Implications of substrate complexity and kelp variability for south-central Alaskan nearshore fish communities

Understanding the interactions between kelp beds and nearshore fish is essential because anthropogenic changes and natural variability in these beds may affect available habitat for fishes. In this study fish communities were investigated in south-central Alaska kelp beds characterized by a range of substrate complexity and varying densities of both perennial understory kelps and annual canopy kelps. Many of the observed fish species, as well as understory and canopy kelps, were positively associated with structurally complex substratum. Targeted canopy and understory kelp beds supported seasonal populations of adult and juvenile Pacific cod (Gadus macrocephalus), rockfishes (Sebastes spp.), and year-round populations of greenlings (Hexagrammos spp.). Monthly changes in kelp and fish communities ref lected seasonal changes; the densities of some species were greatest during periods with higher temperatures. This work illustrates the importance of structurally complex kelp beds with persistent understory kelp populations as important fish habitat for several commercially and recreationally important fishes

Size-frequency distribution, growth, and mortality of snow crab (Chionoecetes opilio) and arctic lyre crab (Hyas coarctatus) in the Chukchi Sea from 2009 to 2013

The snow crab Chionoecetes opilio and Arctic lyre crab Hyas coarctatus are prominent members of the Chukchi Sea epifaunal community. A better understanding of their life history will aid in determining their role in this ecosystem in light of the changing climate and resource development. In this study, the size frequency distribution, growth, and mortality of these two crab species was examined in 2009, 2010, 2012, and 2013 to determine temporal and spatial patterns within the eastern Chukchi Sea, and to identify potential environmental drivers of the observed patterns. Temporally, the mean size of both sexes of C. opilio and H. coarctatus decreased significantly from 2009 to 2013, with the number of rare maximum sized organisms decreasing significantly to near absence in the latter two study years. Spatially, the mean size of male and female crabs of both species showed a latitudinal trend, decreasing from south to north in the investigation area. Growth of both sexes of C. opilio and H. coarctatus was linear over the sampled size range, and mortality was highest in the latter two study years. Life history features of both species related to different environmental parameters in different years, ranging from temperature, the sediment carbon to nitrogen ratio of the organic content, and sediment grain size distribution. Likely explanations for the observed temporal and spatial variability are ontogenetic migrations of mature crabs to warmer areas possibly due to cooler water temperatures in the latter two study years, or interannual fluctuations, which have been reported for C. opilio populations in other areas where successful waves of recruitment were estimated to occur in eight year intervals. Further research is suggested to determine if the spatial and temporal patterns found in this study are part of the natural variability in this system or if they are an indication of long-term trends

Large-scale spatial distribution patterns of echinoderms in nearshore rocky habitats

This study examined echinoderm assemblages from nearshore rocky habitats for large-scale distribution patterns with specific emphasis on identifying latitudinal trends and large regional hotspots. Echinoderms were sampled from 76 globally-distributed sites within 12 ecoregions, following the standardized sampling protocol of the Census of Marine Life NaGISA project ( www.nagisa.coml.org ). Sample-based species richness was overall low (2 cm in 1 m 2 quadrats) was highest in the Caribbean ecoregions and echinoids dominated these assemblages with an average of 5 ind m −2 . In contrast, intertidal echinoderm assemblages collected from clearings of 0.0625 m 2 quadrats had the highest abundance and richness in the Northeast Pacific ecoregions where asteroids and holothurians dominated with an average of 14 ind 0.0625 m −2 . Distinct latitudinal trends existed for abundance and richness in intertidal assemblages with declines from peaks at high northern latitudes. No latitudinal trends were found for subtidal echinoderm assemblages with either sampling technique. Latitudinal gradients appear to be superseded by regional diversity hotspots. In these hotspots echinoderm assemblages may be driven by local and regional processes, such as overall productivity and evolutionary history. We also tested a set of 14 environmental variables (six natural and eight anthropogenic) as potential drivers of echinoderm assemblages by ecoregions. The natural variables of salinity, sea-surface temperature, chlorophyll a , and primary productivity were strongly correlated with echinoderm assemblages; the anthropogenic variables of inorganic pollution and nutrient contamination also contributed to correlations. Our results indicate that nearshore echinoderm assemblages appear to be shaped by a network of environmental and ecological processes, and by the differing responses of various echinoderm taxa, making generalizations about the patterns of nearshore rocky habitat echinoderm assemblages difficult

Aspects of Benthic Decapod Diversity and Distribution from Rocky Nearshore Habitat at Geographically Widely Dispersed Sites

Relationships of diversity, distribution and abundance of benthic decapods in intertidal and shallow subtidal waters to 10 m depth are explored based on data obtained using a standardized protocol of globally-distributed samples. Results indicate that decapod species richness overall is low within the nearshore, typically ranging from one to six taxa per site (mean = 4.5). Regionally the Gulf of Alaska decapod crustacean community structure was distinguishable by depth, multivariate analysis indicating increasing change with depth, where assemblages of the high and mid tide, low tide and 1 m, and 5 and 10 m strata formed three distinct groups. Univariate analysis showed species richness increasing from the high intertidal zone to 1 m subtidally, with distinct depth preferences among the 23 species. A similar depth trend but with peak richness at 5 m was observed when all global data were combined. Analysis of latitudinal trends, confined by data limitations, was equivocal on a global scale. While significant latitudinal differences existed in community structure among ecoregions, a semi-linear trend in changing community structure from the Arctic to lower latitudes did not hold when including tropical results. Among boreal regions the Canadian Atlantic was relatively species poor compared to the Gulf of Alaska, whereas the Caribbean and Sea of Japan appeared to be species hot spots. While species poor, samples from the Canadian Atlantic were the most diverse at the higher infraordinal level. Linking 11 environmental variables available for all sites to the best fit family-based biotic pattern showed a significant relationship, with the single best explanatory variable being the level of organic pollution and the best combination overall being organic pollution and primary productivity. While data limitations restrict conclusions in a global context, results are seen as a first-cut contribution useful in generating discussion and more in-depth work in the still poorly understood field of biodiversity distribution

Sea Ice and Substratum Shape Extensive Kelp Forests in the Canadian Arctic

The coastal zone of the Canadian Arctic represents 10% of the world’s coastline and is one of the most rapidly changing marine regions on the planet. To predict the consequences of these environmental changes, a better understanding of how environmental gradients shape coastal habitat structure in this area is required. We quantified the abundance and diversity of canopy forming seaweeds throughout the nearshore zone (5–15 m) of the Eastern Canadian Arctic using diving surveys and benthic collections at 55 sites distributed over 3,000 km of coastline. Kelp forests were found throughout, covering on average 40.4% (±29.9 SD) of the seafloor across all sites and depths, despite thick sea ice and scarce hard substrata in some areas. Total standing macroalgal biomass ranged from 0 to 32 kg m–2 wet weight and averaged 3.7 kg m–2 (±0.6 SD) across all sites and depths. Kelps were less abundant at depths of 5 m compared to 10 or 15 m and distinct regional assemblages were related to sea ice cover, substratum type, and nutrient availability. The most common community configuration was a mixed assemblage of four species: Agarum clathratum (14.9% benthic cover ± 12.0 SD), Saccharina latissima (13% ± 14.7 SD), Alaria esculenta (5.4% ± 1.2 SD), and Laminaria solidungula (3.7% ± 4.9 SD). A. clathratum dominated northernmost regions and S. latissima and L. solidungula occurred at high abundance in regions with more open water days. In southeastern areas along the coast of northern Labrador, the coastal zone was mainly sea urchin barrens, with little vegetation. We found positive relationships between open water days (days without sea ice) and kelp biomass and seaweed diversity, suggesting kelp biomass could increase, and the species composition of kelp forests could shift, as sea ice diminishes in some areas of the Eastern Canadian Arctic. Our findings demonstrate the high potential productivity of this extensive coastal zone and highlight the need to better understand the ecology of this system and the services it provides.publishedVersio

Spatial Relationships between Polychaete Assemblages and Environmental Variables over Broad Geographical Scales

This study examined spatial relationships between rocky shore polychaete assemblages and environmental variables over broad geographical scales, using a database compiled within the Census of Marine Life NaGISA (Natural Geography In Shore Areas) research program. The database consisted of abundance measures of polychaetes classified at the genus and family levels for 74 and 93 sites, respectively, from nine geographic regions. We tested the general hypothesis that the set of environmental variables emerging as potentially important drivers of variation in polychaete assemblages depend on the spatial scale considered. Through Moran's eigenvector maps we indentified three submodels reflecting spatial relationships among sampling sites at intercontinental (>10000 km), continental (1000–5000 km) and regional (20–500 km) scales. Using redundancy analysis we found that most environmental variables contributed to explain a large and significant proportion of variation of the intercontinental submodel both for genera and families (54% and 53%, respectively). A subset of these variables, organic pollution, inorganic pollution, primary productivity and nutrient contamination was also significantly related to spatial variation at the continental scale, explaining 25% and 32% of the variance at the genus and family levels, respectively. These variables should therefore be preferably considered when forecasting large-scale spatial patterns of polychaete assemblages in relation to ongoing or predicted changes in environmental conditions. None of the variables considered in this study were significantly related to the regional submodel

Current Patterns of Macroalgal Diversity and Biomass in Northern Hemisphere Rocky Shores

Latitudinal gradients in species abundance and diversity have been postulated for nearshore taxa but few analyses have been done over sufficiently broad geographic scales incorporating various nearshore depth strata to empirically test these gradients. Typically, gradients are based on literature reviews and species lists and have focused on alpha diversity across the entire nearshore zone. No studies have used a standardized protocol in the field to examine species density among sites across a large spatial scale while also focusing on particular depth strata. The present research used field collected samples in the northern hemisphere to explore the relationships between macroalgal species density and biomass along intertidal heights and subtidal depths and latitude. Results indicated no overall correlations between either estimates of species density or biomass with latitude, although the highest numbers of both were found at mid-latitudes. However, when strata were examined separately, significant positive correlations were found for both species numbers and biomass at particular strata, namely the intertidal ones. While the data presented in this paper have some limitations, we show that latitudinal macroalgal trends in species density and biomass do exist for some strata in the northern hemisphere with more taxa and biomass at higher latitudes

Large-scale spatial distribution patterns of gastropod assemblages in rocky shores

Gastropod assemblages from nearshore rocky habitats were studied over large spatial scales to (1) describe broad-scale patterns in assemblage composition, including patterns by feeding modes, (2) identify latitudinal pattern of biodiversity, i.e., richness and abundance of gastropods and/or regional hotspots, and (3) identify potential environmental and anthropogenic drivers of these assemblages. Gastropods were sampled from 45 sites distributed within 12 Large Marine Ecosystem regions (LME) following the NaGISA (Natural Geography in Shore Areas) standard protocol ( www.nagisa.coml.org ). A total of 393 gastropod taxa from 87 families were collected. Eight of these families (9.2%) appeared in four or more different LMEs. Among these, the Littorinidae was the most widely distributed (8 LMEs) followed by the Trochidae and the Columbellidae (6 LMEs). In all regions, assemblages were dominated by few species, the most diverse and abundant of which were herbivores. No latitudinal gradients were evident in relation to species richness or densities among sampling sites. Highest diversity was found in the Mediterranean and in the Gulf of Alaska, while highest densities were found at different latitudes and represented by few species within one genus (e.g. Afrolittorina in the Agulhas Current, Littorina in the Scotian Shelf, and Lacuna in the Gulf of Alaska). No significant correlation was found between species composition and environmental variables (r≤0.355, p>0.05). Contributing variables to this low correlation included invasive species, inorganic pollution, SST anomalies, and chlorophyll-a anomalies. Despite data limitations in this study which restrict conclusions in a global context, this work represents the first effort to sample gastropod biodiversity on rocky shores using a standardized protocol across a wide scale. Our results will generate more work to build global databases allowing for large-scale diversity comparisons of rocky intertidal assemblages

Trans-Arctic asymmetries, melting pots and weak species cohesion in the low-dispersal amphiboreal seaweed Fucus distichus

Amphiboreal taxa are often composed of vicariant phylogroups and species complexes whose divergence and phylogeographic affinities reflect a shared history of chronic isolation and episodic trans-Arctic dispersal. Ecological filters and shifting selective pressures may also promote selective sweeps, niche shifts and ecological speciation during colonization, but these are seldom considered at biogeographical scales. Here we integrate genetic data and Ecologic Niche Models (ENMs) to investigate the historical biogeography and cohesion of the polymorphic rockweed Fucus distichus throughout its immense amphiboreal range, focusing on trans-Arctic asymmetries, glacial/interglacial dynamics, and integrity of sympatric eco-morphotypes. Populations were sampled throughout the Pacific and the Atlantic, from southern rear-edges to the high-Arctic. They were genotyped for seven microsatellites and an mtDNA spacer, and genetic diversity and structure were assessed from global to local scales. ENMs were used to compare niche divergence and magnitude of post-glacial range shifts in Pacific versus Atlantic sub-ranges. Haplotypic and genotypic data revealed distinct and seemingly isolated Pacific vs Arctic/Atlantic gene-pools, with finer-scale regional sub-structuring pervasive in the Pacific. MtDNA diversity was highly structured and overwhelmingly concentrated in the Pacific. Regionally, Alaska showed the highest intra-population diversity but the lowest levels of endemism. Some sympatric/parapatric ecotypes exhibited distinct genotypic/ haplotypic compositions. Strikingly, niche models revealed higher Pacific tolerance to maximum temperatures and predicted a much more consolidated presence in the NE Atlantic. Glacial and modern ranges overlapped extensively in the Pacific, whereas the modern Atlantic range was largely glaciated or emerged during the Last Glacial Maximum. Higher genetic and ecogeographic diversity supports a primary Pacific diversification and secondary Atlantic colonization, also likely reflecting the much larger and more stable climatic refugia in the Pacific. The relic distribution and reduced ecological/morphological plasticity in the NE Atlantic are hypothesized to reflect functional trans-Arctic bottlenecks, recent colonization or competition with congeners. Within the Pacific, Alaska showed signatures of a post-glacial melting pot of eastern and southern populations. Genetic/ecotypic variation was generally not sufficiently discontinuous or consistent to justify recognizing multiple taxonomic entities, but support a separate species in the eastern Pacific, at the southern rear-edge. We predict that layered patterns of phylogeographic structure, incipient speciation and niche differences might be common among widespread low-dispersal amphiboreal taxa