A synthesis of genetic connectivity in deep-sea fauna and implications for marine reserve design.

With anthropogenic impacts rapidly advancing into deeper waters, there is growing interest in establishing deep-sea marine protected areas (MPAs) or reserves. Reserve design depends on estimates of connectivity and scales of dispersal for the taxa of interest. Deep-sea taxa are hypothesized to disperse greater distances than shallow-water taxa, which implies that reserves would need to be larger in size and networks could be more widely spaced; however, this paradigm has not been tested. We compiled population genetic studies of deep-sea fauna and estimated dispersal distances for 51 studies using a method based on isolation-by-distance slopes. Estimates of dispersal distance ranged from 0.24 km to 2028 km with a geometric mean of 33.2 km and differed in relation to taxonomic and life-history factors as well as several study parameters. Dispersal distances were generally greater for fishes than invertebrates with the Mollusca being the least dispersive sampled phylum. Species that are pelagic as adults were more dispersive than those with sessile or sedentary lifestyles. Benthic species from soft-substrate habitats were generally less dispersive than species from hard substrate, demersal or pelagic habitats. As expected, species with pelagic and/or feeding (planktotrophic) larvae were more dispersive than other larval types. Many of these comparisons were confounded by taxonomic or other life-history differences (e.g. fishes being more dispersive than invertebrates) making any simple interpretation difficult. Our results provide the first rough estimate of the range of dispersal distances in the deep sea and allow comparisons to shallow-water assemblages. Overall, dispersal distances were greater for deeper taxa, although the differences were not large (0.3-0.6 orders of magnitude between means), and imbalanced sampling of shallow and deep taxa complicates any simple interpretation. Our analyses suggest the scales of dispersal and connectivity for reserve design in the deep sea might be comparable to or slightly larger than those in shallow water. Deep-sea reserve design will need to consider the enormous variety of taxa, life histories, hydrodynamics, spatial configuration of habitats and patterns of species distributions. The many caveats of our analyses provide a strong impetus for substantial future efforts to assess connectivity of deep-sea species from a variety of habitats, taxonomic groups and depth zones.

Post-glacial redistribution and shifts in productivity of giant kelp forests.

Quaternary glacial-interglacial cycles create lasting biogeographic, demographic and genetic effects on ecosystems, yet the ecological effects of ice ages on benthic marine communities are unknown. We analysed long-term datasets to develop a niche-based model of southern Californian giant kelp (Macrocystis pyrifera) forest distribution as a function of oceanography and geomorphology, and synthesized palaeo-oceanographic records to show that late Quaternary climate change probably drove high millennial variability in the distribution and productivity of this foundation species. Our predictions suggest that kelp forest biomass increased up to threefold from the glacial maximum to the mid-Holocene, then rapidly declined by 40-70 per cent to present levels. The peak in kelp forest productivity would have coincided with the earliest coastal archaeological sites in the New World. Similar late Quaternary changes in kelp forest distribution and productivity probably occurred in coastal upwelling systems along active continental margins worldwide, which would have resulted in complex shifts in the relative productivity of terrestrial and marine components of coastal ecosystems.

The relationship between dispersal ability and geographic range size.

There are a variety of proposed evolutionary and ecological explanations for why some species have more extensive geographical ranges than others. One of the most common explanations is variation in species' dispersal ability. However, the purported relationship between dispersal distance and range size has been subjected to few theoretical investigations, and empirical tests reach conflicting conclusions. We attempt to reconcile the equivocal results of previous studies by reviewing and synthesizing quantitative dispersal data, examining the relationship between average dispersal ability and range size for different spatial scales, regions and taxonomic groups. We use extensive data from marine taxa whose average dispersal varies by seven orders of magnitude. Our results suggest dispersal is not a general determinant of range size, but can play an important role in some circumstances. We also review the mechanistic theories proposed to explain a positive relationship between range size and dispersal and explore their underlying rationales and supporting or refuting evidence. Despite numerous studies assuming a priori that dispersal influences range size, this is the first comprehensive conceptual evaluation of these ideas. Overall, our results indicate that although dispersal can be an important process moderating species' distributions, increased attention should be paid to other processes responsible for range size variation.

Reproduction on the edge: large-scale patterns of individual performance in a marine invertebrate.

Reproductive output is a central attribute of life history, providing a measure of individual and population performance. The fields of ecology, biogeography, and evolutionary biology take disparate approaches in addressing spatial variation in reproduction, and thus we lack clear predictions for how reproductive output should vary geographically. We empirically investigate these contrasting theoretical approaches by determining geographic patterns in reproductive output for intertidal populations of the purple sea urchin, Strongylocentrotus purpuratus, at 15 sites spanning a large geographic distance (9 degrees span of latitude) from central California, USA, to Baja California, Mexico. Contrary to predictions from biogeography, some of the highest values of reproductive output are at sites near the species' southern range boundary. Additionally, sea urchins do not exhibit a latitudinal gradient in reproduction, but rather show considerable mesoscale variation in reproductive output. Spatial analyses reveal that this variation is correlated with coastal topographic features that are known to influence the pattern of nearshore nutrient flux and circulation. We hypothesize that urchins' reproductive output may be driven by the spatial distribution of their food supply, drift macroalgae, the abundance of which is influenced by both nutrient supply and alongshore transport processes that are coupled to topographic features. Large-scale studies such as this provide valuable insight into the causes of species' range limits, population connectivity, habitat reserve design, and forecasting the effects of climate change on species' distributions.

Deep-water kelp refugia as potential hotspots of tropical marine diversity and productivity.

Classic marine ecological paradigms view kelp forests as inherently temperate-boreal phenomena replaced by coral reefs in tropical waters. These paradigms hinge on the notion that tropical surface waters are too warm and nutrient-depleted to support kelp productivity and survival. We present a synthetic oceanographic and ecophysiological model that accurately identifies all known kelp populations and, by using the same criteria, predicts the existence of >23,500 km(2) unexplored submerged (30- to 200-m depth) tropical kelp habitats. Predicted tropical kelp habitats were most probable in regions where bathymetry and upwelling resulted in mixed-layer shoaling above the depth of minimum annual irradiance dose for kelp survival. Using model predictions, we discovered extensive new deep-water Eisenia galapagensis populations in the Galápagos that increased in abundance with increasing depth to >60 m, complete with cold-water flora and fauna of temperate affinities. The predictability of deep-water kelp habitat and the discovery of expansive deep-water Galápagos kelp forests validate the extent of deep-water tropical kelp refugia, with potential implications for regional productivity and biodiversity, tropical food web ecology, and understanding of the resilience of tropical marine systems to climate change.

Temperature control of larval dispersal and the implications for marine ecology, evolution, and conservation.

Temperature controls the rate of fundamental biochemical processes and thereby regulates organismal attributes including development rate and survival. The increase in metabolic rate with temperature explains substantial among-species variation in life-history traits, population dynamics, and ecosystem processes. Temperature can also cause variability in metabolic rate within species. Here, we compare the effect of temperature on a key component of marine life cycles among a geographically and taxonomically diverse group of marine fish and invertebrates. Although innumerable lab studies document the negative effect of temperature on larval development time, little is known about the generality versus taxon-dependence of this relationship. We present a unified, parameterized model for the temperature dependence of larval development in marine animals. Because the duration of the larval period is known to influence larval dispersal distance and survival, changes in ocean temperature could have a direct and predictable influence on population connectivity, community structure, and regional-to-global scale patterns of biodiversity.