Steven D. Gaines

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.

Can Catch Shares Prevent Fisheries Collapse

Recent reports suggest that most of the worlds commercial fisheries could collapse within decades. Although poor fisheries governance is often implicated, evaluation of solutions remains rare. Bioeconomic theory and case studies suggest that rights-based catch shares can provide individual incentives for sustainable harvest that is less prone to collapse. To test whether catch-share fishery reforms achieve these hypothetical benefits, we have compiled a global database of fisheries institutions and catch statistics in 11,135 fisheries from 1950 to 2003. Implementation of catch shares halts, and even reverses, the global trend toward widespread collapse. Institutional change has the potential for greatly altering the future of global fisheries.

Species diversity: from global decreases to local increases

Current patterns of global change can strongly affect biodiversity at global, regional and local scales. At global scales, habitat destruction and the introduction of exotic species are contributing to declines in species diversity. At regional and local scales, evidence for declines in diversity is mixed, and recent work suggests that diversity might commonly be increasing. In spite of these trends, considerable research continues to consider explicitly the effects of declines in diversity on processes that operate at regional and local scales (such as ecosystem functioning), without explicitly considering the converse set of questions, namely the effects of increases in diversity. Here, we examine evidence that indicates how species diversity is changing across spatial scales and argue that global decreases in diversity are commonly contrasted by increases in diversity at regional and local 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.

Designing marine reserve networks for both conservation and fisheries management

Marine protected areas (MPAs) that exclude fishing have been shown repeatedly to enhance the abundance, size, and diversity of species. These benefits, however, mean little to most marine species, because individual protected areas typically are small. To meet the larger-scale conservation challenges facing ocean ecosystems, several nations are expanding the benefits of individual protected areas by building networks of protected areas. Doing so successfully requires a detailed understanding of the ecological and physical characteristics of ocean ecosystems and the responses of humans to spatial closures. There has been enormous scientific interest in these topics, and frameworks for the design of MPA networks for meeting conservation and fishery management goals are emerging. Persistent in the literature is the perception of an inherent tradeoff between achieving conservation and fishery goals. Through a synthetic analysis across these conservation and bioeconomic studies, we construct guidelines for MPA network design that reduce or eliminate this tradeoff. We present size, spacing, location, and configuration guidelines for designing networks that simultaneously can enhance biological conservation and reduce fishery costs or even increase fishery yields and profits. Indeed, in some settings, a well-designed MPA network is critical to the optimal harvest strategy. When reserves benefit fisheries, the optimal area in reserves is moderately large (mode ≈30%). Assessing network design principals is limited currently by the absence of empirical data from large-scale networks. Emerging networks will soon rectify this constraint.

Species invasions and extinction: The future of native biodiversity on islands

Predation by exotic species has caused the extinction of many native animal species on islands, whereas competition from exotic plants has caused few native plant extinctions. Exotic plant addition to islands is highly nonrandom, with an almost perfect 1 to 1 match between the number of naturalized and native plant species on oceanic islands. Here, we evaluate several alternative implications of these findings. Does the consistency of increase in plant richness across islands imply that a saturation point in species richness has been reached? If not, should we expect total plant richness to continue to increase as new species are added? Finally, is the rarity of native plant extinctions to date a misleading measure of the impact of past invasions, one that hides an extinction debt that will be paid in the future? By analyzing historical records, we show that the number of naturalized plant species has increased linearly over time on many individual islands. Further, the mean ratio of naturalized to native plant species across islands has changed steadily for nearly two centuries. These patterns suggest that many more species will become naturalized on islands in the future. We also discuss how dynamics of invasion bear upon alternative saturation scenarios and the implications these scenarios have for the future retention or extinction of native plant species. Finally, we identify invasion-motivated research gaps (propagule pressure, time-lags to extinction, abundance shifts, and loss of area) that can aid in forecasting extinction and in developing a more comprehensive theory of species extinctions.

An index to assess the health and benefits of the global ocean

The ocean plays a critical role in supporting human well-being, from providing food, livelihoods and recreational opportunities to regulating the global climate. Sustainable management aimed at maintaining the flow of a broad range of benefits from the ocean requires a comprehensive and quantitative method to measure and monitor the health of coupled human–ocean systems. We created an index comprising ten diverse public goals for a healthy coupled human–ocean system and calculated the index for every coastal country. Globally, the overall index score was 60 out of 100 (range 36–86), with developed countries generally performing better than developing countries, but with notable exceptions. Only 5% of countries scored higher than 70, whereas 32% scored lower than 50. The index provides a powerful tool to raise public awareness, direct resource management, improve policy and prioritize scientific research.

Species Invasions Exceed Extinctions on Islands Worldwide: A Comparative Study of Plants and Birds

Species richness is decreasing at a global scale. At subglobal scales, that is, within any defined area less extensive than the globe, species richness will increase when the number of nonnative species becoming naturalized is greater than the number of native species becoming extinct. Determining whether this has occurred is usually difficult because detailed records of species extinctions and naturalizations are rare; these records often exist, however, for oceanic islands. Here we show that species richness on oceanic islands has remained relatively unchanged for land birds, with the number of naturalizations being roughly equal to the number of extinctions, and has increased dramatically for vascular plants, with the number of naturalizations greatly exceeding the number of extinctions. In fact, for plants, the net number of species on islands has approximately doubled. We show further that these patterns are robust to differences in the history of human occupation of these islands and to the possibility of undocumented species extinctions. These results suggest that species richness may be increasing at subglobal scales for many groups and that future research should address what consequences this may have on ecological processes.

SOME IMPLICATIONS OF DIRECT POSITIVE INTERACTIONS FOR COMMUNITY SPECIES DIVERSITY

Direct positive interactions (mutualisms and commensalisms) are generally accepted as important processes in communities. They appear to be most common in environments with relatively high physical disturbance, stress, or predation, where associated species can increase the growth and survival of other species unable to survive in isolation. Although ecologists have documented direct positive interactions among species for decades, there is less known about how these interactions affect community species diversity patterns. In this paper, we present a qualitative theoretical model that considers how direct positive interactions affect community species diversity. The model uses, as its basis, familiar unimodel species diversity models (i.e., “compensatory mortality” and “intermediate disturbance” hypothesis) to understand where direct positive interactions are likely to be important. Initially, it predicts that direct positive interactions increase species diversity by facilitating species that might not normally survive under very high physical disturbance, stress, or predation. In addition, it suggests that, under intermediate physical disturbance, stress, or predation, facilitator species that might normally be competitively excluded are released from competition. We suggest that facilitator species may then create new interaction webs that would not be possible in their absence. To illustrate these ideas, we describe a case study taken from a New England salt marsh community where a gradient in physical conditions occurs. In this community, direct positive interactions, and their indirect effects, are predicted to increase the species diversity by at least 35%. This empirical case study and model show that by incorporating direct positive interactions into ecological experiments and theory, it is possible to expand our understanding of the mechanisms responsible for community species diversity patterns.

POPULATION MODELS FOR MARINE RESERVE DESIGN: A RETROSPECTIVE AND PROSPECTIVE SYNTHESIS

We synthesize results from existing models of marine reserves to identify key theoretical issues that appear to be well understood, as well as issues in need of further exploration. Models of marine reserves are relatively new in the scientific literature; 32 of the 34 theoretical papers we reviewed were published after 1990. These models have focused primarily on questions concerning fishery management at the expense of other objectives such as conservation, scientific understanding, recreation, education, and tourism. Roughly one-third of the models analyze effects on cohorts while the remaining models have some form of complete population dynamics. Few models explicitly include larval dispersal. In a fisheries context, the primary conclusion drawn by many of the complete population models is that reserves increase yield when populations would otherwise be overfished. A second conclusion, resulting primarily from single-cohort models, is that reserves will provide fewer benefits for species with greater adult rates of movement. Although some models are beginning to yield information on the spatial configurations of reserves required for populations with specific dispersal distances to persist, it remains an aspect of reserve design in need of further analysis. Other outstanding issues include the effects of (1) particular forms of density dependence, (2) multispecies interactions, (3) fisher behavior, and (4) effects of concentrated fishing on habitat. Model results indicate that marine reserves could play a beneficial role in the protection of marine systems against overfishing. Additional modeling and analysis will greatly improve prospects for a better understanding of the potential of marine reserves for conserving biodiversity.