BULLETIN OF MARINE SCIENCE, 61(2): 271285, 1997

REPRODUCTION OF TARPON, MEGALOPS ATLANTICUS, FROM FLORIDA AND COSTA RICAN WATERS AND NOTES ON THEIR AGE AND GROWTH Roy E. Crabtree, Edward C. Cyr, Didiher Chacn Chaverri, William O. McLarney and John M. Dean
ABSTRACT
We examined 1,469 tarpon ranging 102-2,045 mm fork length (FL) that were collected in South Florida waters and 217 tarpon ranging 880-1,860 mm FL that were collected in Costa Rican waters from 1989 to 1994. Among the Florida tarpon, gonads of 737 fish were examined histologically, and gonadosomatic indices were calculated for 409 tarpon. Among Costa Rican tarpon, gonads of 202 tarpon were examined histologically, and gonadosomatic indices were calculated for 178 tarpon. Florida fish were significantly larger than those from Costa Rica, and sexually mature females were significantly larger than sexually mature males in both areas. All Florida males were sexually mature by 1,175 mm FL, and only one smaller male (901 mm) was mature. The smallest sexually mature Florida female was 1,285 mm FL, and all larger females except one were sexually mature. Only one Florida tarpon, a 7-yr-old female, younger than 10 yrs was sexually mature; all tarpon older than 10 yrs were sexually mature. In Costa Rica, only one immature female (880 mm FL) was examined; all other females (1,126-1,860 mm FL) were sexually mature. All Costa Rican males examined (880-1,550 mm FL) were sexually mature. Florida tarpon spawned during April-July and by August, most females were spent or recovering. Spawning in Costa Rican waters was not seasonal, and reproductively active females were caught during all months. Fecundity estimates for Florida tarpon ranged 4.5-20.7 million oocytes and had a significant positive relationship with fish weight. Otoliths of tarpon from Costa Rican waters were difficult to read. We examined thin-sectioned otoliths (sagittae) from 194 Costa Rican tarpon but could estimate ages for only 87 fish (45%). Costa Rican tarpon are long-lived and reach a maximum age of at least 48 yrs. Most (74%) tarpon caught in Costa Rica that we aged were 15 to 30 yrs old.

Tarpon, Megalops atlanticus, are large, migratory, elopomorph fish that frequent coastal and inshore waters of the tropical and subtropical Atlantic Ocean. In the western Atlantic, tarpon regularly occur from Virginias eastern shore to central Brazil and throughout the Caribbean Sea and the Gulf of Mexico (Wade, 1962; Hildebrand, 1963; de Menezes and Paiva, 1966; Zale and Merrifield, 1989). In parts of Central America and south Florida, tarpon are the basis of economically important recreational fisheries. In Florida, the fishery is intensely regulated, and anglers are required to purchase a $50 permit before harvesting a fish. Since the establishment of the permit system in 1989, the harvest of tarpon in Florida has declined to less than 100 fish per yr, and the fishery is now mostly catch-and-release. Tarpon occur in a variety of habitats ranging from freshwater lakes and rivers to offshore marine waters, but large tarpon targeted by Floridas fishery are most abundant in estuarine and coastal waters. In Florida, the fishery is seasonal; most tarpon are caught during May-July, although some fish are caught in all months. The Costa Rican fishery is located principally in the Rio Colorado region, where fishing occurs year-round both in coastal waters and in the river. Most of the tarpon fishing in the
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area is catch-and-release, and anglers usually fish with professional guides employed by local fishing lodges. Little information is available on the reproductive biology of tarpon. Chacn Chaverri and McLarney (1992) documented the occurrence of larvae and juveniles during JulyNovember in Costa Rican inshore waters. No other information on reproduction or age and growth exists for the area. More is known about the biology of tarpon from other areas. Crabtree et al. (1992) and Crabtree (1995) described the early life history of tarpon from Florida waters and found that spawning is seasonal and occurs offshore during MayAugust. Onshore transport of larvae is episodic and affected by environmental events (Shenker et al., 1995). Breder (1944) examined gonads of tarpon from Florida waters but did not adequately describe either temporal spawning patterns or age and length at maturity. Crabtree et al. (1995) described the age and growth of tarpon from Florida waters and found that tarpon are long-lived and can reach ages greater than 50 yrs. De Menezes and Paiva (1966) examined gonads from tarpon caught off the northeast coast of Brazil and found that reproduction occurs during October-January. In this paper, we describe seasonal spawning patterns, length at sexual maturity, and fecundity of tarpon from South Florida and from the Rio Colorado area of Costa Rica. Age estimates are presented for tarpon from Costa Rican waters. MATERIALS AND METHODS
We obtained Florida tarpon from a variety of sources from April 1988 to November 1993. Most large fish (>1,100 mm FL) were obtained from taxidermists in Fort Myers and Fort Lauderdale and were caught in either the Florida Keys or Boca Grande Pass on Floridas Gulf coast (26 43' N, 82 16' W). A second source of large fish was tournaments held in the Keys, Boca Grande Pass, and the Tampa Bay area (27 40' N, 82 35' W). All large tarpon were caught with hook-and-line gear. Small Florida tarpon (<1,100 mm FL) were taken with cast nets, hook-and-line gear, electroshocker, trammel nets, and gill nets at various locations. Tarpon in Costa Rica were caught with hook-andline gear during normal fishing operations conducted by fishing lodges located in the Rio Colorado area on the Caribbean coast of northeastern Costa Rica from May 1989 to January 1994. Standard length (SL), fork length (FL), and total length (TL) were measured to the nearest millimeter (mm). All lengths reported are fork lengths. Large tarpon (>1,100 mm) were weighed to the nearest 0.5 kg, and smaller tarpon were weighed to the nearest gram. Otoliths (sagittae) were removed, cleaned with bleach (5.25% sodium hypochlorite), and rinsed first in water and then in 95% ethanol. Otoliths were stored dry or in 95% ethanol until sectioned. Sex, gonad condition, and gonad weight in grams were recorded. Gonad samples were removed from the fish and preserved in 10% buffered Formalin; they were then soaked in water for 1 h and stored in 70% ethanol. Length-weight regressions were calculated by linear regression of log10-transformed data and were compared with a t-test (Zar, 1984). Histological sections of gonads from 737 Florida tarpon and 202 Costa Rican tarpon were prepared and assessed for reproductive state. Initially, gonad sections were taken from all tarpon. After examining histological sections from 168 Florida tarpon smaller than 400 mm, we concluded that we could not reliably sex small tarpon and subsequently processed sections only from fish larger than 400 mm. Gonad samples were processed histologically with a modification of the periodic acid Schiff s stain for glycol-methacrylate sections, with Weigerts iron-hematoxylin as a nuclear stain and metanil yellow as a counterstain (Quintero-Hunter et al., 1991). Gonads were classified based on the modified maturity criteria of Murphy and Taylor (1990) in Table 1. The presence of widespread atresia in inactive ovaries was interpreted as evidence of past reproductive

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Table 1. Reproductive classes of tarpon gonads of each sex Class 1. Immature 2. Developing Females Few folds of ovigerous lamellae; few or no primary oocytes; oogonia predominate. Ovigerous lamellae fill lumen of entire gonad; abundant primary oocytes; absence of atretic oocytes. Males Only spermatogonia present; no evidence of tubule development. Early spermatogenesis; few scattered crypts of spermatocytes; peripheral tubules differentiating, lumen not developed.

3. Maturing 4. 5.

6.

7.

8.

Early vitellogenesis; oogonia, primary oocytes, and oocytes with yolk vesicles present. Mature Late vitellogenesis; oogonia, primary Late spermatogenesis; few oocytes, and oocytes with yolk spermatogonia; spermatozoa collecting vesicles and yolk globules present. in tubules and central lumen. Ripe Maturation. Oogonia, primary oocytes, and oocytes with yolk vesicles, globules, and migrating nuclei present; hydrated oocytes present in spawning fish. Partially spent Collapsed, empty, atretic, postovulatory follicles present; amorphous hydrated oocytes occasionally present; oogonia, primary oocytes, and vitellogenic oocytes with yolk vesicles and yolk globules present. Spent Oogonia and primary oocytes present in ovigerous lamellae; atretic postovulatory follicles present; ovary with empty areas; atretic oocytes scattered throughout, usually in association with blood vessels. Recovering Prolific recrudescence of ovigerous Network of efferent tubules lined with lamellae with myriad oogonia and spermatogonia; few spermatozoa primary oocytes; PAS-positive present; PAS-positive melanomelano-macrophage centers present macrophage centers present in central in the lamellae; tunica thickened and sinus; tunica thickened and convoluted; ovary greatly reduced in convoluted; testes greatly reduced in size. size.

activity. The presence of postovulatory follicles (POF) was interpreted as evidence of recent spawning. Tarpon with gonads in maturity classes 3-8 were considered sexually mature. Monthly mean oocyte diameters and gonadosomatic indices (GSI) were plotted to show seasonal reproductive patterns. Oocyte size was determined by measuring the minimum and maximum diameter of each oocyte within a random subsample of 100 oocytes from each histological section and then averaging the two measurements. Oocytes were measured only if the nucleus was visible, indicating that the section was approximately through the middle of the oocyte. Measurements were made with a digital image-processing system. In addition, minimum and maximum diameters of 10 of the largest oocytes in each histological section were measured to estimate mean maximum oocyte diameter. GSIs were calculated for 409 sexually mature Florida tarpon and 178 sexually mature Costa Rican tarpon as GSI = (GW/(TW-GW))*100 where GW = total gonad weight (g) and TW = total fish weight (g). Total fecundity (the standing stock of advanced yolked oocytes) of 32 Florida tarpon was estimated gravimetrically from subsamples taken from various locations in each ovary. Twenty-six

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fish were subsampled from anterior, middle, and posterior portions of each ovary to evaluate spatial variations in oocyte size within the ovary and between ovaries. Subsamples were preserved in 10% Formalin and later transferred to 70% ethanol. Subsamples of ovary containing approximately 1,000-1,500 late-vitellogenic oocytes were weighed to the nearest 0.01 mg, and total fecundity was calculated based on the mean number of oocytes per gram of ovary. Gonads from all fish sampled for fecundity were in maturity classes 4 or 5, and only late-vitellogenic oocytes >0.5 mm in diameter were counted. Undamaged otoliths from Costa Rican tarpon were weighed to the nearest 0.01 mg and weights were compared to those of Florida tarpon reported by Crabtree et al. (1995). Weights of left and right otoliths were not significantly different (paired t-test, n = 33, t = 0.8237, P = 0.416), so otolith weights were pooled for analysis. If both left and right otolith weights were available for an individual fish, the mean of the two weights was calculated. Linear regressions were fit to log10transformed otolith weight and length data and were compared with a t-test (Zar, 1984). We prepared otoliths of 194 Costa Rican tarpon for age estimation following the methods of Crabtree et al. (1995). Annuli were counted three times by each of two independent readers using compound microscopes. Because there was no significant difference between the mean counts of each reader (paired t-test, n = 87, t = 1.9724, P = 0.052), ages were pooled for analysis. Annulus counts for individual otoliths usually showed some level of variation among readings. We established criteria for accepting or rejecting individual otoliths by calculating a coefficient of variation (CV) = (s/y x 100%) where s = the standard deviation of counts for a given otolith and y = the mean annulus count for a given otolith. We allowed a maximum CV of 12.16% following Crabtree et al. (1995).

RESULTS The 1,469 Florida tarpon we examined ranged 102-2,045 mm in length; the 217 Costa Rican tarpon we examined ranged 880-1,860 mm in length. Among Florida tarpon, 740 (50.4%) of the fish we examined were young-of-the-year or 1-yr-old fish (<400 mm). Body size in tarpon was sexually dimorphic. Sexually mature Florida females (gonad class 3 or greater) had a significantly larger mean fork length (1,681 mm, SD = 139.3, n = 299) and attained a larger maximum length (2,045 mm) than sexually mature Florida males (mean = 1,450 mm, SD = 131.7, maximum length = 1,884 mm, n = 108; t-test, t = 14.991, df = 405, P < 0.001, Fig. 1). Sexually mature Costa Rican females had a significantly larger (t-test, t = 12.694, df = 200, P < 0.001) mean fork length (1,456 mm, SD = 114.0, n = 113) and attained a larger maximum length (1,860 mm) than sexually mature Costa Rican males (mean = 1,254 mm, SD = 110.3, maximum length = 1,550 mm, n = 89, Fig. 1). Standard length, fork length, and total length relationships for Costa Rican tarpon are presented in Table 2, those for Florida tarpon were presented by Crabtree et al. (1995). The slopes of the length-weight equations for male and female Costa Rican tarpon were significantly different (t-test, t = 2.323, df = 204, P = 0.011); Costa Rican females larger than about 1,150 mm were usually heavier than males of the same length. Tarpon from Costa Rican waters were smaller than tarpon from Florida waters (Fig. 1). The largest Florida males (upper quartile = 1,524 mm, largest = 1,884 mm) and females (upper quartile = 1,776 mm, largest = 2,045 mm) were larger than the largest males (upper quartile = 1,320 mm, largest = 1,550 mm) or females (upper quartile = 1,525 mm, largest = 1,860 mm) we observed in Costa Rica. Furthermore, the mean length of sexually mature Florida males (1,450 mm) was significantly larger than that of sexually mature Costa Rican males (1,254 mm; t-test, t = 11.157, df = 195, P < 0.001), and the mean length of sexually mature Florida females (1,681 mm) was significantly larger (t-test, t =

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Figure 1. Length-frequency distributions of sexually mature male and female tarpon from Florida and Costa Rican waters.

15.311, df = 410, P < 0.001) than that of sexually mature Costa Rican females (1,456 mm). We were usually unable to determine the sex of small tarpon (<400 mm). Only 11 of 179 (6.1%) Florida fish ranging 124-400 mm in length and examined histologically could be sexed. The smallest female that we could sex was 343 mm in length and the smallest male was 237 mm. The smallest female that had primary oocytes was 523 mm. Some immature fish without primary oocytes or signs of spermatogenesis were sexed based on general gonad morphology. Ovaries had many finger-like lobes that contained only oogonia in most class-1 ovaries, but they occasionally contained primary oocytes that had little cytoplasm and a nucleus containing a large basophilic nucleolus. In contrast, testes were typically straight along the distal edge and had fewer invaginations. The smallest sexually mature Florida male in our sample was 901 mm in length; the next-smallest mature male was 1,175 mm in length, and all larger males were sexually mature. The smallest sexually mature Florida female was 1,285 mm in length, and all larger females, except one 1,368-mm female, were mature. Only one Florida tarpon, an exceptionally large 7-yr-old female (1,370 mm), younger than 10 yrs was sexually mature. All Florida tarpon older than 10 yrs were sexually mature. All Costa Rican males examined (880-1,550 mm) were sexually mature. One immature, developing Costa Rican female (880 mm, 7 yrs old) was examined; all other females (1,126-1,860 mm) were sexually mature. The youngest sexually mature male in our Costa Rican sample was 7 yrs old; the youngest sexually mature female was 12 yrs old. Small, previtellogenic oocytes (0.02-0.2 mm diameter) were present in all tarpon with developing ovaries (Fig. 2,3). Females with mature ovaries had a second class of vitellogenic oocytes ranging 0.5-0.9 mm in diameter. The largest oocytes observed were

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Figure 2. Oocyte diameter-frequency distributions for female tarpon from Florida waters.

those with migrating nuclei and diameters of approximately 0.9 mm and a few atretic hydrated oocytes with diameters of approximately 1.0 mm. Only a few atretic, hydrated oocytes were observed in spent or partially spent fish, and we did not examine any spawning females containing many hydrated oocytes. Reproduction in Florida appeared to be seasonal, but because Floridas tarpon fishery is seasonal, we were unable to examine any mature fish collected during October-January. More than 80% of the females examined during April-June contained maturing vitellogenic oocytes (gonad class 4; Fig. 4). Tarpon with migrating nuclei (n = 3), atretic hydrated oocytes (n = 2), and partially spent females (n = 16) with both maturing vitellogenic oocytes and POFs were found during May-August. During July, spent and recovering females increased in frequency and by August, represented more than 90% of the females (Fig. 4). Mean diameters of the largest oocytes present in individual ovaries peaked during May and were much lower in February and August (Fig. 5). Mean GSIs were highest during April-July for both males and females (Fig. 6). The decline in female GSIs and mean diameters of the largest oocytes during July and August corresponded with the increased percentage of spent and recovering fish during late summer (Fig. 4).

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Figure 3. Oocyte diameter-frequency distributions for female tarpon from Costa Rican waters.

Spawning in Costa Rican waters was not seasonal. Males with mature testes were present in all months except July, August, and December, when we did not collect any males. Our sample sizes for these months were low because most of the fishing lodges were closed during these months. Most males had mature testes; only one male with maturing testes and two males with recovering testes were found. Females with mature, ripe, partially spent, or spent ovaries were captured in every month (Fig. 4). Partially spent females with POFs and a second batch of maturing vitellogenic oocytes 0.3-0.7 mm diameter (Figs. 3,7) were present in all months except June and August, when sample sizes were low. Only four females with recovering ovaries were found. Most females had either mature ovaries (35%) and contained vitellogenic oocytes of 0.6-0.9 mm diameter (Fig. 3) or were partially spent (22%) or spent (33%) and contained many POFs. Both mean monthly diameters of the 10 largest oocytes present in ovaries (Fig. 5) and GSIs had no seasonal trends (Fig. 6). Total fecundity estimates for Florida tarpon ranged 4.5-20.7 million oocytes and had a significant positive relationship with fish weight (Table 2). Oocyte development among areas within the ovary was homogeneous. We used a two-factor analysis of variance to

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Figure 4. Monthly distributions of gonad classes for sexually mature female tarpon from Florida and Costa Rican waters.

compare oocyte densities with side and position as the effects. The number of late vitellogenic oocytes per gram of wet ovary weight was not significantly different between left and right ovaries (ANOVA, df = 1, P = 0.988) or among subsamples from anterior, middle, or posterior sections of the ovary (ANOVA, df = 2, P = 0.427). Furthermore, we found no significant interaction between side and position from which subsamples were taken (ANOVA, df = 2, P = 0.999). Weights of left and right ovaries from mature (gonad

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Figure 5. Monthly mean diameters of the largest oocytes present in ovaries with standard errors and number of fish examined for sexually mature female tarpon from Florida and Costa Rican waters.

class 4) females were not significantly different (paired t-test, n = 195, t = 1.727, P = 0.086), but left testes from mature (gonad class 4) males were significantly larger than right testes (paired t-test, n = 94, t = 4.912, P < 0.001). Of 194 otoliths from tarpon collected in Costa Rican waters and examined for age estimation, 95 (49%) were judged unreadable by one or both readers and were not assigned ages, and an additional 12 (6%) otoliths were rejected for having a high variation among reads (CV 12.16%); thus, 87 (45%) otoliths were accepted for age estimates. The length-frequency distribution of fish whose otoliths were rejected because they were unsuitable for age estimation was not significantly different from the length-frequency

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Figure 6. Monthly mean GSI with standard errors and sample sizes for sexually mature female and male tarpon from Florida and Costa Rican waters.

distribution of all fish examined (Kolmogorov-Smirnov two-sided test statistic = 0.397, P = 0.997). Tarpon from Costa Rican waters are long-lived; the oldest fish we examined were a 47yr-old male (1,301 mm) and a 47-yr-old female (1,530 mm). The youngest fish examined were a 7-yr-old male (1,030 mm) and a 7-yr-old female (880 mm). There was no significant difference in the average age of males (22.9 yrs) and females (22.4 yrs; t-test, t = 0.106, df = 84, P = 0.779). Most (74%) tarpon caught in this study were 15 to 30 yrs old. We weighed 96 otoliths from Costa Rican tarpon (range = 0.147-0.558 g, mean = 0.351 g, SD = 0.0746) in order to compare them with those from Florida tarpon (Crabtree et al., 1995). The otolith-weight frequency distribution for sexually mature Costa Rican tarpon was not significantly different from that of sexually mature Florida tarpon (KolmogorovSmirnov two-sided test statistic = 0.837, P = 0.496, Fig. 8). Furthermore, the mean otolith weight for sexually mature Costa Rican tarpon (0.353 g) was not significantly different from that of sexually mature Florida tarpon (0.352 g; t-test, t = 0.159, df = 376, P = 0.874), and the weight range of otoliths from sexually mature Florida tarpon (0.180-

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Figure 7. A histological section from an ovary from a 1,498-mm-FL tarpon from Costa Rican waters showing vitellogenic oocytes (VO) and postovulatory follicles (POF). Scale bar = 200 microns.

0.695 g) included that of otoliths from Costa Rican tarpon (0.208-0.558 g). Weights of otoliths from Costa Rican tarpon had a significant positive relationship with fork length (Table 2). DISCUSSION Tarpon from Florida waters appeared to attain larger sizes than those from Costa Rican waters (Fig. 1), but because we relied on anglers to provide fish, our data are probably biased. Most Florida fish were caught during tournaments or were kept as trophies to be mounted by a taxidermist; presumably in both situations anglers selectively kept larger fish. Sometimes Florida tournaments imposed minimum size requirements of up to 50 kg. Since males were typically smaller than females and rarely exceeded 45 kg, our samples from the Florida recreational fishery contained roughly twice as many females as males. In Costa Rica, anglers kept tarpon as trophies or because they wished to participate in our research program. We assume that anglers in Costa Rica, like those in Florida, selectively kept larger fish, but we do not know the extent of this bias in each data set. As in Florida, we examined more Costa Rican females than males, probably a result of the bias towards large fish, but the difference was less than in Florida. It is possible that behavioral differences between anglers from the two areas affected our comparisons of the size of tarpon from the two areas.

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Figure 8. Otolith weight-frequency distributions for sexually mature tarpon from Florida and Costa Rican waters.

In Costa Rican waters, tarpon reached sexual maturity at a smaller size than tarpon from Brazilian or Florida waters. Costa Ricas tarpon fishery principally targeted sexually mature individuals. We examined only a single immature Costa Rican fish and were unable to estimate length and age at sexual maturity; however, the scarcity of immature fish in our collections suggests that sexual maturity was attained at less than 880 mm by males and at less than 1,126 mm by females. In Florida, where more than 50% of our sample consisted of immature fish, the smallest mature male examined was 901 mm and the smallest mature female examined was 1,285 mm. Breder (1944) reported that the smallest ripe female he observed off Floridas Gulf coast was 1,398 mm, but he suggested that female tarpon probably attain sexual maturity at about 1,200 mm. De Menezes and Paiva (1966) reported that tarpon from Brazilian waters begin to attain sexual maturity at lengths of 950 mm for males and 1,250 mm for females. Unfortunately, because relatively few fish in the critical transitional length range of 800-1,200 mm have been examined from any area, we are unable to accurately estimate the length or age at which 50% of the tarpon population reaches sexual maturity. Tarpon from Costa Rican waters showed no seasonal pattern of reproduction, unlike tarpon from other areas. In Florida, tarpon gonadal development was seasonal and peaked during May-July. By August, most females were spent or recovering. Breder (1944) examined tarpon from Floridas Gulf coast and reported that spawning occurs at least during May-July, but he presented little data to support this assertion. Large Florida tarpon were not available during all months, so our evidence for seasonal spawning based on the examination of gonads is equivocal; however, other evidence also suggests seasonal spawning in Florida waters. Back-calculated hatching dates of tarpon larvae from otolith microstructure analysis and larval growth rates suggest spawning during MayAugust off Floridas Gulf coast (Crabtree et al., 1992; Crabtree, 1995). Smith (1980)

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Table 2. Length-length, length-weight, and otolith weight-length regressions for tarpon from Costa Rican waters and the fecundity-weight regression for tarpon from Florida waters. TL = total length (mm), FL = fork length (mm), SL = standard length (mm), WT = weight (kg), OWT = otolith weight (g). Sample fork-length range for length-length regressions was 880-1,860 mm and for length-weight regressions was 880-1,860 mm for females and 880-1,550 mm for males; the sample otolith weight range for the otolith weight-length regression was 0.147-0.558 g; the sample weight range for the fecundity-weight regression was 30-90 kg. Values in parentheses are standard errors. Y = a + bX b (1 SE) 1.0098 (0.0093) 0.8946 (0.0085) 0.9727 (0.0090) 0.8717 (0.0105) 1.1128 (0.0135) 1.0968 (0.0104) 3.2274 (0.1182) 2.8269 (0.1214) 0.3386 (0.0441) 1.0339 (0.1709)

Y FL FL SL SL TL TL log10WT (females) log10WT (males) log10FL log10FEC

X SL TL FL TL SL FL log10FL log10FL log10OWT log10WT

n 215 214 215 213 213 214 116 92 96 32

a (1 SE) 34.3497 (12.3531) 34.0065 (13.3892) 9.9633 2.3232) 45.0217 (16.5797) 96.8629 (17.8513) 66.7341 (14.3338) -8.6713 (0.3737) -7.4459 (0.3758) 3.2861 (0.0208) 5.2838 (0.2966)

0.982 0.981 0.982 0.970 0.970 0.981 0.867 0.858 0.621 0.550

collected tarpon larvae from the Gulf of Mexico and suggested that spawning occurs during June-July. The presence of partially spent females with ovaries containing POFs and advanced vitellogenic oocytes (Fig. 7) in both Florida and Costa Rican waters suggests that tarpon are multiple spawners. In Costa Rica, partially spent females were more common than in Florida and made up 22% of the females we examined (Fig. 4). Recovering females were not abundant (n = 4) in Costa Rican waters, suggesting that females spawned repeatedly throughout the year and had no extended period of inactivity. In contrast, 74% of the sexually mature Florida females examined near the end of the Florida spawning season (August) were recovering. Spent females comprised 33% of our Costa Rican sample. These females often contained POFs, suggesting recent spawning, but did not have an additional batch of maturing vitellogenic oocytes. Spent females either quickly developed a batch of maturing vitellogenic oocytes or left the Rio Colorado area soon after spawning for a period of recrudescence elsewhere. In Florida, the number of recovering females was similar to the number of spent females, but this was not true in Costa Rica. We did not examine any fully hydrated females and were thus unable to estimate batch

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fecundity. If tarpon are multiple spawners, our estimate of total fecundity for Florida tarpon may not accurately represent total annual egg production. We did not validate the annual nature of annulus formation for Costa Rican tarpon. We assumed that the annuli we counted on otoliths of Costa Rican tarpon were annual marks based on the similar appearance of these annuli to those of Florida tarpon. Crabtree et al. (1995) validated the annual nature of annulus formation for 12 out of 18 oxytetracyclinemarked otoliths from Florida tarpon that were 4-9 yrs old. Otoliths from six fish were either unreadable or did not form the expected number of annuli. The annual nature of annulus formation for older tarpon (> 9 yrs) has not been validated. Otoliths of tarpon from Costa Rican waters were more difficult to interpret than those of tarpon from Florida waters. We rejected a relatively high percentage (55%) of the otoliths from Costa Rican tarpon as unreadable. Rejected otoliths had irregular and poorly defined features, which precluded our making precise annulus counts. Crabtree et al. (1995) were unable to read 20.6% of the otoliths of tarpon from Florida waters. The more tropical environment of Costa Rica and the absence of seasonal reproduction probably contributed to the less seasonal nature of otolith growth in Costa Rican fish than in tarpon from more temperate Florida waters. In Costa Rican waters, tarpon attained ages similar to those of tarpon from Florida waters. The oldest Costa Rican male and female we examined were each estimated to be 47 yrs old. Crabtree et al. (1995) reported similar maximum ages for tarpon from Florida waters. The oldest Florida male was 43 yrs old and the oldest female was 55 yrs old. Crabtree et al. (1995) showed otolith weight to be closely related to age for Florida tarpon. The similarity between otolith weights of Costa Rican tarpon and those of Florida tarpon (Fig. 8) supports our view that tarpon from the two areas attain similar ages. In conclusion, the differences in overall size and size at sexual maturity of tarpon from Costa Rican and Florida waters suggest limited movement of fish between the two areas. Additional research is needed to learn if these differences have a genetic or an environmental basis. An implication of these data is that the movement of tarpon within the western North Atlantic may be sufficiently localized for regional differences in life histories to arise. If so, this emphasizes the need for conservation of tarpon habitat and wise management of the fishery in both Florida and Costa Rica, because movement of tarpon into these fisheries from other areas may be limited. ACKNOWLEDGMENTS
We thank the D. Hawley Foundation, the Pate Foundation, Wildlife Conservation International, and the New York Zoological Society for their support. Thanks also to Capt. M. Collins, B. Pate, and A. Carr, III. We also thank the anglers, tournament participants, and taxidermists who provided the specimens examined; in particular, in Florida we thank I. Shaw Taxidermy, Pflueger Taxidermy, the Boca Grande Fishing Guides Association, the Florida Keys Fishing Guides Association, the Gold Cup Tarpon Tournament, the Suncoast Tarpon Roundup, and Millers Marina; in Costa Rica we thank B. Barnes of Casa Mar fishing lodge and T. Staley of the Rio Colorado lodge. Many people participated in portions of this study including R. Bishop, L. Crabtree, F. Cross and other employees of the Florida Game and Freshwater Fish Commission, C. Harnden, D. Harshany, V . Lopez, G. Matute, V . Neugebauer, D. Snodgrass, and C. Stevens. I. Quintero-Hunter and H. Grier provided advice on histology. J. Colvocoresses, P. Hood, S. Kennedy, J. Leiby, R. McBride, M. Murphy, K. Peters, J. Quinn, and K. Sulak made helpful comments that improved the manuscript. Thanks to the Wildlife Service of Costa Rica and the Barra del Colorado National Wildlife

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Refuge for their cooperation. This project was supported in part under funding from the Department of the Interior, U.S. Fish and Wildlife Service, Federal Aid for Sportfish Restoration Project Number F-59; the Florida Marine Fisheries Commission; the Don Hawley Foundation; and the Pate Foundation. Portions of these data were included in a dissertation submitted by E. C. Cyr as partial fulfillment of the requirements of the Ph.D. degree, University of South Carolina. This paper is contribution No. 1055 of the Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina, Columbia.

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