Does spawning behavior minimize exposure to environmental stressors for encapsulated gastropod embryos on rocky shores?

Adults of motile intertidal invertebrates are able to seek shelter to avoid environmental stress associated with low tides, but embryos within egg masses are effectively sessile for the duration of their encapsulation. Gastropod egg masses from 34 taxa on two rocky shores in SE Australia (34°37′08″S, 150°92′03″E and 34°35′45″S, 150°53′20″E) were surveyed over 2 years (June 2002–May 2004) to test the hypothesis that eggs are deposited in patterns that minimize exposure to environmental stress. Egg masses were expected to be predominantly deposited in shaded habitats not prone to environmental extremes. It was also anticipated that the deposition of egg masses in habitats exposed to UVR, desiccation, and/or extremes in temperature would occur when exposure to these abiotic factors was minimized. Among the taxa investigated, only four species spawned in full sun (Bembicium nanum, Nerita morio, Siphonaria zelandica and S. denticulata). Summer had the highest UVR index, water temperature, and air temperature as well as the lowest daytime tides. Univariate and multivariate analyses confirmed that egg mass abundance was highest during summer, with no change in egg mass size. This study shows that those species depositing egg masses on the surfaces of rock platforms do not adjust the seasonal timing or macrohabitat location of their spawning to avoid physiologically stressful conditions, particularly UVR. Alternate reasons for the evolution of egg mass deposition behavior in apparently sub-optimal habitats are discussed, and it is almost certainly the complex interplay of a variety of highly species-specific factors that is responsible for the patterns observed.     

Recovery after trampling disturbance in a canopy-forming seaweed population

The recovery of Ascophyllum nodosum, and the associated benthic community, was followed after repeated trampling disturbance at different intensities, from 2005 to 2010, at its southern limit (41°41′27″N; 8°50′57″W). The cover of A. nodosum and associated organisms and the demography of A. nodosum individuals in the most disturbed areas were followed over a recovery period of 54 months. Recovery of A. nodosum assemblages was negatively affected by increasing trampling intensity. After 54 months, the highest intensity plots were still significantly different from control plots. Growth and mortality rates of A. nodosum increased shortly after disturbance but progressively recovered to levels close to control plots. The results of this study demonstrate the sensitivity of this habitat-building species to human trampling and also show its plasticity to cope with disturbance events.     

Domoic acid in benthic flatfish on the continental shelf of Monterey Bay,California, USA

Within Monterey Bay, California, USA, the food web transfer of domoic acid (DA), a neurotoxin produced by diatoms of the genus Pseudo-nitzschia, has led to major mortality events of marine mammals and birds. Less visible, and less well known, is whether invertebrates and fish associated with the benthos are also affected by blooms of DA-producing Pseudo-nitzschia spp. This study examines the presence of DA in benthic flatfish offshore of Davenport, California, (37°0′36″N, 122°13′12″W) and within Monterey Bay, California (36°45′0″N, 122°1′48″W), including species that feed primarily in the sediment (benthic-feeding) and species that feed primarily in the water column (benthopelagic-feeding). Flatfish caught between 10 December 2002 and 17 November 2003 at depths of 30–180 m had concentrations of DA in the viscera ranging from 3 to 26 μg DA g⁻¹ of viscera. Although the DA values reported are relatively low, benthic-feeding flatfish were frequently contaminated with DA, especially as compared with the frequency of contamination of flatfish species that feed in the water column. Furthermore, on days in which both benthic-feeding and benthopelagic-feeding flatfish were collected, the former had significantly higher concentrations of DA in the viscera. Curlfin turbot, Pleuronicthys decurrens, the flatfish with both the highest level and frequency of DA contamination, are reported to feed exclusively on polychaetes, suggesting that these invertebrates may be an important vector of the toxin in benthic communities and may pose a risk to other benthic-feeding organisms. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Interpreting seawater temperature range using oxygen isotopes and zooid size variation in Pentapora foliacea (Bryozoa)

This is the first investigation of how two independent proxies for seawater temperature inference (zooid size variation and oxygen isotope ratios of skeletal carbonate) relate to the actual measured ranges of temperature experienced by cheilostome bryozoan colonies. Nine specimens of the bimineralic marine cheilostome bryozoan Pentapora foliacea (Ellis and Solander, 1786) were analysed, collected from ~18-m depth at two localities in Wales, UK—four from Skomer Island (51°42′510″N, 5°13′42.60″W) and five from Porth Ysgaden, Lleyn Peninsula (52°54′6.75″N, 4°38′47.34″W). The annual range of temperature implied by zooid size variability provides a good approximation of the actual range of temperature recorded by a datalogger. However, annual ranges of temperature reconstructed from skeletal oxygen isotope ratios were narrower, typically not showing the lowest temperatures experienced by the colonies. This can be explained by progressive thickening of zooid skeletal walls during the life of the colony that homogenises the temperature signal by time-averaging over the lifetime of the colonies. Our study provides evidence that a combined morphological isotope approach has great potential in the reconstruction of annual ranges in seawater temperatures from historical and fossil bryozoans, particularly for species that lack ontogenetic skeletal wall thickening and bimineralic skeletal composition. As cheilostome bryozoans have been common in benthic communities since the Late Cretaceous, they represent a valuable and underutilised resource for the interpretation of environmental regimes.     

Spatial–temporal variability of epibenthic assemblages on subtidal biogenic reefs in the northern Adriatic Sea

Spatial distribution and temporal variation of epibenthic assemblages of coralligenous biogenic rocky outcrops occurring in the northern Adriatic Sea (45°04′–45°24′N; 12°23′–12°43′E) were investigated by photographic sampling from 2003 to 2006 at 12 randomly selected sites. The dominant reef-forming organisms were the encrusting calcareous algae (Lithophyllum stictaeforme, Lithothamnion minervae and Peyssonnelia polymorpha), while the main bioeroders were boring sponges (Cliona viridis, C. celata, C. thoosina, C. rhodensis, Piona vastifica) and the bivalve Gastrochaena dubia. Composition of the assemblages varied thorough years and among sites. Spatial heterogeneity, at local and regional scale, prevailed over temporal variation. This variability was related both to the geo-morphological features of the outcrops and to environmental variables. Sites clearly differed in the percent cover of reef builder and bioeroder species while only limited temporal variation within site was found. Some taxa revealed complex intra-site temporal trends. These results provide valuable information on the diversity and variability of epibenthic assemblages of the northern Adriatic coralligenous reefs, essential for the management and conservation of these unique biogenic habitats.     

An experimental field test of host-finding mechanisms in a Caribbean gnathiid isopod

Field experiments were conducted from dusk to dawn off St. John (18° 18′ 59.32″ N, 64° 43′ 24.5″ W) and Guana Island (18° 28″ 28.31″ N, 64° 34′ 30.83″ W), Virgin Islands from June through August 2008-2010 to assess the sensory cues used by the nocturnal/crepuscular fish-parasitic gnathiid isopod, Gnathia marleyi, to locate fish hosts. Experimental traps providing both visual and olfactory cues from live French grunts (Haemulon flavioliniatum) attracted significantly more gnathiids than traps providing only visual cues or control traps (empty or with a rock), which were not significantly different from each other. In another experiment, traps providing both cues and only olfactory cues attracted significantly more gnathiids than empty control traps, but were not significantly different from each other. Our findings suggest that during nocturnal and crepuscular periods, visual cues provided by resting or slow-moving fish are not alone sufficient to attract gnathiids, while olfactory cues alone are. The traps designed for this study offer a new method of sampling free-living gnathiid isopods.     

Regional-scale variation in the distribution and abundance of farming damselfishes on Australia’s Great Barrier Reef

Territorial damselfishes that manipulate (“farm”) the algae in their territories can have a marked effect on benthic community structure and may influence coral recovery following disturbances. Despite the numerical dominance of farming species on many reefs, the importance of their grazing activities is often overlooked, with most studies only examining their roles over restricted spatial and temporal scales. We used the results of field surveys covering 9.5° of latitude of the Great Barrier Reef to describe the distribution, abundance and temporal dynamics of farmer communities. Redundancy analysis revealed unique subregional assemblages of farming species that were shaped by the combined effects of shelf position and, to a lesser extent, by latitude. These spatial patterns were largely stable through time, except when major disturbances altered the benthic community. Such disturbances affected the functional guilds of farmers in different ways. Since different guilds of farmers modify benthic community structure and affect survival of juvenile corals in different ways, these results have important implications for coral recovery following disturbances.     

Size and spatial structure in deep versus shallow populations of the Mediterranean gorgonian <Emphasis Type="Italic">Eunicella singularis</Emphasis> (Cap de Creus,northwestern Mediterranean Sea)

In the Western Mediterranean Sea, the gorgonian Eunicella singularis (Esper, 1794) is found at high densities on sublittoral bottoms at depths from 10 to 70 m. Shallow colonies have symbiotic zooxanthellae that deeper colonies lack. While knowledge of the ecology of the shallow populations has increased during the last decades, there is almost no information on the ecology of the deep sublittoral populations. In October and November 2004 at Cap de Creus (42°19′12″ N; 03°19′34″ E), an analysis of video transects made by a remotely operated vehicle showed that shallow populations (10–25 m depth) were dominated by small, non-reproductive colonies, while deep sublittoral populations (50–67 m depth) were dominated by medium-sized colonies. Average and maximum colony heights were greater in the deeper populations, with these deeper populations also forming larger patch sizes and more extensive regions of continuous substrate coverage. These results suggest that shallow habitats are suitable for E. singularis, as shown by the high recruitment rate, but perturbations may limit or delay the development of these populations into a mature stage. This contrasts with the deep sublittoral habitats where higher environmental stability may allow the development of mature populations dominated by larger, sexually mature colonies.     

The reproductive ecology of the invasive ascidian, <Emphasis Type="Italic">Styela clava,</Emphasis> in Auckland Harbour,New Zealand

The ascidian Styela clava, native to the north-west Pacific, is an invasive species affecting New Zealand’s marine ecosystems, biodiversity and aquaculture operations. To provide detailed information on the reproductive biology of S. clava in New Zealand for post-border biosecurity management, long-term seasonal patterns of gametogenesis were determined from May 2006 to May 2008 in Auckland’s Waitemata Harbour (36°49′20″S, 174°45′85″E). Of particular interest was whether the critical 15°C threshold spawning temperature for reproduction observed in the Northern Hemisphere applied here to the first Southern Hemisphere study. S. clava gametogenesis followed a regular seasonal cycle with ripe gametes appearing as early as September and persisting to June; this time frame corresponds to the period when sea surface temperatures in the region first reach 15°C and with spawning occurring mainly during late summer to early autumn. From photoperiod manipulation, it was determined that spawning occurred at approximately 18:20. The extended reproductive period and a short generation time in the Waitemata Harbour provides a lengthy opportunity for S. clava to spread. Findings are discussed in relation to S. clava’s post-border management.     

Relationship between depth, sediment, latitude, and the structure of benthic infaunal assemblages on the mainland shelf of southern California

A regional benthic survey was conducted in 1994, and the data were used to assess the relationship among three habitat factors (depth, sediment grain size, and latitude) and the distribution of benthic infaunal assemblages on the southern California coastal shelf. Benthic samples were collected with a 0.1 m² Van Veen grab from 251 sites on the continental shelf (10–200 m deep) from Point Conception, California, to the United States–Mexico international border. The relationship between habitat and assemblages was investigated by conducting a Q-mode cluster analysis to define groups of stations with similar species composition and then examining whether differences were present in physical habitat attributes among those groups of stations. Analysis of data from 175 uncontaminated sites yielded four habitat-related benthic infaunal assemblages along the southern California coastal shelf: a shallow-water assemblage from 10–32 m, a mid-depth assemblage between 32 and 115 m, and two deep-water (115–200 m) assemblages, one in fine and one in coarse sediments. These empirically defined points in the depth and sediment grain size gradients can be used to define reference habitats for the development of biocriteria. Benthic abundance and diversity were greatest in the mid-depth assemblage, conforming to predictions for benthic assemblages in regions of upwelling. Within the 500 km of coastline examined, latitude was not an important factor in defining assemblages. Received: 3 December 1999 / Accepted: 9 October 2000     

Seasonal variation in the diversity and abundance of pelagic larvae of Antarctic marine invertebrates

Most marine benthic macroinvertebrate species reproduce via a larval phase but attempts to explain the occurrence of different larval strategies (feeding or non-feeding, pelagic or benthic) in different habitats have been largely inconclusive. There have been very few year-round surveys of meroplankton at any latitude and in consequence fundamental data on the diversity, abundance, and timings of larval life history phases are lacking. There has been considerable debate regarding the viability of pelagic larvae in cold waters with highly seasonal primary production but there has been only one year-round study of meroplankton in the Southern Ocean, and that was outside of the Antarctic Circle. We present data from the first year-round survey of meroplankton assemblages at a location within the Antarctic Circle. We surveyed abundances of meroplanktonic larvae over 1.5 year at Rothera Point, West Antarctic Peninsula (67°34′S, 68°07′W). Larvae were collected in monthly diver-towed net samples close to the seabed at 20 and 6 m total water depths at each of three locations and were identified and counted live immediately after sampling. A total of 99 operationally defined taxonomic types representing 11 phyla were recorded but this is likely to be an underestimate of true diversity because of inherent difficulties of identification. Larvae were present in all months of the year and although planktotrophic larvae were more abundant in summer, both feeding and non-feeding types were present in all months. Comparisons of seasonal larval abundances with data from a settlement study at the same sites and from the literature show that larvae of mobile adults settle in summer regardless of developmental type, whereas sessile taxa settle in all seasons. We suggest that this is a consequence of differences in the food requirements of mobile and sessile fauna and that the availability of food for post-larval juveniles is more critical for survival than factors affecting the larval stage itself.     

Flipper beat frequency and amplitude changes in diving green turtles, <Emphasis Type="Italic">Chelonia mydas</Emphasis>

Using a turtle-borne camera system, changing flipper beat frequency and amplitude were measured in five diving green turtles (Chelonia mydas Linnaeus 1758) in the Bahía de los Angeles, Mexico (28°58′N, 113°33′W). These observations were made between June and August 2002. Turtles worked hardest (i.e., had the highest flipper beat frequency and amplitude) at the start of descents when positive buoyancy is predicted to oppose their forward motion. During the later part of descents, turtles worked less hard in line with opposing buoyancy forces being reduced. For example, flipper beat frequency declined from about 60–80 beats min⁻¹ at the start of descent to around 25–40 beats min⁻¹ after 30 s of the descent. At the start of ascents the flipper beat frequency was around 30 beats min⁻¹, lower than on descent, and declined as the ascent progressed with often passive gliding for the final few meters to the surface. This pattern of effort during diving appears to apply across a range of marine reptiles, birds and mammals suggesting that graded effort during descent and ascent is an optimum solution to minimising the cost of transport during diving.     

Macrofaunal responses to structural complexity are mediated by environmental variability and surrounding habitats

Investigating the context that surrounds each habitat is crucial to understand local responses of assemblages of species to habitats. Here, I tested whether responses of benthic macroinvertebrates to the structural complexity of experimental habitats were mediated by the characteristics of their surrounding habitats (i.e. rockpools or emergent-rock surfaces). Each type of surrounding habitat provided particular biotic (e.g. algal growth) and abiotic (e.g. temperature, water movement) conditions that were expected to affect benthic assemblages. The results show that (1) composition of entire assemblages was affected by the matrix and type of habitat; (2) effects of the matrix on the number of species varied depending on the different types of habitats; (3) abundant species showed specific responses to type of habitat, independently of the matrix; and (4) relationships between numbers of species and two major environmental variables (i.e. micro-algal biomass and sediment) varied depending on the type of habitats and the surrounding matrix. Generally, these findings demonstrate that understanding the consequences of the spatial structure of these habitats is essential to advance our knowledge on patterns of abundance and distributions of functionally important species and ultimately the structure of intertidal assemblages.     

Habitat modification affects recruitment of abalone in central New Zealand

The habitat experienced during early life-history stages can determine the number and quality of individuals that recruit to adult populations. In a field experiment, biogenic habitat complexity was manipulated (presence or absence of foliose macroalgae) at two depths (2–3 m and 5–6 m) and the habitat-dependent effects on recruitment of the black foot abalone (Haliotis iris) were examined at three field sites along the south coast of Wellington, New Zealand (41°20′S, 174°47′E), between July and November 2005. Recruit density (<5 weeks post-settlement) was measured on cobbles covered with crustose coralline algae. Habitats of low complexity (barrens treatments) had consistently greater densities of recruits than habitats of high complexity (algae treatments). However, recruits in algae habitats were larger, and for deep habitats, there was greater survival in algae habitats compared with barrens habitats. While depth had no significant effect on early recruit (<2 weeks post-settlement) density, late recruit (<5 weeks post-settlement) density was greater in shallow habitats, and so it seems recruit survival was greater in shallow habitats. In this experiment, algal habitat complexity had strong effects on early recruit abundance, but habitat-dependent variations in recruit growth and survival may modify initial patterns of abundance and determine recruitment to adult abalone populations.     

The effects of symbiotic state on heterotrophic feeding in the temperate sea anemone <Emphasis Type="Italic">Anthopleura elegantissima</Emphasis>

The temperate sea anemone Anthopleura elegantissima is facultatively symbiotic with unicellular algae. Symbiotic A. elegantissima can supplement heterotrophic feeding with excess photosynthate from their algal partners, while asymbiotic individuals must rely solely on heterotrophy. A. elegantissima individuals were collected from Swirl Rocks, Washington (48°25′6″ N, 122°50′58″ W) in July 2010, and prey capture and feeding characteristics were measured to determine whether asymbiotic individuals are more efficient predators. Feeding abilities were then measured again after a 3-week exposure to full sunlight or shaded conditions. Freshly collected asymbiotic anemones had larger nematocysts, but symbiotic individuals showed greater nematocyte sensitivity. Sunlight enhanced digestion and reduced cnida density in all anemones regardless of symbiotic state. Results suggest that the phototropic potential of A. elegantissima, as influenced by symbiotic condition, has little effect on heterotrophic capacity. The anemones appear to maximize heterotrophic energy input independent of the presence or identity of their algal symbionts.     

Erratum to: Patterns of spatial variability of seagrass epiphytes in the north-west Mediterranean Sea

This study aimed to gain insight on patterns of spatial variability of seagrass epiphytes of both leaves and of rhizomes in three different habitats, continental coasts, offshore banks and islands. Moreover, we tried to discriminate between habitat-dependant variability and scale-dependant variability of Posidonia oceanica epiphytic assemblages. Results showed the absence of significant differences in the structure of assemblages of epiphytes both on leaves and on rhizomes among continental coasts, offshore banks and islands, even if the patterns of spatial variability changed among habitats. In fact, although a high variability at small scales appeared a constant pattern in epiphytic assemblages, large-scale variability resulted higher in continental coasts and offshore banks than in islands. In conclusion, epiphytic assemblages of Posidonia oceanica appear homogeneous among habitats, showing a similar structure and species composition in the same geographic area. On the contrary, differences between meadows appeared linked to local differences in environmental factors that are more evident in habitats influenced by human disturbance. An erratum to this article can be found at     

Diversity and cover of a sessile animal assemblage does not predict its associated mobile fauna

Habitat-forming organisms often determine the structural properties and food resources available to a wide diversity of associated mobile species. Sessile invertebrate assemblages on marine hard substrates support an abundant fauna of mobile invertebrates whose associations with traits of their host assemblages are poorly known. To assess how changes to habitat-forming species are likely to affect their associated mobile fauna, the relationships between abundance, diversity and composition of mobile invertebrates and the diversity, cover and composition of the sessile assemblages they use as habitat were quantified in Sydney Harbour, Australia (33°50′S, 151°16′E). Similar compositions of sessile species were more likely to share a similar composition of mobile species, but univariate measures of the habitat (percent cover, species and functional diversity, prevalence of non-indigenous species) did not predict variation in associated mobile assemblages. These results demonstrate that in this habitat it is difficult to predict the diversity of marine assemblages based on common surrogate measures of biodiversity.     

The application of predicted habitat models to investigate the spatial ecology of demersal fish assemblages

Benthic habitats are known to influence the abundance and richness of demersal fish assemblages; however, little is known about how habitat structure and composition influences these distributions at very fine scales. We examined how the benthic environment structures marine fish assemblages using high-resolution bathymetry and accurate predicted benthic habitat maps. Areas characterised by a mosaic of habitat patches supported the highest richness of demersal fishes. A total of 37.4% of the variation in the distribution of the fish assemblage was attributed to 6 significant variables. Depth explained 23.0% of the variation, with the boulders explaining 12.6% and relief 1.4%. The remaining measures (seawhips, light/exposure and solid reef) provided a small (<1.0%) but significant contribution. Identifying components of the benthic environment important in structuring fish assemblages and understanding how they influence the spatial distribution of marine fishes is imperative for better management of demersal fish populations.     

Seasonal distribution patterns of juvenile loggerhead sea turtles (<Emphasis Type="Italic">Caretta caretta</Emphasis>) following capture from a shipping channel in the Northwest Atlantic Ocean

Thirty-four juvenile loggerhead sea turtles captured by trawling from the Charleston, South Carolina (USA), shipping channel (32°42′N; −79°47′W) between May 2004 and August 2007 were tagged with satellite transmitters to assess the extent to which they remained near the capture location given their collection along a seasonal migratory corridor. Seventy-five percent of juveniles were classified as seasonal residents. Migrants predominantly swam north in the spring and nomads wandered south in the summer, but predictive indicators for non-resident status were not identified. All but one juvenile generally remained south of 34°N, within 40 km of shore, and in waters <30 m deep throughout the year. Nine of 14 loggerhead sea turtles monitored during the winter remained exclusively over the continental shelf, three briefly occurred in oceanic habitats, and two foraged extensively in oceanic habitats. Residents distributed >15 km from shore between spring and autumn were three times as likely to occur in oceanic habitats in winter. Modest seasonal movements contrasted with adults tagged at similar latitudes and with juveniles tagged further north and suggest distinct foraging groups within a regional foraging ground.     

Multivariate analysis of the bentho-demersal ichthyofauna along soft bottoms of the Eastern Atlantic: comparison between unvegetated substrates,seagrass meadows and sandy bottoms beneath sea-cage fish farms

By means of multivariate techniques, we studied: (1) the differences in the structure of bentho-demersal, non-cryptic, fish assemblages associated with unvegetated sandy substrates, vegetated meadows constituted by the seagrass Cymodocea nodosa and the bottoms under the influence of sea-cage fish farms; as well as (2) the persistence of these patterns with regard to different scales of spatial variability, across three islands of the Canarian Archipelago (Central East Atlantic). Our sampling strategy (involving three islands, with five locations per island, and two sites within each location) detected significant changes in the composition and structure of the fish assemblages between the three habitats. Fish assemblages associated with the unvegetated and vegetated bottoms were similar among the surveyed islands. In contrast, we observed a significant inter-island variability in the fish populations associated with the sea-cage fish farms. The presence of the sea-cage fish farms increased the overall fish abundance (184.8±49.8 ind 100 m⁻²) as compared to both the vegetated (38.8±9.7 ind 100 m⁻²) and unvegetated habitats (1.1±0.4 ind 100 m⁻²). Differences within and between the habitats were found to be associated with the relative abundance of a few fish species. The most abundant species were Xyrichthys novacula in the unvegetated bottoms and Diplodus annularis, Spondyliosoma cantharus and Mullus surmuletus on the seagrass meadows. Finally, we recorded an increase in the abundance of Heteroconger longissimus, Trachinus draco and Pagellus acarne in the bottoms beneath the sea-cage fish farms. These species, in addition to a group of large benthic chondrichthyes, were responsible for the differences between islands in the composition and structure of the demersal ichthyofauna beneath the sea-cage fish farms.