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Ren L. Eckert and Scott A. Eckert

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Ren L. Eckert and Scott A. Eckert JOURNAL OF CRUSTACEAN BIOLOGY, 7(4): 682-690. 1987 ren L.Eckert and Scott A. Eckert ABSTRACT Leatherback sea turtles (Dermochelys corracea) nesting at regular time intervals on Sandy Point, St. Croix. U.S. Virgin Islands, provided an opportunity TO obtain multipie measure- ments of Conchoderma wrgaium, a pedunculate epibiotic cimped. Mean capnuiar length for gravid C. virgatum was 12.4 mm (SD = 1.8, range 8.8-1 5.9 mm). A growth curve was predicted after fitting paired measurements (capture and recapture) from 43 individuals to a von Bertalanffv growth interval equation. Estimates of asymptotic length and intrinsic growth rate were made using nonlinear least-squares regression procedures. The predicted asymptotic size of 14.6-rnm capitular length is consideraoly less than that reported elsewhere for the species. It is possible that stress associated with the terrestrial nesting phase of the host prevented the barnacles from attaining full growth potential. Considerable plasticity in both maximum size and intrinsic growth rate may exist between populations exposed to different physio-ecological regimes. Conchoderma vzrgatum (Spengler, 1790) is a lepadomorph or pedunculate cir- riped known from tropical and temperate waters (Stubbmgs, 1967). Geographic distribution records for C. virgaium attached to fishes are summarized by Dawson (1 969). Opportunistic settlement on fishes, whales, sea turtles, crabs, ships, buoys, cables, and other artifacts isolated from shore is summarized by Hastings (1972). Additional host records are spread widely throughout the literature and include mako sharks (Williams. 1978), sea turtles (Annandale. 1909: Chevreux and de Guerne. 1893; Crozier, 19 16; Foster, 1978; Hubbs, 1977: Hughes, 1974; Monroe and Limpus, 1979), sea snakes (Annandale, 1909), whales (Clarke, 1966), pelagic crabs (Jerde, 1967; Moazzam and Rizvi. 1979), parasitic invertebrates (Williams, l978), ships (Dalley and Crisp, 198 1: Darwin, 185 1; Evans, 1958; Zullo, l963), buoys (Annandale, 1909; Il'in et a/.,1978; Tsikhon-Lukanina el a/.,1977; Maclntyre. 1966), and oil platforms (Foster and Willan, 1979). Conchodkrma v;¥rgatzis clearly pelagic: there are no records of it from pier pilings or sub- or intertidal substrates. For this reason. populations are rarely encountered and rigorous studies of the species are lacking. Geographic and tem- poral origins are particularly difficult to establish when settlement occurs on wide- ranging ships, fishes, sea turtles, and cetaceans. Nothing is known of population age structure, fecundity, or survivability. Published estimates of growth rate and asymptotic size vary widely, suggesting that different populations may exhibit different growth trajectories, perhaps in response to varying physio-ecological regimes. In this study, we were able to obtain repeated measurements of individual barnacles from a population of known geographic origin. Leatherback sea turtles (Dermochelys coriacea), journeying from temperate foraging zones to tropical nesting grounds. served as hosts to indigenous C. virgatum that colonized the turtles upon their arrival at the nesting area (St. Croix, U.S. Virgin Islands). The gravid turtles came ashore to nest at regular time intervals, enabling us to doc- ument the spatial and temporal colonization by C. virgatum throughout the nesting season. Our objectives were to document size at sexual maturity, growth rate, and asymptotic length for the barnacle population. The divergence of our results from 684 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 7, NO. 4. 1987 Table 1. Morphometric data for Conchoderma virgatuml collected from leatherback sea turtles (Der- mocwvs coriacea) nesting on Sandy Point, St. Croix, U.S. Virgin Islands, 1983. All barnacles Gravid barnacles onlv2 Dimension IV .t-(mm) SD Range .V x(mm) SD Range Total length 93 13.9 5.8 3.5-25.8 37 19.0 2.8 12.8-25.8 Capituiumlength 93 9.0 3.7 2.5-15.9 37 12.4 1.8 8.8-15.9 Capitulum width 93 6.2 2.6 1.5-1 1.5 37 8.6 1.4 5.1-1 1.5 ' Preserved in 70% eihanol. :Ovigerous lamella present. where Lr is the capituiar length at recapture, a is maximum (asymptotic) length, Lc is the capitular length at first capture. e is the base of the natural logarithm, k is the intrinsic growth rate, and d is the capture-recapture interval in days. Fabens' (1 965) adaptation of the von Benalanffy growth model was developed specifically for situations such as this one where the age of the organism remains unknown. The constraint of unknown age is removed by growth interval equations which allow the estimation of growth curves based solely upon observed growth rates between the capture and recapture of individuals of different sizes (Fabens. 1965; Frazer and Ehrhart, 1985). The von Bertalanffy analysis was chosen over other growth models in order that our results wouid be comparable to an earlier study of C. virgaturn by Dalley and Crisp (198 1). We used a Statistical Analysis Systems (SAS) nonlinear (NLIN) least-squares regression procedure (SAS, Inc.. 1985) to obtain estimates of parameters a and k. The estimates allowed us to generate a growth curve using the more general von BenalanKy equation: where L is the capitular length. b is a parameter related to the size of the barnacle at settlement. and t is age in days. Six species of epibiotic cirripeds were present on the turtles during nesting, including three balanomorphs (Stomatoiepas dermochelys, Platylepas hexastylos, Balanus trigonus) and three lepadomorphs {Conchoderma virgatum, C. aurzzurn, Lepas anatifera). Conchoderma virgatum was the most commonly observed cir- riped, proliferating on tags, carapace ridges, epidermal abrasions, balanornorph cirripeds, and, to a lesser extent. the neck. shoulders, flanks, and interridge zones of the carapace. Front and rear flippers were rarely colonized. It was not possible to evaluate settlement on the ventral side of the turtle. Settlement densities of C. virgatum varied widely. By the end of the nesting season, the species numbered in the tens on some turtles and in the hundreds on others. Clumped individuals were more common than solitary ones. In 1983, 37 (39.8%) of the individuals examined were gravid, that is, they contained ovigerous lamellae (Fig. 1). Morphometric data are summarized in Table 1. The mean capitular length : total length ratio (preserved specimens) was 65.1O/o (N = 9 3, range 47.8-80.2°/0) A strong positive correlation (Pearson's prod- uct-moment) was observed between capitular length and total barnacle length (r = 0.965), as well as between capitular length and width (r = 0.974). Since the maximum capitular length has become the standard index of size for pedunculate barnacles, all references hereafter to the "size" of C. virgatum will refer to capitular and not to total length. From paired measurements obtained in situ (Table 2)' nonlinear regression procedures (SAS. Inc., 1 9 8 5) provided estimates of maximum size (a) and intrinsic growth rate (k) (Table 3). Estimates of a and k were inserted into the von Bertalanffy growth interval equation (I): 686 JOURNALOF CRUSTACEAN BIOLOGY, VOL. 7, NO. 4, 1987 Table 3. Estimates of the parameters a and k from nonlinear least-squares regression of the von Benaianifv growth interval equation [Lr = a - (a - Lc)e-kdHFabens,1965) for Conchoderma virgatwn in Caribbean waters. 0.04 1-0.092 nor the time interval over which growth rate was calculated are consistently reported. For this reason. the values are incommensurable, that is, there is the danger of unwittingly comparing rates representing different life history stages (different points along the growth trajectory) to one another. Discrepancies within known age classes would allow the identification of phenotypic plasticity, but such data are not available. Capitular length at maturation is more easily equated between populations. In 1983, 37 (39.8%) of the C. vlrgatum examined were gravid, exhibiting a mean size of 12.4 mm (SD = 1.8. range 8.8-1 5.9). In contrast, Maclntyre (1 966) reported that C. virgaium from eastern Australia did not contain brood lamellae prior to reaching a capitular length of 11 mm (2 = 15.8 mm, SD = 2.0, 1V = 39). The means differ significantly (P < 0.05) and provide evidence that populations may be recognizable on the basis of reproductive as well as morphometric characters. In the present study. a "maximum average" size (Knight. 1968) of 14.6 mrn was predicted. The largest size attained during the course of the study was 16.7 rnm; this particular barnacie had settled 60-70 days earlier, soon after the host turtle nested for the first time. The observed maximum size agrees niceiy with the confidence interval predicted by the model (Table 3). but falls well below the 28-mm asymptotic size predicted by earlier von Benalanffy models (Dalley and Crisp. 198 1) and maximum sizes reported by other investigators for the same species. These include 13.8 mm (Dawson, 1969), 20 mm (Maclntyre, 1966), 22 mm (Il'in et at., 1978). "rather above one inch long" (approximately 30 mrn; Darwin, 185 l), 60-mm total length (approximately 35-mm capitulum: Beckett, l968), and 80 mm (Foster and Willan, 1979). Furthermore, neither our growth curve nor that of Dailey and Crisp (198 1) can account for the growth reported by Annandale (1 909), Il'in el al. (1 978), or Maclntyre (1 966) (Fig. 3). It is possible that the discrepancies are genetic, but considerable ecotypic and/ or ontogenetlc variation has been reported in this species and proposed subspecies have not received wide acceptance (Zevina, 1982). It is unlikely that the latitude of St. Croix accounted for the diminutive size, since larger sizes have been reported from the tropical Atlantic (Dalley and Crisp, 198 1; Il'in et al.,1978). It is doubtful that something inherent in the turtle (e.g.. dermal chemistry) inhibited the growth of C. virgatum. since individuals settling outside of the breeding area (prior to the nesting season) were as large as 28.0 mm when nesting commenced.
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