Summary Dispersal is essential for species to survive the threats of habitat destruction and climate change. Combining descriptions of dispersal ability with those of landscape structure, the concept of functional connectivity has been popular for understanding and predicting species’ spatial responses to environmental change. Following recent advances, the functional connectivity concept is now able to move beyond landscape structure to consider more explicitly how other external factors such as climate and resources affect species movement. We argue that these factors, in addition to a consideration of the complete dispersal process, are critical for an accurate understanding of functional connectivity for plant species in response to environmental change. We use recent advances in dispersal, landscape and molecular ecology to describe how a range of external factors can influence effective dispersal in plant species, and how the resulting functional connectivity can be assessed. Synthesis. We define plant functional connectivity as the effective dispersal of propagules or pollen among habitat patches in a landscape. Plant functional connectivity is determined by a combination of landscape structure, interactions between plant, environment and dispersal vectors, and the successful establishment of individuals. We hope that this consolidation of recent research will help focus future connectivity research and conservation.

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Directed dispersal by animal vectors has been found to have large effects on the structure and dynamics of plant populations adapted to frugivory. Yet, empirical data are lacking on the potential of directed dispersal by rotational grazing of domestic animals to mediate gene flow across the landscape. Here, we investigated the potential effect of large-flock shepherding on landscape-scale genetic structure in the calcareous grassland plant Dianthus carthusianorum, whose seeds lack morphological adaptations to dispersal to animals or wind. We found a significant pattern of genetic structure differentiating population within grazed patches of three nonoverlapping shepherding systems and populations of ungrazed patches. Among ungrazed patches, we found a strong and significant effect of isolation by distance (r = 0.56). In contrast, genetic distance between grazed patches within the same herding system was unrelated to geographical distance but significantly related to distance along shepherding routes (r = 0.44). This latter effect of connectivity along shepherding routes suggests that gene flow is spatially restricted occurring mostly between adjacent populations. While this study used nuclear markers that integrate gene flow by pollen and seed, the significant difference in the genetic structure between ungrazed patches and patches connected by large-flock shepherding indicates the potential of directed seed dispersal by sheep across the landscape.

Response to habitat fragmentation may not be generalized among species, in particular for plant communities with a variety of dispersal traits. Calcareous grasslands are one of the most species-rich habitats in Central Europe, but abandonment of traditional management has caused a dramatic decline of calcareous grassland species. In the Southern Franconian Alb in Germany, reintroduction of rotational shepherding in previously abandoned grasslands has restored species diversity, and it has been suggested that sheep support seed dispersal among grasslands. We tested the effect of rotational shepherding on demographic and genetic connectivity of calcareous grassland specialist plants and whether the response of plant populations to shepherding was limited to species dispersed by animals (zoochory). Specifically, we tested competing dispersal models and source and focal patch properties to explain landscape connectivity with patch-occupancy data of 31 species. We fitted the same connectivity models to patch occupancy and nuclear microsatellite data for the herb Dianthus carthusianorum (Carthusian pink). For 27 species, patch connectivity was explained by dispersal by rotational shepherding regardless of adaptations to zoochory, whereas population size (16% species) and patch area (0% species) of source patches were not important predictors of patch occupancy in most species. [Correction made after online publication, February 25, 2014: Population size and patch area percentages were mistakenly inverted, and have now been fixed.] Microsite diversity of focal patches significantly increased the model variance explained by patch occupancy in 90% of the species. For D. carthusianorum, patch connectivity through rotational shepherding explained both patch occupancy and population genetic diversity. Our results suggest shepherding provides dispersal for multiple plant species regardless of their dispersal adaptations and thus offers a useful approach to restore plant diversity in fragmented calcareous grasslands.

Local adaptation is necessary for population survival and depends on the interplay between responses to selective forces and demographic processes that introduce or retain adaptive and maladaptive attributes. Host-parasite systems are dynamic, varying in space and time, where both host and parasites must adapt to their ever-changing environment in order to survive. We investigated patterns of local adaptation in raccoon populations with varying temporal exposure to the raccoon rabies virus (RRV). RRV infects approximately 85% of the population when epizootic and has been presumed to be completely lethal once contracted; however, disease challenge experiments and varying spatial patterns of RRV spread suggest some level of immunity may exist. We first assessed patterns of local adaptation in raccoon populations along the eastern seaboard of North America by contrasting spatial patterns of neutral (microsatellite loci) and functional, major histocompatibility complex (MHC) genetic diversity and structure. We explored variation of MHC allele frequencies in the light of temporal population exposure to RRV (0-60 years) and specific RRV strains in infected raccoons. Our results revealed high levels of MHC variation (66 DRB exon 2 alleles) and pronounced genetic structure relative to neutral microsatellite loci, indicative of local adaptation. We found a positive association linking MHC genetic diversity and temporal RRV exposure, but no association with susceptibility and resistance to RRV strains. These results have implications for landscape epidemiology studies seeking to predict the spread of RRV and present an example of how population demographics influence the degree to which populations adapt to local selective pressures.

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Distance, environmental heterogeneity and local adaptation can strongly influence population structure and connectivity. Understanding how these factors shape the genomic landscape of threatened species is a major goal in conservation genomics and wildlife management. Herein, we use thousands (6,859) of single nucleotide polymorphism markers and spatial data from hundreds of individuals (n = 646) to re-evaluate the population structure of Agassiz’s desert tortoise (Gopherus agassizii). Analyses resolve from 4 to 8 spatially well-defined clusters across the range. Western, central, and southern populations within the Western Mojave recovery unit are consistent throughout, while analyses sometimes merge other recovery units depending on the level of clustering. Causal modeling consistently associates genetic connectivity with least-cost distance, based on multiple landscape features associated with tortoise habitat, better than geographic distance. Some features include elevation, soil depth, rock volume, precipitation, and vegetation coverage, suggesting that physical, climatic, and biotic landscape features have played a strong evolutionary role restricting gene flow between populations. Further, 12 highly differentiated outlier loci have associated functions that may be involved with neurogenesis, wound healing, lipid metabolism, and possibly vitellogenesis. Together, these findings have important implications for recovery programs, such as translocations, population augmentation, reproduction in captivity and the identification of ecologically important genes, opening new venues for conservation genomics in desert tortoises.

Ecological connectivity networks have been proposed as an efficient way to reconnect communities in fragmented landscapes. Yet few studies have evaluated if they are successful at enhancing actual functional connectivity (i.e. realized dispersal or gene flow) of focal species, or if this enhanced connectivity is enough to maintain genetic diversity and fitness of plant populations. Here we test the efficacy of an ecological connectivity network implemented in southern Germany since 1989 to reconnect calcareous grassland fragments through rotational shepherding. We genotyped 1,449 individuals from 57 populations and measured fitness-related traits in 10 populations of Pulsatilla vulgaris, a flagship species of calcareous grasslands in Europe. We tested if the shepherding network explained functional connectivity in P. vulgaris and if higher connectivity translated to higher genetic diversity and fitness of populations. We found that population-specific Fst was lowest in populations that had high connectivity within the shepherding network, and that well-connected populations within the network had significantly higher genetic diversity than ungrazed and more isolated grazed populations. Moreover, genetic diversity was significantly positively correlated with both seed set and seed mass. Together our results suggest that the implementation of an ecological shepherding network is an effective management measure to maintain functional connectivity and genetic diversity at the landscape scale for a calcareous grassland specialist. Populations with reduced genetic diversity would likely benefit from inclusion, or better integration into the ecological connectivity network. Our study demonstrates the often postulated but rarely tested sequence of positive associations between connectivity, genetic diversity, and fitness at the landscape scale, and provides a framework for testing the efficacy of ecological connectivity networks for focal species using molecular genetic tools.