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POPULATION DYNAMICS and LIFE HISTORY of the Mangrovecrabaratuspisonll (BRACHYURA, GRAPSIDAE) in a MARINE ENVIRONMENT

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POPULATION DYNAMICS and LIFE HISTORY of the Mangrovecrabaratuspisonll (BRACHYURA, GRAPSIDAE) in a MARINE ENVIRONMENT BULLETIN OF MARINE SCIENCE, 45(1): 148-163, 1989 POPULATION DYNAMICS AND LIFE HISTORY OF THE MANGROVECRABARATUSPISONll (BRACHYURA, GRAPSIDAE) IN A MARINE ENVIRONMENT Humberto D[az and Jesus Eloy Conde ABSTRACT Aratus pisonii commonly inhabits the supralittoral zone of roots, branches and canopy of the red mangrove, Rhizophora mangle. Population dynamics and life history of this crab were studied during a 2-year investigation in a marine environment. Adult crab abundance was estimated with a mark-recapture program at 13 sites. Males attain greater carapace widths (CW) than females. The CW frequency distributions were similar through time, being more symmetrical for females, but strongly skewed to the left for males. Females were significantly more abundant. Sex ratios fluctuated within the year; however, there were no significant differences between years or seasons. A V-shaped sex ratio curve as a function of size was found. A continuously low juvenile abundance was detected throughout the study. Ovigerous females were continuously present in the population during the study. Their percentage fluctuated through time, with the highest value in the last trimester, coinciding with the rainy season. Number of eggs per female increased as a linear function of carapace width, having a mean of 11,577 eggs and extremes of 3,724 and 27,134. No conspicuous crab migrations were observed; marked individuals reappeared consistently at the sites where they had been marked even 5 months after marking. Carapace width and length showed a high isometric correlation for both sexes. Females tend to have a longer intermolting time than males, although the difference is not statistically significant. Ara/us pisonii (Milne Edwards) is a grapsid crab commonly found from the supralittoral zone of roots to the canopy of Rhizophora mangle, in the borders of mangrove forests. On the Atlantic coast it is distributed from eastern Florida (USA) to northern Brazil and throughout the Caribbean islands. On the Pacific coast it ranges from Nicaragua to Peru (Rathbun, 1918; Chace and Hobbs, 1969). Despite this species' notorious abundance, wide distribution and potential role as a regulatory agent of the mangrove root fouling community, life history and dynamics of south Caribbean coastal populations have not been adequately stud- ied. Hartnoll (1965) and Warner (1967, 1977) studied most aspects of its life history and some features of the population dynamics at Port Royal, Jamaica. Warner (1967), reported growth rate, fecundity, and reproductive output and proposed a lunar breeding rhythm. Von Hagen (1977) reported additional life history features of this crab in Trinidad, The complete larval development of A. pisonii from the same area was described by Warner (1968), who reported one prezoea, four zoeae and one megalopa; no variability in the number of stages was found. On the contrary, Diaz and Bevilacqua (1987) found distinct larval devel- opmental sequences in A. pisonii larvae from marine and estuarine locations. In such sequences, one and sometimes two zoeal stages had been suppressed. Fur- thermore, Diaz and Bevilacqua (1986) found differential survival of A, pisonii larvae when reared under different salinity conditions, using larvae originating from marine and estuarine mangrove swamps. Warner (1970) studied the behavior of A. pisonii and Young (1972) studied some aspects of its physiological ecology. To partially fill the gap in information on A. pisonii, and at the same time provide a biological data baseline for a national park which sustains an ever increasing tourist pressure, we undertook a long term study of the population dynamics and life history of A. pisonii at Morrocoy National Park, Falcon, Ven- 148 DiAZ AND CONDE: ARATUS PlSONl/ IN MARINE MANGROVES 149 ~""" \ 1 i j / i" i, ..,i ~ MANGROVE FOREST o 1 Km 10"46' 68° 20' Figure I. Locations of study sites at Morrocoy National Park and its relative position on the Ven- ezuelan coast. ezuela. This paper describes the annual and seasonal variations in abundance, sex ratio, fecundity, size distribution, recruitment and biometry of an A. pisonii population over a 2-year span in a marine mangrove system. MATERIAL AND METHODS The Morrocoy National Park, located on the northwest coast of Venezuela (loo52'N-68°l6'W), contains a system of interconnected mangrove lagoons which communicates with the open sea by means of several channels (Fig. I). The surface water salinity throughout the Park fluctuates slightly, 36 ± 2 0/00, during the year (Diaz et aI., 1985). There is no fluvial input and the rain regime has only one peak, during the last trimester of the year (Matteucci and Colma, 1986). During this peak, due to rain runoff, transient estuarine conditions might appear along certain restricted mangrove areas near the mainland. The most conspicuous marine communities of the Park are mangrove forests, coral reefs and Thalassia beds. As along most of the Venezuelan coast, tidal range varies at most 25- 30 em. From October 1977 to September 1979, a mark-recapture program was conducted at 13 sites in the Park, according to the schedule and places indicated in Table I and Figure I. At each site, five cylindric double-entrance traps made of plastic mesh and baited with fish meat were deployed at approximately l-m intervals and fixed on branches and aerial sections of roots of the red mangrove (Rhizophora mangle). Traps were set by the sea margin, where they remained for 5 consecutive days. Each day, traps were inspected for crabs; these were marked by affixing a small, sequentially numbered, autoadhesive paper tag to their carapace. Tags were coated with transparent fingernail hardener. Carapace length and width were measured to the nearest 0.05 mm using a Vernier caliper. Sex was 150 BULLETIN OF MARINE SCIENCE, VOL. 45, NO. I, 1989 Table 1. Collection schedule of Ara/us pisonii at different sites in Morrocoy National Park 1977 1978 ]979 Site Oct Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Nov Jan Mar Jun Sep I X X X X X X X X X X X X X X X X X 2 X X X X X X X X X X X X X X X X X 3 X X X X X X X X X X X 4 X X X X X X X 5 X X X X X X X X X X X X X X X X X 6 X X X X X X X X X X X X X X X X X 7 X X X X X X X X X 8 X X X X X X X X X X X X X X X X X 9 X X X X 10 X X X X II X X X X X X X X X X 12 X X X X 13 X X X X recognized and the presence of eggs was also assessed. Crabs were released at points of capture. Tag numbers were recorded daily. On thc last day of the sampling schedule, ovigerous females were collected in areas close to the sampling sites and preserved in 10% formalin in seawater. At the laboratory, the number of eggs was estimated by means of a volumetric method (Diaz et aI., 1983). Adult population abundance was estimated by means of a modified Peterson estimator, under a Schnabel census sequence (Seber, 1982). To grossly estimate the intermolting times, crabs were picked up haphazardly and confined indi- vidually in cages constructed around roots, including their aerial part. Cages were inspected at irregular time intervals and molting occurrence registered. RESULTS Population Structure. - Throughout the study 2,675 males and 3,480 females were collected and measured for the analysis of population structure. The carapace width (CW) of males ranged from 6.50 to 26.80 mm. For females it ranged from 6.85 to 24.65 mm. The average CW size of males (x ± Sx = 19.45 ± 0.06, n = 2,666) was significantly larger (t = 18.02, P < 0.001) than that of the females (y ± Sy = 18.08 ± 0.04, n = 3,458). The size (CW) distributions of males and females collected during the whole study were significantly different (G = 831.67; P < 0.001). The CW distribution of females had less range and was more symmetrical than that of males (Fig. 2), which was strongly skewed to the left. The mode of the male distribution was shifted more to the right (20-21 mm) than that offemales (17-18 mm). The male and female CW frequency distributions were analyzed month by month by means of Stem-and-Leaf graphs (Tukey, 1977; Conde et aI., 1986). They were, in general, unimodal and slightly asymmetrical (Fig. 3A, B), with more size classes to the left of the median than to the right. Except on January 1978, the size class distributions for males and females for each month were statistically different (G-test, Sokal and Rohlf, 1981). The modal size (CW) class of males fluctuated month by month between 17 and 23 mm, while for females it fluctuated in a more narrow range: 17 to 19 mm. Juveniles were scant in the population (Fig. 3A, B). These graphs, and the boxplots (Fig. 4) reflect the absence of dramatic changes in population structure. Sex Ratio. - The overall female: male ratio (1.30: 1.00) of A. pisonii deviated significantly from the normal Mendelian or 1.00: 1.00 ratio (Gp = 105.59; P < DiAZ AND CONDE: ARATUS PlSONl/ IN MARINE MANGROVES 151 20 FEMALES, n = 3458 18 I 14 12 10 8 <.9w 6 « 4 r- z 2 w u a::: MALES, n = 2670 w 14 0... 12 I 6 4 2 6 7 8 9 10 II 12 13 14 15 16 17 18 1920 21 22 23 24 25 26 CARAPACE WIDTH (mm) Figure 2.
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