Habitat-related differences in the pre-copulatory guarding and copulation behavior of Neohelice granulata (Brachyura,Grapsoidea, Varunidae)

Mating behavior strategies of brachyuran crabs are flexible depending on the habitat and social context. We evaluate pre-copulatory guarding in Neohelice granulata and describe the unusual copulatory behavior. Laboratory experiments were conducted to test whether factors such as male size, presence of burrows, a male-biased operational sex ratio (OSR) and female mating history affect the copulation duration in two different study areas. Females initiated a searching behavior approximately 20–26 h before they became receptive by assessing large male burrows and displaying a courtship behavior (“flirting”) in its entrance. Once the female entered inside the male burrow, pre-copulatory guarding was observed until the female became receptive and successful copulation occurred immediately after. This species shows some very unusual copulatory behavior involving eyestalk grasping not seen in any other varunid crab. The copulation duration was independent of male size and the presence of burrows, although it was dependent on the study area, the OSR and the female mating history. Thus, copulation duration is a flexible character in this species which varies with habitat and biological factors such as mating history and sex ratio.     

Different sperm allocation strategies in two populations of the semiterrestrial crab Neohelice granulata (Brachyura,Grapsoidea, Varunidae)

Sperm reserves are costly to produce and so the development of different sperm allocation strategies may be advantageous. Knowledge of sperm allocation strategies is important from an evolutionary perspective because they contribute to characterize mating systems. However, population differences in sperm allocation strategies of the same species have rarely been assessed. Here, we studied the male sperm allocation strategies in two different populations of Neohelice granulata in Mar Chiquita Lagoon (MCL) and San Antonio Oeste (SAO), Argentina. We found that the quantity of ejaculate transferred by N. granulata males depended on different factors such as male size and female receptivity duration, while it was independent of female size, at both study areas. Regarding the operational sex ratio factor, the amount of ejaculate transferred depended on it in MCL but not in SAO. Thus, N. granulata males may have the capacity to regulate the quantity of ejaculate transferred based on individual factors such as their own size and female receptivity duration, and population factors such as operational sex ratio, which varies in the two populations studied.     

Burrowing behaviour of the paddle crab Ovalipes catharus (White, 1843) (Brachyura: Portunidae)

The burrowing behaviour of Ovalipes catharus (White, 1843) is described and compared with that of some other burrowing crabs. O. catharus is a fast, back burrowing crab which makes temporary burrows in soft sand. The average time from initiation of burrowing to complete submergence was 6.5 s, there was no difference between males and females but burrowing time increased significantly with crab size. Similar morphological adaptations are required for burrowing and swimming: streamlined body shape, smooth lightly constructed exoskeleton, and flattened legs. O. catharus has a reversed respiratory current like O. guadulpensis but it differs in its burrowing behaviour. Compared to some other crabs, burrowing in O. catharus is rapid.     

Population genetics of cold‐water coral associated Pleustidae (Crustacea,Amphipoda) reveals cryptic diversity and recent expansion off Iceland

Amphipoda are a diverse and important faunal component of deep‐sea habitats worldwide. However, relatively little is known about species that are associated with cold‐water corals, in particular concerning their genetic diversity, population genetic processes, population differentiation, and potential host specificity. We exemplarily investigate these questions studying three pleustid species—Stenopleustes aff. malmgreni (Boeck, 1871), Stenopleustes aff. nodifera (Sars, 1883) and Neopleustes aff. boecki (Hansen, 1888)—that are associated with cold‐water corals and polychaete assemblages along the Reykjanes Ridge southwest of Iceland. We sequenced mitochondrial COI and performed ddRAD to study thousands of nuclear loci. These species and a pair of morphologically cryptic species within Stenopleustes aff. nodifera were consistently delimited by COI and ddRAD. We conclude that all of our studied species are new to science, raising the number of pleustids reported from cold‐water corals to six. Two species were collected only once, and these species exhibited high levels of inbreeding, suggesting little exchange with other populations, which might be a result of strong host specificity. The other two species were wider distributed, with very low differentiation among populations, even between populations separated by 130 km. These species showed signs of recent population expansions, possibly in relation to changes in the cold‐water coral associated ecosystem (e.g., due to glacial cycles and/or volcanic or geothermal activities). They had lower levels of inbreeding and were each associated with three different host coral species, which might facilitate dispersal and gene flow. Our results show divergent population genetic patterns for the studied pleustid species. These might be attributed to the species’ host specificity and highlight important differences among species with seemingly similar life‐styles. Species diversities appear to be greatly underestimated, limiting our abilities to truly assess the vulnerability of the fauna living in these threatened cold‐water coral habitats.     

Sex‐specific affinity for redox‐active metals influences antioxidant responses of Callinectes amnicola (blue crab) populations in littoral and open water habitats of a tropical coastal lagoon

The critical role of ecological preferences and opportunity in determining contaminant uptake and adaptive responses of sexes in the wild is still poorly understood. This ecological relationship was investigated by measuring metal bioaccumulation and antioxidant activity in male and female blue crab populations from open water habitat and the littoral/inter‐tidal zone of the Lagos Lagoon. A total of 741 samples of blue crab (littoral zone: 263 females, 137 males; open water zone, 230 females, 111 males) was collected monthly over 24 months (January 2010–January 2012) from each site and the measurements of morphometric features (wet weight, carapace length, carapace width) were recorded; condition index, metal (redox active: Cu, Zn, redox inactive: Pb, Cd) concentration in tissues (gills, hepatopancreas, gonads and muscle) and antioxidant activity (superoxide dismutase, reduced glutathione, glutathione peroxidase, catalase and malondialdehyde) were measured for each sex. Monthly sediment samples for both habitats were also analysed for metals using standard methods. Female crabs were significantly larger (p < .05) with a better condition index than the male crabs across sites and seasons, while higher oxidative damage was recorded in male crabs in the littoral zone compared to the open water zone. The results show that there was a negative association between antioxidant activity and lipid peroxidation; a negative relationship between concentrations of redox‐inactive metals (Pb and Cd) and antioxidant activity in male crab tissues; and a positive relationship between uptake of a redox‐active metal (Cu) and antioxidant activity in female crab tissues. Although these trends suggest sex‐specific toxicity, they also associate redox‐inactive metals with the downregulation of antioxidant activity and oxidative stress. Furthermore, the higher condition index of females corroborates the possibility of sex‐specific toxicity, while the larger‐sized females compared to males suggests size‐sexual dimorphism in the blue crab populations. The site‐specific oxidative damage between sexes may be attributed to the different complexity of both habitats, which affords different ecological opportunities for the sexes.     

Microsatellites reveal fine‐scale genetic structure of the Chinese surf clam Mactra chinensis (Mollusca,Bivalvia, Mactridae) in Northern China

As a commercially important fisheries resource in East Asia, the Chinese surf clam Mactra chinensis has experienced severe population declines in the past decades, probably due to over‐exploitation. To provide scientific bases for fisheries management and artificial breeding, we investigated genetic variation and population structure of Mactra chinensis in Northern China using microsatellites. Samples collected from eight localities throughout natural habitats of the species in Northern China were genotyped. Nine microsatellites revealed high allelic diversity with 14–36 alleles per locus. Observed and expected heterozygosity varied from 0.593 to 0.945 and from 0.638 to 0.958, respectively. Pairwise F_ST values indicated that all population pairs had small but significant genetic differentiation. A Mantel test showed statistically significant correlations between genetic distance and geographical distance, indicating that genetic differentiation of the Chinese surf clam conformed to a pattern of isolation‐by‐distance. Cluster analysis using neighbor‐joining separated the eight populations into three groups. The three areas of low gene flow identified by barrier analysis corresponded with local oceanographic features, suggesting that marine currents and peninsulas play an important role in population structuring of this species.     

Small macro‐benthic invertebrates influence the survival,growth and behaviour of juvenile green sea urchins Strongylocentrotus droebachiensis in the laboratory

Recent settlers of many marine benthic invertebrates are cryptic, which exposes them to a suite of animals that differs from those they may experience as adults, potentially resulting in interactions causing mortality and/or reducing growth. Previous field experiments have indicated that such is the case with small juvenile green sea urchins Strongylocentrotus droebachiensis but which taxa are responsible for the mortality and reduced growth was not determined. A laboratory study was conducted to examine the effects of small macro‐benthic invertebrates, specifically chitons, scaleworms and larger juvenile conspecifics, as well as the full suite of cobble‐dwelling organisms, on the mortality, growth and behaviour of small (<3 mm) juvenile sea urchins. The likelihood of survival of small juvenile sea urchins was lower in the presence of larger juvenile sea urchins or with the full suite of cobble‐dwelling organisms than in the absence of animals. The small juvenile sea urchins survived and grew the best when they were with chitons and scaleworms. The behaviour of small sea urchins with the full suite of cobble‐dwelling organisms was more cryptic than the behaviour of urchins with scaleworms. This study indicates that interactions with the suite of small organisms living amongst cobbles can affect survival, growth and behaviour of small juvenile sea urchins, and that larger juvenile sea urchins can be a source of mortality for smaller conspecifics.     

Spatio‐temporal variation in biomass of the deep‐sea red crab Chaceon affinis in Gran Canaria Island (Canary Islands,Eastern‐Central Atlantic)

The spatial and temporal biomass distribution of Chaceon affinis and its vulnerability to fishing activity in Gran Canaria (Canary Islands) were investigated. The first goal was to assess the influence of the slope steepness and substrate on the size of crab patches, size of the crabs, and crab biomass. The second goal was to evaluate spatial and temporal variation in the biomass over a 15‐month period. The last goal was to assess the influence of fishing activity upon the reduction in the biomass over the same 15‐month period. Only two or three locations in the sampling area generated high‐biomass contour patches. When these patches were superimposed on the isobath lines, they were coincident with the main depth range described for the species in the area. The map of the biomass values clearly showed three structures with cores of the highest biomass in both muddy and rocky‐muddy areas. The biomass was higher on muddy than on rocky‐muddy bottoms. Biomass was twice as high when steepness was reduced to one third between isobaths of 500 and 900 m. The size of crab patches increases linearly with the decrease in slope steepness. The spatial structure of crabs remained fairly stable over time, showing that biomass changes with depth over time. Maps of the estimated biomass values over the 15‐month period showed the same two main patches over time with the cores of highest biomass separated by a distance of between 4.2 and 4.5 km. Although the bathymetric distribution by sexes showed temporal changes, with a displacement to deeper areas made by both sexes over the studied period, only a partial temporal segregation between males and females was observed. During the study period, crabs underwent a significant decline in biomass and this was consistent with the combined catches of both commercial and experimental fishing in the area. Due to its low mobility, C. affinis is highly vulnerable to local depletion by intensive fishing efforts.     

Differentiation in reproductive traits of geminate mangrove crabs of the genus Aratus (Decapoda: Brachyura) across the Central American Isthmus

The mangrove crab Aratus pisonii was considered to have an amphi‐American distribution; however, a recent study revealed that the Eastern Tropical Pacific populations were genetically distinct, thus representing a new species: Aratus pacificus. These species separated by the Isthmus of Panama have diverged under different environmental conditions that may have influenced their reproductive biology. As the available information about this genus concerns almost exclusively the Caribbean species, the aim of the present study was to analyse and compare reproductive aspects of A. pacificus and A. pisonii obtained from both the Pacific and Caribbean coasts of Costa Rica. Females were collected from April 2011 to April 2012, and reproductive features such as breeding season, size distribution of ovigerous females, fecundity, reproductive output, embryo volume and embryo water content were assessed. Both species produced embryos during the entire sampled period. Most females of A. pacificus carrying embryos close to hatching were found during the rainy season. Ovigerous females of A. pisonii were substantially larger and reached sexual maturity at a larger size than females of A. pacificus. Embryo production started in A. pacificus at a smaller female size than in A. pisonii. As fecundity increased with female size, the average fecundity was lower in A. pacificus. Females of A. pisonii produced larger embryos, which might be related to lower food availability: higher energy content in the embryo enhances the chances of larval survival. These data regarding reproductive features of the Pacific and Caribbean species support the conclusion to separate A. pacificus from A. pisonii.     

Midlittoral polychaete communities in the Eastern Mediterranean Sea: new information from the implementation of the Natural Geography in Shore Areas (NaGISA) protocol and comparisons at local and regional scales

The aim of this study was to attempt to fill current knowledge gaps on midlittoral Mediterranean biodiversity at local and regional scales, by using benthic polychaetes as a model taxon. Two different data sets were analysed: (i) a quantitative data set from the two Natural Geography in Shore Areas (NaGISA) study sites in Crete and (ii) a qualitative data set from multiple sites across the Mediterranean. At the local scale, the results provide evidence that (i) discrete species communities are formed in midlittoral Mediterranean habitats, which vary by geographical location and year, depending on the scale of observation; (ii) macrophyte coverage and Chl‐a are the only environmental variables associated, albeit weakly, with the above pattern; (iii) although naturally disturbed, the Cretan NaGISA sites do not seem to experience any anthropogenic stress; (iv) environmental heterogeneity and history seem to be much less important in shaping the polychaete communities than inter‐specific interactions; however, it is not possible to specify at this stage whether local or regional processes or even their interactions may shape the polychaete communities. At the regional scale, the results indicate that (i) the only factor that seems to be involved in the regional pattern is the identity of the study providing the data sets, which implies variability and bias in how research projects are carried out, from the sampling design through to data collection and analysis; (ii) the Cretan NaGISA sites may be considered as representative of the habitat in the Mediterranean, under certain conditions.     

The distribution of meiofauna on back‐reef sandy platforms in the Maldives (Indian Ocean)

The Maldivian archipelago comprises some of the most characteristic and significant world atoll systems, but the meiobenthic assemblages of these islands continue to be largely unknown. To investigate variations in meiofaunal spatial distribution and biodiversity in back‐reef platforms, three transects were studied, two at Felidhoo (the north and east sides) and one at South Malé. The sedimentological features of the samples obtained were also analyzed to further current knowledge on the relationships that exist between sediments and meiofauna. Our results reveal that the meiofaunal assemblage at these locations is well diversified and includes 20 major taxa. Nematodes and copepods are dominant, together forming 68% of the total meiofauna, followed by platyhelminthes, polychaetes and ostracods. The nematode assemblage is very rich and composed of 34 families and 123 genera – 96 of which (78%) set new records for the Maldives. The structures of the meiofaunal and nematode assemblages are relatively similar on the ‘large‐scale’ level (i.e. when the different platforms are compared) and reveal a low β‐diversity. However, significant dissimilarities were detected within each platform, emphasizing that such ‘small‐scale’ differences are the main factors determining the structure of the meiofauna and, in particular, the nematode assemblages. Although significant differences were not detected between the transects, greater levels of dissimilarity were recognized at North Felidhoo. Here, the building of inclined deposit layers plays a significant role in increasing the heterogeneity of the platform habitats and sediments, confirming the great importance of sediment granulometry as an environmental variable. Indeed, a close relationship is observed between meiofauna (especially for the nematodes) and grain size, which appears to control the structure, diversity and trophic composition of the Maldivian meiofauna assemblages, thus highlighting the high biodiversity existing in the medium‐coarse sands.     

Long‐term comparison of structure and dynamics of the red coral metapopulation of the Portofino Promontory (Ligurian Sea): a case‐study for a Marine Protected Area in the Mediterranean Sea

A large amount of data on the precious coral Corallium rubrum has attested to a dramatic shift in the size structure of its over‐harvested Mediterranean populations in recent decades. One of the main problems for the conservation of this species is the substantial lack of data concerning the time span necessary for a population subjected to pluri‐decennial harvesting pressure to return to its pristine status. Here, we present a multi‐annual data set gathered from the Marine Protected Area (MPA) of Portofino, which hosts the most important shallow‐water coral populations in the Ligurian Sea and was subjected to strong harvesting pressure from the 1950s to the early 1970s. Quantitative comparison of the population structure data recorded in 1964, 1990 and 2012 indicated a strong size increase of the colonies (from 3 to 8 g mean weight), resulting in an increase in colony biomass from 300 to the current 1500 g·m^?2. As a consequence, over the same span of time, the density of colonies has slightly decreased. The role of mass mortality phenomena, like that occurred in this area during 1999, as possible biological features triggering the switch of red coral populations from a ‘grass plain‐like’ towards a ‘forest‐like’ structure, is discussed. All these data indicate that the instigation of MPAs is a winning strategy in the conservation of this precious species and similar management plans should also be evaluated for the protection of the deep benthic communities thriving on off‐shore banks.     

Life Cycle of Liocarcinus arcuatus (Brachyura: Portunidae) in the Ría de Arousa (Galicia, NW Spain): Role of Beach and Mussel Raft Culture Areas

Abstract. The population biology of the portunid crab Liocarcinus arcuatus is analyzed over a yearly cycle in beach zones and mussel raft culture areas in the Ria de Arousa.
The breeding cycle of this species has two annual peaks, which result in two annual cohorts that are recruited in different areas. The growth rate is higher in the group settled in spring than in autumn, and, especially, in the culture areas compared with the beach zones. There are movements from the beach zones to the raft polygons related to reproductive and feeding behavior as well as growth. The role of mussel culture in the dynamics of this species is discussed.     

Patterns of genetic and morphometric diversity in the marbled crab (Pachygrapsus marmoratus, Brachyura, Grapsidae) populations across the Tunisian coast

The present study reports on population structure analysis of the marbled crabPachygrapsus marmoratus (Fabricius, 1787) from the Tunisian coast, an appropriate location to study biogeographical process...     

Depth of settlement of two mussel species on suspended collectors in Marlborough Sounds,New Zealand (Note)

The preferred depth of settlement and settlement period of two species of mussel spat were monitored at Yncyca Bay (NZMS1 S15 220460) and Wet Inlet (NZMS1 S16 265432) in the Marlborough Sounds on collectors suspended in 0–4 m of water between 20 October 1975 and 4 April 1976. Peak settlement of Mytilus edulis aoteanus was between 20 October 1975 and 4 January 1976, and decreased with increasing depth. Peak settlement of Perna canaliculus was between 16 February 1976 and 12 April 1976, and showed no preferred depth of settlement.

Capture of wild stocks of Mytilus edulis aoteanus and Perna canaliculus spat for use in aquaculture has become more reliable through accurate forecasting of peak settlement periods, and knowledge of preferred depth of settlement of the two species.

Preferred depth of settlement of spat may affect the shore zonation of mussels.     

Comparison of the biology, dynamics, and secondary production of Talorchestia brito (Amphipoda, Talitridae) in Atlantic (Portugal) and Mediterranean (Tunisia) populations

The biology, population dynamics, and production of Talorchestia brito were studied at two sandy beaches located on the Atlantic (Portugal) and on the Mediterranean (Tunisia) coasts, respectively. The seasonal variation in abundance and the overall densities were similar in both populations. Reproduction occurred from February to September in the Atlantic, and from March to early November in the Mediterranean. The sex ratio was male biased in the Atlantic, and female biased in the Mediterranean. Based on data from the Atlantic population, both abundance and the proportion of reproductive females were positively correlated with temperature, while the proportion of juveniles in the population was positively correlated with temperature and sediment moisture. On average, individuals from the Atlantic were larger than the ones from the Mediterranean. Life span was estimated at six to nine months in the Atlantic, and five to eight months in the Mediterranean. Talorchestia brito was shown to be a semiannual species, with iteroparous females producing two broods per year, and exhibited a bivoltine life cycle. The minimum age required for males' and females' sexual differentiation and for female sexual maturation was shorter in the Mediterranean. Growth production (P) was estimated at 0.19 g m⁻² y⁻¹ ash free dry weight (AFDW; 4.3 kJ m⁻² y⁻¹) in the Atlantic population, and 0.217 g m⁻² y⁻¹ AFDW (4.9 kJ m⁻² y⁻¹) in the Mediterranean one. Elimination production (E) was estimated at 0.35 g m⁻² y⁻¹ AFDW (7.9 kJ m⁻² y⁻¹) in the Atlantic, and 0.28 g m⁻² y⁻¹ AFDW (6.3 kJ m⁻² y⁻¹) in the Mediterranean. The average annual biomass ( ) (standing stock) was estimated at 0.032 g m⁻² in the Atlantic beach, and 0.029 g m⁻² in the Mediterranean one, resulting, respectively, in ratios of 5.9 and 7.5 and ratios of 10.8 and 9.6. Like other talitrids, T. brito exhibited geographic variation in morphometrical characteristics, sex ratio, growth rates, life span, and reproduction period, with the Atlantic population presenting a slower life history.     

Molecular analysis of the commercial deep-sea crabs Chaceon ramosae and Chaceon notialis (Brachyura,Geryonidae) reveals possible cryptic species in the South Atlantic

Deep-sea red crabs Chaceon ramosae and Chaceon notialis are commercially and ecologically important crustaceans in South Atlantic waters. With some overlapping occurring at the edge of the crabs’ distribution, the fisheries are managed separately in Brazil and Uruguay, with some differences in the regulations, although the stock is supposedly shared between the two countries. To assess genetic information on these two species, we examined the nucleotide sequence of the mitochondrial 16S rDNA and CO1 mtDNA genes in samples from the southern region. We also compared these data to sequences from other congeners. The genetic analysis revealed a clear separation between C. notialis and C. ramosae. In addition, we detected the existence of at least two different groups within the species that is currently recognized as C. notialis. This might indicate the existence of cryptic species, leading us to speculate about the presence of two different taxonomic entities, although further research is needed to resolve this issue.     

Long‐term presence and habitat use of Cuvier's beaked whale (Ziphius cavirostris) in the Central Tyrrhenian Sea

Cuvier's beaked whale (Ziphius cavirostris, G. Cuvier 1823) is a poorly known species and many international agreements have asked for a better understanding of its biology for conservation purposes. In the present study, systematic cetacean surveys were carried out from ferries along a trans‐regional fixed transect in the Central Tyrrhenian Sea (Civitavecchia, Latium – Golfo Aranci, Sardinia), just outside the southeastern border of the Pelagos Sanctuary. This research provided long‐term, consistent data on Cuvier's beaked whale during two research periods (1990–1992 and 2007–2011). The objective of the research was to compare the presence, distribution and habitat use of Cuvier's beaked whale between the two investigated periods. Summer data (June–September) from the two periods were compared in terms of frequency of sightings, group size and spatial distribution related to the main ecogeographical features. A presence‐absence model (generalized additive modelling) was performed to predict habitat suitability in the two study periods. The results highlight long‐term site fidelity of Cuvier's beaked whale in the Central Tyrrhenian Sea with encounter rates comparable to the ones reported for other key areas. Separate suitability models based on 1990s and 2000s data appeared to work for each individual time period but differences were evident between the two periods, indicating changes in habitat selection over time. Our findings of the study appear to expand the definition of suitable beaked whale habitat and underline how the temporal scale of the analysis can affect the results in habitat studies. Moreover, this research highlights the importance of the Central Tyrrhenian Sea marine region for Cuvier's beaked whale and the ability of continuous monitoring to identify changes in cetacean frequency and distribution, necessary for adaptive conservation management approaches.     

The genetic structure of the exotic ascidian Styela plicata (Tunicata) from Italian ports,with a re‐appraisal of its worldwide genetic pattern

The pleated ascidian Styela plicata (Lesueur, 1823) is a solitary species commonly found in ports and marinas around the world. It has been recorded in the Mediterranean region since the mid‐19th century. In the present work, the species’ genetic diversity was analysed, employing a 613‐bp portion of the mitochondrial cytochrome c oxidase subunit I (COI) gene from 149 individuals collected in 14 ports along Italian coasts at spatial scales ranging from 1 to approximately 2200 km. Haplotype and nucleotide diversity values were h = 0–0.933 (total h = 0.789) and π = 0–0.145 (total π = 0.0094), respectively. A general southward trend of increasing within‐population genetic diversity was observed. Analysis of molecular variance revealed significant genetic structuring but no significant differences were detected among basins, and no isolation by distance was found. Our data were integrated with the COI sequences available from previous studies and re‐analysed in order to investigate the possible routes of introduction of this ascidian into the Mediterranean Sea. The presence of the two COI haplogroups detected in previous molecular investigations on S. plicata at intercontinental spatial scale was confirmed in the Mediterranean Sea. The results revealed multiple introductions of S. plicata, although some locations appear to have experienced rapid expansion from few founding individuals with reduced genetic diversity. However, continuous introductions would confound the pattern deriving from single founder events and make it difficult to estimate the time needed for gene diffusion into established populations. This mixing of effects creates difficulties in understanding the past and current dynamics of this introduction, and managing this alien invasive ascidian whose genetic structure is continuously shuffled by vessel‐mediated transport.