Brian P. Kinlan

PROPAGULE DISPERSAL IN MARINE AND TERRESTRIAL ENVIRONMENTS: A COMMUNITY PERSPECTIVE

Studies in terrestrial systems suggest that long-distance propagule dispersal is important for landscape pattern and dynamics, but largely inconsequential for local demography. By contrast, in marine systems, dispersal at regional scales may drive local dynamics, because many species may have large mean dispersal distances. To assess var- iation in marine dispersal scales, we estimated mean dispersal distances from genetic iso- lation-by-distance slopes. Estimates ranged widely, from a few meters to hundreds of kilometers. Dispersal differed among taxonomic groups (macroalgae, invertebrates, and fish) and among species in different functional groups (e.g., producers and herbivores). Differences in dispersal scale have important implications for marine community dynamics, reserve design, responses to large-scale perturbations, and evolution of interacting species. To place genetic estimates of marine dispersal in context, we compared them to other measures of dispersal in the ocean and to estimates of dispersal on land. Maximum scales of dispersal by sedentary marine species exceeded maximum estimates of terrestrial plant dispersal by at least one to two orders of magnitude. Direct and genetic estimates of terrestrial plant dispersal were comparable to estimates of marine plant dispersal. Rates of marine macroalgal range expansion, however, far exceeded spread rates of terrestrial plants. Terrestrial plant spread rates were more similar to those of short-dispersing marine organ- isms that lack secondary dispersal by drifting adults. Genetic estimates of dispersal by different functional groups suggest that herbivores typically disperse much farther than their plant resources both on land and in the sea, although the timing, frequency, and consequences of dispersal may differ in the two systems. Terrestrial herbivores have more flexible dispersal behavior than marine organisms that disperse each generation by plank- tonic transport of larvae. Our results validate some long-standing views about the greater dispersal potential of species in the ocean, but also highlight the extreme heterogeneity in dispersal scale among marine species. As a result, development of a community perspective on marine connectivity will require consideration of multiple dispersal mechanisms and scales.

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.

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 km2 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.

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.

Exploration of the Canyon-Incised Continental Margin of the Northeastern United States Reveals Dynamic Habitats and Diverse Communities

The continental margin off the northeastern United States (NEUS) contains numerous, topographically complex features that increase habitat heterogeneity across the region. However, the majority of these rugged features have never been surveyed, particularly using direct observations. During summer 2013, 31 Remotely-Operated Vehicle (ROV) dives were conducted from 494 to 3271 m depth across a variety of seafloor features to document communities and to infer geological processes that produced such features. The ROV surveyed six broad-scale habitat features, consisting of shelf-breaching canyons, slope-sourced canyons, inter-canyon areas, open-slope/landslide-scar areas, hydrocarbon seeps, and Mytilus Seamount. Four previously unknown chemosynthetic communities dominated by Bathymodiolus mussels were documented. Seafloor methane hydrate was observed at two seep sites. Multivariate analyses indicated that depth and broad-scale habitat significantly influenced megafaunal coral (58 taxa), demersal fish (69 taxa), and decapod crustacean (34 taxa) assemblages. Species richness of fishes and crustaceans significantly declined with depth, while there was no relationship between coral richness and depth. Turnover in assemblage structure occurred on the middle to lower slope at the approximate boundaries of water masses found previously in the region. Coral species richness was also an important variable explaining variation in fish and crustacean assemblages. Coral diversity may serve as an indicator of habitat suitability and variation in available niche diversity for these taxonomic groups. Our surveys added 24 putative coral species and three fishes to the known regional fauna, including the black coral Telopathes magna, the octocoral Metallogorgia melanotrichos and the fishes Gaidropsarus argentatus, Guttigadus latifrons, and Lepidion guentheri. Marine litter was observed on 81% of the dives, with at least 12 coral colonies entangled in debris. While initial exploration revealed the NEUS region to be both geologically dynamic and biologically diverse, further research into the abiotic conditions and the biotic interactions that influence species abundance and distribution is needed.

A Metapopulation Perspective on the Patch Dynamics of Giant Kelp in Southern California

Publisher Summary This chapter examines the patch dynamics of Macrocystis in Southern California from a metapopulation perspective and begins by synthesizing existing and new information pertaining to the metapopulation structure and dynamics of Macrocystis in the Southern California Bight. The biological and physical factors that affect colonization are reviewed, and a new empirical and theoretical estimate of spore dispersal distance for varying oceanic conditions is presented. The average kelp bed in Southern California appears to be connected to one to three neighboring kelp beds via spore dispersal for a relatively wide range of oceanographic conditions. More importantly, connectivity, even among nearby patches, seems to be mediated by spores that travel far beyond the median dispersal distance. The persistence of a giant kelp metapopulation depends on the tails of the dispersal curve and cannot be predicted simply from the average or median dispersal distance. Patch size, fecundity, and proximity to neighboring patches undoubtedly exert strong influences on the level of connectivity among patches. Compared with many terrestrial habitats, the aqueous medium through which kelp propagules disperse is relatively unstructured. Moreover, unlike passively dispersed kelp spores and drifters, immigrants in many terrestrial metapopulations exhibit complex behavioral interactions with the heterogenous landscape that influence connectivity between patches. The strong dependence of giant kelp connectivity and patch dynamics on environmental factors, such as geomorphology and oceanography, suggests that the metapopulation structure of this species is likely to differ substantially among regions.

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.