Biological Report 82P1104)--' T R EL.82.4. /c0.-July 1989.

0t

Species Profiles: Life Histories andEnvironmental Requirements of Coastal Fishes DTand Invertebrates (South Florida) P 10 8 1989

LADYFISH AND TARPON

0

N

Coastal Ecology Group*Fish and Wildlife Service Waterways Experiment Station

U.S. Department of the Interior U.S. Army Corps of EngineersS DI. 'T;r'! S ,.'fc,,'A T NNT A

A ~~~ ~ 0 0:~8 ; )87"

Biological Report 82(11.104)TR EL-82-4July 1989

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

LADYFISH AND TARPON

by

Alexander V. Zaleand

Susan G. Merrifield

Oklahoma Cooperative Fish and Wildlife Research UnitDeparbent of Zoology

404 Life Sciences WestOklahoma State University

Stillwater, OK 74078

Project OfficerDavid tbran

National Wetlands Research CenterU.S. Fish and Wildlife Service

1010 Gause BoulevardSlidell, LA 70458

Performed for

Coastal Ecology GroupWaterways Experiment StationU.S. Army Ccrps of Engineers

Vicksburg, MS 39180

and

National Wetlands Research CenterResearch and DevelopmentFish and Wildlife Service

U.S. Department of the InteriorWashington, DC 20240

This series may be referenced as follows:

U.S. Fish and Wildlife Service. 1983-19 . Species profiles: life historiesand environmental requirements of coast a fishes and invertebrates. U.S. FishWildl. 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 andenvironmental requirements of coastal fishes and invertebrates (South Florida)-- ladyfish and tarpon. U.S. Fish Wildl. Serv. Biol. Rep. 82(11.104). U.S.Arnly Corps of Engineers, TR EL-82-4. 17 pp.

PREFACE

This species profile is one of a series on coastal aquatic organisms,principally fish, of sport, commercial, or ecological importance. The profilesare designed to provide coastal managers, engineers, and biologists with a briefcomprehensive sketch of the biological characteristics and environmentalrequirements of the species and to describe how populations of the species may beexpected to react to environmental changes caused by coastal development. Eachprofile has sections on taxonomy, life history, ecological role, environmentalrequirements, and economic importance, if applicable. A three-ring binder isused 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 Engineersand the U.S. Fish and Wildlife Service.

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

Information Transfer SpecialistNational Wetlands Research CenterU.S. Fish and Wildlife ServiceNASA-Slidell Computer Complex1010 Gause BoulevardSlidell, LA 70458

or

U.S. Army Engineer Waterways Experiment StationAttention: WESER-CPost Office Box 631Vicksburg, MS 39180

F on, For

By_. t o n

CONVERSION TABLE

Metric to U.S. Customary

Multiply By To Obtain

millimeters (mm) 0.03937 inchescentimeters (cm) 0.3937 inchesmeters (m) 3.281 feetmeters (m) 0.5468 fathomskilometers (km) 0.6214 statute mileskilometers (km) 0.5396 nautical miles

square meters (m2 ) 10.76 square feetsquare kilometers (km2) 0.3861 square mileshectares (ha) 2.471 acres

liters (1) 0.2642 gallonscubic meters (m

3) 35.31 cubic feet

cubic meters (m3) 0.0008110 acre-feet

milligrams (mg) 0.00003527 ouncesgrams (g) 0.03527 ounceskilograms (kg) 2.205 poundsmetric tons (t) 2205.0 poundsmetric tons (t) 1.102 short tons

kilocalories (kcal) 3.968 British thermal units

Celsius degrees (0C) 1.8(OC) + 32 Fahrenheit degrees

U.S. Customary to Metric

inches 25.40 millimetersinches 2.54 centimetersfeet (ft) 0.3048 metersfathoms 1.829 metersstatute miles (mi) 1.609 kilometersnautical miles (nmi) 1.852 kilometers

square feet (ft2 ) 0.0929 square meterssquare miles (mi2 ) 2.590 square kilometersacres 0.4047 hectares

gallons (gal) 3.785 literscubic feet (ft3 ) 0.02831 cubic metersacre-feet 1233.0 cubic meters

ounces (oz) 28350.0 milligramsounces (oz) 28.35 gramspounds (lb) 0.4536 kilogramspounds (lb) 0.00045 Petric tonsshort tons (ton) U.9072 metric tons

British thermal units (Btu) 0.2520 kilocaloriesFahrenheit degrees (OF) 0.5556 (OF - 32) Celsius degreesiv

CONTENTS

PagePREFACE ............................................................... iiiCONVERSION TABLE ....................................................... ivACKNOWLEDGMENTS........................................................ vi

NOMENCLATURE/TAXONOMY/RANGE .............................................. 1MORPHOLOGY AND IDENTIFICATION AIDS....................................... 3

Ladyfish ............................................................. 3Tarpon............................................................... 4

REASON FOR INCLUSION IN THIS SERIES...................................... 4LIFE HISTORIES ......................................................... 4GROWTH CHARACTERISTICS.................................................. 7

Age and Growth ....................................................... 7Morphometric Relations ................................................. 8

FISHERY................................................................ 8ECOLOGICAL ROLE ........................................................ 9

Feeding Behavior/Food Habits ........................................... 9Predators............................................................ 10Parasites............................................................ 10

ENVIRONMENTAL REQUIREMENTS.............................................. 10Temperature.......................................................... 10Salinity............................................................. 11Dissolved Oxygen ..................................................... 11Contaminants......................................................... 12Turbidity............................................................ 12Wetlands 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, EdmonLow Library, Oklahoma State University, provided exceptioial assistance withliterature searches and inter-library loan. H. Franklin Percival of the FloridaCooperative Fish and Wildlife Research Unit, University of Florida, Gainesville,and 0. Eugene Maughan of the Oklahoma Unit provided administrative assistance.

vi

1A

Figure 1. A: ladyfish; B: tarpon.

LADYFISH AND TARPON

NOMENCLATURE/TAXONOMY/RANGE Geographic range .... western AtlanticOcean from Bermuda and southern New

Scientific name ......... Elops saurus England (but uncommon north of CapeLinnaeus (Robins et al. TW T Hatteras) to Rio de Janeiro, Brazil,

Preferred common name ....... ladyfish and throughout the Gulf of Mexico(Figure A) (Figure 2); also occurs in the In-

Other common names .. bigeyed herring, dian and western Pacific Oceansbony-fish, chiro, Francesca, John (Jordan and Evermann 1896; BigelowMariggle, Liza, matajuelo real, and Schroeder 1953; Briggs 1958;piojo, skipjack, tenpounder (Eldred Berra 1981). Marine and brackishand Lyons 1966; Jordan and Evermann estuarine (Eldred and Lyons 1966;1969) Nelson 1984).

Class ................... OsteichthyesOrder .................... Elopiformes Scientific name .. Megalops atlanticusFamily ...................... Elopidae Valenciennes (Roblns et a--T]B1 -

01

CAPE A TLANT/CCANAVERAL OCEAN

\\70 TAMPA

K ~ ST. PETERSBURG

KEECHOBEE

PALM BEACH

GULF OFFLRDMEXICOFLRD

0MIAMICAPE ROMANOb

THOUSAND ISLANDSjAREA

PAIL ES F .A BA-

KILOMETERS 1-e 00 .~ '

Figure 2. Ladyfish and tarpon are distributed along the entire coast of SouthFlorida in the Continental Shelf and brackish estuarine waters; tarpon ascendrivers into freshwater also.

2

Preferred common name ......... tarpon larger mouth; the jaw extends con-(Figure IB) siderably posterior to the rear edge

Other common names ........ big scale, of the orbit (Bigelow and Schroedercaffum, grande ecaille, grande ecoy, 1953). The belly is not keeled orjewfish, sabalo, sabilo real, serrated as in herrings, but is rela-sadina, savalle, savallo, savalo- tively broad and covered with ordinaryreal, savanilla, silver fish, silver scales (Jordan and Evermann 1969).king, tarpom, tarpum (Gill 1907;Hildebrand 1937; Babcock 1951; Wade1962a; Jordan and Evermann 1969) The following description of the

Class ................... Osteichthyes Elopidae (including the tarpon) isOrder .................... Elopiformes summarized from Jordan and EvermannFamily ....... Elopidae or Megalopidae (1969). Body elongate, somewhat

compressed, and covered with silverycycloid 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. Premaxillarieset al. (1980), whereas Greenwood et short and nonprotactile; maxillariesal. (1966), Forey (1973a, 1973b), and form lateral margins of the upper jaw.Nelson (1984) recognized the Eye relatively large, with adiposeMegalopidae and Elopidae as separate eyelid. Bands of villiform teeth onfamilies within the suborder jaws, vomer, palatines, pterygoids,Elopoidei. The issue is equivocal and tongue, and base of skull. Opercularunlikely to be resolved soon. bones thin, with expanded membranous

margins. Gill membranes entirelyseparate and free from the isthmus;

Geographic range .... Western Atlantic gillrakers long and slender. DorsalOcean from Virginia to Brazil and fin inserted over or slightly behindGulf of Mexico (Figure 2); eastern the pelvics. Caudal fin forked, dor-Atlantic off tropical Africa; chief sal and anal fins depressible intocenters of abundance are the West scaly sheaths. No spines or adiposeIndies, Florida, and Gulf of Mexico; fin. Very long accessory .cales atstragglers recorded from Nova the pectorals and pelvics.Scotia, Bermuda, Argentina, and thePacific terminus of the Panama Canal(Hildebrand 1939; Wade 1962a, 1969;Nelson 1984). No evidence exists to Ladyfishsuggest that tarpon have become es-tablished in the Pacific (Swanson Body very elongate and covered1946; Wade 1962a). Generally marine with small, thin, silvery scales.or brackish estuarine, but often as- Head small and pointed, with verycends rivers into fresh water large terminal mouth; maxillary1951; Wade 1962a;Robins 1978; Nelson reaches far behind eye. Branchios-1984). tegal ray 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 linestraight, with simple pores, 110 to

The ladyfish and tarpon are both 120 scales. Color silvery all overherring-like in general appearance but and bluish dorsally, with lower partsare readily distinguished from of sides and ventral surface yellowishclupeids by the presence of an elon- or white. Dorsal and caudal finsgate bony gular plate between the dusky yellowish and silvery. Pelvicsbranches of the lower jaw and a much and pectorals speckled, yellowish, and

03

dusky. Reaches a maximum length of revenues generated by the fishery areabout 1 m (usually less than 60 cm) formidable. The ladyfish is alsoand weight of several kilograms. sought by anglers; it has the sportingData from Bigelow and Schroeder (1953) attributes of the tarpon, but comes inand Jordan and Evermann (1969). a smaller package suitable for light

tackle. Both species are consideredinedible in the United States becauseof the boniness of the flesh, and

Tarpon therefore do not support commercialfisheries. However, they are eaten in

Body oblong, compressed, and limited quantities elsewherecovered with large, thick, silvery, (Hildebrand 1939; Babcock 1951).cycloid scales. Mouth large and supe-rior. Branchiostegal rays 23. Dorsalfin with 12 rays, inserted con-siderably behind the pelvics. Analdeeply falcate, 20 rays, about twice LIFE HISTORIESas long as dorsal, has greatly elon-gated last ray. Caudal widely forked Spawning locations of ladyfishand scaly. Lateral line nearly are unknown, but have been inferred tostraight, 41 to 48 scales; its tubes be offshore throughout most of theradiate widely over the surface of the range of the species, as judged by thescales. Vertebral counts 53 to 57. locations of capture of early larvaeColor bright silver, with dorsal sur- (Hildebrand 1943; Gehringer 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 spawnData from Bigelow and Schroeder throughout most of their range in off-(1953), Jordan and Evermann (1969), shore waters (Wade 1962a; Hildebrandand Nelson (1984). 1963; Eldred 1967). Eldred (1967,

1968, 1972) inferred from larvalThe two species are easily dis- capture locations that spawning took

tinguished (Jordan and Evermann 1969). place in the Florida Straits, GulfThe ladyfish has large pseudobranchs Stream, and Caribbean. Smith (1980)and small scales. The last ray of the provided strong evidence (based on thedorsal is not elongated, and the anal collection of very young larvae) thatfin is smaller than the dorsal. Con- tarpon spawn off the Caribbean coastversely, the tarpon has large scales of Mexico near Cozumel and Bancoand no pseudobranchs. The last ray of Chinchorro (Yucatan Channel), offthe dorsal is elongated, its free por- west-central Florida, and off thetion 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 offis larger than the dorsal. Georgia (Gehringer 1959b) and North

Carolina (Berrien et al. 1978)indicates that spawning occurs therealso, and probably to some extent

REASON FOR INCLUSION IN THIS SERIES along the entire coast from Florida toCape Hatteras.

The tarpon is the premier inshorebig-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 iscially 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 ladyfish are unknown.

4

Eggs of neither tarpon nor Ladyfish grow to a maximum stand-ladyfish have been described, nor are ard length (SL) of about 40-45 mmyolk-sac larvae of the ladyfish known. during Stage I, shrink to about 18-20Smith (1980) described and illustrated mm SL during Stage II, and metamor-late yolk-sac larvae of tarpon. His phose into juveniles at about 30-35 mmsmallest specimen, 5.7 mm in notochord SL at the end of Stage Ill (Hildebrandlength (NL), retained only trace 1943; Alikunhi and Rao 1951; Gehringeramounts of yolk, indicating that the 1959a; Jones et al. 1978). Durationsyolk-sac stage ends at about 6 mm NL. of about 29 days for Stage II and 42Eggs and yolk-sac larvae that Breder days for Stage III were reported by(1944) believed to be tarpon were er- Gehringer (1959a) for larvae reared inroneously identified (Eldred 1972). the laboratory. Alikunhi and Rao

(1951) reported the duration of StagesPost yolk-sac larval development II and III combined as only 9 days in

in both species progresses through the laboratory; concurrent field col-three distinct stages (terminology lections provided supporting evidencefrom Wade 1962a, modified by Jones et for this rapid rate of change. Noal. 1978). Stage I ib 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 fullyformed leptocephalus larva. The lep-tocephal6s is characterized by a long, Sizes of Stage I tarpon rangeribbon-like, colorless, transparent from 6 mm NL to 28 mm SL (Mercado andbody; large fang-like teeth; a very Ciardelli 1972; Smith 1980). Durationsmall head; and small fins. It lacks of Stage I is estimated to be 2 to 3gills 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 SLnutrients are absorbed through the during Stage II and become juvenilesskin. In Stage II, the larva at about 40 mm SL after Stage IIIdecreases markedly in length and (Wade 1962a). Duration of Stage IIgradually loses the ribbon-like lep- was 20-25 days in the laboratorytocephalic morphology. Stage Ill is a (Mercado and Ciardelli 1972). On thesecond period of length increase that basis of Harrington's (1966) data, weterminates with the beginning of the estimate duration of Stage III to bejuvenile stage. Late in Stage II and about 7-8 weeks.throughout Stage III the larva un-dergoes pronounced changes in bodyform, including increases in body Spawning of ladyfish appears todepth, snout length, head length, dor- extend throughout most of the year,sal and anal fin height, and pectoral perhaps peaking in fall, as judged byfin size. Late in Stage III, the body the occurrence of Stage I larvae.starts to become opaque and silvery. Alikunhi and Rao (1951) collected lateJuveniles resemble adults in general Stage I larvae from October to Decerm-appearance. Early life history stages ber in coastal Indian waters. Hil-of tarpon were described by Hildebrand debrand (1943) collected Stage I(1934), Hollister (1939), Harrington larvae uff Beaufort, North Carolina,(1958), Gehringer (1959b), Wade from October through May; off Texas in(1962a), Eldred (1967, 1968, 1972), February, March, April, and November;Mercado and Ciardelli (1972), Jones et off the Florida Keys in November; andal. (1978), and Smith (1980). )ff Cuba in May. Offshore collectionsDescriptions of larval and juvenile of Stage I larvae were made by Geh-ladyfish were published by Hildebrand ringer (1959a) off Florida and Georgia(1943), Alikunhi and Rao (1951', Geh- in October, off South Carolina in May,ringer (1959a), Eldred and Lyons and off North Caroliri in November.(1966), and Jones et al. (1978). Arnold et al. (1960) ollected lep-

05

tocephali from early March to mid-May Snelson 1983) but may ascend riversnear Galveston, Texas. Tabb and Man- for considerable dis nces (Tagatzning (1961) reported that larvae were 1967).abundant in Florida Bay from Septemberthrough December. Eldred and Lyons(1966) reported collecting Stage I Habitats of Stage I tarpon larvaeleptocephali off Florida in January, are clear, warm, oceanic watersFebruary, May, June, August, October, (Gehringer 1959b; Robins 1978) withinand December. 100 m of the surface (Wade 1962a).

Surface water temperatures at collec-tion sites ranged from 26.0 to 30.0 'C

Summarizing various references on and salinities from 33.6 to 36.0 pptthe occurrences of larval tarpon, (Wade 1962a; Berrien et al . 1978;Robins (1978) and Smith (1980) noted Smith 1980). Estimated temperaturethat Stage I larvae occur from mid-May and salinity ranges at depth of cap-to late August, and Stage II larvae ture were 22.2-28.4 °C and 33.6-36.7from late June to early October; they ppt (Wade 1962a).inferred that spawning occurs in latespring or early summer.

Stage II and III tarpon larvaeand juveniles live in salt marsh and

Early Stage I larvae of ladyfish mangrove ponds, tidal creeks, rivers,were captured offshore (Gehringer ditches, beaches, and mosquito-control1959a) 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 andoccur inshore (Gehringer 1959a) at Randall 1957; Erdman 1960; Harrington26.3-38.5 ppt and 17.5-29.0 0C (Eldred and Harrington 1960, 1961; Wade 1962a,and Lyons 1966). Older early-life 1969; Rickards 1968; Dahlberg 1972;stages (Stage II and III larvae and Tagatz 1973; Gilmore et al. 1981;juveniles) inhabit coastal beaches, Snelson 1983). These habitats arecanals, bayous, lagoons, tidal ponds, typically shallow (<1 m), have a sandycreeks, rivers, and mosquito control mud or mud substrate with no rootedimpoundments (Erdman 1960; Zilberberg submerged vegetation, are lined by1966; Dahlberg 1972; Govoni and Mer- reeds or mangroves, usually have tur-riner 1978; Gilmore et al. 1981; bid or dark-stained waters, and may beThompson and Deegan 1982; Snelson either stagnant or have considerable1983). They live in water of a wide current (Beebe 1927; Breder 1933;range of salinities and tenperatures: Simpson 1954; Wade 1962a, 1969; Rick-14-45 ppt and 24-32 IC (Harrington ards 1968). In such habitats, larvae1958); 17.5-39.0 ppt (Harrington and and juveniles are able to withstandHarrington 1961); 34.3-34.6 ppt and environmental conditions deleterious18-23 °C (Eldred and Lyons 1966); 1.4- to many other fishes. Juvenile tarpon11.2 ppt and 21-30 CC (Herke 1969); have been collected at widely varying0.1-28.7 ppt and >19.9 °C (Dahlberg salinities: 31.8 ppt (Simpson 1954),1972); 10-20 ppt and <35 °C (Rose et 18.8-33.4 ppt (Moffett and Randallal. 1975); 5.6-5.8 ppt (Theiling and 1957), 14-45 ppt (Harrington 1958),Loyacano 197.); 2.2-6.1 ppt (Govoni 17.5-39.0 ppt (Harringt:on and Har-and Merrlner 1978); 0.0-8.8 ppt and rington 1961), 0.0-22.3 ppt (Rickards16-28 'C (Thompso and De-gan 1982). 1968), 0-47 ppt (Wade 1969), and 15-21Rose et al. (197F; ported a pH range ppt (Gilmore et al. 1982). Most lar-of 6.8-8.7 For .;tal impoundment val and juvenile tarpon live at rela-inhabited by - _ ladyfish. Adult tively high temperatures: 36.6 °Cladyfish usually li " in relatively (Moffett and Randall 1957), 36.0 'Copen inshore and tal habitats (Rickards 1968), and 40 0 C (Wade(Dahlberg 1972 Guim .e et al. 1981; 1969). Because tarpon respire

6

aerially (by gulping air) at least as Florida drainage ditch grew an averageearly as the beginning of Stage III of 1.0 mm/day (range, 0.7-1.4 mm/day)(Harrington 1966), low dissolved from 22 August to 20 October (Moffettoxygen concentrations .rc not and Rdndall 1957). Over the samedeleterious to survival. The strong period, modal lengths of tarpon inodor of hydrogen sulfide at capture this population increased by 1.4sites, 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 ofvestigators (e.g., Beebe 1927; Breder about 30 mm/month (Rickards 1968).1933, 1944; Rickards 1968; Wade 1969).Juvenile tarpon are often collectedfrom isolated marsh ponds that are Breder (1944), who determinedconnected to the estuary only during growth rates of captive juvenile tar-spring tides. Wade (1969) collected pon, wrote that 12 fish maintained atjuvenile tarpon at pH's of 6.8 to 8.2. the old New York Aquarium for 113 to

314 days grew an average of 0.088mm/day (range, 0.048-0.186 mm/day);

In eastern Florida marshes, Wade initial and final length ranges were(1969) found Stage III larvae in 94-145 and 110-176 mm TL. Threeditches at the headwaters of small tagged fish (355-365 mm TL) confinedcreeks. Small juveniles (40-80 mm SL) in natural ponds in southern Floridalived in larger ditches and creeks, did not grow in 133 to 152 daysespecially in the deeper pools. Large (August to January). Two othersjuveniles were found in larger canals (tagged in July) grew from 345 to 390and rivers. Juvenile tarpon eventu- and from 370 to 380 mm TL in 258 andally emigrate from marsh and mangrove 167 days, respectively. Four juvenilehabitats and enter coastal waters when tarpon (originally 230-350 mm TL)they reach about 600-800 mm TL (Robins maintained in laboratory pools grew 131978). In Georgia, tarpon are unable to 179 mm (mean increment, 97 mm) into overwinter in marsh habitats; 15 months; a fifth grew from 436 tojuveniles left marshes, presumably to 465 mm TL in 6 months.migrate south, by late October, whenthey had attained about 160 mm SL(Rickards 1968). Adults live in bays, Ten tarpon raised by Harringtonlagoons, and coastal habitats (Breder (1966) in the laboratory from Stage1944; Dahlberg 1972; Gilmore et al. III larvae (18.1-22.7 mm SL; mean,1981; Snelson 1983) or may cruise the 21.4 mm) for 1 year grew to 55.4-105.3open ocean (Robins 1978). mm SL (mean, 67.2 mm SL).

GROWTH CHARACTERISTICS Although scales of adult tarponhave distinct rings resembling annuli

Age and Growth (Breder 1944; Moffett and Randall1957), these marks have not been

Moffett and Randall (1957), who validated as annuli and should be con-examined length-frequency distribu- 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 aremodal lengths increased from 75-80 mm shown in Figure 3. Maximum age basedFL in early September to 110-115 mm FL on these checks was 16 years (Moffettat the end of the month, and inferred and Randall 1957); however, largera length increase of 3bout 1.4 mm/day; fish have been captured.rates declined by about 50% in Oc-tober. Five marked juvenile tarpon Gehringer (1959a) reared ladyfish(301-376 mm FL when tagged) in a south in the laboratory from early Stage II

7

to the juvenile phase. Rates of W = 9 x 10-6 TL3

change in standard length during thefirst part of Stage 11 (from about 35- where W = weight in grams and TL = to-40 mm to 25 mm SL) averaged -1.061 tal length in millimeters.mm/day. Further shrinking to about20-21 mm. SL proceeded at about -0.342 Harrington (1958) derived themm/day. Initial length increase following length-weight relation fordurinq early Stage III, from about 20 154 tarpon, 16.0-45.5 mm SL:to 25 mm SL, averaged 0.140 mm/day.Growth rates of late Stage III larvae W = 0.05514 x 1.06 9SL - 0.15and early juveniles (<60 mm SL) wereabout 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. TheField collections suggested a faster relation is valid only for fish withinrate of growth (about 2 mm/day) under the stated size range.natural conditions (Gehringer 1959a).No information is available on growth On the basis of a graph presentedof adult ladyfish. by Moffett and Randall (1957), we

derived the following relation betweenMorphometric Relations total length (TL) and fork length (FL)

for tarpon:Breder (1944) presented the fol-

lowing length-weight relation for TL = 1.10 FL.adult tarpon in Florida:

Harrington (1958) developed thefollowing conversions between forklength (FL), total length (TL), and

standard length (SL) in millimeters-for tarpon 25-54 mm SL:

/FL = 1.1282SL - 1;TL = 1.3333SL - 2.

£ Sekavec (1974) derived the fol-lowing length-weight formula from 295juvenile ladyfish 45-201 mm FL fromLouisiana:

/a logio W = -5.3295 + 3.1123 loglo FL

where W = weight in grams and FL =

fork length in millimeters. Mean con-dition factor (K, where K = [W/(FL) 3 ]

-- "I COW 11"PdW ", , ,x 106; Lagler 1956) of these fish was-,At(Mole, end Rarodt 1%v 8.1 (range 6.6-8.9).

0 400 COW (M mir Rendell 1167)

FISHERY0

NUMBER OF PUTAnvE ANNUU The tarpon and ladyfish fisheries

are solely recreational; no commercialFigure 3. Back-calculated mean fishery exists for either species inlengths of tarpon from the west and the United States. Neither species iseast coasts of Florida, at putative recorded in the National Marineannuli on scales. Recreational Fisheries Survey and

8

regional catch data are nonexistent copepods and ostracods) and secon-(Grant L. Beardsley, Senior Scientist darily on insects and small fishes;for Recreational Fisheries, National larger juveniles continue to feed onMarine Fisheries Service, Southeast zooplankton, but progressively in-Fisheries Center, Miami, Florida; crease consumption of insects, fishespers. comm.). (especially poeciliids and

cyprinodontids), crabs, and grassTilmont et al. (unpublished) sum- shrimps of the genus Palaemonetes

marized recreational fishery statis- (Beebe 1927; Breder 1933; Mottett andtics for tarpon and ladyfish in Randall 1957; Harrington and Har-Everglades 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 (Robinsmade up an average of 0.2% of the 1978).reported recreational catch annually.Mean annual catch rates varied from In laboratory settings, early0.1 to 0.4 fish/h. Less than 10% of Stage II ladyfish larvae ate livethe tarpon caught were harvested, plankton (Alikunhi and Rao 1951) andprimarily for trophy mounts. Tarpon live brine shrimp (Artemia) naupliiaccounted for 1% of the catch, 0.4% of (Gehringer 1959a). Stage Il1 larvaethe harvest, and 7% of the effort in ate small live Fundulus and Gambusiathe professionally guided fishery in and pieces of shrimp and fish. Underthe park. Reported catch rates sug- natural conditions. Stage II and IIIgested that stocks of tarpon in the ladyfish larvae (<50 mm SL) feed al-park were relatively stable. most exclusively on zooplankton; con-

Ladyfish were commonly caught but sumption of zooplankton by juvenilesinfrequently harvested by anglers in is progressively reduced as ingestionEverglades National Park (Tilmont et of small fishes and shrimps increasesal., unpublished). Ladyfish made up (Harrington and Harrington 1961).about 5% of the total reported catch Ladyfish <100 mm long feed especiallybut less than 0.4% of the harvest, on insects, copepods, and otherFew anglers (0.02%) considered the arthropods (Fyfe 1986).fish a preferred species. Mean annualcatch rates varied from 0.25 to 0.67 Adult tarpon and ladyfish arefish/h. Ladyfish population abun- strictly carnivorous and feeddances in the park were cyclic, peak- primarily on mid-water prey (Hilde-ing about every five years. However, brand 1963; Sekavec 1974). Food isa general increase in ladyfish abun- swallowed whole (Sekavec 1971).dances in the park occurred from thelate 1960's through the early 1980's. Adult ladyfish feed primarily on

fish; fish constituted 94%, 82%, and34% of food items found in ladyfishstomachs by Sekavec (1974), Darnell

ECOLOGICAL ROLE (1958), and Knapp (1949), respec-tively. In Sekavec's (1974) study,

Feeding Behavior/Food Habits juvenile Gulf menhaden (Brevoortiatyrannus) composed 72% of the iden-

Stage I tarpon and ladyfish titiable fish consumed.larvae do not forage; nutrients areobtained directly from seawater by in- Decapod crustaceans are also im-tegumentary absorption (Pfeiler 1986). portant foods of ladyfish. Linton

(1904) reported that diets of 12Stage II and III tarpon larvae ladyfish from North Carolina consisted

and small juveniles (<125 mm SL) feed exclusively of shrimp. Knapp (1949)primarily on zooplankton (e.g., found that 78.2% of stomach contents

09

of laayfish from the Texas coast were reported from the intestine ofcrustaceans. Decapods made up 5.5% ladyfish (Corkum 1959).(Sekavec 1974) and 10% (Darnell 1958)of the diets of ladyfish fromLouisiana. ENVIRONMENTAL REQUIREMENTS

Adult tarpon feed both noctur- Temperaturenally and diurnally (Wade 1962a) on avariety of organisms including mullets The tarpon and ladyfish are dis-(Mugil spp. ) , pinfish (Lagodon tinctly thermophilic fishes. BothrhF-Uides), ariid catfishes, Atlantic have been reported in cold-relatedneedlefi-sh (Strongylura marina), sar- fish kills in Florida (Storey anddines (Harengula spp.), shrimp, and Gudger 1936; Storey 1937; Snelson andcrabs (Babcock 1951; Wade 1962a). Bradley 1978). At Port Aransas,

Texas, annual tarpon abundances arePredators correlated with yearly water tempera-

ture regimes (Moore 1975). TarponPredation by carnivorous concentrate around heated power-plant

zooplankters and small fishes un- effluents during winter in the Indiandoubtedly causes high mortality of eggs River, Florida (Snelson 1983).and larvae of both ladyfish and tarponbefore the larvae enter coastal nurs- Early Stage I larvae of bothery marshes. In marshes, juvenile species occur only in warm oceanictarpon 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 al. 1978; Smithfett and Randall 1957; Rickards 1968; 1980), and it appears probable thatWade 1969). Both species are probably such temperatures are necessary forpreyed upon by piscivorous birds proper development of eggs and early(Beebe 1927; Rickards 1968), and adult larvae.tarpon are occasionally eaten bysharks, porpoises, and alligators Moffett and Randall (1957) ex-(Wade 1962a, 1962b). posed juvenile tarpon (72-130 mm FL)

held at 25-27 0C to high temperaturesParasites in laboratory trials. Fish were

warmed from maintenance to test tem-The di genetic trematode peratures in 3 hours. Fish subjected

Lecithochirium microstomum occurs in to 39.4-39.6 OC survived the 24-hthe stomach of tarpon (Manter 1947). trials; those exposed to 40.5-41.9 °CThe isopods Nerocila acuminata and did not. In a series of cold-Cymothoa oestrum are external tolerance trials, juvenile tarpon (71-parasites (Babcock 1951; Pearse 1952). 130 mm FL) died within 24 h at tem-Causey (1953) reported the copepod peratures of 14.8-18.0 °C; others sur-Paralebion pearsei from tarpon. The vived trials at 14.8-19.7 °C (Moffetttrematode Bivescula tarponis is and Randall 1957). Tabb (personalpresent in the pyoric caecae and communication in Wade 1962a) reportedalong the entire length of the intes- mortality of tarpon in Everglades Na-tine (Sogandares-Bernal and Hutton tional Park when water temperature1959). Though not parasitic, remoras decreased from 24 to 11 °C within a(Remora remora) are commonly observed few hours. Rickards (1968) collectedaTt -a-cedT-6-1arge tarpon (Babcock sluggish juvenile tarpon in a Georgia1951; Wade 1962a). salt marsh in late November when water

temperature was 16.0 ° C. Concur-Trematodes of the genera rently, several juveniles maintained

Bucephalus and Prosorhynchus have been in a steel holding tank died overnight

10

when water temperatures dropped from salinities ranging from 0.0 to 45.021.0 to 12.0 OC. However, Wade (1969) ppt (Harrington 1958; Harrington andcollected juvenile tarpon at tempera- Harrington 1961; Herke 1969; Dahlbergtures as low as 12 °C, and Gilmore et 1972; Govoni and Merriner 1978;al. (1982) collected tarpon at 14 0C Thompson and Deegan 1982).from a mosquito control impoundment ineastern Florida. Robins (1978) stated Adult ladyfish also tolerate athat the lower lethal temperature of wide range of salinities, but appeartarpon is about 100 C. less likely than tarpon to occupy

freshwater. We found no reference inLadyfish appear to be slightly the literature specifically reporting

more tolerant of low temperatures than adults from truly freshwater. One oftarpon, judging by the lower frequency us (A.V.Z.) has captured ladyfish inof 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 ofto 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 evaluating1975). salinity requirements of fishes) to be

freshwater because of its distanceSalinity from the mouth of the river (about 190

km), but numerous salt springs main-Throughout most of their life tain relatively high salinities.

stages, tarpon and ladyfish tolerate a Tagatz (1967) collected ladyfish aswide range of salinities. However, far upstream as Palatka, Florida (135early Stage I larvae of both species km from the mouth) and reportedhave been collected only at oceanic salinity there to be 0 ppt; however,salinites of 28.5-39.0 ppt (Wade he recorded all salinities less than1962a; Eldred and Lyons 1966; Eldred 1.0 ppt as 0 ppt. It is likely that1967, 1968, 1972; Berrien et al. 1978; appreciable salinities (perhaps >0.5Smith 1980), and it is likely that ppt) are, if not required, then atsuch concentrations are required by least preferred by ladyfish.eggs, yolk-sac larvae, and early StageI larvae of both species for proper Dissolved Oxygendevelopment.

Tarpon are obligate air breathersBeyond Stage I, tarpon and (the swimbladder contains alveolar

ladyfish are decidedly euryhaline. tissue; Shlaifer 1941) and areJuvenile tarpon can withstand direct frequently seen "rolling" at the sur-transfer from salt- to freshwater and face gulping air; when prevented fromvice-versa (Breder 1944; Moffett and reaching the surface, they die withinRandall 1957). Habitats occupied by 7 to 128 h, even in highly oxygenatedtarpon range from fresh to hyper- water (Shlaifer 1941). Air breathingsaline, or 0 to 47 ppt (Hildebrand is imitatively mediated by visual1939; Simpson 1954; Moffett and Ran- cues; juveniles in a school come todall 1957; Harrington 1958; Harrington the surface in rapid successionand Harrington 1961; Rickards 1968; (Shlaifer and Breder 1940; ShlaiferWade 1969; Dahlberg 1972; Tagatz 1973; 1941), perhaps to reduce individualTucker and Hodson 1976). susceptibilities 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 dissolvedbrackish water (10.4 ppt) and success- oxygen concentration (Shlaifer andfully transferred them directly to Breder 1940; Shl3ifer 1941). Air-freshwater (0.08 ppt). Juvenile breathing precludes mortality inladyfish have been collected at anoxic waters and allows tarpon to

11

survive under conditions deleterious Wetlands Destruction and Degradationto most fishes. Tarpon have thisability at least as early as the Offshore and coastal habitats ofbeginning of Stage III (Harrington very young and adult tarpon and1966). ladyfish are relatively immune to

human-induced degradation. Con-Air breathing has not been versely, the estuaries, salt 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 ladyfishnot been reported in the literature, and tarpon in Florida are highly vul-Dissolved oxygen requirements of nerable to changes induced by develop-ladyfish are unknown, but it is likely ment. Robins (1978) discussed thethat the species is relatively various activities that are degradingtolerant of hypoxic conditions, as it tarpon nursery grounds in Florida; hisis often found with tarpon in poorly comments are also applicable to earlyoxygenated habitats. Ladyfish in- life history stages of ladyfish, whichhabited a coastal impoundment in share these habitats. Among factorsLouisiana in which dissolved oxygen resulting in the destruction of nurs-concentrations reached a minimum of ery wetlands, he listed filling of1.0 mg/l (Rose et al. 1975). wetlands, canalization, bulkheading,

construction of water-line right-of-ways and steep-sided boat-access

Contaminants finger-canals, and impoundment of wet-lands for mosquito control. Progress

Aerial spraying and ground fog- has recently been made in amelioratingging for nuisance insect control are the effects of impoundment forwidely practiced in Florida's coastal mosquito control because impoundmentzone, and agricultural pesticides and does not necessarily result in theherbicides used in south Florida enter destruction of wetlands. Rather, ir-coastal waters. Robins (1978) pounded wetlands, if properly managed,reported that tarpon are extremely can retain the beneficial characteris-susceptible to contaminants. Applica- tics of natural wetlands while provid-tion of dieldrin pellets in a Florida i ng adequate mosquito controlsalt marsn for the control of larval (Clements and Rogers 1964; Provostsandflies (Culicoides) resulted in 1973). However, access to these wet-mortality of ladyfish and tarpon lands (and subsequent opportunities(Harrington and Bidlingmayer 1958). for egress) by larval and juvenile

tarpon and ladyfish is precluded orseverely curtailed by reduced or non-

Turbidity existent exchange with estuarinewaters (Wade 1969; Gilmore et al.

Stage I larvae of both ladyfish 1982; Harrington and Harrington 1982).and tarpon occur only in clear off- Improved impoundment managementshore waters. Subsequent life history strategies, aimed at enhancing ex-stages appear to be tolerant of high change rates, have been proposed byturbidities. Habitats occupied, espe- Clements and Rogers (1964), Provostcially by juveniles, are generally (1973), Montague et al. (1985), anddescribed as turbid and dark-stained. Lewis et al. (1985).

12

LITERATURE CITED

Alikunhi, K.H., and S.N. Rao. 1951. Breder, C.M., Jr. 1933. Young tarponNotes on the metamorphosis of Elops on Andros Island. Bull. N.Y. Zool.saurus Linn. and MegaTops Soc. 36:65-67.cyprinoides (Broussonet) with obser-vations on their growth. J. Zool. Breder, C.M., Jr. 1944. MaterialsSoc. India 3:99-109. for the study of the life history of

Tarpon atlanticus. ZoologicaArnold, E.L., Jr., R.S. Wheeler, and g: 21 _725.

K.N. Baxter. 1960. Observations onfishes and other biota of East Briggs, J.C. 1958. A list of FloridaLagoon, 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 copepodsBabcock, 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 in30:141-145. Florida, 1958-1963. Mosquito News

24:265-276.Berra, T.M. 1981. An atlas of dis-

tribution of the freshwater fishfamilies of the world. University Corkum, K.C. 1959. Some trematodeof Nebraska Press, Lincoln. 197 pp. parasites of fishes from the Missis-

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 Dolphin Dahlberg, M.D. 1972. An ecologicalsurvey of the Continental3WET study of Georgia coastal fishes.waters between Martha's Vineyard, U.S. Natl. Mar. Fish. Serv. Fish.Massachusetts and Cape Lookout, Bull. 70:323-353.North Carolina, 1965-66. Natl. Mar.Fish. Serv. Sandy Hook Lab. Tech. D&-nell, R.M. 1958. Food habits ofSer. Rep. No. 15. 152 pp. fishes and larger invertebrates of

Lake Pontchartrain, Louisiana, anestuarine community. Publ. Inst.

Bigelow, 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. Megalops atlanticus Valenciennes,

13

(Megalopidae) in Florida waters. Gill, T. 1907. The tarpon and lady-

Fla. Board Conserv. Mar. Lab. Leafl. 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.

larval tarpon, Megalops atlanticus Donohoe. 1982. A comparison of the

Valenciennes, from the Gulf ot 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. NortheastGulf Sci. 5:25-37.

Eldred, B. 1972. Note on larval tar-pon, Megalops atlanticus (Mega- Gilmore, R.G., Jr., C.J. Donohoe, D.W.

lopidae) in the Florida Straits. 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, (Elopia) in F orida Gosline, 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, Honolulu. 208 pp.

Erdman, D.S. 1960. Larvae of tarpon, Govoni, J.J., and J.V. Merriner.Megalops atlantica, from the Anasco 1978. The occurrence of ladyfish,RTiver, Puerto Rico. Copeia Elops saurus, larvae in low salinity

1960:146. waters -and another record forChesapeake Bay. Estuaries 1:205-

Forey, P.L. 1973a. Relationships of 206.elopomorphs. Zool. J. Linn. Soc.53(Suppl. 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 Phyletic studies of teleosteanelopiform fishes, fossil and recent. fishes with a provisional class-Bull. Br. Mus. Nat. Hist. (Geol.), ification of living forms. Bull.

Suppl. 10:1-222. 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, Megalops atlantica Valenciennesa subtropical salt marsh from east from transitional stage through on-central Florida, U.S.A. Fla. Sci. set of scale formation. Copeia49(Suppl .1):18. 1958:1-10.

Gehringer, J.W. 1959a. Early Harrington, R.W., Jr. 1966. Changes

development and metamorphosis of the through one year in the growth rates

ten-pounder, Elops saurus Linnaeus. of tarpon, Megalops atlanticusU.S. Fish. WfTTTServ ish. Bull. Valenciennes, reared from mid-59:619-647. metamorphosis. Bull. Mar. Sci.

16:863-883.Gehringer, J.W. 1959b. Leptocephalus

of the Atlantic tarpon, Megalops at-lanticus Valenciennes, from offsh~e- Harrington, R.W., Jr., and W.L. Bid-waters. Q. J. Fla. Acad. Sci. lingmayer. 1958. Effects of

21:235-240. dieldrin on fishes and invertebrates

14

of a salt marsh. J. Wildl. Manage. India, including a study of the22:76-82. caudal skeleton and a comparison

with the Atlantic species, TarponHarrington, R.W., Jr., and E.S. Har- atlanticus. Zoologica 24:449-17T.?

rington. 1960. Food of larval andyoung tarpon, Megalops atlantica.Copeia 1960:311rT.- -Jones, P.W , F.D. Martin, and J.D.

Harrington, R.W., Jr., and E.S. Har- Hardy, Jr. 1978. Development ofrington. 1961. Food selection fishes of the Mid-Atlantic Bight: anamong fishes invading a high sub- atlas of egg, larval and juveniletropical salt marsh: from onset of stages. Vol. 1: Acipenseridaeflooding through the progress of a through Ictaluridae. U.S. Fishmosquito 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 andpounding a Florida salt marsh to Middle America. Bull. U.S. Natl.prevent breeding by salt marsh Mus., No. 47 (1):1-1240.mosquitos. 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

collection of larval ladyfish, Elops fishes. Dover Publications, Inc.,saurus in Louisiana. Proc.La. New York. 574 pp.Xcad_.-Sci. 32:29-30.

Knapp, F.T. 1949. Menhaden utiliza-Hildebrand, S.F. 1934. The capture tion in relation to the conservation

of a young tarpon, Tarpon atlan- of food and game fishes of the Texasticus, at Beaufort, No-rt 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 social44:239-248. component of respiration in fishes.

Copeia 1976:689-697.Hildebrand, S.F. 1939. The Panama

Canal as a passageway for fishes, Lagler, K.F. 1956. Freshwaterwith lists and remarks on the fishes fishery biology. W.C. Brown Co.,and invertebrates observed. Dubuque, Iowa. 421 pp.Zoologica 24:15-45.

Lewis, R.R., III, 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 fisheryElops saurus Linnaeus. J. Wash. resources of Florida. Pages 281-336AcSa -F g T:90-94. in W.S. Seaman, Jr., ed. Florida

Tq-uatic Habitat and FisheryHildebrand, S.F. 1963. Family Resources. Florida Chapter American

Elopidae. Pages 111-131 in Fishes Fisheries Society, Gainesville.of the western North Atlantic.Sears Found. Mar. Res. Mem. 1(3).

Linton, E. 1904. Parasites of fishesHollister, G. 1939. Young Megalops of Beaufort, North Carolina. Bull.

cyprinoides from Batavia, Dutch East U.S. Bur. Fish. 24:321-428.

* 15

Manter, H.W. 1947. Digenetic Timbers Conf. Ecol. Anim. Controltrematodes of marine fishes. Am. Habitat Manage. 5:5-17.Midl. Nat. 38:339-340.

Rickards, W.L. 1968. Ecology andMcLane, A.J., ed. 1974. McLanes's growth of juvenile tarpon, Megalops

new standard fishing encyclopedia atlanticus, in a Georgia salt marsh.and international angling guide, 2nd TU.1far. Sci. 18:220-239.ed. Holt, Rinehart and Winston,N.Y. 1,156 pp. Robins, C.R. 1978. The tarpon -

unusual biology and man's impactMercado, J.E., and A. Ciardelli. determine its future. Mar. Recrea-

1972. Contribucion a la morfologia tional Fish. 2:105-112.y organogenesis de los leptocefalosdel sabalo Megalops atlanticus Rebins, C.R., R.M. Bailey, C.E. Bond,(Pisces: Meia.opidae7. 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 namesMoffett, 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. 174Lab. 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 ofof salt marsh manageinent on Merritt penaeid shrimp in Louisiana salt-Island National Wildlife Refuge, marsh impoundments. Trans. Am.Florida. Fla. Coop. Fish Wild?. Fish. Soc. 104:296-307.Res. Unit Tech. Rep. 17. Gaines-ville. 92 pp. Sekavec, G.B. 1971. Gross morphology

of the digestive tract of theMoore, R.H. 1975. Occurrence of ladyfish, Elops saurus. Chesapeake

tropical marine fishes at Port Aran- Sci. 12:275-2.sas, Texas 1967-1973, related to seatemperatures. Copeia 1975:170-172. Sekavec, G.B. 1974. Summer foods,

length-weight relationship, and con-Nelson, J.S. 1984. Fishes of the dition factor of juvenile ladyfish,

world, 2nd ed. John Wiley and Sons, Elops saurus Linnaeus, fromNew York. 523 pp. Louisiana coastal streams. Trans.

Am. Fish. Soc. 103:472-476.Pearse, A.S. 1952. Parasitic crus-

tacea from the Texas coast. Publ. Shlaifer, A. 1941. Additional socialInst. Mar. Sci. Univ. Tex. 2:5-42. and physiological aspects of

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planation of the developmentalstrategy in leptocephalous larvae of Shlaifer, A., and C.M. Breder, Jr.marine teleost fishes. Environ. 1940. Social and respiratory be-Biol. Fishes 15:3-13. havior of small tarpon. Zoologica

25:493-512.

Provost, M.W. 1973. Salt marsh Simpson, D.G. 1954. Two small tarponmanagement in Florida. Proc. Tall from Texas. Copeia 1954:71-72.

16

Smith, D.G. 1980. Early larvae of Tagatz, M.E. 1967. Fishes of the St.the tarpon, Megalops atlantica Johns River, Florida. Q. J. Fla.Valenciennes (Pisces: Eiopidaej, Acad. Sci. 30:25-50.with notes on spawning in the Gulfof Mexico and the Yucatan Channel. Tagatz, M.E. 1973. A larval tarpon,Bull. Mar. Sci. 30:136-141. Megalops atlanticus, from Pensacola,

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Snelson, F.F., Jr. 1983. Ichthyo-fauna of the northern part of the Theiling, D.L., and H.A. Loyacano, Jr.Indian River Lagoon system, Florida. 1976. Age and growth of red drumFla. 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 dueto cold on the east coast of Thompson, B.A., and L.A. Deegan.Florida, January, 1977. Fla. Sci. 1982. Distribution of ladyfish41:1-12. (Elops saurus) and bonefish (Albuia

vu7p-es-ep-tocephali in Louisiana.Sogandares-Bernal, F., and R.F. Hut- BlXTlT-ar. Sci. 32:936-939.

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within Everglades 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-metamorphicland, Florida. Ecology 18:10-26. larvae of the tarpon, Megalops at-

lantica, from the Cape F-ear RiverStorey, M., and E.W. Gudger. 1936. estuary, North Carolina, 1973-74.

Mortality of fishes due to cold at Chesapeake Sci. 17:123-125.Sanibel Island, Florida, 1886-1936.Ecology 17:640-648. Wade, R.A. 1962a. The biology of the

tarpon, Megalops atlanticus, and theStorey, M., and L.M. Perry. 1933. A ox-eye, Megalops cypri n-Tdes, with

record of young tarpon at Sanibel emphasis on larval development.Island, Lee County, Florida. 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 juvenileTabb, D.C., and R.B. Manning. 1961. tarpon and effects of dieldrin onA checklist 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 coastalBull. Mar. Sci. Gulf Carib. 11:552- marsh of northwest Florida. Con-649. trib. Mar. Sci. 11:126-134.

*17

30272-101

REPORT DOCUMENTATION I. REPORT N. S3. R cipntes Accession No.

R_PORT DOCUM IOAGE lBiological Report 82(11.104)*I __

4. Title and Subtitle 5. Rep ,rt Date

Species Profiles: Life Histories and Environmental Requirements July 1989

of Coastal Fishes and Invertebrates (South Florida) -- Ladyfish --

and Tarpon

7. Author(%) S. Performing Organization Rapt. No.

Alexander V. Zale and Susan G. Merrifield9. performing Organization Name and Address 10. Project/Task/Work Unit No.

Oklahoma Cooperative Fish and Wildlife Research Unit

404 Life Sciences West 11. Contrect(C) or Grnt(G) No.

Oklahoma State University (C)

Stillwater, OK 74078 (0)

12. Sponsoring organization Name and Address 13. Type of Report & period Co~'red

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 14.

Washington, DC 20240 Vicksburg, MS 39180

is. Supplementary Note

* U.S. Army Corps of Engineers Report No. TR EL-82-4

I& Abstract (Limit: 20 words)

Species profiles are literature summaries of the taxonomy, morphology, distribu-

tion, life history, habitats, and environmental requirements of coastal species

of Fishes and aquatic invertebrates. They are designed to assist in environ-

mental inpact assessment. The tarpon and ladyfish are popular gamefishes.

Adults spawn offshore. Larval and juvenile stages inhabit coastal marshes and

mangroves. Both species are thermaphilic (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. Descriptors

Fishes Fisheries Life cycles

Estuaries TemperatureFeeding habits SalinityGrowth Oxygen

b. Identiflrs/Open.Ended Terms

Tarpon Trophic ecologyMegalops atlanticus Spawning

Ladyfi sh Environmental requirements

Elops saurus

c- COSATI Fileld/Group

I&. Availability Staernan; It. security Class MTIS Report) 21. No. of pages

Unlimited Unclassified 17W0. Security Class MIS Page) 22. Price

Unclassified(See ANSI-ZS I OPTIONAL IOrtM 272 (4-77)

(Formerly NTIS-35)Department of Commerce

-- -- , , i I I II I I

As the Nation's principal conservation agency, the Department ofthe Interior has responsibility for most of our nationally ownedpublic lands and natural resources. This includes fostering thewisest use of our land and water resources, protecting our fishand wildlife, preserving the environmental and cultural values of ournational parks and historical places, and providing for the enjoy-ment of life through outdoor recreation. The Department assessesour energy and mineral resources and works to assure that theirdevelopment is in the best interests of all our people. The Depart-ment also has a major responsibility for American Indian reservationcommunities and for people wno live in island territories under U.S.administration.

U.S. DEPARTMENT OF THE INTERIORFISH AND WILDLIFE SERVICE [

TAKE PRIDEin America

UNITED STATESDEPARTMENT OF THE INTERIOR POSTAGE AND FEES PAID

FISH AND WILDLIFE SERVICE U.A DEPARTMENT OF THE INTERIORNational Wetlands Research Center INT-4

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