Variations in crab claw morphology and diet across contrasting inter‐tidal habitats

Crabs are important predators of inter‐tidal ecosystems, controlling the abundance and distribution of their prey populations. Often the same crab species occupies several habitats and, although their effects on prey have been quantified across habitats, crabs’ dietary and morphological responses to differing environmental influences have been overlooked. Here, we used the crabs Eriphia verrucosa and Pachygrapsus marmoratus as model species to examine differences in claw morphometry – size and wear – and diet between rocky shore and heterogeneous sand flat habitats. We predicted that, intra‐specifically, crabs from rocky shores would consume more hard‐shelled prey owing to their high availability and consequently, would display chelipeds with the following claw characteristics: a higher degree of claw damage, stronger musculature (higher propel height) and increased mechanical advantage (defined as the ratio of input lever length to output lever length) than crabs in the heterogeneous sand flat habitats. Sampling was performed in heterogeneous sand flat habitats and rocky shores of the Central Portuguese coast. For each crab species, carapace width, diet composition and several claw morphometric measures were recorded, revealing significant intra‐specific differences (using multivariate analysis) between shore types. We found that E. verrucosa and P. marmoratus consumed more hard prey on rocky shore than on sand flat habitats, which resulted in rocky shore crabs having more accentuated dentition wear and larger musculature than their sand flat habitat counterparts. We suggest that the strong response of crab claw morphometry to environmentally induced diet variations is an important mechanism in the successful adaptation of crab species to inhabit differing habitats. A major implication is that the impact of the same species on prey may vary largely with habitat type as a result of predation efficiency varying with claw condition.     

The intermingling of benthic macroinvertebrate communities during a period of shifting range: The “East of Nantucket” Atlantic Surfclam Survey and the existence of transient multiple stable states

A survey of the region eastward of Nantucket provided an opportunity to examine the cold temperate–boreal boundary along the high‐energy Great South Channel. Here described are the benthic macroinvertebrate community types encountered, with a focus on the influence of climate change on the range boundaries of the benthic biomass dominants and the potential existence of transient multiple stable states. The survey identified three primary community types. The shallowest sites were occupied by a surfclam‐dominated community, comprising an abundance of large (≥150 mm) surfclams, and a few common attached epibiota primarily attached to exposed surfclam shell. Two communities exist at intermediate depths, one dominated by submarket and small market‐size surfclams (<150 mm) and the other, created by mussel mats and their attendant epibiota, crabs, sea urchins, and other mobile epifauna. Mussels are a foundational species, establishing a hard‐bottom terrain conducive to these other denizens in soft‐bottom habitat. Cobbles were nearly ubiquitous, rocks were routinely recovered, and boulders were encountered occasionally. Slow growing attached epibionts were exceedingly rare and mobile epifauna were not obviously associated with these large sedimentary particles; nor were the surfclam or mussel communities. The frequency of barnacle scars suggests sediment scour under the high‐flow regime characteristic of the surveyed region, which voids the habitat potential of these sedimentary particles. The abundance of surfclam shell indicates that surfclams have inhabited the shoaler depths for an extended time; limited shell at deeper sites supports the inference from the absence of large animals that these sites are relatively newly colonized and represent further evidence of an offshore shift in range brought on by increasing bottom water temperatures. The dichotomous nature of the two primary community types at mid‐depths suggests that these two communities represent multiple stable states brought on by the interaction of an invading cold temperate species with the receding boreal fauna resulting in a transient intermingling of species, which, however, structure the habitat into exclusionary stable states rather than overlapping in a co‐occurrence ecotone.     

Genetic diversity and population structure of the size‐selectively harvested owl limpet,Lottia gigantea

Size‐selective harvesting can elicit a genetic response in target species through changes in population genetic subdivision, genetic diversity and selective regimes. While harvest‐induced genetic change has been documented in some commercially important species through the use of historic samples, many commonly harvested species, such as coastal molluscs, lack historic samples and information on potential harvest induced genetic change. In this study, we have genotyped six microsatellite markers from populations across much of the California mainland range of the size‐selectively harvested owl limpet (Lottia gigantea) to explore the genetic structure and diversity of this species. We found no significant genetic structure or differences in genetic diversity among populations of L. gigantea. Our results suggest high gene flow among populations and that differences in life history, demography, and body size previously observed between protected and exploited populations is largely due to phenotypic plasticity. From a conservation perspective, if proper actions are taken to curb harvesting, then exploited populations should be able to return to their pre‐impact state given sufficient time.     

Shell use by sympatric hermit crab species in a seasonally open coastal lagoon in Northeastern Brazil

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.     

Shell‐shape and morphometric variability in Mytilus galloprovincialis from micro‐tidal environments: responses to different hydrodynamic drivers

Tidal conditions differently influence inter‐tidal organisms in terms of general physiological and metabolic responses. In this study we investigated the morphological response in shells of Mytilus galloprovincialis native to different micro‐tidal coastal environments in the Northern Adriatic Sea. Our purpose was to highlight the ecophenotypic variability across tidal levels and to elucidate how tidal currents and waves produced by anthropogenic activities may play a part in modulating shell morphology. Three sampling sites were selected: an open‐sea area 15 km off‐shore and two sites within the lagoon of Venice, the first near one of its three inlets, and the other one in the proximity of Venice city centre. At each sampling site, organisms were seasonally collected at different depths within their vertical zonation, either in the inter‐tidal zone – i.e. at both the highest and lowest tide zonation limits, and subtidally. The mussel shells were analysed by investigation of their morphometric relationships (height/length and width/length ratios) and by elliptic Fourier analysis of the shell contours. Shell thickness and condition index were also evaluated for a better comprehension of energy allocation/partitioning. Estimates based on long‐term measurements, visual observation, wind statistics and wave growth laws allowed an evaluation of the forces acting on shells. At the open‐sea site, the observed phenotypic variability of both shell shape and thickness was clearly related to the tidal vertical zonation. At the two lagoon sites, the currents generated by tidal flow through the inlet and the waves caused by the frequent passage of boats influenced both shell shape and thickness. A trade‐off between protection and growth was apparent along the tide gradient, as emphasized by the differences in the partitioning and allocation of energy between shell and flesh production.     

Benthic studies in upper Buzzards Bay,Massachusetts: 2011/12 as compared to 1955

Precise biodiversity mapping that can be compared over decades would greatly inform the discussion of climate change impacts on species richness and shifts in dominant benthic species. Buzzards Bay, Massachusetts, USA, has a long history of field studies documenting benthic community structure. This bioassessment compares 1955 benthic infaunal sampling results with 2011/2012 samples to determine whether there has been a compositional change. I also compared structural components including grain size and near‐bottom temperatures. In addition, benthic temperature rise and fall was logged at 15‐min intervals from May to October of consecutive years. There was an observed difference in the benthic compositional structure, substrate and temperature between the 1955 and 2011/2012 station conditions. The 1955 identification of Buzzards Bay as being nutrient poor and having a reported 20.5 °C benthic temperature maximum has changed. Coastal development has brought increased nutrient inputs, and the 2012 bottom temperature maximum was at least 4 °C greater. These conditions contribute to shifts in benthic species dominance, and the substrate itself has been transformed to shell reefs at the stations where the gastropod Crepidula fornicata proliferated.     

Changes in survivorship,behavior, and morphology in native soft‐shell clams induced by invasive green crab predators

Many studies on invasive species show reduced native densities, but few studies measure trait‐mediated effects as mechanisms for changes in native growth rates and population dynamics. Where native prey face invasive predators, mechanisms for phenotypic change include selective predation, or induced behavioral or morphological plasticity. Invasive green crabs, Carcinus maenas, have contributed to declines in native soft‐shell clams, Mya arenaria, in coastal New England, USA. We tested the hypothesis that clam ability to detect chemical cues from predators or damaged conspecifics would induce greater burrowing depth as a refuge from invasive crabs, and greater burrowing would require increased siphon growth. To determine how crab predation affected clam survivorship and phenotypic traits in the field, clams in exclosure, open, and crab enclosure plots were compared. Crab predation reduced clam density, and surviving clams were deeper and larger, with longer siphons. To determine whether the mechanism for these results was selective predation or induced plasticity, phenotypes were compared between clams exposed to chemical cues from crab predation and clams exposed to seawater in laboratory and field experiments. In response to crab predation cues, clams burrowed deeper, with longer siphons and greater siphon mass. Overall, crab predation removed clams with shorter siphons at shallow depths, and crab predation cues induced greater burrowing depths and longer siphons. Longer siphons and greater siphon mass of deeper clams suggests clams may allocate energy to siphon growth in response to crabs. By determining native behavior and morphological changes in response to an invasive predator, this study adds to our understanding of mechanisms for invasive impacts and illustrates the utility of measuring trait‐mediated effects to investigate predator–prey dynamics.     

皱纹盘鲍(Haliotis discus hannai)不同群体贝壳形态和相对生长变异研究

皱纹盘鲍是我国重要的渔业资源,其地理群体也是我国重要的鲍种质资源来源。不同地理群体间形态及相对生长的差异对皱纹盘鲍资源管理和种质资源鉴定等提供重要参考。以我国自然海区采集的7个皱纹盘鲍群体作为研究材料,通过测定9个形态测量指标并定义了这些测量指标间的17个比率变量,分析了群体内和群体间贝壳形态和体重相对生长的变异。除个别变量外,协方差分析揭示形态比率变量在群体间存在显著变异;异速生长研究揭示了壳形态特征与皱纹盘鲍群体中异速生长差异之间的高度线性相关性,异速生长程度呈现一定的群体区别。非度量多维尺度分析(NMDS)和马氏距离聚类分析(CLUSTER)将皱纹盘鲍群体分为2个组,线性判别分析(LDA)与NMDS和CLUSTER结果相符,然而LDA未能较好的将各种群样本分类。研究结果初步表明皱纹盘鲍表现出与我国沿海环境和/或生态条件差异相关的表型可塑性,然而这种表型差异也可能受到人工底播苗种的遗传影响。研究结果将对于皱纹盘鲍渔业资源管理如管理单位的界定,以及为今后全面评价皱纹盘鲍不同地理群体的遗传变异等提供参考。     

Induction of female‐to‐male sex change in a large protogynous fish,Choerodon schoenleinii

Most behavioral studies on hermaphroditic fishes have focused on small‐sized species, which are tractable for research. Although many species of large hermaphroditic fishes are important fishery resources, their proximate mechanisms (visual, chemical and/or behavioral cues) in the social regulation of sex change have not been determined. Determination of these would inform resource management and aquaculture. In order to get closer to understanding the proximate mechanisms underlying the social regulation of female‐to‐male sex change in large hermaphroditic fishes, this study reports situations that induced female‐to‐male sex change in black‐spot tuskfish, Choerodon schoenleinii, a species of large protogynous fish, in massive laboratory tanks. The situations differed in the possibility of male‐to‐female tactile contact and in the group sex ratio, enabling us to infer plausible proximate mechanisms underlying sex change induction. Tactile contact between individuals is suspected to be closely related to the incidence of female‐to‐male sex change in C. schoenleinii. Visual and chemical cues alone may be insufficient to inhibit such sex changes. Male‐to‐female tactile contact may have an important influence on female‐to‐male sex change, i.e., inhibition of this sex change, in this species. The effect of sex ratio of a social group on the incidence of sex change may be due to the relative frequency and intensity of male tactile contact with each female, which may vary with the number of females. In the absence of a dominant male, tactile contact among females may affect the incidence of sex change, as well as determine which individuals change sex.     

The first use of Fulton's K for assessing and comparing the conditions of inter‐tidal fish populations

Fulton's K condition factor was applied, for the first time, to inter‐tidal specimens of the shanny (Lipophrys pholis) and long‐spined scorpion fish (Taurulus bubalis) from two English rocky shore and two Welsh rocky shore sites during summer 2010 and winter 2011. As both species contribute to the diet of commercial species such as cod (Gadus morhua) and near‐threatened species such as the European otter (Lutra lutra), their condition may affect that of these predators. Fulton's K found that inter‐tidal Welsh fish maintained a ‘good’ condition between seasons, whereas the inter‐tidal English fish were in a poorer condition during winter. Although condition also changed amongst the sites on each coast, further studies are needed into fish morphologies, environmental parameters, prey availabilities and abundances, and fish specimen sex and maturities.     

Sexual traits plasticity of the potentially invasive limpet Bostrycapulus odites (Gastropoda: Calyptraeidae) within its natural distribution in South America

The slipper limpet Bostrycapulus odites has recently been reported as an introduced species on the Spanish Mediterranean coast. Another species in the family Calyptraeidae, Crepidula fornicata, is a problematic invasive species in the Northeast Atlantic, which makes the closely related Bostrycapulus introduction a matter of concern. Information on the biology of B. odites in its natural environment can facilitate predictions on where the species could find favourable conditions for settlement and expansion. Size and sex were recorded for individuals in four populations over 2600 km of the Southwestern Atlantic shoreline, in Argentina and Brazil, covering the whole native latitudinal range of this widely distributed species. Animals were collected by hand in the intertidal or by SCUBA in subtidal locations; specimens were examined to determine shell length, sex, and the presence and number of brooded egg capsules in females. Mean shell length for the whole population did not differ significantly among populations. However, this species experiences sequential hermaphroditism (protandrism), and the size at sexual maturity (minimum male size) increased significantly with latitude. The fecundity of females at all sites increased with size, and the number of brooded egg capsules as well as the average size of capsules was larger in bigger individual females; however, mean fecundity varied among sites independent of mean female body size. This limpet species modifies its reproductive traits with local conditions, and sexual characters develop earlier in more temperate localities.     

Latitudinal and bathymetric trends in egg size variation: a new look at Thorson's and Rass's rules

The inverse relationship between egg size in marine animals and water temperature was simultaneously described by two outstanding marine scientists: G. Thorson and T.S. Rass. This rule consists of two different phenomena. Thorson's rule describes ecological processes related to changes in larval biology and morphology that are caused by a selective pressure of natural selection on the different types of larval development. It belongs to the realm of macro‐evolution. Rass's rule describes physiological processes within populations and species, and between closely related species. This is not related to changes in reproductive strategy, and therefore belongs to the realm of micro‐evolution and to the early stages of macro‐evolution. Populations begin to produce larger eggs in colder environments because of phenotypic plasticity. Thorson's rule describes temperature‐dependent changes in the relative abundance of small‐ and large‐egged species, whereas Rass's rule describes a temperature‐dependent relative position of both groups within an adaptive range of reproductive strategies.     

Spatial and temporal variation of diverse inter‐tidal algal assemblages in Southwest O‘ahu

Spatial and temporal variation in tropical inter‐tidal communities is poorly known, making predictions about the effects of climate change and other anthropogenic disturbances difficult. Along Southwest O‘ahu, Hawai'i, local residents are concerned about the environmental effects of coastal development and the perceived loss of targeted algal species, which are collected for human consumption. To describe the coastal benthic community and better understand the processes that form and maintain it, the abundance and composition of macroalgae were sampled in the region's inter‐tidal zone from 2006 to 2015. Sixty‐six macroalgal species and two broad algal assemblages were identified that corresponded to substrate topography and sand influence at a similar tidal elevation. Along flat carbonate benches with a sand beach, Phaeophyceae and Rhodophytes occurred in almost equal proportions, while shores with slightly more topographic relief and angular substrate were dominated by Rhodophytes. Foliose or turf algal forms were most common. Surveys captured the local invasion of an alga, Avrainvillea sp. and significant declines in abundant macroalgae in 2015 after a period of unseasonably warm, calm water. Temporal changes in algal assemblages were related to maximum water temperature and wave height but not precipitation. Thus, algal assemblages appear to be structured by local beach morphology as they interact with sand and wave activity and episodically by unusual weather events. However, manipulation and continuous monitoring of the algal assemblages coupled to sensing of the localized environment is necessary to confirm factors related to assembly maintenance and recent species shifts.     

Interactive effects of pH and temperature on native and alien mussels from the west coast of South Africa

Global warming and ocean acidification influence marine calcifying organisms, particularly those with external shells. Among these, mussels may compensate for environmental changes by phenotypic plasticity, but this may entail trade-offs between shell deposition, growth and reproduction. We assessed main and interactive effects of pH and temperature on four mussel species on the west coast of South Africa (33°48′ S, 18°27′ E) in October 2012 by comparing shell dissolution, shell growth, shell breaking force and condition index of two native species, the ribbed mussel Aulacomya atra and the black mussel Choromytilus meridionalis, and two aliens, the Mediterranean mussel Mytilus galloprovincialis and the bisexual mussel Semimytilus algosus. Live mussels and dead shells were exposed for 42 days to seawater of pH 7.5 or 8.0, at 14 °C or 20 °C. Low pH, high temperature and their combination increased shell dissolution of the two aliens but their growth rates and condition indices remained unchanged. Aulacomya atra also experienced greater shell dissolution at a low pH and high temperature, but grew faster in low-pH treatments. For C. meridionalis, shell dissolution was unaffected by pH or temperature; it also grew faster in low-pH treatments, but had a lower condition index in the higher temperature treatment. Shell strength was not determined by thickness alone. In most respects, all four species proved to be robust to short-term reduction of pH and elevation of temperature, but the native species compensated for greater shell dissolution at low pH by increasing growth rate, whereas the aliens did not, so their invasive success cannot be ascribed to benefits accruing from climate change.     

Limpets as prey: Lipophrys pholis choice for size

Patellid limpets are key species on rocky shores, structuring intertidal assemblages through their grazing. Their role as prey for fish is, however, often overlooked in studies of intertidal ecosystem functioning. The shanny Lipophrys pholis is a common predator of limpets on rocky shores in Northern Europe and, in this study, we examined the characteristics of its predation on limpets based on prey size. The limpet size range that adult L. pholis (10–12.5 cm) can eat was examined in the laboratory using Patella depressa in 0.2 cm classes, varying between 0.5 cm and 3.0 cm in maximum shell length. There was a limpet size refuge above 1.8 cm, while all smaller sizes were readily consumed by the shanny. The predator attacking behaviour was also examined and found to vary with prey size. Limpets up to 0.8 cm were crushed by the jaw‐constricting force and eaten whole, whereas larger prey were prised from the substratum and the body subsequently separated from the shell. To examine whether there was a L. pholis preference for P. depressa size, a two‐stage laboratory experiment was done using two size classes defined as small (0.7–0.8 cm maximum shell length) and large (1.5–1.6 cm maximum shell length). In the first stage, the predator was given each limpet size class separately. In the second stage, the fish was given a choice between the two classes. Lipophrys pholis was shown to have a preference for the large size class (1.5–1.6 cm). The average number of limpets consumed by the shanny was examined for the duration of one high‐tide typical of the central region of Portugal (≈ 4 h). On average, approximately five limpets (up to an eight limpet maximum) were consumed. The present study shows that L. pholis has a feeding preference based on limpet size, which suggests that the limpet population structure and intra‐ and inter‐specific interactions may be influenced by the shanny predation.     

An approach to ecological monitoring in the rocky intertidal: A survey of Bantry Bay and Dunmanus Bay,Ireland

The methodology of a monitoring study of the fauna and flora of the rocky intertidal zone in Bantry Bay and Dunmanus Bay in southwest Ireland is described and some representative results are presented. Five shores, four in Bantry and o one in Dunmanus Bay, were visited monthly from May, 1978 to June, 1980, and more limited monitoring is continuing. On each visit to each shore, four research programmes were undertaken—transect studies, recolonisation of cleared areas, barnacle studies and limpet studies. The transect methods are discussed in relation to other methods in current use. Combining community and key species approaches is shown to be most useful in monitoring programmes. The increased effectiveness of continuous shore monitoring when compared with one-off or before and after impact-type studies is stressed.     

Overview of West African fisheries under climate change: Impacts,vulnerabilities and adaptive responses of the artisanal and industrial sectors

Climate change affects ocean conditions, fish stocks and hence fisheries. In West Africa, climate change impacts on fisheries were projected to be mainly negative through multi-facet ways. However, analysis of adaptation responses of fishers to climate change impacts is scarce. This paper reviews the impacts on climate change on fisheries in West Africa and discusses the potential adaptation strategies adopted by both the artisanal and industrial fishing sectors. Overall, climate change and over-exploitation have altered species composition of fisheries catches in West Africa. The effect of ocean warming on fisheries is indicated by the increase in dominance of warmer water species in the landings, shown from an increase in Mean Temperature of Catch, in the region. Climate change induced changes in potential catch and species composition, which inherently have similar symptoms as over-exploitation, are expected to have repercussions on the economic and social performance of fisheries. Both artisanal and industrial sectors may adapt to these changes mainly through expansion of fishing ground that increases operation costs. Our results highlight that historical changes in target species are more common in industrial than artisanal fisheries. This result challenges the prevailing assumption that artisanal fisheries, given their limited movement capacity, would adapt to climate change by shifting target species and/or gear type.     

Aspects of the ecology of a littoral chiton,Sypharochiton pellisekpentis (Mollusca: Polyplacophora)

On several Auckland shores, a littoral chiton, Sypharochiton pelliserpentis (Quoy and Gaimard, 1835), was widely distributed and common. At Castor Bay it was the commonest chiton, and its density equalled or exceeded that of the commonest limpet (Cellana spp.) over most of the inter‐tidal range. Spot measurements of population density were made at other sites including exposed and sheltered shores. The smallest animals were restricted to the lower shore in pools or on areas of rock which were slow to drain. Exclusive of these small animals, the population structure was similar in pools and water‐filled crevices situated either high or low on the shore. This was also true for populations of animals from drained surfaces, but, at both shore levels, the population structure in the two micro‐habitats (pools and drained surfaces) was distinct.

Single measurements of salinity in the very small volumes of water in which chitons were living gave a range of 13.3–45.8‰. In laboratory experiments at normal temperatures this species could survive these fluctuations for the maximum time possible between tides. Desiccation experiments, in which weight loss was expressed as a percentage of total water content, showed weight loss to depend on size. No significant differences in weight loss were found between populations from the two shore levels and the two micro‐habitats considered, though there were slight survival differences. Desiccation is probably an important factor influencing population structure, but loss of up to 75% of water content could be tolerated before 50% mortality occurred. Osmotic and desiccation stress evoked a characteristic behaviour pattern.     

笠藤壶的形态变异

本文研究了舟山沿海两种笠藤壶（日本笠藤壶Ｔｅｔｒａｃｌｉｔａｊａｐｏｎｉｃａ、鳞笠藤壶Ｔ．ｓｑｕａｍｏｓａｓｑｕａｍｏｓａ）个体的形态变异,分析了壳板形态与生境、潮区、年龄之间的关系以及壳板形态的种间差别。壳口面积与基底面积之比值（Ｓａ／ＳＡ）在各年龄组间变化较小,而壳口面积与个体容积之比值（Ｓａ／Ｖ）随着藤壶年龄增加而减小,藤壶个体亦随年龄增加由矮圆锥形变陡。盖板各测值之比值在不同年龄组间变化不大,受环境影响较小,可以作为分类鉴定的依据。