<I>Cambaroides Japonicus</I>
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BULLETIN OF MARINE SCIENCE, 61(1): 147–157, 1997 SURVIVAL AND GROWTH OF THE JAPANESE CRAYFISH CAMBAROIDES JAPONICUS IN A SMALL STREAM IN HOKKAIDO Tadashi Kawai, Tatsuo Hamano, and Shuhei Matsuura ABSTRACT Growth over the life cycle, including seasonal variation, of the Japanese crayfish, Cambaroides japonicus (de Haan, 1841), was examined for a population found in a stream in Hokkaido, northern Japan. The growth equation estimated for males was Lt = 106.6 (1- exp (-0.02987 (0.08333t + 0.9444 + 0.3279 sin (0.5236t - 0.6947)))) and for females, Lt = 126.2 (1-exp (-0.02413 (0.08333t + 1.213 + 0.2768 sin (0.5236t - 1.548)))), where Lt is the carapace length (mm) measured from the posterior of the eye socket to the posterior margin of the carapace at age t (months after settlement in July). The longevity of males is esti- mated to be 11 yr, and females 10 yr. Both sexes become sexually mature 5 or 6 yrs after hatching. The Japanese crayfish, Cambaroides japonicus, is a species indigenous to northern Ja- pan, found only in Hokkaido, Aomori, Akita, and Iwate Prefectures (Okada, 1933). It reaches approximately 5 - 6 cm in total length, and has been utilized as a food in Tokachi, Hokkaido. However, as many of the natural habitats of C. japonicus have been destroyed by human activities, aquaculture of this crayfish has been proposed for the future. It is therefore important to know the ecology of this species to assess its suitability in an aquaculture program. Previous studies have been limited to the reproductive cycle and molting season, etc. (Kurata, 1962; Kawai et al., 1994b; Kawai et al., 1995a), and information on the ecol- ogy of C. japonicus is scarce. Thus, we have studied the growth characteristics using growth equations and a life table of C. japonicus in its natural habitat, and the data are compared with other Astacoidea. We briefly discuss the possibility of aquaculture for this species in Hokkaido. MATERIAL AND METHODS Specimens of Cambaroides japonicus were collected from June 1990 through to June 1992, from a stream, near Atsuta, Hokkaido (Fig. 1). The stream flows into Ishikari Bay and is fed by many springs which maintain the water level. The stream is approximately 1 km in length, with a maximum depth of 5 cm and maximum width of 1 m. Collections of crayfish were carried out at 3 - 4 wk intervals throughout the period of the investigation. The total number of samples was 36. Twenty five sampling stations were established in the stream, and sampled with a 1-m2 quadrat. Crayfish were collected by hand from underneath boulders or fallen leaves. We measured carapace length, CL, from the posterior portion of the eye socket to the posterior margin of the carapace, to the nearest 0.1 mm, using hand-held calipers. Sex of the crayfish was determined by the presence or absence of the gonopore at the base of the 3rd pereopods and from the morphology of the 1st and 2nd pleopods following Holdich and Reeve (1988). Gonopore and pleo- pods of “juveniles” less than 10 mm in CL were extremely small and could not be used to determine the sex of these specimens. All of the specimens were released after measurment. Yearly fluctuations in the breeding season and individual densities in the study stream are small (Kawai et al., 1994b) and the authors expected little variation in the yearly growth pattern of C. japonicus. Thus, all the sampling data were summarized by calender month. Since no molting occurred during winter, from November to May, and growth resumed from June in the natural habitat (Kawai et al., 147 148 BULLETIN OF MARINE SCIENCE, 61(1): 147–157, 1997 Figure 1. Map showing the study site in Hokkaido, northern Japan. 1994b), data for the period from November through to May were collected. Since C. japonicus live in a small stream, it was difficult to collect large sample numbers and to make a polymodal histogram from a single year’s sampling. Therefore, we treated each year separately. First, we estimated the growth in the field based on growth information derived in a laboratory which had conditions similar to study habitat (Kawai et al., 1995b). Next, we separated modes by two methods: moving averages and by using no average. Both procedures were compared to estimate growth and the moving-average method provided the better fit. Frequency data were classified by size classes as moving means over three CL-class intervals. The most reasonable width of the class interval, 0.4 mm, was adopted after applying several different widths to the data set. Separation of overlapping age groups was accom- plished using a computer analysis based on the maximum likelihood method (Akamine, 1985; Fournier et al., 1990). Juveniles were also analysed by the method described above. Since the sex ratio of this species is 1:1, irrespective of season and body size (Kawai et al., 1995a), each size class of juveniles was divided equally into males and females. The ordinary von Bertalanffy equation and the sine-wave von Berterlanffy equation (Pauly and Gashutz, 1979) were applied to all data. A computer was used to fit the data by the Marquardt method with Akaike’s Information Criterion (AIC), employed as a measure of goodness of fit (Akaike, 1973). During the regular sampling it was difficult to sample smaller crayfish which hide among small stones and gravel. To calculate the actual density of each year class and to construct a life table, additional sampling was conducted on 14 September 1994. Approximately 50 m2 were sampled with a 1-m2 quadrat placed at 20-m intervals along the stream. The top 5-cm layer of the sediment was collected by a shovel and the samples were transported to the laboratory after being fixed in 10% formalin, where the specimens were sorted from the gravel. Sizes and sexes of the crayfish were determined as described above. RESULTS Mean water temperatures of the stream during the recruitment season, May to June, were uniform: 1990, 15.9° C; 1991; 15.5° C; and 1992, 15.4° C. Therefore, recruitment seems to be constant every year, and females carrying eggs were caught during May and June in both 1991 and 1992 (Kawai et al., 1994b). Carnivorous fish and other large animals were not KAWAI ET AL.: SURVIVAL AND GROWTH OF THE JAPANESE CRAYFISH 149 Figure 2. Size distribution of Cambaroides japonicus in a small stream in Hokkaido. Solid triangles are the mean of the Gaussian distributions described by the dotted curves. The numerals above the triangles indicate the number of months after hatching. observed in the stream. The total numbers of C. japonicus sampled during the regular collections were 1454 males, 1453 females and 318 juveniles. Mean individual density was 3.6 inds. m-2 calculated as (1454 males + 1453 females + 318 juveniles)/(1 m2 x 25 stations x 36). The CL of the largest individual was 34.2 mm for males and 31.6 mm for females. The number of normal distributions of overlapping age cohorts was 9 - 11 for males and 9 - 10 for females in each month. A newly settled cohort appeared in July (Fig. 2). The reproductive season of C. japonicus occurs over the summer (Kawai et al., 1994b) and we believe hatching and settlement occur in July. Modes in the monthly samples were determined from growth information derived from laboratory studies. We estimated the growth in the field based on laboratory observa- tions since the laboratory conditions were similar to stream habitat (Kawai et al., 1995b). Molting frequency per year in the laboratory of C. japonicus of each sex is two or three molts per individual for the < 10 mm CL size class, one or two molts for individuals ≥ 10 and ≤ 18 mm, and only one molt for individuals > 18 mm (Kawai et al., 1995b). Their growth factor, calculated as (100 x post-molting CL)/(pre-molting CL) following Mauchline (1977), was approximately 110, irrespective of sex and body size. Based on this growth 150 BULLETIN OF MARINE SCIENCE, 61(1): 147–157, 1997 Figure 3. The fit of the ordinary Bertalanffy growth equation and the Pauly and Gashutz equation to the mean carapace length data of Cambaroides japonicus collected from in a small stream in Hokkaido. factor, the annual CL growth in the natural habitat was estimated as approximately 1.1 for large individuals and 1.3 for small individuals. In any one month, the mean of each cohort in Figure 2 is equivalent to the CL multiplied by the year-younger cohort by 1.1 - 1.3. Therefore, we suggest that the normal distribution (Fig. 2) is representative of single year- class cohorts. Male: Lt = 106.6 (1-exp (-0.02987 (0.08333t + 0.9444 + 0.3279sin (0.5236t - 0.6947)))) (1) KAWAI ET AL.: SURVIVAL AND GROWTH OF THE JAPANESE CRAYFISH 151 Figure 4. Size distribution of Cambaroides japonicus in substrate samples from 50 m2 in a small stream in Hokkaido on 14 September 1994. Solid triangles are the mean of Gaussian distributions described by the dotted curves. The numeral above triangles denote the predicted year of hatching. n = 69, AIC = -62.45 Lt = 55.47 (1-exp (-0.005782 (t + 7.594))) (2) n = 69, AIC = 9.626 Female: Lt = 126.2 (1-exp (-0.02413 (0.08333t + 1.213 + 0.2768sin (0.5236t-1.548)))) (3) n = 69, AIC = -17.16 Lt = 54.46 (1-exp (-0.005844 (t + 8.059))) (4) n = 69, AIC = 35.64 where Lt is CL at age t (months) after settlement (Fig. 3). 152 BULLETIN OF MARINE SCIENCE, 61(1): 147–157, 1997 Figure 5.