(z- C /2e(x-i.,-;-? Fr Science of Biology Journal )."-e a 4 June, 1975 Vol. 1 No 2 R72—vz. c / -7;e4/- Copyright 1975 Shawky Farag The Distribution of the Oviparous Killifishes of

Kenneth carpio and Yucatan populations to F. C. barbouri and F. c. Relyea - Associate Professor of Biology polyommus. In Florida, this species ranges from Cape San T3la1, Jacksonville University Gulf Co. (one record, Kaill, 1967), on Florida's gulf coast southward Jacksonville, Florida 32211 to the Florida Keys and Miami, and northward toLake Worth, Palm Beach Co. (JU 1338) on the Atlantic coast. The species is ° The State of Florida has a remarkably rich representation of egg- common, however, only from about 29 N. Lat., about Cedar Key, laying killifishes of the family Cyprinodontidae. The Fun- southward on the Gulf coast. The species then occurs disjunctly to dulus, with from 10 to 12 species (dependent on one's taxonomic the north on the Atlantic coast from Vero Beach, Co. point of view), is especially well represented. This represents (Kaill, op. cit.) northward to the southern end of , nearly one-half of the species known from the United Volusia Co. (JU 1298). The northern and southern ends of the States. Two other species of Fundulus occur in Yucatan, and these Atlantic disjunct population may not be breeding populations, but are related to, and possibly derived from, Florida forms. In all, 19 only juveniles (Relyea, pers. obs., and Kaill, op. cit., Vero Beach). to 21 species of oviparous killies occur in Florida, and 3 of these are The main populations on the Atlantic coast are in the Titusville and endemic to the state. The total number of killie species for the Melbourne areas (about 29° N. Lat.). Records from Florida's United States is 46 (10 genera, 8 of which occur in Florida). northern Gulf coast (Fowler, 1917:39, Carrabelle; Goode and Bean, This paper will denote the known geographic distribution of 1882:433, Pensacola) under the name Cyprinodon mydrus are Florida killies, and should be useful to students of the difficult, perhaps impossible, to verify due to erroneous labeling zoogeography, ecology, and of Florida fishes. For and possible mixing of collections in the USNM. Miller (1974) has discussions of Cyprinodont genera the ready is referred to Myers made an effort to rectify some of these problems. The species does (1931), Miller (1955a), and Rosen (1964, 1971). Miller (1955b) and not appear to occur in the Carrabelle or Pensacola areas now. Brown (1957) have presented general reviews of the genus Fun- Gunter's (1941:204) and Garman's (1895:27) records of F. carpio dulus, and Moore (1968) provided a complete key to the North from Texas are probably in error and based on C. variegatus. American oviparous killifishes. Fundulus chrysotus (Holbrook) Adinia xenica (Jordan and Gilbert) The golden topminnow occurs in freshwaters throughout Florida. The occurs in protected brackish waters from Its total range is from South Carolina to the southern tip of Florida the Florida Keys (Jacksonville University Collections from Big (but not the Keys) and westward along the Gulf coast to eastern Pine Key, Monroe County, JU 937 and 1034, and Hastings and Texas, and northward to southern Missouri and Tennessee (Miller, Yerger, 1971) to Texas along the coast. The western 1955b; Brown, 1957). This species may be distinguished from the limits of this species along the Texas coast are not known similar F. cingulatus. by its pointed gill rakers (blunt in F. adequately (given as San Patricio Co. by Hastings and Yerger, op. cingulatus). cit. ). Apparently Adinia does not occur north of the Miami area on Fundulus cingulatus (Valenciennes) the Atlantic coast of Florida (JU 870, Dade Co.). The banded topminnow is a freshwater species that ranges from Cyprinodon hubbsi (Carr) southern Georgia southward to at least the in south The Lake Eustis is endemic to at least 6 large (Brown, 1956), and westward through the Florida Florida freshwater lakes (Eustis, JU 1342, and original description, panhandle to the Escambia River (Bailey, Winn, and Smith, 1954). Carr, 1936; Harris, JU 1363; Dora, JU 199 and 1356; Weir, JU 1349; It is certainly not a continuously distributed form, however, and is Griffin, JU 1339; and Yale, JU 1355). Its absence from other similar not especially common at the extremes of its range. This may be lakes such as Lake Placid and Big is an the least studied of Florida freshwater killies. enigma. Pupfish from are probably C. Fundulus confluentus (Goode and Bean) The marsh killifish, a brackish and freshwater species, occurs variegatus. throughout the state of Florida, including the Florida Keys. It is Cyprinodon variegatus (LacOde) more common in freshwater in the southern half of the peninsula. The sheepshead pupfish ranges continuously from The species ranges from Chesapeake Bay (Hildebrand and Massachusetts to Texas, and occurs in both fresh and brackish Schroeder, 1928) southward throughout Florida and westward situations in Florida. The pupfish in Lake Okeechobee is probably along the Gulf of Mexico coast. Bailey, Winn, and Smith (1954) this species, as are pupfish in freshwater situations in the Florida defined the western limit as Baldwin Co., Alabama, with F. con- Keys and (JU 1334, Okeechobee; JU 941, 1031, 1035, Big fluentus being replaced westward from there by F. pulvereus. Pine Key; JU 272, 397, 439, 561, Everglades localities). A disjunct Relyea (1965) regarded F. pulvereus. as conspecific with F. con- subspecies, C. v. artifrons, Hubbs (1936), is known from Yucatan. fluentus, hence F. confluentus has a continuous distribution inthe Populations in the Florida Keys and Cuba have been recognized as Gulf of Mexico To at least the Laguna Madre of Texas (Breuer, another subspecies, C. v. riverendi (Jordan and Evermann, 1896). 1962). For a discussion of West Indian and Bahamian populations of Fundulus grandis (Baird and Girard) Cyprinodon (several species) see Hubbs and Miller (1942), and for The gulf killifish, contrary to the common name, is not restricted western U. S. species, Miller (1948). Ager (1971) and Kushlan and to the Gulf of Mexico. It does occur continuously, ot nearly so, Lodge (1974) briefly discussed Lake Okeechobee and south Florida in the Gulf from Laguna de Tamiahua, Mexico (Miller, 1955b) populations, respectively. •through the northern Gulf and southward to about Floridichthys carpio (Gunther) Marco, Collier Co., Florida (southwestern Florida coast). There The gold-spotted killifish occurs both in Florida and Yucatan, seem to be no records between Marco and Cape Sable (Florida with Florida populations being relegated to the subspecies F. c. Bay, Monroe Co.) (Relyea, 1967). In Florida Bay and the Florida

49 Keys, populations are probably referable to F. E. saguanus (Cuban fja_lamitdu (Agassiz) subspecies, Rivas, 1948). Relyea (1967) suggested that Florida Bay and and Keys populations, and Cuban populations, be recognized as F. Fundulus lineolatus (Agassiz) saguanus. The distribution of this latter form is from Boca Raton The star head topminnows present another complex pattern of ( two known specimens, both juveniles, JU 951) southward through geographic variation. Brown (1958) considered this species com- the Florida Keys to the Tortugas (United States National Museum plex to consist of three subspecies: 1. Fundulus notti lineolatus, Coll., USNM 116897, 2 specimens) and Cuba (Rivas, 1948). On the ranging from Virginia to central Florida and westward to the Atlantic coast, another population of Fundulus grandis occurs in Florida and Georgia. I have specimens of this disjunctly from Gulf and south Florida populations just described. form southward to Lake Chekika, in southern Dade Co., near This Atlantic population occurs from Lake Worth (Univ. of Florida Homestead ( JU 392). It is not known from the Florida Keys. 2. F. n. Coll., UF 1490 and 4662) northward to Amelia Island, Nassau Co. notti ranging from the New River ( intergrades with E. n. (JU 36, 44, 111, and 113). The distribution of Fundulus saguanus lineolatus in the which connects the Ochlockonee between disjunct Gulf and Atlantic F. grandis populations probably and New Rivers) westward to eastern Texas. 3. F. n. dispar, reflects recent splitting of F. grandis populations with recession of ranging throughout the Mississippi Valley northward. Rivas (1966) the Wisconsin glacier. Fundulus saguanus would then represent a awarded specific rank to F. lineolatus. Moore (1968:112-113) gave a glacial relict, in the sense of Walters and Robins (1961), which is general review of the issue. Further taxonomic and distributional now allopatric with respect to F. grandis. The closely related study of this strictly freshwater species is warranted. F'unduhis grandissimus of Yucatan may be another glacial relict in Fundulus olivaceus (Storer) this species complex. The black spotted topminnow is a stream dwelling counterpart of Fundulus heteroclitus (Linnaeus) the pond dwelling F. notti complex. It ranges throughout the middle The mummichog is a temperate species related to F. grandis. It and lower Mississippi Valley southward to the Gulf of Mexico. ranges from Canada (Brown, 1957) southward to Brevard Co., Along the Gulf it ranges from eastern Texas to the Choctawhatchee Florida (3.5 mi. S of Edgewater, JU 876). Fundulus heteroclitus River in the Florida panhandle ( Thomerson, 1966). Since this overlaps the distribution of F. grandis in northeastern Florida species occurs in the in Georgia,it should be between Amelia Island, Nassau Co., and Brevard Co. Hybrids looked for in Florida portions of that river system (Thomerson, op. should be looked for in collections from this area. cit.) Fundulus 'enkinsi (Evermann ) Fundulus seminolis (Girard) The topminnow is a rarely encountered fish in The killifish is an essentially freshwater species that is Florida. The few records are from extreme western Florida restricted to Florida. It ranges throughout the peninsula westward ( Escambia River, Bailey, Winn, and Smith, 1954; a specimen from to Carrabelle, Franklin Co. ( Tulane Univ. Coll., TU 42447; Florida Perdido Bay in the Florida State Univ. Coll.; Pensacola Bay State Univ. Coll., FSU 11944,12059,15669). One specimen from Deer specimens in the Tulane Univ. Coll.). This is a brackish water Point Lake, Bay Co., represents the western known specimen, and species, and is not common throughout its range from Texas may be a bait introduction (the species is probably established in (Galveston? ) to western Florida. the lake) (F'SU 14680). Tabb and Manning (1961) recorded this Fundulus majalis (Walbaum) species from the southern tip of Florida. Although the species is and common in the , no specimens are known from Fundulus similis (Baird and Girard) Georgia tributaries of that drainage. The striped killifish (F. majalis) and long nose killifish (F. Jordanella floridae (Goode and Bean) similis ) will be considered together since some authors consider The common Florida flagfish is endemic to the peninsula of them to be conspecific ( Relyea, 1967; a position shared by Griffith, Florida. It ranges in freshwater from the southern tip of Florida 1974, and not disputed by Chen's, 1971, karyological studies). Taken (no records from the Florida Keys) northward and westward to the together, these forms range from New Hampshire (Jackson, 1953) Ochlockonee River (personal observation). southward to and through the Florida Keys (no records from the Leptolucania ommata (Jordan) Tortugas or Cuba) and westward in the Gulf of Mexico, presumably The ocellated killifish occurs throughout Florida (except the continuously, to a lagoon 35 miles north of Tampico, Mexico Keys) in freshwater circumstances. It is also known from southern ( Miller, 1955b). The related F. persimilis is restricted to Yucatan. Georgia and Alabama (Bailey, Winn, and Smith, 1954). Its western The overall distribution is similar to that for the F. grandis-F. limit is probably the Escambia River. heteroclitus species complex discussed earlier. As with F. grandis, goodei (Jordan) there are no records between Marco and Florida Bay on the Gulf The blue fin killifish occurs in south Georgia and throughout the coast, and no records between Lake Worth (JU 1337) and the Miami Florida peninsula (Moore, 1968). Its western limit in the Florida area on the Atlantic coast. Florida Keys populations may be panhandle appears to be the (Smith-Vaniz, allopatric and worthy of nomenclatural recognition (see Relyea, 1968), but its distribution is spotty west of the . It is 1967). The majalis form grades into the more southern similis form common in the Dead Lakes ( drainage). This is between Matanzas Inlet and Flagler Beach (northeastern Florida), basically a freshwater species. in the same general area of overlap between F. grandis and F. Lucania parvn ( Baird) heteroclitus. Geological phenomena have probably influenced the The rainwater killifish is a brackish, occasionally freshwater, distribution of both of these brackish water species complexes in species which ranges from Massachusetts to northeastern Mexico similar ways. ( Hubbs and Miller, 1965). This distribution is probably not con-

50 tinuous, especially in Florida where a distribution similar to that of 8. Carr, A. F. 1936. A new species of Cyprinodon from Lake Eustis, Fundulus grandis is apparent. Hubbs and Miller (op. cit.) Florida. Copeia 1936(3 ) :160-163. examined no specimens from coastal areas between the mouth of 9. Chen, T. R. 1971. A comparative chromosome study of twenty the St. Johns River and the upper Keys, a distance of over 300 killifish species of the genus Fundulus (Teleostei:Cyprinodon- miles. Specimens do occur in that region. My collections indicate a tidae). Chromosoma 32:436-453. reasonably continuous distribution southward through Brevard 10. Fowler, H. W. 1917. Cold blooded vertebrates from Florida. County. I have been unable to collect the species between there and Copeia 43:38-39. the upper Keys, although based on the distribution of other killies I 11. Garman, S. 1895. The Cyprinodontes. Mem. Mus. Comp. Zool. suspect that it may occur southward to near Lake Worth. Florida 19 : 1-179. Keys populations may be allopatric with respect to both Atlantic 12. Goode, G. B. and T. H. Bean. 1882. Descriptions of twenty-five and Gulf populations of L. parva, and would then represent glacial new species of fish from the southern United States, and three new relicts. genera, Letharcus, Ioglossus, and Chriodorus. Proc. U.S.N.M. Luçnia Qua also occurs as possibly isolated populations in two 5:412-437. Florida springiin the St. Johns River drainage, Alexander- Springs 13. Griffith, R. W. 1974. Environment and salinity tolerance in the (JU 1326) and Juniper Springs (JU 764). Although most abundant at genus F'undulus. Copeia 1974(2) :319-331. the spring "boils", specimens can be found in the spring "runs" 14. Gunter, G. 1941. Death of fishes due to cold on the Texas coast, downstream. Systematic and ecological studies of this species in January 1940. Ecol. 22( 2) : 202-208. Florida are warranted. 15. Hastings, R. W. and R. W. Yerger. 1971. Ecology and life history Rivulus marmoratus ( Poey ) of the diamond killifish, Adinia xenica (Jordan and Gilbert). The rivulus is probably the rarest Florida killifish, although an Amer. Midl. Nat. 86(2) :276-291. extensive population may occur in the Everglades in salt in- 16. Hildebrand, S. F. and W. C. Schroeder. 1928. Fishes of fluenced areas ( Tabb and Manning, 1961). The species .occurs in Chesapeake Bay. Bull. U.S. Bur. Fish. 43(1) :1-388. Cuba and the Bahamas ( Bbhlke and Chaplin, 1968). Its full Florida 17. Hubbs, C. L. Fishes of the Yucatan peninsula. Cam. Inst. distribution is uncertain. Wash. Publ. 457:157-287. 18. and R. R. Miller. 1942. Studies of the fishes of the order Cyprinodontes. XVIII. Cyprinodon laciniatus, a new species from the Bahamas. ACKNOWLEDGEMENTS Occ. Pap. Mus. Zool. Univ. Mich. 458 : 1-11. 19. 1965. Studies of the cyprinodont fishes. IOCH. Variation in I extend thanks to the curators of several fish collections that I Lucania parva, its .establishment in western United have made use of in accumulating distributional records, but States, and description of a new species from an interior basin in Coahuila, Mexico. Misc. especially Dr. Carter R. Gilbert, Florida State Museum, University Publ. Mus. Zool. Univ. Mich. 1271-104. 20. Jackson, C. F. of Florida; Dr. Ralph W. Yerger, Florida State University; and Dr. 1953. Northward occurrence of southern fishes (Fundulus, Mugil, Pomatomus) Royal D. Suttkus, Tulane University. I also wish to thank in coastal waters of New Hamp- shire. Copeia 1953(3):192. Jacksonville University students HarrySahlman, Bruce Sutton and 21. Jordan, D. S. and B. W. Julian Humphries for assistance in the field. Evermann. 1896. Fishes of North and Middle America. Bull U.S. Nat. Mus. Pt. 1,47:1-1240. 22. Kaill, W. M. 1967. Ecology and behavior of the cyprinodontid fishes Jordanella floridae Goode and Bean, Floridichthys carpio LITERATURE CITED Gunther , and Cyprinodon variegatus Lacqede. Unpubl. Ph.D. dissertation, Cornell Univ., 159 p. 1. Ager, L. A. 1971. The fishes of Lake Okeechobee. Quart. Jour. 23. Kushlan, J. A. and T. E. Lodge. 1974. Ecological and Fla. Acad. Sci. 34: 53-62. distributional notes on the freshwater fish of southern Florida. Fla. 2. Bailey, R. M., H. W. Winn, and C. L. Smith. 1954. Fishes from the Sci. 37(2):110-128. Escambia River, Alabama and Florida, with ecologic and 24. Miller, R. R. 1948. The cyprinodont fishes of the Death Valley taxonomic notes. Proc. Acad. Nat. Sci. Phila. 106:109-164. system of eastern California and southwestern Nevada. Misc. Publ. 3. Bohlke, J. E. and C. C. G. Chaplin. 1968. Fishes of the Bahamas. Mus. Zool. Univ. Mich. 68:1-155. Livingston Publ. C., 771 p. 25. 1955a. A systematic review of the Middle American 4. Breuer, J. P. 1962. An ecological survey of the lower Laguna fishes of the genus Profundulus. Misc. Publ. Mus. Zool. Univ. Mich. Madre, 1953-1959. Publ. Inst. Mar. Sci. Univ. Texas 8:153-183. 92: 1-64. 26. 1955b. An annotated list of the American cyprinodont 5. Brown, J. L. 1956. Distinguishing characters of the cyprinodont fishes of the genus Fundulus, with description of Fundulus fishes Fundulus cinaulatus Valenciennes and Fundulus chrysotus similis from Yucatan. Occ. Pap. Mus. Zool. Univ. Mich. 568:1-25. (Gunther). Copeia 1956 (4):251-254. 27. 1974. Designation of a neotype for Cyprinodon 6. 1957. A key to the species and subspecies of the mydrus, species of Floridichthys. Copeia 1974(4):981. cyprinodont genu§ Fundulus in the United States and Canada, east 28. Moore, G. A. 1968. Fishes. Pt. 2. In: Vertebrates of the United of the continental divide. Jour. Wash Acad. Sci. 47 (3):69-77. States by Blair, W. F., Blair, A. P., Brodkorb, P., Cagle, F. R., and 7. 1958. Geographic variation in southeastern Moore, G. A. McGraw-Hill Book Co., pp 21-165. populations of the cyprinodont fish Fundulus notti (Agassiz). 29. Myers, G. S. 1931 . The primary groups of oviparous Amer. Midl. Nat. 59:477-488. cyprinodont fishes. Stanford Univ. Publ. (Biol. Sci.) 6(3):241-254.

51 30. Relyea, K. 1965. Taxonomic studies of the cyprinodont fishes, 35. Rosen, D. E. 197/. Suborder Cyprinodontoidei. In: Fishes of the Fundulus confluentus Goode and Bean, and Western North Atlantic. 6:229-262. (Evermann). Unpubl. Masters Thesis, Florida State Univ., 73 p. 36. Smith-Vaniz, W. F. 1968. Freshwater fishes of Alabama. 31. 1967. Systematic study of two species complexes of Paragon Press, 211 p. brackish water Fundulus (Pisces: Cyprinodontidae). Unpubl. Ph.- 37. Tabb, D. C. and R. B. Manning. 1961. A checklist of the flora and D. Dissertation, Tulane Univ. fauna of northern Florida Bay and adjacent brackish waters of the 32. Rivas, L. R. 1948. Cyprinodont fishes of the genus Fundulus in Florida mainland collected during the period July, 1957 through the West Indies, with description of a new subspecies from Cuba. September, 1960. Bull. Mar. Sci. Gulf. and Carib. 11(4):552-648. Proc. U.S.N.M. 98:215-222. 38. Thomerson, J. E. 1966. A comparative biosystematic study of 33. 1966. The taxonomic status of the cyprinodontid Fundulus notatus and Fundulus olivaceus (Pisces: Cyprinodon- fishes Fundulus notti and F. lineolatus. Copeia 1966(2):353. tidae). Tulane Stud. Zoo!. 13(1) :29-47. 34. Rosen, D. E. 1964. The relationships and taxonomic position of 39. Walters, V. and C. R. Robins. 1961. A new toadfish the halfbeaks, killifishes, silversides, and their relatives. Bull. (Batrachoididae) considered to be a glacial relict in the West In- Amer. Mus. Nat. Hist. 127(5):217-268. dies. Am. Mus. Nov. 2047:1-24.

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