Effects of shell status and social context on the agonistic behavior of the tropical hermit crab,Clibanarius signatus

In the hermit crabClibanarius signatus, the influence of both shell status and social context on agonistic behavior was examined. The experimental hermits were of the same size and sex, but differed in the status of the inhabited shell, which was either optimal, sub-optimal or damaged. The evidence from this experiment suggests that the agonistic behavior ofC. signatus is influenced to a great extent by asymmetries in the resource value; attacks are mainly performed by crabs which inhabit shells in a relatively poor condition and are mostly directed towards opponents with better-fitting shells. However, this effect differs according to the social context, being more pronounced when the potential gain is greater. Shell fights are decidedly more complex than most animal combats; more work is certainly needed to understand the mechanisms underlying resource assessment and decisions made during hermit crabs' interactions.     

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.     

A kinetic study of the gill (Na+, K+)-ATPase, and its role in ammonia excretion in the intertidal hermit crab, Clibanarius vittatus

To better comprehend the role of gill ion regulatory mechanisms, the modulation by Na(+), K(+), NH(4)(+) and ATP of (Na(+), K(+))-ATPase activity was examined in a posterior gill microsomal fraction from the hermit crab, Clibanarius vittatus. Under saturating Mg(2+), Na(+) and K(+) concentrations, two well-defined ATP hydrolyzing sites were revealed. ATP was hydrolyzed at the high-affinity sites at a maximum rate of V=19.1+/-0.8 U mg(-1) and K(0.5)=63.8+/-2.9 nmol L(-1), obeying cooperative kinetics (n(H)=1.9); at the low-affinity sites, hydrolysis obeyed Michaelis-Menten kinetics with K(M)=44.1+/-2.6 mumol L(-1) and V=123.5+/-6.1 U mg(-1). Stimulation by Na(+) (V=149.0+/-7.4 U mg(-1); K(M)=7.4+/-0.4 mmol L(-1)), Mg(2+) (V=132.0+/-5.3 U mg(-1); K(0.5)=0.36+/-0.02 mmol L(-1)), NH(4)(+) (V=245.6+/-9.8 U mg(-1); K(M)=4.5+/-0.2 mmol L(-1)) and K(+) (V=140.0+/-4.9 U mg(-1); K(M)=1.5+/-0.1 mmol L(-1)) followed a single saturation curve and, except for Mg(2+), obeyed Michaelis-Menten kinetics. Under optimal ionic conditions, but in the absence of NH(4)(+), ouabain (K(I)=117.3+/-3.5 mumol L(-1)) and orthovanadate inhibited up to 67% of the ATPase activity. The inhibition studies performed suggest the presence of F(0)F(1), V- and P-ATPases, but not Na(+)-, K(+)- or Ca(2+)-ATPases as contaminants in the gill microsomal preparation. (Na(+), K(+))-ATPase activity was synergistically modulated by NH(4)(+) and K(+). At 20 mmol L(-1) K(+), a maximum rate of V=290.8+/-14.5 U mg(-1) was seen as NH(4)(+) concentration was increased up to 50 mmol L(-1). However, at fixed NH(4)(+) concentrations, no additional stimulation was found for increasing K(+) concentrations (V=135.2+/-4.1 U mg(-1) and V=236.6+/-9.5 U mg(-1) and for 10 and 30 mmol L(-1) NH(4)(+), respectively). This is the first report to detail ionic modulation of gill (Na(+), K(+))-ATPase in C. vittatus, revealing an asymmetrical, synergistic stimulation of the enzyme by K(+) and NH(4)(+), as yet undescribed for other (Na(+), K(+))-ATPases, and should provide a better understanding of NH(4)(+) excretion in pagurid crabs.     

Filter feeding in the hermit crab

Summary The hermit crab, Pagurus bernhardus is able to remove both Artemia nauplii and unicellular algae from suspension. Crabs with wet weights of 1.1–9.2 g consumed all of the 300 Artemia nauplii contained in 200 ml of sea water within 1 h. Crabs weighing 0.7–1.1 g wet weight filtered suspended Dunaliella algae (8 m) from concentrations of 10–350 million cells per liter at a rate of 26% and 47% within 2 and 5 h, respectively. A similar result was obtained with an 11 g crab. During filter feeding activity a water current is generated by the flagella of the exopods of the second and third maxillipeds. Artemia nauplii are caught by grasping movements of the endopods of the third maxillipeds, whereas filtering of unicellular algae is probably achieved by the two maxillae. It is assumed that filter feeding activity depends on the same structures and behavior as in deposit feeding. P. bernhardus is one more example of a benthic marine animal which may use any food source which becomes available in the course of the seasons.     

Morphological and physiological properties of the giant interneuron of the hermit crab (Pagurus pollicaris)

The physiological and morphological properties of the giant interneurons in the hermit crab Pagurus pollicaris are described. The cell bodies are located anteriorly in the supraesophageal ganglion, close to the mid-line. Each cell sends a neurite posteriorly and then laterally, so that they cross over in the center of the ganglion. Each axon then branches: one branch runs laterally while the other travels posteriorly and leaves the ganglion in the circumesophageal connective on the side contralateral to the cell body. The giant axons travel in the circumesophageal connectives and through the thoracic and abdominal ganglia without branching. Each giant axon makes synaptic contact with its ipsilateral giant abdominal flexor motor neuron and with a second flexor motor neuron that has its axon in the contralateral third root. In the supraesophageal ganglion there is a bidirectional synapse between the two giant interneurons. Intracellular recordings from the giant axons show that there is a delay of 0.5 to 0.75 ms that cannot be accounted for by spike propagation along the axons, and may be accounted for by a chemical synapse between the giant interneurons.     

Microsatellite analysis of genetic diversity and population structure of Chinese mitten crab （Eriocheir sinensis）

Chinese mitten crab (Eriocheir sinensis) has higher commercial value as food source than any other species of Eriocheir in China.To evaluate the germplasm resources and characterize the genetic diversity and population structure of the crabs in different water systems,two stocks and two farming populations were assessed with 25 polymorphic microsallite loci available in public GenBank.Basic statistics showed that the average observed heterozygosity (Ho) amongst populations ranged from 0.5789 to 0.6824.However,a remarkable presence of inbreeding and heterozygote deficiencies were observed.To analyze population structure,pairwise FST coefficients explained only ～10.3% variability from the subdivision of mitten crab populations,the remaining variability stems from the subdivision within subpopulations.Although the four populations had slight differentiation,different allelic frequencies resulted in distinct population structures.Two stocks and one farming population were clustered together to the phylogenetic branch of Yangtze crab,with an approximate membership of 95%.Whereas,another fanning population was clustered singly to the phylogenetic branch of the Liaohe crab,with a membership of 97.1%.The tests for individual admixture showed that Yangtze crab had probably been contaminated with individuals from other water systems.Genetic relationships between populations also supported the conclusion that Yangtze crab and Liaohe crab had different gene pools in spite of the origins of the same species.     

Phylogeny and genetic diversity of palolo worms (Palola, Eunicidae) from the tropical North Pacific and the Caribbean

Palolo worms (Palola, Eunicidae) are best known for their annual mass spawnings, or "risings," in the South Pacific. Palola currently contains 14 morphologically similar species, mostly from shallow tropical waters. In this study, 60 specimens of Palola from nine locations in the tropical North Pacific and the Caribbean were sequenced for the two mitochondrial markers cytochrome c oxidase subunit I and 16S ribosomal RNA to infer phylogenetic relationships, genetic diversity, and phylogeography within the taxon. Phylogenetic analysis was performed using Bayesian statistics and parsimony. Vouchers of the same specimens were examined morphologically. Two major clades (A and B) can be distinguished within the monophyletic Palola. A number of individuals in clade B bear rows of ventral eyespots in the posterior body region, typical for swarming P. viridis and probably a synapomorphy for clade B. No morphological synapomorphy was found for clade A. Haplotypes from divergent clades often co-occur in the same location. Some haplotypes are geographically widespread, in one case covering the entire east-west expansion of the tropical Pacific. These results imply that despite the apparent absence of teleplanic larvae in eunicid polychaetes, long-distance dispersal is possible in at least some lineages of Palola. With the first taste of palolo I understood the Samoans' love for it. Certainly it suggested a salty caviar, but with something added, a strong, rich whiff of the mystery and fecundity of the ocean depths. -R. Steinberg. Pacific and Southeast Asian cooking. Time-Life Books, New York, 1970.     

Phylogeography and genetic diversity of East Asian Neolitsea sericea (Lauraceae) based on variations in chloroplast DNA sequences

Neolitsea sericea is an evergreen broad leaved tree in the warm-temperate regions of East Asia. This area is a hotspot for plant species richness and endemism caused by dynamic changes in land configuration during the Quaternary. However, the historical migration of such evergreen tree species is still poorly understood. In an attempt to reconstruct the phylogeographic history of N. sericea during the Quaternary, we identified the chloroplast DNA haplotypes of 287 individuals from 33 populations covering almost all of its geographic range. Analyses were based on sequence data from the trnL–F, psbC–trnS, and rps16 regions. Nine haplotypes were identified. The majority included ancestral types in the southwestern part of the main islands of Japan, with other region-specific haplotypes being found in populations on the Korean Peninsula, Taiwan (Isl. Lanyu), and elsewhere in Japan. A statistical parsimony network revealed two lineages derived from Japanese main islands. One was represented on the Korean Peninsula, the other on Isl. Lanyu. The current distribution of N. sericea has been shaped by colonization via land bridges. During the glacial periods, two primary, but separate migration routes were followed—from the southwestern part of the Japanese main islands to either the Korean Peninsula or Taiwan. In addition, we believe the Zhoushan populations were shaped by post-glacial processes through an ECS land bridge (East China Sea basin) from northern refugia that existed during the late Pleistocene.     

Dramatic mitochondrial gene rearrangements in the hermit crab Pagurus longicarpus (Crustacea, anomura)

The entire mitochondrial gene order of the crustacean Pagurus longicarpus was determined by sequencing all but approximately 300 bp of the mitochondrial genome. We report the first major gene rearrangements found in the clade including Crustacea and Insecta. At least eight mitochondrial gene rearrangements have dramatically altered the gene order of the hermit crab P. longicarpus relative to the putatively ancestral crustacean gene order. These include two rearrangements of protein-coding genes, the first reported for any nonchelicerate arthropod. Codon usage and amino acid sequences do not deviate substantially from those reported for other crustaceans. Investigating the phylogenetic distribution of these eight rearrangements will add additional characters to help resolve decapod phylogeny.     

Structure of seasonal migrations of the littoral hermit crab Clibanarius vittatus (Bosc)

A seasonal migration in the hermit crab Clibanarius vittatus (Bosc) is described and its structure examined for size-related or sexual differences that might lead to differential mortalities. The largest crabs, which were nearly all males, began to leave the shore in early summer, but the smaller females stayed until late autumn. The smaller crabs also returned to the shore at least one month in advance of the large crabs in the spring. No size-related or sexual differences were found in the abilities of the crabs to regulate their volumes when moved from 34.5%. to 10.5%. salinity which is the observed salinity range in their usual habitat.     

Phylogeography of Cycas revoluta Thunb. (Cycadaceae) on the Ryukyu Islands: very low genetic diversity and geographical structure

Variations in the chloroplast and mitochondrial DNA of Cycas revoluta Thunb. (Cycadaceae) were examined in 22 populations distributed across the Ryukyu Islands and southern Kyushu. Among the 14,130 bp of sequence examined, only one site mutation and one indel were polymorphic. The identified polymorphisms were located in the spacers between trnS (UGA) and trnfM (CAU) of the chloroplast DNA and between nad1 exon B and exon C of the mitochondrial DNA, respectively. Three haplotypes were identified from the Ryukyu Islands and southern Kyushu. The areas of distribution of the three haplotypes were highly geographically structured. The boundaries of two of the three haplotypes were demarcated by Okinoerabujima Island in the middle Ryukyus. The northern type and southern types lay north and south of the island, respectively. The third haplotype was almost sympatrically distributed with the southern type. The genetic variation within C. revoluta was estimated to be very low (h = 0.641, π = 0.00071) in comparison to its relative in Taiwan, C. taitungensis, which possesses 97 cpDNA haplotypes and 55 mtDNA haplotypes from two relic populations. A reasonable explanation for the low genetic diversity of the cycad on the Ryukyu Islands could be severe bottleneck effects, resulting from the submersion of low islands and the diminished landmass of islands in the interglacial age in the Quaternary period. The geographically restricted nature of the haplotypes could be attributed to vicariance resulting from the land configuration of the Ryukyu Islands, including changes in geography during the interglacial age in the Quaternary.     

Phylogeography and genetic diversity of the red squirrel (Sciurus vulgaris) in China: Implications for the species' postglacial expansion history

The geographical distribution of Sciurus vulgaris spans much of the Palearctic from western Europe and the UK eastward to the pacific coast of East Asia. S. vulgaris occurs in China only in the far northwest and the northeast. Understanding of the species’ postglacial expansion history in East Asia has been limited by the paucity of molecular data. In this study, we used partial D-loop and cytochrome b gene sequences to assess mitochondrial DNA variation in S. vulgaris in China. Our objectives were to (1) determine phylogeographical patterns of S. vulgaris in China; (2) understand the species’ postglacial expansion history in this region; and (3) quantify genetic diversity levels within S. vulgaris populations in China. We identified a supported phylogenetic group in S. vulgaris from China, and found no tendency for haplotypes to cluster by geographic region. Our analysis of S. vulgaris from China and other regions supports the hypothesis that the Calabria region of southern Italy is a glacial refugium for the species. We tentatively propose a postglacial expansion pattern for the squirrels: migrating from Calabria via Central and Eastern Europe to Russia and from there to China, and firstly to the northwest and then to northeast in China. We found high levels of genetic diversity in S. vulgaris populations across China as a whole, and discussed its influential factors.     

Phylogeography and population genetic structure of double-crested cormorants (Phalacrocorax auritus)

We examined the genetic structure of double-crested cormorants (Phalacrocorax auritus) across their range in the United States and Canada. Sequences of the mitochondrial control region were analyzed for 248 cormorants from 23 breeding sites. Variation was also examined at eight microsatellite loci for 409 cormorants from the same sites. The mitochondrial and microsatellite data provided strong evidence that the Alaskan subspecies (P. a. cincinnatus) is genetically divergent from other populations in North America (net sequence divergence = 5.85 %; Φ_ST for mitochondrial control region = 0.708; F_ST for microsatellite loci = 0.052). Historical records, contemporary population estimates, and field observations are consistent with recognition of the Alaskan subspecies as distinct and potentially of conservation interest. Our data also indicated the presence of another divergent lineage, associated with the southwestern portion of the species range, as evidenced by highly unique haplotypes sampled in southern California. In contrast, there was little support for recognition of subspecies within the conterminous U.S. and Canada. Rather than genetically distinct regions corresponding to the putative subspecies [P. a. albociliatus (Pacific), P. a. auritus (Interior and North Atlantic), and P. a. floridanus (Southeast)], we observed a distribution of genetic variation consistent with a pattern of isolation by distance. This pattern implies that genetic differences across the range are due to geographic distance, rather than discrete subspecific breaks. Although three of the four traditional subspecies were not genetically distinct, possible demographic separation, habitat differences, and documented declines at some colonies within the regions, suggests that the Pacific and possibly North Atlantic portions of the breeding range may warrant differential consideration from the Interior and Southeast breeding regions.     

Phylogeography of Pinus subsection Australes in the Caribbean Basin

Background and Aims

Four species of Pinus subsection Australes occur in the Caribbean Basin: P. caribaea, P. cubensis, P. maestrensis and P. occidentalis. This study analyses the phylogeography of these species to assess possible colonization events from Central America to the islands and subsequent population expansions during glacial periods driven by both drier climate and larger emerged land areas.

Methods

Allele size data were obtained for plastid microsatellites for 314 individuals from 24 populations, covering the distribution range of subsection Australes in the Caribbean Basin.

Key Results

In total, 113 plastid haplotypes were identified. The highest genetic diversity was found in populations of P. caribaea. Overall, Caribbean Basin populations fit the isolation by distance model. Significant phylogeographical structure was found (R_ST = 0·671 > permuted R_ST = 0·101; P < 0·0001). The haplotype network and a Bayesian analysis of population structure (BAPS) indicated different Central American origins for P. caribaea var. bahamensis and P. caribaea var. caribaea plastids, with Central America populations in northern and south-eastern groups. Sudden expansion times for BAPS clusters were close to three glacial maxima.

Conclusions

Central America contains ancestral plastid haplotypes. Population expansion has played a major role in the distribution of genetic diversity in P. caribaea var. hondurensis. Two colonization events gave rise to the P. caribaea var. bahamensis and P. caribaea var. caribaea lineages. Plastid variation in the eastern species (P. cubensis, P. maestrensis and P. occidentalis) evolved independently from that in P. caribaea var. caribaea. Incomplete lineage sorting between P. cubensis and P. maestrensis is apparent. Inferred expansion times for P. caribaea var. bahamensis and for the eastern lineages correspond to glacial maxima, whereas those for P. caribaea var. hondurensis correspond to the beginning of the temperature decrease that led to Marine Isotope Stage 8.     

Phylogeography of the South China field mouse (Apodemus draco) on the southeastern Tibetan Plateau reveals high genetic diversity and glacial refugia

The southeastern margin of the Tibetan Plateau (SEMTP) is a particularly interesting region due to its topographic complexity and unique geologic history, but phylogeographic studies that focus on this region are rare. In this study, we investigated the phylogeography of the South China field mouse, Apodemus draco, in order to assess the role of geologic and climatic events on the Tibetan Plateau in shaping its genetic structure. We sequenced mitochondrial cytochrome b (cyt b) sequences in 103 individuals from 47 sampling sites. In addition, 23 cyt b sequences were collected from GenBank for analyses. Phylogenetic, demographic and landscape genetic methods were conducted. Seventy-six cyt b haplotypes were found and the genetic diversity was extremely high (π = 0.0368; h = 0.989). Five major evolutionary clades, based on geographic locations, were identified. Demographic analyses implied subclade 1A and subclade 1B experienced population expansions at about 0.052-0.013 Mya and 0.014-0.004 Mya, respectively. The divergence time analysis showed that the split between clade 1 and clade 2 occurred 0.26 Mya, which fell into the extensive glacial period (EGP, 0.5-0.17 Mya). The divergence times of other main clades (2.20-0.55 Mya) were congruent with the periods of the Qingzang Movement (3.6-1.7 Mya) and the Kun-Huang Movement (1.2-0.6 Mya), which were known as the most intense uplift events in the Tibetan Plateau. Our study supported the hypothesis that the SEMTP was a large late Pleistocene refugium, and further inferred that the Gongga Mountain Region and Hongya County were glacial refugia for A. draco in clade 1. We hypothesize that the evolutionary history of A. draco in the SEMTP primarily occurred in two stages. First, an initial divergence would have been shaped by uplift events of the Tibetan Plateau. Then, major glaciations in the Pleistocene added complexity to its demographic history and genetic structure.     

Dominance and effects of strange conspecifics on aggressive interactions in the hermit crab Pagurus longicarpus (say)

Dominance orders in the hermit crab Pagurus longicarpus were observed in the laboratory. Groups of four crabs formed loose dominance hierarchies as determined by repeated display and retreat behaviour. Stronger dominance orders were inversely correlated with the frequency of aggressive interactions. Recognition of individuals, as measured by frequency and intensity of aggressive encounters between familiar crabs and introduced strangers, was not important in maintaining dominance orders. Rather, P. longicarpus recognized the aggressive state of conspecifics, as shown by frequencies of aggressive encounters after individuals of different dominance rankings were exchanged between established hierarchies.     

Phylogeography, genetic diversity and conservation of the large copper butterfly Lycaena dispar in Europe

We argue that insect species conservation at large scales should take account of the distribution of genetic diversity among populations. Maintenance of genetic diversity may be vital in retaining a species' adaptive capacity and evolutionary potential. We illustrate the concept using the example of the large copper butterfly Lycaena dispar in Europe. This species has become extinct in parts of its range and is declining rapidly in others, whilst conversely, increasing in many areas. The latter has recently reduced its conservation status. Mitochondrial DNA analysis is used to construct a phylogeography from a preliminary sample set obtained from across Europe. A cytochrome b fragment of 402 base pairs was sequenced and 10 haplotypes were found. Relatedness among populations suggest that those from northern and central Europe are closely related and probably form one evolutionary significant unit (ESU) reflecting post-glacial colonization from southeast Europe. In contrast, the sample from Italy is divergent and should be considered a separate ESU. Our results, combined with ecological data, suggest that conservation action for this species should be targeted on specific regions and populations.     

Phylogeography of the invasive seaweed Asparagopsis (Bonnemaisoniales, Rhodophyta) reveals cryptic diversity

The rhodophyte seaweed Asparagopsis armata Harvey is distributed in the northern and southern temperate zones, and its congener Asparagopsis taxiformis (Delile) Trevisan abounds throughout the tropics and subtropics. Here, we determine intraspecific phylogeographic patterns to compare potential causes of the disjunctions in the distributions of both species. We obtained specimens throughout their ranges and inferred phylogenies from the hypervariable domains D1-D3 of the nuclear rDNA LSU, the plastid spacer between the large and small subunits of RuBisCo and the mitochondrial cox 2-3 intergenic spacer. The cox spacer acquired base changes the fastest and the RuBisCo spacer the slowest. Median-joining networks inferred from the sequences revealed the absence of phylogeographic structure in the introduced range of A. armata, corroborating the species' reported recent introduction. A. taxiformis consisted of three nuclear, three plastid and four mitochondrial genetically distinct, lineages (1-4). Mitochondrial lineage 3 is found in the western Atlantic, the Canary Islands and the eastern Mediterranean. Mitochondrial lineages 1, 2, and 4 occur in the Indo-Pacific, but one of them (lineage 2) is also found in the central Mediterranean and southern Portugal. Phylogeographic results suggest separation of Atlantic and Indo-Pacific lineages resulted from the emergence of the Isthmus of Panama, as well as from dispersal events postdating the closure event, such as the invasion of the Mediterranean Sea by mitochondrial lineages 2 and 3. Molecular clock estimates using the Panama closure event as a calibration for the split of lineages 3 and 4 suggest that A. taxiformis diverged into two main cryptic species (1 + 2 and 3 + 4) about 3.2-5.5 million years ago (Ma), and that the separation of the mitochondrial lineages 1 and 2 occurred 1-2.3 Ma.