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(Linnaeus) . (Brachyura Pacific Science (1976), Vol. 30, No.2, p. 131-145 Printed in Great Britain Sex Ratio, Size at Reproductive Maturity, and Reproduction of the Hawaiian Kona Crab, Ranina ranina (Linnaeus) . (Brachyura, Gymnopleura, Raninidae) I ANN FIELDING2 and SA~1UEL R. HALEy3 ABSTRACT: Sex ratio and size at reproduction of Ranina ranine (Linnaeus) in Hawaii were investigated. A sample of 1596 Kona crabs collected over 1 year in Hawaiian waters was examined to determine sex ratio and size at reproduction. Males constituted 55 percent of the overall samples and a similar proportion in all size classes. Males attain a larger maximum size than do females and have mature spermatozoa when their carapace length exceeds 60 mm. Secondary sexual characteristics in the male develop at a carapace length ofabout 75 mm. Females are ovigerous from May to September. Most ovarian growth occurs between February and May. In May, at the beginning of the spawning season, the number of eggs ovulated is a function of maternal body size: a 25-percent increase in carapace length is associated with a 200-percent increase in number of eggs ovulated. This is not so later in the spawning season (August-September). Larger females appear to ovulate at least twice each season, withthe primary effort going into the first ovulation. The smallest 5-mm size class in which at least 50 percent of the females are ovigerous during the spawning season is 70.0-74.9 mm in carapace length. The mean minimum size of ovigerous females is 86 ± 8 mm in this dimen­ sion. The spermatheca in females is open to the outside at carapace lengths exceed­ ing 60 mm. Eighteen crabs with carapace lengths less than 65 mm were captured. Half (31.9 mm-42.6 mm) were white in color and were all immature; the remaining half (43.6 mm-61.1 mm) were the usual orange color and all of these exhibited active gametogenesis. This correlation of color with size may be of significance for reproductive behavior. Ranina ranina (Linnaeus), commonly referred reproduction, growth, migrations, sex ratio, and to in Hawaii as the Kona crab, is found size frequency of Kona crabs in Waimea and throughout the Indo-Pacific. In Hawaiian Waialua bays on Oahu, Hawaii. For that report, waters, it is found at depths between 6 m and observations were made over a 3-year period 200 m, where it burrows in sandy ocean bot­ (1966-1969) and were tabulated by month but toms. The systematic position of the family not by year. No indication was given that Raninidae, ofwhich R. ranina is the type genus, samples were taken every month over the 3 was recently reviewed by StevCic (1973). years. It would have been useful if the data had Because R. ranina is fished commercially in been tabulated on a yearly basis, because then Hawaii, Onizuka (1972) investigated the any yearly fluctuations in sex ratio or size fre­ quency might have been determined. Also, in­ I This work constitutes part of a thesis presented by the senior author to the graduate school of the Univer­ formation was not included on the frequency of sity of Hawaii in partial fulfillment of the requirements ovigerous females in various size classes, which for the M.A. degree in zoology. Manuscript submitted would have provided a measure of the size at 27 August 1975. reproduction. Onizuka (1972) stated that rear­ 2 University of Hawaii, Department of Zoology, ing studies indicated that female Kona crabs 2538 The Mall, Honolulu, Hawaii 96822. 3 University of Hawaii, Department of Zoology, ovulate twice in summer, but the sizes of 2538 The Mall, Honolulu, Hawaii 96822. females that ovulated twice were not reported. HP S 30 9 131 132 PACIFIC SCIENCE, Volume 30, April 1976 Our inquiry was directed toward the sex capture. Histologically, males were considered ratio, size distribution, annual reproductive to be mature when the sperm ducts contained cycle, general development, size at reproduc­ mature spermatozoa; females were judged to be tion, and sexual dimorphism of R. ranina in mature when vitellogenesis was apparent in the Hawaii. We were limited in the sampling pro­ ovaries. Fresh ovaries were wet-weighed to the gram to collections made aboard a commercial nearest 0.1 g. crab-fishing boat; site, time, and technique Embryo masses (sponges) were removed were determined by that operation. from the abdomens of ovigerous females, blotted, and weighed. Samples ofapproximate­ ly 150 embryos were taken from each of them, weighed to the nearest 0.001 g, and the exact METHODS numbers of embryos counted. In this fashion, Animals used in this study were collected an average weight per embryo was obtained from Penguin Banks, west of the island of and used to calculate the number of embryos Molokai, Hawaiian Islands, an area of approxi­ in each sponge. mately 350 square miles, which is shallow (aver­ age depth 60 m) relative to surrounding water (average depth 600 m), Large numbers ofKona RESULTS crabs were trapped along the outside edge of the banks at depths of 100-200 m. Sex Ratio and Size Distribution Crabs were caught by entanglement in traps Information on population structure was laid along the bottom on a line. Traps were derived from analysis of 1596 crabs, 55 percent made by stretching two layers of 5-cm nylon of which were males (chi-square test at the mesh over l-meter-diameter metal hoops, and 0.01 level indicates a significant deviation from were baited with tuna scraps. Usually, 80 traps a 1:1 sex ratio). Similar sex ratios were reported were attached to a longline at intervals ofabout by Onizuka (1972) and were also significant at 12 m, and were set for 45 minutes. The animals the 0.01 level. The size frequency of male and were held in running seawater in a large deck female crabs in this study is compared to those box until they were transferred to the running reported by Onizuka (1972) in Figure 2. Males seawater system at Kewalo Marine Laboratory attain a greater maximum size than do females in Honolulu. Ordinarily, on a commercial crab­ and predominate in the larger size classes; fishing boat many undersized crabs and all ovig­ whereas females are predominant in the smaller erous females are released upon capture. size classes. The Kolmogorov-Smirnov two­ However, because population data were being sample test (Siegel 1956: 127-136) indicates gathered, all crabs caught were measured and that the Penguin Banks, Waimea Bay, and sexed. Waialua Bay populations have different size Measurements were made with direct-reading distributions. Waimea Bay has the greatest pro­ metric calipers to the nearest 0.1 mm. Carapace portion of small crabs; Penguin Banks, the length was chosen as the reference measure­ greatest proportion of large crabs; and the ment, and was defined as the distance in milli­ Waialua Bay population lies between these two, meters from the posterior margin of the right with the largest proportion of crabs in the orbit to the central posterior margin of the middle size classes. carapace (Figure lA). Onizuka (personal com­ munication) used the same measurement. Sexual Dimorphism Chelar length was measured along the outer surface of the propodus of the right chela, as The male abdomen is narrower than that of illustrated in Figure lB. the female ofa similar carapace size, even in the The data were arranged into 5.0-mm size smallest crabs caught in this study. All males, classes based upon carapace length. When including immature ones, have the first two available, at least five members of each sex in pairs of pleopods modified into a pair of copu­ each size class were dissected within 3 days of latory organs. A gonopore is found on each The H awaiian Kona Crab- FIELDING ANDH ALEY 133 A j 5 cm F IGURE 1. Ranina ranina (L.). Dorsal view of a mature male, illustrat ing the measurements used in th is study. A, carapace length; B, righ t che1ar propodus length. of the coxae of the fifth pair of legs in males, pace spines, larger chelae, and setae on the pal­ and each gonopore is associated with an ex­ mar surface of the dactyli and propoda of th e ternal papilla. All females have visible gono­ chelae. All of these characteristics become more pores on the coxae of the third pair of legs, and prominent with increasing size. This is illus­ matu re females have an external sperm atheca trated for chela length in Figure 4. The chelar located medially on th e (seventh) sternites be­ propodus in both sexes exhibits positive allo­ tween the third and fourth pairs of legs (Gor­ metric growth (allometric coefficient, a = 1.14) don 1963). How fertilization is accomplished up to a carapace length of about 70 mm, at was not determined. which size the growth of the chelar propodus Males attain a larger maximal size than do in females adjusts to a new allometric rate females (Figures 2, 3). Males of at least 75-mm (a = 1.20). For males larger than 70 mm, this carapace length have larger anterolateral cara- structure continues to increase geometrically. .......... ~ MALES (N=882) 0 25 PENGUIN BANKS FEMALES (N =714) - I, >- 20 1 \ U / \ , Z 1 ,, W 15 " \ :::> \ , C 10 . , w ~ , " I " U- 5 ,I .. .. , --- 0 .. ~ 25 MALES (N =637) , ...... ""\ FEMALES (N=499) ->- 20 WAIALUA BAY·' \ \ u , \ 1 " Z 15 1 w 1 1 :::> 1 10 1 CJ 1 w 1 3: 5 1 o .......... ~ 25 MALES (N =2898) - 20 WAIM EA I', FEMALES (N =2430) >- 1, ' .. u BAY 1 .. .. Z 1 , w 15 1 , :::> 1 \ C 10 W ~ LL.
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