Checklist of the continental fishes of the state of Chiapas, Mexico, and their distribution.

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.

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 PageBreakand 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 . 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).

Museum of Zoology at the University of Arts and Sciences of Chiapas

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 (, , , , , , , , , , , and ). Only five species were endemic: the catfish , the cichlids and , the killifish and the molly PageBreak. Based on species richness the most important families were: (35), (29), (18), (17), (16), (12), and (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): and are listed as endangered; , , and are listed as threatened; finally , , , , , and are listed as species under special protection. Based on general salinity PageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreakPageBreaktolerance, 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.

(Primary Freshwater)

(Secondary Freshwater)

(Peripheral Vicarious)

(Peripheral Catadromous)

(Peripheral)

(Exotic)

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 PageBreakthe 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, and appeared in all regions within Chiapas. Other species with widespread distributions were and the exotic cichlid (10 and 11 regions respectively). , , , , and were distributed in nine regions, while , , , , , , and were recorded in eight regions.

Eight marine species were newly recorded as species found in continental waters of Chiapas: , , , , , , , and .

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 () and the Mexican tetra (). 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 by McMahan et al. (2015) and Říčan et al. (2016), the family by Palacios et al. (2016) and the family 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. , , , ), 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, 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 , and 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 unexPageBreakplored. 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: in Usumacinta-Lacantún, in Usumacinta-Lacantún, in Grijalva-Tulijá and Usumacinta-Lacantún, in Grijalva-Teapa and Usumacinta-Jataté, and in Grijalva-Teapa. Of the 12 units, Usumacinta-Lacantún stands out as it houses three endemic species: , , and .

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.