Scott C. Burgess

Beyond connectivity: how empirical methods can quantify population persistence to improve marine protected-area design

Demographic connectivity is a fundamental process influencing the dynamics and persistence of spatially structured populations. Consequently, quantifying connectivity is essential for properly designing networks of protected areas so that they achieve their core ecological objective of maintaining population persistence. Recently, many empirical studies in marine systems have provided essential, and historically challenging to obtain, data on patterns of larval dispersal and export from marine protected areas (MPAs). Here, we review the empirical studies that have directly quantified the origins and destinations of individual larvae and assess those studies relevance to the theory of population persistence and MPA design objectives. We found that empirical studies often do not measure or present quantities that are relevant to assessing population persistence, even though most studies were motivated or contextualized by MPA applications. Persistence of spatial populations, like nonspatial populations, depends on replacement, whether individuals reproduce enough in their lifetime to replace themselves. In spatial populations, one needs to account for the effect of larval dispersal on future recruitment back to the local population through local retention and other connectivity pathways. The most commonly reported descriptor of larval dispersal was the fraction of recruitment from local origin (self-recruitment). Self-recruitment does not inform persistence-based MPA design because it is a fraction of those arriving, not a fraction of those leaving (local retention), so contains no information on replacement. Some studies presented connectivity matrices, which can inform assessments of persistence with additional knowledge of survival and fecundity after recruitment. Some studies collected data in addition to larval dispersal that could inform assessments of population persistence but which were not presented in that way. We describe how three pieces of empirical information are needed to fully describe population persistence in a network of MPAs: (1) lifetime fecundity, (2) the proportion of larvae that are locally retained (or the full connectivity matrix), and (3) survival rate after recruitment. We conclude by linking theory and data to provide detailed guidance to empiricists and practitioners on field sampling design and data presentation that better informs the MPA objective of population persistence.

When is dispersal for dispersal? Unifying marine and terrestrial perspectives

Recent syntheses on the evolutionary causes of dispersal have focused on dispersal as a direct adaptation, but many traits that influence dispersal have other functions, raising the question: when is dispersal ‘for’ dispersal? We review and critically evaluate the ecological causes of selection on traits that give rise to dispersal in marine and terrestrial organisms. In the sea, passive dispersal is relatively easy and specific morphological, behavioural, and physiological adaptations for dispersal are rare. Instead, there may often be selection to limit dispersal. On land, dispersal is relatively difficult without specific adaptations, which are relatively common. Although selection for dispersal is expected in both systems and traits leading to dispersal are often linked to fitness, systems may differ in the extent to which dispersal in nature arises from direct selection for dispersal or as a by‐product of selection on traits with other functions. Our analysis highlights incompleteness of theories that assume a simple and direct relationship between dispersal and fitness, not just insofar as they ignore a vast array of taxa in the marine realm, but also because they may be missing critically important effects of traits influencing dispersal in all realms.

Assessing strategies to minimize unintended fitness consequences of aquaculture on wild populations.

Artificial propagation programs focused on production, such as commercial aquaculture or forestry, entail strong domestication selection. Spillover from such programs can cause unintended fitness and demographic consequences for wild conspecifics. The range of possible management practices to minimize such consequences vary in their control of genetic and demographic processes. Here, we use a model of coupled genetic and demographic dynamics to evaluate alternative management approaches to minimizing unintended consequences of aquaculture escapees. We find that, if strong natural selection occurs between escape and reproduction, an extremely maladapted (i.e., nonlocal‐origin, highly domesticated) stock could have fitness consequences analogous to a weakly diverged cultured stock; otherwise, wild population fitness declines with increasing maladaptation in the cultured stock. Reducing escapees through low‐level leakage is more effective than reducing an analogous number of escapees from large, rare pulses. This result arises because low‐level leakage leads to the continual lowering of wild population fitness and subsequent increased proportional contribution of maladapted cultured escapees to the total population. Increased sterilization efficacy can cause rapid, nonlinear reductions in unintended fitness consequences. Finally, sensitivity to the stage of escape indicates a need for improved monitoring data on how the number of escapees varies across life cycle stages.

Deconstructing environmental predictability: seasonality, environmental colour and the biogeography of marine life histories

Environmental predictability is predicted to shape the evolution of life histories. Two key types of environmental predictability, seasonality and environmental colour, may influence life-history evolution independently but formal considerations of both and how they relate to life history are exceedingly rare. Here, in a global biogeographical analysis of over 800 marine invertebrates, we explore the relationships between both forms of environmental predictability and three fundamental life-history traits: location of larval development (aplanktonic vs. planktonic), larval developmental mode (feeding vs. non-feeding) and offspring size. We found that both dispersal potential and offspring size related to environmental predictability, but the relationships depended on both the environmental factor as well as the type of predictability. Environments that were more seasonal in food availability had a higher prevalence of species with a planktonic larval stage. Future studies should consider both types of environmental predictability as each can strongly affect life-history evolution.

Size-dependent physiological responses of the branching coral Pocillopora verrucosa to elevated temperature and PCO2

ABSTRACT Body size has large effects on organism physiology, but these effects remain poorly understood in modular animals with complex morphologies. Using two trials of a ∼24u2005day experiment conducted in 2014 and 2015, we tested the hypothesis that colony size of the coral Pocillopora verrucosa affects the response of calcification, aerobic respiration and gross photosynthesis to temperature (∼26.5 and ∼29.7°C) and PCO2 (∼40 and ∼1000u2005µatm). Large corals calcified more than small corals, but at a slower size-specific rate; area-normalized calcification declined with size. Whole-colony and area-normalized calcification were unaffected by temperature, PCO2, or the interaction between the two. Whole-colony respiration increased with colony size, but the slopes of these relationships differed between treatments. Area-normalized gross photosynthesis declined with colony size, but whole-colony photosynthesis was unaffected by PCO2, and showed a weak response to temperature. When scaled up to predict the response of large corals, area-normalized metrics of physiological performance measured using small corals provide inaccurate estimates of the physiological performance of large colonies. Together, these results demonstrate the importance of colony size in modulating the response of branching corals to elevated temperature and high PCO2. Summary: Physiological performance of large corals is not accurately estimated by scaling values from small corals, indicating the importance of colony size in the response of branching corals to elevated temperature and PCO2.

Global change, life-history complexity and the potential for evolutionary rescue

Most organisms have complex life cycles, and in marine taxa, larval life‐history stages tend to be more sensitive to environmental stress than adult (reproductive) life‐history stages. While there are several models of stage‐specific adaptation across the life history, the extent to which differential sensitivity to environmental stress (defined here as reductions in absolute fitness across the life history) affects the tempo of adaptive evolution to change remains unclear. We used a heuristic model to explore how commonly observed features associated with marine complex life histories alter a populations capacity to cope with environmental change. We found that increasing the complexity of the life history generally reduces the evolutionary potential of taxa to cope with environmental change. Our model also predicted that genetic correlations in stress tolerance between stages, levels of genetic variance in each stage, and the relative plasticity of different stages, all interact to affect the maximum rate of environmental change that will permit species persistence. Our results suggest that marine organisms with complex life cycles are particularly vulnerable to anthropogenic global change, but we lack empirical estimates of key parameters for most species.

LOCAL ADAPTATION WHEN COMPETITION DEPENDS ON PHENOTYPIC SIMILARITY

Recent work incorporating demographic–genetic interactions indicates the importance of population size, gene flow, and selection in influencing local adaptation. This work typically assumes that density‐dependent survival affects individuals equally, but individuals in natural population rarely compete equally. Among‐individual differences in resource use generate stronger competition between more similar phenotypes (frequency‐dependent competition) but it remains unclear how this additional form of selection changes the interactions between population size, gene flow, and local stabilizing selection. Here, we integrate migration–selection dynamics with frequency‐dependent competition. We developed a coupled demographic‐quantitative genetic model consisting of two patches connected by dispersal and subject to local stabilizing selection and competition. Our model shows that frequency‐dependent competition slightly increases local adaptation, greatly increases genetic variance within patches, and reduces the amount that migration depresses population size, despite the increased genetic variance load. The effects of frequency‐dependence depend on the strength of divergent selection, trait heritability, and when mortality occurs in the life cycle in relation to migration and reproduction. Essentially, frequency‐dependent competition reduces the density‐dependent interactions between migrants and residents, the extent to which depends on how different and common immigrants are compared to residents. Our results add new dynamics that illustrate how competition can alter the effects of gene flow and divergent selection on local adaptation and population carrying capacities.