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Investigations of the Ichthyofauna of Nicaraguan Lakes Papers in the Biological Sciences

1976

Geographical Distribution of Central American Freshwater Fishes

Robert Rush Miller University of Michigan - Ann Arbor

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Miller, Robert Rush, "Geographical Distribution of Central American Freshwater Fishes" (1976). Investigations of the Ichthyofauna of Nicaraguan Lakes. 10. https://digitalcommons.unl.edu/ichthynicar/10

This Article is brought to you for free and open access by the Papers in the Biological Sciences at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Investigations of the Ichthyofauna of Nicaraguan Lakes by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. Published in INVESTIGATIONS OF THE ICHTHYOFAUNA OF NICARAGUAN LAKES, ed. Thomas B. Thorson (University of Nebraska-Lincoln, 1976). Copyright © 1976 School of Life Sciences, University of Nebraska-Lincoln. Made in United States of America Reprinted from COPEIA, 1966, No.4, December 23 pp.77-3-802

Geographical Distribution of Central American Freshwater Fishes

ROBERT RUSH MILLER

The distribution of freshwater fishes in Central America is reviewed. Major parts of the region, especially in Honduras and Nicaragua, have yet to be explored ichthyologically, and systematic revision of important groups, notably the cichlids, is long overdue. The continental area from the Isthmus of Tehuantepec to the Colombian border contains approxi­ mately 456 species, of which over 75% comprise the Cyprinodontidae, Poeciliidae, Cichlidae, and marine invaders (peripheral fishes); about one­ third of the latter (57 species) have taken up more or less permanent residence in fresh water. There are 104 primary species in 10 families, 165 secondary forms in 6 families, and 187 peripheral species distributed among 30 families (nearly half of the latter are ariids, atherinids, gerrids, and gobies). Poeciliids and cichlids are particularly rich and diverse, to­ gether comprising 139 species. Characins are numerous only in the Pana­ manian region, into which they and five South American catfish families have recently penetrated. Except for gars, no North American family has reached beyond northern Guatemala. There are no non-ostariophysan primary fishes in the area. Although our knowledge of the fish fauna approaches adequacy only in the Guatemalan and Panamanian regions, an attempt is made to de­ limit fish provinces. The best known, the U sumacinta assemblage of south­ eastern Mexico and northern Guatemala, has been a strong center of evolution. The marked differences between the fishes of this area and those of the Isthmian province, in Costa Rica and Panama, emphasize that Middle American fishes do not comprise a coherent faunal unit. Other provinces are the impoverished Chiapas-Nicaraguan and the San Juan, the latter notable chiefly for the cichlids of the Great Lakes of Nicaragua. The highlands of southeastern Mexico, Guatemala, and Honduras comprise an ancient land mass with a meager fish fauna in which the primitive cyprinodontid genus Profundulus has evolved. The Middle American fauna, from which most of Panama should be excluded, is characterized by: (1) the adaptive radiation of the proba­ bly autochthonous Poeciliidae; (2) great diversification of the genus Cichlasoma, including the evolution of the derived genera Petenia, Neetroplus, and Hero tilap ia; (3) the likely origin and speciation of Bramocharax; (4) the occurrence of numerous freshwater species of marine origin, a development strongly influenced by (5) the paucity of primary freshwater fishes.

125 774 COPEIA, 1966, NO.4

INTRODUCTION nus of Central America represent a transition between the South and Middle American O~E than half a century has elapsed M smce the last attempt was made by an faunas, all the Panamanian region is arbi­ ichthyologist (Regan, 1906-1908) to summa­ trarily included since the area of faunal rize knowledge of the distribution of fresh­ break has not been clearly delimited. The water fishes in Central America. Regan in­ northern boundary, however, has been de­ cluded all of Mexico in his treatment but to termined by the nature of the fish fauna to do so in this paper would unduly complicate include the Rio Papaloapam on the Atlantic coast and the Rio Tehuantepec on the Pacific and h~ndicap an understanding of the origin and dIspersal of what may be appropriately versant-essentially the Isthmus of Tehuante­ called a Middle American fauna (Miller, pec geological province (Maldonado-Koerdell, 1959:195; Hubbs and Miller, 1960:101). 1964:Fig. 2). This fauna, in marked contrast to that The Rio Coatzacoalcos-the large river just of North America (including the Mexican south of the Papaloapam that drains the Plateau and the coastal lowlands south to northern slope of the Isthmus of Tehuante­ about the Tropic of Cancer), is dominated pec-harbors a fish fauna characteristic of the by the salt-tolerant cyprinodontoids (particu­ ~owland tropical region extending eastward larly the poeciliids) and cichlids, and by a mto the Usumacinta basin of Guatemala. diversity of invaders from the sea. These The Papaloapam contains essentially the same three groups comprise over 75% of the spe­ fauna but with fewer species-including only cies known to occur in fresh water between one ?f northern origin (Ictiobus bubalus) the Isthmus of Tehuantepec and Colombia. that IS not also found in the Coatzacoalcos. Within that area there are no minnows, only In striking contrast is the Rio Panuco, the two suckers, only one North American cat­ major drainage next to the north of the fish, and no perches, darters, or sunfishes­ Papaloapam, which harbors a fish fauna which together comprise the bulk of the strongly transitional between tropical and North American fauna. The abundant ma­ temperate types (Darnell, 1962: 351-359). rine element, consisting of almost 40% of The Rio Tehuantepec likewise marks the the species, include some 57 species that are northern distributional limit for many of more or less permanent residents of fresh the Pacific tropical forms, including the four­ water-a noteworthy feature of the Middle eyed fishes of the family Anablepidae, the American fish fauna. pimelodid catfishes, the Synbranchidae, and Chiefly in the southern part of Middle most of the cichlids. America, in Costa Rica and Panama, there is DISTRIBUTIONAL AND SYSTEMATIC KNOWLEDGE also an abundance of characins and Neo­ tropical catfish families representing, in Alt~ough both our distributional and sys­ greatly diluted fonn, a vanguard from the tematIC knowledge of the fish fauna of Cen­ south of the great Amazonian fauna. One tral America have increased greatly since characin genus (Astyanax) has succeeded in Regan's work was published, there are still crossing the whole length of the Middle large gaps to be filled. Little is known of the American bridge to enter Texas, as has the life history and ecology of any of the species South American cichlid genus Cichlasoma. -a vacuum that must be filled before we can The South American catfish genus Rhamdia, fully understand the fauna. family Pimelodidae, has moved up the bridge Only Mexico and Guatemala have been to southern Mexico (across the Isthmus of comparatively well worked, and even in Tehuantepec), and a few other ostario­ those areas recent collecting has uncovered major ichthyological finds. Most of western physans have progressed part way across Cen­ tral America. No North American primary Honduras and a large part of northern Nica­ freshwater family has succeeded in moving ragua have yet to be examined by ichthyolo­ southward beyond Guatemala, but the salt­ gists, and even in such small countries as tolerant Nearctic gars have managed to reach British Honduras and El Salvador explora­ the Great Lakes basin of Nicaragua-Costa tory work is needed. Lately a number of Rica. ichthyologists have worked in Costa Rica and in the adjoining Great Lakes basin of Nica­ AREA COVERED ragua, and William A. Bussing is completing Although the fishes of the southern termi- a doctoral program on Costa Rican fishes.

126 l\IILLER-CENTRAL AMERICAN FISHES 775

With the very recent establishment in Costa (commonly in the early life-history stages) Rica of the Organization for Tropical and need not concern us here. Studies, we may anticipate that our knowl­ The Middle American fish fauna is marked edge of Costa Rican fishes will be rapidly by great richness and diversity of the Poe­ advanced. Between 1961 and 1965, Horace ciliidae and the Cichlidae, which together G. Loftin collected in those areas of Panama comprise 84% (139 species) of the secondary -chiefly east and west of the Canal Zone­ fishes and more than half of the combined that had previously been not at all or very primary and secondary groups. The Cyprino­ little explored for freshwater fishes. I have dontidae and the pimelodid catfishes of the examined just enough of his material to genus Rhamdia are also numerous in Middle recognize that current concepts of distribu­ America, but the characins, while comprising tion and classification of certain fishes oc­ an impressive total of 20 genera and 51 spe­ curring near the southern terminus of Cen­ cies, number fewer than 10 species north of tral America will need to be considerably Costa Rica. In general there is a dearth of revised. primary fishes north of the Nicaraguan­ The area treated contains about 456 spe­ Costa Rican boundary, the reasons for which cies of fishes known from fresh and brackish are explored by Dr. Myers (1966). water. This includes 104 freshwater or pri­ The Gymnotidae, derived from a characin mary species, 165 secondary species that may stock, are represented by 5 genera and species tolerate varying degrees of salinity, and some south of Nicaragua, but only 2 species of a 187 peripheral species-including such kinds single genus, Gymnotus, penetrate as far as sawfishes, tarpon, anchovies, ariid catfishes, north as Guatemala. atherinids, gerrids, gobies, and flatfishes. Five South American catfish families-the These 187 species are represented by 73 Auchenipteridae, Ageneiosidae, Trichomy­ genera in 30 families, whereas the 165 secon­ cteridae, Callichthyidae, and Loricariidae­ dary fishes include 34 genera in only 6 fami­ have managed to invade the southern part lies and the 104 primary species are assigned of the isthmus as far as Panama or Costa to 43 genera in 10 families (or to 16, if the Rica (Fig. 1); but only the loricariids-the characins are split). It should be stressed largest and probably most specialized of that within some families classified as pri­ existing catfish families-have numerous rep­ mary freshwater fishes (i.e., those regarded as resentatives there (15 species in 8 genera). strictly confined to fresh water) are species The four-eyed fishes of the family Ana­ that may be more salt tolerant than are some blepidae show a curious discontinuous dis­ fishes assigned to the secondary freshwater tribution, with one species (Anableps dowi) groups (see, e.g., Darnell, 1962:354; Schwartz, confined to the Pacific slope from the Rio 1964; Marshall, 1965:333). As Middle Ameri­ Tehuantepec basin of Mexico to the Rio can examples may be cited the ubiquitous Choluteca, Honduras, whereas the other spe­ Astyanax fasciatus, a primary fish that is cies occur along the Atlantic slope of north­ known to tolerate brackish water, [ctalums ern South America (Venezuela to the Ama­ zon). furcatus, a frequent invader of water of low Our knowledge of the systematics of some salinity in Mexico, and Rhamdia wagneri, of the important groups of Middle American the only member of the genus common to freshwater fishes badly needs overhauling. South and Middle America and believed This is especially true of the cichlids, the last capable of dispersing coastwise under favora­ revision of which appeared over 60 years ago ble salinity conditions (Loftin, pers. comm.). (Regan, 1905). I am aware of at least 12 The major peripheral groups comprise nominal species in Mexico and Guatemala only 5 families (Ariidae, Atherinidae, Ger­ alone that belong in synonymy, and of 10 ridae, Eleotridae, and Gobiidae) that con­ in the same area that represent undescribed tribute 85 species or nearly half of the total. species. It may be questioned seriously Although some 31 of the 187 peripheral whether the widespread genus Rhamdia com­ fishes have taken up permanent residence in prises 20 species, and certain characins need fresh water, and about 26 additional species re-evaluation before relationships and dis­ spend much of their lives inland from the tributions are well understood. Not infre­ sea, most members of this group occur only quently, specific names are based solely on sporadically or temporarily in fresh water unique specimens which bear such revealing

127 776 COPEIA, 1966, NO. 4

·40

~32'

28° I 20"1 • IC~TOSTOMIDAE,. ,

I 16°

DISTRIBUTION OF SOME FAMILIES OF FRESHWATER 8° FISHES IN MIDDLE AMERICA

------Southern limits of northern families --- Northern limits of southern families AUCHENIPTERIDAE (P) LORICARIIDAE (CR) 4" ~ ___ ~ ______~ TRICHOMYCTERIDAE (CR) 106° - 98°

Fig. 1. Distributional limits of certain primary and secondary freshwater fishes in Middle Amer­ ica. A single characin (Astyanax fasciatus) and cichlid (Cichlasoma cyanoguttatum), representing the tropical families Characidae and Cichlidae, respectively, reach the U. S. The northern limit of the Neotropical Anablepidae (not shown) is the same as that for the Pimelodidae on the Pacific slope. Of 5 South American catfish families 3 range northward only to Panama (P), the other 2 to Costa Rica (CR). The Ictaluridae (like the Cyprinidae) also occur on the SW coast of Mexico in the Rio Balsas basin. The Nearctic sunfishes (Centrarchidae) and darters (Percidae) range southward into Mexico only to the north of the Tropic of Cancer (just south of 24°); this is as far as non­ ostariophysan families penetrate Middle America.

type localities as "Mexico" or simply "Cen­ fishes occurs in the characins, of which there tral America." Proper assignment of names are only 6 species in 1 phyletic line in therefore often requires examination of type Guatemala but 36 species in 6 phyletic lines specimens and these are scattered among the in Panama. The two countries share only world's museums. Fortunately the poeciliids a single species of this group-the ubiquitous have recently received a comprehensive re­ Astyanax fasciatus (which can disperse view (Rosen and Bailey, 1963) that clears the through brackish water)-although they do way for intensive studies of this prolific and have four genera in common (Astyanax, diversified group. (For a different interpreta­ Brycon, Hyphessobrycon, and Roeboides). tion of the limits and divisions of Gambusia, The other notable difference is the occur­ see Rivas, 1963.) rence of 5 South American catfish families Ichthyologically the two best-known coun­ in Panama, comprising 20 species, none of tries in Central America are Guatemala and which inhabits Guatemala. Gymnotids are Panama and it is instructive to compare the also more diverse in Panama, which has .5 composition of their primary and secondary genera, only 1 of which reaches Guatemala. fishes. Each country contains a comparable Among the secondary fishes, the situation total number of species and families, 90 spe­ is reversed: Guatemala has 70 species in 18 cies in 11 families in Guatemala and 100 spe­ genera and 6 families, whereas Panama has cies in 12 families in Panama. The major only 34 species in 15 genera assigned to 4 difference in composition of the primary families. Here the major difference in COlU-

128 1\IILLER-CENTRAL AMERICAN FISHES 777 position is in the cichlids, of which there are and some species and genera are autoch­ 40 species in Guatemala but only lOin thonous. There are very restricted species Panama. (These 10 are represented by 3 that live in association with widespread ones. genera and 7 species that do not occur in Among the latter are those which, by virtue Guatemala.) Although the number of poe­ of particularly broad ecological tolerances, ciliids in these countries is similar-21 species extend beyond the confines of a single prov­ in Guatemala and 17 in Panama-only 3 are ince and are members of two or more such shared (Gambusia nicaraguensis, Phallichthys assemblages. Of the regions tentatively recog­ amates, and the ubiquitous Poecilia mexi­ nized below, only the Usumacinta Province cana) and only 3 additional genera (Alfaro, seems firmly established. The number and Brachyrhaphis, and Poeciliopsis) have repre­ precise delimitation of other provinces will sentatives in each area. Both Guatemala and require a greater knowledge of the fish fauna Panama have 6 species of cyprinodontids. than is now available. with no species and only the genus Rivulus Usumacinta Province.-The broad hot low­ in common. lands of Tabasco in southeastern Mexico re­ The major difference in cichlid develop­ ceive the waters of the two largest river sys­ ment has both historical and ecological bases. tems of Middle America: the Grijalva and the former discussed by Myers (1966). Cich­ the Usumacinta. Although widely separated lids occupy much the same ecological posi­ in their upper courses, once they break out tion in the tropics as do the temperate North of the Chiapas-Guatemalan highlands they American sunfishes. They are carnivorous. form and are joined by large, slowly-moving herbivorous, or omnivorous, inhabit springs. waters, of rather uniform flow. In their small creeks, and ponds to large rivers and lower courses these rivers comingle in a maze lakes, from coastal lowlands (including man­ of bifurcating channels and distributaries. grove-bordered lagoons) to mountain high­ oxbows, marshes, and swamps-providing a lands, where they are scarce; many of the large area of productive, permanent fish species are important food fishes. As a group habitats. The Usumacinta system is the they have been able to adapt to a large rna jor source of the best-marked fish province variety of habitats. Probably the two most in Middle America. It includes the area important factors limiting their northward from the Rio Papaloapam of southeastern dispersal and altitudinal distribution have Mexico eastward and southward to the Rio been temperature and competition. Their Polochic of eastern Guatemala (Fig. 2). It is maximum flowering in Middle America oc­ separable into two major divisions, the curs in the tropical lowland area of south­ Usumacinta-Grijalva and the Papaloapam­ eastern Mexico and northern Guatemala-a Coatzacoalcos, and two lesser ones, the Yuca­ region containing the most voluminous rivers tan Peninsula and the Polochic-Lake Izabal. of Middle America. Excluding the Pacific drainage, this province comprises approximately the northern half FISH PROVINCES of Regan's (1906-08, map 2) provisional Guatemalan Province. Most primary freshwater fishes usually can The Usumacinta Province contains well disperse overland only by physiographic over 200 species in all categories, with 108 changes of the land, lowering of sea level. or species, 29 genera, and 9 families of primary anastomosing of river systems during the and secondary fishes. Two secondary groups. high water of the rainy season. Conse­ the cyprinodontoids and the cichlids, com­ quently, their distributional patterns as a prise approximately 90 species. including a rule rather closely match those of the rna jor number of endemic and evidently autoch­ hydrographic systems. The presence of a thonous genera and a host of endemic spe­ large and diverse drainage basin-such as the cies. The province is also distinguished by Mississippi in North America or the Amazon the large number of marine derivatives (18 in South America-provides the focus for the species in II genera and 9 families) that have evolution of a major fish fauna. taken up permanent residence in fresh water. As here conceived, a fish province is a Among these are the clupeid Dorosoma anale, region that is characterized by groups of asso­ the ariid catfish genus Potamarius, a half­ ciated species that have similar geographic beak (Hyporhamphus mexicanus), the atheri­ and ecologic ranges. Endemism is marked, nid genera Archomenidia and Xellatherina.

129 778 COPEIA, 1966, NO.4

94 90 86

~ I

16

12

------~.------..-- 12

0 200 i 400 600 800 I I I I ! KILOMETERS 8 8

98 94 90 86 82 78

Fig. 2. Tentative arrangement of major fish provinces of Middle America, showing numbers of families, genera, and species of primary and secondary fishes in each area. Insufficient collecting between the Rio Motagua (Guatemala) and the Rio San Juan (Nicaragua) prevents the delimitation of a province (or provinces) in that region.

the gerrid Diapterus mexican us, eleotrids of south; 2 represent a widespread swordtail the genus Leptophilypnus, the toad fish Batra­ and platyfish belonging to a genus (XiPho­ choids goldmani, and the blind brotulid phorus, Fig. 4) that likely originated to the Typhliasina pearsei. There is also the blind north of the province in the Rio Panuco Furmastix infernal is, of the secondary fresh­ area; 2 are gambusias (G. yucatana, G. nica­ water family Synbranchidae. These latter raguensis), ranging, respectively, north and two cave-inhabiting species are restricted to south of the province; I is the widely distrib­ the Yucatan Peninsula. uted, salt-tolerant Belonesox belizanus (from Among the poeciliids are no fewer than 4 near Veracruz south to Costa Rica, Fig. 5); genera belonging to 2 subfamilies and 3 another the equally ubiquitous Heterandria tribes (Priapella, Carlhubbsia, Xenodexia and (Pseudoxiphophorus) bimaculata (from Vera­ a recently discovered cnesterodontine) that cruz, Mexico to Nicaragua); and 2 are spe­ are confined to and evidently evolved within cies of Poeciliopsis that are more widespread the area. Priapella, with a species each in the on the Pacific coast of Middle America. Papaloapam, Coatzacoalcos, and Usumacinta The cichlids are the most speciose group basins, is the most widespread of these autoch­ in the Usumacinta Province but are not as thonous genera (Fig. 3). Restricted to the diversified as the poeciliids. There are about province are 23 species in 9 genera that rep­ 44 species in 2 genera, constituting more resent all the known tribes of the subfamily than half (53%) of the cichlid fauna of Cen­ Poeciliinae. Included is the distinctive Het­ tral America and nearly 60% of the species erophallus group of the tribe Gambusiini, assignable to the genus Cichlasoma. One containing a species each in the Coatzacoal­ monotypic genus, Petenia, is confined to the cos, Grijalva, and Usumacinta basins. Of the province, and the distinctive Thorichthys 11 remaining species of poeciliids, 3 are mol­ division of Cichlasoma (Miller and Nelson, lies widely distributed both to the north and 1961; Rivas, 1962) has all but 1 of its 8 or 9

130 MILLER-CENTRAL AMERICAN FISHES 779

Fig. 3. Distribution of the poeciliid genera Priapella (3 species) and Alfaro (2 species), modified after Rosen and Bailey (1963). Priapella is an example of an autochthonous endemic in the Usuma­ cinta Province. Except for a crossover in the Tilaran region of Costa Rica, Alfaro is restricted to the Caribbean slope; its known southern limit is near the faunal break between the Middle American and South American faunas. species restricted to the area. Thirty-five of fenestra tum is distributed northward to the the 44 species are not known to range out­ Rio Antigua, just north of Veracruz City; C. side of the province. guttulatum is essentially a Pacific slope spe­ Of the remaining 9 species of cichlids 2 cies, occurring in the province in the Rio (Cichlasoma maculicauda and C. spilurum) Coatzacoalcos basin; and C. aureum (of the occur as far south as Panama and it is note­ Thorichthys group) ranges southward only to worthy that one of these (maculicauda) is the Rio Motagua of southeastern Guatemala usually found near the coast and frequently and adjacent Honduras. in brackish water; 2 (C. friedrichsthali and C. Characins are weakly represented, with urophthalmus) range southward to Costa only 5 species in 4 genera, but I genus Rica and Nicaragua, and urophthalmus oc­ (Bramocharax) is shared with Nicaragua and curs in brackish water and on islands ad­ Costa Rica. jacent to the coast (e.g., Isla Mujeres, Quin­ There are but 3 northern elements repre­ tana Roo); 2 others (octofasciatum and rob­ sented by 2 species of Ictiobus, I of Ictalurus, ertsoni) occur southward into northwestern and a single gar, Lepisosteus tropicus, which Honduras and octofasciatum ranges north to also occurs along the Pacific slope of Central the vicinity of Veracruz City; Cichlasoma America and in the Great Lakes-San J nan

131 780 COPEIA, 1966, NO.4

86°

Xiphophorus

Fig. 4. Distribution of the platyfishes and swordtails, genus Xiphophorus, after Rosen and Bailey (1963). Of the 8 or 9 recognized species, 1 each is confined to the Rio Papaloapam and Rio Coatza­ coalcos basins, 2 (X. helleri and X. maculatus) range westward and southward from the Rio Coatzacoal­ cos, and the remainder occur north of the area treated herein.

basin of Nicaragua and northern Costa Rica. two river systems there are also endemics­ The pimelodid catfishes have 7 species, all e.g., among the poeciliids of the genera in the genus Rhamdia; 5 occur only within Priapella, Xiphophorus, and Poeciliopsis, in the province. the cichlids, and in the atherinid genera There is moderate to strong endemism Archomenidia and Xenatherina. within the subdivisions of the Usumacinta On the Yucatan Peninsula, endemism in­ Province. This is most marked in the Usuma­ volves coastal forms in the families Cyprino­ cinta-Grijalva division, which contains the dontidae, Atherinidae, and Brotulidae (5 spe­ largest and most diversified catchment basin. cies with I endemic genus, Typhliasina) as It has 38 endemics out of a total of 67 species much as it does species whose nearest rela­ of primary and secondary freshwater fishes tives inhabit the adjacent lowlands of south­ -or 57%. Five of the 10 strictly freshwater eastern Mexico (4 species, with 1 endemic marine derivatives are confined to this di­ genus, Furmastix, in the Cyprinodontidae, vision. Poeciliidae, Cichlidae, and Synbranchidae). In the Papaloapam-Coatzacoalcos division The fauna is an impoverished one, largely are 45 species of primary and secondary because of the scarcity of surface waters, par­ fishes, of which 15 (33 %) are endemic; again, ticularly rivers. 5 of 10 freshwater peripheral species are re­ The Polochic-Izabal division has a total of stricted to these basins. Within each of the 28 species of freshwater fishes with 6 en-

132 MILLER-CENTRAL AMERICAN FISHES 781

6°

I'

I 1120r--r------~ ____ 12° II

I

I:

II 98° 90°

Fig. 5. Complementary distribution of the closely related poeciliid genera Belonesox and Brachy­ rhaphis, modified after Rosen and Bailey (1963). The large, predatory Belonesox is a monotypic genus not uncommon in brackish lagoons and may have originated in the Usumacinta Province. Brachyrhaphis is autochthonous in the Isthmian Province, where it is represented by at least 7 spe­ cies; the disjunct B. hartwegi, described from Chiapas, Mexico, has recently been collected by Rosen in Guatemala and may range as far to the southwest as the Gulf of Fonseca, Honduras. demics belonging to the Poeciliidae and Cich­ Not surprisingly, it is marked by an im­ lidae. Four of the 5 peripheral freshwater poverished fauna, with only 8 families, 14 forms are confined to this drainage, includ­ genera, and 30 species of primary and secon­ ing a representative each of the Ariidae, dary fishes (see Hildebrand, 1925; Carr and Atherinidae, Syngnathidae, and Eleotridae. Giovannoli, 1950). The genera Potamarius and Carlhubbsia are Encompassed within the area is the known shared with the Usumacinta-Grijalva drain­ Central American range of Anableps dowi; age. also included is the Pacific distribution of Chiapas-Nicaraguan Province.-Precise de­ the tropical gar, which very nearly matches lineation of other fish provinces is not possi­ that of the cuatro-ojos. ble at this time but some tentative arrange­ There are only 2 characins, Astyanax fas­ ments may be outlined. Extending along the ciatus and Roeboides salvadoris; the latter Pacific slope from the basin of the Rio Te­ represents a genus absent from the Usuma­ huantepec approximately to the Nicoya Pen­ cinta Province. There is a single gym no tid, insula of extreme western Costa Rica (Fig. 2) ranging from Guatemala to the Gulf of Fon­ is an area that is essentially equivalent to the seca. Rhamdia is represented by 4 species, Pacific Mexico-Nicaragua Biotic Province of of which 3 appear to be restricted to the Stuart (1964:356, Fig. 5), except that the area. There are only 2 cyprinodontids, the northern part extends into his Tehuantepec essentially highland genus Profundulus and Province. The fishes of this region are here the peculiar Oxyzygonectes dowi, a salt­ referred to the Chiapas-Nicaraguan Province. tolerant species barely entering the province The area receives much less rainfall than the from the south. Poeciliids also are few, 8 Caribbean lowlands, has a narrower coastal species in 5 genera, of which 3 are confined plain, and its streams are few and of rela­ to the province. The absence of Gambusia, tively small discharge; many are intermittent. so well represented in the Atlantic-Caribbean

133 782 COPEIA, 1966, NO.4

lowlands of Mexico, is noteworthy. Among tico (Bussing, 1963) is presently known only the cichlids are 10 species of Cichlasoma, of from the San Juan drainage in Costa Rica: which 7 are so far only known from this 1 species is possibly restricted to the Grea t area. There are no distinctive freshwater Lakes basin (a Poeciliopsis). derivatives of marine groups. The clupeid genus Dorosoma reaches its The Caribbean fish fauna of Honduras southern limit in the Nicaraguan lakes. and Nicaragua is still too poorly known to where it is represented by the endemic yield adequate interpretations of distribu­ species D. chavesi. A shark, two sawfishes tions in the region between the Rio 1Iotagua, (P. pectinatus rare), and the tarpon also oc­ southeastern Guatemala, and the Rio San cur in Lake Nicaragua. One atherinid (M e­ Juan, Nicaragua. Regan (1906-08) simply laniris sardina) is restricted to the Great placed the fishes of this area (as he did of Lakes. As previously mentioned, this prov­ my Chiapas-Nicaraguan Province) in his ince marks the southern limit of the gars. Guatemala Province. Isthmian Province.-The well-watered San Juan Province.-Regan (1906-08) ten­ tropical lowlands lying south and east of tatively recognized lakes Managua and Nica­ Nicaragua include an area with a distinctive ragua and their outlet, the Rio San Juan. biota that has been called the Costa Rica­ as comprising a region characterized by the Panama Biotic Province (Stuart, 1964:35-1. peculiar cichlid fauna found in the Nica­ Fig. 5). This is essentially equivalent to the raguan lakes (Meek, 1907). Regan called this Isthmus of Costa Rica-Panama Geological the San Juan Province. That it spills south­ Province (Maldonado-Koerdell, 1964:Fig. 2). ward into Costa Rica is indicated by re­ Its fish fauna is here designated as the Isth­ cent collecting near Tortuguero (Caldwell, mian Province, following the terminology of et aI., 1959). The cichlids, though perhaps not quite so Regan (1906-08). When more data become speciose as Regan and Meek believed. are available the boundaries of the province will nevertheless numerous and diversified. Thev need revision since it now includes all of comprise about 16 species in 3 genera, 1 of Panama, of which the central and eastern which-the monotypic Herotilapia-evidently (especially) parts contain fishes with pre­ evolved in the area. The genus Neetroplus. dominantly South American relationships. which may also be autochthonous. is repre­ The Isthmian Province comprises both a sented by one species nearly restricted to the rich and diversified fish fauna. of which the region and by another apparently limited to primary and secondary groups are repre­ Panama. Nine of the 14 species of Cichlasoma sented by 136 species, 55 genera. and 12 fami­ appear to be confined to the province; some lies (or 17 if one splits the Characidae). This are highly specialized, with thick, fleshy lips is more than twice the number of genera and and blunt teeth; others exhibit rubrism 30% more species than are found in the (Meek, 1907). (Possibly the excessive lip de­ Usumacinta Province. Unlike the latter. velopment is sex-limited in one species.) however. the diversity and abundance of spe­ Also characteristic of this province is the cies in the Isthmian area are largely due to genus Bramocharax. with 2 species ranging penetration of South American fishes. espe­ into Costa Rica: but we now know of an cially among the catfishes. characins. gym­ additional species of this genus in the Usumacinta basin of Guatemala. and others notids and, to a lesser extent. the cichlids. mav be discovered in the poorly known inter­ These 4 groups comprise 109 species. 44 veniTl~ region. genera. and 10 families-of which 53 species. Other characins in the Nicaraguan area in­ 33 genera. and 5 families (all catfishes) reach clude 7 species in 6 genera. of which Brycon­ their northern limit in the Panamanian sec­ americus and Rhoadsia have species that tion of the province. The characins. gym­ eithpr range south of NicaraQ'ua or their re1a­ notids. and catfishes together make up 60% tives are southern onlv. Possiblv 4 species of of the total fauna, and more than 60% of the characins are confined to the province. species in these 3 groups are either restricted Poeciliids are not numerous, 9 species in to Panama (east of Bocas del Toro and 6 genera, none of which is distinctive of the Chiriqui provinces) or range southward and region. Most of the species are wide-ranging. eastward into South America. although the recently described Phallichthys Unlike the characins, gymnotids, and cat-

134 MILLER-CENTRAL AMERICA); FISHES 783 fishes (except Rhamdia), the poeciliids show (versus 18 species in 11 genera and 9 fami­ that the area has been a center of note­ lies). (In each case, Agonostomus, Joturus, worthy diversification at various taxonomic and Sicydium were not included.) Most of levels. There are 9 genera and 25 species, of these species, including the more distinctive which 2 genera, 1 subgenus, and 18 species ones, are confined to Pacific streams: a pipe­ are restricted to the province. fish (Syngnathus elcapitanensis) in both coun­ The genus Brachyrhaphis (Fig. 5), closely tries; the goby Guavina micropus in central allied to Gambusia (of which there is only 1 Panama; 2 eleotrids (Hemieleotris latifasci­ species in the Isthmian region), evidently atus and Leptophilypnus panamensis); and evolved in the area, for all but 1 of its ap­ Gobiesox potamius in Costa Rica. On the proximately 9 species are known only from Caribbean slope are the atherinid Melaniris Costa Rica and Panama. Some, like B. casca­ chagresi (both countries) and Gobiesox jalensis, are generally distributed, but most . nudus, which also occurs in the West Indies are restricted to either Costa Rica or Panama, and southward to Venezuela. and often to either the Pacific or Caribbean slopes. The genus is evidently more speciose The region where most typically South than currently recognized. American fishes stop and southern Central N eoheterandria also appears to have origi­ American ones appear has been investigated nated in the Isthmian region, to which 3 of by Horace Loftin. He informed me that the its 4 species are confined (the fourth inhabits transition is approximated by the eastern the Caribbean drainage of Colombia). Two boundaries in western Panama of the prov­ of the species occur only in Panama and 1 inces of Bocas del Toro (near Valiente Pen­ (N. umbratilis) is found on the Caribbean insula) on the Caribbean and Chiriqui (Rio slope of Costa Rica and adjacent Nicaragua Tabasara) on the Pacific. The southern limit (San Juan basin). of the genus Alfaro is very close to the pro­ The subgenus Aulophallus of the genus posed faunal break on the Caribbean slope: Poeciliopsis is represented by 3 species on that boundary also forms the western margin of the Colon-Darien Biotic Province of R van the Pacific slope (P. elongata J P. retropinna J and P. sp.), all of rather restricted distribution. (1963:32), as based on mammal distribution. The genus Priapichthys, distributed from The ranges of approximately 12 of the 44 Costa Rica to Ecuador, has 3 of its 8 species characins, 4 of the 7 species of Rhamdia, 3 in the Isthmian region; it may have origi­ of the 6 kinds of Rivulus, 11 of the 26 poe­ nated there. ciliids, and 6 of the 24 cichlids are essentiallv Cyprinodontids are represented by the restricted to the Costa Rican division as jus't essentially South American genus Rivuius delimited. A careful analysis of the nature with 6 species in the assemblage-4 confined and breadth of the transition zone must await to Panama, 1 in both Costa Rica and Pan­ the completion of studies now in progress. ama, and 1 occurring in Costa Rica and A noteworthy aspect of the distributional Nicaragua. The genus Oxyzygonectes is patterns for freshwater fishes in Panama is nearly restricted to the province. that many of the species of the opposite Nineteen of the 24 species of cichlids be­ slopes of central Panama are identical or han' long to the genus Cichlasoma. Of these, 10 their closest relatives in the two regions. occur only on either the Pacific or Caribbean Also noteworthy is the great similarity of the slopes of Costa Rica, including 3 that spill faunas of the Pacific slope of Panama (east over into the adjacent San Juan basin of of the Azuero Peninsula) and that of the Nicaragua; 2 are confined to Panama; 3 are Caribbean slope of Colombia-particularly generally distributed in both countries; and the likeness of the fishes of the Rio Tuira the remainder occur either to the north (4) and Rio Atrato (Hildebrand, 1938). or the south (1). This again indicates greater Chiapas- Guatemalan Highlands. - Fishes differentiation of Cichlasoma as one moves are rare in the highlands of Central America up Central America from Panama. much above 1500 m. For example, in Panama Freshwater fishes of marine origin number Hildebrand found that streams over this ele­ far fewer than in the Usumacinta assem­ vation were fishless. Probably in this region. blage: 8 species in 7 genera and 6 families to which no north or south temperate fish

135 784 COPEIA, 1966, NO.4

has gained access, temperature provides an PRELI~IINARY DISTRIBCTONAL LIST OF THE effective barrier to the tropical fauna. This FREsHW A TER FISHES OF CENTRAL AMERICA idea gains support by the successful introduc­ From the Isthmus of Tehuantepec (Rio tion of North American trout into these Papaloapam and Rio Tehuantepec) highland Panamanian waters. Farther north to the Panama-Colombia border. in Central America, waters between 1200 and 2500 m are abundantly and chiefly supplied 1. PRBIARY FISHES

with representatives of the prim'itive cyprino­ CHARACIDAE dontid genus Profundulus (Miller, 1955). At CURIMATINAE the higher elevations, generally above 1500 Curimata magdalenae Steindachner m, this is the only native fish. The area oc­ Pacific slope of Costa Rica (Bussing) to cupied comprises part of an ancient land eastern Panama (Rio Tuira) and Atlantic mass (West, 1964:38) extending from the slope of Colombia (Atrato and Magdalena highlands of Chiapas, "Mexico, eastward across basins) to Venezuela (:Maracaibo basin). Guatemala into Honduras and beyond. The river systems in this mountainous area are P ARODONTINAE often precipitous and subject to wide fluctu­ Apareiodon dariensis (Meek and Hildebrand) ations in flow-thus requiring special adapta­ Known only from the Rio Tuira basin. tions for aquatic organisms. Again, no truly Pacific slope of eastern Panama. northern (Nearctic) fishes have succeeded in A pareiodon compressus Breder gaining natural access to this highland. Dis­ Known from the type only; in the Rio tributional and speciational consideratior', Tuira basin, Pacific slope of eastern Pan­ indicate that Profundulus evolved within and ama. spread outward from this land mass. CHARACINAE Characidium marshi Breder COM~IENTS ON DISTRIBUTIONAL LIST Known only from the Rio Tuira basin, The list that follows forms the basis for Pacific slope of eastern Panama. the preceding analysis. It is derived from my Phenagoniates macrolepis (Meek and Hilde­ own manuscript check lists of the fishes of brand) Mexico and Guatemala, the literature, and Pacific slope of eastern Panama (Rio lists or information supplied by others, a<; Tuira) and the Atlantic slope of Colombia often indicated by placing these individuals' (Rio Atrato) to the l\laracaibo drainage of names in parentheses at the ends of the Venezuela. range statements (see also acknowledgments). Compsura gorgonae (Evermann and Golds­ Treated first are the primary freshwater borough) fishes, followed by the secondary and periph­ Both slopes of central Panama and the Rio eral groups. Many brackish-water species are Tuira of eastern Panama, westward on the included; subspecies are not included. It i, Pacific slope to southern Puntarenas Prov­ emphasized that the list is preliminary, since ince, Costa Rica (Bussing). I recognize that those currently working on Pseudocheirodon affinis Meek and Hilde­ Central American fishes will see a need for brand both additions and deletions. For some spe­ Both slopes of central Panama and the Rio cies, documentation is given from the litera­ Tuira on the Pacific slope of eastern Pan­ ture-of which all important references are ama. cited at the end of the paper. An attempt i, Pseudocheirodon sp. made to list the fishes in phylogenetic se­ Pacific slope of Costa Rica (Bussing). quence. Species believed to be new are in­ Astyanax fasciatus (Cuvier) cluded since these were considered in the dis­ Atlantic slope from Argentina to the Rio tributional analysis. A number of cichlid, Nueces, southern Texas, Pacific slope from reported by Behre (1928) from Panama (and Colombia to the Rio Armeria basin, ] alisco, accepted by Hildebrand, 1938) are not con­ Mexico. sidered because they represent known or Astyanax nasutus Meek suspected misidentifications. Great Lakes of Nicaragua (Rivas).

136 '\IILLER-CENTRAL A.\lERICA:\'" FISHES 785

Astyanax ruben-imus Eigenmann Hyphessobrycon milleri Durbin Both slopes of Panama and Colombia. Atlantic slope from the Rio Sarstun U). Astyanax kompi Hildebrand British Honduras to the Rio l\lotagua, Crater lakes at Volcan, Pacific slope of Guatemala. extreme western Panama. Hyphessobrycon tortuguerae Bohlke Astyanax albeolus Eigenmann Atlantic slope of Costa Rica and ':\ icaraguZi Pacific slope of Costa Rica and Panama (including Lake Managua; Astorqui). (Loftin). Hyphessobrycon sp. Astyanax regani Meek Pacific slope of Costa Rica (Bussing, Riyas). Pacific slope of Costa Rica. Bryconamericus emperador (Eigenmann and Bramocharax bransfordi Gill Ogle) Atlantic slope of Nicaragua and Costa Both slopes of central Panama and the Rio Rica. Tuira basin; Pacific slope of Panama: Bramocharax elongatus l\leek Pacific slope of Costa Rica. Atlantic slope of Nicaragua and Costa Rica Bryconamericus zeteki Hildebrand (Bussing). EI Valle, Pacific slope of Panama. Bramocharax sp. Bryconamericus cascajalensis Meek and Hilde­ Rio Usumacinta basin, Peten, Guatemala: brand perhaps the same as one of the aboye Known only from the Atlantic slope of (Rosen). Panama, in the Rio Casca jal, Portobelo. east of ColOn. Creagrutus affinis Steindachner Pacific slope of Panama (Rio Tuira) ami Bryconamericus ricae Eigenmann Atlantic slope of Nicaragua (Lake Nicara­ Colombia (Rio San Juan), and Atlantic gua; Astorqui), Costa Rica, and western slope of Colombia (Rio Atrato, Rio Cauca). Panama (perhaps the same as B. sclero­ Creagrutus notropoides Meek and Hildebrand parius). Known only from the upper Rio Chagres Bryconamericus scleroparius (Regan) basin, Atlantic slope of Panama. Atlantic slope of Costa Rica (Bussing). Brycon argenteus :Meek and Hildebrand Hemibrycon dariensis l\feek and Hildebrand Pacific and Atlantic (San BIas coast) slopes Pacific slope of Panama (Loftin). of Panama (Loftin). Rhoadsia eigenmanni (Meek) Brycon guatemalensis Regan Lake Nicaragua (Riyas) to Atlantic slope Atlantic slope from the Rio Grijalva, of Costa Rica (Bohlke. 1958: 178: Bussing). Tabasco and Chiapas, Mexico south'ward Roeboides guatemalensis (Gunther) to eastern Panama. Atlantic slope from the Rio Chagres. Pan­ Brycon petrosus Meek and Hildebrand ama north to Lake Nicaragua. Atlantic and Pacific slopes of Panama Roeboides occidentalis Meek and Hildebrand (Loftin). Pacific and Atlantic (San BIas coast) slopes Brycon obscurus Hildebrand of Panama (Loftin). Known only from El Valle, Pacific slope Roeboides salvadoris Hildebrand of western Panama. Pacific slope from the Rio Tehuantepec. Brycon striatulus (Kner and Steindachner) Chiapas, :Mexico, to both slopes of Costa Pacific slope of central and eastern Panama. Rica (Rivas): also in far 'western Panama Atlantic and Pacific slopes of Costa Rica (Loftin). (Bussing). Grphyrocharax atricaudatlls l\Ieek and Hildf'­ Brycon chagrensis (Kner and Steindachner) brand Atlantic slope of central Panama (Loftin). Rio Chagres on Atlantic slope and Canal Brycon behreae Hildebrand Zone to Rio Tuira on Pacific slope. Pan­ Pacific slope of western Panama. ama. Hyphessobrycon panamensis Durbin Gephyrocharax intermedius l\Ieek and Hilde­ Atlantic slope of Panama to the Rio Mag­ brand dalena basin, Colombia. Pacific slope of Panama (Rio Chame to Hyphessobrycon compressus (Meek) Rio Chiriqui) and Costa Rica (Bussing). Atlantic slope from the Rio Papaloapam. Gephyrocharax whaleri Hildebrand Mexico to the Rio Polochic, Guatemala. Pacific slope of Panama.

137 786 COPEIA, 1966, NO.4

LEBIASINAE Known only from the Pacific slope of Piabucina festae Boulenger Panama (Loftin). Pacific slope of eastern Panama (Rio Tuira) Hypopomus occidentalis Regan and Colombia (Rio San Juan) and the Both slopes of Panama and Colombia to Atlantic slope of eastern Panama (Loftin) the Maracaibo basin, Venezuela. and Colombia (Rio Atrato). Eigenmannia virescens (Valenciennes) Piabucina panamensis Gill Pacific slope of Panama to the Rio Mag­ Both slopes of Panama excluding the Rio dalena (Atlantic slope) of Colombia. Tuira basin; Pacific slope of Costa Rica thence southward to Buenos Aires. (Bussing). Apteronotus rostratus (Meek and Hilde­ Piabllcina sp. brand) Pacific slope of Costa Rica (Bussing). Pacific slope of eastern Panama and Atlan­ tic slope of Colombia (Rio Magdalena). GASTEROPELECINAE Gasteropelecus maculatus Steindachner CATOSTOMIDAE Pacific slope of Panama from the Rio Ictiobus bubalus (Rafinesque) Chorrera eastward; also on the Pacific Eastern United States, west of the Appala­ (Rio San Juan) and Atlantic (Rio Mag­ chians, southward to the Rio Papaloapam, dalena) slopes of Colombia to Venezuela Veracruz, Mexico. (~f aracaibo basin). Ictiobus meridionalis (Gunther) Atlantic slope of southern Mexico and CTEl\OLUCIINAE Guatemala from the Rio Papaloapam to Ctenolucius beani (Fowler) the Rio Usumacinta. Pacific slope of central and eastern Pan­ ama, southward to the Rio San Juan. ICTALURIDAE Colombia: also from the Rio Atrato basin, Ictalurus meridionalis (Gunther) A.tlantic slope of Colombia. Atlantic slope from Rio Papaloapam. southern 1\fexico, to Rio Usumacinta. ERYTHRININAE northern Guatemala, and Belize River. Hoplias microlepis (Gunther) British Honduras (very close to and per­ Both slopes of Panama and Pacific slope haps the same as I. furcatus) (Lesueur). of Costa Rica (Bussing). Hoplias malabaricus (Bloch) A UCHENIPTERIDAE Pacific slope of eastern Panama, from the Tmchycorystes amblops (Meek and Hildc­ Rio Bayano and Tuira basins, southward brand) to southern Colombia; also from the San Pacific slope of Panama. BIas coast on the Atlantic slope of Panama AGENEIOSIDAE (Loftin) and the Rio A.trato basin In Colombia to Buenos Aires, Argentina. Ageneiosus caucanus Steindachner Pacific slope of Panama (Rio Tuira basin). GYMNOTIDAE Rio Atrato and Rio Magdalena basins, C),17l1lotus campo Linnaeus? Colombia, and Venezuela (Caracas; identity Both slopes of Costa Rica and Nicaragua uncertain). (-:\fyers) and the Atlantic slope of ex­ treme ·western Panama (Behre, 1928: 31 0'\: PIMELODIDAE northern limit of range of this South Imparales sp. American species very uncertain. Atlantic slope (Rio Chagres) and Pacific Cymnotus c)'lindricus LaMonte slope (Rio Santa Maria) of Panam;\ Atlantic slope from Rio 1\fotagua, Guate­ (Loftin). (Imparales Schultz, 1944b.) mala to northwestern Honduras. Rhamdia wagneri (Gunther) Gymnotus sp. Both slopes of Costa Rica and Panam;\ Pacific slope of Guatemala southward at southward to Colombia. least to tributaries of the Gulf of Fonseca, Rhamdia brachypterus (Cope) ~icaragua (Carr and Giovannoli, 1950: 11- Mountain streams of south-central Veracruz, 12, as C. carapo). Mexico (Orizaba, Motzorongo-Rfo Blanco, Sternopygus dariensis Meek and Hildebrand Rio Papaloapam); "probably not distinct

138 ~IILLER-CENTRA.L A;\IERICA.:'\ FISHES 787

from" R. guatemalensis (Regan, 1906-08: Pimelodus clarias (Bloch) 128). Rio Tuira basin, Pacific slope of eastern Rhamdia heteracantha Regan Panama. southward on Atlantic slope to Atlantic slope of Costa Rica. Rio de La Plata (Uruguay-Argentina). Rhamdia rogersi Regan Pimelodella chagresi (Steindachner) Both slopes of Costa Rica and Panama. Pacific slope of Costa Rica (Bussing) and Rhamdia cabrerae Meek both slopes of Panama southward to the Pacific drainage of Guatemala (and El Sal­ Atrato and ;\Iagdalena basins (\tlantic vador?). slope) of Colombia and the ~raracaibo Rhamdia hypselura (Gunther) basin. Venezuela. Pacific slope of southern Mexico. TRICHO:\IYCTERIDAE Rhamdia gllatemalensis (Gunther) Both slopes of ;\fiddle America, from ncar Pygidilll71 striatum Meek and Hildebrand Veracruz City and Rio Tehuantepec, ;\fex­ Pacific slope of Costa Rica near Panaman­ ico, southward to Costa Rica (incIudt>s R. ian border (Bussing) to eastern Panama godmani (Gunther), but Behre's. 1928. (Rio Tuira). Atlantic slope of Panama (Loftin). and both slopes of Colombia. record of godmani from western Panam/i P:vZidiu171 septentrionale Belue represents a different species; specimens Pacific slope of western Panama (Rio examined at UMMZ) . Chiriqui del Tire). Rhamdia salvini (Gunther) Rio San Geronimo, Atlantic drainage of CALLICHTHYIDAE Guatemala. Hoplosternllm thoracatlllll (Valenciennes) Rhamdia polycallllls (Giinther) Pacific slope of central Panama to tlw Rio San Geronimo, Atlantic drainage of Atlantic drainage of Colombia. south'warn Guatemala. to the Orinoco and upper Paraguay. Rlw171dia managuensis (Gunther) Great Lakes of Nicaragua. LORICARIIDAE Rhamdia microptera (Gunther) Astrobleplls longifilis (Steindachner) Rio San Geronimo, Atlantic drainage of Pacific slope of eastern Panama (Rio Tuira Guatemala. basin) to PenL (I follow Gosline. 1947. Rhamdia motaguensis (Gunther) in regarding Astroblepus as a loricariin.) Atlantic slope of British Honduras (Stann HYPOStOlllllS plecostom us (Linnaeus) Creek) and Guatemala (Rio Polochic, Rfo Pacific slope of Costa Rica (Hildebrand. ;\Iotagua) to Honduras. 1938:238) and both slopes of Panam;i. Rhmndia alfaroi Fowler southward to Uruguay. Escobal, on Pacific slope of Costa Rica. Chaetostoma fischeri Steindachner Rha171dia nasuta l\Ieek Both slopes of central Panama ann thf' Pacific slope of Costa Rica. Pacific slope of eastern Panam,i; both slopes Rhamdia laticallda (Heckel) of Colombia, southward into Ecuador. "Mexico." Ancistrus chagresi Eigenmann and Eigen­ Rhamdia nicaraguensis (Gunther)? mann Great Lakes, Rio Prinzapolca, and Rio Both slopes of central Panam,i . Huahuashan, Nicaragua (Astorqui). . -1ncistrus spinoslls ;\Ieek and Hildebrand Rhamdia barbata Meek Pacific slope of central and eastern Pan­ Great Lakes of Nicaragua. ama (Rio Bayano, Rio Tuira). Rhamdia brachycephala Regan Lasiancistrus planiceps (l\Ieek and Hilde­ Pacific slope of Guatemala to Honduras brand) (?). Pacific slope of eastern Panama (Rio Tuira Rhamdia underwoodi Regan basin) . .\tlantic and Pacific slopes of Costa Rica Leptoancistrlls canensis (Meek and Hilde­ and Panama. brand) Rhamdia sp. Pacific slope of Panal11,i east of the Canal Rio Usumacinta basin, Guatemala aile! Zone and the A.tlantic slope (San BIas) of Belize River, British Honduras .. Panama (Loftin).

139 788 COPE lA, 1966, NO.4

Loricaria uracantha Kner and Steindachner Cristobal de las Casas, Chiapas, Mexico. .-\tlantic and Pacific slopes of Panama Profundulus labialis (Gunther) (Loftin) to Venezuela (Schultz, 1944a:325). Atlantic slope of southern .Mexico (Chiapas) Loricaria latiura Eigenmann and Vance into central Guatemala (both slopes). Pacific slope of Panama (Rio Tuira) and Profundulus candalarius Hubbs Atlantic slope of Colombia (Rio Atrato). Atlantic (or interior) drainage of extreme Loricaria variegata Steindachner northwestern Guatemala and adjacent Pacific slope of Panama (Rio Bayano, Rio Chiapas, Mexico (Rio Grande de Comitan, Tuira) and Atlantic slope of Colombia headwaters Rio Grande de Chiapa). (Rio Atrato, Rio Magdalena) to Venezuela. Fundulus grandissimus Hubbs Loricaria capetensis Meek and Hildebrand Confined to the Yucatan Peninsula, 1\Iex­ Known only from the Rio Tuira basin, ico. eastern Panama. Fundulus persimilis Miller Loricaria filamentosa Steindachner Pacific slope (Rio Tuira) of eastern Pan­ Known only from Rio Lagartos, outer tip ama; Atlantic slope of Colombia to the of Yucatan Peninsula. ~I aracaibo basin, Venezuela (Schultz, 1944a: Oxyzygonectes dowi (Gunther) 328). Pacific slope from Poneloya, Nicaragua Loricaria altipinnis Breder (Rivas) to Los Santos Province, western Knmvn only from the Rio Chucunaque Panama (Loftin). (Rio Tuira basin) and confluents, Pacific Rivulus myersi Hubbs slope of eastern Panama. Vicinity of Progreso, Yucatan, Mexico. Sturisoma panamense (Eigenmann and Eigen­ Rivulus isthmensis Garman mann) Both slopes of Costa Rica and Nicaragua Pacific slope of central to eastern Panama (includes R. flabellicauda Regan). (Loftin) through both slopes of Colombia Rivulus volcanus Hildebrand to western Ecuador. Crater lakes near Volcan on Pacific slope St1lrisoma citurense (Meek and Hildebrand) of extreme western Panama. Pacific slope of Panama (Rio Bavano. Rio Rivulus hildebrandi Myers Tuira). Pacific slope of Costa Rica (Bussing) and western Panama (Chiriqui Province). II. SECONDARY FISHES Rivulus tenuis (Meek) LEPISOSTEIDAE Lowland Atlantic streams of Veracruz Lepisosteus tropicus (Gill) (near Veracruz City), Mexico southward Atlantic slope of Middle America in Lake into Honduras (includes R. hendrichsi Nicaragua, Rio San Juan and tributaries Alvarez and Carranza). (Nicaragua, Costa Rica), and the Rio Rivulus brunneus Meek and Hildebrand Usumacinta to Rio Coatzacoalcos basins Atlantic slope of the Canal Zone. of Guatemala and southeastern Mexico: Rivulus montium Hildebrand Pacific slope from southern Chiapas. "!\rex­ Rio Chagres basin, Atlantic slope of Pan­ ico southward to the Gulf of Fonseca, ama. Nicaragua (Rio Negro). Rivulus godmani Regan Rio Motagua basin, Atlantic slope of CYPRljI;ODO~TIDAE Guatemala. Profundulus guatemalensis (Gunther) Rivuius sp. Both slopes of Guatemala southward to Rio Papaloapam and Rio Coatzacoalcos El Salvador and Honduras (Rio Lempa basins, Veracruz and Oaxaca, Mexico. basin). Rivulus chucunaque Breder Profundulus punctatus (Gunther) Both slopes of Middle America from south­ Rio Chucunaque basin (Pacific slope), ern Mexico (Rio Aguacatillo near Aca­ Darien, Panama. pulco,. Rio Coatzacoalcos) southward to El Cyprinodon variegatus Lacepede Salvador. The subspecies C. v. artifrons is confined Profundulus hildebrandi Miller to fresh and brackish waters of the Yucatan Known 'only from the vicinity of San Peninsula.

140 MILLER-CENTRAL Al\IERICA~ FISHES 789

Cyprinodon beltrani Alvarez Poecilia sulphuraria (Alvarez) Known only from Laguna de Chich­ Banos del Azufre, 4 miles W of Teapa, ankanab, Quintana Roo, Mexico. Tabasco, :\Iexico. Floridicthys carpio (Gunther) Poecilia latipinna (Lesueur) Two subspecies (F. c. barbouri and F. c. Coastal waters of the Atlantic slope from polyommus) are confined to the Yucatan South Carolina to Campeche (near Cham­ Peninsula. poton), :\Iexico. Garmanella pulchra Hubbs Poecilia petenensis (Gunther) Yucatan Peninsula to Corozal, British Lake Peten and the Rio Usumacinta basin, Honduras. Guatemala. Poecilia velifera Regan ANABLEPIDAE Northern part of the Yucatan Peninsula, Anableps dowi Gill including Isla Mujeres. Probably confined Pacific drainage from the Rio Tehuantepec, naturally to coastal waters of the outer Oaxaca, Mexico to the Gulf of Fonseca part of the peninsula (Hubbs, 1936:248). drainage of Nicaragua (Astorqui). Priapella bonita (Meek) Upper part of the Rio Papaloapam basin, POECILIIDAEl Atlantic slope of Veracruz, Mexico. Alfaro cultratus (Regan) Priapella interrnedia Alvarez and Carranza Atlantic drainage of western Panama and Upper part of the Rio Coatzacoalcos basin. Costa Rica to Zalaya, Nicaragua (including .-\tlantic slope of Oaxaca. Mexico. tributaries of L. Nicaragua; Astorqui); Priapella compressa Alvarez Pacific drainage in Tilaran region, Costa Palenque (Rio Usumacinta basin) to Rica (Bussing). Rio Teapa Grijalva basin), Chiapas, A lfara huberi (Fowler) ~fexico. Atlantic drainage from southeastern Guate­ Xiphopho1'lls mawlatus (Gunther) mala to Nicaragua (Rio Prinzapolca). Atlantic slope from the Rio .Tamapa basin just south of Veracruz City, l\lexico south­ Poecilia caucana (Steindachner) ward to northern British Honduras. Pacific drainage of Darien, Panama to Rio Xiphophorus milleri Rosen Cauca, Colombia and Lake Maracaibo, Lake Catemaco basin in Rio Papaloapam Venezuela. system, Veracruz, Mexico. Poecilia mexicana Steindachner Xiphopho1'lls clemenciae Alvarez Atlantic slope from the Rio San Juan (in Rio Coatzacoalcos basin on Atlantic slope Rio Grande basin), Nuevo Leon, Mexico of Oaxaca, l\fexico. and Pacific slope from the Rio del Fuenc Xiphopho1'lls helleri Heckel basin, Sonora, Mexico southward to the Atlantic slope from Rio Nautla, Veracruz, Caribbean slope of Colombia, the Pacific ;\fexico southward to northern Honduras. slope of eastern Panama (Rio Tuira), and Genus and species (Cnesterodontini) Rosen: the Netherlands and Colombian 'Vest In­ Rio Usumacinta basin in Peten, Guate­ dies. (See detailed description by Hubbs, mala (two distinct species). 1953, of a Panamanian population; listed Brachyrhaphis terrabensis (Regan) as lHollienesia sphenops gilli.) Pacific slope of Costa Rica and extreme Poecilia sphenops Valenciennes western Panama, at high elevations. Atlantic slope of Middle America from Brachyrhaphis hartwegi Rosen and Bailev lagoon about 30 airline miles N of Vera­ Pacific drainage of Chiapas. l\fexico. and cruz City, Mexico southward to Costa Guatemala (Rosen). Rica; Pacific slope from AguiliIIa, Michoa­ Brachvrhaphis rhabdophora (Regan) can, :\fexico sout1rward to Costa Rica. Both slopes of Costa Rica. Rraclrvrhaphis pfJiscopi (Steindachner)

1 Not including an unidentified genus and species Both slopes of central Panama. from Laguna Ocotal, Mexico, known only from females Brach),rhaphis punctifer (Hubbs) (Miller, 1957:240). P. formosa (Girard), said to range southward to the Rio Papaloapam (Rosen and Atlantic slope of western Panama (Rio Bailey, 1963:54), is also excluded as I know of no Cricamola and vicinity). records south of the Rio Panuco system or the coastal Brach)'rhaphis cascajalensis C\feek and Hilde­ Laguna de Tampamachoco near Tuxpan, northern V('ra­ cruz (specimens kindly loaned by R. M. Darnell). brand)

141 790 COPEIA, 1966, NO.4

.\tlantic slope of Costa Rica to San BIas Priapichthys annectens (Regan) Proyince, Panama. Atlantic and Pacific slopes of Costa Rica. Brachyrhaphis parismina (Meek) Priapichthys panamensis Meek and Hilde­ Atlantic slope of Costa Rica. brand Brachyrhaphis sp. Pacific slope from Rio San Juan, Nicara­ Chiriqui Proyince on Pacific slope of west­ gua (Rivas) to Panama in provinces of ern Panama (Loftin). Veraguas, Cocle, and Panama (Loftin). Brachyrhaphis sp. Priapichth)lS darienensis (Meek and Hilde­ Atlantic slope of Costa Rica (Bussing). brand) Gambusia nicaraguensis Gunther Pacific slope of central and eastern Panama Atlantic slope from Lake Izabal, Guate­ (Loftin) including Isla San Jose (Hilde­ mala southward to the Rio Chagres basin, brand, 1946). Panama (where possibly introduced; not Xeoheterandria tridentiger (Garman) taken by Loftin in his suney, 1961-1965). Both slopes of central Panama and Isla Gambusia yucatana Regan Taboga. Atlantic slope from lower part of Rio X eoheterandria cana (Meek and Hildebrand) Coatzacoalcos, Veracruz, Mexico southward Rio Satigante at Cana, tributary of Rio to northern British Honduras including Tuira, in Darien Province, Pacific slope of Yucatan Peninsula. extreme eastern Panama (same as .?\,T. tri­ Gambusia sexradiata Hubbs dentiger ?). Atlantic slope from Rio Nautla, northern Neoheterandria umbratilis (Meek) Veracruz, Mexico southward to northern Atlantic slope of Costa Rica; Lake Nicara­ Guatemala (V sumacinta basin) and gua (Rivas, 1965; Astorqui). northern British Honduras (Belize River Heterandria bimaculata (Heckel) drainage, un catalogued material at VM­ Atlantic slope of Middle America from a stream 20 miles north of San Jose Cardel ~JZ); absent from most of the Yucatan Peninsula. (UMMZ 184512), northwest of Veracruz City, Mexico southward to northeastern Gambusia sp. Nicaragua (Rio Prinza polca). Vicinity of Teapa, Tabasco, Mexico. Heterandria sp. Gambusia luma Rosen and Bailey Head"raters of Rio Lempa, Pacific slope of Atlantic slope of British Honduras and Guatemala and El Salvador. Guatemala (Belize River to Puerto Barrios). Poeciliopsis turrubaTensis (Meek) Heterophallus rachowi Regan Pacific slope from near N uxco (60 miles Known only from the Rio Coatzacoalcos N'V of Acapulco), Guerrero, Mexico south­ basin, Veracruz, Mexico. ward to Rio Dagua, Colombia; entering Heterophallus echeagarayi (Alvarez) the sea. Known only from Palenque, Chiapas, Poeciliopsis gracilis (Heckel) Mexico. Pacific slope from Rio Verde basin near H eterophallus sp. Oaxaca City, Oaxaca, Mexico eastward to Rio Teapa, Teapa, Tabasco, ~rexico. tributaries of Gulf of Fonseca in Honduras Belonesox belizanus Kner (Rio Choluteca) and probably Nicaragua. Atlantic slope of Middle America from On Atlantic slope of Mexico from 20 miles Laguna San Julian, northwest of Veracruz N of Veracruz City southward to Rio Coat- (Suttkus), Mexico, southward into Costa 7acoalcos; also in upper Rio Grijaln Rica (Caldwell, et al., 1959: 17) . (Chiapas and adjacent Guatemala), Rio Carlhubbsia stuarti Rosen and Bailey Motagua (Guatemala), Rio Vlua (Hon­ Basin of the Rio Polochic and Lake l7ahal. duras), and Lake Nicaragua (Suttkus). Atlantic slope of eastern Guatemala. Poeciliopsis sp. Cnrlhubbsia kidderi (Hubbs) Lake Catemaco, in Rio Papaloapam hasin. ,-\ tlaptic slope from the Rio San Pf'aro Veracruz, Mexico. ;\fartir and Rio de la Pasion (Vsumacinta Poeciliopsis sp. basin), Peten, and Alta Vera Paz, Guate­ Laguna Pajaritos near Coatzacoalcos, Vera­ mala, northward to the Rio Champoton, cruz, Mexico. Campeche and eastward to the Rio Poeciliopsis fasciata Meek Grijalva 'basin, Tabasco. Mexico. Pacific slope of Mexico (:\'uxco, 60 miles

142 MILLER-CENTRAL AMERICAN FISHES 791

NW of Acapulco) to northwestern Guate­ Cichlasoma fenestratum (Gunther) mala; Atlantic headwaters of Rio Coat­ Atlantic slope of Veracruz (Rio Chacha­ zacoalcos near crest of Isthmus of Tehuan­ lacas basin) and Oaxaca, Mexico, not tepec, Oaxaca, Mexico. east of the Rio Coatzacoalcos. Poeciliopsis elongata (Gunther) Cichlasoma sexfasciatum Regan Pacific slope of western to central Panama. Known only from "Guapote, Mexico." Poeciliopsis retropinna (Regan) Cichlasoma bifasciatum (Steindachner) Pacific slope of Costa Rica and western Rio Usumacinta basin in Peten, Guate­ Panama. mala and adjacent parts of Mexico. Poeciliopsis sp. Cichlasoma heterospilum Hubbs Pacific slope of Costa Rica (Bussing). Rio Usumacinta basin in Guatemala and Phallichthys amates (Miller) Mexico. Atlantic slope from Rio Motagua basin to Cichlasoma guttulatum (Gunther) western Panama, Pacific slope of central Pacific slope of Middle America from Costa Rica. Rio Tehuantepec, Oaxaca, Mexico, into Phallichthys fairweatheri Rosen and Bailey Guatemala; Atlantic slope only in Rio Northern British Honduras (N of Belize Coatzacoalcos basin, Mexico (includes C. River) and northern Guatemala (Rio Usu­ zona tum Meek). macinta basin). Cichlasoma godmani (Gunther) Phallichthys tico Bussing Atlantic slope of Guatemala (Rio Polo­ Atlantic slope of western Costa Rica (Rio chic basin, Lake Izabal, Sulphur River San Juan basin). near Puerto Barrios). Xenodexia ctenolepis Hubbs Cichlasoma microphthalmum (Gunther) Atlantic drainage of central Guatemala, in Rio Motagua basin on the Atlantic slope upper part of Rio Usumacinta basin. of Guatemala (includes C. oblongum (Gunther), C. milleri Meek, and C. caeru­ CICHLIDAE leogula Fowler). Aequidens coeruleopunctatus (Kner and Cichlasoma gadowi Regan Steindachner) Rio Tonto and tributaries, Rio Papa­ Atlantic slope of central and eastern Pan­ loa pam basin, Mexico. ama, Pacific slope of Costa Rica (Bussing) Cichlasoma intermedium (Gunther) and Panama, thence (doubtfully) to Co­ Basins of the Rio Usumacinta in Guate­ lombia. mala and Mexico and of the Rio Gri­ Geophagus crassilabris Steindachner jalva, Mexico (includes C. anguiliferum Both slopes of central and eastern Panama. (Gunther) ). Cichlasoma, section Theraps: Cichlasoma sieboldi (Kner and Steindach­ Cichlasoma eigenmanni Meek ner) Rio Tonto and its tributaries in the Rio Pacific slope of Costa Rica to cen tra] Papaloapam basin, Mexico. Cichlasoma nebuliferum (Gunther) Panama; possibly Atlantic slope of Pan­ Atlantic slope of southern Mexico (Rio ama (includes Herichthys underwoodi Coatzacoalcos to Rio Grijalva ?). Regan; Bussing). Cichlasoma maculicauda Regan Cichlasoma irregulare (Gunther) Atlantic slope from the Rio Usumacinta Upper tributaries of the Rio Usumacinta basin, Peten, Guatemala, to Panama in Mexico and Guatemala and the Rio (Rio Chagres). Polochic drainage, Guatemala. Cichlasoma melanurum (Gunther) Cichlasoma lentiginosum (Steindachner) Rio de la Pasion and Lake Peten and Rio Usumacinta basin in Guatemala and adjacent lakes, Guatemala, and Belize Mexico. River, British Honduras. Cichlasoma balteatum (Gill and Bransford) Cichlasoma synspilum Hubbs Great Lakes of Nicaragua (synonym of Atlantic slope from Rio Usumacinta C. nicaraguense ?). basin, Guatemala to Belize River, British Cichlasoma nicaraguense (Gunther) Honduras (may not be distinct from Great Lakes of Nicaragua (includes C. C. melanurum; includes C. hicklingi balteatum ?). Fowler). Cichlasoma sp.

143 792 COPEIA, 1966, NO.4

Rio de la Pasion (Rio Vsumacinta basin), Cichlasoma hetemdontum (Vaillant and northern Guatemala. pellegrin) Cirhlasoma, section Archocentrus: Pacific slope streams of the Isthmus of Cichlasoma spilurum (Gunther) Tehuantepec, Oaxaca and Chiapas, Atlantic slope from Belize River, British Mexico (includes C. evermanni Meek). Honduras, southward to Bocas Province, Cichlasoma altifrons (Kner and Steindach­ western Panama; also on Pacific slope of ner) Costa Rica (includes C. cutteri Fowler). Pacific slope of western Panama and Cichlasoma nigrofasciatum (Gunther) Costa Rica. Pacific slope of Guatemala to Costa Cichlasoma mstratum (Gill and Bransford) Rica; both slopes of Costa Rica. Great Lakes of Nicaragua to the Atlantic Cichlasoma octofasciatum (Regan) and Pacific slopes of Costa Rica (Bussing). Atlantic slope from Rio Paso San Juan Cichlasoma popenoei Carr and Giovannoli (20 miles W of Veracruz City), Veracruz. Rio Choluteca basin, Pacific slope of Mexico to Honduras (Rio Vlua basin); Honduras. also in Yucatan Peninsula (includes C. Cichlasoma margaritiferum (Gunther) hedricki Meek). Known only from the type, from the Cichlasoma centrarchus (Gill and Brans­ Peten region of Guatemala. ford) Cichlasoma citrinellum (Gunther) Atlantic slope from the Great Lakes and Atlantic slope of Nicaragua (including Rio San Juan, Nicaragua into Costa the Great Lakes basin) and Costa Rica. Rica (Caldwell, et al., 1959:26). Cichlasoma erythraeum (Gunther) Cichlasoma immaculatum Pellegrin Great Lakes of Nicaragua (possibly the Rio Polochic basin, Atlantic slope of female of C. labiatum or the same as C. Guatemala. citrinellum). Cichlasoma spinosissimum (Vaillant and Cichlasoma lobochilus (Gunther) Pellegrin) Great Lakes of Nicaragua. Rio Polochic basin, Atlantic slope of Cichlasoma alfaroi Meek Guatemala. Both slopes of Costa Rica, Atlantic slope Cichlasoma, section "Herichthys": of western Panama (includes C. lethrinus Cichlasoma bocourti (Vaillant and Pelle­ Regan; Bussing). grin) Cichlasoma labiatum (Gunther) Lake Izabal and lower Rio Polochic. At­ Great Lakes of Nicaragua (possibly the lantic slope of Guatemala. female is C. erythraeum). Cichlasoma geddesi Regan Cichlasoma tuyrense Meek and Hildebrand Atlantic slope of southern Mexico. Pacific slope of eastern Panama (Rio Cichlasoma pearsei (Hubbs) Bayano, Rio Tuira). Atlantic slope of Mexico and northern Cichlasoma sp. Guatemala in the Rio Vsumacinta basin. including Rio Champoton, Campeche, Pacific slope of Guatemala; relative of Mexico. C. heterodontum. Cichlasoma, section Amphilophus: Cichlasoma sp. Cichlasoma robertsoni Regan Rio de la Pasion and Rio San Pedro, in Rio Coatzacoalcos basin, Veracruz, Mex­ Rio Vsumacinta basin, Peten, Guate­ ico southward along the Atlantic slope mala. to 30 miles E of Tela, Honduras (in­ Cichlasoma, section "Paraneetmplus": cludes C. acutum Miller). Cichlasoma bulleri (Regan) Cichlasoma longimanus (Gunther) Atlantic slope of southern Mexico from Great Lakes of Nicaragua. the Rio Papaloapam to the Rio Grijalva Cichlasoma macracanthum (Gunther) basins. Pacific slope of Mexico (Rio Tehuan­ Cichlasoma, section Parapetenia: tepec basin) southward to El Salvador Cichlasoma mento (Vaillant and Pellegrin) (Rio de Paz basin). Southern Mexico (Rio Negro). Cichlasoma guija Hildebrand Cichlasoma umphthalmus (Gunther) Rio Lempa basin of El Salvador and ex­ Atlantic slope of Middle America from treme southeastern Guatemala. the Rio Coatzacoalcos basin southward

144 MILLER-CENTRAL AMERICAN FISHES 793

into Nicaragua (including Yucatan Pen­ Cichlasoma ellioti Meek. insula and Isla Mujeres). Upper tributaries of the Rio Tonto in Cichlasoma hogaboomorum Carr and Gio­ the Rio Papaloapam basin, Veracruz, vannoli Mexico. (Possibly the same as C. hel­ Lower part of the Rio Choluteca, Pacific leri). slope of Honduras. Cichlasoma helleri (Steindachner) Cichlasoma trimaculatum (Gunther) Rio Grijalva basin, Atlantic slope of Pacific slope of Middle America from southeastern Mexico (range and status Laguna Coyuca NW at Acapulco, l\Iex­ not clear). ico southward to Rio Lempa, El Salva­ Cichlasoma champotonis Hubbs dor (includes C. mojarra Meek, C. cen­ Rio Grijalva-Usumacinta and Rio Cham­ trale Meek, C. gordonsmithi Fowler, and paton basins in northern Guatemala and C. cajali Alvarez and Gutierrez). southeastern Mexico. Cichlasoma tenue Meek Cichlasoma affine (Gunther) Rio Papaloapam basin, Atlantic slope Lake Peten and nearby lakes in Peten, of Mexico. Guatemala to Rio Grijalva basin, Mexico. Cichlasoma salvini (Gunther) Cichlasoma meeki (Brind) Atlantic slope from Rio Papaloapam, Northern part of the Yucatan Peninsula. Veracruz, Mexico southward to Sulphur Cichlasoma hyorhynchum Hubbs River near Puerto Barrios, Guatemala. Rio Usumacinta basin, northern Guate­ Cichlasoma friedrichsthali (Heckel) mala and Belize River, British Honduras. Atlantic slope of southern Mexico (east Cichlasoma pasionis Rivas of the Rio Coatzacoalcos) to Costa Rica. Rivers and lakes of Peten (Usumacinta Cichlasoma managuense (Gunther) basin), northern Guatemala. Great Lakes of Nicaragua and Atlantic Cichlasoma, incertae cedis: slope of Costa Rica (Rivas). Cichlasoma calobrense Meek and Hilde­ Cichlasoma dowi (Gunther) brand Great Lakes of Nicaragua and both Pacific slope of the eastern half of Pan­ slopes of Costa Rica. ama (Rio Bayano, Rio Tuira). Cichlasoma motaguense (Gunther) Cichlasoma spilotum Meek Atlantic slope of Guatemala (Rio Mo­ Atlantic slope of Costa Rica. tagua), Pacific slope of Guatemala, El Cichlasoma umbriferum Meek and Hilde­ Salvador, and Honduras. (Synonym of brand C. friedrichsthali?, see, e.g., Hubbs, Pacific slope of eastern Panama (Rio 1935:19.) Tuira) and Atlantic slope of Colombia Cichlasoma sp. (Rio Atrato, Rio Magdalena). Rio Grande de Chiapa basin (Rio Gri­ Cichlasoma terrabae Jordan and Evermann jalva system), Chiapas, in Atlantic drain­ Pacific slope of Costa Rica. age of Mexico. Cichlasoma tuba Meek Cichlasoma sp. Rio Comitan and adjoining lakes, Chia­ Atlantic slope of Costa Rica (includes pas, Mexico, an interior stream draining Tomocichla underwoodi Regan). toward the Rio Usumacinta basin. Cichlasoma sp. Cichlasoma, section Thorichthys: Upper part of the Rio Coatzacoalcos Cichlasoma callolepis (Regan) basin, Atlantic slope of the Isthmus of Upper tributaries of the Rio Coatzacoal­ Tehuantepec, Mexico. cos basin on the Atlantic slope of the Cichlasoma sp. Isthmus of Tehuantepec, Mexico. Rio Seniso near Caban, in Rio Usuma­ Cichlasoma aureum (Gunther) cinta basin, Guatemala (Hubbs, 1950: Atlantic slope, from the basin of the Rio II). Chachalacas, northwest of Veracruz City, Cichlasoma sp. Mexico, eastward and southward to the Upper tributaries of Rio de la Pasion Rio Polochic-Lake Izabal and Rio Mo­ (Usumacinta basin), Guatemala. tagua basins, Atlantic slope of Guate­ Petenia splendida Gunther mala and adjacent Honduras; possibly Atlantic slope from the Rio Grijalva basin also in British Honduras. to the Rio Usumacinta and the Belize

145 794 COPEIA, 1966, NO.4

River, in southeastern Mexico, northern Elops saurus Linnaeus Guatemala, and British Honduras. Ascending lagoons and river mouths of the N eetroplus nematopus Gunther Atlantic slope from Cape Cod to Brazil. Atlantic slope of Nicaragua (Great Lakes) Elops affinis Regan and Costa Rica (Rivas). Ascending rivers of the Pacific slope from N eetroplus panamensis Meek and Hilde­ the Colorado River (including Salton Sea), brand Arizona-California southward to Peru. Atlantic (Rio Chagres) and Pacific (Rio Tuira) slopes of Panama. CHANIDAE Herotilapia multispinosa (Gunther) Chanos chanos (Forskal) . Atlantic slope of Nicaragua (Great Lakes) Western and eastern Pacific; recorded by and Costa Rica (Caldwell, et al., 1959; Regan (1906-08: 179) from brackish lagoons Bussing). at Chiapam, Pacific coast of Guatemala.

SYNBRANCHIDAE CLUPEIDAE Synbranchus marmoratus Bloch Harengula pensacolae Goode and Bean Both slopes of tropical America from Florida southward to Brazil; taken in Rio southern Mexico (Rio Papaloapam, Rio Champoton at mouth, Yucatan Peninsula Tehuantepec) to Brazil and Peru (includ­ (Hubbs, 1936). ing Yucatan Peninsula to Islas Cozumel Harengula thrissina (Jordan and Gilbert) and Mujeres) . Gulf of California to Peru, entering brack­ Furmastix infernalis (Hubbs) ish to nearly fresh water. Known only from a cienaga in Yucatan, Lile stolifera (Jordan and Gilbert) Entering fresh water along the Pacific Mexico. slope from Baja California to Peru. Ilisha furthi (Steindachner) III. PERIPHERAL FISHES Pacific coast from Panama Bay, Panama to CARCHARHINIDAE the Gulf of Guayaquil, Peru, entering the Carcharhinus leu cas (Valenciennes) locks of the Panama Canal. Atlantic, from New York to southern Dorosoma petenense (Gunther) Kentucky and southern Indiana to Florida Brazil, entering large tropical rivers (Papa­ and Texas, thence southward to the Belize loapam, Usumacinta, San Juan) and lakes River, British Honduras. (Izabal, Nicaragua). Dorosoma anale Meek PRISTIDAE Atlantic slope from the Rio Papaloapam, Veracruz and Oaxaca, Mexico southward Pristis pectinatus Latham to the Rio Usumacinta basin, northern Widespread in the tropical-subtropical At­ Guatemala. lantic to the Mediterranean and in the Dorosoma chavesi Meek tropical eastern Pacific; in brackish water Great Lakes of Nicaragua. of lower reaches of large rivers in the tropics. ENGRAULIDAE Pristis perotteti Muller and Henle Anchovia macrolepidota (Kner and Stein­ Both sides of the Atlantic. In the Ameri­ dachner) cas, from southern Florida to Brazil, enter­ Gulf of California to Ecuador, entering ing lakes (Izabal and Nicaragua) and coastal rivers. lagoons. Anchoa spinifer (Valenciennes) Pristis microdon Latham Atlantic coast from Panama to Santos, Recorded from strictly fresh water in the Brazil; Pacific coast from Panama to Guay­ Rio Chucunaque basin, Pacific slope of aquil, Ecuador. Recorded from fresh water eastern Panama, by Breder (1927:99). in locks of the Panama Canal. A nchoa panamensis (Steindachner) ELOPIDAE Pacific coast from MazatIan, Mexico to Megalops atlanticus Valenciennes Puerto Pizarro, Peru, entering nearly fresh North Carolina to Brazil, entering larger water in Panama. rivers and lakes, (Usumacinta, San Juan, Anchoa mundeoloides (Breder) Chagres; Izabal and Nicaragua). Pacific coast from San Felipe, Baja Cali-

146 ?\IILLER-CENTRAL AMERICAN FISHES 795

fornia to Rio Los Pescadores above Masa­ Netuma oscula (Jordan and Gilbert) chapa, Nicaragua (Rivas). Pacific coast of Panama, entering the Rio Anchoa hepsetus (Linnaeus) Tuira. Atlantic slope from Maine to Uruguay, Arius melanopus Gunther entering rivers. In rivers and lakes of the Atlantic slope A nchoa lamprotaenia Hildebrand from the Rio Papaloapam, Veracruz, Mex­ Florida south through the West Indies to ico southward to Panama. Panama, entering brackish lagoons. Arius taylori Hildebrand A nchoa parva (Meek and Hildebrand) Rio Lempa, El Salvador, and possibly West Indies and Atlantic coast from Gua­ other Pacific slope streams. temala to Venezuela, entering fresh water Arius multiradiatus Gunther (Lake Izabal, Guatemala, Rio Chagres, Pacific coast from Panama Bay to Paita, Panama). Peru, entering rivers. Anchoa curta (Jordan and Gilbert) r1.rius furthi Steindachner Pacific coast from the mouth of the Rio Pacific coast of Costa Rica (Bussing) and Yaqui, Sonora, Mexico to Puerto Pizarro, Panama, entering rivers. Peru, entering nearly fresh water in Pan­ {1rius felis (Linnaeus) ama. Atlantic slope of North America from Cape Anchoa mitchilli (Valenciennes) Cod to Yucatan, entering rivers in the Cape Cod to Yucatan, entering rivers (e.g., southern part of its range. Soto la Marina) and coastal lagoons of Arius guatemalensis Gunther Mexico. Pacific slope of Middle America, from Anchoa lucida (Jordan and Gilbert) Mazatlan, Mexico to Panama, ascending Pacific coast from the mouth of the Rio rivers. Yaqui, Sonora, Mexico to Ecuador, enter­ Arius seemani Gunther ing brackish water. Pacific coast from l\;fexico to Ecuador, Anchoviella elongata (Meek and Hildebrand) en tering rivers. Atlantic slope from Rio Escondido W of Arius caerulescens Gunther Bluefields, Nicaragua (Rivas) to Panama. Pacific coast from Sonora, Mexico to Gua­ Lycengraulis poeyi (Kner and Steindachner) temala, entering lagoons (Huamuchal, Gua­ Pacific slope of Panama, from tidal streams temala). and the Rio Bayano. Arius assimilis Gunther Atlantic coast of Central America, ascend­ ARIIDAE2 ing streams and lakes (Lake Izabal, Guate­ Bagre marinus (Mitchill) mala). Atlantic coast from Cape Cod to Lake Iza­ Arius tuyra Meek and Hildebrand bal, Guatemala, entering rivers and lakes. Rio Tuira, Pacific slope of Panama, in fresh and brackish water. Bagre filamentosus (Swainson) Potamarius nelsoni (Evermann and Golds­ Atlantic coast of Central America, entering borough) rivers and lakes (Nicaragua). Rivers and lakes in the Rio Usumacinta Bagre panamensis (Gill) basin in northern Guatemala and adjacent Pacific coast from Mazatlan, Mexico to parts of southeastern Mexico. Restricted Ecuador, entering river mouths. to fresh water. Bagre pinnimaculatus (Steindachner) Potamarius izabalensis Hubbs and Miller Pacific coast from western Mexico (Mazat­ Lake Izabal and the lower Rio Polochic, Ian) to Ecuador, entering river mouths. Atlantic slope of Guatemala. Restricted to Selenaspis dowi (Gill) fresh water. Pacific coast of Panama to Ecuador, ascend­ ing streams. OPHICHTHIDAE ?\: etuma planiceps (Steindachner) Myrophis punctatus Lutken Pacific slope from Sinaloa, Mexico to Pan­ West Indies and Gulf coast of the United ama (Rio Sabana, Rio Cianati, Darien). States southward to Brazil, entering tidal streams (Caldwell, et al., 1959: 15). 2 I do not recognize Galeichthys and Arius, as de­ Pisodonophis daspilotus Gilbert fined by Meek and Hildebrand (1923: 96), as distinct genera. Pacific coast of Panama, ascending streams.

147 796 COPEIA, 1966, NO.4

ANGUILLIDAE Archomenidia bolivari Alvarez and Carranza Anguilla rostrata (Lesueur) Confined to fresh water in the Rio Coatza­ Cape Cod to Colombia, entering rivers. coalcos basin, Atlantic slope of Oaxaca and Veracruz, Mexico. HEMIRAMPHIDAE Archomenidia sp. Hyporhamphus roberti hildebrandi Jordan Rio Teapa near Teapa, Tabasco, Mexico. and Evermann Membras martinica (Valenciennes) Atlantic coast from the Rio Sarstun, Guate­ Gulf of Mexico, entering coastal lagoons. mala-British Honduras border southward Melaniris meeki (Miller) to Panama (Collette). Atlantic slope of Guatemala (Rio Motagua) Hyporhamphus snyderi Meek and Hildebrand and Honduras. Tidal streams on the Pacific coast of cen­ Melaniris chagresi (l\feek and Hildebrand) tral Panama (Rio Anton, Miraflores Locks) Fresh and brackish water of the Atlantic northward to Baja California. slope of Costa Rica and Panama. Hyporhamphus mexicanus Alvarez M elan iris sardina Meek Confined to freshwater streams and la­ Great Lakes of Nicaragua. gunas of the Rio Usumacinta basin in Melaniris guija (Hildebrand) Peten, Guatemala westward to the Rio Rio Lempa basin, Guatemala and El Coatzacoalcos, Mexico (Collette) . Salvador, and Rio Choluteca, Honduras. Chriodorus atherinoides Goode and Bean Melaniris guatemalensis (Gunther) Florida Keys, Bahamas, Cuba, and Yuca­ Pacific coast of Central America; described tan Peninsula (mouth of the Rio Cham­ from coastal lagoons at Huamuchal, Guate­ poton, Hubbs, 1936:209). mala. M elaniris sp. BELONIDAE Rio Coatzalcoalcos basin, Atlantic slope of Strongylura notata (Poey) Mexico. Bahamas, Florida. West Indies (Cuba and ll,f elan iris sp. Jamaica), and bordering mainland of south­ Lake Peten, Peten, Guatemala. ern Mexico (recorded by Hubbs, 1936:207, M elan iris sp. from brackish-water cienagas near Pro­ Rio Polochic basin, Atlantic slope of Guate­ greso, Yucatan). mala. Strongylura marina (Walbaum) Xenatherina lisa (Meek) Atlantic coast of America from Cape Cod Rio Papaloapam basin, Atlantic slope of to Brazil including the West Indies, enter­ Mexico. ing rivers (includes S. timucu (\Valbaum)). Xenatherina schultzi Alvarez and Carranza Strongylura exilis (Girard) Upper part of the Rio Coatzacoalcos. Pacific coast from California to Peru, en­ Oaxaca, Mexico. tering rivers occasionally (e.g., Rio Anton, SYNGNATHIDAE Panama Bay). Syngnathus scovelli (Evermann and Kendall) Strongylura scapularis (Jordan and Gilbert) Atlantic coast from Florida to the Yucatan Tidal streams on the Pacific slope of cen­ Peninsula, chiefly in salt and brackish tral Panama southward to Ecuador. water but penetrating river mouths. A THERINIDAE3 Syngnathus elcapitanensis (Meek and Hilde­ Menidia sp. brand) Rio Lagartos, northern tip of Yucatan Freshwater streams on the Pacific slope of Peninsula. Costa Rica and Panama. Menidia colei Hubbs Oostethus lineatus (Kaup) Brackish-water cienagas about Progreso, Atlantic coast from South Carolina to Pan­ Yucatan, Mexico. ama, entering fresh water. Archomenidia sallei (Regan) Oostethus sp. Confined to freshwater in the lower Rio Lake Izabal, Atlantic slope of Guatemala. Papaloapam basin, Atlantic slope of Vera­ Pseudophallus starksi (.Jordan and Culver) cruz, Mexico. Pacific coastal streams of mainland Mexico (near Mazatlan) and San Jose del Cabo,

3 I regard Thyrinops as a synonym of Melaniris. Ba ja California southward to Ecuador.

148 MILLER-CENTRAL AMERICAN FISHES 797

Pseudophallus mindii (Meek and Hilde­ Pacific coast from Baja California to Peru, brand) entering fresh water (Caballo Blanco, Atlantic coastal streams from Venezuela Guatemala, Meek, 1908: 140). northward to Honduras and, possibly, to Lutjanus colorado Jordan and Gilbert the Rio Sarstun, Guatemala-British Hon­ Pacific coast from Guaymas, Sonora, Mex­ duras boundary. ico to Panama, entering tidal streams.

CENTROPOMIDAE CARANGIDAE Centropomus undecimalis (Bloch) Caranx latus Agassiz Atlantic coast from Florida to Rio de Atlantic coast of tropical America, entering Janeiro, Brazil, ascending streams. rivers. Centropomus nigrescens Gunther Caranx marginatus (Gill) Pacific coast from Rio Yaqui, Sonora, and Pacific coast from MazatIan, Mexico to Baja California, Mexico to Peru, entering Panama Bay, Panama, entering fresh water. streams. Caranx hippos (Linnaeus) Centropomus parallelus Poey Both coasts of tropical America, entering Atlantic slope from near Veracruz City, tidal streams. Mexico southward to Brazil, entering fresh Oligoplites saurus (Bloch and Schneider) water. Both coasts of tropical America, entering Centropomus poeyi Chavez tidal streams. Gulf of Mexico from Tampico, Tamaulipas Oligoplites mundus Jordan and Starks to Frontera, Tabasco, Mexico, entering Pacific coast from Gulf of California to river mouths (Chavez, 1961). Peru, in tidal streams and coastal lagoons. Centropomus pectinatus Poey Both coasts of tropical America, entering GERRIDAE brackish water. Eucinostomus argenteus Baird and Girard Centropomus armatus Gill Atlantic coast from New Jersey to Brazil, Pacific coast from Chiapam, Guatemala to Pacific coast from California to Ecuador. Guayaquil, Ecuador, entering tidal streams. entering brackish water. Centropomus robalito Jordan and Gilbert Eucinostomus pseudogula Poey Pacific coast from Rio Yaqui, Sonora, Bermuda, Florida, West Indies, and south­ Mexico to Peru, entering rivers. eastern Mexico to Brazil (taken in Rio Centropomus ensiferus Poey Champoton, Yucatan; Hubbs, 1936:252). Atlantic coast from Florida to Brazil; taken (Same as E. melanopterus?) in Lake Izabal, Guatemala. Eucinostomus gracilis (Gill) Pacific coast from Baja California to the LUTJANIDAE Galapagos Islands (taken in fresh water in Lutjanus novemfasciatus Gill western Mexico). Pacific coast from Baja California to the Eucinostomus melanopterus (Bleeker) Galapagos Islands, entering tidal streams. Atlantic coast and tidal streams (occasion­ Lutjanus cyanopterus (Cuvier) ally in strictly fresh water) from Louisiana Atlantic coast of tropical America, entering and Texas to Brazil including the West brackish water. Indies; West Africa. Lutjanus griseus (Linnaeus) Gerres cinereus (Walbaum) Atlantic coast from Massachusetts to Brazil, Atlantic and Pacific slopes from Bermuda entering fresh water (Rio Champoton, and Florida to Venezuela and from Baja Yucatan, Hubbs, 1936:248). California to Peru and the Galapagos Lutjanus jocu (Bloch and Schneider) Islands, entering river mouths. Cape Cod (as strays) southward to Florida, Diaptems rhombeus (Cuvier) the West Indies, and mainland Middle West Indies and Atlantic slope from America to Brazil, entering brackish and Campeche, Mexico to Brazil, entering fresh water (Caldwell, et al., 1959:24). coastal lagoons. Lutjanus apodus (Walbaum) Diapterus evermanni Meek and Hildebrand Atlantic Ocean from Massachusetts to Atlantic coast from Veracruz, Mexico to Bahia, Brazil, entering brackish water. Venezuela, in coastal lagoons and river Lutjanus argentiventris (Peters) mouths.

149 798 COPEIA, 1966, NO.4

Diapterus axillaris Gunther to Brazil, taken in Rio Prinzapolca, Pacific coast from Mazatlan, l\Jexico to Nicaragua (Astorqui). Guatemala, entering coastal lagoons. 1\1 ieropogon turn ieri (Desmarest) Diapterus peruvianus (Cuvier) 'Vest Indies and Atlantic coast of Middle Pacific coast from Mazatlan, Mexico to America to northern South America, en­ Peru, entering tidal streams. tering fresh water (Rio Champot6n, Yuca­ Diapterus olithostomus (Goode and Bean) tan Penins:ula; Lake Izabal, Guatemala). Atlantic coast from Florida to Brazil, enter­ M ieropogon altipinnis Gunther ing rivers. Pacific coast from Guatemala to Ecuador, Diapterus lineatus (Humboldt) entering coastal lagoons. Pacific coast from Acapulco, Mexico to Aplodinotus grunniens Rafinesque Colombia, entering brackish lagoons. Lakes and rivers in the A tlan tic drainage Diapterus brevimanus (Gunther) from northeastern North America south­ Pacific coast from Guatemala to Panama, ward to the Rio Usumacinta basin, Guate­ entering coastal lagoons. (Chiapam, Guate­ mala. mala). Cynoseion arenarius Ginsburg Diapterus plumieri (Cuvier) Gulf coast of the United States and Mexico Atlantic coast from Florida to Panama, southward at least to Laguna de Terminos, en tering coastal lagoons. Campeche. Diapterus mexieanus (Steindachner) Restricted to fresh water on the Atlantic SPARIDAE slope of southern Mexico and northern Arehosargus aries (Valenciennes)? Guatemala (Rio Coatzacoalcos to Rio Usu­ Atlantic coast from British Honduras to macinta basins). Brazil, entering rivers and lakes (Lake Izabal, Guatemala). POMADASYIDAE

Pomadasys leueiseus (Gunther) MUGILIDAE Pacific coast from Mazatlan, Mexico to i\1ugil liza Valenciennes Peru, entering coastal lagoons. "Vest Indies and tropical Atlantic Amer­ Pomadas)'s bayanus Jordan and Evermann ican mainland southward to Brazil. Pacific coast from Baja California to Peru, lHugil eephalus Linnaeus entering rivers. Worldwide, entering rivers. Pomadas)ls maeraeanthus (Gunther) Mugil eurema Valenciennes Pacific coast from Mazatlan, Mexico to Both coasts of the Americas and Panama, entering coastal lagoons. ,,,Test Africa, entering rivers. Pomadasys emero (Cuvier) Atlantic coast of tropical America, entering Mugil triehodon Poey Atlantic coast from Florida to Brazil, en­ rivers. Pomadas),s branieki (Steindachner) tering rivers. Pacific coast from Mazatlan, Mexico to Afugil sp. Peru, entering rivers. Rio Coatzacoalcos near mouth, Veracruz, Pomadasys boueardi (Steindachner) Mexico. Atlantic coast of tropical America, entering Agonostomus monticola (Bancroft) rivers (Rio Papaloapam) and lakes (Lake Freshwater streams of both slopes of North Nicaragua). and Middle America and the West Indies, entering rivers, northward to Florida and SCIAENIDAE Sonora, Mexico. Bairdiella ronehus (Cuvier) Joturus pichardi Poey Atlantic coast from the West Indies and West Indies and Atlantic slope of Amer­ eastern Mexico to Brazil, entering coastal ican mainland from northern Veracruz, lagoons. l\Iexico to Rio Chagres, Panama, restricted Stelliter laneeolatus (Holbrook) to swiftly flowing fresh water. Chesapeake Bay, Maryland southward to Laguna de Terminos, Campeche, Mexico. DACTYLOSCOPIDAE Stelliter rastriter (Jordan and Evermann) Dactyloscopus thysanotus Bohlke Atlantic slope of southern Central America Pacific slope from southern Guatemala

50 MILLER-CENTRAL AMERICA1\' FISHES 799

(Miller and Briggs, 1962:9) to Chame Point, Euleptoeleotris clarki Hildebrand), enter­ Panama entering brackish and fresh water. ing fresh water. Erotelis smaragdus (Valenciennes) ELEOTRIDAE Florida, the West Indies, and Panama (as Gobiornorus dorrnitor Lacepede Euleptoeleotris shropshirei Hildebrand) Streams of the West Indies and of southern to Venezuela, entering brackish water. Florida and Texas southward to Surinam. Gobiornorus rnaeulatus (Gunther) GOBIIDAE Pacific streams from Rio Yaqui, Sonora, Bathygobius mystaeiurn Ginsburg Mexico southward to Peru. West Indies and vicinity of Veracruz City. Dorrnitator rnaeulatus (Bloch) Veracruz, Mexico southward to Colon, Pan­ Atlantic slope from North Carolina to ama, entering river mouths. Brazil, in sluggish streams. Evorthodus lyrieus (Girard) Dormitator latifrons (Richardson) Atlantic coast from Chesapeake Bay to Pacific slope from southern California to Surinam; West Indies; entering fresh Peru, in sluggish streams. water. Guavina guavina (Valenciennes) Gobionellus boleosorna (Jordan and Gilbert) Atlantic coast from Cuba and l\Iexico North Carolina to Venezuela (possibly to (near Veracruz City) to Rio de Janeiro, Brazil); West Indies; entering rivers. entering river mouths. Gobionellus shufeldti (Jordan and Eigen­ Guavina rnieropus Ginsburg mann) Miraflores Locks, Panama Canal. Gulf coast of the United States and north­ Leptophilypnus fluviatilis Meek and Hilde­ eastern Mexico, entering river mouths. brand Gobionellus daguae (Eigenmann) In fresh and brackish water of the Panama Pacific slope of Panama (Rio Tuira basin) Canal Zone (originally Atlantic only); also to Colombia. near Tortuguero, Costa Rica (Caldwell et Gobionellus claytonii (Meek) al., 1959:28, as Mieroeleotris mindii). Fresh and brackish waters along the Atlan­ Leptophilypnus panarnensis (Meek and Hil­ tic slope from northern Veracruz, ~Iexico debrand) to Venezuela, including Trinidad; 'Vest Streams of the Pacific slope of central Indies. Panama (Rio Chorrera, Rio Juan Diaz). Gobionellus hastatus Girard Leptophilypnus sp. Gulf coast of the United States and ~Jex­ Rapids of the Rio de la Pasion (Usumacinta ico, entering brackish water. basin), Peten, Guatemala. Gobionellus sagittula (Gunther) Leptophilypnus sp. Gulf of California to Peru, in brackish Rio Polochic, Atlantic slope of Guatemala. estuaries and lagoons. Eleotris pisonis (Gmelin) Gobionellus mierodon (Gilbert) Atlantic slope from Florida and Texas to Pacific coast from Sonora, Mexico to Pan­ Brazil, entering fresh water. ama, entering fresh water. Eleotris pieta Kner and Steindachner Awaous tajasiea (Lichtenstein) Pacific slope from the Colorado River, Atlantic coast streams from Florida and California to Peru, in lower stream courses. the Rio Grande to Bahia, Brazil; 'Vest Eleotris arnblyopsis (Cope) Indies. Atlantic coast and streams from South Awaous transandeanus (Gunther) Carolina to Surinam; 'Vest Indies. Pacific coast streams from Rio Yaqui, Eleotris isthmensis Meek and Hildebrand Sonora, Mexico to Peru. Atlantic slope of Panama, in or near Mierogobius rniraflorensis Gilbert and Starks brackish water. Tidal streams and coastal lagoons of the Hemieleotris latifaseiatus C~Ieek and Hilde­ Pacific slope from Laguna Coyuca, near brand) Acapulco, Mexico to central Panama Streams of the Pacific slope from Costa (Miller, 1958). Rica to southern Colombia. Garmannia hildebrandi Ginsburg Erotelis armiger (Jordan) Gatun Locks, Panama Canal. Baja California to Panama (includes Garrnannia homoehroma Ginsburg

151 800 COPEIA, 1966, NO.4

Both slopes of the Panama Canal, entering A.tlantic slope of Central America (Panama fresh water. to Nicaragua), entering fresh water (Dead­ Garmannia spes Ginsburg man's Creek, Bluefields, Nicaragua; Rivas). Atlantic slope from Costa Rica to Ven­ Batrachoides goldmani Evermann and Golds­ ezuela, entering coastal lagoons (Caldwell, borough et al., 1959:28). Restricted to the basin of the Rio Gobioides broussonneti Lacepede Usumacinta in Chiapas, Tabasco, and Atlantic coast from Florida to Brazil, enter­ Campeche, l\Iexico, and Peten, Guatemala. ing fresh water. Opsanus beta (Goode and Bean) Sicydium salvini Ogilvie-Grant Periphery of the Gulf of Mexico from the Streams of both slopes of central Panama Yucatan Peninsula (including Campeche southward to Colombia and Ecuador. Bank) through the Florida Keys and north­ Sicydium gymnogaster Ogilvie-Grant ward to at least Palm Beach, Florida Rivers of the Atlantic slope from l\Iisantla, (Walters and Robins, 1961:12); entering Veracruz, "Mexico southward at least to rivers. Honduras. Sicydium altum :Meek GOBIESOCIDAE Atlantic slope of Costa Rica. Gobiesox potamius Briggs Sic),dium mllltipunctatum Regan Rivers of the Pacific slope of Costa Rica. Pacific slope streams from near Mazatlan, Gobiesox nudus (Linnaeus) Mexico southward at least to the Rio Rivers of the Atlantic slope from Costa Choluteca, Honduras (Carr and Giovan­ Rica to Venezuela; West Indies. noli, 1950:35). TETRAODONTIDAE Sic),dillm pittieri Regan Streams of the Pacific slope of Costa Rica Sphoeroides testudineus (Linnaeus) and western Panama. Atlantic slope from Massachusetts to Brazil; West Indies; entering river mouths (Rio BROTULIDAE Champoton, Hubbs, 1936:248; Rio Prin­ Typhliasina pearsei (Hubbs) zapolca, Nicaragua; Astorqui) . Cenotes of the Yucatan Peninsula. Sphoeroides annulatus (J enyns) Pacific coast from Gulf of California to BOTHIDAE northern Peru and the Galapagos Islands; Citharichthys gilberti Jenkins and Evermann entering rivers (two records in Nicaragua Pacific coast from Guaymas, Sonora and by Rivas). Laguna Santa Maria, Baja California, Mexico southward to Peru, entering rivers. ACKNOWLEDGMENTS Citharichthys spilopterus Gunther In preparing this paper, much time and A tlan tic coast from New Jersey to Brazil. effort were devoted to compiling a distribu­ entering rivers southward. tional list of the families, genera, and species. I would like to acknowledge the considerable SOLEIDAE assistance received from Donn E. Rosen (for Achirus lineatus (Linnaeus) Guatemala), Padre Ignacio Astorqui. (Nica­ Florida and Texas to Brazil, sometimes ragua), William A. Bussing and Luis R. ascending river mouths. Rivas (Costa Rica), George S. Myers (Nica­ Achirus mazatlanus Steindachner ragua and Panama), and Horace Loftin Pacific coast from Sonora, Mexico to Peru, (Panama). Bruce B. Collette commented on occasionally entering fresh water but more the Hemiramphidae and Belonidae. Al­ frequently in brackish lagoons. though the distributional list is neither com­ Trinectes fonsecensis (Gunther) plete nor wholly accurate, it provides a re­ Pacific coast from Rio Yaqui, Sonora, liable general picture of the nature of the Mexico to Peru, entering fresh water. Middle American fish fauna and is intended Trinectes maculatus (Bloch and Schneider) to stimulate the appearance of additions and Atlantic coast from Massachusetts to Vene­ corrections. zuela, entering rivers southward. Support for field work is gratefully acknowl­ edged to the Horace H. Rackham School of BATRACHOIDIDAE Graduate Studies of The University of Michi­ Batrachoides gilberti Meek and Hildebrand gan and to the National Science Foundation

152 ~IILLER-CENTRAL AMERICA~ FISHES 801

(G-4854, 12904, 24129; GB-3271), which also book of Middle American Indians, vol. I. R. has partly subsidized publication costs. Paul Wauchope and R. C. West, eds., pp. 3-32. Univ. Texas Press, Austin, Texas. M. Earl drafted Fig. 1 and John Tottenham MARSHALL, N. B. 1965. The life of fishes. prepared Fig. 2. Weidenfeld and Nicolson, London, England. ~fEEK, S. E. 1907. Synopsis of the fishes of the LITERATURE CITED Great Lakes of Nicaragua. Field Colum. Mus. BEHRE, E. H. 1928. A list of the fresh water Pub. 121, Zoo I. Ser. 7(4):97-132. fishes of western Panama between long. 81 0 1908. Notes on fresh-water fishes from 45' and 83 0 15' W. Ann. Carnegie Mus. 18(2): Mexico and Central America. Ibid. 124, Zool. 305-328. Ser. 7(5):133-157 (Oct. 1907-Jan. 8, 1908). BOHLKE, J. E. 1958. Studies on fishes of the --- A:-JD S. F. HILDEBRAND. 1923. The marine family Characidae. No. 16-A new Hyphes­ fishes of Panama. Part I. Zool. Ser., Field Mus. SOb1)lCOn from Costa Rica. Bull. Fla. State l\iat. Hist. 15:1-330. Mus. BioI. Sci. 3(4):173-178. MILLER, R. R. 1955. A systematic review of the BREDER, C. M., JR. 1927. The fishes of the Rio Middle American fishes of the genus Pro­ Chucunaque drainage, eastern Panama. Bull. fundulus. Misc. Pub. ~Ius. Zool., Univ. Mich. Am. Mus. Nat. Hist. 57:91-176. Ko. 92, 64 pp. BUSSING, "\V. A. 1963. A new poeciliid fish, ---. 1957. Fishes from Laguna Octotal. In: Phalliehthys tieo, from Costa Rica. Los Angeles Biological investigations in the Selva Lacan­ Co. Mus. Contrib. Sci. 77:1-13. dona, Chiapas, Mexico. R. A. Paynter, Jr., CALDWELL, D. K., L. H. OGREN, AND L. GIOVAN:\,OLl. (ed.). Bull. Mus. Camp. Zool., Harvard II6(4): 1959. Systematic and ecological notes on some 238-241. fishes collected in the vicinity of Tortuguero, ---. 1958. A Panamanian gobiid fish, Mi­ Caribbean coast of Costa Rica. Rev. BioI. erogobius miraflorensis, from western Mexico, Trap. 7(1):7-33. with taxonomic and ecologic notes. Copeia CARR, A. F., JR. AND L. GIOVA:-Jl'."OLI. 1950. The 1958(3) :235-237. fishes of the Choluteca drainage of southern ---. 1959. Origin and affinities of the fresh­ Honduras. Occ. Pap. Mus. ZooI., Univ. Mich. water fish fauna of western North America. No. 523, 38 pp. In: Zoogeography, C. L. Hubbs, ed., pp. 187- CHAVEZ, U. 1961. Estudio de una nueva especie 222. Am. Assoc. Adv. Sci. Symp. Vol. 51. de robalo del Golfo de Mexico y redescripci6n --- and J. C. Briggs. 1962. Daetyloseopus de Centropomus undecimalis (Bloch). Ciencia amnis, a new sand stargazer from rivers of the 21(2):75-83. Pacific slope of southern ~fexico. Oce. Pap. DARNELL, R. M. 1962. Fishes of the Rio Tamesi Mus. Zool., Univ. Mich. No. 627, II pp. and related coastal lagoons in east-central --- \ND B. C. NELSON. 1961. Variation, life Mexico. Pub. Inst. Mar. Sci., Univ. Texas colors, and ecology of Ciehlasoma eallolepis, a 8:299-365. cichlid fish from southern Mexico, with a dis­ GOSLINE, W. A. 1947. Contributions to the cussion of the Thoriehthys species group. Occ. classification of the loricariid catfishes. Arq. Pap. Mus. Zool., Univ. Mich. No. 622, 9 pp. Mus. Nac. (Brasil) 41 :79-144. MYERS, G. S. 1966. Derivation of the freshwater HILDEBRAND, S. F. 1925. Fishes of the Republic fish fauna of Central America. Copeia 1966 of El Salvador. Bull. U. S. Bur. Fish. 41 :238- (4):766-773. 287. REGAN, C. T. 1905. A revision of the fishes of 1938. A new catalogue of the fresh­ the American cichlid genus Ciehlasoma and of water fishes of Panama. ZooI. Ser., Field Mus. the allied genera. Ann. Mag. Nat. Hist., Ser. Nat. Hist. 22(4):219-359. 7, 16:60-77, 225-243, 316-340, 433-445. 1946. A list of fresh-water fishes from 1906-08. Pisces. In: Biologia Centrali­ San Jose Island, Pearl islands, Panama. Smith. Americana 8:1-203. Misc. Call. 106(3): 1-3. RIVAS, L. R. 1962. Ciehlasoma pasion is, a new HUBBS, C. L. 1935. Fresh-water fishes collected species of cichlid fish of the Thoriehthys group, in British Honduras and Guatemala. Misc. from the Rio de la Pasion, Guatemala. Quart. Pub. Mus. ZooI., Univ. Mich. No. 28, 22 pp. J. Fla. Acad. Sci. 25(2):147-156. 1936. Fishes of the Yucatan Peninsula. ---. 1963. Subgenera and species groups in Carnegie Inst. Wash. Pub. 457:157-287. --_.. 1950. Studies of cyprinodont fishes. the poeciliid fish genus Gambusia Poey. Copeia XX. A new subfamily from Guatemala, with 1963(2) :331-347. ctenoid scales and a unilateral pectoral clasper. 1965. Review of: The poeciliid fishes Misc. Pub. Mus. Zool., Univ. Mich. No. 78, 28 (Cyprinodontiformes), their structure, zoogeog­ pp. raphy, and systematics. By D. E. Rosen and 1953. Geographic and systematic status R. M. Bailey. Copeia 1965(1):1l7-118. of the fishes described by Kner and Steindachner ROSEN, D. E. AND R. M. BAILEY. 1963. The in 1863 and 1865 from fresh waters in Panama poeciliid fishes (Cyprinodontiformes), their and Ecuador. Copeia 1953(3):141-148. structure, zoogeography, and systematics. Bull. --- AND R. R. MILLER. 1960. Potamarius, a Am. Mus. Nat. Hist. 126(1):1-176. new genus of ariid catfishes from the fresh RYAN, R. M. 1963. The biotic provinces of waters of Middle America. Copeia 1960(2): Central America. Acta Zool. Mex. 6(2-3): I-55. 101-II2. SCHULTZ, L. P. 1944a. The catfishes of Vene­ MALDONADO-KoERDELL, M. 1964. Geohistoryand zuela, with descriptions of thirty-eight new paleogeography of Middle America. In: Hand- forms. Proe. U. S. Xat. Mus. 94:173-338.

153 802 COPEIA, 1966, ~O. 4

1944b. A new genus and species of toadfish (Batrachoididae) considered to be a pimelodid catfish from Colombia. J. ·Wash. glacial relict in the West Indies. Am. Mus. Acad. Sci. 34(3):93-95. Noyit. No. 2047, 24 pp. SCHWWTZ, F. J. 1964. 1\' atural salinity toler­ 'VEST, R. C. 1964. Surface configuration and ances of some freshwater fishes. Underwater associated geology of Middle America. In: ~atur. 2(2):13-15. Handbook of Middle American Indians, vol. 1. STUART, L. C. 1964. Fauna of ~Iiddle America. R. Wauchope and R. C. West, eds., pp. 33-83. In: Handbook of Middle American Indians, Univ. Texas Press, Austin, Texas. vol. 1. R. Wauchope and R. C. West, eds., pp. 316-362. Vniy. Texas Press, Austin, Texas. ~luSEu::\f OF ZOOLOGY, THE UNIVERSITY OF "-\LTERS, Y ..\:\D C. R. ROBI:\S. 1961. A new ~IICHIGAK) ANN ARBOR) MICHIGAN 48104.

ADDE:\TDL'M

During the eight years since this list was Roeboides salvadoris is a synonym of R. published, the following major changes and guatemalensis (Miller and Carr, 1974, additions have been noted by me or others. Copeia: 121). P. 785. Compsura and Pseudocheirodon were both Pterobrycon myrnae Bussing, 1974 (Rev. Bio. synonymized with Cheirodon by Fink and Trop. 22(1): 138-144, figs. 1-3). Known Weitzman (1974, Smith. Contrib. Zool. only from the Pacific slope of southern 172); hence C. gorgonae becomes Cheirodon Costa Rica. Insert at bottom, p. 785, col. 1. gorgonae Evermann and Goldsborough, P. Piabucina boruca Bussing, 1967 (op. cit.). Rio affinis becomes Cheirodon affinis (Meek and Grande de Terraba and Rio Coto basins, Hildebrand), and P. terrabae (see below) Pacific slope of Costa Rica. Add to p. 786. becomes Cheirodon terrabae (Bussing). P. Gymnotus campo is unknown from Costa Rica 784. (Bussing, 1967, op. cit.). P. 786. Pseudocheirodon sp. = P. terrabae Bussing, Gymnotus cylindric us occurs also in Costa Rica 1967 (Rev. BioI. Trop. 14 (2); 1966). (Bussing, 1967, op. cit.). P. 786. Known only from Rio Grande de Terraba Imparales sp. = I. panamensis Bussing, 1970 basin, Pacific slope of Costa Rica. P. 784. (L. A. Co. Mus. Contrib. Sci., 196). Pacific Cheirodon dialepturus Fink and Weitzman (op. and Atlantic (Rio Chagres) slopes of cit.). Pacific slope of Panama and Costa Panama. P. 786. Rica, from Code Province to Rio Coto. Rhamdia heteracaniha is also on Pacific slope Add to p. 784. of Costa Rica (Bussing, 1967, op. cit.). P. Cheirodon mitopterus Fink and Weitzman (op. 787. cit.). Atlantic slope of central Panama (Rio Rhamdia nasuta is also on Atlantic slope of Code del ~orte basin). Add to p. 784. Costa Rica (Bussing, 1967, op. cit.). P. 787. Bramocharax sp. = Bramocharax dorioni Rosen Nannorhamdia lineata Bussing, 1970 (op. cit.). (as B. bransfordi dorioni, 1970, Am. Mus. Pacific slope of Costa Rica (Puntarenas Novit. 2435). Upper tributaries of Rio Prov.). Add to p. 787, col. 1. U sumacinta basin in Alta Verapaz, Rivulus myersi is probably a synonym of R. Guatemala. (Now regarded as a full marmoratus Poey; that species is known in species, Rosen, pers. comm., 1974.) P. 785. Middle America from Mexico and Belize Bramocharax baileyi Rosen, 1972 (Am. Mus. (material at USNM and UMMZ). P. 788. ~ovit. 2500). Upper reaches of Rio de la Rivulus godmani is a synonym of Rivulus Pasion and Rio Salinas, Alta Verapaz, tenuis (Miller and Carr, op. cit.). P. 788. Guatemala. Add to p. 785. Rivulus sp. = R. robustus Miller and Hubbs, Hyphessobrycon sp. = H. savagei Bussing, 1967 1974 (Copeia: 865-869). P. 788. (op. cit.). Known only from Rio Grande de Xiphophorus signum Rosen and Kallman (as X. T erraba basin, Pacific slope of Costa Rica. helleri signum, 1969, Am. Mus. N ovit. P.785. 2379). Rio Chajmaic, Rio Usumacinta Rhoadsia eigemnanni is now called Carlana basin, Alta Verapaz, Guatemala. (Now re­ eigenmanni (Fink and Weitzman, op. cit.). P. garded as a full species, Rosen, pers. 785. comm., 1974.) Add to p. 789.

154 Genus and species (Cnesterodontinae) Costa Rica (Bussing, 1967, op. cit.). P. 792. Rosen = Scolichthys iota Rosen and S. Cichlasoma guij"a, regarded as perhaps only a greenwayi Rosen (1967, Am. Mus. Novit. race of C. macracanthum by Kahsbauer 2303). Both inhabit upper tributaries of (1972, Ann. ;..Jaturhistor. ~lus. Wien, 72: Rio Usumacinta in Alta Verapaz, 165), is herewith synonymized with that Guatemala. P. 789. species since comparison of the holotype Brachyrhaphis sp. = B. holdridgei Bussing, (CS~~1 87301, C. meeki) with C. macracan­ 1967 (op. cit.). Atlantic slope of Costa Rica. thum fails to support specific separation. P. P. 790, col. 1, line 8 down. 792. Poeciliopsis sp. = P. paucimaculata Bussing, Cichlasoma heterodontum is a synonym of C. 1967 (op. cit.). P. 791. macracanthum (~liller, ~IS). P. 792. Phallichth)1s fairweatheri Rosen and Bailey. Cichlasoma bulleri. Range should read Range includes the lower Rio L'sumacinta "C pper tributaries of the Rio Coatzacoal­ basin (Chiapas-Tabasco, L'MMZ 196422) cos basin, Oaxaca, ~lexico." P. 792. and Caobas Laguna (Campeche-Quintana Cichlasoma atromaculatum Regan. Atlantic Roo, UMMZ 196466), Mexico. P. 791. slope of Panama (San BIas Prov., material Cichlasoma maculicauda. Although Rivas in US~~I) east to Atrato and San Juan ba­ (1962: 154) recorded this species from the sins, Colombia. Insert under Parapetenia, U sumacinta basin, Guatemala, examina­ p.792. tion of his specimen (now number 5430, Cichlasoma managuense. Add to range "into Gulf Coast Res. Lab.) shows it to be a eastern Honduras" (material at CM~IZ). juvenile of C. heterospilum Hubbs. An adult P.793. male (GCRL 5698) from 36 km WNW of Cichlasoma dowi. Add to range "northward Sayaxche, El Peten, Guatemala, also rep­ into Honduras" (Miller and Carr, op. cit.). resents this species. One distinguishing P.793. character is the length of the pectoral fin, Cich/asoma motaguense. Also Atlantic slope of the tip of which extends to the vent in Costa Rica (Bussing, 1967, op. cit.). P. 793. maculicauda but to beyond the anal origin Cichlasoma sp. = C. mento (Vaillant and Pel­ in heterospilum. C. maculicauda is known legrin). P. 793, col. 1, line 21 up. northward to the basin of Golden Stream, Cichlasoma affine. For "Rio Grijalva basin, southern Belize (UMMZ 195944). P. 791. Mexico" substitute "Laguna Bacalar, Cichlasoma lyonsi Gosse, 1966 (Bull. Inst. R. Quintana Roo, Mexico" (material at Sci. ;..Jat. Belg. 42: 16). Tributary to Rio U~IMZ). P. 793. Coto, Pacific slope of Costa Rica (Punt­ Cichlasoma sp. = C. regani Miller, 1974 (Proc. arenas Prov.). Insert after C. heterospilum, BioI. Soc. Wash., 87(40):465--472). P. 793, p. 791. col. 2, line 14 up. Cichlasoma nicaraguense (Gunther). For ac­ Chanos chanos. Unpublished record for count of variation, coloration, and sys­ Corinto, northwest coast of ~icaragua, tematic status. see Lopez, 1974 (Rev. BioI. Meek, 1906 (FMNH 5830). P. 794. Trop. 22(1): 161-185, figs. 1-11). P. 791. Anchoa ca~orum (Fowler). Recorded from the Cichlasoma sajica Bussing, 1974 (Rev. BioI. fresh ~aters of Belize bv Thomerson and Trop. 22(1)30--37, fig. 1). Pacific slope of Greenfield (1975, Cope{a: 50--52). P. 795. southern Costa Rica from just south of A nchoviella belizensis Thomerson and Green­ Punta Mala to the Rio Esquinas basin. In­ field, 1975 (op. cit.). Lowland fresh waters sert after C. nigrofasciatum, p. 792. of Belize. P. 795. Cichlasoma diquis Bussing, 1974 (op. Arius melanopus as listed is a complex (W. R. cit. :37-44, fig. 2). Pacific slope of southern Taylor, pers. comm., 1974). A. aguadulce Costa Rica from just south of Punta Mala (Meek), Rio Papaloapan basin, Veracruz, to the Rio Coto drainage. Insert after C. ~fexico, southeastward to Rio Polochic alfaroi, p. 792. basin, Guatemala. A. melanopus (Gunther), Cichlasoma spilurum. Range should read "At­ Rio ~lotagua basin, Guatemala, into lantic slope from eastern Campeche, northwestern Honduras. P. 795. Mexico, southward to .... " (material at Strongylura timucu (Walbaum). Atlantic VMMZ and Gulf Coast Res. Lab.). P. 792. slope, from Florida and Veracruz, Mexico Cichlasoma robertsoni. Range emended and (F~I~H 4568), southward and eastward extended to eastern Honduras (Miller and through the Antilles to Brazil, entering Carr, op. cit.). P. 792 rivers (Collette, 1968, Copeia: 190). Add Cichlasoma longimanus. Also Pacific slope of to p. 796.

155 Strongylura hubbsi Collette, 1974 (Copeia: thority of Gilbert and Kelso (op. cit.). Add 612). Atlantic slope streams from basins of to p. 798, before Dactyloscopidae. Rio Papaloapan, ,\1exico, to Rio Bathygobius sop orator (Valenciennes). Florida U sumacinta, Guatemala. Add to p. 796. and Gulf Coast southward to Rio de Melaniris meeki. Confined to' Rio Motagua Janeiro; recorded from fresh water in basin, Guatemala (Miller and Carr, op. Costa Rica (Gilbert and Kelso, op. cit.). cit.). P. 796. Add to p. 799. i'Welaniris alvarezi Diaz-Pardo, 1972 (Anal. Gobionellus clay toni is restricted to Gulf of Esc. Nac. BioI. 19 (1-4): 145-152). Basin Mexico drainage in ,\1exico (see next of lower Rio Grijalva, Tabasco, ,\1exico. entry). P. 799. Add to p. 796. Gobionellus fasciatus (Gill). Southern Carib­ Centropomus poe), Chavez. Recorded from Be­ bean basin from Costa Rica to Dominica, lize by Greenfield (1975, Copeia: in press). entering fresh water (Gilbert and Kelso, P.797. op. cit.). Add to p. 799. Oligoplites palometa (Cuvier). Atlantic slope of Gobionellus pseudofasciatus Gilbert and Ran­ Central America to northern South dall, 1971 (in Gilbert and Kelso, op. cit.). America, entering fresh water (e.g., Lake Atlantic slope of Belize to Panama and Yzabal, Guatemala-Regan, 1906, BioI. Trinidad (see Thomerson and Greenfield, Centrali-Amer., 8: 15). Add to p. 797. op. cit.). Add to p. 799. Diapterus olisthostomus (Goode and Bean) is a Citharichthys uhleri Jordan. West Indies and synonym of D. auratus Ranzani (Deckert, Atlantic coast of ,\1iddle America where it in press, Bull. So. Calif. Acad. Sci.); D. enters rivers (as in Mexico, UMMZ evermanni Meek and Hildebrand is the 184533, and Costa Rica-Gilbert and juvenile of D. olisthostomus (C. L. Hubbs, Kelso, op. cit.). Add to p. 800. pers. comm., 1967). Pp. 797-798. Trinectes paulistanus ,\1iranda-Ribeiro. South­ Diapterus (or Eugerres) brasilianus eastern Yucatan, Mexico, to Brazil, enter­ (Valenciennes). Recorded from the fresh ing fresh water (Gilbert and Kelso, op. cit.; waters of Belize by Thomerson and USNM 114281, Rio Sauce, L. Yzabal, Greenfield (1975, op. cit.). P. 798. Guatemala). Add to p. 800. Larimus breviceps Cuvier. West Indies to Batrachoides gilberti Meek and Hildebrand. Brazil (see Gilbert and Kelso, 1971, Bull. Recorded from fresh water in Belize by Fla. St. Mus. 16). Add to p. 798. Greenfield and Greenfield (1973, Copeia: l'Wenticirrhus americanus (Linnaeus). Atlantic 564). P. 800. slope from Chesapeake Bay southward to Costa Rica, entering lagoons (Gilbert and Kelso, op. cit.). Add to p. 798. I am grateful to William A. Bussing and Echeneis naucrates Linnaeus. The addition of David W. Greenfield for providing a num­ this marine fish (Echeneidae) is on the au- ber of new records.

156