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Composition and Derivation of the Freshwater Fish Fauna of Mexico

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Composition and Derivation of the Freshwater Fish Fauna of Mexico Composition and Derivation of the Freshwater Fish Fauna of Mexico ROBERT RUSH MILLER Museum of Zoology The University of Michigan Ann Arbor, Michigan 48109 U. S. A. MILLER, ROBERT RUSH, 1986. Composition and derivation of the freshwater fish fauna of Mexico. An. Esc. nac. Cienc. biol.. Mex.. 301 121-153. SUMMARY: The 375 species of freshwater fishes herein treated are primary (132). se- condary (186) and vicarious (57). The species are listed along with their general habitats and distributions and whether they are endemic to Mexico or to those drainages not wholly within Mexico. Centers of endemism are briefly discussed. The derivation of the fauna is: American tropics, 16 percent; northern, 40 percent; Middle American. 19 percent (all poe- ciliids); autochthonous, 10 percent (Goodeidae); and vicarious. 15 percent. This summary, especially the information presented in Table 1, has been made possible through the accumulation of knowledge built up by scientists over a long time. My friend and colleague, Jose Alvarez del Villar, has contributed to our understanding of Mexican ichthyology over a period exceeding three decades (1946-1977). His contributions have varied from descriptions of Recent and fossil taxa to philosophical consideration of speciation and of the correlation between earth history and present fish distribution. It is a pleasure to acknowledge his work. INTRODUCTION With an area only about one-fifth that of the continental United States. Nle- xico possesses a rich and diversified freshwater fish fauna comprising about 300 species in 47 families (Miller, in prep.). This is nearly 60 percent of the fresh- water fishes inhabiting the United States and Canada. The species composi- lion is now well known although new discoveries continue to be made and much information on life history and ecology is badly needed. In the present account only 375 species. all virtually confiner' to fresh water. are considered in treating the makeup and origin of the fauna. Diversification' of the Mexican freshwater fish fauna stems from many factors. among which are: (1) the highly varied physical geography (at least three- ° fifths of Mexico is of high relief) ; (2) great latitudinal extent (32 30' N in the NVV to 14° 30' N in the SE): (3) isolation of the large tropical highland (Mesa Central) that contains the important Rio Lerma fauna (Miller & Smith. 1986) ; (4) adaptation by many marine groups to fresh water: and (5) the presence in the southeast of the largest river system in Middle America. the Usu.- rnacinta-Grijalva basin, that lies well within the tropics. It will be shown that the Mexican fish fauna is derived about equally from Nearctic/Holarctic sources 121 122 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS. VOL. 30 and Neotropical or Middle American 1 elements, and that species of marine ancestry and the autochthonous family Goodeidae are also nearly equally re- presented. More than half of Mexico includes desert and semi-desert areas that comprise a water-deficient, ecologically unstable, and frequently harsh environment conta- ining a fascinating fish fauna (Smith & Miller 1986). Geologically recent arid- ity (during the past 8,000 to 10,000 years) has reduced this fauna by extinction of taxa. Three groups of freshwater fishes are included in Table 1 (marked I. II, Ill). as defined by Myers (1938, 1949): Primary, those that are almost exclusively restricted to fresh water: Secondary, species that can tolerate some salinity (even up to 35 ppt., as in Cichlasoma fenestratum and C. urophthalmus) and are able to cross narrow sea barri7;_i and Vicarious, those with marine affinities but now largely or exclusively adapted to fresh water. Some secondary freshwater fishes are known only from fresh water (e.g., the entire family Goodeidae) and may not be able to tolerate salinities greater than 5 ppt., whereas a few primary freshwater fishes (e.g., Astyanax) are known to inhabit brackish water. Some mullets (Agonostomu,s and Joturus) and gobies (Sicydium), may spend their larval and early post- larval stages in salt or brackish water, but are here in- cluded in the vicarious group (III). All three groups are listed in Table 1. along with their habitats and distributions. Variations in salt tolerance should be considered when discussing zoogeography. COMPOSITION AND ORIGIN OF FAUNA Primary freshwater fishes are represented by eight families with 37 genera and 132 species. Within these families the most speciose genera are: Cyprinidae. Notropis with 25 species; Catostomidae. Catostomus with 10 species: Ictaluri- dae, Ictaturus with 10 species. Secondary fishes are also represented by eight families, but have more genera (45) and species (186). The large genera in these families are: Cyprinodontidae. Cyprinodon with 18 species: Poeciliidae. Gambusia with 19 species; and Cichlidae. Cichlosoma with 40 species. Marine derivatives comprise 15 phylogenetically diverse families (lampreys to gobies) with 22 genera and 57 species: the most prolific genus is Chirostoma (Atheri- nidae) with 19 species. Among the primary and secondary fishes, which have a combined total of 319 species, six genera in six families account for 131 species or 41 percent of the total. The earliest known fossil record for Primary fishes is in the Pliocene Chapala Formation, and includes one living species. Notropis sallei, and one extinct species. Micro pterus relictus. For Secondary fishes the goodeid Tapatia occiden- tails occurs in the late Miocene Santa Rosa Formation. For a marine invader. the earliest known fossils occur in the Pliocene Chapala Formation, species of 1 Middle America is defined as comprising Mexico and Central America. Miller, R. Robert, FRESHWATER FISH OF MEXICO 123 the genus Citrostorna. Fossil salmonids are known from late Pleistocene depos- its only (Miller and Smith, 1986). The derivation of the Mexican freshwater fish fauna is broadly categorized as follows: (1) 61 species from the American tropics. comprising 14 primary and 47 secondary fishes, or 16 percent; (2) 148 species of northern origin (Nearctic or Holarctic). including 118 primary and 30 secondary fishes, or 40 percent; (3) Middle American. the 71 species of the Poeciliidae, 19 percent; (4) autoch- thonous. the 36 goodeids, 10 percent; and (5) the 57 vicarious species. represent- ing marine invaders, or 15 percent. CENTERS OF ENDEMISM The great geographic diversity and isolation of drainages have led to a high degree or endemism in the Mexican fish fauna. The major centers are: (1) the Rio Lerma-Santiago basin, with 58 percent for the entire drainage. but 66 percent when considering only the fishes of the Lerma basin above Salto de Juanacatlan (thus eliminating a single species each of Gobiesocidae, Atherinidae, Cichlidae. Eleotridae. and Gobiidae). (2) Rio Usumacinta-Grijalva basin. 36 percent, with major endemism among the poeciliids and cichlids. (3) Rio Pimlico basin. 30 percent, with a "species flock" in the genus Dioncla and seven endemic poeciliids. (4) Rio Balsas basin, 35 percent. including 7 endemic species in 10 families (of which 3 are goodeids and 2 poeciliids). (5) Rio Ameca basin, 32 percent. with all of the endemic species (cyprinids and goodeids) confined to the upper part of the river, east of 104° 45' N (above barrier falls?). (6) Rio Papaloapan basin. 21 percent. with 6 endemic species of poeciliids; the adjacent Rio Coatza- coalcos has only 13 percent endemism but shares 4 species with the PapaToenail that are not found elsewhere (Riaulus robustus, Atherinella. sallei, Cichlasoma ellioti, and C. fenestratum). (7) Rio Conchos basin, although tributary to Rio Grande, has a surprising number of -endemics. 21 percent (7 of 34 species). (8) Rio del Tunal (the upper part of Rio Mezquital near Durango City) has 8 of 13 species (62 %) endemic. chiefly cyprinids. There are three remarkable basins, two of them endorheic, that show high degrees of local endemism and deserve special mention because of their cons- iderable scientific interest. Two of them, unfortunately, have already been stressed by man's activities. In terms of diversity (not only of fishes, but of plants and invertebrates), the Cuatro Clinegas bolson is exceptional (Minckley. 1969, 1978), with about 50 percent of its fishes endemic (8 species in 8 families). The second endorheic basin, Laguna Chichancanab in Yucatan, is unique for its endemic species flock of 5 Cyprinodon (Humphries, 1084b), with only one other fish (a poecilikl) inhabiting the basin. The third localized area is La Media Luna and environs, in the upper Rio Verde. San Luis Potosi. This remarkable eco- system of spring-fed lagunas and marshes, contains 6 endemic species among the cyprinkls. cyprinodontoids, and cichlids, or 67 percent (Williams et cd., 1985). All three of these areas need to be protected against loss of their diversity. 124 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS, VOL. 30 EXPLANATION OF TABLE I Table I lists the families of native Mexican freshwater fishes, according to the groupings (1-M) previously discussed, gives their general habitats, their dis- tributions by drainages or regions, and indicates by an E in column one whether Axe species is endemic (restricted) to Mexico. Those species that occur only in the Rio Grande.basin (labelled E in column three) but also live in the United States are not listed as endemic to Mexico. Examples are: Cyprinidae: Dioncla Notropis braytoni, Notropis chihualtua; Cyprinodontidae: Cyprinodon exiinius; Poedliidae: Gambusia senilts; and Percidae: Etheostoma grahami. The symbol Ex means that the species is extinct, either throughout its range (Evarra bustamantei, E. eigenmanni, E. tialtuacensis, Notropis orca, Stypoclon signifer, Cyprinoclon latifasctatus, Skiffia francesae) or only in Mexico (Scaphir- hynclucs platorynchas, Gila elegans, G. intermeclia, Hybopsis gractlis, Notropis simus. Ptychochedus /ucius, Rhinichthys osculus, Tiaroga cobitis, Catostomus c:larki, Xyrauchen texanus). The symbol XI (Dioncta episcopa) means that I believe this species was errone- ously recorded from Parras, Coahuila (see Miller, Williams, and Williams. in Prep.). I follow Wiley (1976) in recognizing two genera of gars.
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