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.     

Phytoplankton subsidies to the inter‐tidal zone are strongly affected by surf‐zone hydrodynamics

The size of bottom‐up subsidies, food and larvae, from the ocean has a profound impact on inter‐tidal communities and populations. Alongshore variations in subsidies are attributed to variation in coastal conditions, but also might be due to variations in surf‐zone hydrodynamics. We tested this hypothesis by comparing surf‐zone phytoplankton concentrations to surf‐zone hydrodynamics as indicated by the width of the surf zones. To minimize the potential effects of alongshore variation in phytoplankton abundance in the coastal ocean, we sampled closely spaced sites (median separation 1 km) over short periods (3 days). Surf‐zone concentrations of coastal phytoplankton taxa (e.g. Chaetoceros spp., Pseudo‐nitzschia spp. and dinoflagellates) varied with surf‐zone width; 65% to 94% of the variability in their concentration is explained by surf‐zone width. Where surf zones were narrow and more reflective, phytoplankton concentrations were one to several orders of magnitude lower than in wider, more dissipative surf zones. The most closely spaced stations were 30 m apart, but represented distinct habitats – a reflective and a more dissipative surf zone. Phytoplankton concentrations in the reflective surf zone were an order of magnitude or more lower than in the adjacent more dissipative surf zone. A reanalysis of a published study on phytoplankton subsidies found similar results. Alongshore variation in surf‐zone hydrodynamics appears to be an important driver of phytoplankton subsidies to the inter‐tidal zone.     

Diets of Steller sea lions off the coast of Hokkaido,Japan: An inter‐decadal and geographic comparison

Inter‐decadal and geographic variations in the diets of Steller sea lion, Eumetopias jubatus, were examined based on the contents of 408 stomachs collected from coastal areas around Hokkaido Island during the periods 1994–1998 and 2005–2012. The most important prey species in the 1990s were gadid fishes (walleye pollock [Gadus chalcogrammus], Pacific cod [Gadus microcephalus] and saffron cod [Eleginus gracilis]). The frequency of occurrence and gravimetric contribution of gadids decreased in the 2000s latter period at three study sites (Rausu, Shakotan and Rebun) and were replaced by Okhotsk Atka mackerel (Pleurogrammus azonus) and smooth lumpsucker (Aptocyclus ventricosus). However, analysis based on gravimetric composition indicated that the dietary diversity of prey showed only a slight inter‐decadal difference, reflecting the wide diversity of prey ingested during both study periods. These results indicate that Steller sea lions along the Hokkaido coast are opportunistic feeders that utilize a wide variety of prey, and appear to feed mainly upon prey that is easily obtained.     

Towards a standard measure of sea anemone size: assessing the accuracy and precision of morphological measures for cantilever‐like animals

Our capacity to detect and interrogate patterns in nature depends on the use of standard methods for measuring biological units. Consensus methods to quantify the size of individual animals and characteristics of biological communities are critical for comparisons across time and space. Nowhere is this more important than when dealing with organisms such as sea anemones that display high plasticity in body shape. Despite the need for accurate measures of anemone size for ecological comparisons, there is little consensus on the accuracy and precision of size inferences for these animals. We assessed several morphological parameters to determine which in‐field measure accurately and reliably reflects the reference size of an anemone measured in the laboratory: (i) column height, (ii) column diameter, (iii) limbus diameter, (iv) pedal disc diameter, (v) pedal disc area or (vi) pedal disc perimeter. The results revealed large variability in the accuracy and precision amongst measures, which have implications for their suitability as a standard method for in situ measurements. In general, measures of diameter were preferable to those of height, area and perimeter; and those associated with attachment (i.e. the limbus and pedal disc diameter) performed the best. Overall, considering concurrence with measures obtained from two differing but useful reference states, pedal disc diameter was the most accurate parameter (mean percentage difference = 0.6) with which to estimate the size of sea anemones in the field, and we thus recommend its use as an effective, non‐destructive means of gaining insights into their behavioural and evolutionary ecology.     

Cryptic diversity and phylogeography of the island‐associated barnacle Chthamalus moro in Asia

Pleistocene glaciations were among the important historic events that shaped the population structures of marine organisms. Genetic studies of different marine fauna and flora have demonstrated the effect of Pleistocene glaciations on taxa that reside in marginal seas. However, how marine island species responded to Pleistocene glaciations remains relatively unstudied, especially in Asia. Genetic analyses based on the island‐associated barnacle Chthamalus moro collected from 14 sites in Asia reveal that C. moro comprises three distinct lineages, with COI divergence ranging from 3.9 to 8.3%. Population genetic analyses on respective lineages reveal signs of demographic expansion within the Pleistocene epoch at different times. The Ogasawara lineage, which has a more oceanic distribution, expanded the earliest, followed by the population expansion of the Ryukyu and Southern lineages that inhabit islands closer to the continent. The data suggest that the inhabitants of outer islands may have been less affected by Pleistocene glaciations than those that reside closer to the continent, as the former were able to maintain a large, stable, effective population size throughout the late Pleistocene.     

Habitat modification in tidepools by bioeroding sea urchins and implications for fine‐scale community structure

By creating novel habitats, habitat‐modifying species can alter patterns of diversity and abundance in marine communities. Many sea urchins are important habitat modifiers in tropical and temperate systems. By eroding rocky substrata, urchins can create a mosaic of urchin‐sized cavities or pits separated by exposed, often flat surfaces. These microhabitats appear to harbor distinct assemblages of species. We investigated how a temperate rocky intertidal community uses three small‐scale (<100 cm²) microhabitats created by or adjacent to populations of the purple sea urchin (Strongylocentrotus purpuratus): pits occupied by urchins, unoccupied pits, and adjacent flat spaces. In tidepools, flat spaces harbored the highest percent cover of algae and sessile fauna, followed by empty pits and then occupied pits. The Shannon diversity and richness of these sessile taxa were significantly higher in flat spaces and empty pits than in occupied pits. The composition of these primary space holders in the microhabitats also varied. Unlike primary space holders, mobile fauna exhibited higher diversity and richness in empty pits than in flat spaces and occupied pits, although results were not significant. The protective empty pit microhabitat harbored the highest densities of most trophic functional groups. Herbivores, however, were densest in flat spaces, concordant with high algal coverage. These results suggest the habitats created by S. purpuratus in addition to its biological activities alter community structure at spatial scales finer than those typically considered for sea urchins.     

Inter‐ and intra‐colonial genotypic diversity in hermatypic hydrozoans of the family Milleporidae

Genetic diversity is the basis for adaptation and therefore of primary scientific interest, especially in species that are threatened by anthropogenic challenges, e.g. climate change and/or pollution. Coral reefs are among the most threatened but also the most diverse ecosystems and have therefore been studied quite extensively. So far, most investigations have focused on scleractinian corals while the equally important reef builders, the hydrozoans, have been less considered. Here we provide the first study of genotypic variability as well as intra‐colonial genetic variability, the co‐occurrence of more than one genotype within a single colony, in Milleporidae based on microsatellites. We analysed two geographically distinct populations from the Millepora dichotoma complex, one from the Red Sea and one from the Great Barrier Reef. Additionally, a population of Millepora platyphylla was analysed from French Polynesia. We compared microsatellite multilocus genotypes and cytochrome c oxidase subunit I haplotypes for each of the three field sites to detect levels of genotypic diversity at the intra‐ and inter‐specific levels. Furthermore, we examined all species for the occurrence of intra‐colonial genetic variability, a recently described mechanism in scleractinian corals that might enhance the adaptive potential of sessile organisms. We found both species and all field sites to be genotypically variable. Twelve mitochondrial haplotypes and 27 multilocus microsatellite genotypes were identified. In addition, intra‐colonial genetic variability was detected in the M. dichotoma complex from the Great Barrier Reef as well as in M. platyphylla from French Polynesia. All of the intra‐colonial genetically variable colonies consisted of one main genotype and a second divergent genotype caused by somatic mutations (mosaicism). Our study proves that Milleporidae are genetically variable and that the phenomenon of intra‐colonial genetic variability also occurs in this important reef‐building family.     

Size‐frequency distributions of Joania cordata and Argyrotheca cuneata (Brachiopoda: Megathyrididae) from the Central Tyrrhenian Sea

Size‐frequency distributions can support reliable inferences concerning population dynamics of brachiopods, but only a few data are available so far. In this study, length and width frequency distributions of dead specimens of the Recent brachiopods Joania cordata and Argyrotheca cuneata from the Marine Protected Area ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, Italy (41°35′ N, 12°20′ E), are reported in order to add new data about size‐frequency distributions of brachiopods. The studied specimens came from death assemblages in the coralligenous substrate, in the Posidonia oceanica meadows, and in the sand channels. The observed patterns vary from left‐skewed (J. cordata) to right‐skewed (A. cuneata), indicating respectively a low and high mortality of smaller individuals. Significant differences between the coralligenous substrate and the P. oceanica meadow were observed for both species, revealing a variation among different habitats. All length and width distributions are clearly polymodal, but the biological meaning of the peaks is difficult to interpret, as the two species seem to have a 2‐year life span. A biometric analysis of shell sizes revealed that length and width are the most variable parameters during the growth of the animal.     

A comparison of growth patterns between non‐indigenous Halophila nipponica and the native sympatric Zostera marina on the southern coast of the Korean peninsula

The growth dynamics of two co‐occurring seagrass species, Zostera marina and Halophila nipponica, were examined on the southern coast of the Korean peninsula. Zostera marina is a native dominant seagrass species in Korean coastal waters, whereas H. nipponica is a non‐native tropical and subtropical species that has extended its distributional range to the temperate coastal areas of Korea. To examine the differences in the growth dynamics of H. nipponica and Z.  marina, their morphology, density, productivity and biomass, as well as local environmental conditions, were monitored monthly from January 2008 to July 2009. Underwater irradiance at the study site was the highest in April 2009 and the lowest in January 2008. Water temperature ranged from 10.4°C in January 2009 to 24.8°C in September 2008. Significant differences in growth dynamics were observed between the species, due to the effect of water temperature at the study site. Density and areal productivity were the highest in April 2008 and June 2008, respectively, for Z.  marina but the highest in July 2008 for H. nipponica. Leaf size, shoot height and shoot weight were the highest in July 2008 for Z.  marina but the highest in August 2008 or September 2008 for H. nipponica. The productivity of both species was strongly correlated with water temperature at the study site. However, the productivity of these species was not strongly correlated with underwater irradiance or the nutrient availability of either the water column or sediment pore water. Zostera marina exhibited the ecological characteristics of a temperate seagrass, whereas H. nipponica retained the features of a subtropical/tropical seagrass, even after adapting to the temperate coastal waters of Korea.     

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.     

Distribution and population structure of deep‐dwelling red coral in the Northwest Mediterranean

Commercially harvested since ancient times, the highly valuable red coral Corallium rubrum (Linnaeus, 1758) is an octocoral endemic to the Mediterranean Sea and adjacent Eastern Atlantic Ocean, where it occurs on rocky bottoms over a wide bathymetric range. Current knowledge is restricted to its shallow populations (15–50 m depth), with comparably little attention given to the deeper populations (50–200 m) that are nowadays the main target of exploitation. In this study, red coral distribution and population structure were assessed in three historically exploited areas (Amalfi, Ischia Island and Elba Island) in the Tyrrhenian Sea (Western Mediterranean Sea) between 50 and 130 m depth by means of ROV during a cruise carried out in the summer of 2010. Red coral populations showed a maximum patch frequency of 0.20 ± 0.04 SD patches·m^?1 and a density ranging between 28 and 204 colonies·m^?2, with a fairly continuous bathymetric distribution. The highest red coral densities in the investigated areas were found on cliffs and boulders mainly exposed to the east, at the greatest depth, and characterized by medium percentage sediment cover. The study populations contained a high percentage (46% on average) of harvestable colonies (>7 mm basal diameter). Moreover, some colonies with fifth‐order branches were also observed, highlighting the probable older age of some components of these populations. The Ischia population showed the highest colony occupancy, density and size, suggesting a better conservation status than the populations at the other study locations. These results indicate that deep dwelling red coral populations in non‐stressed or less‐harvested areas may diverge from the inverse size‐density relationship previously observed in red coral populations with increasing depth.     

Sexual reproduction and early life‐history traits of the Mediterranean soft coral Alcyonium acaule

Understanding processes that contribute to a better comprehension of the population dynamics of long‐lived species is critical for the maintenance and potential recovery of such species. Despite the abundance of soft corals in Mediterranean rocky reefs, little information exists on their life histories and reproductive patterns. In this study, we assessed the main reproductive characteristics and early life‐history traits of the long‐lived soft coral Alcyonium acaule. The sex ratio was 1:1; the smallest fertile colonies were one finger in size (2.1 ± 0.6 cm in height), and both colony and polyp fertility increased with colony size. Likewise, the number of eggs and spermary sacs per polyp increased significantly with colony size, whereas the diameter of the female and male sexual products did not. Over 6 years of observations (2007–2012), spawning occurred primarily in July, after the seawater reached 20 °C, in a single spawning episode per year. Approximately 80% of female colonies released eggs, which were retained on the surface of the mother colony by mucous strings for up to a few days. High fertilization rates were observed during spawning in 2008 and 2009 (94.9% and 87.0%, respectively). The timing of development was ~24 h for the blastulae, ~48–72 h for the planulae and 8–22 days for metamorphosis into primary polyps. Survivorship of planulae was relatively high (~50% at 45 days after release), but only 24% of larvae metamorphosed into primary polyps, and their survivorship was moderate after 2 months (65% in 2008 and 74% in 2009). Asexual reproduction was negligible, indicating that sexual reproduction is the main mechanism supporting the maintenance and recovery of populations.     

Fitness‐related consequences shed light on the mechanisms of covering and sheltering behaviors in the sea urchin Glyptocidaris crenularis

We know of no comparative assessment on the benefits and costs of long‐term covering and sheltering behaviors in sea urchins. The present study investigated the long‐term effects of conditions suitable for sheltering and covering behaviors on fitness‐related traits of sea urchins Glyptocidaris crenularis. In general, conditions suitable for covering and sheltering behaviors significantly affected the fitness‐related traits of G. crenularis in a long‐term laboratory study of 31 months. Glyptocidaris crenularis kept in conditions suitable for sheltering behavior (bricks with openings) showed significantly lower test size, body weight, organ (test, lantern, gonad and gut) weights, gonad index and slower gonad development than those kept in conditions suitable for covering behavior (presence of shells) and the control conditions (without conditions for covering and sheltering). However, the index of maximum pressure resistance of the test was significantly higher in G. crenularis kept in the sheltering conditions than those in the covering and control conditions. The present study provides new insight into the mechanisms of covering and sheltering behaviors and has implications for the conservation and aquaculture of sea urchins.     

Growth and population dynamics of the non‐indigenous species Branchiomma luctuosum Grube (Annelida,Sabellidae) in the Ionian Sea (Mediterranean Sea)

The population dynamics and gametogenesis of the non‐indigenous polychaete species Branchiomma luctuosum Grube, 1869 (Annelida, Sabellidae) has been investigated at three sites in the Taranto Seas (Ionian Sea, Mediterranean Sea). The species, probably introduced from the Red Sea, has been reported in the Mediterranean Sea since 1983. The species is hermaphrodite, and the reproductive season is between June and October when the largest mean size of oocytes was recorded together with the presence of mature spermatozoa. Small oocytes are present in specimens reaching about 20–25 mm in length. Therefore oogenesis seems to begin early during the first year of life, but the first reproduction can occur when the worms attain a larger size and are at least 6 months of age. Although most of the individuals reproduce seasonally within a discrete period, some individuals can reproduce in different periods during the year, the oogenesis of individuals not being synchronous. A life span of at least 2 years is highlighted, with a faster growth rate during the first months (about 20 mm per month) decreasing to about 10 mm from the 3rd to the 8th months and slowing down again after the worm reaches 100 mm in size. Some differences in growth performance are discussed, enhanced by comparing the sites located at greater depth (5–7 m) and those located at 0.5 m depth, together with the possible interactions of this alien species with the autochthonous sabellid Sabella spallanzanii.     

The putative impacts of the non‐native seaweed Sargassum muticum on native communities in tidepools of Southern California and investigation into the feasibility of local eradication

The ecological impacts of introduced seaweeds have been relatively understudied. Current research suggests that seaweed invasions often result in alterations of native marine communities and disruptions of normal ecosystem functioning, but the effects on native communities can vary among invasive seaweed species, among habitats and over small and large spatial scales. In this study, the impacts of Sargassum muticum, a non‐native brown alga introduced into southern California, USA, several decades ago, were examined by comparing community structure in rocky inter‐tidal pools with and without the seaweed. Sargassum muticum appeared to have little impact on the native community despite measures revealing changes in the abiotic conditions of pools, with S. muticum presence reducing light penetration and ameliorating pool temperature changes during low tides. In other regions and habitat types, S. muticum presence often, but not consistently, resulted in declines in macrophyte diversity and/or abundance and increases in faunal assemblages. The lack of effects of S. muticum in this study, combined with variable impacts by S. muticum and other invasive seaweeds worldwide, suggests that predicting the effects of introduced seaweeds is problematic and warrants further research. Regardless of the effects on native communities, there is often a desire to eradicate or control the spread of non‐native seaweeds. In this study, localized S. muticum eradication attempts, including manipulations of a native canopy and herbivorous urchins, proved unsuccessful as full recovery occurred in ~9 months. While eradication efforts conducted worldwide have resulted in mixed success, there is a trend that early detection and rapid response can increase success, highlighting a need for systematic monitoring and establishment of regional rapid response plans.