Function of the major cheliped in male–male competition in the hermit crab <Emphasis Type="Italic">Pagurus nigrofascia</Emphasis>

Functions of the major cheliped in pagurid hermit crabs have been studied in fights for shells. The major cheliped often shows sexual size dimorphism, suggesting that sexual selection favors the development of the male major cheliped. The function of the major cheliped in male–male competition was examined in Pagurus nigrofascia collected from April to June 2009 on the intertidal rocky shore in southern Hokkaido, Japan (41°N, 140°E). Sexual size dimorphism of the major cheliped was observed, and precopulatory guarding males had larger major chelipeds than solitary ones. Guarding males used the major cheliped to deter intruders during competitive interactions. Males without a major cheliped were disadvantaged even if they were larger than opponents and had ownership. Cheliped size affected the outcomes of contests between similar sized males. This suggests that the male major cheliped in P. nigrofascia protects mates from competitors and, consequently, enhances male mating success. Sexual selection may favor the development of the major cheliped in male pagurids.     

Comparative life-history study on sympatric hyperiid amphipods (<Emphasis Type="Italic">Themisto pacifica</Emphasis> and <Emphasis Type="Italic">T</Emphasis>. <Emphasis Type="Italic">japonica</Emphasis>) in the Oyashio region,western North Pacific

Life-history features of the sympatric amphipods Themisto pacifica and T. japonica in the western North Pacific were analyzed based on seasonal field samples collected from July 1996 through July 1998, and data from laboratory rearing experiments. T. pacfica occurred throughout the year, with populations peaking from spring to summer. In contrast, T. japonica were rare from autumn to early winter, but became abundant in late winter to spring. Mature T. pacifica females and juveniles occurred together throughout the year, indicating year-round reproduction. Mature T. japonica females were observed only in spring, and juveniles occurred irregularly in small numbers, suggesting limited, early-spring reproduction in this study area. Size composition analysis of T. pacifica identified a total of eight cohorts over the 2 years of the study. Due to the smaller sample size and rarity of mature females (>9.6 mm) and males (>7.1 mm), cohort analyses of T. japonica were not comparable. Laboratory rearing of specimens at 2°C, 5°C, 8°C and 12°C revealed that a linear equation best expressed body length growth by T. pacifica, while a logistic equation best expressed body length growth by T. japoncia. Combining these laboratory-derived growth patterns with maturity sizes of wild specimens, the minimum and maximum generation times of females at a temperature range of 2–12°C were computed as 32 days (12°C) and 224 days (2°C), respectively, for T. pacifica, and 66 days (12°C) and 358 days (2°C), respectively, for T. japonica. The numbers of eggs or juveniles in females marsupia increased with female body length and ranged from 23 to 64 for T. pacifica and from 152 to 601 for T. japonica. Taking into account the number of mature female instars, lifetime fecundities were estimated as 342 eggs for T. pacifica and 1195 eggs for T. japonica. Possible mechanisms for the coexistence of these two amphipods in the Oyashio region are also discussed.Communicated by O. Kinne, Oldendorf/Luhe     

Is <Emphasis Type="Italic">Hippolyte williamsi</Emphasis> gonochoric or hermaphroditic? A multi-approach study and a review of sexual systems in <Emphasis Type="Italic">Hippolyte</Emphasis> shrimps

Sexual systems vary considerably among caridean shrimps and while most species are gonochoric, others are described as sequential protandric hermaphrodites or simultaneous hermaphrodites with an early male phase. At present, there is confusion about the sexual system exhibited by several species mostly because those studies attempting to reveal their sexual system draw inferences solely from the distribution of the sexes across size classes. Here we investigated the sexual system of the shrimp Hippolyte williamsi from Chile to determine if the species is protandric or gonochoric with sexual dimorphism (males smaller than females). Morphological identification and size frequency distributions indicated that the population comprised small males, small immature females, and large mature females, which was confirmed by dissections. No transitional individuals were found. Males maintained in the laboratory molted 1–8 times, and many grew up to reach sizes observed in only a small fraction of males in the field. No indication of sex change was recorded. Our results indicate that H. williamsi is a sexually dimorphic gonochoric species and emphasizes the importance of using several kinds of evidence (size measurements, growth experiments, morphological dissections, and histological studies) to reveal the sexual system of Hippolyte species. Whether the observed size dimorphism between males and females in many species of Hippolyte is expression of contrasting sexual and natural selection, and whether divergent sexual fitness functions can contribute to the evolution of hermaphroditism remains to be revealed in future studies.     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Effects of temperature on activity,food consumption rates,and gut passage times of seaweed-eating<Emphasis Type="Italic"> Tegula</Emphasis> species (Trochidae) from California

Previous feeding studies on herbivorous marine snails rarely have focused on temperature effects on food intake. If temperature affects food intake, ectothermic snails may experience difficulty obtaining sufficient nutritional resources, limiting their ability to sustain populations at suboptimal temperatures. We hypothesized that the feeding responses of Tegula species would correspond with temperatures characteristic of their geographic distributions. We determined activity, consumption rates, and gut passage times at 11°C, 15°C, 19°C, and 23°C for three Tegula species with distinct thermal distributions: T. brunnea (cold water), T. aureotincta (warm water), and individuals from warm- and cold-water populations of T. funebralis, a broadly distributed species. Activity and consumption rates of T. aureotincta increased with increasing temperature, but were highest for T. brunnea at 19°C, a temperature rarely achieved in habitats occupied by this species, and lowest at 11°C. Warm-water T. funebralis showed significantly lower activity and consumption rates at 11°C, whereas cold-water T. funebralis consumed food fastest at 15°C and were most active at 23°C. Temperature affected gut passage time only in T. aureotincta. These data suggest that temperature might influence the northern limit of T. aureotincta by affecting activity and food consumption rates. T. brunneas activity and ability to consume food were not hindered by warmer temperatures despite the present day restriction of this species to colder waters. Also, widely separated (>300 km) T. funebralis populations may be adapted to regional conditions based on the different temperature responses of northern and southern snails.Communicated by P.W. Sammarco, Chauvin     

Genetic variation among life-history stages of mussels in a<Emphasis Type="Italic"> Mytilus edulis–M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis> hybrid zone

The mussel species Mytilus edulis L. and M. trossulus Gould coexist and hybridize throughout a large area that includes the north coast of Maine and Atlantic Canada. Previous studies provided genetic evidence for limited hybridization between the two species for mussels >15 mm. The present study used two genetic markers (ITS, Glu-5) to examine the genetic composition of early life-history stages by sampling veliger and pediveliger larvae, juveniles (<2.0 to 15.0 mm shell length) and adults (>15 mm shell length) in Trinity Bay, Newfoundland, during three consecutive years (1995–1997) to determine if differential mortality limits the relative abundance of hybrids. The relative frequency of the two species and the different hybrid genotypes was similar among the larvae, juveniles and small adult mussels. The double hybrid genotype (F₁-like) was the rarest genotype observed. There was no evidence for differential mortality during the early life-history stages, and factors limiting production of hybrids appear to operate before the late larval stage. The observed frequency of hybrids is probably due to a combination of pre- and postzygotic reproductive isolating mechanisms operating early in the mussels life history. M. trossulus dominated the early life-history stages, possibly due to a higher population density and a greater reproductive output than M. edulis. Differential mortality may explain the observed decrease in frequency of M. trossulus and increase in frequency of M. edulis with increasing shell length. A similar frequency of hybrid mussels from larvae to the size class of 55 mm shell length may indicate a rate of mortality intermediate between the two parental species. The M. edulis–M. trossulus hybrid zone appears to be maintained by reproductive isolating mechanisms limiting the production of hybrids and life-history differences that allow the two species to coexist.Communicated by J.P. Grassle, New Brunswick     

Molecular and morphological analyses of the cuttlefish<Emphasis Type="Italic"> Sepia apama</Emphasis> indicate a complex population structure

The giant Australian cuttlefish Sepia apama Gray, 1849 annually forms a massive and unique spawning aggregation in northern Spencer Gulf, South Australia, which has attracted commercial fishing interests in recent years. However, many basic life-history characteristics of S. apama are unknown, and anecdotal evidence suggests that there is more than one species. The present study assessed the population structure and species status of S. apama using data from allozyme electrophoresis, microsatellite loci, nucleotide sequences of the mitochondrial COXIII gene, multivariate morphometrics and colour patterns. Analyses of allozyme and microsatellite allele frequencies revealed two very divergent but geographically separated populations consisting of specimens from the east coast and southern Australia. However, the presence of a heterozygote in a putative contact zone between the east coast and southern Australia suggested that these populations were not reproductively isolated. Mitochondrial haplotypes seem to have introgressed further north into the contact zone than have nuclear alleles. Differences in colour patterns that previously had been attributed anecdotally to different geographic populations were, in fact, correlated with sexual dimorphism. These data are most consistent with S. apama being one species the populations of which were geographically isolated in the past (historical vicariance) and have come into secondary contact. Comparison of microsatellite allele frequencies among four South Australian samples indicated significant deviations from panmixia. South Australian samples were also reliably diagnosed by means of multivariate morphometrics. Significant differences in mantle length were observed among populations.Communicated by M.S. Johnson, Crawley     

Effect of sexual maturation on the tissue biochemical composition of<Emphasis Type="Italic"> Octopus vulgaris</Emphasis> and<Emphasis Type="Italic"> O</Emphasis>.<Emphasis Type="Italic"> defilippi</Emphasis> (Mollusca: Cephalopoda)

Changes in the protein, lipid, glycogen, cholesterol and energy contents, total amino acid and fatty acid profiles of Octopus vulgaris and O. defilippi tissues (gonad, digestive gland and muscle) during sexual maturation (spermatogenesis and oogenesis) were investigated. Both species showed an increase of amino acids and protein content in the gonad throughout sexual maturation (namely in oogenesis), but allocation of these nitrogen compounds from the digestive gland and muscle was not evident. The major essential amino acids in the three tissues were leucine, lysine and arginine. The major non-essential amino acids were glutamic acid, aspartic acid and alanine. With respect to carbon compounds, a significant increasing trend (P<0.05) in the lipid and fatty acid contents in the three tissues was observed, and, consequently, there was also little evidence of accumulated lipid storage reserves being used for egg production. It seems that for egg production both Octopus species use energy directly from food, rather than from stored products. This direct acquisition model contrasts with the previous model for Octopus vulgaris proposed by ODor and Wells (1978: J Exp Biol 77:15–31). Most of saturated fatty acid content of the three tissues was presented as 16:0 and 18:0, monounsaturated fatty acid content as 18:1 and 20:1 and polyunsaturated fatty acid content as arachidonic acid (20:4n-6), eicosapentaenoic acid (20:5n-3) and docosahexaenoic acid (22:6n-3). Though cholesterol is an important precursor of steroid hormones, this sterol content exhibited variations that do not seem to be related with the maturation process. Moreover, significant differences (P<0.05) were obtained between genders, suggesting that perhaps there is a greater physiological demand for cholesterol during spermatogenesis than oogenesis. If the component sterols of octopus are of a dietary origin, considerable variation in the cholesterol content between species might be expected on the basis of the sterol composition of their prey. The glycogen reserves increased significantly in the gonad and decreased significantly (P<0.05) in the digestive gland and muscle of O. vulgaris (these trends were not evident in O. defilippi). Glycogen may play an important role in the maturation process and embryogenesis of these organisms, because carbohydrates are precursors of metabolic intermediates in the production of energy. It was evident that sexual maturation had a significant effect upon the gonad energy content, but the non-significant energy variation (P>0.05) in the digestive gland and muscle revealed no evidence that storage reserves are transferred from tissue to tissue. The biochemical composition of digestive gland and muscle may not be influenced by sexual maturation, but rather by other biotic factors, such as feeding activity, food availability, spawning and brooding.Communicated by S.A. Poulet, Roscoff     

Factors affecting the burrowing behaviour of Helice tridens (Grapsidae) and Macrophthalmus japonicus (Ocypodidae) in an estuary of northeast Japan

Helice tridens (De Haan) (Grapsidae) and Macrophthalmus japonicus (De Haan) (Ocypodidae) build separate burrows in reed marshes and muddy tidal flats, respectively, in the brackish-water estuaries of northeast Japan (38°11N, 141°48E). Habitat segregation between burrows of these two species was analysed by comparing the density of burrows of both species in their natural habitats, and on tidal flats subjected to various types of artificial treatment, in the summer of 1981 and 1982. Baskets, containing many stones, were placed on the ground in the reed marsh, tidal flat and creek and attracted many individuals of H. tridens, but not M. japonicus. When stones were placed on an area of tidal flat, H. tridens frequently formed burrows at the border between the stones and mud, suggesting that burrowing of H. tridens was related to the presence of solid substances lying on the ground, such as stones and shoots of the reed Phragmites australis (Cav.) Trin. In closed systems on the tidal flat and reed marsh, H. tridens and M. japonicus were able to construct burrows in both substrata regardless of high or low frequency. Moreover, it was recognized that H. tridens prevented the burrowing of M. japonicus in these closed systems. These results suggested that habitat segregation between H. tridens and M. japonicus burrows was caused primarily by an exclusive interaction between individuals of these two species.     

Reproductive biology of a filter-feeding amphipod, Leptocheirus pinguis , with extended parental care

Leptocheirus pinguis (Stimpson, 1853) is a widely distributed, abundant, endobenthic amphipod that engages in extended parental care, i.e. females host their juveniles in their burrows for extended time periods. I examined reproduction and population biology of L. pinguis at mean low water (MLW) in muddy sediment in Lowes Cove, Maine, USA. Cores around individual burrows were taken monthly in 1994 and 1995, and four seasonal samples were taken at different tidal heights. During the major reproductive periods in spring/early summer and in the fall, females produced several consecutive broods and hosted growing offspring in their burrows. Juveniles remained in their mothers' burrows until they reached a length of 5 mm (approximately one-third adult size) or more. At the study site, the majority of amphipods in individual burrows were adult females. Following the main reproductive periods, subadult individuals were found in their own burrows, but densities did not increase following the reproductive period in spring/early summer 1994, probably because large numbers of L. pinguis emigrated via the water column between June and December. L. pinguis is an annual species. Many members of the cohort born in spring/early summer start reproducing in the fall, and survive until the following spring when they produce several broods. Members of the cohort born in the fall start reproducing the following spring and also produce several consecutive broods. Both the spring/early summer and fall cohorts die off after the major reproductive period in the following spring/early summer. High standing stocks of microphytobenthos occur in soft-bottoms at MLW, and I conclude that L. pinguis can engage in extended parental care there because its food is abundant year-round. The limited expandability and low stability of burrows in soft-bottoms at MLW do not permit long persistence of parent–offspring groups in L.␣pinguis. Received: 27 March 1997 / Accepted: 30 July 1997     

Gonopore sexing technique allows determination of sex ratios and helper composition in eusocial shrimps

An evaluation of the social organization and sexual system of eusocial species of Synalpheus has been hindered because it has not been possible to determine the sexual composition of colony helpers (workers). The external sexual characters typically used to sex caridean shrimps are lacking in Synalpheus. We used SEM sexing technique to determine the sexual composition of helpers in colonies of Synalpheus regalis, S. rathbunae, S. chacei, S. rathbunae A (see Morrison et al. Mol Phylogen Evol 30:563–568, 2004), and S. filidigitus. Colonies consisted of both sexes and sex ratios of helpers generally conformed to 50:50 female to male. Females were characterized by gonopores with U-shaped slits on the coxae of the third pereopods (first walking legs) while males had oval gonopore openings on the coxae of the fifth pereopods (third or last walking legs). In S. chacei, S. filidigitus, and S. rathbunae A, a few helpers were found that had both male and female gonopores (intersexes). All three reproductive females (queens) of S. filidigitus examined were intersexes. Sexing of helpers allowed us to test some hypotheses about sexual differences in helper morphology that might indicate task specialization (division of labor). Male helpers were not different from female ones in body size (except in S. regalis: males somewhat larger) and in fighting chela size. The lack of sexual dimorphism in these characters suggests no male–female specialization in colony tasks such as defense. The presence of male and female helpers similar in size suggests that the sexual system of these eusocial species is gonochoristic, although protandry of some sort in S. filidigitus can not be ruled out. The intersexuality observed in a few individuals may be due to developmental anomalies, protandry, or even simultaneous hermaphroditism. Finally, the sexing technique allowed us to establish that new colonizers of unoccupied sponges in S. rathbunae are a single male and female of helper size.     

Reproductive biology of a deposit-feeding amphipod, Casco bigelowi, with extended parental care

The reproductive traits of a deposit-feeding amphipod that engages in extended parental care were examined. At the study site in Lowes Cove, Maine, USA, Casco bigelowi (Blake, 1929) occurred in highest densities in soft sediments just below mean low water (MLW). During most months, the sex ratio was ≃1. Many females hosted males in their burrows throughout the summer, but after fertilization of females in September, all adult males disappeared from the study area. In October almost 80% of the females were ovigerous, and in November about the same percentage was parental, i.e. caring for juveniles in their burrows. The females produced only one brood each in late fall which they accommodated in their burrows for 2 mo or longer. The average number of juveniles per female was ∼20 in November, and continuously decreased until January. Juveniles reached sizes >10 mm length in the maternal burrows. In early December the first juveniles were found in their own burrows, but major recruitment took place in late December and January. It is concluded that for C. bigelowi, the delay of recruitment into the winter months with low predation pressure and the large offspring size at this time are major advantages gained by extended parental care. C. bigelowi is host to the peritrich ciliate Cothurnia sp. on its gills, and during the summer months >70% of all amphipods had ciliates on their gills. Juveniles still living in their mother's burrows showed infestation rates similar to that of the parent; those of highly infested mothers were more heavily infested than those of “clean” mothers. Facilitated epibiont transmission during intimate and long-lasting (2␣mo) parent–offspring associations may be a consequence of extended parental care. Received: 25 November 1997 / Accepted: 14 April 1998     

Optimal investment in sons and daughters when parents do not know the sex of their offspring

Optimal parental investment usually differs depending on the sex of the offspring. However, parents in most organisms cannot discriminate the sex of their young until those young are energetically independent. In a species with physical male–male competition, males are often larger and usually develop sexual ornaments, so male offspring are often more costly to produce. However, Onthophagus dung beetles (Coleoptera; Scarabaeidae) are highly dimorphic in secondary sexual characters, but sexually monomorphic in body size, despite strong male–male competition for mates. We demonstrate that because parents provide all resources required by their offspring before adulthood, O. atripennis exhibits no sexual size dimorphism irrespective of sexual selection pressure favoring sexual dimorphism. By constructing a graphic model with three fitness curves (for sons, daughters, and expected fitness return for parents), we demonstrate that natural selection favors parents that provide both sons and daughters with the optimal amount of investment for sons, which is far greater than that for daughters. This is because the cost of producing small sons, that are unable to compete for mates, is far greater than the cost of producing daughters that are larger than necessary. This theoretical prediction can explain sexual dimorphism without sexual size dimorphism, widely observed in species with crucial parental care such as dung beetles and leaf-rolling beetles, and may provide an insight into the enigmatic relationship between sexual size dimorphism and sexual dimorphism.     

Population dynamics, sex ratio and size at sex change in a protandric simultaneous hermaphrodite, the spiny shrimp Exhippolysmata oplophoroides

One of the main goals of sex allocation theory is understanding sex ratio evolution. However, theoretical studies predicting sex ratios in species with unusual sexual systems, such as protandric simultaneous (PS) hermaphroditism, are rare. In PS hermaphrodites, juveniles first develop into functional males that mature into simultaneous hermaphrodites later in life. Here, we report on the sex ratio (males/males + hermaphrodites) in the PS hermaphroditic shrimp Exhippolysmata oplophoroides. A 2-year study demonstrated that hermaphrodites dominated the population in two different bays. This skewed sex ratio may be explained by limited encounter rates among conspecifics. In agreement with this idea, the density of shrimps was extremely low (≤1 shrimp km⁻²) at the two study sites. Size at sex phase change and sex ratios remained relatively stable through time at the two bays. The stability of these parameters might be explained by the rather steady population structure of this species during the study period. A review of sex ratios in PS hermaphroditic shrimps (Lysmata and Exhippolysmata) revealed considerable variation; some species have male- and others hermaphrodite-skewed sex ratios. The conditions explaining inter- and intra-specific sex ratio variation in protandric simultaneous hermaphroditic species remain to be addressed.     

Antifouling adaptations of caridean shrimps: Cleaning of the antennal flagellum and general body grooming

Structural specializations of chelipeds used by caridean shrimp in two kinds of grooming activity are described. In general body grooming, the chelipeds, and in some species, the last walking legs, nip, pick or brush material from the exoskeleton. When the cleaning chelipeds are the second pair, the carpal segment is multisegmented, increasing distal flexibility which aids in grooming. Tufts of compound setae and setal chela locks are characteristic of cleaning chelae. In representatives from 13 of 15 caridean families surveyed, brushes of serrate setae surround the carpal-propodal joint of the first cheliped. This setal structure is used in the specific task of cleaning the chemotactile antennal flagellum. Cleaning brushes on the last walking legs of some species are involved in general body cleaning. Experiments on Heptacarpus pictus showed that when the cleaning chelipeds were ablated, body parts became fouled with epizoites and particulate debris in experimental shrimps, while control shrimps showed little fouling. Suggestions on the adaptive role of general body cleaning in these natant animals are discussed. A survey of cleaning characters in representatives from 15 caridean families suggest that such characters are rather constant within a family. A possible correlation between the taxonomic success of a family and the degree of development of general body cleaning is suggested.     

Burrow morphology of the ghost shrimp<Emphasis Type="Italic"> Nihonotrypaea petalura</Emphasis> (Decapoda: Thalassinidea: Callianassidae) from western Kyushu,Japan

The callianassid shrimp Nihonotrypaea petalura (Stimpson, 1860) is a common member on boulder beaches in Japanese waters. Its burrow morphology was investigated, based on 28 resin casts collected from a steeply sloping beach with dense boulders and 30 from a more gently sloping beach with less dense boulders in Ariake Sound, southern Japan. The structure and dimensions of the burrows from the two beaches were basically the same, and thus a combined mean value could be given to most of their constituent elements. In its entire dimensions, the burrow is greater in lateral extent than depth, with a mean maximum horizontal extension of 145 mm and a mean maximum depth of 119 mm for the shrimp with a mean total length of 31.8 mm. The burrow winds along boulders or cobbles and consists of, from top to bottom, a single surface opening with an ejected mound, a top shaft leading to the uppermost chamber at a mean depth of 48–56 mm, a passage with a regular cross section that is wider than that of the top shaft, and bulbous chambers (mean no.=4.7) with an irregular cross section associated with branches (mean no. per burrow=1.2). Bulbous chambers are much larger than the uppermost chamber and are usually connected by passages, with some directly attached to each other. The combined architecture of these features is unique and relatively simple among the burrows of all callianassid species, the majority of which inhabit bare soft sediments. The structure and function of the N. petalura burrow are discussed in relation to lifestyle.Communicated by T. Ikeda, Hakodate     

Habitat use by sponge-dwelling brittlestars

Cryptic organisms often associate with sessile invertebrates for refuge in space-limited environments. To examine interspecific habitat associations on coral reefs, tube- and vase-shaped sponges were surveyed for associated brittlestars at six sites on the coral reefs off Key Largo, Florida. Of 179 sponges encountered, Callyspongia vaginalis was the most abundant (43.0%), followed by Niphates digitalis (39.7%), and Callyspongia plicifera (4.5%). Three of eight sponge species surveyed did not differ from C. vaginalis in two physical refuge characteristics: oscular diameter and inner tube surface area. Brittlestars (416 total), all of the genus Ophiothrix, were only found in C. vaginalis, N. digitalis, and C. plicifera. The most abundant brittlestar, O. lineata (326), occurred on C. vaginalis (99.0%) and N. digitalis (1.0%), while O. suensonii (67) occurred on C. vaginalis (79.1%), N. digitalis (19.4%), and C. plicifera (1.5%). There was no pattern of co-occurrence of O. lineata and O. suensonii on C. vaginalis. The abundance of O. lineata increased with surface area of C. vaginalis. Differential habitat use was observed in O. lineata, with small individuals (<5 mm disk diameter) located inside and on the surface of sponge tubes and large individuals (5 mm) solely inside tubes. The number of large O. lineata in C. vaginalis never exceeded the number of tubes per sponge, and tagged O. lineata remained in the same sponge for at least 3 weeks. In density manipulations, no pattern of intraspecific competition among large O. lineata was observed; however, there was evidence for interaction between size-classes. Brittlestars selected live sponge habitat over a non-living refuge, suggesting a mechanism for sponge habitat recognition. Sponge-dwelling brittle stars prefer some tube- and vase-shaped sponge species despite similar oscular diameters and surface areas. Surprisingly, these preferred sponge species are known from previous studies to be chemically undefended against generalist fish predators; therefore, brittlestars that inhabit these sponges do not gain an associational chemical defense. Sponge habitat use by O. lineata may be governed by intraspecific interactions to maintain habitat and access to food. While past studies have suggested that O. lineata is an obligate sponge commensal, the present study suggests that O. lineata has a species-specific association with the tube-sponge C. vaginalis.Communicated by O. Kinne, Oldendorf/Luhe     

Role of the bivalveCorbicula japonica in the nitrogen cycle in a mesohaline lagoon

The effect of the bivalveCorbicula japonica on the nitrogen cycle in Lake Shinji, a mesohaline brackish lagoon in Japan, was examined quantitatively based on field surveys and laboratory experiments carried out in the summer of 1982 and 1983. The biomass of the flesh ofC. japonica comprised 97% of the total biomass of the macrozoobenthos in summer. Total biomass ofC. japonica in the lake was estimated 30 986 t fresh wt. The concentration of suspended solids immediately above the lake bottom, whereC. japonica filters the water, was 1.5 to 4 times higher than that in the surface water. In laboratory experiments, the filtration rate was 5.0 litres g dry flesh wt^-1 h^-1, the excretion rates of ammonia and of feces and pseudofeces ofC. japonica were 200×10^-6 g N g dry flesh wt^-1 h^-1 and 33.4 mg dry wt g dry flesh wt^-1 h^-1, respectively at 27°C, the average summer water temperature in the lake. From our study, we estimated thatC. japonica filters almost the same amounts of particulate organic nitrogen produced in the lake by phytoplankton and that supplied from rivers, and that it excretes 30% of filtered nitrogen as feces or pseudofeces and 18% as ammonia.     

Reproductive strategy of<Emphasis Type="Italic"> Calanoides carinatus</Emphasis> and<Emphasis Type="Italic"> Calanus helgolandicus</Emphasis> during a summer upwelling event off NW Spain

Reproductive activity and production of the calanoid copepods Calanus helgolandicus and Calanoides carinatus were measured during a summer upwelling event off the coast of NW Spain. The upwelling pattern affected the distribution and fecundity of both species in the study area. The demographic composition of both populations and the stage of gonad maturation (e.g. the high abundance of fertilised females with mature ova) indicated active reproduction. C. carinatus, a highly fecund species associated with the African upwelling zones and considered as an upwelling specialist, showed low production rates (overall means of 15 eggs female^–1 day^–1 and 3% body C day^–1), despite the fact that the food conditions (high phytoplankton biomass dominated by diatoms) seemed to be optimal for this species. By contrast, C. helgolandicus, a temperate species that shows a strong link between spring phytoplankton blooms and reproduction time, seems to be flexible enough to take full advantage of shorter-term, enhanced feeding conditions associated with the pulsed nature of the summer coastal upwelling. Both the egg and carbon-specific production rates attained by this species (overall means of 26 eggs female^–1 day^–1 and 12% body C day^–1) were similar to values reported for a spring bloom situation. This high production would imply a long spring–summer recruitment event of C. helgolandicus in these waters. For both species the stage of gonad maturation was significantly correlated with their egg production rates and likely influenced by the food conditions; a species-specific nutritional requirement for final oogenesis is suggested. The carbon condition factor (carbon weight/prosome volume) of C. carinatus females was higher than that of C. helgolandicus, suggesting differential use of the carbon ingested; C. helgolandicus seems to use all ingested carbon to produce eggs at a high rates, whereas C. carinatus seems to store part of the ingested carbon as lipid reserves to ensure female survival and to support production during subsequent unfavourable food conditions.Communicated by S.A. Poulet, Roscoff     

Some aspects of the ecology of the echiuran worm Bonellia viridis and associated infauna

In inshore waters around Malta, Bonellia viridis inhabit burrows with multiple exits in calcareous rocks, and are most abundant in areas bordering Posidonia oceanica meadows. The associated epiflora and fauna are typical of the Mediterranean hard-substrate infralittoral zone. The larger infaunal species associated with B. viridis include a poriferan, Cliona sp.; an unidentified nemertean worm; various polychaetes; the sipunculans Phascolosoma granulatum and Aspidosiphon muelleri; two species of molluscs-Lithophaga lithophaga and Lepidopleurus cajetanus; 8 species of decapod crustaceans; and the teleost Gobius geniporus. Four types of burrows are found in rocks containing B. viridis. The larger burrows (referred to herein as UBA burrows) contain 3 main species-the decapods Upogebia deltaura and Alpheus dentipes and B. viridis itself. Experimental evidence suggests that U. deltaura excavates the UBA burrows mechanically, although B. viridis may secondarily modify them by secretion of an acidic mucus and gentle mechanical action. There is a definite community of organisms living as commensals in the burrow of U. deltaura. The assemblage of organisms bears a striking parallelism to the assemblages inhabiting the burrows of species of Upogebia, Callianassa and the echiuran Urechis caupo of the Pacific coast of N. America. A food web for the UBA burrow community is suggested.