BULLETIN OF MARINE SCIENCE, 66(2): 513–521, 2000

EVIDENCE FOR PROTOGYNOUS HERMAPHRODITISM IN THE SERRANID FISH DRUMMONDHAYI (: ) FROM THE CAMPECHE BANK IN THE SOUTHERN

Thierry Brulé, Teresa Colás-Marrufo, Armin Tuz-Sulub and Christian Déniel

The speckled hind, or Kitty Mitchell , Epinephelus drummondhayi (Goode and Bean, 1879), with a maximum total length of about 110 cm and a maximum weight of 30 kg (Matheson and Huntsmann, 1984; Bullock and Smith, 1991; Heemstra and Randall, 1993), occurs in , along the U. S. Atlantic coast from North Carolina to Florida, and in the eastern and northern Gulf of Mexico (Bullock and Smith, 1991; Heemstra and Randall, 1993). Ross (1988), however, emphasized that most biological data on the spe- cies was from the Carolinas, and that information on speckled hind life history and distri- bution from Bermuda and the Gulf of Mexico was scarce. This grouper is found south of Cape Fear on the continental shelf of North and South Carolina at depths of 50 to 100 m (Chester et al., 1984; Huntsman et al., 1994). Juveniles have been caught at depths as shallow as 30 m, and adults commonly inhabit offshore rocky bottom at 60 to 120 m (Bullock and Smith, 1991; Heemstra and Randall, 1993). Since the early 1970s the speckled hind has been relatively rare off the southeastern U.S. (Manooch, pers. comm.). For instance, North and South Carolina headboats have landed an annual average of 11,908 kg (1972–1980), and South Carolina hook and line commer- cial vessels an annual average of 7488 kg (1976–1981) (Matheson and Huntsman, 1984). Speckled hind are rare or absent on many eastern and northern Gulf of Mexico reefs (Smith et al., 1975; Smith, 1976; Hoese and Moore, 1977). On the west Florida shelf, most commercial catches are obtained from depths of 50 m or more (Bullock and Smith, 1991). Records of E. drummondhayi from the southern Gulf of Mexico have been reported only by Springer and Bullis (1956) and Roe (1976) who collected some specimens on the Campeche Bank, off the northern coast of the Yucatán Peninsula. However, subsequent contributions to the identification, distribution and biology of from western Atlantic waters have overlooked Springer and Bullis’, and Roe’s reports of speckled hind from the Campeche Bank (Smith,1971, 1997; Smith, 1976; Fischer, 1978; Bullock and Smith, 1991; Heemstra and Randall, 1993). This area, together with the Texas, Louisiana, and western Florida coasts, have the widest continental shelves in the Gulf of Mexico and major reef fish fisheries (Stevenson, 1981). Fish checklists for catches of the Mexican commercial longline fishing boats of Yucatán sometimes name this species (SEPESCA- INP, 1994; Ríos-Lara et al., 1996). However, an abundance index for speckled hind could not be calculated from these statistics as the Mexican catches lists normally do not distin- guish the various grouper species, combining them all into the category “mero”. As part of a general grouper offshore survey program carried out between 1996 and 1999 on the Campeche Bank, various speckled hind specimens, locally called “lenteja”, were identified in Mexican commercial catches. The purpose of this study was to provide data on the reproduction and regional distribution of E. drummondhayi from the southern Gulf of Mexico. 513 514 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000

Figure 1. Map of the Campeche Bank showing the geographic distribution of fishing locations sampled, and records for Epinephelus drummondhayi observed during the period 1996-1999. Fishing locations: 1— April 1996; 2—April and May 1996; 3, 4—May 1996; 5—November 1996; 6, 7— December 1996; 8—January 1997; 9—January and February 1997; 10, 11—February 1997; 12— May 1997; 13—June 1997; 14—June and July 1997; 15—July 1997; 16—July and August 1997; 17, 18—August 1997; 19—September 1997; 20—September and October 1997; 21—October and November 1997; 22—November 1997; 23—December 1997; 24, 25—January 1998; 26—January and February 1998; 27—February and March 1998; 28, 29—March 1998; 30, 31—June 1998; 32—July 1998; 33—August 1998; 34—August and September 1998; 35—September 1998; 36— April 1999; 37, 38—May 1999.

MATERIAL AND METHODS

Between April 1996 and April 1999, 68 speckled hind from the Campeche Bank were analyzed (size range = 31–96 cm TL). A total of 66 specimens, from 38 fishing locations off the northern coast of the Yucatán Peninsula, were collected from the commercial catches of three offshore fish- ing boats (Fig. 1). The fish were caught with longlines on rocky bottoms. Data on trip duration, capture depth and fishing location were obtained from the captain of each boat. Two gutted speci- mens (33 and 38 cm TL; 0.56 and 1.0 kg GW) were also purchased from the market at the port of Progreso in June and August 1998; the date, location and depth of capture of these fish could not be determined. During onboard collection, speckled hind were easily identified by its distinctive color pattern. According to Rivas (1964), Smith (1971) and Bullock and Smith (1991), this coloration is sufficient to distinguish this species from all other western Atlantic groupers. Fish were measured for total length (TL) and standard length (SL), and weighed for whole weight (WW) and gutted weight (GW). The gonads of 47 specimens were removed for histological study. These were weighed to the nearest gram (gW), preserved in Bouin’s fluid, embedded in paraffin, sectioned to 6 μm thickness, and stained using Gabe and Martoja’s triple stain (Gabe, 1968). Sex and reproductive condition were determined from histological sections of the preserved gonads. They were then classified according to sexual maturity criteria defined by Brulé et al. (1999) for the red grouper, E. morio. The reproductive condition of specimens analyzed through histological study was further examined by calculating a gonosomatic index (GSI = 100 × gW/GW). NOTES 515

Table 1. Records of speckled hind from the southern Gulf of Mexico (Campeche Bank) during the period 1996−1999. n = number of fish sampled.

Dgate Fhishin DNept LWato Lnongo Size rang e l)ocation ()m T)L (cm WW (kg 22 Apr−426May 1996 1'4 2'3° 00 867° 15 1 406−86 1.7−14. 20−217 Feb 1997 135'2'2° 48 847° 32 382−34 0.6−0. 3−138 Jun 1997 146'2'2° 25 837° 30 337−67 0.8−5. 2n7 Ju −145 Jul 1997 195'2'2° 56 837° 29 400−68 1.2−4. 6−293 Sep 1997 116'4-7 2'2° 35 867° 20 301−39 0.5−1. 13−229 Oct 1997 286'2'2° 55 847° 35 313−41 0.7−1. 27−391 Mar 1998 2-2'3° 20 '818° 15 681 3. 5−101 Jun 1998 344'2'3° 15 817° 49 375 0. 242 Aug-4 Sep 1998 351'6 2'3° 37 857° 45 1 407−87 1.5−14. 11−163 Apr 1999 351'6 2'3° 38 837° 40 1 539−96 3.7−17.

RESULTS

The 66 speckled hind from offshore commercial catches were caught at 10 fishing locations situated in the northeastern part of the Campeche Bank, between 22o25'N– 23o38'N and 87o15'W–88o15'W (Fig. 1). The species’ depth range extended from 44 to 165 m (Table 1), with the largest specimens (TL > 70 cm) caught only at the deepest fishing locations, between 146 and 165 m depth (locations 2, 34 and 36). Of the 47 histologically-examined speckled hind, 33 were female, two were classified as transitional individuals, and 12 were male. Size and weight ranges were 33–77 cm TL and 0.67–8.0 kg WW for females and 64–96 TL and 4.6–16.0 kg WW for males. Both transitional individuals measured 73 cm TL and weighed 7.2 and 7.8 kg WW (Fig. 2). In May 1996, August 1998 and April 1999, 12 females ranging from 56 to 77 cm TL, were found to be ripening with vitellogenic oocytes in their ovaries (GSI range = 0.19–6.64%)

Figure 2. Size frequency distribution for females, transitional individuals, males and unsexed fish (i.e., specimens for which gonads could not be examined by histology) for Epinephelus drummondhayi caught on the Campeche Bank between April 1996 and April 1999 (n = 66). 516 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000

Figure 3. Photomicrographs of sections from Epinephelus drummondhayi gonads. a- ripening female collected 4 May 1996, TL = 66 cm, with stage V oocytes (late secondary vitellogenesis) in ovary. b- Ripe-running male collected 28 August 1998, TL = 87 cm, showing some spermatocyte cysts and intralobular sinuses full of spermatozoa in testis; note presence of lamellae and lumen. c- Ripe- running male collected 25 August 1998, TL = 82 cm, with all stages of spermatogenesis in testis; note presence of lamellae, lumen and sperm sinus full of spermatozoa in tunica. d- Transitional collected 25 August 1998, TL = 73 cm, showing stages I and II oocytes (previtellogenic) and scattered spermatocyte cysts. Scale bar = 0.1 mm. G, granulosa; IL, intralobular sinus; L, lumen; LA, lamellae; N, nucleus; PO, previtellogenic oocyte; SPC, spermatocytes; SPZ, spermatozoa; SS, sperm sinus; T, tunica; VO, vitellogenic oocyte; YG, yolk globule; YV, yolk vesicle; Z, zona radiata. NOTES 517

(Fig. 3A). Two ripe-running females with hydrated oocytes (69 cm TL; GSI = 3.32%) or vitellogenic oocytes and postovulatory follicles (74 cm TL; GSI = 6.06%) in ovaries were observed in April 1999. The remaining females, ranged in size from 33 to 71 cm TL, were found to be immature or resting with previtellogenic oocytes in their ovaries in April and May 1996, June 1997, March, June and August 1998, and April 1999 (GSI range = 0.03– 0.74%). All males were observed to be ripening or ripe-running in April and May 1996, August 1998, and April 1999, with cysts of spermatocytes, intralobular sinuses full of spermatozoa, a sperm sinus in the testicular tunica and a membrane-lined central cavity in the testes (GSI range = 0.04–0.21%) (Fig. 3B,C). The gonads of transitional individu- als, caught in August 1998 (GSI = 0.08%) and April 1999 (GSI = 0.16%), contained previtellogenic oocytes, some undergoing atretic, and some nests of spermatocytes (Fig. 3D). Sexually active females and males, and transitional individuals, were all caught at a depth greater than 146 m (locations 2, 34 and 36).

DISCUSSION

No evidence of hermaphroditism in E. drummondhayi has been published to date. A total absence of males in small size categories (<60 cm TL) for Campeche Bank speckled hind is not (by itself) strong evidence for hermaphroditism. However, the identification of features that are strongly indicative of protogyny (Sadovy and Shapiro, 1987) in the gonadal morphology of the Campeche Bank specimens, such as transitional individuals, a membrane-lined central cavity in the testes and a sperm sinus in the testicular tunica, suggests that E. drummondhayi is protogynous. Other western Atlantic groupers for which reasonable evidence of protogyny has been published are: fulva and C. cruentata (Smith, 1959; 1965), E. guttatus (Smith, 1959; Sadovy et al., 1992; Shapiro et al., 1993; Sadovy et al., 1994), E. flavolimbatus (Bullock et al., 1996), E. morio (Moe, 1969; Valdés and Padrón, 1980; Johnson et al., 1998; Brulé et al., 1999), bonaci (Smith, 1959; García-Cagide and García, 1996; Crabtree and Bullock, 1998), M. interstitialis (Bullock and Murphy, 1994), M. microlepis (McErlean and Smith, 1964; Collins et al., 1987; Hood and Schlieder, 1992), M. tigris (Smith, 1959; García-Arteaga et al., in press), and M. venenosa (García-Cagide and García, 1996). According to Sadovy and Colin (1995), sexuality in E. striatus is essentially gonochoristic, with potential for sex change. In the present study, based on the observation of sexually active males and females in the catches, spawning appears to occur in the Spring (April, May) and Summer (August), which is consistent with information in previous studies. For example, Bullock and Smith (1991) collected ripe speckled hind (50–80 cm SL) along the West Florida Shelf in Au- gust, and Heemstra and Randall (1993) stated that spawning for this species occurs from July to September. Gilmore and Jones (1992) observed ripe individuals of E. drummondhayi, off the east coast of Florida, between January to March and between late August and November. This grouper was syntopic with M. microlepis and M. phenax aggregations which occur on the shelf edge reef formations at depths greater than 70 m. Recorded size at first sexual maturity also agrees with previous reports. The smallest active female sampled on the Campeche Bank was 56 cm TL, which is within the 45 to 60 cm range of total female length at maturation reported by Heemstra and Randall (1993). 518 BULLETIN OF MARINE SCIENCE, VOL. 66, NO. 2, 2000

Age and growth data reported by Matheson and Huntsman (1984) for speckled hind on the United States Atlantic coast were used to estimate age of the Campeche Bank speci- mens, with results of from 2 to 12 yrs of age for females, from 8 to at least 15 yrs for males, and 11 yrs for transitionals. All specimens exhibited the distinctive coloration of dark reddish brown with white spots described by Rivas (1964), Smith (1971) and Hoese and Moore (1977). Xanthic speckled hind specimens, such as those described by Schwartz (1978), Smith and Bul- lock (1979) and Ross (1988), were not observed. Ross (1988) noted that all speckled hind smaller than 18 cm TL were xanthic, all specimens larger than 34 cm TL exhibited the typical darker color, and that both yellow and reddish brown individuals occurred be- tween 18 and 34 cm TL. The individuals analyzed in this study ranged in size from 31 to 96 cm TL, perhaps explaining the lack of xanthic specimens. Before the present study, few speckled hind had been reported in the Campeche Bank area. Smith (1971; 1997), Fischer (1978), Bullock and Smith (1991) and Heemstra and Randall (1993) reported no record of speckled hind from the southern Gulf of Mexico. Smith (1976), while compiling available reef fish geographical distribution data from the southeastern United States’ Altantic coast, Bermuda, the Gulf of Mexico, and the Carib- bean , noted that E. drummondhayi was not reported in the southwestern Gulf shelf (Cabo Rojo to Cabo Catoche, Mexico), the western Caribbean shelf (Cabo Catoche, Mexico to the Gulf of Venezuela) and the West Indies (i.e., insular localities within the Caribbean Sea, including the Bahamas). Exceptions include the speckled hind specimens identified by Springer and Bullis (1956) in collections from the Gulf of Mexico and adjacent . These had been caught by handlines on the Campeche Bank (23o39’N and 88o01’W), on rocky bottoms at a depth of 82 m. Also, Roe (1976), in a review of 24 yrs of exploratory fishery and resource assessment activities in the Gulf of Mexico and Caribbean Sea re- ported the capture of four speckled hind (size range: 0.45 to 11.36 kg) from the Campeche Bank (25oN and 85o–93oW), at depths ranging from 47 to 180 m. The depth distribution of captured specimens (44–165 m) in this study agrees with available bathymetric distri- bution data for speckled hind from the western Atlantic, Gulf of Mexico (Springer and Bullis, 1956, Roe, 1976, Miller and Richards, 1980, Chester et al., 1984, Bullock and Smith, 1991; Heemstra and Randall, 1993; Huntsman et al., 1994), and Bermuda (Smith in Bullock and Smith, 1991). Also of note is that the mean size separation for Campeche Bank E. drummondhayi from shallow and deep waters suggests a size-depth distribution similar to that of red grouper, E. morio, from the eastern and southern Gulf of Mexico (Moe, 1969; Brulé et al., 1999). Although, as observed by Chester et al. (1984) on the continental shelf of the South Atlantic Bight, speckled hind from the Campeche Bank tend to occur in slightly deeper waters than other groupers (E. adscensionis, E. guttatus and M. phenax). Groupers are finfish of considerable commercial interest in Yucatán (Brulé and Colás- Marrufo, 1997). Analysis of mexican grouper catches (1996–1997) showed that E. drummondhayi was relatively uncommon on the Campeche Bank and that in importance this species followed E. morio, M. bonaci, M. microlepis, M. phenax, and M. interstitialis (Colás-Marrufo et al., 1998). However, data reported in this work will be useful to design an adequate fishery management strategy for grouper stocks from the southern Gulf of Mexico. NOTES 519

ACKNOWLEDGMENTS

This research was supported by the National Council for Science and Technology (CONACYT) grant no. 2184P-B9507, and the SEMARNAP/S.S.S. 24 DE FEBRERO/CINVESTAV agreement for use of the fishing vessel UNICAP VII. For their assistance during the course of this study, we are grateful to R. Robles de Benito, V. Alcantar-Cárdenas and M. Garduño-Andrade from SEMARNAP/INP-CRIPY; E. González-Gaudiano; J. Peraza-Menéndez and M. Castillo-Martínez from CECADESU/CREDES- Yucalpetén; J. Rodríguez-Félix from S.S.S. 24 DE FEBRERO; A. M. Pech from CONYUC; and L. Contreras- García, the Port Captain of Progreso. For helping with all aspects of the field collections, we thank M. Sánchez-Crespo, V. Duarte-García, S. Mena-González, C. Ureña-Chio, J. Hernández-Viguegas, T. Ramírez-Hernández and P. Mina-Coello. Finally we wish to express our sincere thanks to P. C. Heemstra, C. S. Manooch, III, and C. L. Smith for reviewing the manuscript. We also wish to thank L. Gus- Peltinovich for assistance with photography.

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Date Received: August 11, 1999. Date Accepted: December 2, 1999.

ADRESSES: (T.B., T.C.M. , A.T.S.) CINVESTAV IPN Unidad Mérida, Antigua Carretera a Progreso Km 6, A.P. 73 Cordemex, C.P. 97310, Mérida, Yucatán, México. (C.D.) Université de Bretagne Occidentale, Institut Universitaire Européen de la Mer, Ressources Halieutiques-Poissons marins, Place Nicolas Copernic 29820 Plouzane, Brest, France. CORRESPONDING A UTHOR (T.B.) CINVESTAV IPN Unidad Mérida, Antigua Carretera a Progreso Km 6, A.P. 73 Cordemex, C.P. 97310, Mérida, Yucatán, México. Email .