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1.
There are gaps in what is known about the patterns of gastropod shell use by hermit crabs in Brazilian lagoon areas, especially in Northeastern Brazil. However, this is important because the understanding of selection patterns provides information on life history and eco‐evolutionary conceptions of paguroids. The present study investigated the use of gastropod shells occupied by Pagurus criniticornis and Clibanarius sclopetarius in a coastal lagoon seasonally connected to an estuary and to the sea, correlating eco‐evolutionary aspects and hydrodynamic characters. The study was carried out between February 2013 and January 2014. Hermit crabs and their shells were identified and measured (hermit crabs’ cephalothorax shield length and width, major propodus length and height, weight, shell total length and width, shell aperture length and width). Partial least squares regression was used to analyse the morphometric data. Additionally, a multinomial proportions test was performed to infer patterns (inter‐/intra‐specific) of shells’ occupation. Hermit crab species occupied a total of 13 types of shells, predominantly those of Neritina virginea (67.83%). The shell weight was the most important determinant of shell occupancy in the morphometric model (variable importance in projection >1). The proportions of N. virginea shells used were similar in both species of hermit crabs, except between the sexes of P. criniticornis. Presumably, the high utilization rate of N. virginea shells is related to its abundance and dispersal, and to the shells’ suitability for hermit crabs, which is reflected by the morphometric model, as well as by the hydrodynamics of the lagoon‐estuarine environment. The relative occupation of different types of shells for each species of hermit crabs studied appears to be associated with regulation by inter‐/intra‐specific competition, which fosters the co‐occurrence of those populations. These results endorse coastal lagoons as a refuge and recruitment area for aquatic fauna. In addition, it highlights a challenge in the management and conservation of paguroid species whose population dynamics depend on the resources coming from the Gastropoda community.  相似文献   

2.
Abstract. Hermit crab shells are often encrusted with diverse communities of epibionts. To explore the effects of hermit crabs on the interactions of these encrusting species, two species of hydractiniid hydroids, Hydractinia symbiolongicarpus and Podocoryna carnea , were examined in the presence and absence of Pagurus longicarpus hermit crabs. Colonies growing on Littorina littorea snail shells with hermit crab hosts were compared to those growing on shells without crabs. These experiments suggest that hermit crabs variably affect colony polyp number, colony morphology and the outcome of interspecific competition, according to the size and number of crabs; other factors include water temperature, food availability and hydroid clonal genotype. In order to allow image analysis of perturbations of colony morphology, hydroids were grown on glass surfaces both exposed and unexposed to hermit crabs. Relative to colonies growing on unexposed surfaces, colonies growing on exposed surfaces exhibited larger within-colony patches, thus decreasing the total encrusted surface area and polyp number of colonies. Further, on glass surfaces, hermit crabs accelerated the time to the resolution of interspecific competition ( i.e. , the overgrowth of one colony by the other). While a number of factors may contribute to the relative abundance of these encrusting species, under most circumstances P. longicarpus have large effects on hydractiniid hydroids. Mechanisms underlying these effects likely include hydrodynamics and mechanical disturbance.  相似文献   

3.
Odours of different sources can indicate to hermit crabs the availability of empty shells, crucial resources for the life cycle of almost all of them. Here, we compared Clibanarius erythropus and Pagurus bernhardus for the intensity of investigative behaviour exhibited towards an empty, well-fitting shell in the presence of (1) plain seawater as control and seawater conditioned by (2) dead and live snails, (3) dead and live conspecifics, (4) live predators, and (5) food. During 10 min of observation, we recorded latency (the time until the first contact with the shell), and the number and duration of shell investigation bouts. The two species behaved similarly when exposed to the odours of food, live snails, and predators, while a more intense shell investigation was induced by dead snail odour in C. erythropus and by dead or live conspecific odour in P. bernhardus. Further studies should investigate the influence of phylogeny and ecology on this interspecific difference.  相似文献   

4.
Epibiont communities of Pagurus bernhardus and its inhabiting shell collected in the same month of two consecutive years near the Isle of Cumbrae (Scotland) were analyzed. The epibionts found were: (1) the protozoans Acineta, Conchacineta, Corynophrya, Zoothamnium, Cothurnia mobiusi, Cothurnia longipes, Chilodochona, Cryptacineta, Ephelota gemmipara and Ephelota plana; (2) the hydrozoans Leuckartiara, Clytia and Phialella; (3) the entoproct Barentsia; (4) the cirripeds Balanus balanus and Balanus crenatus; (5) the polychaetes Pomatoceros triqueter, Circeis amoricana paguri and Hydroides norvegica. Among these epibionts, a protozoan (E. plana), two hydrozoans (Leuckartiara, and Phialella), and the entoproct (Barentsia) have not been described previously as epibionts on crustaceans. The protozoan, entoproct and hydrozoan epibiont species found on P. bernhardus represent the first mention of their presence on this hermit crab. The protozoan epibionts were found only on the crab, while the hydrozoan epibionts were observed on the shell and on the crab. The cirriped species were observed only on the external surface of the shell, and the polychaete species were observed on the external and internal surfaces of the shell, although Circeis was found only internally. An analysis of the distribution and density of each epibiont species on the anatomical units of the crab, as well as on the different shell areas were made. The comparison between the data of the two years showed differences with respect to the size of crabs, diversity of epibiont groups and density of epibionts. There was a significant difference between the two years with respect to the distribution of epibionts on the anatomical units of the crab, both in terms of density and biomass. The influence of some environmental parameters on these differences is considered.  相似文献   

5.
Hermit crabs of the family Coenobitidae comprise the land hermit crabs, with 16 Coenobita species, and the coconut crab Birgus latro. They are terrestrial but spend their marine life as planktonic larvae. Some coenobitid crabs are widely distributed in the Indo‐West Pacific region, and some species occupy narrower ranges. To improve our knowledge of coenobitid crab speciation and geographical distribution patterns, we examined the phylogenetic relationship between Coenobita purpureus, which has a narrow distribution in the Northwestern Pacific, and its more widely distributed coenobitids including B. latro, Coenobita brevimanus, Coenobita cavipes, Coenobita perlatus, Coenobita rugosus and Coenobita violascens based on the mtDNA cytochrome c oxidase subunit I (COI) and 16S rDNA genes. We also assessed the phylogeography of Co. purpureus populations based on the COI gene. Our phylogenetic and phylogeographical analyses revealed that (i) Co. purpureus clustered with Co. rugosus and (ii) the Co. purpureus population in the Ogasawara Islands was genetically distinct from those in other regions, which showed genetic panmixia. It has been hypothesized that Co. purpureus evolved in the isolated landmasses of the Ryukyu region during the Pliocene, and that its population expanded and colonized the Ogasawara Islands in the late Pleistocene. Further phylogeographical studies on Coenobita species with relatively narrow distributions coupled with characterization of their phylogenetic relationships with widely distributed congeners will advance our knowledge of the speciation and geographical distribution history of coenobitid crabs.  相似文献   

6.
Female blue crabs (Callinectes sapidus Rathbun) migrate from low salinity estuarine regions to high salinity regions near the ocean to release larvae. During this migration, ovigerous females use ebb-tide transport, a vertical migratory behavior in which they ascend into the water column during ebb tides, to move seaward to larval release areas. In order to determine the relationship of ebb-tide vertical migrations to local currents and the influence of these vertical migrations on the horizontal transport of blue crabs in the estuary, ovigerous females with mature embryos (1–3 days from hatching) were tracked near Beaufort Inlet, North Carolina (USA), in July and August 2001 and 2002. Crabs were tagged and tracked using ultrasonic telemetry, and currents near the crabs were measured simultaneously with a shipboard acoustic Doppler current profiler.During the two seasons, eight crabs were successfully tracked for periods ranging from 3.9–37.0 h and for distances ranging from 1.9–10.6 km. All crabs migrated seaward during the tracking periods. Crabs moved episodically during all tidal phases with periods of movement on the order of minutes to an hour. They moved with local currents in terms of both speed and direction during ebb tides, consistent with ebb-tide transport, and moved down-estuary (seaward) in opposition to local currents during flood tides. The percentage of time that crabs were active was higher during night ebb tides than during day ebb tides or flood tides and increased with increasing ebb-tide current speed. Mean migratory speeds were 0.11, 0.04, 0.08 and 0.02 m s−1 during night ebb, night flood, day ebb and day flood tides, respectively, and net migratory speeds were on the order of 5 km day−1. Due to the episodic nature of the crabs' movements, the total distances that crabs traveled during ebb tides ranged from 10–40% of the distances that passive particles could have traveled under the same conditions.  相似文献   

7.
A series of over 6 000 research-trawl samples collected along the west and south coasts of South Africa between 1987 and 2014 were analysed for the presence and biomass of two parapagurid hermit crabs, Sympagurus dimorphus and Parapagurus bouvieri. The percentage of trawls that landed S. dimorphus and P. bouvieri and the mean caught biomass were higher on the west than on the south coast for both the more-abundant S. dimorphus (30.59 vs 5.81% success and 287.88 vs 31.37 kg km–2, respectively) and for the less-abundant P. bouvieri (13.76 vs 3.58% success and 38.56 vs 16.32 kg km–2, respectively). Very few parapagurids were caught shallower than 150 m; thereafter, the proportion of trawls containing hermit crabs increased, peaking over the depth range 201–250 m for S. dimorphus (54%) and 401–450 m for P. bouvieri (51%), and declining steadily thereafter. On the west coast, the relative caught biomass of S. dimorphus increased significantly from north to south, but there was no apparent latitudinal trend in relative biomass for P. bouvieri. Similarly, there was a significant decline in caught biomass of S. dimorphus with increasing longitude along the south coast, but no apparent trend for P. bouvieri. Although this represents by far the most comprehensive global analyses of distribution and abundance patterns for parapagurids to date, extremely little remains known about the biology and ecological relationships of these species, or indeed of other members of the group.  相似文献   

8.
The habitat in which predator–prey interactions take place may have a profound influence on the outcome of those interactions. Cannibalism is an intriguing form of predation whereby foraging by predators may contribute to the regulation of their own populations.This is particularly interesting in the case of invasive species, like the widely distributed European green crab (Carcinus maenas). This study explores how habitat complexity influences cannibalism rates in green crab populations of Prince Edward Island, Atlantic Canada. Both laboratory and field experiments were conducted to measure feeding rates by individual adult green crabs on a standard number of smaller conspecifics. In the laboratory, experimental treatments mimicked unstructured to increasingly structured habitats: water, sandy bottom, oyster shells, mussel shells, oyster shells with sandy bottom and mussel shells with sandy bottom. In those trials, adult green crabs consumed several times more juveniles on unstructured habitats than on the most structured ones, with a gradual decrease in predation rates across increasingly complex habitats. Field inclusion experiments used the same approach and were conducted in sandy bottoms, sandy bottoms with a layer of oyster shells and sandy bottoms with a layer of mussel shells. These trials showed similar patterns of decreasing feeding rates across increasingly complex habitats, but differences among treatments were not significant. These results support the idea that complex habitats have the potential to mediate predator–prey interactions, including adult–juvenile cannibalism in green crabs.  相似文献   

9.
Despite rapidly growing interest in the effects of ocean acidification on marine animals, the ability of deep-sea animals to acclimate or adapt to reduced pH conditions has received little attention. Deep-sea species are generally thought to be less tolerant of environmental variation than shallow-living species because they inhabit relatively stable conditions for nearly all environmental parameters. To explore whether deep-sea hermit crabs (Pagurus tanneri) can acclimate to ocean acidification over several weeks, we compared behavioral “boldness,” measured as time taken to re-emerge from shells after a simulated predatory attack by a toy octopus, under ambient (pH ～7.6) and expected future (pH ～7.1) conditions. The boldness measure for crab behavioral responses did not differ between different pH treatments, suggesting that future deep-sea acidification would not influence anti-predatory behavior. However, we did not examine the effects of olfactory cues released by predators that may affect hermit crab behavior and could be influenced by changes in the ocean carbonate system driven by increasing CO2 levels.  相似文献   

10.
Post-settlement processes are a major focus in the study of the dynamics of marine populations and communities. Post-settlement movement of juveniles is an important, but often ignored, process which affects local predator–prey and competitive interactions. We used benthic suction sampling and pitfall traps to examine density and locomotory activity of Carcinus maenas juveniles in different intertidal habitat types in the Rio Mira Estuary, Portugal, to better understand intra-specific interactions in a system where density-dependent processes are known to regulate population dynamics. As expected, significantly higher densities of juvenile shore crabs were found from bare mud compared to densely vegetated habitats. At the time of sampling, small and intermediate stages together outnumbered by far the larger juveniles. Conversely, larger crabs were much more frequent than smaller ones in traps. A locomotory index (LI), i.e. the ratio between crab catch in pitfall traps and their density within their moving range, is proposed as a measure of movement. LI analyses indicated that: (1) movement is an order of magnitude higher in large than small juveniles and much higher in sparse than dense vegetation cover; (2) activity of small juveniles is mostly crepuscular, regardless of vegetation cover; and (3) movement of large juveniles is very limited in dense Zostera patches, but very high in sparsely vegetated areas, during the day and night. These results suggest that small juveniles are relatively protected under dense vegetation cover due to lower mobility of larger crabs, and provide evidence of temporal segregation of activity windows between juvenile crabs of different sizes, which may be a key mechanism to reduce cannibalism and therefore increase the carrying capacity of nursery habitats.  相似文献   

11.
The sea anemones Calliactis conchicola Parry, 1952 and Paracalliactis rosea Hand, 1975, which live associated with gastropods, hermit crabs, and true crabs, possess mounting responses similar to those known for several species of Calliactis. In some areas of New Zealand Calliactis conchicola occurs most abundantly on living gastropods and in others on gastropod shells occupied by hermit crabs. No behaviour by either the gastropod or the hermit crabs was observed which explains this differential distribution. One hermit crab stimulates Paracalliactis rosea to mount its shell by a regular tapping of the anemone with its chelae or dactyls, but ignores Calliactis conchicola. Several other hermit crabs, which may be found carrying these anemones, do not stimulate them to mount. The spider crab Leptomithrax longipes may be found in nature carrying specimens of four kinds of anemones: Calliactis conchicola, Paracalliactis rosea, Bunodactis chrysobathys, and Phellia aucklandica. This crab prises these anemones from most substrates with its chelae, cleans them with its mouth parts, and places them on its legs and carapace with its chelae. From observation, Calliactis conchicola apparently protects this crab from octopus predatiori, but not from other predators such as small sharks. Though associations of anemones with hermit crabs and other invertebrates are usually considered to be commensal, it seems likely that these associations may be mutualistic.  相似文献   

12.
Pagurus maclaughlinae is the most common hermit in the Indian River Lagoon System. Wide variations in lagoonal salinity make it likely that P. maclaughlinae is euryhaline and that other hermit species in the area are more stenohaline, at least in some stages of their life histories. In a study of salinity tolerance, crabs were held unfed at salinities of 5–50 (25 control) for up to 30 days. Based on survivorship curves, P. maclaughlinae tolerated acute exposure to salinities of 10–45 for up to 18 days, and survivorship up to 30 days at 20–45 equaled or exceeded survivorship of the control. In a study of acclimation, the osmotic pressure of hemolymph was measured after crabs were held in the laboratory for 12, 48, and 96 h acutely exposed to salinities of 10–45. Paired t-tests revealed that the crabs weakly hyperregulated their hemolymph at 45–154 mOsmol above the external medium at all salinities and sampling times, and the osmotic differential of their hemolymph was fully acclimated by 96 h. In a third study, acclimatization of hemolymph was studied on crabs at four field sites that differed in their recent salinity histories. Field-collected crabs weakly regulated their hemolymph 72–84 mOsmol above the external medium at all sites sampled. Performance did not differ by site. The range of salinity tolerance and acclimation of hemolymph of P. maclaughlinae partly explain their wide distribution, and the consistent osmotic differential of its hemolymph indicates that the osmoregulatory ability of this small-bodied species is conserved in populations throughout the lagoon. Although some other larger-bodied hermit species in the region are euryhaline as adults, their tendency to hyperregulate strongly at low salinities possibly adds an energetic burden that, along with their less euryhaline long-lived larvae, might exclude them from the lagoon. Salinity tolerance of larval P. maclaughlinae has yet to be studied.  相似文献   

13.
Pagurus hermit crabs have a well‐developed right cheliped (major cheliped) and in some species the major cheliped of males is longer than that of females. This paper describes sex‐related differences in major cheliped length and regeneration pattern of the major cheliped in the hermit crab Pagurus filholi. We also examined the function of the major cheliped in male–male competition. Major cheliped length of males was longer than that of females in P. filholi. Males regenerated larger chelipeds than females at the first molt after experimentally induced autotomy. Body size growth in males of the regeneration group was less than that in intact males of the control group while there was no significant difference in body size growth of females between regeneration and control groups. Major cheliped length was included in the best model to explain the outcome of male–male competition and thus sexual selection appears to be a causative factor in the sex‐related difference of the major cheliped length. Sex‐related differences in the regeneration pattern may reflect differences in evolutionary pressures on males for large major chelipeds and females for large body size.  相似文献   

14.
For epibiotic or symbiotic marine invertebrates, alternative host species may differ substantially in quality, and under some circumstances such differences in host quality may lead to the evolution of increased host specificity. However, the fitness consequences of alternative hosts for epibiotic or symbiotic marine invertebrates have rarely been quantified. In Southern California, the gastropod Crepidula onyx is often found as an epibiont on either bay mussels (Mytilus galloprovincialis) or cone snails (Conus californicus). These hosts differ greatly in maximum size, with possible effects on size at sex change and final size in Cr. onyx, and thus on fecundity. Further, Cr. onyx on the two hosts differ in shell shape, possibly affecting the size of the ctenidium, which Cr. onyx uses for suspension feeding. We examined these potential effects of host use on fitness components in Cr. onyx. Epibionts on mussels reached much larger average sizes than did those on cone snails; further, epibionts on mussels often completed sex change at much larger sizes than did those on cone snails. On average, mussel epibionts had threefold higher average fecundities than did cone snail epibionts. Although there was a slight difference in shell shape between epibionts on the two host species, there was no difference in the scaling of ctenidium area with body size for Cr. onyx from the two hosts. The large average differences in fecundity in epibionts associated with the two alternative hosts suggests that there may be strong selection on host choice at larval settlement.  相似文献   

15.
Abstract. During 1986 and 1987 a decapod crustacean taxocoenosis of Posidonia oceanica beds from the southeast of Spain has been studied.
Data on abundance-dominance of the 50 species found and other information such as reproduction and the selective use of gastropod shells by the dominant hermit crabs arc given.
Differences in richness and cquitability between shallow (3–4 m) and deeper (5–7 m) samples arc also shown.
The taxocoenosis presents a geometrical distribution in which only 9 species have a relative abundance higher than 1 % and in which the structure is basically controlled by the two dominant hermit crabs, Cestopagitius limidus and Calcium tubularis , and hydrodynamic conditions.  相似文献   

16.
Pagurus exilis (Benedict, 1892) is an endemic South Atlantic hermit crab with a distribution ranging from Rio de Janeiro State, Brazil, to Buenos Aires Province, Argentina. The present study analyzed the reproduction of two populations at the extremes of this geographical distribution, and compared their reproductive period, fecundity, and changes in egg size and egg volume during the incubation period in order to assess the variability over this latitudinal range. Hermit crabs were collected monthly over a 2‐year period. In total, 108 females were analyzed for Brazil (44 non‐ovigerous and 64 ovigerous), and 141 for Argentina (87 non‐ovigerous and 54 ovigerous). Reproduction in Brazil occurs year‐round, with peaks in the fall and winter seasons; in Argentina reproduction occurs only in spring and summer. The Brazilian ovigerous females were significantly larger than the Argentina ones (Brazil: SL = 5.33 ± 1.45 mm; Argentina: SL = 4.15 ± 0.52 mm; P < 0.001). The fecundity was 1447 ± 831 eggs (317 to 2885) in Brazil and 987 ± 711 eggs (114 to 2665) in Argentina, with a trend towards higher fecundity in Brazil. Eggs in the Argentina population were larger than those in Brazil for all the three stages investigated, and no changes in egg volume were found during egg development for both populations. The reproductive traits of the two populations showed some important differences, which may reflect adaptations to local environmental conditions, demonstrating a high plasticity of reproductive features of the species in Brazilian and Argentine waters. The strategy adopted by the Argentina population involves a lower production of larger eggs compared to the population in Brazil; this lower production is associated with reproduction in cold‐water regions.  相似文献   

17.
Ecosystems are fundamentally connected to one another, although empirical documentation of the links between them, especially the links between marine and terrestrial ecosystems, has been limited. This study examined the foraging behavior and ecology of an understudied invertebrate, the hermit crab Coenobita compressus, which is remarkably abundant at the nexus of marine, terrestrial, and freshwater ecosystems. Data collected between 2008 and 2011 in the Osa Peninsula of Costa Rica revealed that C. compressus plays a generalist scavenger role across ecosystems. Beach almond fruits (Terminalia catappa) were a staple of the crabs’ diet, but the crabs also commonly consumed a variety of other fruit species and various parts of plants (nuts, seeds, flowers, husk, stems, buds) as well as algae, fungus, animal feces, and the carcasses of insects, jellyfish, conspecifics, other crustacean species, freshwater and saltwater fish, sea turtles (eggs and adults), and mammals. The crabs fed on these resources in prominent social aggregations, also forming such aggregations as they exchanged the aquatically derived gastropod shells that they use as homes while roaming the land. The size of the crabs’ aggregations around plant versus animal resources and around terrestrial versus aquatic resources did not differ, underscoring the species’ generalist foraging behavior. Broadly, these results suggest that C. compressus has one of the most diverse diets of any crustacean and plays a critical role in recycling organic matter and nutrients. This species might also mediate the passage of resources across different ecosystems, although further research is needed to explore this possibility.  相似文献   

18.
Changes in the concentration of lipovitellin, a lipoprotein that is the major protein in mature oocytes of crustaceans, were determined during ovarian development in female blue crabs, Callinectes sapidus. To test the effects of reproductive toxins on vitellogenesis, crabs were fed food containing cadmium or injected with the juvenile hormone antagonist, precocene II. Monoclonal antibodies prepared against one of the lipovitellin peptides (peptide A—107 kDa) were used in an enzyme linked immunosorbent assay to determine lipovitellin concentrations. Recently molted female crabs fed food containing cadmium (0.02 mg/g) for 18 days had oocytes with the same lipovitellin concentration as controls, but after 24 days the oocytes in cadmium treated crabs had significantly lower lipoprotein contents than controls (30 ± 4 ng/oocyte—cadmium treated; 45 ± 4 ng/oocyte—control). Precocene II-treated crabs had significantiy lower lipovitellin concentrations after 18 and 24 days. Diameters of oocytes were the same in both treated and control crabs. The results suggest that some pollutants may affect reproduction by acting during secondary vitellogenesis i.e., period when there is a large increase in lipovitellin synthesis. Since lipovitellins serve many of the nutritional needs of recently emerged crab embryos, poor larval survival may occur as a result of lower lipovitellin content.  相似文献   

19.
There is a growing awareness of the role that indirect interactions play in influencing food webs and ecosystem structure. In this study, the hypothesis that crustose algal epibionts provide gastropods associational resistance from predation was investigated through field surveys and laboratory feeding assays. In rocky low intertidal/shallow subtidal systems in the northeast Pacific, several species of crustose algae (the red alga Peyssonnelia meridionalis and crustose corallines) can colonize the shells of living Tegula brunnea snails. The growth patterns of these epibiontic crustose algae allow them to cover their host’s surface completely, which may, in turn, protect their hosts from predation. A multi‐site field survey of T. brunnea revealed that >60% of snails were at least 75% covered with one or more species of crustose algae, with 35% fully covered, indicating that this is common in the field. Laboratory feeding assays revealed that sea stars, a primary predator of T. brunnea, distinguished among snails with different shell coverings; Pisaster consumed nearly three times as many bare (i.e. no crustose algae) snails as those covered with Peyssonnelia, while Pycnopodia consumed four times as many bare snails as those covered with crustose corallines. These results suggest that epibiont crustose algae can benefit their hosts via associational resistance; this finding may have implications for the role of associational resistance in trophic interactions.  相似文献   

20.
Mangrove ecosystems are sites with high biodiversity of benthic fauna, and fiddler crabs (genus Uca) are common benthic fauna in mangroves. The North Sulawesi in Indonesia has a good condition of mangrove while the information of the fiddler crabs is still limited. Manual samplings were conducted in wet, dry and transient seasons at a mangrove in Kema, North Sulawesi to investigate the species composition, density and distribution pattern of fiddler crabs. A total of 168 individuals, subjected to eight species of genus Uca crabs were collected at the mangrove, with U. triangularis having the highest abundance and U. annulipes having the lowest abundance. The densities of fiddle crabs were 27.56 ind./m2, 32.89 ind./m2 and 14.22 ind./m2 at the seaward, middle and landward zones, respectively, and the density was higher in dry and wet seasons than in transient season.  相似文献   

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