Dispersal connectivity and reserve selection for marine conservation

Although larval dispersal is crucial for the persistence of most marine populations, dispersal connectivity between sites is rarely considered in designing marine protected area networks. In particular the role of structural characteristics (known as topology) for the network of larval dispersal routes in the conservation of metapopulations has not been addressed. To determine reserve site configurations that provide highest persistence values with respect to their connectivity characteristics, we model nine connectivity topological models derived from graph theory in a demographic metapopulation model. We identify reserve site configurations that provide the highest persistence values for each of the metapopulation connectivity models. Except for the minimally connected and fully connected populations, we observed two general ‘rules of thumb’ for optimising the mean life time for all topological models: firstly place the majority of reserves, so that they are neighbours of each other, on the sites where the number of connections between the populations is highest (hub), secondly when the reserves have occupied the majority of the vertices in the hub, then select another area of high connectivity and repeat. If there are no suitable hubs remaining then distribute the remaining reserves to isolated locations optimising contact with non-reserved sites.     

Larval supply and juvenile recruitment of coral reef fishes to marine reserves and non-reserves of the upper Florida Keys,USA

For marine organisms, decoupling between the planktonic larval stage and the benthic-associated juvenile stage can lead to variable patterns of population replenishment, which have the potential to influence the effectiveness of marine reserves. We measured spatial and temporal variability in larval supply and recruitment of fishes to coral reefs of different protection levels and tested whether protection level influenced the relationship between supply and recruitment. We sampled pre-settlement larvae and newly settled recruits from four reefs (two reserves and two non-reserves) in the Florida Keys National Marine Sanctuary, USA. Replicate point measures of larval supply over 14 months and 17 monthly measurements of recruitment varied significantly among months and sites. Sites with the same protection level had significantly different patterns of larval supply as well as larval and recruit diversity, but recruitment magnitude differed only by protection level, where densities were greater at reserves. Differences in larval supply among sites included two particularly large peaks in larval abundance at one site, possibly associated with the observed passage of small-scale oceanographic features. To examine whether relationships between larval supply and recruitment varied by protection level, we selected one species that was present in both the light trap samples and the monthly recruitment surveys. Recruitment of the bicolor damselfish Stegastes partitus was significantly and positively related to larval supply at three of the four sites thus, protection level did not influence this linkage. Since local variability among sites can lead to spatial differences in population replenishment, characterization of larval supply and recruitment to potential marine reserve sites may help to identify optimal locations in a region and contribute to more effective reserve design.     

Evaluating the Effects of Three Forms of Marine Reserve on Northern Abalone Populations in British Columbia, Canada

Abstract: Marine reserves have been suggested as tools for assisting the management of fisheries by protecting vulnerable marine species from overexploitation. Although there is a theoretical basis for believing that marine reserves may serve as management tools, there are few marine reserves in the world in which to test their effectiveness. My research evaluated three forms of marine reserve on the south coast of Vancouver Island, British Columbia, Canada. I used northern abalone ( Haliotis kamtschatkana ), a severely depleted shellfish in this region, as an indicator of the effectiveness of the reserves. Abalone populations in eight sites receiving different degrees of spatial protection were counted and measured in situ during the spring of 1996 and 1997. In all sites with enforced harvest closures, populations of abalone were greater, and one site with nearly 40 years of protection had on average much larger (older) abalone. Reproductive output, as a function of abundance and size, was also greater in the enforced reserve areas. Larval dispersal from reserves, and hence the benefit to exploited areas, was not formally surveyed. Nevertheless, the results of my study, combined with knowledge of present abalone populations, life history, and regional hydrodynamics, suggest that establishment of reserves is justified in the absence of perfect knowledge of larval dispersal.     

Integrating larval connectivity into the marine conservation decision-making process across spatial scales

Larval dispersal connectivity is typically integrated into spatial conservation decisions at regional or national scales, but implementing agencies struggle with translating these methods to local scales. We used larval dispersal connectivity at regional (hundreds of kilometers) and local (tens of kilometers) scales to aid in design of networks of no-take reserves in Southeast Sulawesi, Indonesia. We used Marxan with Connectivity informed by biophysical larval dispersal models and remotely sensed coral reef habitat data to design marine reserve networks for 4 commercially important reef species across the region. We complemented regional spatial prioritization with decision trees that combined network-based connectivity metrics and habitat quality to design reserve boundaries locally. Decision trees were used in consensus-based workshops with stakeholders to qualitatively assess site desirability, and Marxan was used to identify areas for subsequent network expansion. Priority areas for protection and expected benefits differed among species, with little overlap in reserve network solutions. Because reef quality varied considerably across reefs, we suggest reef degradation must inform the interpretation of larval dispersal patterns and the conservation benefits achievable from protecting reefs. Our methods can be readily applied by conservation practitioners, in this region and elsewhere, to integrate connectivity data across multiple spatial scales.     

Marine Ecological Research in New Zealand: Developing Predictive Models through the Study of No-Take Marine Reserves

Abstract:  New Zealand established its first no-take marine reserve more than 25 years ago. Twenty no-take marine reserves have now been created, although few of these are considered comparable. We considered whether existing conceptual models of population and community structure based only on data from exploited systems lack the baseline information of natural states necessary to make accurate predictions for new reserves. Three of the oldest and best-studied reserves are situated on the northeastern coast of New Zealand. These reserves are considered broadly comparable replicates, and research has shown the recovery of previously exploited predator populations and the reestablishment of trophic controls over community structure and productivity. None of the major changes was predicted when the reserves were created. All the observations from and experimental tests of hypotheses in these three ecologically comparable reserves have provided predictive models for future reserves. Recent surveys in newly created reserves, however, suggest that these models are bioregion and habitat specific. In these new reserves the recovery of previously exploited predators was predicted but did not always occur. Where trends were correctly predicted, the speed and amplitude of the changes were not accurately predicted. Research in New Zealand suggests that it is not yet possible to predict explicit outcomes for newly created reserves and less possible to predict detailed results for systems of reserves. Results from a representative system of reserves, including all major habitats within all bioregions and broadly comparable reserves, are needed. Such a system will enable the range and variety of natural ecosystem dynamics to be investigated and provide the controls necessary to measure the effects of exploitation.     

Spatial variability in habitat associations of pre- and post-settlement stages of coral reef fishes at Ishigaki Island,Japan

Successful settlement of pelagic fish larvae into benthic juvenile habitats may be enhanced by a shortened settlement period, since it limits larval exposure to predation in the new habitat. Because the spatial distribution of marine fish larvae immediately prior to settlement versus during settlement was unknown, field experiments were conducted at Ishigaki Island (Japan) using light trap sampling and underwater visual belt transect surveys to investigate the spatial distribution patterns of selected pre- and post-settlement fishes (Acanthuridae, Pomacentridae, Chaetodonidae and Lethrinidae) among four habitats (seagrass bed, coral rubble, branching coral and tabular coral). The results highlighted two patterns: patterns 1, pre- and post-settlement individuals showing a ubiquitous distribution among the four habitats (Acanthuridae) and pattern 2, pre-settlement individuals distributed in all habitats, but post-settlement individuals restricted to coral (most species of Pomacentridae and Chaetodontidae) or seagrass habitats (Lethrinidae). The first pattern minimizes the transition time between the larval pelagic stage and acquisition of a benthic reef habitat, the latter leading immediately to a juvenile lifestyle. In contrast, the second pattern is characterized by high settlement habitat selectivity by larvae and/or differential mortality immediately after settlement.     

Effects of protected areas on survival of threatened gibbons in China

Establishing protected areas (PAs) is an essential strategy to reduce biodiversity loss. However, many PAs do not provide adequate protection due to poor funding, inadequate staffing and equipment, and ineffective management. As part of China's recent economic growth, the Chinese government has significantly increased investment in nature reserves over the past 20 years, providing a unique opportunity to evaluate whether PAs can protect threatened species effectively. We compiled data from published literature on populations of gibbons (Hylobatidae), a threatened taxon with cultural significance, that occurred in Chinese reserves after 1980. We evaluated the ability of these PAs to maintain gibbon habitat and populations by comparing forest cover and human disturbance between reserves and their surrounding areas and modeling the impact of reserve characteristics on gibbon population trends. We also assessed the perspective of reserve staff concerning PA management effectiveness through an online survey. Reserves effectively protected gibbon habitat by reducing forest loss and human disturbance; however, half the reserves lost their gibbon populations since being established. Gibbons were more likely to survive in reserves established more recently, at higher elevation, with less forest loss and lower human impact, and that have been relatively well studied. A larger initial population size in the 1980s was positively associated with gibbon persistence. Although staff of all reserves reported increased investment and improved management over the past 20–30 years, no relationship was found between management effectiveness and gibbon population trends. We suggest early and emphatic intervention is critical to stop population decline and prevent extinction.     

When are no-take zones an economically optimal fishery management strategy?

Discussions on the use of marine reserves (no-take zones) and, more generally, spatial management of fisheries are, for the most part, devoid of analyses that consider the ecological and economic effects simultaneously. To fill this gap, we develop a two-patch ecological-economic model to investigate the effects of spatial management on fishery profits. Because the fishery effects of spatial management depend critically on the nature of the ecological connectivity, our model includes both juvenile and adult movement, with density dependence in settlement differentiating the two types of dispersal. Rather than imposing a reserve on our system and measuring its effect on profits, we ask: "When does setting catch levels to maximize system-wide profits imply that a reserve should be created?" Closing areas to fishing is an economically optimal solution when the value derived from spillover from the reserve outweighs the value of fishing in the patch. The condition, while simple to state in summary form, is complex to interpret because it depends on the settlement success of the dispersing organisms, the nature of the costs of the fishing, the economic and ecological heterogeneity of the system, the discount rate, and growth characteristics of the fish population. The condition is more likely to be satisfied when the closed area is a net exporter of biomass and has higher costs of fishing, and for fish populations with density-independent settlement ("adult movement") than with density-dependent settlement ("larval dispersal"). Rather surprisingly, there are circumstances whereby closing low biological productivity areas, and even sometimes low cost areas to fish, can result in greater fishing profits than when both areas are open to fishing.     

Modeling the impacts of hunting on the population dynamics of red howler monkeys (Alouatta seniculus)

Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey (Alouatta seniculus) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km², increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.     

Reserve design to optimize functional connectivity and animal density

Ecological distance-based spatial capture–recapture models (SCR) are a promising approach for simultaneously estimating animal density and connectivity, both of which affect spatial population processes and ultimately species persistence. We explored how SCR models can be integrated into reserve-design frameworks that explicitly acknowledge both the spatial distribution of individuals and their space use resulting from landscape structure. We formulated the design of wildlife reserves as a budget-constrained optimization problem and conducted a simulation to explore 3 different SCR-informed optimization objectives that prioritized different conservation goals by maximizing the number of protected individuals, reserve connectivity, and density-weighted connectivity. We also studied the effect on our 3 objectives of enforcing that the space-use requirements of individuals be met by the reserve for individuals to be considered conserved (referred to as home-range constraints). Maximizing local population density resulted in fragmented reserves that would likely not aid long-term population persistence, and maximizing the connectivity objective yielded reserves that protected the fewest individuals. However, maximizing density-weighted connectivity or preemptively imposing home-range constraints on reserve design yielded reserves of largely spatially compact sets of parcels covering high-density areas in the landscape with high functional connectivity between them. Our results quantify the extent to which reserve design is constrained by individual home-range requirements and highlight that accounting for individual space use in the objective and constraints can help in the design of reserves that balance abundance and connectivity in a biologically relevant manner.     

Catastrophe management and inter-reserve distance for marine reserve networks

We consider the optimal spacing between marine reserves for maximising the viability of a species occupying a reserve network. The closer the networks are placed together, the higher the probability of colonisation of an empty reserve by an occupied reserve, thus increasing population viability. However, the closer the networks are placed together, the higher the probability that a catastrophe will cause extinction of the species in both reserves, thus decreasing population viability. Using a simple discrete-time Markov chain model for the presence or absence of the species in each reserve we determine the distance between the two reserves which provides the optimal trade-off between these processes, resulting in maximum viability of the species.     

Open and closed seascapes: where does habitat patchiness create populations with high fractions of self-recruitment?

Which populations are replenished primarily by immigrants (open) and which by local production (closed) remains an important question for management with implications for response to exploitation, protection, and disturbance. However, we lack methods for predicting population openness. Here, we develop a model for openness and show that considering habitat isolation explains the existence of surprisingly closed populations in high-dispersal species, including many marine organisms. Relatively closed populations are expected when patch spacing is more than twice the standard deviation of a species'. dispersal kernel. In addition, natural scales of habitat patchiness on coral reefs are sufficient to create both largely open and largely closed populations. Contrary to some previous interpretations, largely closed marine populations do not require mean dispersal distances that are unusually short, even for species with relatively long pelagic larval durations. We predict that habitat patchiness has strong control over population openness for many marine and terrestrial species with a highly dispersive life stage and relatively sedentary adults. This information can be used to make initial predictions about where populations will be more or less resilient to local exploitation and disturbance.     

A Decision Framework for the Adaptive Management of an Exploited Species with Implications for Marine Reserves

Abstract:  Marine reserves have both conservation and fishery benefits. Nevertheless, there are no general criteria about when and where to establish new reserves, how to evaluate their efficacy, and how to conduct adaptive management to achieve conservation goals. We applied a decision-theory framework to optimally allocate conservation resources between improving data on population status and establishing a reserve for species conservation. Our goal was to maximize reserve benefits given the constraints of a population growth rate that would permit sustainability of resources. We illustrate our decision framework with a retrospective analysis of a 7-year time series on abundance of the leopard grouper ( Mycteroperca rosacea ) in the Sea of Cortés, Mexico. We used the lower bound of the distribution of the population growth rate ( λ ) as a decision rule for determining how many years of monitoring are needed to detect reserve effects. We determined the minimum time frame needed to estimate λ based on a stated level of risk tolerance for four sites. As expected, the coefficient of variation for the λ declined with the number of years of data. This increased precision with additional years of data resulted from the high degree of annual variability in the system. Where populations were slow to respond to reserves, more data were needed to detect a positive λ value. For the leopard grouper case study, confidence in the estimate of λ increased with the number of years of data. Our decision framework may be used to identify the minimum number of years of data needed before a management decision about reserve establishment could be made that is reasonably likely to meet its management objectives.     

Oceanic variability and coastal topography shape genetic structure in a long-dispersing sea urchin

Understanding the scale of marine population connectivity is critical for the conservation and sustainable management of marine resources. For many marine species adults are benthic and relatively immobile, so patterns of larval dispersal and recruitment provide the key to understanding marine population connectivity. Contrary to previous expectations, recent studies have often detected unexpectedly low dispersal and fine-scale population structure in the sea, leading to a paradigm shift in how marine systems are viewed. Nonetheless, the link between fine-scale marine population structure and the underlying physical and biological processes has not been made. Here we show that patterns of genetic structure and population connectivity in the broadcast-spawning and long-distance dispersing sea urchin Centrostephanus rodgersii are influenced by physical oceanographic and geographic variables. Despite weak genetic differentiation and no isolation-by-distance over thousands of kilometers among samples from eastern Australia and northern New Zealand, fine-scale genetic structure was associated with sea surface temperature (SST) variability and geography along the southeastern Australian coast. The zone of high SST variability is characterized by periodic shedding of eddies from the East Australian Current, and we suggest that ocean current circulation may, through its influence on larval transport and recruitment, interact with the genetic consequences of large variance in individual reproductive success to generate patterns of fine-scale patchy genetic structure. If proven consistent across species, our findings suggest that the optimal scale for fisheries management and reserve design should vary among localities in relation to regional oceanographic variability and coastal geography.     

Fishing the line near marine reserves in single and multispecies fisheries.

Throughout the world "fishing the line" is a frequent harvesting tactic in communities where no-take marine reserves are designated. This practice of concentrating fishing effort at the boundary of a marine reserve is predicated upon the principle of spillover, the net export of stock from the marine reserve to the surrounding unprotected waters. We explore the consequences and optimality of fishing the line using a spatially explicit theoretical model. We show that fishing the line: (1) is part of the optimal effort distribution near no-take marine reserves with mobile species regardless of the cooperation level among harvesters; (2) has a significant impact on the spatial patterns of catch per unit effort (CPUE) and fish density both within and outside of the reserve; and (3) can enhance total population size and catch simultaneously under a limited set of conditions for overexploited populations. Additionally, we explore the consequences of basing the spatial distribution of fishing effort for a multispecies fishery upon the optimality of the most mobile species that exhibits the greatest spillover. Our results show that the intensity of effort allocated to fishing the line should instead be based upon more intermediate rates of mobility within the targeted community. We conclude with a comparison between model predictions and empirical findings from a density gradient study of two important game fish in the vicinity of a no-take marine-life refuge on Santa Catalina Island, California (USA). These results reveal the need for empirical studies to account for harvester behavior and suggest that the implications of spatial discontinuities such as fishing the line should be incorporated into marine-reserve design.     

Assessing the current state of ecological connectivity in a large marine protected area system

The establishment of marine protected areas (MPAs) is a critical step in ensuring the continued persistence of marine biodiversity. Although the area protected in MPAs is growing, the movement of individuals (or larvae) among MPAs, termed connectivity, has only recently been included as an objective of many MPAs. As such, assessing connectivity is often neglected or oversimplified in the planning process. For promoting population persistence, it is important to ensure that protected areas in a system are functionally connected through dispersal or adult movement. We devised a multi-species model of larval dispersal for the Australian marine environment to evaluate how much local scale connectivity is protected in MPAs and determine whether the extensive system of MPAs truly functions as a network. We focused on non-migratory species with simplified larval behaviors (i.e., passive larval dispersal) (e.g., no explicit vertical migration) as an illustration. Of all the MPAs analyzed (approximately 2.7 million km²), outside the Great Barrier Reef and Ningaloo Reef, <50% of MPAs (46-80% of total MPA area depending on the species considered) were functionally connected. Our results suggest that Australia's MPA system cannot be referred to as a single network, but rather a collection of numerous smaller networks delineated by natural breaks in the connectivity of reef habitat. Depending on the dispersal capacity of the taxa of interest, there may be between 25 and 47 individual ecological networks distributed across the Australian marine environment. The need to first assess the underlying natural connectivity of a study system prior to implementing new MPAs represents a key research priority for strategically enlarging MPA networks. Our findings highlight the benefits of integrating multi-species connectivity into conservation planning to identify opportunities to better incorporate connectivity into the design of MPA systems and thus to increase their capacity to support long-term, sustainable biodiversity outcomes.     

Barnacle larvae actively select flow environments supporting post-settlement growth and survival

Many marine dispersive propagules select specific settlement sites based on a range of environmental cues. However, the link between larval choice and post-settlement growth and survival is still poorly understood. Here we show that cypris larvae of the barnacle Balanus improvisus actively reject surfaces exposed to local flow speeds exceeding 5-10 cm/s. Field experiments show that post-settlement growth and survival decline in freestream flows above 15 cm/s. Moreover, studies in flume flow at local speeds exceeding 10 cm/s reveal that early juveniles show reduced feeding rates caused by deformation of the cirral fan, reduced retention efficiency, and a decrease in time spent feeding. We conclude that cypris larvae actively reject flow environments that will be suboptimal for suspension feeding in the early post-settlement phase. Our study suggests that larval choice can be adaptively connected to a specific part of the life cycle, in this case the very sensitive time after metamorphosis.     

Poaching, enforcement, and the efficacy of marine reserves.

Marine reserves are promoted as an effective supplement to traditional fishery management techniques of harvest quotas and effort limitation. However, quantitative fishery models have ignored the impact of noncompliance (poaching). Here we link a model of a harvested fish population to a game-theoretic representation of fisherman behavior to quantify the effect of poaching on fishery yield and the enforcement effort required to maintain any desired level of reserve effectiveness. Although higher fish densities inside reserves will typically entice fishermen to poach, we show that the initial investment in enforcement efforts provides the greatest return on maintaining the benefits of the reserve to the fishery. Furthermore, we find that poaching eliminates the positive effect of fish dispersal on yield that is predicted by traditional models that ignore fisherman behavior. Our results broaden a fundamental insight from previous models of marine reserves, the effective equivalence of the harvest quota and reserve fraction, to the more realistic scenario in which fishermen attempt to maximize their economic payoffs.     

Role of Patch Size, Disease, and Movement in Rapid Extinction of Bighorn Sheep

Abstract: The controversy (  Berger 1990, 1999 ; Wehausen 1999 ) over rapid extinction in bighorn sheep ( Ovis canadensis ) has focused on population size alone as a correlate to persistence time. We report on the persistence and population performance of 24 translocated populations of bighorn sheep. Persistence in these sheep was strongly correlated with larger patch sizes, greater distance to domestic sheep, higher population growth rates, and migratory movements, as well as to larger population sizes. Persistence was also positively correlated with larger average home-range size ( p = 0.058, n = 10 translocated populations) and home-range size of rams ( p = 0.087, n = 8 translocated populations). Greater home-range size and dispersal rates of bighorn sheep were positively correlated to larger patches. We conclude that patch size and thus habitat carrying capacity, not population size per se, is the primary correlate to both population performance and persistence. Because habitat carrying capacity defines the upper limit to population size, clearly the amount of suitable habitat in a patch is ultimately linked to population size. Larger populations (250+ animals) were more likely to recover rapidly to their pre-epizootic survey number following an epizootic ( p = 0.019), although the proportion of the population dying in the epizootic also influenced the probability of recovery ( p = 0.001). Expensive management efforts to restore or increase bighorn sheep populations should focus on large habitat patches located ≥23 km from domestic sheep, and less effort should be expended on populations in isolated, small patches of habitat.