Rev. BioI. Trop., 41 (1): 89-92, 1993

Shell length at sexual maturity and spawning cycle of guyanensis (: ) from Costa Rica

Rafael A. Cruz' y Carlos R. Villalobos' • Escuela de Ciencias Biológicas, Laboratorio de Manglares y Ecología Estuarina. Universidad Nacional. Heredia, Costa Rica. • Escuela de Biología y Centro de Investigación en Ciencias del Mar y Limnología. Universidad de Costa Rica, Costa Rica.

(Rec.25-X-1991. Acep. 20-VilI-1992)

Abstract: A population of Myte/la guyanensis from Costa Rica was studied for 12 months in order to determine shel1 length at sexual maturity, frecuency and duration of the spawning cycle under natural conditions. The smallest mature individual was 18 mm long. The musseIs spawned from September through November and from February through April. Sexes were distinguishable in al1 individuals with a shell length greater than 30 mm. No evidence of protandry or hermaphroditism was observed. Sex ratios oí M. guyanensisdid not differ significantly of 1:1.

Key words: Sexual maturity, spawning cycle, Bivalvia, Mylella guyanensis.

Mussels have been a subjet of great interest MATERIALAND METHODS in recent years due to their increasing commer­ cial importance. Most studies are related 10 the Specimens of were co­ reproductive characteristics of the genus llected from Ochoa Estuary, Colorado de Mytilus (Seed 1976), a species widely distribu­ Abangares, Golfo de Nicoya, Costa Rica ted in temperate waters. In tropical regions, (lOoll' N, 85c6' W). Monthly collections were with the exeption of the works by Carvajal made from May, 1988 through April, 1989. (1969), Lunetta (1969) and Mandeli & Acuña Sample size varied from 46 to 90 mussels. (1975), very liule information does exist with A total of 793 mussels were examined, regard to tropical mussels. of whích 154 were used 10 determine length Mytell a guyanensis Lamark, 1918 is a at first maturity. Four length categoríes we­ common mussel that inhabits the mangrove re used (1) 13.9 - 17.9 mm, (2) 18.0 - 22.0 ecosy stem from Baja California to Perú mm, (3) 22.1 - 26.1 mm and (4) 26.2 - 30.2 (Keen 1971). Individuals are found on the mm. The remaining individuals were em­ muddy slopes of small channels, stuck to ployed to establish sex ratio and the repro­ mangrove tree roots or half buried in mudo To ductive cyele. date, in spíte of íts importance, the only infor­ Hístologíeal sections of the viceral mass and mation available is Sibaja's (1986) macrosco­ mantle were fixed with Bouin. After inelusion pical observatíons on sexual maturity and in parafín, sections were sliced at 7 mn and Bolaños (1988) work on seed collection and staíned with hematoxilín-eosine. A microsco­ growth. The goals of the present study were píe examination of the gonadal tissue was ma­ (1) to define shell length at sexual maturity de before assigning individuals to any of the and (2) to determine the frecuency and dura­ categories of gonadal condition, namely indíf­ tion of the spawning cyele. ferent and mature. 90 REVISTA DE BIOLOGIA 1ROPICAL

Forthe his tologica1 study of the spawning ey­ TABLE 1 ele of adults (>30 mm in length), aIl individuals NumbertVId perce1ÚQge of els.muss MyteDa guyanensis, were assigned 10 a certain category of gonadal in indifferenlor gOlllldicmodwation. stage development, detennined by the presenceand de­ gree of matuIation of gametogeniecells in thefo­ llieles. These eategories are a modifieation of Size Indifferent Mamre (mm) males females those usedforother species (Ropes 1968, Lozada and Reyes 1982, Brousseau 1984). 13.9-17.9 18(90.0%) 2 (10.0%) 0(0.0%) Categoriesof gonadal conditions. Gametogenesis: Stage of multiplieation, 18.9-220 16 (33.3%) 30(60.3%) 2(6.4%) eell arrangement and progressive gametoge­ nesis. In male the alveoli walls appear thin 22.1-26.1 22(4.2%) 24 (49.8%) 22(46.�) and show darkly stained spermatogonia, or 26.2-30.2 0(0%) 16 (42.1%) 22 (57.9%) with most follieles filled with spermatozoa. In females the alveoli eontains a large num­ ber of elongated, staIked oocytes free in the were distinguishable in all size-classes studied alveolar lumen and bases attaehed to the al­ (>30 mm). veolar wall. Annual cycle: The gonad structure of M . Maximun maturity: Gonads are composedof guyanensis fits well the one described by darkly stained spermatozoa with their tails Lozada and Reyes (1982) in Perumytilus pur­ pointing toward the eenter of the lumen, for­ puratus (Lamark 1819). ming eoneentrie bands. Females show ripe The macroscopic and histological study re­ oocytes layingfree in the lumen of the folliele. veals that the annual cycle with periods of They adopta more globular shape. growth, maturity and discharge of gonadic pro­ Spent: Alveoli eontain few spermatozoa in ducts occurssimultaneously in both sexes. the center. the alveolar walls appear thiekened. Mature individuals (gametogenesis, ripe, In females, few oocytes are still attached 10 the and spawning) were observed throughout the alveola and few residual ripe ova may remain year. The largest numberof ripeindividuals oc­ in the alveolar lumen. In both sexes a lot of corred from July through September and from connective interfollicular tissue is presento Decemberthrough January. Although spawning occurred year round it was possible 10 identify RESULTS two heavy spawning periods: one from August through November and another from February Shell length and sexual maturity: Individuals throughApril (Fig. 1). of size-l (Thble 1) weredetermined as inmature in 90 % of eases. The mantIe appears thin and B GAMETOGENESis DRiPE .SPAWNiNG transparent with only eonnective tissue cells; 100 § § § � � � � E � hence, no sexes were identified. In the remai­ E E E ning 10 %, sexual produets were observed at F 17.3 mm in two males. In individuals of size 2 80 (18.0 - 22.0 mm) sexes were identified in 66.7 % of the cases, showing different levels of go­ 60 nadal development Hence, it is possible 10 as­ sert that M. guyanensis mature and spawn at a length between 18 and 22 mm. In individuals of size 3 (22.1 - 26.1 mm) only 4.2 % were mma­ ture, while all individuals of size 4 exhibited different degreesof gonadaldevelopment 449 individulas examined, : �I..-.... . I�----...... Sex ratio: Out of ' 1_____....a.J 227 (50.6 %) were females and 222 (49.4 %) MJ J ASONDJ FMA were males. In the sample, the proportion of fe­ MONTHS males in all size-elasses did not differ signifi­ Fig. 1. Mentbly variatien of deve10pmem of the MytelÚl cantIy from one-half. Mate and female gonads guyan.ensisgenad. CRUZ& vaLALOBOS: Reproductianof Mytella 91

DISCUSSION RESUMEN

Mytella guyanensis is a dioeCious bivalve as Se estudió una población de M. guyanensis indicated by Sibaja (1986). As in other mussels durante doce meses desde el punto de vista re­ (Sastry 1979), males and females can be ma­ productivo.El ejemplar máspequeño clasifica­ croscopically distinguished by. the color of the do como maduro midió 18 mm. Se determina­ gonads. Female gonads exhibit a yellow colora­ ron dos picos de desove al año; uno de setiem­ rlon while the gonads of males are brown, var­ bre a noviembre y el otro de febreroa abril. No yíng in both from light ro dark, depending on se encontró evidencia de hermafroditismo. La the amount of eggs or sperm. proporción sexualfue 1: 1. M. guyanensis is a smct gonocoric species as many other mussels (Seed 1976). As in other REFERENCES species, such as Artica islandica Linnaeus (Ropes and Murawski 1980) and Anadara Bolaños, J. 1988. Estudio preliminar sobre el comporta­ miento del mejillón Mytella guyanensis (L) (Bivalvia: grandis Broderip (Cruz 1986), females of M . Mytilidae), en un parque fijo y una estructura flotante guyanensis require a longer gonadal develop­ en la Isla Chira. Costa Rica. Tesis de Licenciatura, ment period than males. Universidad de Costa Rica. 117 p. Length at fIrSt maturation is similar ro that reported for other mUssels by Lozada et al. Boyden, C.R. 1971. A comparativesludy of Ihe reproducti­ ve cycles of Ihe cockles CerastOlÚrma edule and C . (1971), Griffiths (1977) and Suckaneck (1981) glaucum. J. Mar. Biol. Ass. U. K. 51:605-622. and for mactrids by Ropes (1979). The later author has indicated that length at first matura­ Brousseau, D.J. 1983. Aspects of Ihe reproduction of Ihe tion corresponds ro a fourth or fIfth of fue ma­ mussel Mytill4. edulis (pelecypoda: Mytilidae) in Long Island Sound. Fish. Bull. 81:733-739. ximum total length. In Mytella guyanensis in occurs 000 fact, first maturation at fourth of to­ Brousseau, D. J. 1984. Reproductive cycle of Anomia sim­ tal length, i.e. at between 18 and 22 mm (maxi­ ple;¡; (Pelecypoda: Anomiidae) from Cape Cod, ' muro total lengthis around 79 mm). Massachusetts. Veliger 26:299-304. The sex ratio (1: 1) follows that reported Carvajal, J.R. 1969. Fluctuación de las larvas del mejillón for other tropical mussels (Velez and Perflil perflil (L). Bol. Inst. Oceanogr. Univ. Oriente Martínez 1967) as weU as for temperate spe­ 8:13-20. cies (Sunila 1981, Kautsky 1982, Brousseau 1983). Cruz, R.A. 1986. The reproductive cycle of Ihe mangrove cockle AlU/dara grandis (Bivalvia: Arcidae) in Costa In general, the results of our work are not in Rica. Brenesia27/28:37-47. accordance with those reported by Sibaja (1986). This author proposeda 30 -35 mm total Griffilhs, R.J. 1977. Reproductive cycles in littoral popula­ length for the species to reach first maturation, tions oC Choromylilus meridionalis (Kr.) and a sex ratio of 3.47 females for each male, anda Aulacoeya ater (MoJina) wilha quantitative assesment of gamete production in Ihe fonner. J. Exp. Mar. Biol. single spawning period (April). These differen­ Ecol. 30:53-71. ces are a consequense of Sibaja's method (1986), which was based on macroscopic ob­ Kautsky, N. 1982. Quantitative studies on ganad cycle, fe­ servations of the colour and appearance of the cundity, reproductive output and recruiment in Balthic Mytilusedulis populations.Mar. Biol. 68:143-160. gonads. The method is quite unreliable as has it been pointed out by Kinsgtons (1974), who Keen, M.A. 1971. Sea shells of Tropical west America. compared the results of Boyden (1971) on Stanford University, Stanford.1064 p. Cardium edule (Linnaeus) and Cardium glau­ P. CUln, (poiret) using histological techniques. Kingston, 1974. Studies on Ihe reproductive cycles of Cardium edule and C. glaucum. Mar. Biol. 28:317-323.

Lozada, E, J. Rolleri & R. Yanes. 1971. Consideraciones ACKNOWLEDGMENTS biológicas de Choromylilus chorus en dos sustratos di­ ferentes. Biol. Pesq. Chile. 5:61-108. We thanlcJorge Rodríguez for bissuggestions, Lozada, E. & P. Reyes. 1982. Reproductive biology of a Jorge Alfaro for the critica1 review of the manus­ population of Perumytilus purpuratus at El tabo, Chile cript, and EstrellaLizano for her assistance. (: Bivalvia: Mytilidae).Velíger 24: 147-154. 92 REVISTADE BIOLOGIATROPICAL

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