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Keiji Iwasaki Institute for Natural Science, Nara University enthos Research Vol. 51, No. 2 : 21- 32 (1996) BENTHOS RESEARCH The Japanese Association of Benthology Vertical Distribution and Life Cycle of Two Isopod Crustaceans within Intertidal Mussel Beds Keiji Iwasaki Institute for Natural Science, Nara University ABSTRACT The vertical distribution and life cycle of two isopod crustaceans, Dynoides dentisinus Shen and Cirolana harf oldi japonica Thielemann, were studied at Shirahama, Wakayama Prefec- ture, central Japan, where two mytilids Septif er virgatus (Wiegmann) and Hormomya mutabilis (Gould) formed vertically contiguous mussel beds. The abundance of the isopods tended to increase downshore within the S. virgatus bed in the upper and middle intertidal zones but decreased abruptly within the H. mutabilis bed in the lower intertidal zone. Very few individuals were found within the H. mutabilis bed throughout the year. The size struc- ture of both species of isopods did not differ greatly among five shore levels within the S. virgatus bed. Multiple linear regression analyses against density of D. dentisinus revealed that the amount of sediment within the mussel beds tended to be negatively correlated while the density and volume of mussels both tended to be positively correlated. Both species re- produced twice a year. However, seasonal changes in size structure suggested that the lon- gevity of the two generations born during the year differed between the species: spring- autumn and autumn-spring generations for D, dentisinus, and spring-next spring and autumn-next autumn generations for C. h. japonica. The association between isopods and mussels is discussed with a focus on the presence or absence of sediment within mussel beds. Key words: intertidal isopods, life cycle, mussel beds, vertical distribution INTRODUCTION Salemaa 1979, 1987; Healy & O'Neill 1984; Tuomi et al. 1988; Kroer 1989) and supralittoral Isopod crustaceans are common in a broad isopods of the family Ligiidae (e. g., Willows range of marine habitats, from supralittoral 1987a, b) . However, relatively few studies have shores to deep-sea environments (e.g., been conducted on the ecology of other rocky Harrison 1988; Brown & Odendaal 1994) . Infor- shore species belonging to the Cirolanidae and mation on their biology and the important roles Sphaeromatidae (Wieser 1962, 1963; Glynn they play in demersal communities has been 1965; Holdich 1968, 1970, 1971, 1976; Johnson gathered progressively (e.g., Shafir & Field 1976). 1980; Stepien & Brusca 1985; Brown & Odendaal In Japan, activity rhythms and behaviour 1994) . In particular, on rocky shores there have been investigated for some supralittoral have been a great number of ecological studies isopods of the genera Tylos and Ligia (e. g., on boreal isopods of the genus Idotea (e. g., Ondo 1958a, b, 1959; Ondo & Mori 1956; Imaf uku 1976) . However, information on the Received November 2, 1995 : Accepted April 10, 1996 distribution and ecology of isopods, par- 21 Vol. 51, No. 2 Benthos Research November, 1996 ticularly cirolanids and sphaeromatids, has bed, and the shore heights of the sampling sites been fragmental and scanty (Sekiguchi 1982, were noted (mean height for the replicates: 1985; Sekiguchi et al. 1981, 1982; Mori & Tanaka +65, +38, + 9 and - 6 cm within the S. virgatus 1989; Iwasaki 1995a), and there have been no bed, - 29 and - 56 cm within the H. mutabilis quantitative studies on the life history or popu- bed) . Mussels, isopods, other invertebrates lation dynamics of intertidal isopods. and sediment within the quadrats were all col- Two isopod crustaceans, Dynoides dentisinus lected. In 1994, sampling was conducted at five Shen (Sphaeromatidae) [Japanese name: levels in the S. virgatus bed and two levels in the Shiriken-umisemi] and Cirolana harf ordi ja- H. mutabilis bed. Triplicate samples were ob- ponica Thielemann (Cirolanidae) [Nise- tained at each level (mean height for the repli- sunahorimushi] , are common species on the cates: +67, +48, +26, +7, -10 cm within the S. intertidal shores of Japan and have been re- virgatus bed, - 31 and - 59 cm within the H. ported to occur abundantly within mussel beds mutabilis bed) . In the laboratory the mussels, (Tsuchiya 1979; Tsuchiya & Nishihira 1985, isopods, other invertebrates, and sediment 1986; Iwasaki 1995a) and among barnacles were separated from one another. Surface (Mori & Tanaka 1989) . The present paper re- water was blotted from the mussels, and their ports the vertical distribution and life cycle of volume was measured by putting them into a the two species inhabiting mussel beds graduated cylinder with sea water. All animals consisting of Septif er virgatus (wiegmann) were counted and weighed, and the size of all [Murasaki-inkogai] and Hormomya mutabilis isopods (width of 1st pereonite) was measured (Gould) [Hibarigai-modoki] . Information on to the nearest 0.05 mm. Accurate and prompt the breeding season and brood size of gravid measurements of body lengths in both isopod females is also given and compared with that species were difficult because specimens fixed for other rocky shore isopods. in formaldehyde solution were rolled up. Since the width of the 1st pereonite was highly corre- STUDY STITE AND METHODS lated with body length for both species (April 1994; D. dentisinus: BL (body length) = 2.152 The study site was a sandstone rocky reef on a Pw (1st pereonite width) -0.045, r - 0.988, p < moderately wave-exposed shore near the Seto 0.001, n = 50, C. h. japonica: BL = 2.977 Pw - Marine Biological Laboratory, Kyoto Univer- 0.139, r = 0.967, p < 0.001, n- 50), the former sity, wakayama Prefecture (33° 42'N,135° 21'E). could represent body size adequately. Addi- The rocky reef had a very smooth, flat surface tionally, males and females were discriminated which sloped gently from the upper intertidal from each other and from juveniles by the pres- (80 cm above mean tide level) to subtidal zones ence of penes and oostegites (or oostegite with ca. 5° inclination. Shore height is hereafter buds), respectively. Males of D. dentisinus expressed in cm preceded by + or -, indicating have a median process on the dorsal surface of above or below mean tide level. The tidal range the pleons, and the lengths of these processes at this site extends from +110 to -110 cm. The were measured because the length relative to mussel S. virgatus formed a bed from the upper body size changed seasonally, supposedly re- to middle intertidal zone, and H. mutabilis in flecting their reproductive maturity. Eggs or the lower intertidal zone. The two mussel beds juveniles within the marsupia of mature females were vertically contiguous with a very narrow were counted. Sediment was dried under the zone of overlap. A detailed map of this study sun for two days and weighed after sifting out site and the zonal distribution of mussels and mussel shell fragments with a sieve with 1 mm other sessile organisms are given in Iwasaki mesh openings. (1994: site B). Monthly sampling was carried out at the Cross-shore sampling was carried out in late middle levels of the respective mussel beds April of 1982 and 1994. In 1982, two quadrats of (+35 cm for S. virgatus, - 40 cm for H. 50 cm2 each were haphazardly tossed onto the mutabilis) from early April, 1982, to early mussel bed at each of four levels in the S. April, 1983, and afterwards, sampling was con- virgatus bed and two levels in the H, mutabilis tinued at four-month intervals until early April, 22 Life cycle of intertidal isopods 1985. The same sampling method as for cross- zone are shown in Table 1. In April, 1982, no shore sampling was employed, and duplicate predictor variable was correlated significantly samples were obtained in each mussel bed. In with the density of D. dentisinus. The coeffi- the laboratory, the same procedures as those in cient of multiple determination was also not the cross-shore samplings were followed. significant. In April, 1994, however, sediment To detect factors affecting the vertical distri- weight was negatively correlated, and density bution of isopods, multiple linear regression and volume of mussels positively correlated, analyses were made for isopod densities in 12 with the density of D. dentisinus. The coeffi- quadrats at six shore levels in April, 1982, and cient of multiple determination was also signifi- in 21 quadrats at seven shore levels in April, cant. The amount of sediment also tended to be 1994. Predictor variables were shore height, negatively correlated with the density of C. h. volume and number of mussels, amount of sedi- japonica, although the correlation was not sig- ment, and density of the predatory polychaetes nificant on either survey date. Arabella iricolor (Montagu) and Nereis To examine the relationship between the den- nichollsi Kott. sities of the two isopod species in April, 1994, residuals of the regressions of the density of D. RESULTS dentisinus on the amount of sediment and den- sity and volume of mussels were calculated (see Vertical change in abundance and size struc- Table 1) , and the correlation between the re- ture siduals and the raw densities for C. h. japonica Figures 1 & 2 show the vertical density pro- were analysed within the S. virgatus bed. No files of D, dentisinus and C. h. japonica. In significant correlation was detected between April, 1982, and April, 1994, densities of both the values ( r = 0.238, n = 15, p > 0.05), so no species tended to increase with decreasing specific association between the distributions shore height within the S. virgatus bed, but a of the two species is indicated. Isopod densities significant negative correlation was detected in April, 1982, could not be analysed in this way only for D, dentisinus in April, 1994 ( r = 0.623, because of the small number of samples. p < 0.05) . Within the H, mutabilis bed, very few The size structure of D.
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