An analysis of competitive interactions between 3 hermit crab species

Summary Competition for empty gastropod shells in a group of three sympatric hermit crabs (Pagurus hirsutiusculus, Pagurus granosimanus, and Pagurus beringanus) was studied in the San Juan Archipelago, Washington State. Estimates of the competitive effects of each species on the others' shell supplies were derived using field data on shell utilization and the results of laboratory experiments to determine rates of acquisition and exchange of shells and preferences for different shell species. Each species experienced approximately an order of magnitude more intraspecific competition than interspecific competition for empty shells. This resulted from differences in preference for shell shapes, shell size use, and habitat use between P. hirsutiusculus and P. granosimanus, and largely from differences in habitat use between P. beringanus and the other two species. Experiments involving the release and recensusing of marked empty shells were used to estimate competitive effects more directly for the interaction between P. hirsutiusculus and P. granosimanus. Results were consistent with the estimates derived from data on resource partitioning. Possible causes of the low levels of interspecific competition are discussed, and results are compared with studies of other organisms that estimated both inter- and intra-specific competition.     

A new genus and two new species of Peltogastridae (Crustacea: Cirripedia: Rhizocephala) parasitizing hermit crabs from Okinawa Island (Ryukyu Archipelago, Japan), and their DNA-barcodes

A new genus and two new species of Peltogastridae, Peltogaster postica sp. nov. and Ommatogaster nana gen. et sp. nov., are described from Okinawa Island, Ryukyu Islands, southwestern Japan. The two new rhizocephalans were found to be parasitic on the estuarine hermit crabs, Pagurus minutus Hess, 1865 and Diogenes leptocerus Forest, 1956, respectively. Peltogaster postica sp. nov. is allied to P. curvata Kossmann, 1874, P. paguri Rathke, 1842 , and P. reticulata Shiino, 1943 , but is distinguished by its relative length and internal and external structures of the mature externa. Ommatogaster gen. nov. is established for the present new species O. nana based on the morphologies of the visceral mass of the externa and the presence of a nauplius eye in the larvae. Partial COI sequences were obtained from the two new species and one known species, Dipterosaccus indicus Van Kampen and Boschma, 1925, to test the possible usefulness of the sequences as tags for species identification.     

The species composition and pattern of distribution of hermit crabs (Decapoda: Paguroidea) in Vostok Bay,Sea of Japan

The species composition, abundance, and vertical distribution of hermit crabs (Decapoda: Paguroidea) were investigated in Vostok Bay, Sea of Japan, from September 2014 to September 2015. A total of seven species from two families were found in the monitoring area of approximately 5000 m² located on the bottom within a depth range of 0–11 m: Pagurus minutus, P. proximus, P. brachiomastus, P. pectinatus, P. ochotensis, P. middendorffii (Paguridae), and Areopaguristes hirsutimanus (Diogenidae). Their population density varied from 22 ± 13 to 26 ± 2 ind./m²; the biomass varied from 55.76 ± 26.10 to 93.38 ± 27.00 g/m². The species composition remained invariable throughout the study period. The proportions of species abundance changed between seasons, as this parameter is largely determined by seasonal migrations of hermit crabs. Pagurus minutus predominated in number in winter and spring (up to 71%); P. proximus, in summer and autumn (up to 47%). In autumn and spring, P. ochotensis and P. pectinatus constituted a major part of biomass (18–25% and 19–20%, respectively). The proportion of a previously abundant species, P. middendorffii, was very small (no larger than 1.3%) during the study period. Another four species of hermit crabs occurred in other parts of Vostok Bay; thus, the fauna of the bay currently includes 11 species of this group.     

Larval,megalopal and early juvenile development in Pagurus granosimanus (Stimpson, 1859) (Decapoda: Anomura: Paguridae) reared in the laboratory

Summary

The larval, megalopal and early juvenile stages of Pagurus granosimanus are described, illustrated and compared with other North Pacific species of the genus. Morphologically, the zoeae of P. granosimanus appear most similar to the Japanese P. brachiomastus in the majority of characters, but share the endopodal setation of the third maxilliped with a second Japanese species, P. pectinatus. The megalopae of P. granosimanus are unlike those of other North Pacific species in having 5+5 marginal setae on the telson, rather than the customary 4+4, or less frequent 3+3. Comparison of juvenile characters is limited to pleopodal changes among the regional species for which data are available. P. granosimanus is unusual in undergoing complete pleopodal loss at the second crab stage with return of left pleopods in the fourth stage.     

The Parasitic Fauna of Common Species of Hermit Crabs of Vostok Bay (Sea of Japan)

The parasitic fauna of common species of hermit crabs was investigated in Vostok Bay (the northwestern Sea of Japan). Pagurus proximus and Areopaguristes hirsutimanus had no parasites. The average prevalence of rhizocephalan infestation in other five species of hermit crabs ranged from 5.67 to 20.83%. The prevalence of the isopod Parapseudione lata on hermit crabs did not exceed 1%, the prevalence of infestation with the isopod Athelges takanoshimensis was significantly higher, reaching 14.58% in Pagurus middendorffii. Parasitism by the isopod P. lata on the hermit crabs Pagurus minutus and P. ochotensis, as well as parasitism by the rhizocephalan Peltogaster reticulata on P. minutus were recorded for the first time. Several variants of simultaneous infestation with two parasitic species were found. The decline in the abundance of P. middendorffii accompanied by the high prevalence of parasites, primarily the colonial rhizocephalan Peltogasterella gracilis, suggests that parasite invasion may play a more significant role in regulating the abundance of hermit crabs than was previously assumed.     

Activities of two British species of Pagurus (Crustacea,Decapoda, Paguroidea)

Two series of experiments were conducted to investigate the activities of two British species of hermit crab, Pagurus bernhardus (L.) and Pagurus prideauxi Leach. The first series showed that P. bernhardus was more active during day than night whereas the opposite was true of P. prideauxi. In the second series, conducted under periods of 24 hours constant illumination, the activity pattern of P. bernhardus persisted to a lesser degree, but P. prideauxi showed no activity pattern. Differences in activity were observed among individuals of the same species, and also between individuals kept in the laboratory for different lengths of time.     

Hermit crab biocoenoses: a worldwide review of the diversity and natural history of hermit crab associates

The symbiotic associates of hermit crabs (excluding parasites and flora) are reviewed worldwide. The review includes species found on the shells occupied by hermit crabs (epibiotic species), species boring into these shells (endolithic species), species living within the lumen of the shell (either free-living or attached to the shell), species attached to the hermit crabs themselves, and hypersymbionts. In total over 550 invertebrates, from 16 phyla are found associated with over 180 species of hermit crabs. Among these associates, 114 appear to be obligate commensals of hermit crabs, 215 are facultative commensals, and 232 are incidental associates. The taxa exhibiting the highest number of associates are arthropods (126), polychaetes (105), and cnidarians (100). The communities of species associated with Dardanus arrosor, Paguristes eremita, Pagurus bernhardus, Pagurus cuanensis, and Pagurus longicarpus are the best studied and harbor the most diverse assemblages of species. While trends in biodiversity of hermit crab assemblages do not follow predicted patterns (e.g., hermit crabs within the Indo-West Pacific do not harbor more species than those from temperate regions), this is suggested to reflect a lack of sampling rather than a true representation of the number of associates. Hermit crabs date to at least the Cretaceous and provided a niche for a number of groups (e.g., hydractinians, bryozoans, polydorids), which were already associates of living gastropods. Apparently hermit crab shells initially supplied a substrate for settlement and then these symbiotic relationships were reinforced by enhanced feeding of symbionts through the activity of the hosts. Through their use and recycling of gastropods shells, hermit crabs are important allogenic ecosystem engineers in marine habitats from the intertidal to the deep sea. Hermit crabs benefit from some symbionts, particularly cnidarians and bryozoans, through extension of shell apertures (alleviating need to switch into new shells) and by providing protection from predators. However, hermit crabs are also negatively impacted (e.g., decreased reproductive success, increased predation) by some symbionts and a review of egg predators is provided. Thus, the symbiotic relationships between hermit crabs and many associates are difficult to characterize and often exhibit temporal changes depending on environmental and biological factors. Research on the biology of these symbionts and the costs/benefits of their associations with hermit crabs are analyzed. While some associates (e.g., Hydractinia spp.) have been studied in considerable detail, for most associations little is known in terms of the impacts of symbionts on hosts, and future experimental studies on the multitude of relationships are suggested.     

An analysis of inter‐ and intraspecific aggregations in two sympatric species of hermit crab (Decapoda,Anomura, Paguridae)

A series of experiments was conducted on the relationship between population density and spacing out in two sympatric species of crab, Pagurus bernhardus (L.) and Pagurus prideauxi Leach. A second series of experiments investigated the two species’ aggressiveness and activity by observing the behaviour of pairs of animals.

In the first series of experiments, animals became more aggregated (Clarke and Evans’ R) as the population density (p) was increased. The rate of aggregation (dR/dp) was greater when the species were mixed than when they were separate.

In the second series of experiments, activity, total encounters, and resolved encounters, were highest in paired P. bernhardus, lower in mixed pairs of the two species, and lowest in pairs of P. prideauxi. Larger individuals were more active than smaller ones in P. bernhardus and P. prideauxi pairs. In the mixed pairs, P. bernhardus were more active. Paired P. bernhardus were slightly more active than isolated P. bernhardus but this difference was not statistically significant. The activities of paired and isolated P. prideauxi were not significantly different. The distances between individuals in pairs were similar for all combinations.

The results are discussed in relation to the distribution of sympatric species.     

Comparative brain architecture of the European shore crab <Emphasis Type="Italic">Carcinus maenas</Emphasis> (Brachyura) and the common hermit crab <Emphasis Type="Italic">Pagurus bernhardus</Emphasis> (Anomura) with notes on other marine hermit crabs

The European shore crab Carcinus maenas and the common hermit crab Pagurus bernhardus are members of the sister taxa Brachyura and Anomura (together forming the taxon Meiura) respectively. Both species share similar coastal marine habitats and thus are confronted with similar environmental conditions. This study sets out to explore variations of general brain architecture of species that live in seemingly similar habitats but belong to different major malacostracan taxa and to understand possible differences of sensory systems and related brain compartments. We examined the brains of Carcinus maenas, Pagurus bernhardus, and three other hermit crab species with immunohistochemistry against tyrosinated tubulin, f-actin, synaptic proteins, RF-amides and allatostatin. Our comparison showed that their optic neuropils within the eyestalks display strong resemblance in gross morphology as well as in detailed organization, suggesting a rather similar potential of processing visual input. Besides the well-developed visual system, the olfactory neuropils are distinct components in the brain of both C. maenas and P. bernhardus as well as the other hermit crabs, suggesting that close integration of olfactory and visual information may be useful in turbid marine environments with low visibility, as is typical for many habitats such as, e.g., the Baltic and the North Sea. Comparing the shape of the olfactory glomeruli in the anomurans showed some variations, ranging from a wedge shape to an elongate morphology. Furthermore, the tritocerebrum and the organization of the second antennae associated with the tritocerebrum seem to differ markedly in C. maenas and P. bernhardus, indicating better mechanosensory abilities in the latter close to those of other Decapoda with long second antennae, such as Astacida, Homarida, or Achelata. This aspect may also represent an adaptation to the “hermit lifestyle” in which competition for shells is a major aspect of their life history. The shore crab C. maenas, on the other hand seems to rely much less on mechanosensory information mediated by the second antennae but in water, the visual and the olfactory senses seem to be the most important modalities.     

Hermit crabs Pagurus parvispina Komai, 1997 and Discorsopagurus maclaughlinae Komai, 1995 (Decapoda: Paguridae): New records for Russian waters of the Sea of Japan

The hermit crab Pagurus parvispina Komai, 1997 (Decapoda: Paguridae) was found in Vostok Bay (Peter the Great Bay, Sea of Japan) at a depth of 50?C65 m. Earlier, this species was recorded from the type locality in northern Japan at a depth of 150?C200 m. The finding of P. parvispina in Vostok Bay is its first record in Russian waters and substantially expands the distribution area of this species in the Sea of Japan. Discorsopagurus maclaughlinae Komai, 1995 inhabits Russian waters and was earlier identified as Orthopagurus schmitti (Stevens, 1925); its taxonomical position is discussed.     

Multigene molecular systematics confirm species status of morphologically convergent Pagurus hermit crabs

Introduction

In spite of contemporary morphological taxonomy appraisals, apparent high morphological similarity raises uncertainty about the species status of certain Pagurus hermit crabs. This is exemplified between two European species, Pagurus excavatus (Herbst, 1791) and Pagurus alatus (Fabricius 1775), whose species status is still difficult to resolve using morphological criteria alone.

Methodology/Principal Findings

To address such ambiguities, we used combinations of Maximum Likelihood (ML) and Bayesian Inference (BI) methods to delineate species boundaries of P. alatus and P. excavatus and formulate an intermediate Pagurus phylogenetic hypothesis, based upon single and concatenated mitochondrial (cytochrome oxidase I [COI]) and nuclear (16S and 28s ribosomal RNA) gene partitions. The molecular data supported the species status of P. excavatus and P. alatus and also clearly resolved two divergent clades within hermit crabs from the Northeast Atlantic Ocean and the Mediterranean Sea.

Conclusions/Significance

Despite the abundance and prominent ecological role of hermit crabs, Pagurus, in North East Atlantic Ocean and Mediterranean Sea ecosystems, many important aspects of their taxonomy, biology, systematics and evolution remain poorly explored. The topologies presented here should be regarded as hypotheses that can be incorporated into the robust and integrated understanding of the systematic relationships within and between species of the genus Pagurus inhabiting the Northeast Atlantic Ocean and the Mediterranean Sea.     

红眼寄居蟹在实验室和野外条件下对贝壳的利用

占据适宜的贝壳对于寄居蟹的发育、繁殖和存活至关重要。尽管很多研究探讨了蟹类对贝壳的选择,但寄居蟹选择多大的贝壳仍不清楚。在实验条件下,本文作者用巴西Anchieta岛上野生红眼寄居蟹( Pagurusbrevidactylus)最常利用的黑衣蟹守螺( Cerithium atratum)和节桑椹螺( Morula nodulosa)进行了贝壳选择实验,通过回归分析确定目标贝类及其大小。观察到寄居蟹对一种独特贝类的选择具有性别意义,从而验证了野外观察结果。雄性明显地倾向选择黑衣蟹守螺的贝壳,而产卵和非产卵的雌性个体对腹足类贝壳的选择差异不显著。尽管两性之间对适宜贝壳的选择存在差异,贝壳适宜度指数(SAI)表明,种群占据那些足够大的贝壳(SAI =1·20±0·23)。红眼寄居蟹对贝壳利用的这种模式可能是为了避免与体型相似同域物种的竞争,从而在后来的生长中减少频繁地更换贝壳。根据目前的数据可以得到以下结论:红眼寄居蟹对贝壳的选择不仅取决于贝壳的参数,而且还与寄居蟹的个体和性选择有关[动物学报51 (5) : 813 -820 , 2005]。     

Intraspecific relationships of the hermit crab Diogenes pugilator: predation and competition

The population of the hermit crab Diogenes pugilator is numerically quite substantial in the examined area, therefore it is analysed if the population was subject to any pressure by predators and by other species of hermit crabs living in that zone. It was found there is a potential pressure by predators which might influence it. Only Liocarcinus depurator is able to keep the population trend of D. pugilator under control. With regard to the relationships with the other species of hermit crab, D. pugilator dominates only over Pagurus anachoretus, while in the other cases, there was just competition for food.     

Gastropod shells: A potentially limiting resource for hermit crabs

The availability of gastropod shells to hermit crabs in the Newport River Estuary, Beaufort, N.C. has been assessed by determining the numbers of usuable shells occurring in characteristic subtidal habitats and by measuring shell size adequacy. The proportion of useable shells occupied by hermit crabs ranged from 58–99 % and many of the shells not used by hermit crabs were judged unavailable because they were occupied by sipunculids or only uncovered by the dredge. The shell adequacy index (shell size occupied/shell size preferred) was significantly below 1.0 for the largest species (Pagurus pollicaris Say) in the one location where sufficient numbers were collected and for the next largest species (P. longicarpus Say) in three of the four locations where it was collected. The shell size adequacy index for the smallest species (P. annulipes Stimpson) did not differ significantly from 1.0 in either of the two locations in which it was found. These observations suggest that the availability of gastropod shells plays a significant rôle in limiting the abundance of at least the larger hermit crabs.     

The influence of some symbionts on the shell-selection behaviour of the hermit crabs, Pagurus pollicarus and Pagurus longicarpus

The influence of some symbionts on the shell-selection by the hermit crabs Pagurus pollicarus and P. longicarpus was examined by placing individual hermit crabs with two similar shells in a choice situation and recording the shell occupied after 12 hr. One shell contained a symbiont species and the other did not. The results indicated that organisms normally found on or in empty shells influence the shell-section by these species of hermit crab. P. pollicarus preferred shells occupied by the sea anemone Calliactis tricolor or by the hydroid Hydractina echinata as opposed to bare shells. P. longicarpus also preferred shells with H. echinata. Both crab species rejected shells with the barnacle Balanus amphitrite. Shells containing the molluscs Crepidula fornicata or C. plana were rejected by the smaller hermit crab P. longicarpus. These molluscs appeared to exert no influence on P. pollicarpus unless they were large or abundant, at which point their weight or occlusion of available space possibly has negative effects on the crab.     

Effect of gastropod shell characteristics and hermit crabs on shell Epifauna

The epifauna on gastropod shells occupied by the hermit crabs Pagurus pollicaris (Say) and P. longicarpus (Say) was examined, as was the utilization of shells by these two hermit crabs. In the study area in Tampa Bay, Florida, shells were not a limiting factor to the hermit crab population, and there apparently was little competition for shells. Interspecific competition for shells was limited because the two hermit crab species differed in size and hence occupied shells of different sizes. The total number and density of most epifaunal species were higher on shells occupied by hermit crabs than on unoccupied shells, possibly because hermit crabs prevent their shells from being buried and hence lengthen the time the epifaunal community can grow and develop. The hermit crab species also appeared to affect the epifaunal community, for the total number and density of most epifaunal species were larger on shells occupied by P. pollicaris than P. longicarpus. With increasing shell size, the populations of most epifaunal species, also were larger but not their density. Least influential in affecting the epifaunal community was the species of shells.     

The species of Phora (Diptera: Phoridae) of Costa Rica

The Phora of Costa Rica were reviewed and three species, all belonging to Schmitz's Group III, were recognized: P. americana Schmitz and Wirth, P. truncata new species and P. paramericana new species. Of the three species, P. americana is by far the most abundant, with the other two species being rarely collected. All specimens were collected in the central highlands of the country. The previous record of the Holarctic Region species P. stictica Meigen from Costa Rica was found to be based on a misidentification of P. truncata.     

Structural and functional characterization of haemocyanin from the anemone hermit crab Dardanus calidus

Oxygen-binding to haemocyanin (Hc) is generally an exothermic process, with overall enthalphy of oxygenation varying from species to species. A number of crustacean Hcs showed a null or reduced enthalphy of oxygenation, among others, the anomuran Pagurus bernhardus and Paralithodes camtscaticae possess a completely temperature-independent oxygen-binding in a wide range of temperature and pH. Functional analysis performed on purified native, hexameric and dodecameric Hc forms of the anemone hermit crab Dardanus calidus allowed to calculate the enthalphy of oxygenation values that resulted equal to -36.2, -33.8 and -26.8 kJ/mol, respectively. Thus, the temperature sensitivity of oxygen binding of D. calidus Hc is in contrast with the temperature independence reported for P. bernhardus and P. camtscaticae, suggesting a high Hc functional heterogeneity within Anomura. Functional characterization also evidenced a strong oxygen affinity modulation by protons (DeltalogP(50)/DeltapH = -0.97) and lactate [DeltalogP(50)/Deltalog(lactate) = -0.38], and a significant decrease in cooperativity by physiological concentration of lactate (n(50) from 2.8 to 1.7 at pH 7.5).     

Periclimenes speciosus, a new species of anthozoan associated shrimp (Crustacea: Decapoda: Palaemonidae) from southern Japan

A new species of palaemonid shrimp, Periclimenes speciosus sp. nov., is described and illustrated on the basis of 20 specimens collected from warm-temperate and subtropical waters of southern Japan. Periclimenes speciosus belongs to the "P. aesopius species group", and is associated with sea anemones and a scleractinian coral as well as behaving as a fish cleaner. Morphologically, the new species appears closest to P. holthuisi Bruce, 1969, but can be distinguished from P. holthuisi by the form and armature of the cutting edges of dactylus and fixed finger of the second pereiopod. The coloration in life of both species clearly discriminates one species from the other. The taxonomic status of some specimens previously reported as P. holthuisi is briefly discussed.     

Seasonal variation in the occurrence of planktonic larvae of sympatric hermit crabs

Patterns of seasonal change in the abundances of the planktonic larvae of four sympatric hermit crabs were examined at five fixed stations over 18 months in Christmas Bay, Texas. Clibanarius vittatus (Bosc) was present in the plankton from April through October, Pagurus longicarpus Say from September through May and P. pollicaris Say from December through February. Pagurus annulipes (Stimpson) zoeae were the most frequently encountered zoeae and were found throughout the year, but were most abundant in late spring and summer. Zoea I of P. annulipes and P. pollicaris were found to be reliably distinguishable on the basis of their carapace lengths. Clibanarius zoea I were very abundant (3149) and 42 megalopa but no zoea II–V were found. Inadequate sampling and the short durations of these missing stages do not seem sufficient to explain their absence.