Abstract Abstract
An updated checklist of the distribution of fishes that inhabit the continental waters of the Mexican state of Chiapas is presented. The state was compartmentalized into 12 hydrological regions for the purpose of understanding the distribution of fish fauna across a state with large physiographic variance. The ichthyofauna of Chiapas is represented by 311 species distributed in two classes, 26 orders, 73 families, and 182 genera, including 12 exotic species. The families with the highest number of species were Cichlidae, Poeciliidae, Sciaenidae, Carangidae, Ariidae, Gobiidae, and Haemulidae. This study attempts to close gaps in knowledge of the distribution of ichthyofauna in the diverse hydrological regions of Chiapas, Mexico.
Keywords: Distribution, endemism, fish diversity, ichthyology, southern Mexico
Introduction
The hydrological wealth of Chiapas is manifested through its 72 perennial rivers and abundant streams, lakes, and ponds. The presence of large hydroelectric dams has significantly increased the surface area of the state’s bodies of water (Velasco-Colín 1976). Chiapas has a coastline of 270 km and more than 70,000 hectares of estuaries and coastal lagoons (Contreras-Espinosa 2010), which favors the presence of rich fish diversity (Velasco-Colín 1976, Lozano-Vilano and Contreras-Balderas 1987, Rodiles-Hernández et al. 2005). Much of the state is located in the Usumacinta ichthyographic province/area of endemism (Miller et al. 2005, Matamoros et al. 2015), which means that its continental waters host a high number of endemic species, making Chiapas a freshwater biodiversity hotspot (Hudson et al. 2005, Matamoros et al. 2015).
Several attempts have been made to record continental water fish diversity in Chiapas through numerous works such as checklists, annotated checklists, books and scattered records in the literature (e.g. Velasco-Colín 1976, Lozano-Vilano and Contreras-Balderas 1987, Lazcano-Barrero and Vogt 1992, Tapia-García et al. 1998, Rodiles-Hernández et al. 1999, Rodiles-Hernández 2005, Rodiles-Hernández et al. 2005, 2013, Lozano-Vilano et al. 2007, González-Díaz et al. 2008, Espinosa-Pérez et al. 2011, Velázquez-Velázquez et al. 2013, Gómez-González et al. 2012, 2015). The first comprehensive publication on continental fishes of Chiapas was made by Velasco-Colín (1976), who reported 74 species distributed across 28 families. He also included brief information about the ecology, biology and distribution of several species and, in some cases, added relevant fishing information.
Subsequently Lozano-Vilano and Contreras-Balderas (1987) published an annotated checklist in which they registered 135 species belonging to 38 families in the state’s continental waters. In addition to an increase in the number of data records, for the first time the distribution of fishes was associated with seven of the state’s physiographic regions. Eighteen years later Rodiles-Hernández (2005) and Rodiles-Hernández et al. (2005) recorded 205 species in 44 families and 207 species in 45 families respectively. In the first study, distributions were reported at the level of the two main Chiapas river basins, the Grijalva-Usumacinta and the Coast of Chiapas, whereas, in the second study, the distributional geographic units were the Atlantic and the Pacific slope. Velázquez-Velázquez et al. (2013) was the last published attempt to summarize continental fishes of Chiapas. They reported 262 species across 57 families, and once again the geographic distribution units were the Grijalva and the Usumacinta River basins and the coast of Chiapas.
Two interesting trends emerge about the continental fishes of Chiapas. First, the number of recorded species has continued to increase over time likely due to an increase in sampling localities, implementation of new sampling techniques, new records and species descriptions. The second trend is related to the geographic units in which the state has been divided. For instance, Lozano-Vilano and Contreras-Balderas (1987) divided the state into seven physiographic regions, based on terrestrial relief. Most studies used broad delineations limited to the three major hydrologic regions (coast of Chiapas and the Grijalva and Usumacinta River basins) masking detailed information on finer distributional patterns like localized endemism and drainage interconnections.
Therefore, the aim of this study is to provide an updated checklist of the continental fishes of Chiapas, including distribution data, based on extensive literature research and complemented with material deposited in the ichthyological collection of the (MZ-P-UNICACH). For the first time, we use finer scale geographic divisions for the state, implemented at the sub-basin level, following the National Institute of Statistics and Geography (INEGI 2010).
Materials and methods
The bulk of records came from the material of 204 species deposited in the ichthyological collection of the MZ-P-UNICACH Museum of Zoology (MZ-P-UNICACH, SEMARNAT: CHIS-PEC-210-03-09). In addition, we performed an extensive literature review for records of continental fishes of Chiapas. The checklists previously published by Lozano-Vilano and Contreras-Balderas (1987), Rodiles-Hernández (2005), Rodiles-Hernández et al. (2005), Espinosa-Pérez et al. (2011), and Velázquez-Velázquez et al. (2013) were taken as the basis for this work and were supplemented with publications by Lazcano-Barrero and Vogt (1992), Tapia-García et al. (1998), Rodiles-Hernández et al. (1999), Lozano-Vilano et al. (2007) and Gómez-González et al. (2012, 2015) who developed lists for particular regions of the state. We also included Castro-Aguirre et al. (1999) and Miller et al. (2005).
Species were systematically arranged by order and family following Nelson (2006). Genera and species were arranged alphabetically; scientific names and authorities were corroborated following Eschmeyer et al. (2016). Tolerance to salinity was based on Myers (1938).
The 12 geographical units for Chiapas (Figure 1) were utilized to determine the distribution of each species across the state. These 12 units were based on existing hydrological sub-basins of the state (INEGI 2010). The main rivers, ponds, lakes and coastal lagoons of each sub-basin are listed in Table 1.
Figure 1.
Geographical units for the study of the distribution of the fish fauna of the state of Chiapas: I (Usumacinta-Chixoy) II (Usumacinta-Lacantún) III (Usumacinta-Catazajá) IV (Usumacinta-Jataté) V (Grijalva-Tulijá) VI (Grijalva-Teapa) VII (Grijalva-Peñitas) VIII (Grijalva-Malpaso), IX (Grijalva-Chicoasén) X (Grijalva-La Angostura) XI (Costa-Itsmo) XII (Costa-Soconusco).
Table 1.
Geographic units utilized to study the distribution of the fish fauna of Chiapas and sub-basins that form them.
| Hidrological region | Basin | Sub-basin | Geographic unit |
|---|---|---|---|
| COSTA DE CHIAPAS | R. SUCHIATE AND OTHERS | R. Suchiate | Costa-Soconusco |
| R. Cozoloapan | |||
| R. Cahuacán | |||
| Puerto Madero | |||
| R. Coatán | |||
| R. Huehuetán | |||
| R. HUIXTLA AND OTHERS | R. Huixtla | ||
| R. Despoblado | |||
| L. del Viejo y Tembladeras | |||
| R. Cacaluta | |||
| R. Sesecapa | |||
| R. Novillero | |||
| R. PIJIJIAPAN AND OTHERS | R. Margaritas y Coapa | Costa-Istmo | |
| R. Pijijiapan | |||
| R. San Diego | |||
| El Porvenir | |||
| R. Jesús | |||
| L. de la Joya | |||
| MAR MUERTO | R. Zanatenco | ||
| Mar Muerto | |||
| R. La Punta | |||
| R. Las Arenas | |||
| R. Tapanatepec | |||
| GRIJALVA - USUMACINTA | R. USUMACINTA | R. Usumacinta | Usumacinta-Catazajá |
| R. Chacamax | |||
| R. Chacaljáh | |||
| R. CHIXOY | R. Chixoy | Usumacinta-Chixoy | |
| R. Negro | |||
| R. GRIJALVA - VILLAHERMOSA | R. Viejo Mezcalapa | Grijalva-Peñitas | |
| R. Mezcalapa | |||
| R. Tzimbac | |||
| R. Zayula | |||
| R. Platanar | |||
| R. Paredón | |||
| R. Pichucalco | |||
| R. Tacotalpa | |||
| R. Samaria | |||
| R. de la Sierra | Grijalva-Teapa | ||
| R. Almendro | |||
| R. Plátanos | |||
| R. Chacté | Grijalva.Tulijá | ||
| R. Puxcatán | |||
| R. Macuspana | |||
| GRIJALVA - USUMACINTA | R. Shumulá | ||
| R. Yashijá | |||
| R. Tulijá | |||
| R. Bascá | |||
| R. Chilapa | |||
| R. GRIJALVA - TUXTLA GUTIÉRREZ | P. Nezahualcóyotl | Grijalva-Malpaso | |
| R. La Venta | |||
| R. Encajonado | |||
| R. Cintalapa | |||
| R. de Zoyatenco | |||
| R. Alto Grijalva | Grijalva-Chicoasén | ||
| R. Hondo | |||
| R. Chicoasén | |||
| R. Suchiapa | |||
| Tuxtla Gutiérrez | |||
| El Chapopote | |||
| R. Santo Domingo | |||
| R. GRIJALVA - LA CONCORDIA | P. La Angostura | Grijalva-La Angostura | |
| R. Selegua | |||
| R. Lagartero | |||
| R. Aguacatenco | |||
| R. San Pedro | |||
| R. La Concordia | |||
| R. Grande o Salinas | |||
| R. Aguazurco | |||
| R. San Miguel | |||
| R. Yahuayita | |||
| R. Zacualpa | |||
| R. Tapizaca | |||
| R. Comitan | |||
| R. LACANTÚN | R. Lacantún | Usumacinta-Lacantún | |
| R. Ixcán | |||
| R. Chajul | |||
| R. Lacanjá | |||
| R. San Pedro | |||
| L. Miramar | |||
| R. Perlas | |||
| R. Jataté | |||
| R. Azul | Usumacinta-Jataté | ||
| R. Tzaconejá | |||
| R. Margaritas | |||
| R. Santo Domingo | |||
| R. Seco | |||
| R. Caliente | |||
| R. Euseba |
Results
The continental fishes of the state of Chiapas are represented by two classes, 26 orders, 73 families, 182 genera and 311 species (Table 2), including 12 exotic species (Ctenopharyngodon idella, Cyprinus carpio, Micropterus salmoides, Oncorhynchus mykiss, Oreochromis aureus, Oreochromis mossambicus, Oreochromis niloticus, Parachromis managuensis, Pterygoplichthys disjunctivus, Pterygoplichthys multiradiatus, Pterygoplichthys pardalis, and Tilapia zilli). Only five species were endemic: the catfish Lacantunia enigmatica, the cichlids Rocio ocotal and Thorichthys socolofi, the killifish Tlaloc hildebrandi and the molly Poecilia thermalis. Based on species richness the most important families were: Cichlidae (35), Poeciliidae (29), Sciaenidae (18), Carangidae (17), Ariidae (16), Gobiidae (12), and Haemulidae (11). Almost all of these families, except the first two, contains peripheral species. These eight families represented 44.37% (138) of the state’s total species richness. Thirteen species are included in risk categories under Mexican law (NOM-059-SEMARNAT-2010; SEMARNAT 2010): Poecilia sulphuraria and Tlaloc hildebrandi are listed as endangered; Priapella compressa, Thorichthys socolofi, Vieja hartwegi and Xiphophorus clemenciae are listed as threatened; finally Chiapaheros grammodes, Gambusia eurystoma, Hippocampus ingens, Potamarius nelsoni, Priapella intermedia, Rhamdia guatemalensis and Chuco intermedium are listed as species under special protection. Based on general salinity tolerance, and excluding exotic species, 16 are primary freshwater fishes, 65 secondary freshwater fishes, and the rest of the species are peripheral (Table 2).
Table 2.
Systematic list of the continental waters ichthyofauna of Chiapas. Ecological classification: PF, SF, PF, PC, P, Ex.
| No | Taxon | Ecological classification | Grijalva- La Angostura | Grijalva-Chicoasén | Grijalva-Malpaso | Grijalva-Peñitas | Grijalva- Teapa | Grijalva-Tulijá | Usumacinta-Jataté | Usumacinta-Lacantún | Usumacinta- Chixoy | Usumacinta-Catazajá | Costa-Istmo | Costa-Soconusco |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Order Carcharhiniformes | ||||||||||||||
| I Family Carcharhinidae | ||||||||||||||
| 1 | Carcharhinus leucas (Müller & Henle, 1839) | P | x | |||||||||||
| 2 | Carcharhinus limbatus (Müller & Henle, 1839) | P | x | x | ||||||||||
| 3 | Carcharhinus cerdale Gilbert, 1898 | P | x | |||||||||||
| 4 | Rhizoprionodon longurio (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 5 | Negaprion brevirostris (Poey, 1868) | P | x | |||||||||||
| II Family Sphyrnidae | ||||||||||||||
| 6 | Sphyrna tiburo (Linnaeus, 1758) | P | x | |||||||||||
| Order Pristiformes | ||||||||||||||
| III Family Pristidae | ||||||||||||||
| 7 | Pristis pectinata Latham, 1794 | P | x | |||||||||||
| 8 | Pristis microdon Latham, 1794 | P | x | |||||||||||
| Order Rhinobatiformes | ||||||||||||||
| IV Family Rhinobatidae | ||||||||||||||
| 9 | Pseudobatos glaucostigma (Jordan & Gilbert, 1883) | P | x | |||||||||||
| Order Myliobatiformes | ||||||||||||||
| V Family Urotrygonidae | ||||||||||||||
| 10 | Urotrygon aspidura (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 11 | Urotrygon chilensis (Günther, 1872) | P | x | |||||||||||
| 12 | Urotrygon munda Gill, 1863 | P | x | |||||||||||
| 13 | Urotrygon nana Miyake & McEachran, 1998 | P | x | |||||||||||
| 14 | Urotrygon rogersi (Jordan & Starks, 1895) | P | x | |||||||||||
| VI Family Dasyatidae | ||||||||||||||
| 15 | Hypanus longus (Garman, 1880) | P | x | x | ||||||||||
| 16 | Himantura pacifica (Beebe & Tee-Van, 1941) | P | x | x | ||||||||||
| VII Family Myliobatidae | ||||||||||||||
| 17 | Aetobatus laticeps Gill, 1865 | P | x | x | ||||||||||
| VIII Family Rhinopteridae | ||||||||||||||
| 18 | Rhinoptera steindachneri Evermann & Jenkins, 1891 | P | x | x | ||||||||||
| Order Lepisosteiformes | ||||||||||||||
| IX Family Lepisosteidae | ||||||||||||||
| 19 | Atractosteus tropicus Gill, 1863 | PF | x | x | x | x | x | x | ||||||
| Order Elopiformes | ||||||||||||||
| X Family Elopidae | ||||||||||||||
| 20 | Elops affinis Regan, 1909 | P | x | x | ||||||||||
| XI Family Megalopidae | ||||||||||||||
| 21 | Megalops atlanticus Valenciennes, 1847 | P | x | x | ||||||||||
| Order Albuliformes | ||||||||||||||
| XII Family Albulidae | ||||||||||||||
| 22 | Albula esuncula (Garman, 1899) | P | x | |||||||||||
| Order Anguilliformes | ||||||||||||||
| XIII Family Ophichthidae | ||||||||||||||
| 23 | Myrichthys xysturus (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 24 | Ophichthus zophochir Jordan & Gilbert, 1882 | P | x | x | ||||||||||
| Order Clupeiformes | ||||||||||||||
| XIV Family Pristigasteridae | ||||||||||||||
| 25 | Pliosteostoma lutipinnis (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 26 | Odontognathus panamensis (Steindachner, 1876) | P | x | |||||||||||
| 27 | Opisthopterus dovii (Günther, 1868) | P | x | |||||||||||
| XV Family Engraulidae | ||||||||||||||
| 28 | Anchoa argentivittata (Regan, 1904) | P | x | |||||||||||
| 29 | Anchoa curta (Jordan & Gilbert, 1882) | P | x | x | ||||||||||
| 30 | Anchoa ischana (Jordan & Gilbert, 1882) | P | x | x | ||||||||||
| 31 | Anchoa lucida (Jordan & Gilbert, 1882) | P | x | x | ||||||||||
| 32 | Anchoa mitchilli (Valenciennes, 1848) | P | x | |||||||||||
| 33 | Anchoa mundeola (Gilbert & Pierson, 1898) | P | x | x | ||||||||||
| 34 | Anchoa walkeri Baldwin & Chang, 1970 | P | x | |||||||||||
| 35 | Anchoa starksi (Gilbert & Pierson, 1898) | P | x | x | ||||||||||
| 36 | Anchovia macrolepidota (Kner, 1863) | P | x | x | ||||||||||
| XVI Family Clupeidae | ||||||||||||||
| 37 | Dorosoma anale Meek, 1904 | P (V) | x | x | x | x | x | x | x | x | x | |||
| 38 | Dorosoma petenense (Günther, 1867) | P (V) | x | x | x | x | x | x | x | x | x | |||
| 39 | Harengula thrissina (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 40 | Lile gracilis Castro-Aguirre & Vivero, 1990 | P | x | x | ||||||||||
| 41 | Lile nigrofasciata Castro-Aguirre, Ruiz-Campos & Balart, 2005 | P | x | x | ||||||||||
| 42 | Opisthonema libertate (Günther, 1867) | P | x | x | ||||||||||
| 43 | Opisthonema medirastre Berry & Barret, 1964 | P | x | |||||||||||
| Order Gonorynchiformes | ||||||||||||||
| XVII Family Chanidae | ||||||||||||||
| 44 | Chanos chanos (Forsskål, 1775) | P | x | x | ||||||||||
| Order Cypriniformes | ||||||||||||||
| XVIII Family Cyprinidae | ||||||||||||||
| 45 | Ctenopharyngodon idella (Valenciennes, 1844)Ex | Ex | x | x | x | x | x | x | ||||||
| 46 | Cyprinus carpio (Linnaeus, 1758)Ex | Ex | x | x | x | x | ||||||||
| XIX Family Catostomidae | ||||||||||||||
| 47 | Ictiobus meridionalis (Günther, 1868) | PF | x | x | x | x | x | x | x | |||||
| Order Characiformes | ||||||||||||||
| XX Family Characidae | ||||||||||||||
| 48 | Astyanax aeneus (Günther, 1860) | PF | x | x | x | x | x | x | x | x | x | x | x | x |
| 49 | Bramocharax sp. | PF | x | x | x | x | ||||||||
| 50 | Brycon guatemalensis Regan, 1908 | PF | x | x | x | x | x | x | x | x | x | |||
| 51 | Hyphessobrycon compressus (Meek, 1904) | PF | x | x | x | x | ||||||||
| 52 | Roeboides bouchellei Fowler, 1923 | PF | x | x | ||||||||||
| Order Siluriformes | ||||||||||||||
| XXI Family Lacantuniidae | ||||||||||||||
| 53 | Lacantunia enigmatica Rodiles-Hernández, Hendrickson & Lundberg, 2005 | PF | x | |||||||||||
| XXII Family Loricariidae | ||||||||||||||
| 54 | Pterygoplichthys disjunctivus (Weber, 1991) Ex | Ex | x | |||||||||||
| 55 | Pterygoplichthys multiradiatus (Hancock, 1828) Ex | Ex | x | |||||||||||
| 56 | Pterygoplichthys pardalis (Castelnau, 1855) Ex | Ex | x | x | x | x | x | |||||||
| XXIII Family Heptapteridae | ||||||||||||||
| 57 | Rhamdia guatemalensis (Günther, 1864) | PF | x | x | x | x | x | x | x | x | x | x | x | x |
| 58 | Rhamdia laluchensis Weber, Allegrucci & Sbordoni, 2003 | PF | x | |||||||||||
| 59 | Rhamdia laticauda (Kner, 1858) | PF | x | x | x | x | x | x | x | |||||
| 60 | Rhamdia parryi Eigenmann & Eigenmann, 1888 | PF | x | |||||||||||
| XXIV Family Ictaluridae | ||||||||||||||
| 61 | Ictalurus meridionalis (Günther, 1864) | PF | x | x | x | x | x | x | x | x | x | |||
| XV Family Ariidae | ||||||||||||||
| 62 | Bagre panamensis (Gill, 1863) | P | x | |||||||||||
| 63 | Bagre pinnimaculatus (Steindachner, 1876) | P | x | |||||||||||
| 64 | Cathorops dasycephalus (Günther, 1864) | P | x | |||||||||||
| 65 | Cathorops cf. fuerthii | P | x | |||||||||||
| 66 | Cathorops kailolae Marceniuk & Betancur-R., 2008 | P (V) | x | x | x | x | x | x | ||||||
| 67 | Cathorops liropus (Bristol, 1897) | P | x | x | ||||||||||
| 68 | Cathorops steindachneri (Gilbert & Starks, 1904) | P | x | x | ||||||||||
| 69 | Notarius kessleri (Steindachner, 1876) | P | x | |||||||||||
| 70 | Notarius planiceps (Steindachner, 1876) | P | x | |||||||||||
| 71 | Notarius troschelii (Gill, 1863) | P | x | |||||||||||
| 72 | Potamarius nelsoni (Evermann & Goldsborough, 1902) | P (V) | x | x | x | x | x | x | x | |||||
| 73 | Potamarius usumacintae Betancourt-R. & Willink, 2007 | P (V) | x | x | x | |||||||||
| 74 | Sciades dowii (Gill, 1863) | P | x | |||||||||||
| 75 | Sciades felis (Linnaeus, 1766) | P | x | |||||||||||
| 76 | Sciades guatemalensis (Günther, 1864) | P | x | x | ||||||||||
| 77 | Sciades seemanni (Günther, 1864) | P | x | x | ||||||||||
| Order Gymnotiformes | ||||||||||||||
| XXVI Family Gymnotidae | ||||||||||||||
| 78 | Gymnotus maculosus Albert & Miller, 1995 | PF | x | |||||||||||
| Order Salmoniformes | ||||||||||||||
| XXVII Family Salmonidae | ||||||||||||||
| 79 | Oncorhynchus mykiss (Walbaum, 1972)Ex | Ex | x | |||||||||||
| Order Aulopiformes | ||||||||||||||
| XXVIII Family Synodontidae | ||||||||||||||
| 80 | Synodus scituliceps Jordan & Gilbert, 1881 | P | x | x | ||||||||||
| Order Batrachoidiformes | ||||||||||||||
| XXIX Family Batrachoididae | ||||||||||||||
| 81 | Batrachoides boulengeri Gilbert & Starks, 1904 | P | x | |||||||||||
| 82 | Batrachoides goldmani Evermann & Goldsborough, 1902 | P (V) | x | x | x | x | x | x | x | |||||
| 83 | Batrachoides waltersi Collette & Russo, 1981 | P | x | x | ||||||||||
| 84 | Porichthys greenei Gilbert & Starks, 1904 | P | x | |||||||||||
| Order Mugiliformes | ||||||||||||||
| XXX Family Mugilidae | ||||||||||||||
| 85 | Agonostomus monticola (Bancroft, 1834) | P (Ca) | x | x | x | x | x | x | ||||||
| 86 | Joturus pichardi Poey, 1860 | P (Ca) | x | x | ||||||||||
| 87 | Mugil cephalus Linnaeus, 1758 | P | x | x | ||||||||||
| 88 | Mugil curema Valenciennes, 1836 | P | x | x | x | x | ||||||||
| 89 | Mugil hospes Jordan & Culver, 1895 | P | x | x | ||||||||||
| Order Atheriniformes | ||||||||||||||
| XXXI Family Atherinopsidae | ||||||||||||||
| 90 | Atherinella guatemalensis (Günther, 1864) | P | x | x | ||||||||||
| 91 | Atherinella alvarezi (Díaz-Pardo, 1972) | P (V) | x | x | x | x | x | x | x | x | x | |||
| 92 | Atherinella panamensis Steindachner, 1875 | P | x | |||||||||||
| 93 | Atherinella schultzi (Alvarez & Carranza, 1952) | P (V) | x | x | x | x | ||||||||
| 94 | Membras gilberti (Jordan & Bollman, 1889) | P | x | x | ||||||||||
| Order Beloniformes | ||||||||||||||
| XXXII Family Hemiramphidae | ||||||||||||||
| 95 | Hyporhamphus mexicanus Alvarez, 1959 | P (V) | x | x | x | x | x | x | x | |||||
| 96 | Hyporhamphus snyderi Meek & Hildebrand, 1973 | P | x | x | ||||||||||
| 97 | Hyporhamphus naos Banford & Collette, 2001 | P | x | x | ||||||||||
| XXXIII Family Belonidaex | ||||||||||||||
| 98 | Strongylura hubbsi Collette, 1974 | P (V) | x | x | x | x | x | x | x | |||||
| 99 | Strongylura exilis (Girard, 1854) | P | x | |||||||||||
| 100 | Tylosurus fodiator Jordan & Gilbert, 1882 | P | x | |||||||||||
| Order Cyprinodontiformes | ||||||||||||||
| XXXIV Family Rivulidae | ||||||||||||||
| 101 | Cynodonichthys tenuis Meek, 1904 | SF | x | x | x | x | x | |||||||
| XXXV Family Profundulidae | ||||||||||||||
| 102 | Profundulus punctatus (Günther, 1866) | SF | x | x | x | |||||||||
| 103 | Tlaloc candalarius (Hubbs, 1924) | SF | x | |||||||||||
| 104 | Tlaloc hildebrandi Miller, (1950) | SF | x | x | ||||||||||
| 105 | Tlaloc labialis (Günther, 1866) | SF | x | x | x | x | x | x | x | |||||
| XXXVI Family Anablepidae | ||||||||||||||
| 106 | Anableps dowei Gill, 1861 | SF | x | x | ||||||||||
| XXXVII Family Poeciliidae | ||||||||||||||
| 107 | Belonesox belizanus Kner, 1860 | SF | x | x | x | x | x | |||||||
| 108 | Brachyrhaphis hartwegi Rosen & Bailey, 1982 | SF | x | |||||||||||
| 109 | Carlhubbsia kidderi (Hubbs, 1936) | SF | x | x | x | |||||||||
| 110 | Gambusia eurystoma Miller, 1975 | SF | x | |||||||||||
| 111 | Gambusia sexradiata Hubbs, 1936 | SF | x | x | x | x | x | x | x | x | ||||
| 112 | Gambusia yucatana Regan, 1914 | SF | x | x | x | |||||||||
| 113 | Heterophallus echeagarayi (Alvarez, 1952) | SF | x | x | ||||||||||
| 114 | Heterophallus milleri Radda, 1987 | SF | x | |||||||||||
| 115 | Phallichthys fairweatheri Rosen & Bailey, 1959 | SF | x | x | x | |||||||||
| 116 | Poecilia kykesis Poeser, 2002 | SF | x | |||||||||||
| 117 | Poecilia mexicana Steindachner, 1863 | SF | x | x | x | x | x | x | x | x | ||||
| 118 | Poecilia nelsoni (Meek, 1904) | SF | x | x | ||||||||||
| 119 | Poecilia sphenops Valenciennes, 1836 | SF | x | x | x | x | x | x | x | x | x | x | ||
| 120 | Poecilia sulphuraria (Alvarez, 1948) | SF | x | |||||||||||
| 121 | Poecilia thermalis Steindachner, 1863 | SF | x | |||||||||||
| 122 | Poeciliopsis fasciata (Meek, 1904) | SF | x | x | x | x | x | |||||||
| 123 | Poeciliopsis hnliickai Meyer & Vogel, 1981 | SF | x | x | x | |||||||||
| 124 | Poeciliopsis pleurospilus (Günther, 1868) | SF | x | x | x | x | x | x | ||||||
| 125 | Poeciliopsis turrubarensis (Meek, 1912) | SF | x | x | ||||||||||
| 126 | Priapella intermedia Alvarez & Carranza, 1952 | SF | x | |||||||||||
| 127 | Priapella chamulae Schartl, Meyer & Wilde, 2006 | SF | x | x | ||||||||||
| 128 | Priapella compressa Alvarez, 1948 | SF | x | |||||||||||
| 129 | Priapella lacandonae Meyer, Schories & Schartl, 2011 | SF | x | |||||||||||
| 130 | Pseudoxiphophorus bimaculatus (Heckel, 1848) | SF | x | x | x | x | x | x | x | x | ||||
| 131 | Xenodexia ctenolepis Hubbs, 1950 | SF | x | |||||||||||
| 132 | Xiphophorus alvarezi Rosen, 1960 | SF | x | |||||||||||
| 133 | Xiphophorus clemenciae Álvarez, 1959 | SF | x | |||||||||||
| 134 | Xiphophorus hellerii Heckel, 1848 | SF | x | x | x | x | x | x | x | |||||
| 135 | Xiphophorus maculatus (Günther, 1866) | SF | x | x | x | x | ||||||||
| Order Syngnathiformes | ||||||||||||||
| XXXVIII Family Syngnathidae | ||||||||||||||
| 136 | Hippocampus ingens Girard, 1859 | P | x | x | ||||||||||
| 137 | Pseudophallus starksii (Jordan & Culver, 1895) | P | x | x | ||||||||||
| XXXIX Family Fistulariidae | ||||||||||||||
| 138 | Fistularia commersonii Rüppell, 1838 | P | x | |||||||||||
| Order Synbranchiformes | ||||||||||||||
| XL Family Synbranchidae | ||||||||||||||
| 139 | Ophisternon aenigmaticum Rosen & Greenwood, 1976 | PF | x | x | x | x | x | x | x | x | ||||
| 140 | Synbranchus marmoratus Bloch, 1795 | PF | x | x | ||||||||||
| Order Perciformes | ||||||||||||||
| XLI Family Centropomidae | ||||||||||||||
| 141 | Centropomus armatus Gill, 1863 | P | x | x | ||||||||||
| 142 | Centropomus medius Günther, 1864 | P | x | x | ||||||||||
| 143 | Centropomus nigrescens Günther, 1864 | P | x | x | ||||||||||
| 144 | Centropomus robalito Jordan & Gilbert, 1882 | P | x | x | ||||||||||
| 145 | Centropomus undecimalis (Bloch, 1792) | P | x | x | x | x | ||||||||
| 146 | Centropomus parallelus Poey, 1860 | P | x | x | ||||||||||
| 147 | Centropomus poeyi Chávez, 1961 | P | x | |||||||||||
| 148 | Centropomus unionensis Bocourt, 1868 | P | x | |||||||||||
| 149 | Centropomus viridis Lockington, 1877 | P | x | x | ||||||||||
| XLII Family Serranidae | ||||||||||||||
| 150 | Dermatolepis dermatolepis (Boulenger, 1895) | P | x | |||||||||||
| 151 | Alphestes multiguttatus (Günther, 1867) | P | x | x | ||||||||||
| 152 | Epinephelus labriformis (Jenyns, 1840) | P | x | |||||||||||
| 153 | Epinephelus analogus Gill, 1863 | P | x | |||||||||||
| 154 | Epinephelus quinquefasciatus (Bocourt, 1868) | P | x | |||||||||||
| 155 | Mycteroperca xenarcha Jordan, 1888 | P | x | |||||||||||
| 156 | Rypticus nigripinnis Gill, 1861 | P | x | |||||||||||
| XLIII Family Centrarchidae | ||||||||||||||
| 157 | Micropterus salmoides (Lacepéde, 1802)Ex | Ex | x | x | ||||||||||
| XLIV Family Nematistiidae | ||||||||||||||
| 158 | Nematistius pectoralis Gill, 1862 | P | x | x | ||||||||||
| XLV Family Carangidae | ||||||||||||||
| 159 | Carangoides otrynter (Jordan & Gilbert, 1883) | P | x | |||||||||||
| 160 | Carangoides vinctus (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 161 | Caranx caballus Günther, 1868 | P | x | |||||||||||
| 162 | Caranx caninus Günther, 1867 | P | x | x | ||||||||||
| 163 | Caranx sexfasciatus Quoy & Gaimard, 1825 | P | x | |||||||||||
| 164 | Chloroscombrus orqueta Jordan & Gilbert, 1883 | P | x | |||||||||||
| 165 | Gnathanodon speciosus (Forsskål, 1775) | P | x | |||||||||||
| 166 | Hemicaranx leucurus (Günther 1864) | P | x | |||||||||||
| 167 | Hemicaranx zelotes Gilbert, 1898 | P | x | x | ||||||||||
| 168 | Oligoplites altus (Günther, 1868) | P | x | x | ||||||||||
| 169 | Oligoplites saurus (Bloch & Schneider, 1801) | P | x | x | ||||||||||
| 170 | Selene brevoortii (Gill, 1863) | P | x | |||||||||||
| 171 | Selene oerstedii Lütken, 1880 | P | x | x | ||||||||||
| 172 | Selene peruviana (Guichenot, 1866) | P | x | x | ||||||||||
| 173 | Trachinotus kennedyi Steindachner, 1876 | P | x | x | ||||||||||
| 174 | Trachinotus paitensis Cuvier, 1832 | P | x | |||||||||||
| 175 | Trachinotus rhodopus Gill, 1863 | P | x | x | ||||||||||
| XLVI Family Lutjanidae | ||||||||||||||
| 176 | Hoplopagrus guentherii Gill, 1862 | P | x | x | ||||||||||
| 177 | Lutjanus argentiventris (Peters, 1869) | P | x | x | ||||||||||
| 178 | Lutjanus colorado Jordan & Gilbert, 1882 | P | x | x | ||||||||||
| 179 | Lutjanus guttatus (Steindachner, 1869) | P | x | x | ||||||||||
| 180 | Lutjanus novemfasciatus Gill, 1862 | P | x | x | ||||||||||
| XLVII Family Lobotidae | ||||||||||||||
| 181 | Lobotes pacificus Gilbert, 1898 | P | x | |||||||||||
| XLVIII Family Gerreidae | ||||||||||||||
| 182 | Diapterus brevirostris (Sauvage, 1879) | P | x | x | ||||||||||
| 183 | Eucinostomus currani Zahuranec, 1980 | P | x | x | ||||||||||
| 184 | Eucinostomus dowii (Gill, 1863) | P | x | x | ||||||||||
| 185 | Eucinostomus gracilis (Gill, 1862) | P | x | |||||||||||
| 186 | Eugerres axillaris (Günther, 1864) | P | x | x | ||||||||||
| 187 | Eugerres lineatus (Humboldt, 1821) | P | x | |||||||||||
| 188 | Eugerres mexicanus (Steindachner, 1879) | P (V) | x | x | x | x | x | x | ||||||
| 189 | Gerres simillimus Regan, 1907 | P | x | x | ||||||||||
| XLIX Family Haemulidae | ||||||||||||||
| 190 | Conodon serrifer Jordan & Gilbert, 1882 | P | x | |||||||||||
| 191 | Genyatremus pacifici (Günther, 1864) | P | x | x | ||||||||||
| 192 | Haemulopsis axillaris (Steindachner, 1869) | P | x | x | ||||||||||
| 193 | Haemulopsis elongatus (Steindachner, 1879) | P | x | |||||||||||
| 194 | Haemulopsis leuciscus (Günther, 1864) | P | x | x | ||||||||||
| 195 | Haemulopsis nitidus (Steindachner, 1869) | P | x | |||||||||||
| 196 | Orthopristis chalceus (Günther, 1864) | P | x | |||||||||||
| 197 | Pomadasys bayanus Jordan & Evermann, 1898 | P | x | |||||||||||
| 198 | Pomadasys branickii (Steindachner, 1879) | P | x | |||||||||||
| 199 | Pomadasys macracanthus (Günther, 1864) | P | x | x | ||||||||||
| 200 | Pomadasys panamensis (Steindachner, 1876) | P | x | |||||||||||
| L Family Polynemidae | ||||||||||||||
| 201 | Polydactylus approximans (Lay & Bennett, 1839) | P | x | x | ||||||||||
| 202 | Polydactylus opercularis (Gill, 1863) | P | x | x | ||||||||||
| LI Family Sciaenidae | ||||||||||||||
| 203 | Aplodinotus grunniens Rafinesque, 1819 | P (V) | x | x | x | x | x | x | x | x | ||||
| 204 | Bairdiella armata Gill, 1863 | P | x | |||||||||||
| 205 | Bairdiella ensifera (Jordan & Gilbert, 1882) | P | x | x | ||||||||||
| 206 | Bairdiella icistia (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 207 | Cynoscion albus (Günther, 1864) | P | x | x | ||||||||||
| 208 | Cynoscion stolzmanni (Steindachner, 1879) | P | x | |||||||||||
| 209 | Cynoscion xanthulus Jordan & Gilbert, 1882 | P | x | |||||||||||
| 210 | Elattarchus archidium (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 211 | Isopisthus remifer Jordan & Gilbert, 1882 | P | x | |||||||||||
| 212 | Larimus effulgens Gilbert, 1898 | P | x | |||||||||||
| 213 | Menticirrhus elongatus (Günther, 1864) | P | x | |||||||||||
| 214 | Menticirrhus nasus (Günther, 1868) | P | x | x | ||||||||||
| 215 | Menticirrhus panamensis (Steindachner, 1876) | P | x | |||||||||||
| 216 | Micropogonias altipinnis (Günther, 1864) | P | x | x | ||||||||||
| 217 | Micropogonias megalops (Gilbert, 1890) | P | x | |||||||||||
| 218 | Nebris occidentalis Vaillant, 1897 | P | x | |||||||||||
| 219 | Paralonchurus goodei Gilbert, 1898 | P | x | |||||||||||
| 220 | Stellifer cf. walkeri | P | x | |||||||||||
| LII Family Mullidae | ||||||||||||||
| 221 | Pseudupeneus grandisquamis (Gill, 1863) | P | x | |||||||||||
| LIII Family Kyphosidae | ||||||||||||||
| 222 | Kyphosus elegans (Peters, 1869) | P | x | x | ||||||||||
| LIV Family Chaetodontidae | ||||||||||||||
| 223 | Chaetodon humeralis Günther, 1860 | P | x | x | ||||||||||
| LV Family Cichlidae | ||||||||||||||
| 224 | Amphilophus trimaculatus (Günther, 1867) | SF | x | x | x | x | ||||||||
| 225 | Astatheros macracanthus (Günther, 1864) | SF | x | x | x | x | ||||||||
| 226 | Chiapaheros grammodes (Taylor & Miller, 1980) | SF | x | x | ||||||||||
| 227 | Cincelichthys pearsei (Hubbs, 1936) | SF | x | x | x | x | x | x | x | |||||
| 228 | Chuco intermedium (Günther, 1862) | SF | x | x | x | x | x | x | x | |||||
| 229 | Cribroheros robertsoni (Regan, 1905) | SF | x | x | ||||||||||
| 230 | Kihnichthys ufermanni Allgayer, 2002 | SF | x | x | x | |||||||||
| 231 | Maskaheros argenteus (Allgayer, 1991) | SF | x | x | x | x | ||||||||
| 232 | Maskaheros regani (Miller, 1974) | SF | x | x | ||||||||||
| 233 | Mayaheros urophthalmus (Günther, 1862) | SF | x | x | x | x | x | |||||||
| 234 | Oreochromis aureus (Steindachner, 1864) Ex | Ex | x | |||||||||||
| 235 | Oreochromis mossambicus (Peters, 1852) Ex | Ex | x | x | x | |||||||||
| 236 | Oreochromis niloticus (Linnaeus, 1758) Ex | Ex | x | x | x | x | x | x | x | x | x | x | x | |
| 237 | Oscura heterospila (Hubbs, 1936) | SF | x | x | x | |||||||||
| 238 | Parachromis friedrichsthalii (Heckel, 1840) | SF | x | x | x | |||||||||
| 239 | Parachromis managuensis (Günther, 1867) Ex | Ex | x | x | x | x | x | x | x | x | ||||
| 240 | Paraneetroplus gibbiceps (Steindachner, 1864) | SF | x | x | ||||||||||
| 241 | Petenia splendida Günther, 1862 | SF | x | x | x | x | x | x | x | |||||
| 242 | Rocio ocotal Schmitter-Soto, 2007 | SF | x | |||||||||||
| 243 | Rocio octofasciata (Regan, 1903) | SF | x | x | x | x | x | |||||||
| 244 | Theraps irregularis Günther, 1862 | SF | x | x | x | x | ||||||||
| 245 | Thorichthys meeki Brind, 1918 | SF | x | x | x | x | ||||||||
| 246 | Thorichthys pasionis (Rivas, 1962) | SF | x | x | x | |||||||||
| 247 | Thorichthys socolofi (Miller & Taylor, 1984) | SF | x | x | ||||||||||
| 248 | Thorichthys helleri (Steindachner, 1864) | SF | x | x | x | x | x | x | x | x | ||||
| 249 | Trichromis salvini (Günther, 1862) | SF | x | x | x | x | x | x | X | |||||
| 250 | Tilapia zilli (Gervais, 1848) Ex | Ex | x | x | x | |||||||||
| 251 | Rheoheros coeruleus (Stawikowski & Werner, 1987) | SF | x | |||||||||||
| 252 | Rheoheros lentiginosus (Steindachner, 1864) | SF | x | x | x | x | x | x | ||||||
| 253 | Vieja bifasciata (Steindachner, 1864) | SF | x | x | x | x | x | x | ||||||
| 254 | Vieja breidohri (Werner & Stawikowski, 1987) | SF | x | |||||||||||
| 255 | Vieja guttulata (Günther, 1864) | SF | x | |||||||||||
| 256 | Vieja hartwegi (Taylor & Miller, 1980) | SF | x | x | x | x | ||||||||
| 257 | Vieja melanura (Günther, 1862) | SF | x | x | x | x | x | x | x | |||||
| 258 | Wajpamheros nourissati (Allgayer, 1989) | SF | x | x | x | |||||||||
| LVI Family Pomacentridae | ||||||||||||||
| 259 | Abudefduf troschelii (Gill, 1862) | P | x | x | ||||||||||
| 260 | Stegastes flavilatus (Gill, 1862) | P | x | |||||||||||
| LVII Family Labridae | ||||||||||||||
| 261 | Halichoeres aestuaricola Bussing, 1972 | P | x | |||||||||||
| 262 | Halichoeres dispilus (Günther, 1864) | P | x | |||||||||||
| LVIII Family Scaridae | ||||||||||||||
| 263 | Nicholsina denticulata (Everman & Radcliffe, 1917) | P | x | |||||||||||
| LIX Family Dactyloscopidae | ||||||||||||||
| 264 | Dactyloscopus lunaticus Gilbert, 1890 | P | x | x | ||||||||||
| 265 | Dactyloscopus amnis Miller & Briggs, 1962 | P | x | |||||||||||
| LX Family Eleotridae | ||||||||||||||
| 266 | Dormitator latifrons (Richardson, 1844) | P | x | x | ||||||||||
| 267 | Eleotris picta Kner, 1863 | P | x | x | ||||||||||
| 268 | Erotelis armiger (Jordan & Richardson, 1895) | P | x | x | ||||||||||
| 269 | Gobiomorus dormitor Lacepéde, 1800 | P | x | x | x | x | x | |||||||
| 270 | Gobiomorus maculatus (Günther, 1859) | P | x | x | ||||||||||
| 271 | Guavina micropus (Ginsburg, 1953) | P | x | |||||||||||
| 272 | Leptophilypnus guatemalensis Thacker & Pezold, 2006 | P (V) | x | x | ||||||||||
| LXI Family Gobiidae | ||||||||||||||
| 273 | Aboma etheostoma Jordan & Starks, 1895 | P | x | x | ||||||||||
| 274 | Awaous transandeanus (Günther, 1861) | P (Ca) | x | |||||||||||
| 275 | Barbulifer mexicanus Hoese & Larson, 1985 | P | x | |||||||||||
| 276 | Bathygobius andrei (Sauvage, 1880) | P | x | x | ||||||||||
| 277 | Ctenogobius sagittula (Günther, 1862) | P | x | x | ||||||||||
| 278 | Evorthodus minutus Meek & Hildebrand, 1928 | P | x | x | ||||||||||
| 279 | Gobioides peruanus (Steindachner, 1880) | P | x | |||||||||||
| 280 | Gobionellus liolepis (Meek & Hildebrand, 1928) | P | x | |||||||||||
| 281 | Gobionellus microdon (Gilbert, 1892) | P | x | x | ||||||||||
| 282 | Microgobius miraflorensis Gilbert & Starks, 1904 | P | x | x | ||||||||||
| 283 | Parrella lucretiae (Eigenmann & Eigenmann, 1888) | P | x | |||||||||||
| 284 | Sicydium salvini Ogilvie-Grant, 1884 | P (Ca) | x | |||||||||||
| LXII Family Microdesmidae | ||||||||||||||
| 285 | Microdesmus dorsipunctatus Dawson, 1968 | P | x | x | ||||||||||
| 286 | Microdesmus suttkusi Gilbert, 1966 | P | x | |||||||||||
| LXIII Family Ephippidae | ||||||||||||||
| 287 | Chaetodipterus zonatus (Girard, 1858) | P | x | x | ||||||||||
| 288 | Parapsettus panamensis (Steindachner, 1876) | P | x | |||||||||||
| LXIV Family Acanthuridae | ||||||||||||||
| 289 | Acanthurus xanthopterus Valenciennes, 1835 | P | x | |||||||||||
| LXV Family Sphyraenidae | ||||||||||||||
| 290 | Sphyraena ensis | P | x | |||||||||||
| LXVI Family Trichiuridae | ||||||||||||||
| 291 | Trichiurus nitens Garman, 1899 | P | x | |||||||||||
| LXVII Family Scombridae | ||||||||||||||
| 292 | Scomberomorus sierra Jordan & Starks, 1895 | P | x | x | ||||||||||
| Order Pleuronectiformes | ||||||||||||||
| LXVIII Family Paralichthydae | ||||||||||||||
| 293 | Citharichthys gilberti Jenkins & Evermann, 1889 | P | x | x | ||||||||||
| 294 | Cyclopsetta panamensis (Steindachner, 1876) | P | x | |||||||||||
| 295 | Etropus crossotus Jordan & Gilbert, 1882 | P | x | |||||||||||
| 296 | Syacium latrifons (Jordan & Gilbert, 1882) | P | x | |||||||||||
| 297 | Syacium ovale (Günther, 1864) | P | x | |||||||||||
| LXIX Family Achiridae | ||||||||||||||
| 298 | Achirus mazatlanus (Steindachner, 1869) | P | x | x | ||||||||||
| 299 | Achirus scutum (Günther, 1862) | P | x | x | ||||||||||
| 300 | Achirus zebrinus Clark, 1936 | P | x | |||||||||||
| 301 | Trinectes fimbriatus (Günther, 1862) | P | x | |||||||||||
| 302 | Trinectes fonsecensis (Günther, 1862) | P | x | x | ||||||||||
| LXX Family Cynoglossidae | ||||||||||||||
| 303 | Symphurus chabanaudi Mahadeva & Munroe, 1990 | P | x | |||||||||||
| 304 | Symphurus elongatus (Günther, 1868) | P | x | |||||||||||
| 305 | Symphurus melanurus Clark, 1936 | P | x | |||||||||||
| Order Tetraodontiformes | ||||||||||||||
| LXXI Family Balistidae | ||||||||||||||
| 306 | Pseudobalistes naufragium (Jordan & Starks, 1895) | P | x | x | ||||||||||
| LXXII Family Tetraodontidae | ||||||||||||||
| 307 | Arothron meleagris (Bloch & Schneider, 1801) | P | x | |||||||||||
| 308 | Sphoeroides annulatus (Jenyns, 1842) | P | x | x | ||||||||||
| 309 | Sphoeroides rosenblatti Bussing, 1996 | P | x | x | ||||||||||
| LXXIII Family Diodontidae | ||||||||||||||
| 310 | Diodon holocanthus Linnaeus, 1758 | P | x | |||||||||||
| 311 | Diodon hystrix Linnaeus, 1758 | P | x | x | ||||||||||
| Total species by geographical units | 23 | 31 | 45 | 55 | 36 | 46 | 11 | 63 | 54 | 72 | 174 | 153 | ||
Of the 12 geographical units (Fig. 1), the region with the highest number of species was Costa-Itsmo with 174 species, followed by Costa-Soconusco with 153 species and the third was Usumacinta-Catazajá with 72 species. The region with the lowest recorded species was Usumacinta-Jataté with only 11 species. Numbers of species from other geographical units are presented in Table 2. Spatially, Astyanax aeneus and Rhamdia guatemalensis appeared in all regions within Chiapas. Other species with widespread distributions were Poecilia sphenops and the exotic cichlid Oreochromis niloticus (10 and 11 regions respectively). Atherinella alvarezi, Brycon guatemalensis, Dorosoma anale, Dorosoma petenense, and Ictalurus meridionalis were distributed in nine regions, while Aplodinotus grunniens, Gambusia sexradiata, Ophisternon aenigmaticum, Parachromis managuensis, Poecilia mexicana, Pseudoxiphophorus bimaculatus, and Thorichtys helleri were recorded in eight regions.
Eight marine species were newly recorded as species found in continental waters of Chiapas: Acanthurus xanthopterus, Atherinella panamensis, Fistularia commersonii, Halichoeres dispilus, Nicholsina denticulata, Orthopristis chalceus, Stegastes flavilatus, and Sphyraena ensis.
Discussion
Knowledge of the species richness of continental fishes in Chiapas has increased significantly over recent years compared to previous assessments (e.g. Rodiles-Hernández et al. 2005, Velázquez-Velázquez et al. 2013). The increasing number of known species is the result of collections in new localities, improvement in sampling effort, and larger systematic and taxonomic reviews. For instance, an extensive literature search provided many reports of marine species, principally elasmobranchs, in continental waters of Chiapas by Castro-Aguirre et al. (1999). The large increment in the checklist is due to the inclusion of many elasmobranchs fishes that were included previously in the work of Castro-Aguirre et al. (1999), but that for some reason these records were ignored in more recent accounts of fishes in the continental waters of Chiapas. Castro-Aguirre et al. (1999) reported 41 species of marine fishes including an important number of sharks and sting-rays in the state continental water.
Two species previously reported were removed from the list of species in Chiapas in this study: the American eel (Anguilla rostrata) and the Mexican tetra (Astyanax mexicanus). The American eel was mentioned in the pioneering work of Velasco-Colín (1976), and since then listed in subsequent publications (Lozano-Vilano and Contreras-Balderas 1987, Rodiles-Hernández 2005, Rodiles-Hernández et al. 2005, Espinosa-Pérez et al. 2011, Velázquez-Velázquez et al. 2013). However, these works do not offer precise geographical locations for these species and there are no vouchered specimens from Chiapas in national or international collections. Records of the Mexican tetra in Chiapas probably contain misidentifications as mentioned by Lozano-Vilano and Contreras-Balderas (1987) and Ornelas-García et al. (2008), thus supporting the absence of this species in Southern Mexico. We have included Important and recent taxonomic changes made in the family Cichlidae by McMahan et al. (2015) and Říčan et al. (2016), the family Poeciliidae by Palacios et al. (2016) and the family Profundulidae by Morcillo et al. (2016).
More than 1000 species of fishes have been reported in the continental waters of Mexico, including freshwater and estuarine fishes (Espinosa-Pérez 2014). The continental fish fauna of the state of Chiapas represents approximately 29% of the continental fish fauna of the entire country of Mexico. This highlights the great diversity of fishes inhabiting continental environments of Chiapas as a result of the region’s hydrological wealth. Our results are comparable with those from other southern Mexican states such as Quintana Roo (Schmitter-Soto 1998), Oaxaca (Martínez-Ramírez et al. 2004) and Tabasco (Espinosa-Pérez and Daza-Zepeda 2005).
The native obligate freshwater (primary and secondary) species of Chiapas accounted for only 26% (81) of the state’s total species richness. The communities are dominated by peripheral species, many of them permanent (vicarious) residents of the Grijalva-Usumacinta basin (e.g. Aplodinotus grunniens, Eugerres mexicanus, Hyporhamphus mexicanus, Strongylura hubbsi), but the majority are distributed in brackish environments of the Costa-Itsmo and Costa-Soconusco sub-basins. Some of these communities also permeate nearby rivers. In terms of slopes, the Pacific slope houses 68% of the state fish fauna while the Gulf slope houses 33%, and in terms of regional diversity the Usumacinta region is considered one of the most diverse areas of endemism for freshwater fishes in Central America; however, from a biogeographical perspective the entire Central American region has a depauperate freshwater fish fauna compared with the vast diversity of ostariophysan fishes found in North and South America (Miller 1966, Myers 1966, Bussing 1985, Chakrabarty and Albert 2011, Matamoros et al. 2015). This could explain the presence of a great number of peripheral species recorded in the continental environments of Chiapas. This pattern is comparable with other countries of Central America such as Guatemala (Kihn-Pineda et al., 2006), Honduras (Matamoros et al. 2009) and El Salvador (McMahan et al. 2013).
Mexican law protects thirteen freshwater species; however, Rhamdia guatemalensis is quite abundant in Chiapas and possesses a wide distribution through other geographic areas of Mexico and Central America (Miller et al. 2005, Hernández et al. 2015). Its inclusion should be reconsidered in the NOM-059-SEMARNAT-2010. Conversely, we suggest that Mexican laws should consider including Lacantunia enigmatica, Rhamdia laluchensis and Vieja breidohri as protected species on the grounds of their restricted distribution.
Since the pioneering work of Lozano-Vilano and Contreras-Balderas (1987), this is the first time the state of Chiapas has been regionalized in a more detailed scale than the three great basins (Grijalva, Usumacinta and Costa). Lozano-Vilano and Contreras-Balderas (1987) proposed seven physiographic regions; however, their proposal was based on physiographic characteristics of landscape relief rather than hydrology. In this study we present a zonation based on the level of hydrological regions (sub-basins), which provides a more robust delineation of the geographical areas for fish species and facilitates a closer examination of the distribution of endemic species. This approach demonstrates that gaps in knowledge of the distribution of species is still quite large and indicates that some portions of the territory remain moderately sampled or unexplored. For instance, the Usumacinta-Jataté sub-basin, with only 11 species recorded, remains largely unexplored. The detailed regionalization of Chiapas highlights the necessity of increasing sampling efforts in certain zones.
Although hydrological regions Grijalva, Usumacinta and Costa of Chiapas have been used in previous studies to discover endemism in the state (Rodiles-Hernández 2005, Rodiles-Hernández et al. 2005, Velázquez-Velázquez et al. 2013), the zonation of our study allows identification of smaller geographic units, permitting us to be more specific in studies of endemism. Thus, the distribution of endemic species in Chiapas includes: Lacantunia enigmatica in Usumacinta-Lacantún, Rocio ocotal in Usumacinta-Lacantún, Thorichthys socolofi in Grijalva-Tulijá and Usumacinta-Lacantún, Tlaloc hildebrandi in Grijalva-Teapa and Usumacinta-Jataté, and Poecilia thermalis in Grijalva-Teapa. Of the 12 units, Usumacinta-Lacantún stands out as it houses three endemic species: Lacantunia enigamatica, Rocio ocotal, and Thorichthys socolofi.
Forty years of scientific research on the continental fish fauna of Chiapas has gone a long way since the work of Velasco-Colín (1976). However, this does not seem nearly enough time to completely finish to record the real extend of the state species richness with its distribution. In this work we present distributional data at 12 geographic units. However, although this is the finest distributional scale for the state, a major goal should be to complete distributional data for the 92 existing sub-drainages in the state. Many of these water bodies have never been sampled either for lack of financial resources or because they are located in remote areas of the state.
Acknowledgements
This work has been possible due to several research grants to MZ-P-UNICACH from (CONACYT), (CONABIO) and (CONANP). Recent ichthyological explorations in Chiapas have been possible by grants provided by the (CFE). We are very grateful to all members of the UNICACH Museum of Zoology for their field and lab contributions. We are in particular debt to UNICACH for their unconditional support of our research initiatives. Thanks to Allison M. Matamoros and Caleb D. McMahan for proofreading the manuscript.
Citation
Velázquez-Veláquez E, López-Vila JH, Gómez-González AE, Romero-Berny EI, Lievano-Trujillo JL, Matamoros WA (2016) Checklist of the continental fishes of the state of Chiapas, Mexico, and their distribution. ZooKeys 632: 99–120. doi: 10.3897/zookeys.632.9747
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