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This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute.

Notice: © 1983 Florida Academy of Sciences. This manuscript is an author version with the final publication available and may be cited as: Gilmore, R. G. & Hastings, P. A. (1983). Observations on the ecology and distribution of certain tropical peripheral fishes in Florida. Florida Scientist, 46(1), 31-51.

Biological Sciences

OBSERVATIONS ON THE ECOLOGY AND DISTRIBUTION OF CERTAIN TROPICAL PERIPHERAL FISHES IN FLORIDA

(l)R. GRANT GILMORE AND 12 lPHILIP A. HASTINGS

'"Harbor Branch Foundation, Inc., R.R. l, Box 196, Fort Pierce, Florida 33450, and '"Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85720

ABSTRACT: Maximum densities of east-central Florida population~· of the tropical peripheral JL~hes, Oostethus brachyurus lineatus, Gobiomorus dormitor, Awaous tajasica and Gobionellus pseudofasciatus occurred during the winter and spring month~ in the lower portiom of freshwater tributaries to the Indian River lagoon. All populations declined during the wet season. reaching minimum leveL~ during peak rainfall periods. Specific microhabitat requirements were observed for each species. These species have a limited Florida range which i.~ di.~cn~sed relative to various environmental parameters and dispersal mechanisms, such as water temperature regimes, oceanic current patterns and particular microhabitat preferences.

THE freshwater fish fauna of peninsular Florida has no indigenous primary freshwater representatives of tropical American origin. Floridian primary freshwater fishes have been recruited from temperate North American families (i.e., Centrarchidae, Percidae, Catostomidae and Cyprinidae; Carr and Coin, 1955; Briggs, 1958) and comprise 29-46% of the freshwater fish fauna depending on the latitude along the Florida peninsula. Therefore, approximately 54-71 o/o of the fishes occurring in the fresh waters of peninsular Florida may be classified as secondary freshwater or euryhaline marine forms (Kushlan and Lodge, 1975; Gilmore, 1977a). Marine fishes occurring in fresh water have been known collectively as peripheral species ("diadromous", vicarious", "complementary" or "sporadic" forms after Meyers, 1951). In Florida, many of these peripheral species are of tropical origin. Four tropical peripheral species, the opposum pipefish, Oostethus • brachyurus lineatus (Kaup), river goby, Awaous tajasica (Lichtenstein), slashcheek goby, Gobionellus pseudojasciatus Gilbert and Randall, and the bigmouth sleeper, Gobiomorus dormitor Lacepede, have been considered rare in Florida waters and have received little study (Gilmore, 1977b; Gilbert, 1978; Hastings, 1978). Recent collections of these species have pro­ duced enough information for a preliminary analysis of the temporal and spatial distribution of east-central Florida populations. These peripheral tropical species are exceptional in being among the few examples of euryhaline tropical freshwater species (either "vicarious", "diadromous" or "complementary") occurring in the fresh waters of continental North America. In contrast, the continental North American marine ichthyofauna contains numerous examples of tropical Caribbean marine forms. 32 FLORIDA SCIENTIST [Vol. 46

METHODS- Specimens were captured utilizing a variety of techniques (Gilmore, 1977 a). Col­ lections were made from November 1971 to December 1980 in a wide varietv of neritic, estuarine and freshwater habitats from 26°58'N to 29°0.5'N along the Florida east c,;ast (Fig. 1; Gilmore, 1977a; Gilmore, 1977b; Hastings, 1978). All fish length measurements are standard lengths (SL) unless otherwise noted. Inlet +N

0 () "'y.,.

Jupiter Inlet

FIG. l. Indian River lagoon of east central Florida and associated freshwater tributaries.

DISTRIBUTION AND NATURAL HISTORY OBSERVATIONS-Most specimens of the species discussed below were collected in low gradient freshwater rivers, creeks and canals flowing into the Indian River lagoon in east-central Florida. These freshwater tributaries have floodgates, salinity barriers or locks at various locations along their course. In all cases except one (0. brachyurus), the species discussed were collected from the downstream side of an artificial barrier. Additional captures of juvenile Oostethus brachyurus were made in saline estuarine waters and nearshore neritic waters.

Oostethus brachyurus lineatus (Kaup) (Opossum Pipefish) Distribution: (Fig. 2A, 3) Oostethus brachyurus (Bleeker) is widely distributed in tropical regions, occurring in the Atlantic, Indian and Pacific oceans (Dawson, 1979). The western Atlantic subspecies 0. brachyurus No. l, 1983) GILMORE AND HASTINGS- TROPICAL FLOIUDA FISHES 33

A 8

Fzc. 2. Caribbean-Western Atlantic distribution of (A) Oostcthus brachyurus lincatus (hypothetical oceanic distribution is stippled); (B) Gobiomon'"~ domzitor; (C) Au:aon~ ta;a~ica; (D) Gobionellns pseudofasciatus (hypothetical continental distribution of G. pseudojasciatus is hatched).

lineatus occurs from South Carolina to Sao Paulo, Brazil and commonly enters fresh water in Central America, the Greater Antilles (Black and Montego rivers, Jamaica, Caldwell, 1966; Rio San Juan, Cuba, Eigenmann, 1902) and east-central Florida (Gilmore, 1977b). It has not been collected from freshwater tributaries in extreme southern Florida (vicinity of Florida Bay), western Florida or the northern Gulf of Mexico. Extralimital (i.e., beyond the normal geographic range of breeding populations) salt marsh captures have been made in Mississippi, but these populations did not over­ winter (Dawson, 1970). One individual was collected in Sargassum in the northeastern Gulf of Mexico (Hastings and Bortone, 1976). Darnell (1955a, 1962) did not take this species in the Rio Tamesi drainage in northeastern • Mexico. However, Jordan and Dickerson (1908) record 3 specimens of 0. brachyurus (1 female, 2 males, as Doryhamphus lineatus) from lagoons at Tampico, Tamaulipas, Mexico. It is possible that permanent breeding populations may occur in fresh waters of this region of Mexico (22 o ION), but this has not been documented. Specimens have been collected recently from the lower reaches of the Rio Grande River on the Texas-Mexican border (Frank Pezold, pers. comm.). Ecology: Between 1972 and 1981 235 specimens (64-175 mm SL) of the opossum pipefish were captured from all months of the year. Specimens 34 FLORIDA SCIENTIST [Vol. 46

FIG. 3. Florida collection locations for Oostethu~ brachyurus lineatu~. ( t) ); Gobiomorn~ dormitor ( •); AwaouY tajasica (.A.); and all 3 species combined with Gobionellus pseudoja.sciatu1· (.). .------

30

~------28 --=-6 8 9

26 10

24

p 22 E 11 ~ 8. 20 ~~~12 ....E / .. /,/ / ------18 ------k1

16

14_~---.o.---~----~----~---~.---~----~6-----~------~--~1~0---- 1

Salinity ppt. FIG. 4. Temperature-salinity hydroclimograph for 0. lineatus using monthly means for col­ lections made from 1972 to 1981. Dark bars arc proportional to the total number of individuals collected during that month. No. 1, 198.3] GILMORE AND HASTINGS- THOPICAL FLOHIDA FISHES 35 301~ ---l

60 1 1 I pseudofasc atus ~ :: _._I ~· J I l. ------" ,> c Oostethus 0 I • u_.____ll_ __ brach:,:_ __ _ ~ .0 Gobiomorus E, 20 z dormitor 10 • - _ _. ·---· ------40 30 Awaous 20 tajasica 10 ___L ______---- JFMAMJJASOND

MONTH Fl(;_ 5. Comparative numerical seasonal distribution of total monthly collections of G. pseudofa~ciatus, 0. brachyurw·, G. dormitor and A. taia~ica with mean monthly rainfall (6.5 yr. mean from Fellsmere Station 7 SSW, Indian River Co., NOAA Climatological Data, Annual Summary, Florida, 1977).

were collected from the St. Lucie River (Martin Co.), and Sebastian Creek drainages, and the North, Main and South Relief canals in Indian River County, Florida. Specimens from the Indian River lagoon were collected at 2 sea grass bed locations in northern St. Lucie County. Salinities at the time of capture ranged from 0.0-37.0 ppt, though most specimens, 98 o/o, came from fresh water ( < 1.0 ppt). Water temperatures at capture ranged from 15.0 (at 0.0 ppt) to 30.5°C (at 0.0 ppt). A climagraph (Fig. 4) shows mean salinity and temperature values for monthly collections of opossum pipefish and the estuarine and neritic occurrence of juveniles during March and June. Up to 25 individuals were taken in a single collection (Nov. 1979) with 145 individuals, 64 o/o of all captured, collected during the winter months, November through March (Figs. 4 and 5). Few specimens were collected • during the wet season, June through October, with a seasonal population low during the peak rainfall months of August, September and October (Figs. 4 and 5). The smallest specimens (five, 64-86.3 mm SL) were taken at higher salinities (maximum of 37 ppt) in seagrass beds of the Indian River lagoon and in neritic waters of the Atlantic Ocean. These specimens were con­ sidered juveniles, as pouch development normally takes place at lengths greater than 100 mm. Our observations indicate that 0. brachyurus does not breed in neritic or 36 FLORIDA SCIENTIST [Vol. 46 pelagic waters, but is anadromous. Suitable biotopes for breeding are ap­ parently limited as this species has been found breeding throughout the year only in tropical America, the Antilles, and in east-central Florida. To-date, the only permanent breeding population reported north of Mexico occurs in patches of the emergent aquatic plants Polygonum hydropiperiodes and Panicum spp. in freshwater tributaries to the Indian River lagoon (Christensen, 1965; Gilmore, 1977b). Extralimital captures of 0. brachyurus in breeding condition have been made at the mouth of the St. Johns River, Florida (McLane, 1955), and from coastal South Carolina (Fowler, 1945, 2 specimens, 1 a male with eggs). Hildebrand (1939) in­ dicated that a permanent breeding population occurs in Gatun Lake, Panama, and Gilbert and Kelso (1971) provided evidence that this species breeds in tributaries to the Tortuguero estuary, Caribbean Costa Rica.

Gobiomorus dormitor Lacepede (Bigmouth Sleeper) Distribution: (Figs. 2B, 3) Gobiomorus dormitor occurs from the Aransas River, Texas and east-central Florida to Surinam (Miller, 1959; Briggs, 1958; Gilmore, 1977; Hoese and Moore, 1977). The bigmouth sleeper is common in the Rio Grande and in the Rio Tamesi drainage in Mexico (Darnell, 1955a, 1962; Haese and Moore, 1977) south through Central America (Nordlie, 1979, 1981) and in Jamaica (White River, Caldwell, 1966), Cuba (Poey, 1866; Eigenmann, 1902, Rio San Juan and Vento Springs), Puerto Rico (abundant, Rio Loiza, Rio de Caguitas, Rio Bayamon, Evermann and Marsh, 1900), and Martinique in the Lesser Antilles (Schultz, 1949). Current fish surveys suggest a limited Florida distribution as Gobiomorus dormitor has not been collected from freshwater tributaries of Florida Bay and the Gulf of Mexico (Tabb and Manning, 1961; Gunter and Hall, 1965; Odum and Heald, 1971; Barnett, 1972; Kushlan and Lodge, 1974; Brockmann, 1974). It has not been recorded from the Bahama Islands (Bohlke and Chaplin, 1968) nor from tributaries of the northern or north­ eastern Gulf of Mexico (Bailey et al., 1954; Hellier, 1967; Yerger, 1977). Ecology: Ninety-one specimens of the bigmouth sleeper (16-317 mm SL) have been collected in freshwater tributaries to the Indian River lagoon (South and North Relief canals, Indian River County, and Sebastian Creek, Indian River and Brevard counties) in salinities of 0.0-13.0 ppt and at temperatures of 10.5-29.5°C. Two specimens were collected at temperatures as low as 10.5°C. However, most individuals were captured from March to July at mean temperatures above 22°C (Fig. 7). Specimens were collected during all months of the year but were most common from January through July, with the largest single collections oc­ curring in January and July. Like 0. brachyurus, G. dormitor experienced a No. 1, 1983] GILMORE AND HASTINGS- TROPICAL FLORIDA FISHES 37

FIG. 6. (A) South Relief Canal at S.R. 605 bridge, Indian River County. Awaous taiasica was captured under the bridge, 0. brachyurn~ in Polygonum hydropiperoides (Poly) and Panicum purpurascens (Pan). (B) North Relief Canal under U.S. Hwy 1 bridge where largest col­ lections of A. taiasica have been made (mud and sand) along with Gobionellus pseudoja~ciatu1· (sand). (C) Polygonurn hydropiperoides. the principal microhabitat of 0. lineatus. (D) Deep undercut bank, a typical Gobiomorus dormitor microhabitat.

30

28

5

26 10

24 4

.u 22 ~ • •~ a. 20 •E 1-•

• 18

16

14 0 3 4 5 6 8 9

Salinity ppt.

FIG. 7. Temperature-salinity hydroclimograph for G. dormitor using monthly means for collections made from 1972 to 1981. Dark bars are proportional to the total number of in­ dividuals collected during that month. 38 FLOIUDA SCIENTIST [Vol. 46 population decline during the peak wet season months, August through Oc­ tober (Figs. 5 and 7). Juveniles were most common in late fall and spring col­ lections. Most large adults were collected near stream banks by hand, hook-and­ line, seine and with gill nets. Juveniles (16-50 mm) were collected over sand bottoms and along heavily vegetated shallow shorelines (Fig. 6). No specimens were taken in extensive, higher-salinity (17 -40 ppt) collections made along mangrove shorelines and in marshes, seagrass beds, open sand bottoms and lagoon reefs within the Indian River lagoon (Gilmore, 1977 a). East-central Florida populations of G. dormitor were found throughout the year only in freshwater tributaries, canals and natural stream beds. Our studies and the observations of others (e.g., Darnell, 1962; Nordlie, 1981) show the biotope preference of Gobiomorus to be quiet silt or sand bottoms in relatively shallow waters near vegetated banks of sluggish streams. All tributaries in which we collected this species have a relatively low gradient with currents varying from 0.2-1.0 m/sec. However, currents may increase temporarily (to 1.6 m/sec) when floodgates are opened, par­ ticularly in small canals. The benthic solitary, semi-burrowing nature of adult G. dormitor was verified by our collections. Juveniles (16-40 mm SL) were observed in groups of 4 to 5 individuals loosely dispersed, resting on a sand bottom and occasionally rising into the water column. Our collections do not show a preference for upstream or downstream locations within the freshwater tributary as G. dormitor has been collected at all stream collec­ tion sites and upstream as far as dam or floodgate barriers. However, it has not been collected from tributary mouths, or from the Indian River lagoon. The periodically opened locks on the St. Lucie River may allow the fish to pass further upstream as G. dormitor has been collected in Lake Okeechobee (Lee et al., 1980). Some lacustrine biotope preferences appear to be quite different than preferred stream habitats (e.g., rock bottom to depths below 15 m in Lake Jiloa, Nicaragua) and reveal the opportunistic and adaptive nature of this species (McKaye et al., 1979; Nordlie, 1981). Although G. dormitor appears to prefer epibenthic locations and stream banks during diurnal periods, there are several nocturnal observations in­ dicating that the species migrates to the surface, and occasionally out of the water (semiterrestrial activity) along stream banks. Christensen (1965) observed ... "15 to 20 individuals among rocks ... at night ... in water less than 4 inches deep" below a floodgate in the Loxahatchee River, Palm Beach County, Florida. Darnell (1955a, 1955b) observed nocturnal semiterrestrial activity in the Rio Tamesi and Rio Sabinas, Tamaulipas, Mexico, and Koenig, et al., (in Thorson, 1976) described G. dormitor as" ... often seen at night at the surface of shallow waters ... "in Lake Nicaragua. The bigmouth sleeper has been found to prey upon crustaceans, fishes, various insects and arachnids (Meek and Hildebrand, 1916; Darnell, 1965; Zaret and Rand, 1971; Nordlie, 1981). The species is a strict carnivore whose maximum size (610 mm SL; Hoese and Moore, 1977) allows it to be con- 1\'o. 1, 1983] GILMORE AND HASTINGS -TROPICAL FLORIDA FISHES .39

sidered a top predator in many freshwater habitats. Loftin (196.5) considered it to be one of the "dominant large predators in most of the freshwater streams of Panama". Darnell (1962) reported that G. dormitor may be somewhat "catadromous", spawning downstream in "brackish-water lagoons and estuaries". Large sexually mature adults and the smallest individuals (to 21 mm) were collected in the lower estuary at Tortuguero (Nordlie, 1981; Costa Rica), therefore, indicating downstream spawning activity. However, observations of reproduction in freshwater Lago de Yojoa in Honduras (Darnell, 1962) and Lake Jiloa, Nicaragua (McKaye et al., 1979) demonstrate that estuarine migrations are not necessary for effective reproduction in this species. In our collections the youngest specimens (16-23.5 mm) occurred in fresh to brackish water on the downstream side of salinity barriers, at salinities of 0.0-13.0 ppt. Insufficient specimens have been captured to allow isolation of specific spawning locations, but the cap­ ture of young-of-year as well as the capture of a ripe female in Sebastian Creek indicates that this species apparently spawns within this tributary.

Awaous tajasica (Lichenstein) (River Goby) Distribution: (Figs. 2C, 3). Awaous tajasica ranges from Florida south to Cuba (Poey, 1866; Rio Cuyaguateje, Eigenmann, 1902; Yerger, 1978), Puerto Rico (Bayamon River, Rio de Caguitos, Evermann, 1899; Anasco River, Austin, 1971), Jamaica (Port Antonio, Caldwell, 1966), Costa Rica (Gilbert and Kelso, 1971), Panama (Brockmann, 1965), Venezuela (Schultz, 1949) and Brazil (Brockmann, 1965). Awaous tajasica has not been recorded from the Lesser Antilles, Bahama Islands where major freshwater drainages are absent (Bohlke and Chaplin, 1968) or from the drainages of the western Gulf of Mexico (Darnell, 1955a, 1962; Roese and Moore, 1978). The only North American records for this species are from Florida. Currently the only documented occurrence of A. tajasica in peninsular Florida is from captures made in the Oklawaha-St. Johns drainage (McLane, 1955) and tributaries to the Indian River lagoon (Gilmore, 1977a, b; Yerger, in Gilbert, 1978). This species was also collected from Garnier's Creek, a tributary to Choctawhat­ chee Bay in the panhandle region of west Florida (Yerger, in Gilbert, 1978). However, the permanency of a northwest Florida population at this latitude (30°30N) is unknown. Awaous tajasica has not been recorded from fresh or marine waters south of the St. Lucie River, Martin County (27°10N) and was not listed by Kushlan and Lodge (1974) as a member of the south Florida ichthyofauna. Ecology: Forty-three specimens (29.5-148.0 mm SL) of the river goby have been collected from the St. Lucie River (Martin County), North and South Relief canals (Indian River County) and the N. Fork of Sebastian Creek (Brevard County). No specimens were collected at tributary mouths or in the Indian River lagoon itself. Salinities and temperatures at capture 40 FLORIDA SCIENTIST [Vol. 46 varied from 0.0-4.0 ppt and from 20-28°C, respectively, with most specimens coming from freshwater. Specimens were collected from March to November, with most from March through May. The largest single collec­ tion was made in March 1979 when 32 individuals (29.5-75.2 mm SL) wen captured on a sand bottom under the U.S. Route 1 bridge over the Nortt Relief Canal. Ninety-six percent of the captures have occurred on open sane bottoms below bridges (Fig. 6) which may indicate an association wit! shaded portions of the stream and/or sediment deposited around bridg pilings. Both maximum numbers and the smallest individuals occur i March, indicating an early spring recruitment. This species has been ol: served to burrow into sand bottoms and may account for its disappearanc from winter seine samples. The genus Awaous is represented by several species distribut{ throughout the world tropics (Brockmann, 1965). All occur in fresh water adults and little is known of their spawning habits and early life history. Tl occurrence of larval and small juvenile A. tajasica in the mouths of estuari implies that early development takes place under brackish estuarine condi­ tions and not in freshwater. Both Gilbert and Kelso (1971) and Schultz (1949) collected their smallest specimens (12.5-19 mm SL) from the mouths of estuaries in Costa Rica and Venezuela. Central American specimens (adults and juveniles) have been collected in various shallow lotic freshwater habitats usually with no aquatic vegeta­ tion and over sand, mud or rocky bottoms (Loftin, 1965; Gilbert and Kelso, 1971). Loftin (1965) collected the species at altitudes up to 610 m above mean sea level along the Atlantic slope in Panama. Currents in the Pana­ manian streams collected by Loftin varied from "fast" to "slow" demonstrating that current flow rate was not a major barrier to the upstream migrations to higher elevations. As there are no records of lentic populations of A. tajasica, this species may prefer flowing water in lotic habitats, hence its limited distribution in Florida.

Gobionellus pseudojasciatus Gilbert and Randall (Slashcheek Goby) Distribution: (Figs. 2D, 3) the slashcheek goby, Gobionellus pseudojasciatus, may be considered a tropical continental species having been recorded from Trinidad, Panama, Costa Rica, Guatemala, Belize, east-central Florida, the Bahamas, Greater Antilles, and Lesser Antilles (Hastings, 1978). It has not been recorded in Florida south or north of tributaries to the Indian River lagoon nor from tributaries of the Gulf of Mexico. Ecology: Three hundred and fifty specimens (17-44 mm SL) of G. pseudojasciatus were collected from the South Relief Canal and the Sebas­ tian Creek drainage. Specimens were captured in shallow water, primarily over sand bottom throughout the year and at salinities and temperatures No. 1, 1983] GILMORE AND HASTINGS~ TROPICAL FLORIDA FISHES 41

--I 4 ~3 24 0~1 ·­\

i 22 ~ E 20 ~

18 16 _j 14 ~-,.o----..---.-----.3--4..-----.s~--;cs--~---. 9 10

Salinity ppt

Fie. 8. Temperature-salinity hydroclimograph for G. pseudojasciatn~ using monthly means for collections made from 1972 to 1981. Dark bars are proportional to the total number of in­ dividuals collected during that month. ranging from 0.0-13.0 ppt and 17.0-32.0°C, respectively. This species is most common during the spring dry season, increasing in numbers as water temperatures increase from March through June (Figs. 5, 8 and 9). Stream populations were least abundant at our stations during the peak of the wet season from August through November. A comparison of upstream and downstream stations suggests that G. pseudojasciatus move into downstream localities during December and into upstream localities during January, reaching maximum numbers upstream from March through May (Fig. 9). Upstream populations decline from May through September while downstream populations increase during May and June, but then decline from July through August. PHYSICAL PARAMETERS AND BIOGEOGRAPHY- The 4 species discussed ex­ emplify a basically continental distribution limited to subtropical and tropical fresh waters, estuaries and nearshore neritic waters (Fig. 2). This basic distribution pattern is also seen in 54 other tropical fish species occur­ ring in east-central Florida, most of which are estuarine or marine forms (Table 1). Many of these species have a Florida distribution limited to the coastline from the vicinity of Cape Canaveral and Sebastian Inlet south to Biscayne Bay. The following discussion considers several environmental variables which may create the conditions allowing Oostethus brachyurus lineatus, Gobiomorus dormitor, Awaous tajasica and Gobiomorus 42 FLORIDA SCIENTIST [Vol. 46

pseudojasciatus and possibly other tropical species to be distributed as they are.

TABLE l. Tropical marine fishes showing a disjunct Caribbean-east Florida distribution. that are common to abundant in the Indian Hivt·r lagoon or adjacent continental shelf. i.e .. not recorded or rarely occurring on continental shelf of west Florida (Springer and VVoodhurn. 1960: Springer, 1961; Moe and Martin, 196.5; Smith, HJ76), northern Gulf of Mexico (Hastings. HJ7fl). and Texas (Hoese and Moore, 1978). Fishes following this same distribution pattern hut con­ sidered rare in east-central Florida have been omitted from this list.

Species

Engraulidae Apogonidae Arzchoa lamprotaerzia Apogon hinotatus' Synodontidae A. planifrom Synodus saurus A. stellatus' Ophidiidae Lutjanidae Lepophidium cervinum' Lutjm11~1· mwli~' Scorpaenidae L. apodu~· Scorpaerza grandicornis' L. jocu Centropomidae Gerreidac Centropomus pectinatus'Eu Diapterus auratr~~·Eu Serranidae Eucinostomn1· lejroyi * Hypoplectrus gemma E. mclanoptcrus' Prontogrammus aueroruhen~ (;CTTCS cirU'TClJ.,\' * Grammistidae Rypticus hL~trispirzu~' R. saponaceus' Pomadasyidac Clinidae Anisotrernus surinarnensis * Lahriwnnu~ gohio' Haemulorz parrai' L. nuchipirmi~' H. sciurus' Malococtem~s macrop1~1·' Sparidae M. triangu/atus' Archosargus rhomboidalis' Paraclim~1· rzigripirmis' Diplodu~ argenteus' Blenniidae Sciaenidae Blermius cristatus' Bairdiella sanctaeluciae' Hypleurochilus aequipinnis Equetes acuminatus' Gohiidae Umhrina coroides' Crwtholepis thomp.wmi Pempheridae Cohionellus occmlicnl' Pempheris schomhurgki * G. stigmaturis' Labridae Lophogohius cyprirwides 'Eu Halichoeres maculipinna * Bothidae Scaridae Citharichthys arerzaccus· Scan~s guacamaia Mono!eiU' arztillarum' Spariwnna chrysopterum ' Paralichthys dentatus' S. ruhripinne' P. ohlongu1·' Opistognathidae Scophtha/mus aquosus Opistognathus aurifrons' Tetraondontidae 0. rohimi' Sphoeroides testudineus' Dactyloscopidae Dactyloscopu~ crossotus·' Gillellus greyae' G. ruhrocinctu1·'

* = form permanent breeding populations in east central Florida. Eu ~ euryhaline tropical spt•cies. No.1, 1983] GILMORE AND HASTINGS ~TROPICAL FLOIUDA FISHES 43

Spillway Downstream Site soom 2ooom J F M A M J J A s .....

0 0 .... 0 0 •• . . . . . N D

FIG. 9. Spatial and temporal distribution of G. pseudofasciatus in the north fork of Sebastian Creek and the Fellsmere Canal, 1979-1980. [ = 1-4 individuals, L__j = .5-10,

= 11-15, mill = 16-19, - = 20 + 0

Temperature and Biogeography: Past and present temperature regimes have had a critical affect on the distribution of tropical species in the climatically transitional Florida peninsula. The presence of permanent breeding populations of tropical peripheral species in tributaries to the In­ dian River lagoon indicates that a survivable temperature regime occurs in these streams today. The minimum water temperature at which these fishes have been collected has ranged from 10.5°C for Gobiomorus dormitor, to ' l5°C for 0. brachyurus, l7°C for Gobionellus pseudojasciatus, to 20°C for A. ta;asica. The 10.5°C water temperature record for G. dormitor was ex­ ceptional as 9 previous years of winter temperature records from these streams reveal that temperature minima do not usually fall below l7°C. No hypothermal stress and mortality has been observed in these 4 species though it has been documented for other resident tropical species (Gilmore et al., 1978). The range of local freshwater temperatures is not as great as the temperature range for local estuarine waters. Our data show a water temperature minimum in local freshwater streams 4-5°C higher than in the Indian River lagoon. Movement of shallow ground water through canal 44 FLORIDA SCIENTIST [Vol. 46 banks may account for some of these thermal differences (Beck, 1965). In Florida, winter freshwater temperatures are generally higher than those of adjacent estuarine waters because of isothermal spring and shallow ground­ water sources. Isothermal spring effects on minimum water temperatures can be seen in various higher latitude tributaries such as the Suwannee River, 29°21 to 30°32N latitude, with a minimum of 15°C (Beck, 1965), the Santa Fe River, 29°53' to 29°55N with l7°C (Hellier, 1967), and the Hillsborough River, 27°53' to 28°21 N, with 19°C (Barnett, 1972). The use of these freshwater sources by fishes as refugia during periods of potential thermal stress has been documented (Gilmore et al., 1978). The absence of the 4 species studied from freshwater tributaries elsewhere in peninsular Florida may be due to seasonally low neritic and estuarine water temperatures, particularly if the life history of these fishes includes a marine or estuarine phase (as seen in 0. hrachyurus). Minimum marine and estuarine temperatures can be quite different between locations at the same latitude on the east and west coasts of peninsular Florida. The east central and southeast coasts of Florida are generally warmer (Fig. 10; Rivas, 1968; Clark et al., 1970; Ichiye et al., 1973; Gilmore et al., 1978). Awaous tajasica, 0. hrachyurus and G. pseudojasciatus have not been recorded from fresh water in Florida south of 26°58N. Gohiomorus dar-

FrG. 10. Mean February sea surface temperatures for the western North Atlantic and Carib­ bean (redrawn from Fuglister, 1947, with conversion of original Fahrenheit temperatures to ap­ proximations of degrees Celsius). No.1, HJ83] GILMORE AND HASTINGS- TROPICAL FLOIUIJA FISHES 45 mitor has been recorded from freshwater canals of southeast Florida in Palm Beach and Broward counties, south to 26°00'N (Herrema, 1974). As freshwater drainages of extreme southern Florida are seasonally \Varmer than those of the Indian River lagoon, temperature may not be the only major factor affecting the distribution of these fishes in extreme south Florida and southwest Florida. Postglacial dispersal of tropical peripheral species from the Caribbean to Florida would be a plausible theory of distribution if water temperatures in Florida during the Wisconsin glaciation had been below the thermal tolerance levels of these species. Emiliani (1970) describes a 6°C water temperature oscillation for the Caribbean Sea between glacial and in­ terglacial times. Our observations during the present interglacial period show water temperatures as low as l0 ..5°C at locations where these fishes were captured. The minimum water temperatures during the last glaciation were lower and possibly beyond the thermal tolerance levels of tropical peripheral species. It is also possible that at least the gobioid tropical peripheral species are relicts of a more widespread interglacial distribution which included the northern Gulf of Mexico and southeastern Piedmont. A vicariant event (i.e., glaciation) could have then separated the Central American and Florida populations. A similar vicariance model for the distribution of Antillean­ Central American tropical species to the Florida peninsula has been presented by Rosen (1975) for other organisms (e.g., Wagner's mastiff bat, Eumops glaucinus jloridanus). The continental distribution of the fishes of the freshwater poeciliid genus Heterandria follow the generalized "North American-Caribbean track" described by Rosen (1975) with Florida and Central American forms representing 2 distinct species. Heterandria has a discontinuous generic distribution pattern demonstrating possible allopatric speciation (vicariance). The Heterandria distribution pattern is similar to that of the tropical peripheral fish species discussed here (excluding the An­ tilles and northern coast of South America). Florida fish relicts may have survived the last glaciation through warm isothermal spring refugia. Temperature tolerance levels for these species may also have changed since the last glaciation and there are no available data on their present hypothermal tolerances. Behavioral adaptations such as burrowing (observed in G. dormitor and A. tajasica) and aestivation (not observed) during adverse environmental periods may have also enabled the gobioid species to survive glaciation. However, past and present water temperature regimes are but one of several environmental parameters affect­ ing the present distribution of these species. This is demonstrated by their discontinuous distribution in warm water regions today. Even if east Florida may have a relict tropical peripheral fish population, because of oceanic dispersal these populations may not have lost contact with southern populations. In contrast to the probable allopatric speciation in Heterandria varying degrees of taxonomic uniformity is found among 46 FLORIDA SCIENTIST [Vol. 46 these species throughout their range (Miller, 1959; Brockmann, 1965; Gilbert and Kelso, 1971; Gilbert and Randall, 1980). All western Atlantic populations of 0. brachyurus show taxonomic continuity at the subspecific level as defined by Dawson (1979). This indicates gene flow rather than isolation and differentiation.

FIG. 11. (A) Tropical Western Atlantic current patterns during July (redrawn from lchiye et al., 1973); (B) Loop Current and cross-hatched cyclonic gyre (cross-hatched region) on the west Florida continental shelf (redrawn from Ichiye et al., 1973).

Oceanic Currents and Biogeography: Oceanic currents are known to play an important role in the distribution of 0. brachyurus (Dawson, 1979). Oostethus brachyurus has been captured at points of landfall for major warm water currents such as the Loop Current in the Gulf of Mexico and the Florida Current along the southeast coast of North America (Figs. 2A, 11 and 12). Juvenile 0. brachyurus have been taken in neritic and pelagic col­ lections (Gilbert and Kelso, 1971; Dawson, 1979) and have been found in association with floating Sargassum (Bohlke and Chaplin, 1968; Hastings and Bartone, 1976). Oceanic dispersal may occur in the gobioid species as well as in 0. brachyurus as all of these tropical peripheral species have demonstrated varying degrees of euryhalinity. At Tortuguero, Costa Rica, A. tajasica and No. 1, 1983] GILMORE AND HASTINGS- TROPICAL FLORIDA FISHES 47

)

FIG. 12. (A) Drift card returns from a 24-27 October 1972 release in the Caribbean south of Puerto Rico (redrawn from Duncan et al., 1977. Fig. 2). (B) Drift bottle reco\'ery sites after release in the Mona Passage west of Puerto Rico (redrawn from Metcalf et al., 1977, Fig. 2). (C) and (D) 2 studies of drift bottle trajectories after release from the northern portion of the Loop Current in the northern Culf of Mexico (redrawn from Ichiye et al., 1973).

G. dormitor are found in the lower estuary and the former species was col­ lected from the "tismiche" occurring in the inlet to Tortuguero Lagoon (Gilbert and Kelso, 1971). These observations and the occurrence of A. ta­ jasica and G. dormitor in the Antilles suggests that these 2 species may also be subject to oceanic dispersal. There is no present evidence that G. pseudojasciatus may also be capable of oceanic distribution as it has only been recorded from continental locations and the continental island of Trinidad. ., Oceanic distribution may be more plausible after the examination of the dispersal mechanism of tropical species geographically sympatric with these 4 tropical peripheral species. There are 54 marine and euryhaline estuarine fishes (Table 1) which have a discontinuous tropical distribution similar to the tropical peripheral species discussed here. Many of these 54 species have larvae capable of being carried by major oceanic currents. Studies of drift bottle trajectories reveal potential drift routes for wafted juveniles of several tropical peripheral and marine species capable of oceanic transport by surface currents (e.g., 0. brachyurus). Drift bottles released in the Gulf of Mexico near the apex of the Loop Current were carried to the 48 FLORIDA SCIENTIST [Vol. 46 Florida east coast, not the Gulf coast of the Florida peninsula (Ichiye et al., 1973). Drift bottle movement to the central west coast of Florida was blocked by the presence of a cyclonic gyre located on the broad west Florida shelf (Figs. 11, 12). Once bottles reached the Florida Current they were car­ ried by this strong and steady current to various locations along the Florida east coast. Drift bottle releases in the Caribbean (Duncan et al., 1977; Met­ calf et al., 1977) also show a trajectory which, after passage through the Yucatan Straits (by the Yucatan Current) either end up in the western or northern Gulf of Mexico or east coast of Florida, not the west coast of penin­ sular Florida (Fig. 12). The lack of drift bottle returns from the west coast of peninsular Florida may provide an explanation for the absence of 0. brachyurus records from that coast and possibly the absence of other tropical species (Table 1). Oceanic current transit times may in part account for the discrepancy between the east and west Florida distribution of 0. brachyurus. If penin­ sular west coast pipefish populations could be recruited from Central America via the Loop Current and associated gyres, transit time would be longer than for the east coast populations recruited from the more direct and swift Florida Current. Trans.it time in oceanic currents would be important if wafted individuals could only tolerate higher salinities for short periods of time. If the wafted individuals are small, in transit mortality rates resulting from predation could be very high. HABITAT AND COMMUNITY RELATIONSHIPS- Oostethus brachyurus, A. ta­ jasica, G. dormitor and G. pseudojasciatus have been collected together at various locations throughout the American tropics. The first 3 species have been collected at the same date and location in Venezuela (Schultz, 1949), Panama (Loftin, 1965), Costa Rica (Gilbert and Kelso, 1971) and Tampico, Mexico (Jordan and Dickerson, 1908). Gobionellus pseudojasciatus has been collected with the other species in Costa Rica (Gilbert and Kelso, 1971) and Belize (Thomerson and Greenfield, 1975, excluding A. tajasica). However, observations in both Central America and east-central Florida show that although these species may be sympatric in various stream systems their specific biotope preferences are quite different. In Panama and Costa Rica adult A. tajasica were collected in flowing lotic water up to altitudes of 610 m and over sand, mud or rock bottoms (Loftin, 1965; Gilbert and Kelso, 1971), while G. dormitor was found in a variety of lotic and lentic habitats (Loftin, 1965; Gilbert and Kelso, 1971; McKaye et al., 1979; Nordlie, 1981). In east-central Florida adult G. dormitor were collected near stream banks and under bank overhangs with no apparent preference for floral types. Juvenile G. dormitor were observed on sandy bottoms near the bank adja­ cent to emergent or floating vegetation; they may hover above the sand or in the vegetation, but are most often seen resting on the bottom. Florida populations of A. tajasica occurred on shaded sand to mud bottoms and prin­ cipally under bridges. East-central Florida freshwater collections of 0. brachyurus were limited to emergent bank vegetation, specifically No. 1, 1983] GILMOHE AND HASTINGS- THO PICA!. FLOHIDA FISHES 49

Polygonurn hydropiperiodes, Panicurn repens and P. purpurascens. Go­ bionellus pseudojasciatus was captured on shallow open sand bottoms with little or no vegetative cover. These biotopes, including the specific floral species, are found in streams throughout peninsular Florida. This indicates that other environmental conditions are required by these particular species in addition to specific substrata. Stream hydrography, water chemistry and biological parameters such as ) food resources (e.g., invertebrate fauna and microflora) and predators may vary considerable around the Florida peninsula (Beck, 1965). These factors may also play a vital role in the successful habitation of tropical peripheral fishes in east Florida streams. None of these parameters has been adequately studied relative to these species. The study of faunas occurring within the same biotope and their interac­ tions with tropical peripheral fish species may expose other mechanisms in­ fluencing the restricted Florida distribution of 0. brachyurus, G. dormitor, A. tajasica and G. pseudojasciatus. These interactions may be both com­ petitive and predatory, particularly for larvae and juveniles. Controlled ex­ perimental and in situ field observations of interspecific interactions may reveal biological factors affecting the distribution of these species. In Florida, a study of freshwater and peripheral faunal interactions is complicated by the erection of artificial water flow barriers on most streams, canals and rivers in coastal regions of the lower peninsula south of 28°00'N. Many of these barriers do not permit peripheral tropicals to migrate into in­ land streams where potential interactions with primary freshwater fish faunas may occur. Primary freshwater faunas and communities in turn, are best developed and most diverse on the upstream side of these barriers \vhere these tropical peripheral species do not occur. The data presented in this study are coincidental to major faunal studies of freshwater fish communities in east-central Florida. Further analysis of these data and that of sympatric fishes will reveal additional biological in­ formation on these particular tropical peripheral species and their in­ terspecific interactions within the freshwater fish community. AcKNOWLEDCMENTS- We thank LaVergne Williams, George Kulczycki, Wavne Magley, Christopher Donohoe, Douglas Cooke and Mike Clark for aid in collecting specimens. Several } specimens of Oostethus brachyurns were captured by Applied Biology, Inc. at the Florida Power & Light power plant on Hutchinson Island, St. Lucie Co., and at the Indiantown power plant on the St. Lucie Canal. RobertS. Jones and Carter H. Gilbert kindly reviewed early versions of this manuscript. This constitutes contribution no. 24H from the Harbor Branch Foundation, Inc.

LITERATURE CITED

BAILEY, H. M., H. E. WINN, AND C. L. SMITH. HJ54. Fishes from Escambia River, Alabama and Florida, with ecologic and taxonomic notes. Proc. Acad. Nat. Sci. Philadelphia. Vol. 61, 1954:109-164. BARNETT, B. S. 1H72. The freshwater fishes of the Hillsborough River drainage, Florida. M.S. thesis. Univ. of South Florida, Tampa. . BEcK, W. M., Jn. 1965. The streams of Florida. Bull. Florida St. Mus. 10:91-126. 50 FLOI\IDA SCIENTIST [Vol. 46

BoHLKE. J. E .. A~D C.C.C. CHAPLI~. l\J6R. Fishc•s of the Bahamas and Adjacent Tropical \\"aters. Livingston Pub!. Co .. \\'nme\\·oocl. PennsYlvania. Bmccs. J. C. 195R. A list of Florida fishes and their distribution. Bull. Florida St. \lm. :2::22:3- :JIR. BRoCK'\IA~"'- F.\\". 196.5. A taxonomic stuch of the Panamanian fn·sll\\·ater gobies of the .l(encra Atraous and Sicydium. \1.S. tlwsis. Florida St. Unh .. Tallahassee. _____ . 1914. SeasonalitY of fishes in a South Florida brackish canal. ()uart. J. Florida Acad. Sci. 3/:65-10. CALDWELL D. K. Hl66 . .\1arine and freshwater fishes of Jamaica. Bull. Inst. Jamaica. Sci. Scr. No. l'i. CARR. A .. A~D C. J. Gmr-:. HJ5S. Guide to the Reptiles. Amphibians and Fresln,·ater Fishes of Florida. Univ. Florida Press. Gainesville. CHICKFJU~c. A . .\1. 1930. An Atlantic pipefish caught in transit through the Panama Canal. Copeia. 19:30:8.5-86. CHRISTEl':SE~. R. F. 196.5. An ichthyological surwy of Jupiter Inlet and Loxahatchee RiH'r. Florida. linpubl. .\lasters Thesis. Florida St. Univ .. Tallahassee. CLARK. J .. \\'. G. S'\IITIL A. \\'. KE:-;DALL. Al':D \1. 1'. FAHEY. 1910. Studies of estuarine de­ pendence of Atlantic coastal fishes. Data lkp. I. L'.S. Burc•au of Sport Fish. and \\'ildl.. Tech. Paper .59. DAR~ ELL R. M. 19.53. An ecological stuch· of the H.io Sabinas and related waters in the southern Tamaulipas. Mexico with special reference to the fishes. Ph.D. dissert. Uni\· . .\linnesota . .\linneapolis. Minnesota. _____ . 195.5. Nocturnal terrestrial habits of the tropical gobioid fish Gohiomorus dormitor. with remarks on its ecolO!-,'Y· Copeia. 1955:2:31-2:38. _____ . 1962. Fishes of the Rio Tamesi and related coastal lagoons in east-central .\texico. Pub!. Inst. Mar. Sci. Vol. 8:299-:36.'5. DA wsor-:. C. E. 1910. A Mississippi population of the opossum pipefish. Oostcthus linea/us (Smgnathidae). Copeia. 19/0:'i/2-'i/:3. _____ . HJ/9. Review of the polvtypic dornhamphinc' pipefish Oostcthus brachyurus (Bleeker). Bull. Mar. Sci. 29:465-480. Dt·~cA~. C. P .. D. K. ATwooD. J. R. Dt·~cAr-:. Al':D P. 1\. FHOELICII. 19/'i. Drift bottle returns from the Caribbean. Bull . .\1ar. Sci. 27:580-.'586. EIGE1':'\1Ar-:~. C. H. 1902. The freshwater fishes of western Cuba. Bull. Fish. Comm.: 211-236. E'\IILIA~I. C. 1970. Pleistocene paleotemperatures. Science. 168(:39:3:3):822-R2S. EvER'\!Al':~. B.\\' .. Al':D \1. C. MAHSH. HJOO. The fishes of Puerto Rico. U.S. Fish. Comm. Bull. 1900. FowLER. H.\\'. 1945. A study of the fishes of the Southern Piedmont and coastal plain. Acad. Nat. Sci. Philadelphia \lonogr. 1:1-408. FucusTER. F. C. 1947. Average monthly sea surface temperatures of the western north Atlantic ocean. Pap. Phys. Ocean and Meteorology. Massachusetts Inst. Tech .. \\'nods Hole Ocean. Inst. 10(2): 1-2.5. GILBERT. C. R. 1978. Opossum pipefish. Pp :38-:39. /11: Gilbert. C. R. (eel.). Hare and Endan­ gered Biota of Florida: Vol. 4. Fishes. Univ. Presses Florida. Gainesville. _____ , AKD D. P. KELSO. 1971. Fishes of the Tortuguero area. Caribbean Costa H.ica. Bull. Florida St. Mus. 16:1-.53. -----· AND J. E. RAKDALL. 1979. Two new western Atlantic species of the gobioid fish genus GobionclltL~. with remarks on characteristics of the genus. :-\orthcast Gulf Sci. 3:27-47. GIL'\IOHE. R. C. 1977a. Fishes of the Indian River lagoon and adjacent waters. Florida. Bull. Florida St. .\1us .• 22:101-147. _____ . 1977b. Notes on the opossum pipefish Oostl'fhus liwatus. from the Indian River lagoon and vicinity, Florida. Copeia. 1977:181-18:3. GILMORE. R. G .. L. H. BuLLOCK. A~D F. H. BERRY. 197R. Hypothermal mortalitv in marine fishes of south-central Florida. January. 1977. Northeast Gulf Sci. 2:/'i-97. HASTINGS. P. A. 1918. First North American continental records of Gobione//u.~ pscudojasciatu.s (Pisces: Cobiidae). Northeast Gulf Sci. 2:140-144. _____, Al\D S. A. BoRTONE. 1976. Additional notes on tropical marine fishes in the northern Gulf of Mexico. Florida Sci. :39:12:3-125. HELLIER. T. R .. JR. 1967. The fishes of the Santa Fe River system. Bull. Florida St. :\ius. 11. HERREMA, D. J. 1914. Marine and brackish water fishes of southern Palm Beach and northern Broward counties. Florida. Masters Thesis, Florida Atlantic Univ .. Boca Raton. No.1, 1983] GILMORE AND HASTINGS- TROPICAL FLORIDA FISHES 51

HILDEBRAND, S. F. 1939. The Panama Canal as a passageway for fishes, with lists and remark~ on the fishes and invertebrates observed. Zoologic a. 24:1.5-45. HoEsE, H. D., AND R. H. MooRE. 1977. Fishes of the Gulf of Mexico, Texas, Louisiana and ad­ jacent waters. Texas A & M Univ. Press, College Station. lcmYE, T., H. H. Kuo, AND M. R. CARRES. 1973. Assessment of currents and hydrography of the eastern Gulf of Mexico. Texas A & M Univ. Contrib. No. 601: p. 0-1 through ll-24. JoRDAN. D. S., AND M. C. DICKERSON. 1908. Notes on a collection of fishes from the Gulf of Mexico at Vera Cruz and Tampico. Proc. U.S. Nat. Mus. 34(1.592):11-22. KoENIG, K. W., R. J. BEATTY, AND S. MARTINEZ. 1976. Species diversity and distribution of fish in Lake Nicaragua. Pp. 321-325. In Thorson, T. B. (ed.). Investigations of the iehthy­ • ofauna of Nicaraguan Lakes. School of Life Sciences, Univ. of Nebraska, Lincoln. KusH LAN, J. A., AND T. -E. Lom:E. 1974. Ecological and distributional notes on the freshwater fish of southern Florida. Florida Sci ..37:110-128. LEE, D. S., C. R. GILBERT, C. H. HocuTT, R. E. JENKINS, D. E. McAI.I.ISTEH, AND J. R. STAUF­ FER, JR. 1980. Atlas of North American freshwater fishes. North Carolina St. Mus. Nat. Hist. McLANE, VV. M. 1955. The fishes of the St. Johns River system. Ph.D. dissert., Univ. of Florida, Gainesville. McCosKER, J. E., AND C. E. DAWSON. 197.5. Biotic passage through the Panama Canal, with particular reference to fishes. Mar. Bioi. 30:.343-3.51. METCALF, \V. G., M. C. STALCUP, AND D. K. ATwooD. 1977. Mona Passage drift bottle study. Bull. Mar. Sci. 27(3):.'586-.591. MILLER. R. R. 19.59. Variation, distribution, and relationships of the Mexican eleotrid fish Gohiornoru~ polylepis. Occ. Pap. Mus. Zoo. Univ. Michigan No. 607. _____ . 1966. Geographical distribution of Central American freshwater fishes. Copeia. 1966:12.5-1.56. Mm:, M.A., AND G. T. MAHTIN. 196.5. Fishes taken in monthly trawl samples offshore Pinellas County, Florida, with new additions to the fish fauna of the Tampa Bay area. Tulane Studies in Zoo!. 12:129-1.51. MYEHS, G. S. 19.51. Freshwater fishes and East Indian zoogeography. Stanford Ichthyol. Bull. 4:11-21. _____ . 1966. Derivation of the freshwater fish fauna of Central America. Copeia. 196(i: 766-802. NoHDLIE, F. G. 1979. Niche specificities of eleotrid fishes in a tropical estuary. Hev. Bioi. Trop. 27:3.5-.50. 1981. Feeding and reproductive bioloh'Y of eleotrid fishes in a tropical estuary. J. Fish. Bioi. 1981:97-110. PoEY, F. 1868. Fisico-natural de Ia isla de Cuba. Torno. 2. RIVAS, L. P. HJ68. Fishermen's atlas of monthly sea surface temperatures for the Gulf of Mexico. U.S. Fish Wildlife Serv. Bureau Connn. Fish. Contrib. No ..5\J: 1-33. RosEN, D. W. 197.5. A vicariance model of Caribbean biogeographv. Sys. Zoo!. 24:431-464. Sun·LTZ. L. P. 1949. A further contribution to the ichthyoloh'Y of Venezuela. !'roc. U.S. Nat. Mus. 9D(.'323.5). SMITH, G. B. 1976. Ecology and distribution of eastern Gulf of Mexico reef fishes. Florida Dept. Nat. Res., Florida Mar. Res. Pub!. no. 19. SPRIN<:EH, V. G. 1961. Notes on and additions to the fish fauna of the Tampa Bay area in Florida. Copeia. 1961:480-482. _____ ,AND K. D. WooDBUHN. 1960. An ecological study of the fishes of the Tampa Bay ., area. Florida St. Bd. Conserv. Prof. Pop. Ser. no. 1:1-104. THoMEHSON. J. E .. AND D. \V. GHEENFIELD. HJ7.5. Anchoddla helizensi~. a new species of an­ chovy from Belize, Central America, with records of associated freshwater species. Copeia. 197.5:.50-.52. Ym<:>J<. H. \\'. UJ78. Hiver Coby. Pp. 46-47. In: Gilbert, C.H. (eel.). Rare and Endangered Biota of Florida: Vol. 4. Fisht's, Univ. Presses Florida. Gainesville.

Florida Sci. 46(1):31-.51. 1983.