Richness,abundance and shell use of subarctic and arctic hermit crabs

Hermit crabs are an obvious, common and abundant feature of global shallow water environments but almost nothing is known of their ecology at their extreme latitudinal ranges, the Arctic and southern South America. In this study, we report the first investigation on hermit crabs and their use of a key resource, gastropod shells, in a subpolar and a polar environment—amongst the most rapidly changing places on earth. Hypothesised low levels of richness were confirmed by surveys—only Pagurus pubescens was found in western Spitsbergen and only three pagurids were found in northern Norway. At the northern-most of their extent, hermit crabs were fairly common but Arctic abundances (1–5 m⁻²) were an order of magnitude lower than in many warm temperate or tropical localities. Along the open coast of Spitsbergen, the occurrence of P. pubescens was infrequent and very patchy, but it was more abundant in the fjords. In Isfjorden, the largest studied fjord, the population of P. pubescens was mainly represented by small individuals. Spitsbergen P. pubescens used few shell types and >87% just occurred in Margarites or Buccinum shells. The proportion of the gastropod shells, used by P. pubescens, which were damaged, was high and increased with shell size. These hermit crabs are at the edge of the range for both their species and Anomura. The extremes of their location are reflected by: They probably represent the least rich assemblage, with the lowest and most patchy typical abundances reported to date and are amongst the smallest individuals using the least diverse and most damaged gastropod shells.     

The past ownership of a resource affects the agonistic behavior of hermit crabs

Resource value is often considered the most important nonstrategic variable in the fighting behavior of invertebrates. In our study, we tested whether the experience of shells of a given quality, occupied in the recent past, might affect the agonistic behavior of the hermit crab Pagurus longicarpus. We analyzed the fights battled by 84 test crabs against size-matched unknown rivals before (premanipulation phase) and after (postmanipulation phase) having modified the quality of the domicile shell of the contestants. Specifically, we compared the behavior of crabs that had been subject to either a worsening or an improvement in the quality of their shell with crabs that, although being similarly subject to a shell manipulation, occupied a shell of the same quality as in the premanipulation phase. We found that the crabs subject to a worsening in the quality of their shell were more aggressive than those subject to its improvement and that the former were even more aggressive than those that occupied a bad quality shell also in the premanipulation phase. Crabs seemed not to gather information about the opponent’s shell during fights or not to use this information, most often behaving in accordance with the quality of the domicile shell. These results are clear in showing the role played by the experience of a previously occupied shell, also confirming that the agonistic behavior of P. longicarpus is mainly based on decision rules of the type “own resource value dependency.”     

Ecology of subtropical hermit crabs in SW Madagascar: short-range migrations

Many mobile animals migrate because of the different benefits provided by different localities in time and space. For hermit crabs, such benefits include resource (shell, water, food) acquisition and gamete release. One of the more successful crustacean land-invaders, Coenobita hermit crabs, undertake complex short-range migrations in SW Madagascar. Number of active hermit crabs was inversely related to wind strength and positively related to tidal range, emphasising that movement would conserve water. A circadian component was also recorded in the locomotory activity of Coenobita pseudorugosus and C. rugosus. Path linearity varied with many of the same parameters, but also with beach slope. Movement was primarily perpendicular to shore in small individuals, but the parallel proportion increased with hermit crab size and tidal range, probably driven mostly by shell and food searching. Despite the costs of movement and shell carriage in the terrestrial environment, C. pseudorugosus and C. rugosus were as fast as their marine counterparts. Their speeds varied principally with individual size and were approximately 20% faster without shells and about 20% slower when climbing up a 20° slope, compared to horizontal or downhill travel. Hermit crabs, which are highly numerous and speciose in SW Madagascar, do not seem to partition niches by differential movement patterns. Aside from provision of shells in middens and capturing large adults for bait or pets, human activity may have a profound effect on hermit crab movement: observations at rare uninhabited marine reserves like Nosy Ve show that considerable diurnal activity may take place despite the apparent hostility of the environment to an essentially marine animal.     

Mating success of male hermit crabs in shell generalist and shell specialist species

Summary The reproductive behavior of two species of diogenid hermit crabs was studied in Hawaii. In the shell generalist, Clibanarius zebra, male reproductive success varied little with size, although the largest males were less successful in obtaining copulations than were medium-large males. Male and female size were positively correlated, in successful pairs, thus larger males had the potential to fertilize more eggs when they were successful in obtaining a copulation. Female fecundity in C. zebra was not affected by species of gastropod shell inhabited once female size was taken into account. Male copulatory success was very strongly influenced by the species of gastropod shell inhabited. Males in Trochus or Nerita shells had greatly reduced reproductive success compared to males in Turbo or Nassarius shells. This result was due both to (1) males in Trochus especially dropping and otherwise poorly handling females during precopulatory behavior and (2) females not responding to precopulatory behavior patterns executed by males in Trochus and Nerita. Transferring males from good to bad shapes of shells and vice versa showed that male success was a function of shell type inhabited and not some correlated feature of the crabs. In the shell specialist, Calcinus seurati, which is found primarily in Nerita shells as an adult, males in Nerita shells were quite successful in obtaining copulations.     

The breeding biology of two sympatric species of tropical intertidal hermit crabs,Clibanarius chapini and C. senegalensis

A one-year study of breeding in the two intertidal hermit crabs Clibanarius chapini Schmitt and C. senegalensis Chevreux and Bouvier showed that both species maintain a high reproductive activity throughout the year. Brief low reproductive activities, found in both species, occurred in different months: C. chapini in November, C. senegalensis in May. While these drops cannot be attributed to any recognisable environmental factor, it is suggested that such pattern may result from competition between the two species. Sex-ratio studies indicated slightly more females than males in both species. The maximal size attained by males of both species is considerably greater than that of the females; in C. senegalensis this is very marked. It is suggested that it may be selectively advantageous for a female to attain a smaller maximal size since her brood must also be accomodated within the shell. As expected for continuously breeding species, recruitment of young into the population is also continuous.     

Temporal and spatial settlement patterns of sympatric hermit crabs and the influence of shell resource availability

Larvae of marine organisms often need specific resources or environments at settlement, and their success at settlement might be strongly influenced by the abundance and distribution of such specific resources. The larvae of hermit crabs need small shells to settle, so it is thought that the distribution and abundance of small shells influence the settlement pattern of hermit crabs. To investigate the influence of small shell distribution on the settlement of pagurid hermit crab larvae, we conducted a field experiment at an intertidal rocky shore in Hakodate Bay, Japan. From the line-transect sampling in the field, we found that Pagurus middendorffii settled extensively in the offshore side of the intertidal zone while P. nigrofascia settled in the uppermost area of the intertidal zone. Small shells were most abundant in a narrow shallow trough, slightly offshore from the uppermost area of the intertidal zone. For both species, settler abundance was high where adults were abundant, but settler abundance did not appear to be related to shells abundance. An experiment to clarify settlement patterns showed that larval recruits tended to be similar to those in the line-transect sampling of settlers. Thus shells may not be a primary factor affecting settlement patterns at relatively large scale within the intertidal flat. However, when we analyzed the relationship of settlers and shells separately within each transect, the distribution of settlers was well explained by shell resource availability. Therefore on a smaller scale, shell availability may influence the number of settlers. Settlement periods of P. middendorffii and P. nigrofascia fully overlapped, so their larvae probably were affected by similar transport factors, such as current and tidal movement. Nevertheless they showed different spatial patterns of settlement.Communicated by T. Ikeda, Hakodate     

Ecology of subtropical hermit crabs in S.W. Madagascar: cluster structure and function

The Madagascar coast (both) has a higher density and diversity of hermit crabs than is known from any other locality in the western Indian Ocean. Of the 20 species occurring at Anakao (S.W. Madagascar), 11 aggregated into clusters, including all but one of the species above the subtidal zone. The mean number of hermit crabs and species in clusters varied with several spatial parameters and time. Over 80% of the community clustered in certain habitats in particular tidal zones, whilst as low as 3% clustered in others. The highest intensity of clustering with shore zone was coincident with peak numbers of hermit crabs. The initiation and duration of clusters of hermit crabs above the eulittoral was driven by circadian rhythms, whilst those in the eulittoral were governed by tidal state. Clusters above and below the eulittoral were longer in duration, and those in the subtidal were more temporally variable than those above it. Certain pairs of species showed positive correlations of occurrence and (more rarely) of abundance, and all the correlations (of occurrence) of one, Clibanarius eurysternus, were negative. Positive correlations of occurrence were related to the degree of shell-use commonality between species pairs. Eulittoral species clustered with other individuals of approximately similar size and exchanged shells upon cluster disintegration. There was evidence of a dehydration-reduction function to shell clustering in addition to shell-exchange facilitation. The highly variable species-specific strategies of clustering may be important in alleviating both intra- and interspecific competition in assemblages of similar and highly abundant species.     

Fighting behavior in hermit crabs: the combined effect of resource-holding potential and resource value in Pagurus longicarpus

A study was conducted to determine whether asymmetries in both resource-holding potential (RHP) and resource value (RV) influence dominance and fighting behavior in the hermit crab Pagurus longicarpus. A total of 120 groups of three crabs were observed for 10 min in four experiments that tested all diverse combinations of equal/different RHP (i.e. ‘body size’) and equal/different RV (i.e. ‘shell size’ and ‘shell quality’). In a fifth experiment, dominant and subordinate individuals of the same size category (26 groups) were forced to enter shells of opposite quality than those previously occupied, and then the behavior of the reconstituted original groups was observed for additional 10 min. As expected, crabs in lower quality shells were more willing to initiate and to escalate fights. However, their attacks were directed to any crab of the group, independently of the defender's shell quality, and the fight duration did not vary with the different value of the resources at stake. This may indicate that P. longicarpus is unable to assess the quality of the shells available in its social environment but bases its tactical decisions during fights solely on its own resource. This suggestion was confirmed by the change in the fighting behavior of crabs whose shell quality was experimentally altered. This manipulation induced an overall increase in the intensity of aggression, drastic modification of crab behavior, and inversion of the hierarchy even though these crabs have had previous experiences of wins/losses and were familiar to the other members of the group. In this species, large crab size and/or the occupancy of adequate (and oversized) shells appeared to be the most likely determinant of contest resolution. Individuals seemed to retain a memory of the previously held resource and behaved accordingly.     

Effect of salinity on hypoxia tolerance of resting green crabs, Carcinus maenas, after feeding

Mechanisms that can influence the tolerance of hypoxia in brackish waters were studied in resting and fed crabs, Carcinus maenas, at 15?°C. Mortality, blood oxygenation, acid-base status and lactate concentration were analysed in fed crabs held in full-strength normoxic seawater (32.5‰?S) and then transferred for 24?h to a partial pressure of oxygen (P_o2) of 3?kPa (1.4?mg?l^?1) and various salinities (17, 12.5, 10, 8‰?S). At salinity levels >10‰, fed crabs tolerated P_o2 values as low as 3?kPa in the ambient water and 0.5?kPa in their arterial blood for 24?h without switching to anaerobic metabolism. Only below 10‰?S did their blood-lactate content rise, leading to their death despite the fact that their blood O₂-content was twice the control value measured in full-strength normoxic seawater and their blood P_o2 did not decrease below values recorded at higher salinity levels. Addition of CO₂ to 8‰?S water (CO₂ partial pressure increasing from 0.1 to 0.3?kPa) decreased blood-lactate production and mortality, suggesting that at 10‰?S impairment of the O₂ supply is limited by an excessive blood O₂-affinity. The results are discussed in terms of the distribution (?10‰?S) of C. maenas along salinity gradients in estuaries and bays.     

Prey, feeding rates, and asexual reproduction rates of the introduced oligohaline hydrozoan Moerisia lyonsi

Moerisia lyonsi Boulenger (Hydrozoa) medusae and benthic polyps were found at 0 to 5‰ salinity in the Choptank River subestuary of Chesapeake Bay, USA. This species was introduced to the bay at least 30 years before 1996. Medusae and polyps of M. lyonsi are very small and inconspicuous, and may occur widely, but unnoticed, in oligohaline waters of the Chesapeake Bay system and in other estuaries. Medusae consumed copepod nauplii and adults, but not barnacle nauplii, polychaete and ctenophore larvae or tintinnids, in laboratory experiments. Predation rates on copepods by medusae increased with increasing medusa diameter and prey densities. Feeding rates on copepod nauplii were higher than on adults and showed no saturation over the range of prey densities tested (1 to 64 prey l⁻¹). By contrast, predation on copepod adults was maximum (1 copepod medusa⁻¹ h⁻¹) at 32 and 64 copepods l⁻¹. Unexpectedly, M. lyonsi colonized mesocosms at the Horn Point Laboratory during the spring and summer in 4 years (1994 to 1997), and reached extremely high densities (up to 13.6 medusae l⁻¹). Densities of copepod adults and nauplii were low when medusa densities were high, and estimated predation effects suggested that M. lyonsi predation limited copepod populations in the mesocosms. Polyps of M. lyonsi asexually produced both polyp buds and medusae. Rates of asexual reproduction increased with increasing prey availability, from an average total during a 38 d experiment of 9.5 buds polyp⁻¹ when each polyp was fed 1 copepod d⁻¹, to an average total of 146.7 buds polyp⁻¹ when fed 8 copepods d⁻¹. The maximum daily production measured was 8 polyp buds and 22 medusae polyp⁻¹. The colonizing potential of this hydrozoan is great, given the high rates of asexual reproduction, fairly wide salinity tolerance, and existence of a cyst stage. Received: 29 October 1998 / Accepted: 3 March 1999     

Quantitative differences in mitochondrial ultrastructure of a thiobiotic and an oxybiotic turbellarian

Specimens of the thiobiotic acoel Solenofilomorpha funilis Crezée, 1975 and the oxybiotic acoel Otocelis sp. were collected from Mount Desert Island, Maine, USA in spring and summer of 1982, and the ultrastructure and distribution of their mitochondria were compared. The volume density and the surface density of the mitochondria are significantly higher in the epidermis and parenchyma of S. funilis than in the corresponding tissues of Otocelis sp. Surface densities of mitochondrial cristae are higher in all tissues studied (epidermis, muscle, and parenchyma) of S. funilis than in Otocelis sp. The differences in mitochondrial ultrastructure of these two species suggest that mitochondria in anaerobic tissues differ in function from their aerobic counterparts.     

Role of feeding time and food amount in the reproduction of spotted Munia

It was observed that food restriction significantly increase gonadal development for about two months indicating that hypothalamo hypophyseal gonadal axis becomes more sensitive to feeding stress during the early breeding period than the other reproductive phases. Normal testicular development was also not seen in the spotted munia in which food was restricted for two times each of 3 hour duration, despite the fact that daily food consumption was not significantly less (only by 6.60%) compared to the control group. This suggests that daily restriction in feeding for a longer period (total 6 h or more) may also alter the reproductive function in wild birds.     

Evolutionary significance of prenuptial molting in female <Emphasis Type="Italic">Pagurus</Emphasis> hermit crabs

Molting and breeding entail major energetic costs for female crustaceans. However, females of some hermit crabs perform a molt immediately prior to copulation (prenuptial molt). The evolutionary significance of the prenuptial molt was examined in Pagurus hermit crabs, and two hypotheses were tested: (1) prenuptial molt might enhance the success of the present clutch by cleaning the pleopods of females and thereby preventing eggs from being dislodged from the pleopods, and (2) prenuptial molt might function for growth and increase future fecundity at the cost of energetic expenditure on the present brood. Although these hypotheses are not mutually exclusive, our results rejected the former hypothesis and supported the latter hypothesis. All four Pagurus species examined showed significant negative relationships between prenuptial molting and continuity of breeding; i.e., they showed high molting frequency after they had a long rest period from breeding. Females of P. minutus increased their size through the prenuptial molt, and showed a decreased clutch size due to the molt. The number of dislodged eggs increased if females molted in P. minutus. These results suggest that hermit crabs undergoing a prenuptial molt might not gain any clear immediate advantage of enhanced survival of eggs in the present clutch, and that the prenuptial molt would mainly contribute to growth.     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000     

Unravelling host and symbiont phylogenies of halichondrid sponges (Demospongiae, Porifera) using a mitochondrial marker

We present the first comparative phylogenetic analysis of a selected set of marine sponges and their bacterial associates. The Halichondrida form an important order in demosponge systematics and are of a particular interest due to the production of secondary metabolites. We sequenced a fragment of the cytochrome oxidase subunit 1 (CO1) gene of the sponges and their bacterial associates, compared the reconstructed phylogenies and found evidence for radiation in coevolution. The tree of six host-species associations showed four supported cospeciation events between the sponges and the bacteria. In addition, we present the first gene tree of sponges based on a mitochondrial marker. The tree shows major congruences with previous morphological studies and suggests the applicability of a mitochondrial marker in sponge molecular systematics.     

Effects of food ration and feeding regime on growth and reproduction of the sea urchin Strongylocentrotus droebachiensis

To determine the effects food ration and feeding regime on growth and reproduction of Strongylocentrotus droebachiensis (Müller), sea urchins in laboratory aquaria were fed kelp (Laminaria longicruris) supplied at either a high (H, ad libidum daily) or a low (L, ad libidum 1 d wk⁻¹) ration in two successive 12-wk intervals during the reproductive period. After 24 wk, urchins fed the high ration continuously (HH) or for the last 12 wk only (LH) had a significantly greater mean gonad index [(gonad weight/total body weight) × 100] and body weight than urchins fed the low ration continuously (LL) or for the last 12 wk only (HL). Urchins in the HL treatment had a significantly greater gonad index than those in the LL treatment; there was no significant difference in gonad index between the LH and HH treatments. Females had a greater gonad index than males in the low ration (LL and HL) treatments at the end of the experiment; there was no significant difference between sexes in the high ration (LH, HH) treatments. Gametogenesis proceeded to maturation in all treatments and some individuals spawned at the end of the experiment. Females in the high ration (HH and LH) treatments had a greater proportion of nutritive phagocytes in their ovaries than females in the low ration treatments, but there was no effect of feeding treatment on oocyte or ovum size. Feeding treatment had no effect on the relative abundance of nutritive phagocytes in the testes, although the proportion of spermatocytes was higher (and that of spermatozoa lower) in the high ration than in the low ration treatments. Urchins in the high ration treatments had a lower mean jaw height index [(jaw height/test diameter) × 100] and greater mean test diameter than those in the low ration treatments at the end of the experiment, although these differences were not statistically significant. Feeding rate on kelp at the end of the experiment was significantly greater for urchins in the low ration than in the high ration treatments. Our experimental results show that even relatively low rations of kelp support somatic and gonadal growth in S. droebachiensis. Increasing the supply of kelp, particularly during the period of active gametogenesis, results in maximal rates of growth and reproduction. These results suggest that populations of S. droebachiensis in barrens may derive a substantial proportion of their nutrition from drift kelp, which may contribute to their persistence in these habitats. They also explain the large body size, high reproductive effort and fecundity of urchins grazing on kelp beds. These findings have important implications for understanding the dynamics of natural populations of S. droebachiensis and for development of effective aquacultural practices. Received: 17 February 1997 / Accepted: 5 March 1997     

Obligate commensalism of <Emphasis Type="Italic">Curvemysella paula</Emphasis> (Bivalvia: Galeommatidae) with hermit crabs

Curvemysella paula is a markedly crescent-shaped bivalve that lives inside snail shells occupied by hermit crabs. Here, we describe the unique symbiotic life, growth pattern, and reproductive biology of this bivalve, based on specimens collected from the shallow, muddy bottom of the Seto Inland Sea, Japan. C. paula was found attached to columellae in the siphonal canal, mainly of nassariid snail shells occupied by two types of hermit crabs: Diogenes edwardsii (Diogenidae) and Spiropagurus spiriger (Paguridae). The crescent-shaped shell of C. paula is an adaptation to symbiotic life in the narrow interspace between the snail shell and the hermit-crab abdomen. C. paula is a protandric hermaphrodite. In our samples, each host snail shell harbored one (or rarely a few) large female and several males. All the female bivalves settled on the host shells with their anterior end facing outward and benefited from currents created by the hermit crab when feeding. In the muddy bottom, snail shells are a limited resource for both the hermit crabs and symbiotic bivalves. The bivalves benefit from the mobility of the hermit crabs, which prevent the shells from becoming buried in the mud. C. paula represents the only example of obligate commensalism with hermit crabs found in Bivalvia.     

The gill symbiont of the hydrothermal vent mussel Bathymodiolus thermophilus is a psychrophilic,chemoautotrophic, sulfur bacterium

Certain hydrothermal vent invertebrates, e.g. Riftia pachyptila and Calyptogena magnifica, are clearly established as harboring dense populations of chemoautotrophic sulfur bacteria in specialized tissues. By contrast, the physiological characteristics of the abundant intracellular gill symbiont of the vent mussel Bathymodiolus thermophilus have been questioned. The low activities of enzymes diagnostic for CO₂ fixation (Calvin cycle) and for sulfur-driven energy generation, as measured by other investigators, have been attributed to bacterial contamination of the gill surface. Based on research at the Galápagos Rift hydrothermal vents in 1988 and subsequent laboratory experiments, the current study confirms that the B. thermophilus symbiont is a psychrophile for which thiosulfate and sulfide stimulate CO₂ fixation. It strongly indicates that the symbiont is a chemoautotroph by establishing the following: (1) Sulfide and thiosulfate can stimulate CO₂ fixation by partially purified symbionts by up to 43-fold and 120-fold, respectively; (2) the ribulose-1,5-bisphosphate carboxylase/oxygenase activity of the symbiont is sufficient to account for its sulfide- or thiosulfate-stimulated CO₂ incorporation; (3) the symbiont's molar growth yield on thiosulfate, as judged by CO₂ incorporation, is indistinguishable from that of free-living chemoautotrophs. Due to the high protein-degrading activity of B. thermophilus gill lysate, it is also suggested that host lysis of symbionts plays a more important role in the nutrition of the vent mussel than in R. pachyptila or C. magnifica, for which no comparable protein-degrading activity was found.     

Shell preference in a hermit crab: comparison between a matrix of paired comparisons and a multiple-alternative experiment

The preference of the hermit crab, Calcinus californiensis, among six species of shells, was tested by two different experiments. The first experiment used pair-wise trials, analyzing the preference by Chi-square tests using two different constructions of the null hypothesis. One hypothesis was based on a no-preference among shell species, the second on comparing the number of crabs changing for a particular shell species when two options were given versus the changing when no options were offered. The second experiment was a multiple-alternative test based on a rank ordering of the shell preference. This method has both statistical and resource-saving advantages over the traditional pair-wise comparisons. The sequence of shell preference was similarly independent of the procedure used. The preferred shell species are heavy and might be associated with hydrodynamic advantages and with the protection against predation. The shell preference matches with the pattern of shell occupancy indicating that the shell use in nature is determined by the crab’s preference. The information generated may be used for further research on shell preference as a methodological alternative.