Observational evidence that maladaptive gene flow reduces patch occupancy in a wild insect metapopulation

Theory predicts that dispersal throughout metapopulations has a variety of consequences for the abundance and distribution of species. Immigration is predicted to increase abundance and habitat patch occupancy, but gene flow can have both positive and negative demographic consequences. Here, we address the eco‐evolutionary effects of dispersal in a wild metapopulation of the stick insect Timema cristinae, which exhibits variable degrees of local adaptation throughout a heterogeneous habitat patch network of two host‐plant species. To disentangle the ecological and evolutionary contributions of dispersal to habitat patch occupancy and abundance, we contrasted the effects of connectivity to populations inhabiting conspecific host plants and those inhabiting the alternate host plant. Both types of connectivity should increase patch occupancy and abundance through increased immigration and sharing of beneficial alleles through gene flow. However, connectivity to populations inhabiting the alternate host‐plant species may uniquely cause maladaptive gene flow that counters the positive demographic effects of immigration. Supporting these predictions, we find the relationship between patch occupancy and alternate‐host connectivity to be significantly smaller in slope than the relationship between patch occupancy and conspecific‐host connectivity. Our findings illustrate the ecological and evolutionary roles of dispersal in driving the distribution and abundance of species.     

The strength of migratory connectivity for birds en route to breeding through the Gulf of Mexico

The strength of migratory connectivity is a measure of the cohesion of populations among phases of the annual cycle, including breeding, migration, and wintering. Many Nearctic‐Neotropical species have strong migratory connectivity between breeding and wintering phases of the annual cycle. It is less clear if this strength persists during migration when multiple endogenous and exogenous factors may decrease the cohesion of populations among routes or through time along the same routes. We sampled three bird species, American redstart Setophaga ruticilla, ovenbird Seiurus aurocapilla, and wood thrush Hylocichla mustelina, during spring migration through the Gulf of Mexico region to test if breeding populations differentiate spatially among migration routes or temporally along the same migration routes and the extent to which within‐population timing is a function of sex, age, and carry‐over from winter habitat, as measured by stable carbon isotope values in claws (δ¹³C). To make quantitative comparisons of migratory connectivity possible, we developed and used new methodology to estimate the strength of migratory connectivity (MC) from probabilistic origin assignments identified using stable hydrogen isotopes in feathers (δ²H). We found support for spatial differentiation among routes by American redstarts and ovenbirds and temporal differentiation along routes by American redstarts. After controlling for breeding origin, the timing of American redstart migration differed among ages and sexes and ovenbird migration timing was influenced by carry‐over from winter habitat. The strength of migratory connectivity did not differ among the three species, with each showing weak breeding‐to‐spring migration MC relative to prior assessments of breeding‐wintering connectivity. Our work begins to fill an essential gap in methodology and understanding of the extent to which populations remain together during migration, information critical for a full annual cycle perspective on the population dynamics and conservation of migratory animals.     

Is realised connectivity among populations of aquatic fauna predictable from potential connectivity?

1. Effective management of aquatic fauna requires knowledge of the ways in which populations in different catchments and sub‐catchments are connected. A powerful way to estimate this is using genetic markers, which provide information on the average amount of genetic connectivity among populations over generations. Although many studies of genetic connectivity have appeared in the literature, there are innumerable species that have not been studied. 2. This study explores whether it is possible to make broad generalisations about population connectivity, based on readily available information in the form of species life history and architecture of the aquatic habitat. 3. A number of models have been proposed to explain the pattern of connectivity shown by aquatic species with different life‐history characteristics, for example, the stream hierarchy model, Isolation by Distance, the Death Valley Model, the headwater model and panmixia. 4. In this study, we propose a dichotomous key to assign species to different models of potential connectivity. The key is based on a few very simple questions about the life history of the species and the geographical arrangement of study sites. We then assessed the performance of the key with 109 data sets of Australian fish and macroinvertebrates, using genetic data to provide an estimate of realised connectivity. 5. The realised connectivity fitted the proposed potential connectivity model in over 70% of cases, and we suggest this might be a useful initial approach for managers where empirical data are lacking.     

Grassland connectivity explains entomophilous plant species assemblages in an agricultural landscape of the Pampa Region,Argentina

The Pampa grassland of Argentina is one of the most highly threatened biomes in the world. A high proportion of the original grassland cover has been transformed into land for agriculture or degraded. In the southern part of the region, fragmented semi‐natural grasslands over exposed rock still persist and connectivity between them is assumed to be crucial for maintaining viable populations. We quantified overall connectivity of grassland patches in a sector of the Southern Pampa region, and investigated the degree to which landscape connectivity explains entomophilous plant species assemblages in a subset of patches. We characterized each of the 301 patches in the landscape by their degree of intra‐patch and inter‐patch connectivity based on graph theory, and considering threshold dispersal distances from 100 to 1000 m. We surveyed entomophilous plant species in 39 grassland patches and classified the species in three categories (annual herbs, perennial herbs and shrubs) considering their different growth form and longevity. The influence of connectivity variables on entomophilous plant species assemblages variation was explored using Canonical Correspondence Analysis. Although grassland patches were poorly connected at all threshold distances, some of them were found to be critical for global connectivity. Connectivity significantly explained total, annual‐biennial and shrub assemblages for all threshold dispersal distances (6–13% of total variation). Variation in annual species assemblages was associated with intra‐patch and inter‐patch connectivity at short distance (100 m), while variation in shrub species assemblages was explained by intra‐patch and inter‐patch connectivity for distances between 100 m and 1000 m. This study evidenced the low connectivity of the study system, allowed the identification of critical areas for conservation, and provided valuable information to develop management strategies in increasingly human‐dominated landscapes.     

Predicting global population connectivity and targeting conservation action for snow leopard across its range

Movements of individuals within and among populations help to maintain genetic variability and population viability. Therefore, understanding landscape connectivity is vital for effective species conservation. The snow leopard is endemic to mountainous areas of central Asia and occurs within 12 countries. We assess potential connectivity across the species’ range to highlight corridors for dispersal and genetic flow between populations, prioritizing research and conservation action for this wide‐ranging, endangered top‐predator. We used resistant kernel modeling to assess snow leopard population connectivity across its global range. We developed an expert‐based resistance surface that predicted cost of movement as functions of topographical complexity and land cover. The distribution of individuals was simulated as a uniform density of points throughout the currently accepted global range. We modeled population connectivity from these source points across the resistance surface using three different dispersal scenarios that likely bracket the lifetime movements of individual snow leopard: 100 km, 500 km and 1000 km. The resistant kernel models produced predictive surfaces of dispersal frequency across the snow leopard range for each distance scenario. We evaluated the pattern of connectivity in each of these scenarios and identified potentially important movement corridors and areas where connectivity might be impeded. The models predicted two regional populations, in the north and south of the species range respectively, and revealed a number of potentially important connecting areas. Discrepancies between model outputs and observations highlight unsurveyed areas of connected habitat that urgently require surveying to improve understanding of the global distribution and ecology of snow leopard, and target land management actions to prevent population isolation. The connectivity maps provide a strong basis for directed research and conservation action, and usefully direct the attention of policy makers.     

Migratory connectivity in a declining bird species: using feather isotopes to inform demographic modelling

Aim Conservation programmes for endangered migratory species or populations require locating and evaluating breeding, stopover and wintering areas. We used multiple stable isotopes in two endangered European populations of wrynecks, Jynx torquilla L., to locate wintering regions and assess the degree of migratory connectivity between breeding and wintering populations. Location Switzerland and Germany. Methods We analysed stable nitrogen (δ¹⁵N), carbon (δ¹³C) and hydrogen (δD) isotopes from wing feathers from two populations of wrynecks to infer their wintering origins and to assess the strength of migratory connectivity. We tested whether variation in feather isotopic values within the Swiss population was affected by bird age and collection year and then considered differences in isotopic values between the two breeding populations. We used isotopic values of summer‐ and winter‐grown feathers to estimate seasonal distributions. Finally, we calculated a species‐specific δD discrimination factor between feathers and mean annual δD values to assign winter‐grown feathers to origin. Results Bird age and collection year caused substantial isotopic variation in winter‐grown feathers, which may be because of annually variable weather conditions, movements of birds among wintering sites and/or reflect asynchronous moulting or selection pressure. The large isotopic variance in winter‐grown feathers nevertheless suggested low migratory connectivity for each breeding population, with partially overlapping wintering regions for the two populations. Main conclusions Isotopic variance in winter‐grown feathers of two breeding populations of wrynecks and their geographical assignment point to defined, albeit overlapping, wintering areas, suggesting both leapfrog migration and low migratory connectivity. On this basis, integrative demographic models can be built looking at seasonal survival patterns with links to local environmental conditions on both breeding and wintering grounds, which may elucidate causes of declines in migratory bird species.     

Individual dispersal,landscape connectivity and ecological networks

Connectivity is classically considered an emergent property of landscapes encapsulating individuals' flows across space. However, its operational use requires a precise understanding of why and how organisms disperse. Such movements, and hence landscape connectivity, will obviously vary according to both organism properties and landscape features. We review whether landscape connectivity estimates could gain in both precision and generality by incorporating three fundamental outcomes of dispersal theory. Firstly, dispersal is a multi‐causal process; its restriction to an ‘escape reaction’ to environmental unsuitability is an oversimplification, as dispersing individuals can leave excellent quality habitat patches or stay in poor‐quality habitats according to the relative costs and benefits of dispersal and philopatry. Secondly, species, populations and individuals do not always react similarly to those cues that trigger dispersal, which sometimes results in contrasting dispersal strategies. Finally, dispersal is a major component of fitness and is thus under strong selective pressures, which could generate rapid adaptations of dispersal strategies. Such evolutionary responses will entail spatiotemporal variation in landscape connectivity. We thus strongly recommend the use of genetic tools to: (i) assess gene flow intensity and direction among populations in a given landscape; and (ii) accurately estimate landscape features impacting gene flow, and hence landscape connectivity. Such approaches will provide the basic data for planning corridors or stepping stones aiming at (re)connecting local populations of a given species in a given landscape. This strategy is clearly species‐ and landscape‐specific. But we suggest that the ecological network in a given landscape could be designed by stacking up such linkages designed for several species living in different ecosystems. This procedure relies on the use of umbrella species that are representative of other species living in the same ecosystem.     

Designing connected marine reserves in the face of global warming

Marine reserves are widely used to protect species important for conservation and fisheries and to help maintain ecological processes that sustain their populations, including recruitment and dispersal. Achieving these goals requires well‐connected networks of marine reserves that maximize larval connectivity, thus allowing exchanges between populations and recolonization after local disturbances. However, global warming can disrupt connectivity by shortening potential dispersal pathways through changes in larval physiology. These changes can compromise the performance of marine reserve networks, thus requiring adjusting their design to account for ocean warming. To date, empirical approaches to marine prioritization have not considered larval connectivity as affected by global warming. Here, we develop a framework for designing marine reserve networks that integrates graph theory and changes in larval connectivity due to potential reductions in planktonic larval duration (PLD) associated with ocean warming, given current socioeconomic constraints. Using the Gulf of California as case study, we assess the benefits and costs of adjusting networks to account for connectivity, with and without ocean warming. We compare reserve networks designed to achieve representation of species and ecosystems with networks designed to also maximize connectivity under current and future ocean‐warming scenarios. Our results indicate that current larval connectivity could be reduced significantly under ocean warming because of shortened PLDs. Given the potential changes in connectivity, we show that our graph‐theoretical approach based on centrality (eigenvector and distance‐weighted fragmentation) of habitat patches can help design better‐connected marine reserve networks for the future with equivalent costs. We found that maintaining dispersal connectivity incidentally through representation‐only reserve design is unlikely, particularly in regions with strong asymmetric patterns of dispersal connectivity. Our results support previous studies suggesting that, given potential reductions in PLD due to ocean warming, future marine reserve networks would require more and/or larger reserves in closer proximity to maintain larval connectivity.     

Species are hypotheses: avoid connectivity assessments based on pillars of sand

Connectivity among populations determines the dynamics and evolution of populations, and its assessment is essential in ecology in general and in conservation biology in particular. The robust basis of any ecological study is the accurate delimitation of evolutionary units, such as populations, metapopulations and species. Yet a disconnect still persists between the work of taxonomists describing species as working hypotheses and the use of species delimitation by molecular ecologists interested in describing patterns of gene flow. This problem is particularly acute in the marine environment where the inventory of biodiversity is relatively delayed, while for the past two decades, molecular studies have shown a high prevalence of cryptic species. In this study, we illustrate, based on marine case studies, how the failure to recognize boundaries of evolutionary‐relevant unit leads to heavily biased estimates of connectivity. We review the conceptual framework within which species delimitation can be formalized as falsifiable hypotheses and show how connectivity studies can feed integrative taxonomic work and vice versa. Finally, we suggest strategies for spatial, temporal and phylogenetic sampling to reduce the probability of inadequately delimiting evolutionary units when engaging in connectivity studies.     

Estimating migratory connectivity of birds when re‐encounter probabilities are heterogeneous

Understanding the biology and conducting effective conservation of migratory species requires an understanding of migratory connectivity – the geographic linkages of populations between stages of the annual cycle. Unfortunately, for most species, we are lacking such information. The North American Bird Banding Laboratory (BBL) houses an extensive database of marking, recaptures and recoveries, and such data could provide migratory connectivity information for many species. To date, however, few species have been analyzed for migratory connectivity largely because heterogeneous re‐encounter probabilities make interpretation problematic. We accounted for regional variation in re‐encounter probabilities by borrowing information across species and by using effort covariates on recapture and recovery probabilities in a multistate capture–recapture and recovery model. The effort covariates were derived from recaptures and recoveries of species within the same regions. We estimated the migratory connectivity for three tern species breeding in North America and over‐wintering in the tropics, common (Sterna hirundo), roseate (Sterna dougallii), and Caspian terns (Hydroprogne caspia). For western breeding terns, model‐derived estimates of migratory connectivity differed considerably from those derived directly from the proportions of re‐encounters. Conversely, for eastern breeding terns, estimates were merely refined by the inclusion of re‐encounter probabilities. In general, eastern breeding terns were strongly connected to eastern South America, and western breeding terns were strongly linked to the more western parts of the nonbreeding range under both models. Through simulation, we found this approach is likely useful for many species in the BBL database, although precision improved with higher re‐encounter probabilities and stronger migratory connectivity. We describe an approach to deal with the inherent biases in BBL banding and re‐encounter data to demonstrate that this large dataset is a valuable source of information about the migratory connectivity of the birds of North America.