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REFERENCE COW Do Not Remove from the Library U. S. Fish and Wildlife Service Biological Report 82(11.1()4) National Wetlands Kescarcn .. :nrer 'TR EL-82.4 JUIY 1989 700 Cajun Dome Eoulc-tr.;yri

Species Profiles: Life Histories and Environmental Requirements of Coastal Fishes and Invertebrates (South Florida)

LADYFISH AND

Coastal Ecology Group Fish and Wildlife Service Waterways Experiment Station U.S. Department of the Interior U.S. Army Corps of Engineers Biological Report 82(11.104) TR EL-82-4 July 1989

Species Profiles: Life Histories and Environmental Requirements of Coastal Fi shes and Invertebrates (South Florida)

LADYFISH AND TARPON

Alexander V. Zale and Susan G. Merrifield

Oklahoma Cooperative Fish and Wildlife Research Unit Department of Zoology 404 Life Sciences West Oklahoma State University Stillwater, OK 74078

Project Officer David bran National We tl ands Research Center U.S. Fish and Wildlife Srvice 1010 Gause Boulevard Slide1 1, LA 70458

Performed for

Coastal Ecology Group Waterways Experiment Station U.S. Army Corps of Engineers Vicksburg, MS 39180

and

National Wetlands Research Center Research and Development Fi sh and Mi1 dli fe Service U .S. Department of the Interior Washington, DC 20240 This series may be referenced as follows: U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life histories and environmental requirements of coasm fishes and invertebrates. U.S. Fish Wildl. Serv. Biol. Rep. 82(11). U.S. Army Corps of Engineers, TR EL-82-4.

This profile may be cited as follows:

Zale, A.V., and S.G. Merrifield. 1989. Species profiles: life histories and environmental requirements of coastal fishes and invertebrates (South Florida) -- ladyfish and tarpon. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.104). U.S. Army Corps of Engineers, TR EL-82-4. 17 pp. PREFACE

This species profile is one of a series on coastal aquatic organisms, principal ly fish, of sport, commercial, or ecological importance. The profiles are designed to provide coastal managers, engineers, and biologists with a brief comprehensive sketch of the biological characteristics and environmental requirements of the species and to describe how populations of the species may be expected to react to environmental changes caused by coastal development. Each profile has sections on taxonomy, 1ife history, ecological role, environmental requirements, and economic importance, if appl icable. A three-ring binder is used for this series so that new profiles can be added as they are prepared. This project is jointly planned and financed by the U. S. Army Corps of Engineers and the U.S. Fish and Wildlife Service.

Suggestions or questions regarding this report should be directed to one of the following addresses.

Information Transfer Specialist National Wetlands Research Center U.S. Fish and Wildlife Service NASA-Sl idel 1 Computer Complex 1010 Gause Boulevard Slidell, LA 70458

U. S. Army Engineer Waterways Experiment Station Attention: WESER-C Post Office Box 631 Vicksburg, MS 39180 CONVERSION TABLE

Metric to U.S. Customary

To Obtain millimeters (mm) inches centimeters (cm) inc hes meters (m) feet meters (m) fathoms kilometers (km) statute miles kilometers (km) nautical miles square meters (m2) 10.76 square feet square kilometers (km2) 0.3861 square miles hectares (ha) 2.471 acres liters (1) gal 1ons cubic meters (m3) cubic feet cubic meters (m3) acre-feet milligrams (mg) ounces grams (g) ounces ki1 ograms (kg) pounds metric tons (t) pounds metric tons (t) short tons kilocalories (kcal) British thermal units Celsius degrees (OC) Fahrenheit degrees

U.S. Customary to Metric inches 25.40 millimeters inches 2.54 centimeters feet (ft) 0.3048 meters fathoms 1.829 meters statute miles (mi) 1.609 ki1 ometers nautical miles (nmi) 1.852 ki1 ometers square feet (ft2) square meters square miles (mi2) square ki1 ometers acres hectares gallons (gal) 1i ters cubic feet (ft3) cubic meters acre-feet cubic meters ounces (oz) milligrams ounces (02) grams pounds (lb) ki1 ograms pounds (lb) metric tons short tons (ton) metric tons British thermal units (Btu) kilocalories Fahrenheit degrees (OF) Cel sius degrees CONTENTS

Page PREFACE ...... iji CONVERSION TABLE ...... lv ACKNOWLEDGMENTS ...... vi NOMENCLATURE/TAXONOMY/RANGE ...... 1 MORPHOLOGY AND IDENTIFICATION AIDS ...... 3 Ladyfish ...... 3 Tarpon ...... 4 REASON FOR INCLUSION IN THIS SERIES ...... 4 LIFE HISTORIES ...... 4 GROWTH CHARACTERISTICS ...... 7 Age and Growth ...... 7 Morphometric Relations ...... 8 FISHERY ...... 8 ECOLOGICAL ROLE ...... 9 Feeding Behavior/Food Habits ...... 9 Predators ...... 10 Parasites ...... 10 ENVIRONMENTAL REQUIREMENTS ...... 10 Temperature ...... 10 Salinity ...... 11 Dissolved Oxygen ...... 11 Contaminants ...... 12 Turbidity ...... 12 Wetlands Destruction and Degradation ...... 12 LITERATURE CITED ...... 13 ACKNOWLEDGMENTS

Drafts of this Species Profile were critically reviewed by Ed Rutherford, National Park Service, Everglades National Park, Florida; C. Richard Robins, University of Miami, Miami, Florida; and Paul H. Eschmeyer, Editorial Office, U.S. Fish and Wildlife Service, Fort Collins, Colorado. Sheila G. Johnson, Edmon Low Library, Oklahoma State University, provided exceptional assistance with 1i terature searches and inter-1 ibrary 1oan. H. Frank1i n Percival of the Florida Cooperative Fish and Wildl ife Research Unit, University of Florida, Gainesvil le, and 0. Eugene Maughan of the Oklahoma Uni t provided administrative assistance. Figure 1. A: ladyfish; B: tarpon.

LADYFISH AND TARPON

Geographic range .... western Atlantic Ocean from and southern New Scientific name ...... England (but uncommon north of Cape Linnaeus (Robins et a1 . TWOT - Hatteras) to Rio de Janeiro, , Preferred common name ...... 1adyf ish and throughout the (Figure 1A) (Figure 2); also occurs in the In- Other common names . . bigeyed herring, dian and western Pacific Oceans bony-fi sh, chi ro, Francesca, John (Jordan and Evermann 1896; Bigelow Mari ggle, Liza, matajuelo real, and Schroeder 1953; Briggs 1958; piojo, skipjack, tenpounder (El dred Berra 1981). Marine and brackish and Lyons 1966; Jordan and Evermann estuarine (Eldred and Lyons 1966; 19691 Nel son 1984). Class ...... Osteichthyes Order ...... Scientific name .. Megalops at1anticus Family ...... Elopi dae Valenciennes (Rob-. 1980) T. PETERSBURG

PALM BEACH

FLORIDA

CAPE ROMAN

THOUSAND ISLANDS

MILES

KILOMETERS

Figure 2. Ladyfish and tarpon are distributed along the entire coast of South Florida in the Continental Shelf and brackish estuarine waters; tarpon ascend rivers into freshwater also. Preferred common name ...... tarpon larger mouth; the jaw extends con- (Figure 1B) siderably posterior to the rear edge Other common names ...... big scale, of the orbit (Bigel ow and Schroeder caffum, grande ecail ley grande ecoy, 1953). The belly is not keeled or jewfish, sabalo, sabi lo real, serrated as in herrings, but is rela- sadina, saval 1e, saval 1o, saval o- ti vely broad and covered with ordinary real , savani 11 a, si1 ver fish, si1 ver scales (Jordan and Evermann 1969). king, tarpom, tarpum (Gill 1907; Hildebrand 1937; Babcock 1951; Wade 1962a; Jordan and Evermann 1969) The following description of the Class ...... Osteichthyes El opi dae (incl udi ng the tarpon is Order ...... Elopiformes summarized from Jordan and Evermann Family ...... or Megalopidae 11969). Body elongate, somewhat compressed, and covered with silvery cycl oid scales. No scales on head. The tarpon was placed in the Lateral line present. Mouth broad, Elopidae by Gosline (1971) and Robins lower jaw prominent. Premaxill aries et a1 . (1980) , whereas Greenwood et short and nonprotactile; maxi1 laries al. (1966), Forey (1973a, 1973b1, and form lateral margins of the upper jaw. Nelson (1984) recognized the Eye re1ati vely 1arge, with adipose Megalopidae and Elopidae as separate eyelid. Bands of villiform teeth on families within the suborder jaws, vomer, palatines, pterygoids, Elopoidei. The issue is equivocal and tongue, and base of skull. Opercular unl ikely to be resol ved soon. bones thin, with expanded membranous margins. Gill membranes entirely separate and free from the isthmus; Geographic range .... Western Atlantic gillrakers long and slender. Dorsal Ocean from to Brazil and fin inserted over or slightly behind Gulf of Mexico (Figure 2); eastern the pelvics. Caudal fin forked, dor- Atlantic off tropical Africa; chief sal and anal fins depressible into centers of abundance are the West scaly sheaths. No spines or adipose Indies, Florida, and Gulf of Mexico; fin. Very long accessory scales at stragglers recorded from Nova the pectorals and pelvics. Scotia, Bermuda, Argentina, and the Pacific terminus of the Panama Canal (Hi1debrand 1939; Wade 1962a, 1969; Nelson 1984). No evidence exists to Ladyfi sh suggest that tarpon have become es- tablished in the Pacific (Swanson Body very elongate and covered 1946; Wade 1962a). Generally marine with small, thin, silvery scales. or brackish estuarine, but often as- Head small and pointed, with very cends rivers into fresh water 1 arge terminal mouth; maxi 11 ary 1951; Wade 1962a; Robins 1978; Nelson reaches far behind eye. Branchios- 1984). tegal rays 30. Dorsal fin inserted slightly behind the pelvics. Dorsal, anal, and pelvic fin ray counts, 20, 13, and 15, respectively. Caudal MORPHOLOGY/IDENTIFICATION AIDS lobes long and slender. Lateral line straight, with simple pores, 110 to 'The ladyfish and tarpon are both 120 scales. Color silvery a1 1 over herring-1 ike in general appearance but and bluish dorsally, with lower parts are readily distinguished from of sides and ventral surface yellowish clupeids by the presence of an elon- or white. Dorsal and caudal fins gate bony gular plate between the dusky ye1 1owi sh and si1 very. Pel vics branches of the lower jaw and a much and pectoral s speckled, ye1 1owi shy and dusky. Reaches a maximum length of revenues generated by the fishery are about 1 m (usually less than 60 cm) formidable. The ladyfish is also and weight of several kilograms. sought by anglers; it has the sporting Data from Bigelow and Schroeder (1953) attributes of the tarpon, but comes in and Jordan and Evermann (1969). a smaller package suitable for light tackle. Both species are considered inedible in the United States because of the boniness of the flesh, and therefore do not support commercial fisheries. However, they are eaten in Body oblong, compressed, and 1 imited quanti ties elsewhere covered with large, thick, silvery, (Hi1debrand 1939; Babcock 1951). cycloid scales. Mouth large and supe- rior. Branchiostegal rays 23. Dorsal fin with 12 rays, inserted con- siderably behind the pelvics. Anal deeply falcate, 20 rays, about twice LIFE HISTORIES as long as dorsal, has greatly elon- gated last ray. Caudal widely forked Spawning 1ocations of ladyfish and scaly. Lateral line nearly are unknown, but have been inferred to straight, 41 to 48 scales; its tubes be offshore throughout most of the radiate widely over the surface of the range of the species, as judged by the scales. Vertebral counts 53 to 57. locations of capture of early larvae Color bright si1 ver, wi th dorsal sur- (Hi ldebrand 1943; Gehri nger 1959a; face somewhat darker than ventral. Eldred and Lyons 1966). Similarly, Reaches 2 to 2.6 m and over 90 kg. tarpon are believed to Data from Bigel ow and Schroeder throughout most of their range in off- (19531, Jordan and Evermann (1969), shore waters (Wade 1962a; Hi 1debrand and Nelson (1984). 1963; Eldred 1967). Eldred (1967, 1968, 1972) inferred from larval The two species are easily dis- capture locations that spawning took tinguished (Jordan and Evermann 1969). place in the Florida Straits, Gulf The 1adyf i sh has 1arge pseudobranchs Stream, and . Smith (1980) and small scales. The 1ast ray of the provided strong evidence (based on the dorsal is not elongated, and the anal collection of very young larvae) that fin is small er than the dorsal . Con- tarpon spawn off the Caribbean coast versely, the tarpon has large scales of Mexico near Cozumel and Banco and no pseudobranchs. The last ray of Chinchorro (Yucatan Channel ) , off the dorsal is elongated, its free por- west-central Florida, and off the tion being as long as, or longer than, southern part of Veracruz, Mexico. the height of the fin. The anal fin The presence of small larvae off is larger than the dorsal. Georgia (Gehringer 1959b) and North Carolina (Berrien et al. 1978) indi ca tes that spawn ing occurs there also, and probably to some extent REASON FOR INCLUSION IN THIS SERIES along the entire coast from Florida to Cape Hatteras. The tarpon is the premier inshore big-game fish of the Florida coast Fecundity of a tarpon 2 m long (McLane 1974; Robins 1978). Esteemed was estimated to be about 12,200,000 for its stamina, strength, and espe- (Babcock 1951). Sexual maturity is cially its leaping prowess, it is attained at a total length (TL) of avidly sought by anglers. Numerous about 120 cm (Breder 1944). Fecundity annual tournaments are directed and size at sexual maturity of specifically at this species. Tourist 1adyf ish are unknown. Eggs of neither tarpon nor Ladyfish grow to a maximum stand- ladyfish have been described, nor are ard length (SL) of about 40-45 mm yo'l k-sac 1arvae of the 1adyf ish known. during Stage I, shrink to about 18-20 Smith (1980) described and ill ustrated mm SL during Stage 11, and metamor- late yo1 k-sac larvae of tarpon. His phose into juveniles at about 30-35 mm smallest specimen, 5.7 mm in notochord SL at the end of Stage I11 (Hildebrand length (NL), retained only trace 1943; Alikunhi and Rao 1951; Gehringer amounts of yolk, indicating that the 1959a; Jones et a1 . 1978). Durations yolk-sac stage ends at about 6 mm NL. of about 29 days for Stage I1 and 42 Eggs and yo1 k-sac 1arvae that Breder days for Stage I11 were reported by (1944) believed to be tarpon were er- Gehringer (1959a) for larvae reared in roneously identified (El dred 1972). the 1 aboratory. Alikunhi and Rao (1951) reported the duration of Stages Post yo1 k-sac 1arval devel opment I1 and I11 combined as only 9 days in in both species progresses through the 1 aboratory; concurrent field col- three di stinct stages ( termi no1 ogy 1ecti ons provided supporting evidence from Wade 1962a, modified by Jones et for this rapid rate of change. No al. 1978). Stage I is an initial records of water temperature accom- period of length increase that cul- panied the data for either study. minates in the development of a fully formed 1eptocephal us 1arva. The 1ep- tocephal us is characterized by a 1ong, Sizes of Stage I tarpon range ribbon-1 ike, colorless, transparent from 6 mm NL to 28 mm SL (Mercado and body; large fang-1 ike teeth; a very Ciardell i 1972; Smith 1980). Duration small head; and small fins. It lacks of Stage I is estimated to be 2 to 3 gills and red blood cells, and its gut months in the ocean (Smith 1980). is not open (Robins 1978). Oxygen and Larval tarpon shrink to about 14 mm SL nutrients are absorbed through the during Stage I1 and become juveniles skin. In Stage 11, the larva at about 40 mm SL after Stage I11 decreases markedly in length and (Wade 1962a). Duration of Stage I1 gradually loses the ribbon-1 ike lep- was 20-25 days in the laboratory tocephalic morphology. Stage I11 is a (Mercado and Ciardelli 1972). On the second period of length increase that basis of Harrington's (1966) data, we terminates with the beginning of the estimate duration of Stage I11 to be juvenile stage. Late in Stage I1 and about 7-8 weeks . throughout Stage I11 the larva un- dergoes pronounced changes in body form, including increases in body Spawni ng of 1adyfi sh appears to depth, snout 1ength, head length, dor- extend throughout most of the year, sal and anal fin height, and pectoral perhaps peaking in fall, as judged by fin size. Late in Stage 111, the body the occurrence of Stage I larvae. starts to become opaque and silvery. Alikunhi and Rao (1951) collected late Juveniles resemble adul ts in general Stage I 1arvae from October to Decem- appearance. Early life history stages ber in coastal Indian waters. Hil- of tarpon were described by Hildebrand debrand (1943) collected Stage I (1934), Holl ister (1939), Harri ngton 1arvae off Beaufort, North Carol ina, (1958), Gehringer (1959b1, Wade from October through May; off Texas in (1962a1, Eldred (1967, 1968, 19721, February, March, April, and November; Mercado and Ciardell i (19721, Jones et off the Florida Keys in November; and al. (19781, and Smith (1980). off Cuba in May. Offshore collections Descriptions of 1arval and juvenile of Stage I larvae were made by Geh- ladyfish were published by Hildebrand ringer (1959a) off Florida and Georgia (19431, Alikunhi and Rao (19511, Geh- in October, off South Carolina in May, ringer (1959a1, Eldred and Lyons and off North Carolina in November. (1966), and Jones et a1 . (1978). Arnold et al. (1960) collected lep- tocephali from early March to mid-May Snelson 1983) but may ascend rivers near Gal veston, Texas. Tabb and Man- for considerable distances (Tagatz ning (1961) reported that larvae were 1967 1. abundant in Florida Bay from September through December. El dred and Lyons (1966) reported col lecting Stage I Habitats of Stage I tarpon larvae leptocephali off Florida in January, are clear, warm, oceani c waters February, May, June, August, October, (Gehringer 1959b; Robins 1978) withi n and December. 100 m of the surface (Wade 1962a). Surface water temperatures at col 1ec- tion sites ranged from 26.0 to 30.0 "C Summarizing various references on and salinities from 33.6 to 36.0 ppt the occurrences of larval tarpon, (Wade 1962a; Berrien et al. 1978; Robins (1978) and Smith (1980) noted Smith 1980). Estimated temperature that Stage I larvae occur from mid-May and salinity ranges at depth of cap- to late August, and Stage I1 larvae ture were 22.2-28.4 "C and 33.6-36.7 from late June to early October; they ppt (Wade 1962a 1. inferred that spawning occurs in late spring or early summer. Stage I1 and I11 tarpon larvae and juveniles live in and Early Stage 1 larvae of ladyfish mangrove ponds, tidal creeks, rivers, were captured offshore (Gehringer di tches, beaches, and mosqui to-con trol 1959a) at 28.5 ppt and 28.1 "C (Eldred impoundments (Storey and Perry 1933; and Lyons 1966). Late Stage I larvae Breder 1944; Simpson 1954; Moffett and occur inshore (Gehri nger 1959a) at Randal 1 1957; Erdman 1960; Harri ngton 26.3-38.5 ppt and 17.5-29.0 "C (Eldred and Harrington 1960, 1961; Wade 1962a, and Lyons 1966). 01 der early-life 1969; Rickards 1968; Dahlberg 1972; stages (Stage I1 and I11 larvae and Tagatz 1973; Gilmore et al. 1981; juveniles) inhabit coastal beaches, Snelson 1983). These habitats are canal s, bayous, 1agoons, tidal ponds, typically shallow (19.9 " C (Dahlberg salinities: 31.8 ppt (Simpson 19541, 1972); 10-20 ppt and <35 "C (Rose et 18.8-33.4 ppt (Moffett and Randall al. 1975); 5.6-5.8 ppt (Theiling and 1957), 14-45 ppt (Harri ngton 19581, Loyacano 1976); 2.2-6.1 ppt (Govoni 17.5-39.0 ppt (Harri ngton and Har- and Merriner 1978); 0.0-8.8 ppt and rington 19611, 0.0-22.3 ppt (Rickards 16-28 "C (Thompson and Deegan 1982). 19681, 0-47 ppt (Wade 19691, and 15-21 Rose et a1 . (1975) reported a pH range ppt (Gilmore et a1 . 1982). Most lar- of 6.8-8.7 for a coastal impoundment val and juvenile tarpon live at rela- inhabited by juvenile ladyfish. Adult tively high temperatures: 36.6 "C ladyfish usually live in relatively (Moffett and Randall 19571, 36.0 "C open inshore and coastal habitats (Rickards 19681, and 40 "C (Wade (Dahl berg 1972; Gilmore et a1 . 1981; 1969). Because tarpon respire aerially (by gulping air) at least as Florida drainage ditch grew an average early as the beginning of Stage I11 of 1.0 mm/day (range, 0.7-1.4 mm/day) (Harrington 1966 1, 1ow dissol ved from 22 August to 20 October (Moffett oxygen concentrations are not and Randall 1957). Over the same deleterious to survival. The strong period, modal lengths of tarpon in odor of hydrogen sulfide at capture this population increased by 1.4 sites, indicative of poor tidal flush- mm/day. In a Georgia salt marsh, ing, has been reported by various in- juvenile tarpon grew at a rate of vesti gators (e.g., Beebe 1927; Breder about 30 mm/month (Rickards 1968). 1933, 1944; Rickards 1968; Wade 1969). Juvenile tarpon are often collected from isolated marsh ponds that are Breder (1944), who de termi ned connected to the estuary only during growth rates of captive juvenile tar- spring tides. Wade (1969) collected pon, wrote that 12 fish maintained at juvenile tarpon at pH's of 6.8 to 8.2. the old New York Aquarium for 113 to 314 days grew an average of 0.088 mm/day (range, 0.048-0.186 mm/day); In eastern Florida marshes, Wade initial and final length ranges were (1969) found Stage I11 larvae in 94-145 and 110-176 mm TL. Three ditches at the headwaters of small tagged fish (355-365 mm TL) confined creeks. Small juveniles (40-80 mm SL) in natural ponds in southern Florida lived in larger ditches and creeks, did not grow in 133 to 152 days especially in the deeper pools. Large (August to January). Two others juveniles were found in 1arger canals (tagged in July) grew from 345 to 390 and rivers. Juvenile tarpon eventu- and from 370 to 380 mm TL in 258 and ally emigrate from marsh and mangrove 167 days, respectively. Four juvenile habitats and enter coastal waters when tarpon (originally 230-350 mm TL) they reach about 600-800 mm TL (Robins maintained in laboratory pools grew 13 1978). In Georgia, tarpon are unable to 179 mm (mean increment, 97 mm) in to overwi nter in marsh habitats; 15 months; a fifth grew from 436 to juveni 1es left marshes, presumably to 465 mm TL in 6 months. mi grate south, by 1ate October, when they had attained about 160 mm SL (Rickards 1968). Adults live in bays, Ten tarpon raised by Harrington 1agoons, and coastal habitats (Breder (1966) in the laboratory from Stage 1944; Dahl berg 1972; Gilmore et a1 . I11 larvae (18.1-22.7 mm SL; mean, 1981; Snelson 1983) or may cruise the 21.4 mm) for 1 year grew to 55.4-105.3 open ocean (Robins 1978). mm SL (mean, 67.2 mm SL).

GROWTH CHARACTER1 ST1CS A1 though scales of adul t tarpon have distinct rings resembl ing annul i Age and Growth (Breder 1944; Moffett and Randall 19571, these marks have not been Moffett and Randall (19571, who validated as annuli and should be con- examined length-frequency distri bu- sidered with extreme caution. Back- tions of juvenile tarpon from a south calculated mean lengths at the forma- Florida mangrove pond, reported that tion of these putative annuli are modal lengths increased from 75-80 mm shown in Figure 3. Maximum age based FL in early September to 110-115 mm FL on these checks was 16 years (Moffett at the end of the month, and inferred and Randal 1 1957) ; however, 1arger a length increase of about 1.4 mm/day; f ish have been captured. rates declined by about 50% in Oc- tober. Five marked juvenile tarpon Gehringer (1959a reared 1adyfish (301-376 mm FL when tagged) in a south in the laboratory from early Stage I1 to the juvenile phase. Rates of change in standard length during the first part of Stage I1 (from about 35- where W = weight in grams and TL = to- 40 mn to 25 mm SL) averaged -1.061 tal length in millimeters. mm/day . Further shri nking to about 20-21 mm SL proceeded at about -0.342 Harrington (1958) derived the mm/day. Initial length increase following length-wei ght relation for during early Stage 111, from about 20 154 tarpon, 16.0-45.5 mm SL: to 25 mn SL, averaged 0.140 mn/day. Growth rates of late Stage I11 larvae and early juveniles (~60mm SL) were about 0.626 mm/day. Larger juveniles where W = weight in grams and SL = grew an average of 0.628 mm/day. standard length in millimeters. The Field collections suggested a faster relation is valid only for fish within rate of growth (about 2 mn/day) under the stated size range. natural conditions (Gehringer 1959a). No information is available on growth On the basis of a graph presented of adult 1adyfi sh. by Moffett and Randall (19571, we derived the fol 1owing relation between Mor~hometricRe1 ations total length (TL) and fork length (FL) for tarpon: Breder (1944) presented the fol- 1owing 1ength-wei ght relation for TI- = 1.10 FL. adult tarpon in Florida: Harrington (1958 1 developed the fol lowing conversions between fork length (FL), total length (TL), and standard 1ength (SL in mi11 imeters for tarpon 25-54 mn SL:

Sekavec (1974) derived the fol- 1owing length-weight formula from 295 juvenile ladyfi sh 45-201 mn FL from Louisiana:

where W = weight in grams and FL = fork length in millimeters. Mean con- dition factor (K, where K = [W/(FL)'] x 106; Lagler 1956) of these fish was n met eort (Moffon md kndal 1957) 8.1 (range 6.6-8.9). o out mrrt (Monen mna Rmndmll 1857)

FISHERY

0 10 15 NUMBER OF PUTATIVE ANNULI The tarpon and ladyfish fisheries are solely recreational ; no commercial Figure 3. Back-calcul ated mean fishery exists for either species in lengths of tarpon from the west and the United States. Neither species is east coasts of Florida, at putative recorded in the National Marine annuli on scales. Recreational Fisheries Survey and regional catch data are nonexistent copepods and os tracods) and secon- (Grant L. Beardsley, Senior Scientist darily on insects and small fishes; for Recreational Fisheries, National larger juveniles continue to feed on Mari ne Fisheri es Service, Southeast zooplankton, but progressively in- Fisheries Center, Miami, Florida; crease consumption of insects, fishes pers. comm. 1. (especially poecil iids and cyprinodontids), crabs, and grass Ti1mont et a1 . (unpubl ished) sum- of the genus Palaemonetes marized recreational fishery statis- (Beebe 1927; Breder 1933; Mottett and tics for tarpon and ladyfi sh in Randall 1957; Harrington and Har- Evergl ades National Park, Florida, rington 1960, 1961; Rickards 1968). from 1958 through 1984. Tarpon were Juvenile tarpon are typically crepus- sought by less than 3% of anglers and cular and nocturnal foragers (Robi ns made up an average of 0.2% of the 1978). reported recreational catch annual ly. Mean annual catch rates varied from In laboratory settings, early 0.1 to 0.4 fish/h. Less than 10% of Stage I1 ladyfish larvae ate 1ive the tarpon caught were harvested, plankton (Alikunhi and Rao 1951) and primarily for trophy mounts. Tarpon 1ive brine (Artemia) naupl ii accounted for 1% of the catch, 0.4% of (Gehringer 1959a). Stage I11 larvae the harvest, and 7% of the effort in ate small live Fundulus and Gambusia the professional ly guided fishery in and pieces of shrimp and fish. Under the park. Reported catch rates sug- natural conditions, Stage I1 and I11 gested that stocks of tarpon in the ladyfish larvae (<50 mm SL) feed al- park were re1atively stable. most exclusively on zooplankton; con- Ladyfish were commonly caught but sumption of zooplankton by juveniles infrequently harvested by anglers in is progressively reduced as inges tion Everglades National Park (Tilmont et of small fishes and shrimps increases al., unpublished). Ladyfish made up (Harri ngton and Harrington 1961). about 5% of the total reported catch Ladyfish

Adult tarpon feed both noctur- nally and diurnally (Wade 1962a) on a variety of organisms including mu1 lets The tarpon and ladyfish are dis- (Mugil spp.), pinfish ( ti nctly thermophil ic fishes. Both rwdes), ariid catfishes, At1 antic have been reported in col d-re1 ated need1 efi sh (Strongyl ura marina) , sar- fish kills in Florida (Storey and dines (Harengu la spp. ),shrimp, and Gudger 1936; Storey 1937; Snelson and crabs (Babcock 1951; Wade 1962a). Bradley 1978). At Port Aransas, Texas, annual tarpon abundances are Predators correlated with yearly water tempera- ture regimes (Moore 1975). Tarpon Predation by carnivorous concentrate around heated power-pl ant zooplankters and small fishes un- effluents during winter in the Indian doubtedly causes high mortality of eggs River, Florida (Snel son 1983). and 1arvae of both ladyfish and tarpon before the larvae enter coastal nurs- Early Stage I larvae of both ery marshes. In marshes, juvenile species occur only in warm oceanic tarpon are probably immune to piscine waters (22.2-30.0 "C; Wade 1962a; predation other than that by juvenile Eldred and Lyons 1966; Eldred 1967, ladyfish or tarpon (Beebe 1927; Mof- 1968, 1972; Berrien et a1 . 1978; Smith fett and Randall 1957; Rickards 1968; 1980), and it appears probable that Wade 1969). Both species are probably such temperatures are necessary for preyed upon by pisci vorous birds proper development of eggs and early (Beebe 1927; Rickards 19681, and adult 1arvae. tarpon are occasionally eaten by sharks, porpoi ses, and alligators Moffett and Randall (1957) ex- (Wade 1962a, 1962b). posed juvenile tarpon (72-130 mm FL) he1 d at 25-27 " C to high temperatures Parasites in laboratory trials. Fish were warmed from maintenance to test tem- The digenetic trematode peratures in 3 hours. Fish subjected Leci thochirium microstomum occurs in to 39.4-39.6 "C survived the 24-h the stomach of tarpon (Manter 1947). trials; those exposed to 40.5-41.9 "C The isopods Neroci 1a acumi nata and did not. In a series of cold- Cymothoa oestrum are external to1erance trial s, juvenile tarpon (71- parasites (Babcock 1951; Pearse 1952). 130 mm FL) died within 24 h at tem- Causey (1953) reported the cope pod peratures of 14.8-18.0 "C; others sur- Para1 ebi on pearsei from tarpon. The vived trials at 14.8-19.7 " C (Moffett trematode Bivescula tarponi s i s and Randall 1957). Tabb (personal present in the pyloric caecae and communication in Wade 1962a) reported along the entire length of the intes- mortal ity of tarpon in Everglades Na- ti ne ( Sogandares-Bernal and Hutton ti onal Park when water temperature 1959). Though not parasitic, remoras decreased from 24 to 11 "C within a ( Remora remora) are commonly observed few hours. Rickards (1968) col lected attache- arge tarpon (Babcock a sluggish juvenile tarpon in a Georgia 1951; Wade 1962a). salt marsh in late November when water temperature was 16.0 " C. Concur- Trematodes of the genera rently, several juveni 1es mai ntained Bucephal us and Prosorhynchus have been in a steel holding tank died overnight when water temperatures dropped from salinities ranging from 0.0 to 45.0 21.0 to 12.0 OC. However, Wade (1969) ppt (Harrington 1958; Harrington and collected juvenile tarpon at tempera- Harrington 1961; Herke 1969; Dahl berg tures as low as 12 OC, and Gilmore et 1972; Govoni and Merriner 1978; al. (1982) collected tarpon at 14 "C Thompson and Deegan 1982). from a mosquito control impoundment in eastern Florida. Robins (1978) stated Adult ladyfish also tolerate a that the lower lethal temperature of wide range of sali nities, but appear tarpon is about 10 O C. less likely than tarpon to occupy freshwater. We found no reference in Ladyfish appear to be slightly the 1i terature specifically reporting more tolerant of low temperatures than adults from truly freshwater. One of tarpon, judging by the 1ower frequency us (A.V.Z.) has captured ladyfish in of fish-kills (Storey 1937); they have the Lake George section of the St. been collected at temperatures of 11.0 Johns River, Florida, at salinities of to 35.0 OC (Harrington 1958; Tagatz 0.5-2.0 ppt. This area is often as- 1967; Dahlberg 1972; Rose et al. sumea [by compilers evaluating 1975). salinity requirements of fishes) to be freshwater because of its distance Sal ini ty from the mouth of the river (about 190 km) , but numerous salt springs main- Throughout most of their life tain re1atively high salinities. stages, tarpon and ladyfish tolerate a Tagatz (1967) collected ladyfish as wide range of salinities. However, far upstream as Palatka, Florida (135 early Stage I larvae of both species km from the mouth) and reported have been collected only at oceanic salinity there to be 0 ppt; however, sal inites of 28.5-39.0 ppt (Wade he recorded all salinities less than 1962a; Eldred and Lyons 1966; Eldred 1.0 ppt as 0 ppt. It is likely that 1967, 1968, 1972; Berrien et a1 . 1978; appreciable salinities (perhaps >0.5 Smith 1980), and it is likely that ppt) are, if not required, then at such concentrations are required by least preferred by ladyfish. eggs, yo1 k-sac larvae, and early Stage I larvae of both species for proper Dissolved Oxvaen development. Tarpon are obligate air breathers Beyond Stage I, tarpon and ( the swimbl adder contains a1 veol ar 1adyf i s h are decidedly euryhal ine. tissue; Shlaifer 1941) and are Juvenile tarpon can withstand direct frequently seen "rolling" at the sur- transfer from salt - to freshwater and face gulping air; when prevented from vice-versa (Breder 1944; Moffett and reaching the surface, they die within Randal 1 1957 1. Habitats occupied by 7 to 128 h, even in highly oxygenated tarpon range from fresh to hyper- water (Shl aifer 19411. Air breathing saline, or 0 to 47 ppt (Hildebrand is imitatively mediated by visual 1939; Simpson 1954; Moffett and Ran- cues; juveniles in a school come to dall 1957; Harrington 1958; Harrington the surface in rapid succession and Harrington 1961; Rickards 1968; (Shl aifer and Breder 1940; Shl aifer Wade 1969; Dahlberg 1972; Tagatz 1973; 19411, perhaps to reduce indi vidual Tucker and Hodson 1976). suscepti bi1 i ties to predation by fish- eating birds (Kramer and Graham 1976). Alikunhi and Rao (1951) collected The frequency of air breathing is in- late Stage I ladyfish larvae from versely correlated with dissolved brackish water (10.4 ppt) and success- oxygen concentration (Shl aif er and ful ly transferred them directly to Breder 1940; Shlaifer 1941). Air- freshwater (0.08 ppt). Juvenile breathing precludes mortal ity in 1 adyfi sh have been collected at anoxic waters and allows tarpon to survive under conditions deleterious Wet1 ands Destruction and Dearadati on to most fishes. Tarpon have this ability at least as early as the Offshore and coastal habi tats of begi nni ng of Stage II1 (Harrington very young and adult tarpon and 1966). ladyfish are relatively immune to human-i nduced degradation. Con- Air breathing has not been versely, the estuaries, sal t marshes, reported for ladyfish, and it is un- and coastal mangroves used as nur- likely that it occurs. "Rolling" has series by larval and juvenile ladyfish not been reported in the literature. and tarpon in Florida are highly vul- Di ssol ved oxygen requirements of nerable to changes induced by develop- ladyfish are unknown, but it is likely ment. Robins (1978) discussed the that the species is relatively various activities that are degradi ng tolerant of hypoxic conditions, as it tarpon nursery grounds in Florida; his is often found with tarpon in poorly comnents are also applicable to early oxygenated habitats. Ladyfi sh in- 1ife history stages of ladyfish, which habited a coastal impoundment in share these habitats. Among factors Louisiana in which di ssol ved oxygen resulting in the destruction of nurs- concentrations reached a minimum of ery wetlands, he listed filling of 1.0 mg/l (Rose et a1 . 1975). wetl ands, canal ization, bulkheading, construction of water-1 ine right-of- ways and steep-si ded boat-access Contami nants finger-canal s, and impoundment of wet- 1ands for mosquito control. Progress Aerial spraying and ground fog- has recently been made in ameliorating ging for nuisance insect control are the effects of impoundment for widely practiced in Florida ' s coastal mosquito control because impoundment zone, and agricultural . pesticides and does not necessarily result in the herbicides used in south Florida enter destruction of wetl ands. Rather, im- coastal waters. Robins (1978) pounded wetl ands, if properly managed, reported that tarpon are extremely can retain the beneficial characteri s- susceptible to contaminants. Appl ica- tics of natural wetl ands whi 1e provid- tion of dieldrin pellets in a Florida i ng adequate mosquito control salt marsh for the control of 1arval (Clements and Rogers 1964; Provost sandfl ies (Cul icoides) resul ted in 1973). However, access to these wet- mortality of l adytl sh and tarpon lands (and subsequent opportunities (harri ngton and Bid1 ingmayer 1958). for egress) by larval and juvenile tarpon and 1adyfish is precluded or severely curtailed by reduced or non- Turbi di ty exi stent exchange with estuari ne waters (Wade 1969; Gi1 more e t a1 . Stage I larvae of both ladyfish 1982; Harri ngton and Harri ngton 1982). and tarpon occur only in clear off- Improved impoundment management shore waters. Subsequent 1ife history strategies, aimed at enhancing ex- stages appear to be tolerant of high change rates, have been proposed by turbidi ties. Habitats occupied, espe- Clements and Rogers (1964), Provost cially by juveniles, are generally (1973), Montague et a1 . (19851, and described as turbid and dark-stai ned. Lewis et al. (1985). Alikunhi, K.H., and S.N. Rao. 1951. Breder, C.M., Jr. 1933. Young tarpon Notes on the of Elops on Andros Is1and. Bull . N.Y. Zool . saurus Linn. and Mega- SOC. 36 :65-67. cypri noides (Broussonet) wi th obser- vations on their growth. J. Zool. Breder, C.M., Jr. 1944. Materials Soc. India 3:99-109. for the study of the life history of Tarpon at1anticus. Zoologica Arnold, E.L., Jr ., R.S. Wheeler, and K.N. Baxter. 1960. Observations on fishes and other biota of East Briggs, J.C. 1958. A list of Florida Lagoon, Galveston Bay. U.S. Fish fishes and their distribution. Wildl. Serv. Spec. Sci. Rep. No. Bull. Fla. State Mus. 2:223-318. 344. 30 pp. Causey, D. 1953. Parasitic copepods Babcock, L.L. 1951. The tarpon, 5th of Texas. Publ. Inst. Mar. Sci. ed. Privately printed, Buffalo, Univ. Tex. 3:5-16. N.Y. 157 pp.

Clements, B.W., Jr., and A.J. Rogers. Beebe, W. 1927. A tarpon nursery in 1964. Studies of impounding for the Haiti. Bull. N.Y. Zool. Soc. control of salt marsh mosquitos in 30 :141- 145. Florida, 1958-1963. Mosquito News 24: 265-276. Berra, T.M. 1981. An atlas of dis- tri bution of the freshwater fish families of the world. University Corkum, K.C. 1959. Some trematode of Nebraska Press, Lincoln. 197 pp. parasites of fishes from the Missi s- sippi gulf coast. Proc. La. Acad. Sci. 22:17-29. Berrien, P.L., M.P. Fahay, A.W. Ken- dall , Jr., and W.G. Smith. 1978. Ichthyoplankton from the RV Do1 phin Dahl berg, M. D. 1972. An ecol ogi cal survey of the ContinentalTiTT study of Georgia coastal fishes. waters be tween Martha ' s Vineyard, U.S. Natl. Mar. Fish. Serv. Fish. Massachusetts and Cape Lookout, Bull . 70 :323-353. North Carol ina, 1965-66. Natl . Mar. Fish. Serv. Sandy Hook Lab. Tech. Darnell, R.M. 1958. Food habits of Ser. Rep. No. 15. 152 pp. fishes and 1 arger invertebrates of Lake Pontchartrain, Louisiana, an estuarine community. Publ . Inst. Bigel ow, H.B., and W.C. Schroeder. Mar. Sci. Univ. Tex. 5:354-416. 1953. Fishes of the Gulf of Maine. U.S. Fish Wildl. Serv. Fish. Bull. Eldred, B. 1967. Larval tarpon, 53. 577 pp. Mega1 ops at1 anticus Valenciennes, (Mega1 opidae) in Florida waters. Gill, T. 1907. The tarpon and 1ady- Fla. Board Conserv. Mar. Lab. Leaf1 . fish and their relatives. Smith- Ser. 4(4). 9 pp. sonian Misc. Coll . 48:31-46.

Eldred, B. 1968. First record of a Gilmore, R.G., D.W. Cooke, and C.J. 1arval tarpon, Mega1 ops at1anticus Donohoe. 1982. A comparison of the Valenciennes, from the Gult of fish populations and habitat in open Mexico. Fla. Board Conserv. Mar. and closed salt marsh impoundments Lab. Leafl. Ser. 4(7). 2 pp. in east-central Florida. Northeast Gulf Sci. 5:25-37. Eldred, B. 1972. Note on larval tar- pon, Mega1 ops at1anticus (Mega- Gilmore, R.G., Jr., C.J. Donohoe, D.W. 1 opidae) in thetraits. Cooke, and D.J. Herrema. 1981. Fla. Dep. Nat. Resour. Mar. Res. Fishes of the Indian River Lagoon Lab. Leafl. Ser. 4(22). 6 pp. and adjacent waters, Florida. Har- bor Branch Found. Tech. Rep. No. 41. Eldred, B., and W.G. Lyons. 1966. 36 PP. Larval ladyfish, Elops saurus Lin- naeus 1766, (El opidaelTTFIorida Gosl ine, W.A. 1971. Functional mor- and adjacent waters. Fla. Board phology and classification of Conserv. Mar. Lab. Leafl . Ser. 4(2). teleostean fishes. University Press 6 PP. of Hawaii , Honol ulu. 208 pp. Erdman, D.S. 1960. Larvae of tarpon, Govoni, J.J., and J.V. Merriner. Mega1 ops at1antica, from the Anasco 1978. The occurrence of ladyfish, River, Puerto Rico. Copeia Elops saurus, 1arvae in low salinity 1960:146. watermd another record for Chesapeake Bay. Estuaries 1:205- Forey, P.L. 1973a. Re1 ationships of 206. elopomorphs. Zool. J. Linn. Soc. 53( Supple 1) :351-368. Greenwood, P.H., D.E. Rosen, S.H. Weitzman, and G.S. Myers. 1966. Forey, P.L. 1973b. A revision of the Phyl etic studies of teleostean elopiform fishes, fossil and recent. fishes with a provisional class- Bull. Br. Mus. Nat. Hist. (Geol.), ification of living forms. Bull. Suppl . 1O:l-222. Am. Mus. Nat. Hist. 131:339-455. Harrington, R.W., Jr. 1958. Mor- Fyfe, J.L. 1986. Trophic analysis of phometry and ecology of small tar- six species of fishes collected from pon, Mega1 ops at1antica Valenciennes a subtropical salt marsh from east from transitional stage through on- central Florida, U.S.A. Fla. Sci. set of scale formation. Copeia 49(S~ppl.1) :18. 1958: 1-10.

Gehringer, J. W. 1959a. Early Harri ngton, R.W., Jr. 1966. Changes development and metamorphosis of the through one year in the growth rates ten-pounder, El ops saurus Linnaeus. of tarpon, Mega1 ops at1anticus U.S. Fish. WiiServ.ish. Bull. Valenciennes, reared trom mi d- 59:619-647. metamorphosis. Bul l. Mar. Sci . 16 ~863-883. Gehringer, J .W. 1959b. Leptocephal us of the At1 antic tarpon, Mega1 ops at- 1anticus Valenciennes, fTEi3TZhore Harrington, R.W., Jr., and W.L. Bid- waters. Q. J. Fla. Acad. Sci. 1 ingmayer. 1958. Effects of 21 :235-240. dieldrin on fishes and invertebrates of a salt marsh. J . Wildl . Manage. India, including a study of the 22:76-82. caudal skeleton and a comparison with the Atlantic species, Tarpon Harrington, R.W., Jr., and E.S. Har- at1anticus. Zool ogica 24:449-475. rington. 1960. Food of 1arval and young tarpon, Mega1 ops at1antica. Copeia 1960:311-319. Jones, P.W., F.D. Martin, and J.D. Harrington, R.W., Jr., and E.S. Har- Hardy, Jr. 1978. Development of rington. 1961. Food selection fishes of the Mid-Atlantic Bight: an among fishes invading a high sub- atlas of egg, larval and juvenile tropical salt marsh: from onset of stages. Vol. 1: Acipenseridae flooding through the progress of a through Ictaluridae. U.S. Fish mosquito brood. Ecology 42:646-666. Wildl. Serv. Biol. Serv. Program FWS/OBS-78/12. 366 pp. Harrington, R.W., Jr., and E.S. Har- rington. 1982. Effects on fishes Jordan, D.S., and B.W. Evermann. and their forage organisms of im- 1896. The fishes of North and pounding a Florida salt marsh to Middle America. Bull. U.S. Natl. prevent breeding by salt marsh Mus., NO. 47 (1):l-1240. mosqui tos. Bull . Mar. Sci . 32:523- 531. Jordan, D.S., and B.W. Evermann. Herke, W.H. 1969. An unusual inland 1969. American food and game col 1ec ti on of 1arval 1adyf ish, El ops fishes. Dover Pub1 ications, Inc., saurus in Louisiana. Proc.7 New York. 574 pp. Acad.ci . 32 :29-30. Knapp, F.T. 1949. Menhaden utiliza- Hildebrand, S.F. 1934. The capture ti on in relation to the conservation of a young tarpon, Tarpon atlan- of food and game fishes of the Texas ticus, at Beaufort, North Carolina. gulf coast. Trans. Am. Fish. Soc. Copeia 1934:45-46. 79 :137-144.

Hildebrand, S.F. 1937. The tarpon in Kramer, D.L., and J.B. Graham. 1976. the Panama Canal. Sci. Month. Synchronous air breathing, a social 44 :239-248. component of respiration in fishes. Copei a 1976 :689-697. Hildebrand, S.F. 1939. The Panama Canal as a passageway for fishes, Lagler, K.F. 1956. Freshwater with 1ists and remarks on the fishes fishery biology. W.C. Brown Co., and i nvertebrates observed. Dubuque, Iowa. 421 pp. Zool ogica 24: 15-45. Lewis, R.R., 111, R.G. Gilmore, Jr., Hildebrand, S.F. 1943. Notes on the D.W. Crewz, and W.E. Odum. 1985. affinity, anatomy and development of Mangrove habitat and fishery El ops saurus Linnaeus. J. Wash. resources of Florida. Pages 281-336 E'ZX ET33:90-94. in W.S. Seaman, Jr., ed. Florida muatic Habitat and Fishery Hi1 debrand, S.F. 1963. Family Resources. Florida Chapter American Elopidae. Pages 111-131 in Fishes Fisheries Society, Gainesville. of the western North mantic. Sears Found. Mar. Res. Mem. l(3). Linton, E. 1904. Parasites of fishes Hol li ster, G. 1939. Young Megalops of Beaufort, North Carolina. Bull. cypri noi des from Batavi a, Dutch tast U.S. Bur. Fish. 24:321-428. Manter, H.W. 1947. Digenetic Timbers Conf. Ecol. Anim. Control trematodes of marine fishes. Am. Habitat Manage. 5:5-17. Mid1 . Nat. 38:339-340. Rickards, W.L. 1968. Eccrlogy and McLane, A.J., ed. 1974. McLanes's growth of juvenile tarpon, Mega1 ops new standard fishing encycl opedia atlanticus, in a Georgia salt marsh. and international angling guide, 2nd Bull . Mar. Sci . 18:220-239. ed. Hol t, Rinehart and Winston, N.Y. 1,156 pp. Robins, C.R. 1978. The tarpon - unusual bi01 ogy and man's impact Mercado, J .E., and A. Ciardell i. determine its future. Mar. Recrea- 1972. Contri bucion a 1a morfologia tional Fish. 2:105-112. y organogenesis de 1os 1eptocefal os del sabal o Mega1 ops at1anticus Rebins, C.R., R.M. Bailey, C.E. Bond, (Pisces: Megalopidae). Bull. Mar. J.R. Brooker, E.A. Lachner, R.N. Sci. 22:153-184. Lea, and W.B. Scott. 1980. A list of the common and scientific names Moffett, A.W., and J.E. Randall. of fishes from the United States and 1957. The Roger Firestone tarpon Canada, 4th ed. Am. Fish. Soc. investigation. Univ. Miami Mar. Spec. Publ. 12. Bethesda, Md. 174 Lab. Progr. Rep. 57-22. 18 pp. PP

Montague, C.L., A.V. Zale, and H.F. Rose, C.D., A.H. Harris, and B. Wil- Percival . 1985. A conceptual model son. 1975. Extensive culture of of salt marsh management on Merritt penaeid shrimp in Louisiana salt- Island National Wildlife Refuge, marsh impoundments. Trans. Am. Florida. Fla. Coop. Fish Wi1 dl . Fish. Soc. 104:296-307. Res. Unit Tech. Rep. 17. Gaines- ville. 92 pp. ~ekavec, G.B. 1971. Gross morphology of the digestive tract of the Moore, R.H. 1975. Occurrence of ladyfish, Elops saurus. Chesapeake tropical marine fishes at Port Aran- Sci . 12:275-276. - sas, Texas 1967-1973, related to sea temperatures. Copei a 1975 :170-172. Sekavec, G.B. 1974. Summer foods, length-wei ght re1ationshi p, and con- Nelson, J.S. 1984. Fishes of the di ti on factor of juvenile ladyfish, world, 2nd ed. John Wiley and Sons, El ops saurus Linnaeus, from New York. 523 pp. LouTs'iana coastal streams. Trans. Am. Fish. Soc. 103:472-476. Pearse, A.S. 1952. Parasitic crus- tacea from the Texas coast. Publ . Shl aifer, A. 1941. Addi ti onal social Inst. Mar. Sci. Univ. Tex. 2:5-42. and physi 01 ogi cal aspects of respiratory behavior in small tar- Pfeiler, E. 1986. Towards an ex- pon. Zoo1 ogi ca 26 :55-60. planati on of the devel opmental strategy in leptocephalous larvae of Shlaifer, A., and C.M. Breder, Jr. marine fishes. Envi ron. 1940. Social and respiratory be- Bi 01. Fishes 15:3-13. havior of small tarpon. Zoologica 25 ~493-512.

Provost, M.W. 1973. Salt marsh Simpson, D.G. 1954. Two small tarpon management in Florida. Proc. Tall from Texas. Copei a 1954:71-72. Smith, D.G. 1980. Early larvae of Tagatz, M.E. 1967. Fishes of the St. the tarpon, Mega1 ops at1antica Johns River, Florida. Q. J. Fla. Valenciennes [Pisces: Elopidael , Acad. Sci. 30:25-50. with notes on spawning in the Gulf of Mexico and the Yuca.tan Channel. Tagatz, M.E. 1973. A larval tarpon, Bull . Mar. Sci . 30: 136-141. Megalops at1anticus, from Pensacola, k l orida. Copei a 19/3 :140- 141.

Snelson, F.F., Jr. 1983. Ichthyo- fauna of the northern part of the Theil ing, D.L., and H.A. Loyacano, Jr. Indian River Lagoon system, Florida. 1976. Age and growth of red drum Fla. Sci . 46 :187-206. from a saltwater marsh impoundment in South Carolina. Trans. Am. Fish. SOC. 105 ~41-44. Snelson, F.F., Jr., and W.K. Bradley, Jr. 1978. Mortality of fishes due to cold on the east coast of Thompson, B.A., and L.A. Deegan. Florida, January, 1977. Fla. Sci. 1982. Distribution of ladyfish 41 :1-12. (El ops saurus and bonefi sh (A1 bul a vulpes)ocephal i in Louisiana. Sogandares-Bernal , F ., and R. F. Hut- Bull.Rar. Sci . 32 :936-939. ton. 1959. Bivescula tarponis, a new trematode in the tarpon Mega1ops Ti lmont, J., E. Rutherford, R. Dawson, atlanticus (Cuv. and Val. ) 'from the and E. Thue. Unpublished. An anal - west coast of Florida. J. ysis of the recreational and comner- Parasi to1 . 45:114-118. cia1 estuarine fisheries harvest wi thin Evergl ades National Park. Storey, M. 1937. The relation be- tween normal range and mortality of Tucker, J.W., Jr., and R.G. Hodson. fishes due to cold at Sanibel Is- 1976. Early and mid-metamorphic 1 and, Flori da. Ecology 18 :10-26. 1 arvae of the tarpon, Mega1 ops at- lantica, from the Cape kear Rim Storey, M., and E.W. Gudger. 1936. estuary, North Carol ina, 1973-74. Mortality of fishes due to cold at Chesapeake Sci . 17: 123-125. Sani be1 Is1and, Florida, 1886-1936. Ecol ogy 17 :640-648. Wade, R.A. 1962a. The biology of the tarpon, Mega1 ops at1anticus, and the Storey, M., and L.M. Perry. 1933. A ox-eye, Mega1 ops cyprinoides, with record of young tarpon at Sanibel emphasi s on 1 arva l development. Island, Lee County, Flori da. Bull. Mar. Sci. Gulf Caribb. 12: 545- Science 78 :284-285. 622.

Swanson, P.L. 1946. Tarpon in the Wade, R.A. 1962b. 'The elusive tar- Pacific. Copeia 1946:175. pon. Sea Frontiers 8:258-267.

Wade, R.A. 1969. Ecology of juvenile Tabb, D.C., and R.B. Manning. 1961. tarpon and effects of dieldrin on A check1 ist of the flora and fauna two associated species. U.S. Fish. of northern Florida Bay and adjacent Wildl. Serv. Tech. Pap. 41. 85 pp. brackish waters of the Florida main- land collected during the period Zilberberg, M.H. 1966. Seasonal oc- July, 1957 through September, 1960. currence of fishes in a coastal Bull. Mar. Sci. Gulf Carib. 11:552- marsh of northwest Florida. Con- 649. trib. Mar. Sci. 11:126-134. so272 - 101 3. Rulplent's Acenslon No. REPORT WCUMENTAllON 1. RE*m t PAGE I Biological Report 82(11.104)*I 4. Title and Subtle 5. Report Dote Species Profiles: Life Histories and Environmental Requirements July 1989 of Coastal Fishes and Invertebrates (South Florida) -- Ladyfish and Tarpon 7. Author(,) 8. Performing Or#onlration Rept. No. Alexander V. Zale and Susan G. Merrifield 9. Performing Omniratlon Name and Mdms 10. Rol.ct/TasklWorlc Unit No. Oklahoma Cooperative Fish and Wild1 ife Research Unit 404 Life Sciences Nest 11. Mnet(C) or GrantfG) No. Oklahoma State University (C) Stillwater, OK 74078 (GI

12 Sponsoring Omnlzation Name and Addms 1% Typa of Report & Period Cowred National Wetlands Research Center U.S. Army Corps of Engineers Fish and Wildlife Service Waterways Experiment Station U.S. Dept. of the Interior P.O. Box 631 Washington, DC 20240 Vicksburg, MS 39180 IS. Supplementary Not.. * U.S. Army Corps of Engineers Report No. TR EL-82-4

16. Abrtract (Limit: 200 word.)

Species profiles are literature summaries of the taxonany, morpholoqv, distribu- tion, life history, habitats, and environmental requiremnts of coastal species of fishes and aquatic invertebrates. They are designed to assist in environ- mental impact assessment. The tarpon and ladyfish are popular gamfishes. Adults spawn offshore. Larval and juvenile stages inhabit coastal marshes and mangroves. Both species are thermophilic (preferring warm water), euryhaline (tolerant of a wide range of salinity), and are capable of surviving at low oxygen concentrations. Wetlands destruction and degradation negatively affect these species by reducing nursery areas.

17. Document Analysis a. Dorripton Fi shes Fisheries Life cycles Estuaries Temperature Feeding habits Sal ini ty Growth Oxygen b. Identitienl0p.n-Ended Terms Tarpon Trophic ecology Megalops atlanticus Spawni ng Ladyfish Envi ronmental requi rements Elops saurus

I la Availablllty Stakmen: I 19. Suurlty Class (This Report) 1 21. No. of PWW I 17 Unlimited Unclassified 20. Suurlty Class (This Pago) n. PAC. Unclassified OFrIONAL FORM 272 (4-77) (Formerly NTiS35) Dopartm.nt of Commerce As the Nation's principal conservation agency, the Department of the Interior has responsibility for most of our nationally owned public lands and natural resources. This includes fostering the wisest use of our land and water resources, protecting our fish and wildlife, preserving the environmental and cultural values of our national parks and historical places, and providing for the enjoy- ment of life through outdoor recreation. The Department assesses our energy and mineral resources and works to assure that their development is in the best interests of all our people. The Depart- ment also has a major responsibility for American Indian reservation communities and for people who live in island territories under U.S. administration.

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