gobiomorus Dormitator Gobiomorus dormitor 206. Dormitator latifrons

1006 Angulo et al. | Freshwater !shes of Costa Rica

Distribution: Atlantic slope: Lago de Nicaragua, Río Frío, 05, n=12), Pirrís drainage, and the Sábalo River (UCR et al. (2006) in Matina and Sixaola. 0-60 m. the redescription of the genus included 215. Gobiomorus maculatus Peripheral. Aboma 216. Gobiomorus polylepis 222. Aboma etheostoma boy, gobio escamoso. Addition to fauna. Guavina Remarks: Two specimens have been collected in Guavina guavina drainage. guavina. Addition to fauna. Distribution: Atlantic slope: Sixaola. 0-10 m. Awaous Remarks: This species has been reported in the 223. Awaous banana 2010). Two specimens have been collected in freshwaters Sarapiquí, Tortuguero, Parismina, Matina and Sixaola. Guavina micropus guavina. Addition to fauna. Awaous transandeanus Remarks: Three specimens have been collected in drainage. 0-120 m. Hemieleotris Bathygobius

Hemieleotris latifasciata (Meek and Hidebrand, 225. Bathygobius andrei 5-100 m. Remarks: This species has been reported in the ). Leptophilypnus One hundred eleven specimens have been collected in Barranca drainage, the Quebrada Aserradero River (UCR 220. Meek and Hildebrand, Distribution: Atlantic slope: Tortuguero. 0-6 m. Remarks: Five specimens have been collected in species in freshwaters in the Quebrada Camaronal River Thacker et al. (2006) in the redescription of the genus recorded this species in freshwaters in the Hatillo Nuevo included material of this species from the drainage drainage. 226. Bathygobius ramosus 221. (Meek and Hildebrand, Addition to fauna. Garita, pers. comm.). 0-10 m. 0-32 m. coast of Costa Rica by ). One specimen has been collected in freshwaters in an Isla del Coco drainage.

1007 Angulo et al. | Freshwater !shes of Costa Rica

Bathygobius soporator Gobionellus pseudofasciatus, Addition to fauna. collected four specimens in freshwaters in Tortuguero Distribution: Atlantic slope: Tortuguero, Parismina, Lagoon, Tortuguero drainage. In the present account we Matina and Sixaola. 0-12 m. Ctenogobius following Remarks: This species has been reported in the (2003) and Van Tassell (2011). 2010). Five specimens have been collected in freshwaters 231. Ctenogobius sagittula Addition to fauna. one specimen in freshwaters in the Tortuguero Lagoon, Tortuguero drainage. coast of Costa Rica by ). Forty- four specimens have been collected in freshwaters in the Ctenogobius Ctenogobius boleosoma Darter goby, madrejuile. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-12 m. Remarks: This species has been reported in the Gobionellus sagittula, recorded this species in freshwaters in the Quebrada Camaronal G. sagittula, recorded this species in freshwaters in the Naranjal River, Tárcoles drainage, the Punta Mala River and the Coronado Ctenogobius following Gobionellus et al. boleosoma, recorded this species in freshwaters in the Van Tassell (2011). Tortuguero Lagoon, Tortuguero drainage. In the present Ctenogobius following 232. Ctenogobius smaragdus McEachran and Fechhelm (2005), Matamoros et al Emerald goby, gobio tranquilo. Addition to fauna. Page and Burr (2011) and Van Tassell (2011). Remarks: This species has been reported in the Ctenogobius fasciatus gobio caramarcada. Addition to fauna. 2010). Three specimens have been collected in freshwaters Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-12 m. Remarks: This species has been reported in the Evorthodus 233. Evorthodus lyricus tismiche. Addition to fauna. Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-6 m. Remarks: This species has been reported in the 2010). Nine specimens have been collected in freshwaters Gobionellus fasciatus, recorded this species in freshwaters in the Tortuguero Lagoon, Tortuguero 06, n=1), Matina drainage, and the Sixaola River (UCR in Ctenogobius following McEachran and Fechhelm (2005), Matamoros et al Tassell (2011). in freshwaters in the Tortuguero Lagoon, Tortuguero drainage. 230. Ctenogobius pseudofasciatus (Gilbert and Randall, Evorthodus minutus Distribution: Atlantic slope: Tortuguero. 0-12 m. Remarks: This species has been reported in the 2010). One specimen has been collected in freshwaters in the Tortuguero Lagoon (UF 13516), Tortuguero drainage. coast of Costa Rica by ). Forty-

1008 Angulo et al. | Freshwater !shes of Costa Rica eight specimens have been collected in freshwaters in coast of Costa Rica by ). Two specimens have been collected in freshwaters in the drainage.

Gobioides Sicydium

235. Gobioides broussonnetii Sicydium adelum goby, gobio violeta. Addition to fauna. Distribution: Atlantic slope: Tortuguero and Parismina. Distribution: Atlantic slope: Parismina, Matina and 0-3 m. Remarks: This species has been reported in the Remarks: This species is endemic to the Parismina, Matina and Sixaola River drainages, Costa Rica (Bussing Sicydium altum Distribution: Atlantic slope: Sarapiquí, Tortuguero, Gobionellus

236. Gobionellus microdon Sicydium cocoensis goby, gobio de esteros. Addition to fauna. River goby, chupapiedras. Addition to fauna. Remarks: This species has been reported in the Remarks: This species has been reported in the saltwaters of the Cocos Island National Park, Costa Rica by ). have been collected in freshwaters in several rivers and streams of unknow name in the Chatam and Wafer Bays drainage. Gobiosoma species is endemic to the Isla del Coco drainage, Costa Rica Gobiosoma spes Vermiculated 2011). goby, gobio jaspeado. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-3 m. Sicydium salvini Remarks: This species has been reported in the 2010). Two specimens have been collected in freshwaters in the Tortuguero Lagoon (UCR 0260-10), Tortuguero Tortuguero Lagoon, Tortuguero drainage. Microdesmus

Lophiogobius Microdesmus dorsipunctatus to fauna. crested-goby, gobio crestado. Addition to fauna. Remarks: This species has been reported in the Remarks: This species has been reported in the ). Two specimens have been collected in freshwaters in Six specimens have been collected in freshwaters in the drainage.

Microgobius

1009 Angulo et al. | Freshwater !shes of Costa Rica

Peripheral. Distribution: Atlantic slope: Tortuguero and Matina. 0-5 m. Citharichthys Remarks: This species has been reported in the Citharichthys arenaceus Evermann and Marsh, 2010). One specimen has been collected in freshwaters Distribution: Atlantic slope: Tortuguero, Parismina, Matina and Sixaola. 0-60 m. this species in freshwaters in the Tortuguero Lagoon, Remarks: Eleven specimens have been collected in Tortuguero drainage. 251. Achirus mazatlanus 0-32 m. Citharichthys gilberti Remarks: This species has been reported in the Bigmouth sanddab, lenguado tapadero. Addition to fauna. Remarks: This species has been reported in the 10, n=2), Pirrís drainage. Trinectes 252. lenguado manchado. Addition to fauna. Pirrís drainage, the Esquinas River (UCR 1312-06, n=1) 2011). Two specimens have been collected in freshwaters drainage, the Tulín River, Tárcoles drainage, the Parrita 253. Trinectes fonsecensis Citharichthys spilopterus Distribution: Atlantic slope: Tortuguero, Parismina, Trinectes inscriptus Matina and Sixaola. 0-60 m. suela reticulada. Addition to fauna. Distribution: Atlantic slope: Matina. 0-5 m. Citharichthys uhleri Voodoo whiff, Remarks: This species has been reported in the Distribution: Atlantic slope: Tortuguero, Parismina, 2010). Four specimens have been collected in freshwaters Matina and Sixaola. 0-60 m. Peripheral. 255. Trinectes paulistanus Achirus . Distribution: Atlantic slope: Tortuguero, Parismina, Achirus declivis Matina and Sixaola. 0-10 m. Distribution: Atlantic slope: Tortuguero and Parismina. 0-3 m. Remarks: This species has been reported in the Peripheral. 2010). Two specimens have been collected in freshwaters Sphoeroides in the Tortuguero Lagoon (UCR 0260-16, n=1), Tortuguero 256. Sphoeroides annulatus Parismina drainage. puffer, timboril anillado. Addition to fauna. 250. Achirus lineatus m. suela listada. Addition to fauna. Remarks: This species has been reported in the

1010 Angulo et al. | Freshwater !shes of Costa Rica

history and composition of the Central American With the exception of Panama, all Central American recorded this species in freshwaters in the Quebrada Salvador (McMahan et al. 2013), Honduras (Matamoros et alet al Paquita River, Pirrís drainage. Sphoeroides rosenblatti Compared with the other Central American countries, puffer, timboril imitador. Addition to fauna. except Panama, Costa Rica appears to have the greatest m. Remarks: This species has been reported in the as Sphoeroides ). Twenty-nine specimens have been collected in freshwaters in the Lagarto River (UCR number of shared primary species decreases gradually: et al. 2006; Matamoros et al Sphoeroides testudineus Froese and Pauly 2013; McMahan et al. 2013). Checkered puffer, botete sapo. Addition to fauna. Distribution: Atlantic slope: Tortuguero. 0-12 m. species from the country; our results indicate the presence Remarks: This species has been reported in the of a total of twenty-four endemic species. In this account, we exclude Gobiesox potamius reported by McMahan et al. (2013) from El Salvador, but include three endemic in freshwaters in the Tortuguero Lagoon (UCR 0260-06, species of the Cocos Island (Eleotris tubularis, Gobiesox fulvus and Sicydium cocoensis), the recently described Poeciliopsis santaelena, and Cynodonichthys siegfriedi herein two specimens in freshwaters in the Tortuguero Lagoon, least seven exotic species from the country; in this account Tortuguero drainage. in natural water bodies of the country. The results of this checklist provide a framework for for the country, 22.5% (32 species) of which, based on particularly by providing up-to-date knowledge on distributions. In addition, data provided in this checklist future due to human introductions and global change and the presence of a total of 250 species, 33 (13.2%) of which will aid in conservation decisions concerning Costa Rican

RF SJ Carcharhinus leucas X X X X X X X 0-35 Per Nat Pristis pectinata X X X X 0-35 Per Nat X Pristis pristis X X X X X X X 0-35 Per Nat Atractosteus tropicus X X X X X X X Sec Nat X 0-10 Per Nat X Megalops atlanticus X X X X X X Per Nat

1011 Angulo et al. | Freshwater !shes of Costa Rica

RF SJ Albula vulpes X 0-2 Per Nat X Anguilla rostrata X X X X X X X X X 0-20 Per Nat Myrophis punctatus X 0-3 Per Nat X Myrophis vafer X Per Nat X Dorosoma chavesi X X X X X X 0-50 Per Nat X 0-66 Per Nat X X 0-10 Per Nat X Sardinella brasiliensis X 0-3 Per Nat X Anchoa curta X 0-12 Per Nat X Anchoa lucida X 0-32 Per Nat X Anchovia macrolepidota X Per Nat X Anchoviella elongata X X 0-5 Per Nat X X 0-3 Per Nat X Carassius auratus X Pri Exo Cyprinus carpio X 112-1516 Pri Exo X Cyphocharax magdalenae X 12-100 Pri Nat Hoplias microlepis X Pri Nat X X X 10-1000 Pri End Astyanax aeneus X X X X X X X X X X X X X X X X X 0-1000 Pri Nat Astyanax cocibolca X X X X X X 5-20 Pri Nat Astyanax orthodus X 1-60 Pri Nat Bramocharax bransfordii X X X X X X X 5-530 Pri Nat Brycon behreae X X X Pri Nat Brycon costaricensis X X X X X X X X 0-600 Pri Nat Bryconamericus gonzalezi X Pri Nat X Bryconamericus scleroparius X X X X X X X X X X 12-600 Pri Nat Bryconamericus terrabensis X Pri End Carlana eigenmanni X X X X X X X Pri Nat Hyphessobrycon panamensis X Pri Nat Hyphessobrycon savagei X X Pri End Hyphessobrycon tortuguerae X X X X X X 0-50 Pri Nat Odontostilbe dialeptura X Pri Nat Pseudocheirodon Térrabae X X Pri End Pterobrycon myrnae X X Pri End Roeboides bouchellei X X X X X X X X X X X X 0-610 Pri Nat Roeboides bussingi X Pri Nat Roeboides ilseae X X 10-660 Pri End Cathorops steindachneri X X 0-55 Per Nat X Cathorops tuyra X X 0-55 Per Nat X Notarius cookei X 0-6 Per Nat X Sciades assimilis X Per Nat X Sciades guatemalensis X 0-11 Per Nat Sciades seemanni X X X X X X 0-32 Per Nat X Pri End Pimelodella chagresi X X 20-660 Pri Nat Rhamdia guatemalensis X X X X X X X X X X X X X X X X X Pri Nat Rhamdia laticauda X X X X X X X X X X X X X X X X 35-1350 Pri Nat Rhamdia nicaraguensis X X X X X X X X X X 20-1160 Pri Nat

1012 Angulo et al. | Freshwater !shes of Costa Rica

RF SJ Trichomycterus striatus X X 20-660 Pri Nat Fonchiiichthys uracanthus X 20-160 Pri Nat Hemiancistrus aspidolepis X X X X 20-560 Pri Nat Pterygoplichthys pardalis X 20-110 Pri Exo X Gymnotus cylindricus X X X X X X X X X 1-50 Pri Nat Gymnotus maculosus X X X X X X X X X Pri Nat Brachyhypopomus occidentalis X 1-60 Pri Nat Oncorhynchus mykiss X X Per Exo Gobiesox fulvus X Per End Gobiesox nudus X X X X X X Per Nat Gobiesox potamius X X X X X X X Per Nat Atherinella argentea X X X X X X X X Per Nat Atherinella blackburni X 0-3 Per Nat X Atherinella chagresi X X X X Per Nat Atherinella hubbsi X X X X X X X X Per Nat Atherinella milleri X X X X X X X X X Per Nat Atherinella sardina X Per Nat Membras gilberti X X X X Per Nat X Cynodonichthys fuscolineatus X X Sec End Cynodonichthys glaucus X X Sec End Cynodonichthys hildebrandi X X Sec Nat Cynodonichthys isthmensis X X X X X X X X X X X 0-1500 Sec Nat Cynodonichthys rubripunctatus X Sec Nat Cynodonichthys siegfriedi X Sec End X X Sec Nat Oxyzygonectes dovii X X X X X X X X 0-15 Sec Nat Alfaro cultratus X X X X X X X X X X 0-300 Sec Nat Belonesox belizanus X X X X X X X X Sec Nat Brachyrhaphis cascajalensis X 5-125 Sec Nat Brachyrhaphis holdridgei X X X X X X X 35-620 Sec Nat Brachyrhaphis olomina X X X X X X X Sec End Brachyrhaphis parismina X X X X X X X X 5-50 Sec Nat Brachyrhaphis rhabdophora X X Sec End Brachyrhaphis roseni X 20-100 Sec Nat Brachyrhaphis terrabensis X X X Sec Nat Gambusia nicaraguensis X 0-15 Sec Nat Phallichthys amates X X X X X X X X X X X X 0-1120 Sec Nat Phallichthys quadripunctatus X Sec End Phallichthys tico X X X X 35-600 Sec Nat Poecilia gillii X X X X X X X X X X X X X X X X X 0-1220 Sec Nat Poecilia mexicana X X X X X X X X Sec Nat Poecilia reticulata X X 30-1250 Sec Exo Poeciliopsis elongata X X X X X X X X Sec Nat Poeciliopsis paucimaculata X X Sec End Poeciliopsis retropinna X X Sec Nat Poeciliopsis santaelena X Sec End X Poeciliopsis turrubarensis X X X X X X X X 0-120 Sec Nat Priapichthys annectens X X X X X X X X X X X X X X Sec End Priapichthys panamensis X X X X X X X X Sec Nat Xenophallus umbratilis X X X X X X X X Sec Nat Xiphophorus variatus X Sec Exo

1013 Angulo et al. | Freshwater !shes of Costa Rica

RF SJ Strongylura marina X 0-3 Per Nat X Strongylura timucu X X 0-13 Per Nat X Hemiramphus brasiliensis X 0-2 Per Nat X Hyporhamphus roberti X 0-2 Per Nat X Microphis lineatus X X X 0-12 Per Nat Pseudophallus elcapitanensis X X X X X X X X 0-20 Per Nat Pseudophallus mindi X X X X X 0-35 Per Nat Pseudophallus starksi X X X X X X X X 0-155 Per Nat Syngnathus scovelli X 0-3 Per Nat X Synbranchus marmoratus X X X X X X X X X X X X X X X X X Sec Nat Ophisternon aenigmaticum X Sec Nat X Centropomus armatus X 0-6 Per Nat X Centropomus ensiferus X X X X 0-11 Per Nat X Centropomus medius X X X X X X X X 0-250 Per Nat Centropomus mexicanus X X X X Per Nat X Centropomus nigrescens X X X X X X X X Per Nat X Centropomus parallelus X X X X X X X X Per Nat Centropomus pectinatus X X X X 0-12 Per Nat Centropomus robalito X X X X X X X X 0-50 Per Nat Centropomus undecimalis X X X X 0-50 Per Nat Centropomus viridis X X Per Nat X Centropomus unionensis X X Per Nat X Epinephelus guttatus X 0-3 Per Nat X Micropterus salmoides X Pri Exo Carangoides vinctus X Per Nat X Caranx caballus X 0-55 Per Nat X Caranx caninus X X X 0-55 Per Nat X Caranx hippos X X 0-5 Per Nat X Caranx latus X X X X 0-20 Per Nat Hemicaranx amblyrhynchus X 0-3 Per Nat X Oligoplites altus X X 0-10 Per Nat X Oligoplites palometa X X 0-10 Per Nat Oligoplites saurus X Per Nat X X X 0-3 Per Nat X X X X X X X X X 0-20 Per Nat X X X Per Nat X X 0-3 Per Nat X X X X X Per Nat X X Per Nat X X X X Per Nat X Diapterus auratus X 0-3 Per Nat X Diapterus peruvianus X X X 0-10 Per Nat X Diapterus rhombeus X X 0-12 Per Nat X Eucinostomus argenteus X 0-6 Per Nat X Eucinostomus currani X X X X X X X X Per Nat Eucinostomus dowii X 0-10 Per Nat X Eucinostomus melanopterus X X X 0-10 Per Nat X Eugerres brevimanus X X X X X X X X Per Nat Eugerres plumieri X X X X 0-31 Per Nat Gerres cinereus X 0-12 Per Nat X Gerres simillimus X X X Per Nat X

1014 Angulo et al. | Freshwater !shes of Costa Rica

RF SJ Conodon nobilis X X X X 0-51 Per Nat X Haemulopsis corvinaeformis X 0-5 Per Nat X Haemulopsis leuciscus X Per Nat X Pomadasys bayanus X X X X X X X Per Nat Pomadasys branickii X Per Nat X Pomadasys crocro X X X X X Per Nat Pomadasys empherus X 0-10 Per Nat X Pomadasys macracanthus X 0-32 Per Nat X Bairdiella ronchus X 0-5 Per Nat X Cynoscion albus X 0-6 Per Nat X X 0-3 Per Nat X Micropogonias furnieri X X 0-3 Per Nat X Ophioscion typicus X Per Nat X Polydactylus virginicus X X X 0-6 Per Nat X Kyphosus sectatrix X 0-3 Per Nat X Agonostomus monticola X X X X X X X X X X X X X X X X X X 0-650 Per Nat Joturus pichardi X X X X X 0-600 Per Nat Mugil cephalus X 0-3 Per Nat X Mugil curema X X X X X X X X X X X X Per Nat X Amatitlania kanna X 0-65 Sec Nat Amatitlania siquia X X X X X X X X X X X X X Sec Nat Amphilophus alfari X X X X X X X X X X X 2-1150 Sec Nat Amphilophus altifrons X Sec Nat Amphilophus bussingi X Sec End Amphilophus citrinellus X X X X X X X X 0-12 Sec Nat Amphilophus diquis X X X Sec End Amphilophus longimanus X X X X X X X X X 0-100 Sec Nat Amphilophus lyonsi X 10-20 Sec Nat Amphilophus rhytisma X Sec Nat Amphilophus rostratus X X X X X X X X 0-200 Sec Nat Andinoacara coeruleopunctatus X 20-100 Sec Nat Archocentrus centrarchus X X X X X X X X Sec Nat Archocentrus multispinosus X X X X X X X X X X X Sec Nat Cryptoheros myrnae X Sec End Cryptoheros sajica X X X Sec End Cryptoheros septemfasciatus X X X X X X X X 12-600 Sec Nat Hypsophrys nematopus X X X X X X X 35-535 Sec Nat Hypsophrys nicaraguensis X X X X X X X X X 5-200 Sec Nat Oreochromis niloticus X X X X X X X X X X X X X 0-1250 Sec Exo X Parachromis dovii X X X X X X X X X X X 0-600 Sec Nat Parachromis loisellei X X X X X X X X X X X Sec Nat Parachromis managuensis X X X X X X X X X X Sec Nat Tomocichla sieboldii X X X X Sec Nat Tomocichla tuba X X X X X X X X X Sec Nat Vieja maculicauda X X X X 0-5 Sec Nat Halichoeres aestuaricola X X Per Nat X Halichoeres notospilus X 0-15 Per Nat X Dactyloscopus amnis X X 0-10 Per Nat X Hypsoblennius maculipinna X 0-10 Per Nat X Dormitator latifrons X X X X X X X X Per Nat Dormitator maculatus X X X X 0-12 Per Nat

1015 Angulo et al. | Freshwater !shes of Costa Rica

RF SJ Eleotris amblyopsis X X X X 0-15 Per Nat Eleotris picta X X X X X X X X X 0-100 Per Nat Eleotris pisonis X X X X 0-60 Per Nat Eleotris tecta X X Per Nat Eleotris tubularis X 0-53 Per End X Erotelis armiger X 0-10 Per Nat X Gobiomorus dormitor X X X X X X X X X 0-60 Per Nat Gobiomorus maculatus X X X X X X X X Per Nat Gobiomorus polylepis X 2-15 Per Nat Guavina guavina X 0-10 Per Nat X Guavina micropus X 0-2 Per Nat X Hemieleotris latifasciata X X X X X X X X 5-100 Per Nat X 0-6 Per Nat X X X X 0-32 Per Nat X Aboma etheostoma X 0-32 Per Nat X Awaous banana X X X X X X X X 0-300 Per Nat Awaous transandeanus X X X X X X X X 0-120 Per Nat Bathygobius andrei X X X X 0-32 Per Nat X Bathygobius ramosus X X 0-10 Per Nat X Bathygobius soporator X X X X 0-12 Per Nat X Ctenogobius boleosoma X X X X 0-12 Per Nat X Ctenogobius fasciatus X X X X 0-12 Per Nat X Ctenogobius pseudofasciatus X 0-12 Per Nat X Ctenogobius sagittula X X X X X X X 0-32 Per Nat X Ctenogobius smaragdus X Per Nat X Evorthodus lyricus X X X X 0-6 Per Nat X Evorthodus minutus X X X X X X X 0-32 Per Nat X Gobioides broussonnetii X X 0-3 Per Nat X Gobionellus microdon X X X X X X X 0-32 Per Nat X Gobiosoma spes X 0-3 Per Nat X X Per Nat X X 0-32 Per Nat X Sicydium adelum X X X Per End Sicydium altum X X X X X Per Nat Sicydium cocoensis X 0-62 Per End X Sicydium salvini X X X X X X X 0-660 Per Nat Microdesmus dorsipunctatus X 0-3 Per Nat X Citharichthys arenaceus X X X X 0-60 Per Nat X Citharichthys gilberti X X X X X X X Per Nat X Citharichthys spilopterus X X X X 0-60 Per Nat Citharichthys uhleri X X X X 0-60 Per Nat Achirus declivis X X 0-3 Per Nat X Achirus lineatus X X 0-5 Per Nat X Achirus mazatlanus X X X 0-32 Per Nat X X 0-32 Per Nat X Trinectes fonsecensis X X X X 0-55 Per Nat Trinectes inscriptus X 0-5 Per Nat X Trinectes paulistanus X X X X 0-10 Per Nat Sphoeroides annulatus X X 0-5 Per Nat X Sphoeroides rosenblatti X X 0-32 Per Nat X Sphoeroides testudineus X 0-12 Per Nat X

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We thank to the staff of the Museo de Zoología, y Limnología (CIMAR) of the Universidad de Costa Rica, particularly to In the Freshwater Fishes of South and Central America. Porto Alegre: facilities offered. Many thanks to Bernald Pacheco and all AquaBiolab S.A. EDIPUCRS. very grateful to everyone who has contributed to this project with their Stanford Ichthyological Bulletin Bührnheim, C.M. and L.R. Malabarba. 2006. Redescription of the type species of Odontostilbe Wilfredo Matamoros, Caleb McMahan, Michael Arroyo and Carlos Cheirodontinae), and description of three new species from the Neotropical Ichthyology an anonymous reviewer for their valuable comments to the manuscript and their constructive suggestions. A preliminary survey of the Siluriformes. Neptune City: T.F.H. Burgess, W.E. 2000. The Cichlasoma story. Herichthys, the break-up. Tropical Fish Hobbyist Miscellaneous Rivulus in Publications of the Museum of Zoology, University of Michigan Costa Rica, with descriptions of new endemic species. Brenesia Peces de las aguas continentales de Costa Rica/ In L.R. Malabarba, R.E. Reis, R.P. Vari, Z.M.S. Lucena and C.A.S. Lucena (ed.). Eleotris tecta . Porto Alegre: EDIPUCRS. slope streams of tropical America (Pisces: Eleotrididae). Revista de Alda, F., A.G. Reina, I. Doadrio and E. Bermingham. 2013. Phylogeny Biologia Tropical and biogeography of the Poecilia sphenops species complex Sicydium adelum (Actinopterygii, Poeciliidae) in Central America. Molecular Gobiidae) from Atlantic slope streams of Costa Rica. Revista de Phylogenetics and Evolution 66(3): 1011-1026. Biologia Tropical Peces de las aguas continentales de Costa Rica/ Bathurst: Crawford House Press. 332 p. Allgayer, R. 2001. Description d’un genre nouveau, Cryptoheros, d’Amerique central et d’une espece nouvelle du Panama (Pisces: Poeciliopsis santaelena, Cichlidae). 1: 13-20. Allgayer, R. 2002. Un cichlide nouveau du genre Tomocichla Rica. Revista de Biologia Tropical (Perciformes: Labroidei) du Rio Guarumo, Panama. 2: Astyanax cocibolca, a new characid (Pisces: 32-36. Ostariophysi) from Lake Nicaragua, Central America. Revista de Biologia Tropical Brenesia Peces demersales y pelagicos costeros de Costa Rica. (accepted). Brycon an illustrated guide. (Teleostei: Characiformes: Characidae) from Nicaragua and Costa Rica, with a key to the lower Mesoamerican species of the genus. Peces de la Isla del Coco y peces Zootaxa (accepted). , una Lebiasinidae). Revista Unimetro , una nueva especie de In L.S. Revista Marine biodiversity of Costa Rica, Unimetro 13(5): 1-10. Central America , una nueva especie de Peces costeros del Caribe de Revista Unimetro, Separata Especial 2(25): 1-16. . San una nueva especie de Revista Peces demersales y pelagicos costeros Unimetro , una nueva especie de Dahlia an illustrated guide. 166 p. (Pisces: Characiformes, Oreochromis Lebiasinidae), una nueva especie. Dahlia niloticus (Pisces: Cichlidae) en las lagunas estacionales del Refugio (Characiformes: Brenesia Lebiasinidae), nueva especie, sistema del río Cauca, Colombia. Dahlia . Balderas: Editorial (Characiformes: Dahlia Guía de Campo de las Especies Comerciales Marinas y de Aguas Salobres de la Peces del Alto río Sinu 1: 1-20. Costa Septentrional de Sur America. Rome: FAO. 513 p. , una nueva especie Middle American cichlids with special emphasis on the section Lebiasininae). Peces del Departamento del Chocó 1: 1-20. ‘Nandopsis’ sensu Regan. Miscellaneous Publications, Museum of Rivulus-Arten von Panamá. Zoology, University of Michigan Rivulus wassmanni spec. nov., eine neue Bachlingsart der Rivulus birkhahni-kuelpmanni cichlid genus Hypsophrys Nordost-Panamá. Aquaristik aktuell diagnoses for the genus and its species. Zootaxa n. sp. and redescription of N. Insculptus Corcovado National Park, Costa Rica. Proceedings of the Academy of Natural Sciences of Philadelphia limits of Notarius. Zootaxa In R.E. Reis, S.O. Betancur-R, R. and A.P. Acero. 2006. A new species of Notarius Zootaxa South and Central America. Porto Alegre: EDIPUCRS.

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Costa Rica. Brenesia y diversidad de peces en un cuerpo de agua temporal en el Refugio In K.E. Revista de Biología Carpenter (ed.). The living marine resources of the Western Central Tropical Scharpf, C. 2006. Annotated checklist of North American freshwater Rome: FAO. Catostomidae through Mugilidae. American Currents Archocentrus morphological and molecular data, with the description of a new (Perciformes: Cichlidae), with the description of two new genera and genus. Journal of Zoological Systematics and Evolutionary Research six new species. Zootaxa - Phylogeny of species formerly assigned to the genus Archocentrus (Perciformes: Cichlidae). Zootaxa Bijdragen tot de Dierkunde 315-322. . Stockholm: Department of Zoology, Stockholm University and Department of Vertebrate America. Copeia Zoology, Swedish Museum of Natural History. 156 p. Fishes of the world Inc., Hoboken. 601 p. Eocene carangid tribe, Paratrachinotini, and a survey of anterior , a new Lebiasininae (Characiformes: Lebiasinidae) Bulletin Neotropical Ichthyology of the Biological Society of Washington Venezuela Astyanax Baird and Girard Neotropical eleotrid genus (Teleostei: Gobioidei), morphological homoplasies. BMC Evolutionary Biology including a new species and comments on Microphilypnus. Copeia Oreochromis niloticus (Pisces: Cichlidae) en el Caribe norte de Costa Rica. Revista de Biología Tropical Pristis perotteti, in Lake Nicaragua, with Recommendations for its Ortega-Lara, A., N. Milani, C. DoNascimiento, F. Villa-Navarro and In T.B. Thorson (ed.). Investigations of the . Lincoln: University of Nebraska- of Lincoln. Heptapteridae) from Colombia. Neotropical Ichthyology Copeia Page, L.M. and B.M. Burr. 2011. 622. North America north of Mexico. Freshwater Fishes of North America. Poecilia reticulata (Pisces: Poeciliidae), en un estanque en Heredia, Costa Rica. Revista de Biología Tropical Smithsonian Contributions to Zoology The biology Perdices, A., E. Bermingham, A. Montilla and I. Doadrio. 2002. Evolutionary of gobies. CRC Press, Science Publishers. history of the genus Rhamdia (Teleostei: Pimelodidae) in Central America. Molecular Phylogenetics and Evolution (Oncorhynchus mykiss) producida en Costa Rica. Agronomía Mesoamericana Eleotris (Teleostei: Eleotridae), of the Western Hemisphere, with Peces nicaraguenses de agua dulce. Managua: Editorial comparisons to the west African species. Tulane Studies in Zoology and Botany Rhamdia macuspanensis: a new species of Oreochromis niloticus en la desembocadura del río Bebedero, Golfo de Nicoya, Costa Rica. Revista in Tabasco, Mexico. Copeia de Biología Tropical Weber, A., G. Allegrucci and V. Sbordoni. 2003. Rhamdia laluchensis, a mollies (Poeciliidae: Poecilia: Mollienesia) based on mitochondrial Chiapas, Mexico. Ichthyological Exploration of Freshwaters DNA sequences. Journal of Fish Biology Xenophallus Huubs (Poeciliidae, Osteichthyes). Aquaria In F.D. Goldman and O. Salvin (ed.). special references to subtribe Nannostomina. Proceedings of the Biologia Centrali-Americana. London: Porter, R.H. United States National Museum International introductions of inland aquatic Revista de Biología species Tropical un sein Formenkreis. Aquarien Magazin BryconamericusRevista de Biología Tropical of Corcovado National Park, Costa Rica. Brenesia Bramocharax and a description of a second, more primitive, species in Guatemala. complex freshwater-marine ecotone. Environmental Biology of Fishes American Museum Novitates 2500: 1-21. Parque Nacional Corcovado luego del cese de las actividades mineras. systematics. Bulletin of the American Museum of Natural History 126: Brenesia of synbranchid eel and the phylogeny and systematics of Bulletin of the American Museum of Natural History Asbana : September 2013 the surf-inhabiting atheriniform (Pisces: Isonidae). Records of : October 2013 the Western Australian Museum : Tiago Pinto Carvalho

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