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. Among the Characidae. I tentatively separate three species of Astyanax and agree with Salvador Contreras-Balderas that the "Astyanax" of Laguna Cate- maco. Veracruz, is an undescribed species best referrable to Bramochanuc. Placement of Notropis ornatus in the genus Cocloma is explained in Miller (1978). Separation of Hybopsis from Notropis follows U. S. usage (Robins et at, 1980). Recognition of Notropis formosus as a species distinct from N. lutrensis is based on Chernoff and Miller (1982). • Catostonms conchos Meek is regarded as a synonym of C. bernardini Girard (Miller, In prep.); thus the range of the latter species includes the Rio Grande (Rio Conchos). Ictiobus mericlionalis Gunther is only tentatively recognized (it may be a synonym of I. bubatus (Rafinesque)). The Mexican species of Moxos- toma (under review by R. E. Jenkins and M. L. Smith) are poorly understood. . In the lctaluridae. the American catfishes commonly called bullheads are re- ferred to Amturus since Ictalurus is diphyletic (Lundberg. 1982) and preliminary biochemical studies (Yates et al., 1984) suggest that Amiurus has major genetic differences from Ictidurus. Ictalurus australis (Meek) may be the same as I. punc- talus (Rafinesque); the Mexican species of this genus need much further study. • I follow Taylor and Menezes (in Fischer. 1978) for generic assignments among the Arliclae. Mexican species of Rhamclia (family Pimelodidae) have recently been revie- wed (Miller, 1984).' • Hyporhamphus paws Miller is a synonym of H. rosae (Jordan & Gilbert) [Collette: pers. comm.. 19831: and since that species occurs as well in brackish and salt water it is not treated here. Miller, R. Robert. FRESHWATER FISH OF MEXICO 125

Cyprinodon pachycephalus is being described by Minckley and Minchley from hot springs and their outflows near San Diego de /Ocala, Chihuahua. Cypri- notion verecundus was recently described by Humphries (1984a) as part of the species flock of five pupfishes inhabiting Laguna Chichancanab, Yucatan. Chapalichthys peraticus, family Goodeidae, is listed as endemic to the Rio Lerma basin although it inhabits endorheic Laguna San Juanico (Alvarez, 19153). just outside of the Lerma-Santiago system (other Lerma fishes in that basin are Alloophorus, Goodea, Poeciliopsis infans, and two species of Chinostoma). In the Poeciliidae. Poecilia pallicla de Buen may be an older name for the re- cently described P. maylancli Meyer (1983). A key to Mexican species of Poecilia (except P. pallic/a) was published recently (Miller, 1983). Poeciliopsis hnilickai was described by Meyer and Vogel (1981) from the upper Rio Grijalva basin. The genus XenocIezia is known from a single Guatemalan species (Hubbs. 1950) but an undescribed species (D. E. Rosen. pers. comm.. 1984) is known from streams in Guatemala that flow into Mexico where they surely contain this fish. The classification by Rosen (1979) is followed for Xiphophorus, and of Chernoff (1983) for the Atherinidae. Chirostoma urge, listed as a single species, is a complex of at least two speci,es under study by Barbour and Chernoff. In the Cichlidae. Cichlasoma socolofi is being described by Miller and Taylor (1C84). Taylor a'nd Miller (1980, 1983) earlier described Cicblasoma grammo- ch,s. C. hartwegi, and C. pantostict urn. In the Mugilidae, the bobo or jotura. Joturus pichardoi, was named by Poey for Don Esteban Picharclo; hence the name is hereby emended. The eleotrid genus Leptaphilypnus has not been taken in Mexico yet but un- doubtedly occurs there. Most of the drainages listed in the table are self evident. On the Atlantic slope, Veracruzan Coastal inclucks (from north to south) Rios Tuxpan, Cazo- nes, Tecolutla, Nautla, and Misantla; the latter stream marks the southern coastal limit of the Cyprinidae. The Grijalva-Usumacinta includes the lowland region west of the Grijalva as far as Rio Tonala. and the base of the Yucatan Penin- sula, north to about the latitude of Escarcega (18° 30' N). The Yucatan Pe- ninsula is all the area north of 18° 30' N. On the Pacific slope. Lower Colorado includes the Mexican part of the Rio Colorado and its distributaries and the parts of the Rios Santa Cruz and San Pedro that lie in Sonora. Baja California includes all of the Peninsula plus those streams draining into the Gulf of Cali- fornia that lie north of 'Rio Yaqui and east of Rio Colorado. Sineloan Coastal encompasses the streams between Rio Yaqui and Rio Grande de Santiago. Ja- liscoan Coastal includes those streams between Rios Ameca and Armeria, and Guerreroan Coastal those streams between Rios Balsas and Verde. Among the interior (endorheic) basins, the last six listed in.Table 1. Guzman Complex. includes Rios Cases Grandes. Santa Maria, del Carmen. and Laguna Bustillos. El Salado has fish lifee only at El Potosi and near Galeana, west and south of Cerro Potosi. Nuevo Leon. The Oriental region or Puebla Plateau is the high. arid region that lies east and northeast of Puebla (Alvarez. 1950, 1972). 126 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS, VOL. 30

RESUMEN

Las 375 especies de peces de agua dulce que se consideran son: primarias (132). secundarias (185) y vicarias (57). Las especies son ,mencionadas con su clistribucion y habitat general, seiialando las endemicas o aquellas cuya distribu- cion no esta restringida a Mexico. Los centros de endemism° son brevemente discutidos. La procedencia de la Ictiofauna es: tropical. 16 %; del norte, 40 %; Centroamerica 19 % (todos poecilidos); autoctonos, 10 % (Goodeidae) y ' vica- rios. 15 %. Estetrabajo. y principalmente la tabla 1, ha sido posible gracias al conoci- miento del trabajo de los cientificos. especialmente la de mi amigo y colega. José Alvarez del Villar, quien ha contribuido grandemente a la comprension de la Ict:ofauna mexicana pr un periodo de 30 alios (1946-1977). Sus contribucio- ties incluyen desde descripciones de especies, recientes y f6siles, hasta considera- ciones filosoficas sobre especializacion y correlaciones de la historia de la tierra . y la distribucion actual de los peces. Es un placer par ml reconocer su trabajo.

REFERENCES

ALVAREZ, J., 1950. Contribucieni al conocimiento de los peces de la regi6n de Los Llanos, estado de Puebla (Mexico). An. Esc. nac. Cienc. biol., Mex. 6 (1-4): 81-107. -- 1963. Ictiologia Michoacana. III.-Los peces de San Juanico y de Tocumbo, Mich. An. Esc. nac. Cienc. biol., Mex. 12 (1-4): 111-138. — 1972. Algunos ejemplos de especiacion en peces mexicanos. Acta Politec. Mex., 13 (60): 81-89. CHERNOFF, B., 1983. Revision of the American atherinid fishes, subfamily Menidiinae, and systematics of the nominal subgenus Atherinella. Unpubl. Ph. D. assert., Univ. Michi- gan, Ann Arbor, Mich., U. S. A. 569 pp. CHERNOPF, B., AND R. R. MILLER., 1982. Notropis bocagrande, a new cyprinid fish from Chihuahua, Mexico, with comment; on Notropis formosus. Copeia 1982 (3 ) : 514-522. — 1984. Atherinella ammophila, a -new atherinid fish from eastern Mexico. Notulae Naturae, 462: 1-12, Figs. 1-6. FISCHER, W., 1978. FAO Species Identification Sheets for Fishery Purposes. Western Cen- tral Atlantic ( Fishing Area 31) , Vol. 1. FAO, United Nations, Rome. HUBBS, C. L., 1950. Studies of cyprinodont fishes. XX. A new subfamily from Guatemala. with ctenoid scales and a unilateral pectoral clasper. Misc. Publ. Mus. Zoo!. Univ. Michigan, 78: 1-28, pls. 1-4. HUMPHRIES, J. M., JR. 1984a. Cyprinodon verecundus, n. sp., a fifth species of pupfish from Laguna Chichancanab. Copeia, 1984 (1): 58-68, figs. 1-5. — 1984b. Genetics of speciation in pupfishes from Laguna Chichancanab, Mexico, pp. 129-139. In: Evolution of Fish Species Flocks. A. A. Echelle and I. Kornfield, eds. Univ. Maine at Orono Press. LUNDBERG, J. G., 1982. The comparative anatomy of the toothless blindcat. Trogloglanis pattersoni Eigenmann, with a phylogenetic analysis of the ictalurid catfishes. Misc. Publ. Mus. Zoo!. Univ. Michigan 163: 1-85. MEYER, M. K., 1983. Une nouvelle espece de Poecilia du Guerrero, Mexique ( Pisces: Poeciliidae). Rev. fr. Aquaria, 10 (2 ) : 55-58, figs. 1-7. 'MEYER, M. K., AND D. VOGEL., 1981. Em n neuer Poeciliopsis aus Chiapas, Mexiko ( Pisces: Poeciliidae ). SENCKENBERG. Biol., 61 ( 1980) ( 5/6) : 357-361, figs. 1-5. Miller, R. Robert, ,FRESHWATER FISH OF MEXICO 127

MILLER, R. It., 1978. Compositon and derivation of the native fish fauna of the Chihuahuan Desert region, pp. 365-381. /n: Transactions of the Symposium of the Biological Reso- urces of the Chihuahuan Desert Region,, UnitedUnited States and Mexico, R. H. Wauer and D. H. Riskind, eds. U. S. Natl. Park Trans. and Proc. Ser. 3 (1977). -- 1983. Checklist and key to the mollies of Mexico (Pisces: Poeciliidae: Poecilia, sub- genus Mollienesia). Copeia, 1983 (3): 817-822. - 1984. Rhamdia redclelli, new species, the first blind pimelodid catfish from Middle America, with a key to the Mexican species. Trans. San Diego Soc. Nat. Hist., 20 (8): .135-143, figs. 1-4. - In prep. Guide to the Freshwater Fishes of Mexico. MILLER, R. R., AND M. L SMITH., 1986. Origin and geography of the fish fauna of Central Mexico, pp. 491-519. /n: The Zoogeography of North American Freshwater Fishes, C. R. Hczutt and E. 0.. Wiley, eds. Wiley-Interscience, New York. MILLER, R. R., AND J. N. TAYLOR., 1984. Cichlasoma soco!ofi, a new species of cichlid fish of the Thorichthys group, from northern Chiapas. Mexico. Copeia. 1984 (4): 933-940, Fig. 1. MILLER, R. R., J. E. WILLIAMS, AND J. D. WILLIAMS. In prep. Extinction in North Ame- rican Fishes During the Past Century. MINCKLEY, W. L., 1969. Environments of the Bolson of Cuatro Cienegas, Coahuila. Mexico. with spnial reference to the aquatic biota. Univ. Texas, El Paso, Sci. Ser. 2: 3-65. figs. 1-27. - 1978. Endemic fishes of the Cuatro Cienegas Bcsin, northern Coahuila, Mexico, pp. 383-404, figs. 1-6. In: Transactions of the Symposium of the Biological Resources of the Chihuahuan Desert Region, United States and Mexico, R. H. Wauer and D. H. Riskind, eds. U. S. Natl. Park Serv. Trans. and Proc. Ser. 3 (1977). MYERS, G. S., 1938. Fresh-water fishes and West Indian zoogeography. Ann. Rept. Smith- sonian Inst., 1937: 339-364. - 1949. Salt-tolerance of fresh-water fish groups in relation to zoogeographic problems. Bud. tot de Dierkunde. 28: 315-322. ROBINS, C. R., R. M. BAILEY, C. E. BOND, J. R. BROOKER, E. A. LACHNER, R. N. LEA. AND W. B. Scorr, 1980. A list of common and scientific names of fishes from the United States and Canada. Amer. Fish. Soc. Spec. Bull. 12: 1-174. ROSEN, D. E., 1979. Fishes from the uplands and intermontane basins of Guatemala: Re- visionary studies and comparative geography. Bull. Amer. Mus. Nat. Hist., 162: 269- 376. SMITH, M. L., AND R. R. MILLER, 1986. The evolution of the Rio Grande basin as inferred from its fish fauna, pp. 458-485. /n: The Zoogeography of North American Freshwater Fishes, C. R. Hocutt and E. 0. Wiley, eds. Wiley Interscience, New York. TAYLOR, J. N. AND R. R. MILLER, 1980. Two new fishes, genus Cichrasoma, frcm Mexico. Occ. Pap. Mus. Zool. Univ. Michigan 693: 1-16. - 1983. Cichlid fishes (genus Cichlasoma) of the Rio Panucc basin, easte-n Mc-:;co. with description of a new species. Occ. Pap. Mus. Nat. Hist. Univ. Kansas. 104: 1-24. WILEY, E. 0., 1976. The phylogeny and biogeography of fossil and recent gars ( Act- inopterygii: Lepisosteidae). Univ. Kansas. Mus. Nat. Hist. Misc. Publ. 64: 1-11l. WILLIAMS, J. E., D. B. BowmAN, A. A. ECHELLE, R. J. EDWARDS, D. A. HENDRICKSON, AND J. J. LANDYE. 1985. Endangered aquatic ecosystems in North American deserts with a list of vanishing fishes of the region. Jour. Arizona-Nevada Acad. Sci. YATES. T. L., M. A. LEWIS, AND M. D. HATCH, 1984. Biochemical syste-mtics of three species of catfish (genus Ictalurus) in New Mexico. Copia 1934 (1): 97-101.

El presente articulo fue recibido para su publicacion en febrero de 1985. 128 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS' , VOL. 30

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TABLE 1. (CONTINUED.) I = PRIMARY; II = SECONDARY; III = VICARIOUS; X = PRESENCE; E = ENDEMIC; E X = EXTINCT.

Habitats Distributions by Drainages

0 M C M g

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V Valle de 3 % Spring Isolate P. >. CA •S mst • g U Algansea lacustris-E X X Algansea monticola-E X X Algansea popoche-E X X Algansea tincella-E X X X X X X Campostora anomalum X X Camoostora otnatum X X X X Codoma ornata-E X X X X Dionda catostomops-E X X Dionda diaboli X X X Dionda dichroma-E X X X Dionda episcopa X X X X XI Dionda erimyzonops-E X X Dionda ipni-E X X Dionda mandibularis-E X X Dionda rasconis-E X X Dionda sp.-E X X X X E Evarra bustamantei-E X X X eigenmanni-E X X E Evarra X X STIEAT 4S9P S 04 oixJi

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X X X

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A-PSJADSTIOO EI TO >< X

„Lowland El Tehuantepec-Guatemala Soto laMarina Sinaloan Panuco Papaloapan Guzman CUatro Lower Cbatzacoalcos Jaliscoan Yaqui-Mayo Yucatan Peninsula Tunal-Santiaguillo Balsas Baja California Guerreroan Lerma-Santiago Grijalva-Usumacinta Armerfa-Coahuayana Ameca-Magdalena Valle de Verde-Atoyac Oriental (Puebla) Nazas-Aguanaval Cavernicole Lacustrine Creek Spring Isolate Stream Upland Veracruzan Rio Grande Big River Montane Salado %WHIN

Colorado

Complex Cienegas

Coastal M6xico

Coastal

Coastal Coastal

132 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS, VOL. 30

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=low X EUTIEW ET OQOS X X APUERJ OP; 03 X 41 >C R.1 41 41 ATOOTUAATIE3 A4ETCSI BUTIDS AUTIQSNOE7

NAAA0 X X X X WEAAQS X X X X X X X X X X X X X X X AAATH BTS X X X X X ATREQUOW X X X

X X X X X X X X X X X X PUETMORI X X X X X X X X X

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2 22 22222222222221°1 11

TABLE 1., CONTINUED

HABITATS DISTRIBUTIONS BY DRAINAGES

TAXA §11.1 i Fg M GG'EI! R.

NOTROPIS SALADONIS-E X X ROBERT

NOTROPIS SALLEI-E X X X X NOTROPIS SIMUS , NOTROPIS STRAMINEUS FRESHWATER FISH OF MEXICO NOTROJOIS TROPICUS-E NOTROPIS XANTHICARA-E NOTROPIS SP. -E NOTROPIS SP. -E NOTROPIS SP. -E PIMEPHALES PRCMELAS

PIMEPHALES VIGILAX X X PTYCHOCHEILUS LUCIUS RHINICHTHYS CATARACTAE X RHINICHTHYS OSCULUS E X STYPODON SIGNIFER-E X TIAROGA OOBITIS E X YURIRIA ALTA-E CATOSIOMIME-I CARPIODES CARPIO X X X X X X

TABLE 1. (Continued.) I = Primary; II = Secondary; III = Vicarious; X = Presence; E = Endemic; Ex = Extinct.

Habitats Distributions by Drainages

Taxa 1 S311VNIV 1 1 11 a Agl 4 14 1 Y, VI 30 L i 0001 u klw 1111 4' 1 d Miig A A Ag41P,11 gifi i1 1111A NN gP F.Cg 8 Catostanus bernardini X X X

Cab:1st:mils clarki EX Catostamus insignis X Catosbanus plebeius X X X Catostcmus cf. plebeius-E Catostcmus wigginsi-E Catostamus sp.-E Catostcmus sp.-E

Catostanus sp.-E Catostanus sp.-E Cycleptus elongatus Ictiobus bubalus 'SV31001010

Ictiobus labiosus-E Ictiobus *10A

meridionalis-E X

Ictiobus niger x x x x OE Ictiobus sp.-E Mbxostcma austrinumrE x x x x x x

Moxostcme congestum X X MILLER, R. ROBERT, FRESHWATER FISH OF MEXICO 135

(ETCPRID) TE4UAT20 °° T X9W ETO EITEA OPETEG TA

TENEUENCV-SEMSI XETDOUP UPWZRE SEBEUPTD OAQEM ETEMPENG-OEDEQUENTIPY OEICOW OPAL\ TEGSSOP UVOAEAAANG SEEM EUEAERMIPOO-PLAAWAV

TEWEEO UPOOSTRUR EUETVPBESQ-EOEWV °BE-PUGS-MAW' OTTTRIBETWES-TEUNI TE4SE010 UEOREUTS OAEK-TNBEA ERUAORTTED EFTS OPEAOTOP SAW,' EINSUPAD UPEONA EQUTOEWNSAEATECTIG SOOTEMEZ4EOP UEDEOTEDED TE4SEO3 UEZNAPEAPTI OONUPD EUTAEW ET cows APUWS ORG

ETOOLUAEARD E4ETOSI BUTADS AUTAGSNOEU

XUA-13 WEEAQS XX X X

AETITTI EGG X X X X X AUEQUOM

PUEIDN X X X X PUETMAL X X X X

19

III 136 ANALES DE LA ESCIJELA NACIONAL DE CIENCIAS BIOLOGICAS, VOL. 30

(' IELAND) TOLTLATJCI ooTnw OP OTTEA APATOS T3 TOARUMBY-SOZEN XEIDWOD UYWZNP SOBAU9T3 =GEM

EIGWOLVN0-AADA4AUNERALL AEA03Y-OCAAA TELSEOP UPOAAAAAND SESTEG EUEAVNGEOP-EPAURI TOQSEOP UEOOSTIVR ETRAISPBEW-EOMM O6ETQUES-EURA7 AUMBETIMS-IEUNY

TEQSEOP MOTEUTS ORTVW-RNBRA URTUOYMP ECEE OPEJOLOO SAMOL PRNSUTUED UPEONA EQUT3M1SN-EATECTZ3 41 4.1 SPORECOENO00 X TREDVOTEDVD TTLGEOP UPZI110219A OONUPA EUTIVW I coos OPUE/D OPI OTOOTUAAARD

ALETOSI BUTIDS AUTAQSNOE7 X XA.A.13 X MMAIS X X X X X X AAATH BTS X X AUOLUCU X WEIDA X X X X X PUOIMORI X Table 1., Continued

Habitats Distributions by Drainages

1 Taxa g 1 dill 010 1. 0 8 111 01000 1 s m4a >4 tn CYPRINDDCNTICRE-II Cualac tessellatus-E X X X X Cyprinodon alvarezi-E X X Cyprinodon atrorus-E X X Cyprinodon beltrani-E X X Cyprinodon bifasciatus-E X X X Cyprinodon eximius X X Cyprinodon fontinalis-E X X CVprinodon labiosus-E X X Cyprinodon latifasciatus-E X X E x Cyprinodon macrolepis7E X Cyprinodon macularius X X X X X Cyprinodon mayaE X X r:f Cyprinodon meeki-E X X X

138 ANALES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOOICAS, VOL. 30

(ERFAND) 'AQUAT ANX9114 AP WIRE OPPETAS TRAPUER6V-SEZE

XAIDWPD X SEBAUGTO CAQ EPAWAWNO-OEDE4UUN MABILV-APAA TALSE03 1190.183.1 SEW( EUEDCENTIPOO.NA ,24SECOO UMOSTIE EUEINTLSW-U ABATWAS-EULA ARRRABAT4112S- TEWECO MOTAAT OAEIG-TRIBE

ETULAITTPO EC OPEAOLco A AJNSUPLAD 4,4=1 E4UTOTUNSN-RATPCT C4 X X SOORECOEZWAD X X X UUDTORPDS X X TELSROP UEZNAOSAA CONU9 EUTAVW VI AGOG APUEADOPI APOOTUIAARD AVEIOSI BUR ADS X X AUTAQSMEA X X X X NAEAD X X MEWLS XX X X X XX X 8 TEF AUUQUOLG PUVIDN X X X X X X X PURIMOU N X X

1 /O 4 43 F4 FA 11 11 TO TO FRESHWATER FISH OF MEXICO

0

M1111 -su 11

X 1

1, 0

by Drainages by Drainages nith sliAMI 0

WWT11 4 8 Distributions Distributions 111 .1

m Od11;

Habitats Continued Continued WM A 1., -E

sp. Table Table

sp. -E

hubbsi-E dugesi-E

Taxa maculata-E -II robustus-E zonistius-E tamazulae-E splendens-E

pardalis-E ateralis-E encaustus-E peraticus-E Allodontichthys Alloophorus Allotoca Allcdontichthys Allodontichthys Allodontichthys Allotoca Ameca Chapalichthys Ataeniobius toweri-E Chapalichthys Chapalichthys Charaoodon Characodon GOODEILAE

1 40 ANAI.ES DE LA ESCUELA NACIONAL DE CIENCIAS BIOLOGICAS. VOL. 30

(elciand) Muni() ooTxow alp elm

opts Ta Tetteuenfoi-sezeN xaTdWoo upuznp sebau9T3 01401D eTewewro-oadaqueugaI peAbow-atuali Telseop uearaliatio SESTEll x W X w eupAenueop-eliewAr x Telseco uroosT Ter pueTetabeweosav thpuE9-w1a' X 4.1 ri

DITIDEePUBS-TBDRL Tequeop ueoTeuTs apw-rnbeA eluaonTeD e Leg opeaoToD Jamog eTnsuT ued upeonA equToewnsn-RATecTa9 sooTeopezwoo uedeoTeded Ie4geop ueznaocueli conuyd euTiew eT 01405 apuelp ond aTooTuiaARD ageTosi buTids euTzienoeri X X X X X Neal3 XXXX mewls X X XXXX X X X X .X zaATB EPTS auequow X XXXXX XXXXXX X X tleTdC1 XX X PueTmal

9 6 4 1 n 44 4.3 z 118 44 r§ ••-I (1* I-1 U

MILLER. R. ROBERT. FRESHWATER FISH OF MEXICO 141

MCRAE() RALMAT20 OOTXPW AP KIRA

OTORELES TH TAIUMANBV-SEZAN XELDWIPP UPPEND SEBAUWO OAVRO EISUARWRO-OADAQUERTGAI, DEABW-APAPTI TALSECO MORAIZAND SPARES

EUEKERTYPOOL-UPAWAV X X TEWSECO UPOOSNER WAREPAM-APAWV X X OBETQUES-MMIA7 X XX X ORANBAPTRES-TEURL TELSEOP UEOLLEUTS

AWKAW-YRIBEA

ELTUOMAD ENG OPEACYROP AAMON UINSUTUAD UPEONX EQURATUNSN-EXPCTIP SOOTROOEZWOO UMIACTEDED

TEQAACOL UZNAPEAPA OCCALVD AUTIQW EL am

APUEAD X ATOPTURATITO

A4ATCSI BUTIDS X AUTAQSNAEU X XX X X X X EALD X X X UMALAQS X X X X X X X X X X X JAATH BTSIL X X X ATMQUOW X XX X

PUALDN X X X X X PUEIMAL X

IJ

TABLE 1. (Continued.) I = Primary; II = Secondary; III = Vicarious; X = Presence; E = Endemic; Ex = Extinct.

Habitats Distributions by Drainages k•.1 ANALES

- Taxa I DE LA ESCUELA NACIONAL 8 ' •51 E al " § II a Gambusia eurystama-E X X Gambusia hurtadoi-E X X Gambusia krumholzi-E X Gambusia longispinis-E Gambusia marshi-E X X panuco-E ? E ? Gambusia CIENCtAS Gambusia rachowi-E Gambusia regani-E X X X E ?

Ganhusia senilis X X BIOLOGICAS, GaMbusia cf. senilis-E Gambusia sexradiata X XX X X Gambusia s)eciosa X X

Gambusia vittata-E X X Gambusia yucatana XXX 8 Heterandria tri bimaculata X X XXXXX Heterandria onesi-E X X XXX Heterandriz sp.-E XXX Heterandria sp.-E X X. X X X

MILLER, R. ROBERT. FRESHWATER FISH OF MEXICO 143

( EVEND) TEQUATAO

UJTX9W AP AITEA OPEIES 12 TEAEUENHV-SEZEN XAIDWOO swam SE/BE-TWO OSAGEND ETEWA4ERD-OADAQUENTTAY PEACCRI-ATAXM

TEQSEOLD UEOLAAAAND

SESTEG EUERENYEOP-ELASWAV

TE4SEPD MANNER ETTATEP6V14-E3EW OFERQUES-EWAARL

0ITTSBET4SES-TEUNI TEASEGP UEOTESTS ORCEW-TNBEX

ETUA04T M SCER OPEACTOO ASMORI

EYNSUTUAD UPEONX X • EQUIPEWNSN-EATECTIO W X SOOTEMEZAEC0 UUDEOTEDED

1EAS2C0 UEZNIMEAEA WALLED X X W X EUTAGPI EL cows X X SPUEAÐ 0TH X X • ATOPTUAAAEO

242ICSI BUTACT X AUTAGSNOERI X X X X X X X 'PESO X X X

=SAW X X X X X X X X X X X X X AAAPD EGG X X X ASEQUOW X X PURIDN X X X X X X X X X X PUETMAL X X X X X X X X X

N-1 al TABLE 1. (Continued.) I = Primary = Secondary; III = Vicarious; X = Presence; E = Endemic; Ex = Extinct.

Habitats Distributions by Drainages I lilpi 1,

Poecillopsis gracilis X X X X X X X Poeciliopsis hnilickai-E X X X Poeciliopsis infans-E XX XXX X X X X Poeciliopsis latidens-E XX XXX X Poeciliopsis lucida-E XX XXX Poeciliopsis monachar.E X X X X X Poeciliopsis occidentals XX XXX Poeciliopsis presidionis-E XX XX Poeciliopsis prolifica-E X X X X X X X Poeciliopsis turneri-E X X Poeciliopsis turrubarensis X X X XXX Poeciliopsis viriosa-E X X X X X X X X MILLER, R. ROBERT, FRESHWATER FISH OF MEXICO 145

(ETCLEND) TE4U8TIO OARXIM 8P ETTEA APETES TH TETIEUENBV-SEZETI XAIDWOD URN%

SEBAUGTO OALENO 4.1 EIMAGENEL-OEDEWENTIEY OEACW-APAELL

TE489:0 USOLAZIAND SESREG

EUVICENWOO.EPATUMI TEQSEOP TRECOSNER EUEREPFIEW-EOEUI THE-PUN-MAW' OTT716EPUES-TEUNS, TEGSEOP UEOTEUTS 0A144,-TNBEA ET =3FTE0 ECEG OPEACQOP EINSUTUED UPEONA EQUTOEUMEN-EARECTAD SOOREOCEZLEOP UEDEAREDED TE/SEOP 1=FLA3EAELL CANTRGA EUTAVVI ET 0.440S EPUE4D ant

ATCOTUZAIRGP

ALETOEI BUTZDS AURALSNOSQ

3FAAZO X X 11129-T4S X XXX X X X XX X X X FRGIG X

AUEQUCW X X PMTDFL X X X X X X XX X X X PUEINICRI TABLE 1. (Continued.) I = Primary; II = Secondary; Ill = Vicarious; X = Presence; E = Endemic; Ex = Extinct.

Habitats Distributions by Draynages

11 , M § 11 52 1! iii00 viihrdAN Xiphophorus pygmaeus-E Xiphophorus variatus-E X X Xiphophorus xiphidium-E ATHERINIDAE-III Atherinella alvarezi-E X X X Atherinella ammophila-E' Atherinella balsana-E Atherinella crystallina-E Atherinella lisa-E Atherinella marvelae-E Atherinella sallei-E Atherinella schultzi X X X Atherinella sp. - E Miller. R. Robe,-i. FRESHWATER FISH OF MEXICO 147

(2-n3nd) TEQUATIO

00TX94 AP ATTEA X X OPVIES 13 TERIEUENBY-SEZEN XAIDTUOP TRYWZNO

SEBATIGTO NEWAWND-OADALTRENTRAL 3E/C0W-APLAIL TELSEOLO UMAAAJANO SESIES EU2IRT2M42OD-E-PAW1Y - 1v422(1D UGOOITTER EUAREPBW-POATUY X X OBETQUES-SULAQ ra ga wrara 4.1 41 14 xx 14 14 OTTT NEFFPUES-TEUNI 14 Mew° USOTEUTS OR2W-TNB2A ETUAOMEO ECES OPAZOTOP JAMOQ ETNSUTUAD UPE-ONA YQUTORANSN-ETTIECTAP

SOO1EMEZQE00 UEDEOIEDVD REQSVOO UEZNIOEXPA OONUYD EUTIVW 21 ows AFOURIP 0TH ATOOTUIAAPO AQVIOSI arpols AUTAQSNOEQ C XXX X X X X X X X X NEASO AMEAQS X X X X X 4.1 IAATH LTE X -0 EUEQUOW X X X X X X X X X X X X X X PUEIDN X X PUETMORI

4 14 .. 04

k 1 6 6 8

TABLE 1. (Continued.) I = Primary; II = Secondary; III = Vicarious; X = Presence; E = Endemic; Ex = Extinct.

Habitats Distributions by Drainages

• A 1 A g 1 Taxa 8 STIPINV M 6 M 4 113 i° Ml8O 3CI

.31 4tu ,$) .3 03 . m HM111111 110111 4,1 ,n Chirostoma rl patzcuaro-E X E' Chirostama promelas-E E g Chirostama riojai-E E Chirostama r sphyraena-E g Poblana alchichica-E E 0 Poblana ferdebueni-E Poblana letholepis-E SYNBRANCHIDAE-II Cphisternon aeniqmaticum Ophisternon infernale-E X Synbranchus marmoratus X CENTRARCH1DAE-I Lepamis cyanellus X Lepomis qulosus X Table 1., COntinued

Habitats Distributions by Drainages

Tama

11 11111 mq111 11 Miller, 8 0§ !14.111 d„ i I N A MP, EillhAll 814mgg R. Robert, Lepomis macrochirus X X Lepamis megalotis X X X X MicropterUs salmoides X X X X X FRESHWATER FISH OF MEXICO

PERCIDAE-I Etheostama australe-E Etheostoma graham'. Etheostoma pottsi-E Etheosfama sp. -E X Percina macrolepida GERREIDAE-III Diapterus mexicanus-E SCIAENIDAE-III Aplcdinotus grunniens

Cichlasoma bartoni-E kto

Extinct. TABLE 1. (Continued.) I = Primary; II = Secondary; III Vicarious; X = Presence; E = Endemic; E. =

Habitats Distributions by Drainages

1 4 'b' Twat 101 hi ll i IV OiM14 lidai ; ›.i4m llidi ›t81N Cichlascma beani-E xx xx xx xx Cichlasama bifasciatun X xx Cichlasama bulleri-E Cichlasoma callolepis-E

Cichlasana X cyanoguttatum Cichlascma xx Cichlasana fenestratum-E xx Cichlascma friedrichsthali X Cichlascma qrsarcdes-E Cichlasana quttulatum Cichlasana hartwegi-E Cichlasoma helleri xx Cidnlasana heterspilum ? X X MILLER, R. ROBERT, FRESHWATER FISH OF MEXICO 151

(EICIAND) TEQUATIO OPTX9141 AP PUPA

OPPTPS T3 TENEUENHOI-SEPPN XAIDMOO UPAND SE6AU9T0 0114END WPMALUNP-OADAQUEMPL X

DEABOKLI-APIPA REQSEOP UMAAXIAND SESIVE ETWERUPOO-EPAWAY

TELSOCIO UPOOSTTPC EUATELOBEW,EPAOL OLOWQMS-ALUAL ALUNBENMS-TEUNY TEWEAGO MATEUTS OARK-TNBE4

ETULCGTTPD PCPS OLOMIOTOP AAMAL ETNSUTUAD UPGEONA

E4UTOEURNSN-ENTECT2D X 43 SOAPOORZWAD IREDELOTEDTD

TE4SECO UEZRUMAAA OMUYA EUTAUM ET 040S APURAP am ATOPTUAAAPO AWICSI BUTADS AURA4SNMA XAAAP UMAALS X X X X X X X X X X X X X OATH BM X X X X AUEQUQK X X X PUEIDN X X X X X X X X X X PUEIMAJ X X X X X X X X X X TABLE 1. (Continued.) I = Primary; II = Secondary; III = Vicarious; X = Presence; E = Endemic; EX = Extinct.

Habitats Distributions by Drainages

iiv Tawa '# , saavxv ga 0 Ik§d u 8 § 8 I 1 1: I i In Baja California Cichlasoma rabertponi X X X X Cichlasma salvini X X Cichlasoma socolofi-E X X Cichlasoma spilurum X X X Cichlasama steindachneri-E X X Cichlasama synspilum XX XXX Cichlasama teapae-E X X Cichlasoma trimaculatum X X X Cidhlasoma urophthalmus X X X X Cichlasoma sp. X X Cichlasoma sp. X X X Cichlasama usumacinta XXXX Petenia splendida XX X X X MUGILIDAE-III Auonostcmus monticola X X X X X X X X XX X X X X X X .X X X X X Joturus pichardoi X X X X ? X Table 1., COntinued

Habitats Distributions by Drainages

gl Taxa , A --°4 R M 2 giil 8 g g 1:1 5 PI p_g. III 1.4 4 g 0,2 11M1.1 11 do 1,6

HA 1 4 Baja California ELEIDTRILAE-III Gobiomorus dormitor XX XX X X XX Gobicmorus maculatus X X XXX X XX Gobicmorus polylepis-E Leptophilypnus sp. GOBIIDAE-III Gobionellus atripinnis Sicydium 21eMiera X X Sicydium multipunctatum

1. This species is endemic to a spring in Rio Atoyac, east of C6rdoba,Veracruz. 2. This species is endemic to the Laguna Sontecomapan drainage Veracryz,(Chernoff and Miller,1984).