Genome-wide investigation of the WRKY transcription factor gene family in weeping forsythia: expression profile and cold and drought stress responses.

Weeping forsythia is a wide-spread shrub in China with important ornamental, medicinal and ecological values. It is widely distributed in China's warm temperate zone. In plants, WRKY transcription factors play important regulatory roles in seed germination, flower development, fruit ripening and coloring, and biotic and abiotic stress response. To date, WRKY transcription factors have not been systematically studied in weeping forsythia. In this study, we identified 79 WRKY genes in weeping forsythia and classified them according to their naming rules in Arabidopsis thaliana. Phylogenetic tree analysis showed that, except for IIe subfamily, whose clustering was inconsistent with A. thaliana clustering, other subfamily clustering groups were consistent. Cis-element analysis showed that WRKY genes related to pathogen resistance in weeping forsythia might be related to methyl jasmonate and salicylic acid-mediated signaling pathways. Combining cis-element and expression pattern analyses of WRKY genes showed that more than half of WRKY genes were involved in light-dependent development and morphogenesis in different tissues. The gene expression results showed that 13 WRKY genes were involved in drought response, most of which might be related to the abscisic acid signaling pathway, and a few of which might be regulated by MYB transcription factors. The gene expression results under cold stress showed that 17 WRKY genes were involved in low temperature response, and 9 of them had low temperature responsiveness cis-elements. Our study of WRKY family in weeping forsythia provided useful resources for molecular breeding and important clues for their functional verification.

A new mechanism of flowering regulation by the competition of isoforms in Osmanthus fragrans.

The regulation of flowering time is typically governed by transcription factors or epigenetic modifications. Transcript isoforms can play important roles in flowering regulation. Recently, transcript isoforms were discovered in the key genes, OfAP1 and OfTFL1, of the flowering regulatory network in Osmanthus fragrans. OfAP1-b generates a full-length isoform of OfAP1-b1 as well as an isoform of OfAP1-b2 that lacks the C-terminal domain. Although OfAP1-b2 does not possess an activation domain, it has a complete K domain that allows it to form heterodimers. OfAP1-b2 competes with OfAP1-b1 by binding with OfAGL24 to create non-functional and functional heterodimers. As a result, OfAP1-b1 promotes flowering while OfAP1-b2 delays flowering. OfTFL1 produces two isoforms located in different areas: OfTFL1-1 in the cytoplasm and OfTFL1-2 in the nucleus. When combined with OfFD, OfTFL1-1 does not enter the nucleus to repress AP1 expression, leading to early flowering. Conversely, when combined with OfFD, OfTFL1-2 enters the nucleus to repress AP1 expression, resulting in later flowering. Tissue-specific expression and functional conservation testing of OfAP1 and OfTFL1 support the new model's effectiveness in regulating flowering. Overall, this study provides new insights into regulating flowering time by the competition of isoforms.

Entropy, Ecology and Evolution: Toward a Unified Philosophy of Biology.

Darwin proposed that the capacity of organisms to produce more offspring that can be supported by the environment would lead to a struggle for existence, and individuals that are most fit for survival and reproduction would be selected through natural selection. Ecology is the science that studies the interaction between organisms and their environment within the context of Darwinian evolution, and an ecosystem is defined as a community of living organisms in conjunction with the nonliving components of their environment, interacting as a system. One topic that has been very much understudied and largely ignored in evolutionary biology is the overarching context of thermodynamics in controlling all biological processes and the evolution of life. Most fundamentally, organisms are self-replicating dissipative structures. Evolution is the process whereby variation in the structure of organisms have differential fitness in terms of their effectiveness at building and maintaining their structure, efficiently consuming free energy, and effectively reproducing and passing on those heritable variations, leading to change in the frequency of genetic variation and associated change in the characteristics in the population. The central process is dissipation of free energy according to the second law of thermodynamics, and evolution therefore is better conceptualized as the emergence of self-replicating dissipative structures that through natural selection become increasingly more efficient at degrading free energy. Ecosystems are linked series of dissipative structures with heat engine dynamics driven by random dissipation of energy and increasing entropy. The structure and composition of ecosystems across scales are emergent dissipative structures driven by the flow of energy and the increase in entropy. Communities and ecosystems are emergent properties of a system that has evolved to most efficiently dissipate energy and increase entropy. By focusing on the fundamental entity (energy), and the fundamental process (dissipation and disordering of energy and increasing of entropy), we are able to have a much clearer and powerful understanding of what life is, from the level of biochemistry, to evolution, to the nature of the organism itself, and to the emergent structures of ecosystems, food webs and communities.

Contrasting effects of human settlement on the interaction among sympatric apex carnivores.

In the face of a growing human footprint, understanding interactions among threatened large carnivores is fundamental to effectively mitigating anthropogenic threats and managing species. Using data from a large-scale camera trap survey, we investigated the effects of environmental and anthropogenic variables on the interspecific interaction of a carnivore guild comprising of tiger, leopard and dhole in Bhutan. We demonstrate the complex effects of human settlement density on large carnivore interactions. Specifically, we demonstrate that leopard-dhole co-occupancy probability was higher in areas with higher human settlement density. The opposite was true for tiger-leopard co-occupancy probability, but it was positively affected by large prey (gaur) abundance. These findings suggest that multi-carnivore communities across land-use gradients are spatially structured and mediated also by human presence and/or the availability of natural prey. Our findings show that space-use patterns are driven by a combination of the behavioural mechanism of each species and its interactions with competing species. The duality of the effect of settlement density on species interactions suggests that the benefits of exploiting anthropogenic environments are a trade-off between ecological opportunity (food subsidies or easy prey) and the risk of escalating conflict with humans.

Synthesizing habitat connectivity analyses of a globally important human-dominated tiger-conservation landscape.

As ecological data and associated analyses become more widely available, synthesizing results for effective communication with stakeholders is essential. In the case of wildlife corridors, managers in human-dominated landscapes need to identify both the locations of corridors and multiple stakeholders for effective oversight. We synthesized 5 independent studies of tiger (Panthera tigris) connectivity in central India, a global priority landscape for tiger conservation, to quantify agreement on landscape permeability for tiger movement and potential movement pathways. We used the latter analysis to identify connectivity areas on which studies agreed and stakeholders associated with these areas to determine relevant participants in corridor management. Three or more of the 5 studies' resistance layers agreed in 63% of the study area. Areas in which all studies agree on resistance were of primarily low (66%, e.g., forest) and high (24%, e.g., urban) resistance. Agreement was lower in intermediate resistance areas (e.g., agriculture). Despite these differences, the studies largely agreed on areas with high levels of potential movement: >40% of high average (top 20%) current-flow pixels were also in the top 20% of current-flow agreement pixels (measured by low variation), indicating consensus connectivity areas (CCAs) as conservation priorities. Roughly 70% of the CCAs fell within village administrative boundaries, and 100% overlapped forest department management boundaries, suggesting that people live and use forests within these priority areas. Over 16% of total CCAs' area was within 1 km of linear infrastructure (437 road, 170 railway, 179 transmission line, and 339 canal crossings; 105 mines within 1 km of CCAs). In 2019, 78% of forest land diversions for infrastructure and mining in Madhya Pradesh (which comprises most of the study region) took place in districts with CCAs. Acute competition for land in this landscape with globally important wildlife corridors calls for an effective comanagement strategy involving local communities, forest departments, Appendix 1 and infrastructure planners. This article is protected by copyright. All rights reserved.

Transcriptomic responses to drought stress among natural populations provide insights into local adaptation of weeping forsythia.

BACKGROUND: Understanding the genetic mechanisms of local adaptation is an important emerging topic in molecular ecology and evolutionary biology. RESULTS: Here, we identify the physiological changes and differential expression of genes among different weeping forsythia populations under drought stress in common garden experiments. Physiological results showed that HBWZ might have higher drought tolerance among four populations. RNA-seq results showed that significant differential expression in the genes responding to the synthesis of flavonoids, aromatic substances, aromatic amino acids, oxidation-reduction process, and transmembrane transport occured among four populations. By further reanalysis of results of previous studies, sequence differentiation was found in the genes related to the synthesis of aromatic substances among different weeping forsythia populations. CONCLUSIONS: Overall, our study supports the hypothesis that the dual differentiation in gene efficiency and expression increases among populations in response to heterogeneous environments and is an important evolutionary process of local adaptation. Here, we proposed a new working model of local adaptation of weeping forsythia populations under different intensities of drought stress, which provides new insights for understanding the genetic mechanisms of local adaptation for non-model species.

Genetic connectivity of two marine gastropods in the Mediterranean Sea: seascape genetics reveals species-specific oceanographic drivers of gene flow.

Oceanographic features such as currents, waves, temperature and salinity, together with life history traits, control patterns and rates of gene flow and contribute to shaping the population genetic structure of marine organisms. Seascape genetics is an emerging discipline that adopts a spatially explicit approach to examine biotic and abiotic factors that drive gene flow in marine environments. In this study, we examined factors that contribute to genetic differentiation in two coastal Mediterranean gastropods whose geographical ranges overlap but which inhabit different environments. The two species differ in several life history traits and in their dispersal capabilities. Genetic differentiation was relatively low for the trochid species Gibbula divaricata (FST  =0.059), and high for the vermetid species Dendropoma lebeche (FST  =0.410). Salinity emerged as the most important variable explaining the genetic structure of both species; sea surface temperature was also important for G. divaricata. For the more sessile D. lebeche, the coastline was predicted to provide important pathways for stepping-stone connectivity and gene flow. Our results provide a greater understanding of the factors influencing marine population connectivity, which may be useful to guide marine conservation and management in the Mediterranean.

Evidence of spatial genetic structure in a snow leopard population from Gansu, China.

Understanding the spatial structure of genetic diversity provides insights into a populations' genetic status and enables assessment of its capacity to counteract the effects of genetic drift. Such knowledge is particularly scarce for the snow leopard, a conservation flagship species of Central Asia mountains. Focusing on a snow leopard population in the Qilian mountains of Gansu Province, China, we characterised the spatial genetic patterns by incorporating spatially explicit indices of diversity and multivariate analyses, based on different inertia levels of Principal Component Analysis (PCA). We compared two datasets differing in the number of loci and individuals. We found that genetic patterns were significantly spatially structured and were characterised by a broad geographical division coupled with a fine-scale cline of differentiation. Genetic admixture was detected in two adjoining core areas characterised by higher effective population size and allelic diversity, compared to peripheral localities. The power to detect significant spatial relationships depended primarily on the number of loci, and secondarily on the number of PCA axes. Spatial patterns and indices of diversity highlighted the cryptic structure of snow leopard genetic diversity, likely driven by its ability to disperse over large distances. In combination, the species' low allelic richness and large dispersal ability result in weak genetic differentiation related to major geographical features and isolation by distance. This study illustrates how cryptic genetic patterns can be investigated and analysed at a fine spatial scale, providing insights into the spatially variable isolation effects of both geographic distance and landscape resistance.

Generalizing Boltzmann Configurational Entropy to Surfaces, Point Patterns and Landscape Mosaics.

Several methods have been recently proposed to calculate configurational entropy, based on Boltzmann entropy. Some of these methods appear to be fully thermodynamically consistent in their application to landscape patch mosaics, but none have been shown to be fully generalizable to all kinds of landscape patterns, such as point patterns, surfaces, and patch mosaics. The goal of this paper is to evaluate if the direct application of the Boltzmann relation is fully generalizable to surfaces, point patterns, and landscape mosaics. I simulated surfaces and point patterns with a fractal neutral model to control their degree of aggregation. I used spatial permutation analysis to produce distributions of microstates and fit functions to predict the distributions of microstates and the shape of the entropy function. The results confirmed that the direct application of the Boltzmann relation is generalizable across surfaces, point patterns, and landscape mosaics, providing a useful general approach to calculating landscape entropy.

Thermodynamic Consistency of the Cushman Method of Computing the Configurational Entropy of a Landscape Lattice.

There has been a recent surge of interest in theory and methods for calculating the entropy of landscape patterns, but relatively little is known about the thermodynamic consistency of these approaches. I posit that for any of these methods to be fully thermodynamically consistent, they must meet three conditions. First, the computed entropies must lie along the theoretical distribution of entropies as a function of total edge length, which Cushman showed was a parabolic function following from the fact that there is a normal distribution of permuted edge lengths, the entropy is the logarithm of the number of microstates in a macrostate, and the logarithm of a normal distribution is a parabolic function. Second, the entropy must increase over time through the period of the random mixing simulation, following the expectation that entropy increases in a closed system. Third, at full mixing, the entropy will fluctuate randomly around the maximum theoretical value, associated with a perfectly random arrangement of the lattice. I evaluated these criteria in a test condition involving a binary, two-class landscape using the Cushman method of directly applying the Boltzmann relation (s = klogW) to permuted landscape configurations and measuring the distribution of total edge length. The results show that the Cushman method directly applying the classical Boltzmann relation is fully consistent with these criteria and therefore fully thermodynamically consistent. I suggest that this method, which is a direct application of the classical and iconic formulation of Boltzmann, has advantages given its direct interpretability, theoretical elegance, and thermodynamic consistency.

Entropy in Landscape Ecology: A Quantitative Textual Multivariate Review.

This paper presents a multivariate textual analysis of more than 1300 papers on entropy in ecology. There are six main insights that emerged. First, there is a large body of literature that has addressed some aspect of entropy in ecology, most of which has been published in the last 5-10 years. Second, the vast majority of these papers focus on species distribution, species richness, relative abundance or trophic structure and not landscape-scale patterns or processes, pe se. Third, there have been few papers addressing landscape-level questions related to entropy. Fourth, the quantitative analysis with hierarchical clustering identified a strongly nested structure among papers that addressed entropy in ecology. Fifth, there is clear differentiation of papers focused on landscape-level applications of entropy from other papers, with landscape focused papers clustered together at each level of the hierarchy in a relatively small and closely associated group. Sixth, this group of landscape-focused papers was substructured between papers that explicitly adopted entropy measures to quantify the spatial pattern of landscape mosaics, often using variations on Boltzmann entropy, versus those that utilize Shannon entropy measures from information theory, which are not generally explicit in their assessment of spatial configuration. This review provides a comprehensive, quantitative assessment of the scope, trends and relationships among a large body of literature related to entropy in ecology and for the first time puts landscape ecological research on entropy into that context.

An Axiom SNP genotyping array for Douglas-fir.

BACKGROUND: In forest trees, genetic markers have been used to understand the genetic architecture of natural populations, identify quantitative trait loci, infer gene function, and enhance tree breeding. Recently, new, efficient technologies for genotyping thousands to millions of single nucleotide polymorphisms (SNPs) have finally made large-scale use of genetic markers widely available. These methods will be exceedingly valuable for improving tree breeding and understanding the ecological genetics of Douglas-fir, one of the most economically and ecologically important trees in the world. RESULTS: We designed SNP assays for 55,766 potential SNPs that were discovered from previous transcriptome sequencing projects. We tested the array on ~ 2300 related and unrelated coastal Douglas-fir trees (Pseudotsuga menziesii var. menziesii) from Oregon and Washington, and 13 trees of interior Douglas-fir (P. menziesii var. glauca). As many as ~ 28 K SNPs were reliably genotyped and polymorphic, depending on the selected SNP call rate. To increase the number of SNPs and improve genome coverage, we developed protocols to 'rescue' SNPs that did not pass the default Affymetrix quality control criteria (e.g., 97% SNP call rate). Lowering the SNP call rate threshold from 97 to 60% increased the number of successful SNPs from 20,669 to 28,094. We used a subset of 395 unrelated trees to calculate SNP population genetic statistics for coastal Douglas-fir. Over a range of call rate thresholds (97 to 60%), the median call rate for SNPs in Hardy-Weinberg equilibrium ranged from 99.2 to 99.7%, and the median minor allele frequency ranged from 0.198 to 0.233. The successful SNPs also worked well on interior Douglas-fir. CONCLUSIONS: Based on the original transcriptome assemblies and comparisons to version 1.0 of the Douglas-fir reference genome, we conclude that these SNPs can be used to genotype about 10 K to 15 K loci. The Axiom genotyping array will serve as an excellent foundation for studying the population genomics of Douglas-fir and for implementing genomic selection. We are currently using the array to construct a linkage map and test genomic selection in a three-generation breeding program for coastal Douglas-fir.

Habitat amount mediates the effect of fragmentation on a pollinator's reproductive performance, but not on its foraging behaviour.

Agricultural intensification, with its associated habitat loss and fragmentation, is among the most important drivers of the ongoing pollination crisis. In this quasi-experimental study, conducted in intensively managed vineyards in southwestern Switzerland, we tested the separate and interdependent effects of habitat amount and fragmentation on the foraging activity and reproductive performance of bumblebee Bombus t. terrestris colonies. Based on a factorial design, we selected a series of spatially replicated study sites across a dual gradient of habitat amount (area of ground-vegetated vineyards) and fragmentation (density of ground-vegetated vineyard fields) in a landscape predominantly consisting of vineyards with bare grounds. The foraging activity of individual bumblebees was measured using the radio frequency identification (RFID) technology, and we assessed final colony size to measure reproductive performance. We found an interactive effect of habitat amount and fragmentation on colony size. More specifically, the degree of fragmentation had a negative effect on bumblebee colony size when the amount of habitat was low, while it had a weak positive effect on colony size in landscapes with high amounts of habitat. At the level of individual vineyard fields, ground vegetation cover exerted a positive effect on bumblebee colony size. Fragmentation, but not habitat amount, significantly influenced foraging activity, with more foraging trips in sites with lower degrees of fragmentation. Our results emphasise the importance of studying the separate and interdependent effects of habitat amount and fragmentation to understand their influence on pollinators, providing guidance for optimising the spatial configuration of agricultural landscapes from a biodiversity viewpoint.

The role of hybridization during ecological divergence of southwestern white pine (Pinus strobiformis) and limber pine (P. flexilis).

Interactions between extrinsic factors, such as disruptive selection and intrinsic factors, such as genetic incompatibilities among loci, often contribute to the maintenance of species boundaries. The relative roles of these factors in the establishment of reproductive isolation can be examined using species pairs characterized by gene flow throughout their divergence history. We investigated the process of speciation and the maintenance of species boundaries between Pinus strobiformis and Pinus flexilis. Utilizing ecological niche modelling, demographic modelling and genomic cline analyses, we illustrated a divergence history with continuous gene flow. Our results supported an abundance of advanced generation hybrids and a lack of loci exhibiting steep transition in allele frequency across the hybrid zone. Additionally, we found evidence for climate-associated variation in the hybrid index and niche divergence between parental species and the hybrid zone. These results are consistent with extrinsic factors, such as climate, being an important isolating mechanism. A build-up of intrinsic incompatibilities and of coadapted gene complexes is also apparent, although these appear to be in the earliest stages of development. This supports previous work in coniferous species demonstrating the importance of extrinsic factors in facilitating speciation. Overall, our findings lend support to the hypothesis that varying strength and direction of selection pressures across the long lifespans of conifers, in combination with their other life history traits, delays the evolution of strong intrinsic incompatibilities.

Calculation of Configurational Entropy in Complex Landscapes.

Entropy and the second law of thermodynamics are fundamental concepts that underlie all natural processes and patterns. Recent research has shown how the entropy of a landscape mosaic can be calculated using the Boltzmann equation, with the entropy of a lattice mosaic equal to the logarithm of the number of ways a lattice with a given dimensionality and number of classes can be arranged to produce the same total amount of edge between cells of different classes. However, that work seemed to also suggest that the feasibility of applying this method to real landscapes was limited due to intractably large numbers of possible arrangements of raster cells in large landscapes. Here I extend that work by showing that: (1) the proportion of arrangements rather than the number with a given amount of edge length provides a means to calculate unbiased relative configurational entropy, obviating the need to compute all possible configurations of a landscape lattice; (2) the edge lengths of randomized landscape mosaics are normally distributed, following the central limit theorem; and (3) given this normal distribution it is possible to fit parametric probability density functions to estimate the expected proportion of randomized configurations that have any given edge length, enabling the calculation of configurational entropy on any landscape regardless of size or number of classes. I evaluate the boundary limits (4) for this normal approximation for small landscapes with a small proportion of a minority class and show it holds under all realistic landscape conditions. I further (5) demonstrate that this relationship holds for a sample of real landscapes that vary in size, patch richness, and evenness of area in each cover type, and (6) I show that the mean and standard deviation of the normally distributed edge lengths can be predicted nearly perfectly as a function of the size, patch richness and diversity of a landscape. Finally, (7) I show that the configurational entropy of a landscape is highly related to the dimensionality of the landscape, the number of cover classes, the evenness of landscape composition across classes, and landscape heterogeneity. These advances provide a means for researchers to directly estimate the frequency distribution of all possible macrostates of any observed landscape, and then directly calculate the relative configurational entropy of the observed macrostate, and to understand the ecological meaning of different amounts of configurational entropy. These advances enable scientists to take configurational entropy from a concept to an applied tool to measure and compare the disorder of real landscapes with an objective and unbiased measure based on entropy and the second law.

Editorial: Entropy in Landscape Ecology.

Entropy and the second law of thermodynamics are the central organizing principles of nature, but the ideas and implications of the second law are poorly developed in landscape ecology. The purpose of this Special Issue "Entropy in Landscape Ecology" in Entropy is to bring together current research on applications of thermodynamics in landscape ecology, to consolidate current knowledge and identify key areas for future research. The special issue contains six articles, which cover a broad range of topics including relationships between entropy and evolution, connections between fractal geometry and entropy, new approaches to calculate configurational entropy of landscapes, example analyses of computing entropy of landscapes, and using entropy in the context of optimal landscape planning. Collectively these papers provide a broad range of contributions to the nascent field of ecological thermodynamics. Formalizing the connections between entropy and ecology are in a very early stage, and that this special issue contains papers that address several centrally important ideas, and provides seminal work that will be a foundation for the future development of ecological and evolutionary thermodynamics.

Conserving threatened riparian ecosystems in the American West: Precipitation gradients and river networks drive genetic connectivity and diversity in a foundation riparian tree (Populus angustifolia).

Gene flow is an evolutionary process that supports genetic connectivity and contributes to the capacity of species to adapt to environmental change. Yet, for most species, little is known about the specific environmental factors that influence genetic connectivity, or their effects on genetic diversity and differentiation. We used a landscape genetic approach to understand how geography and climate influence genetic connectivity in a foundation riparian tree (Populus angustifolia), and their relationships with specieswide patterns of genetic diversity and differentiation. Using multivariate restricted optimization in a reciprocal causal modelling framework, we quantified the relative contributions of riparian network connectivity, terrestrial upland resistance and climate gradients on genetic connectivity. We found that (i) all riparian corridors, regardless of river order, equally facilitated connectivity, while terrestrial uplands provided 2.5× more resistance to gene flow than riparian corridors. (ii) Cumulative differences in precipitation seasonality and precipitation of the warmest quarter were the primary climatic factors driving genetic differentiation; furthermore, maximum climate resistance was 45× greater than riparian resistance. (iii) Genetic diversity was positively correlated with connectivity (R2  = 0.3744, p = .0019), illustrating the utility of resistance models for identifying landscape conditions that can support a species' ability to adapt to environmental change. From these results, we present a map highlighting key genetic connectivity corridors across P. angustifolia's range that if disrupted could have long-term ecological and evolutionary consequences. Our findings provide recommendations for conservation and restoration management of threatened riparian ecosystems throughout the western USA and the high biodiversity they support.

Isolation by distance, resistance and/or clusters? Lessons learned from a forest-dwelling carnivore inhabiting a heterogeneous landscape.

Landscape genetics provides a valuable framework to understand how landscape features influence gene flow and to disentangle the factors that lead to discrete and/or clinal population structure. Here, we attempt to differentiate between these processes in a forest-dwelling small carnivore [European pine marten (Martes martes)]. Specifically, we used complementary analytical approaches to quantify the spatially explicit genetic structure and diversity and analyse patterns of gene flow for 140 individuals genotyped at 15 microsatellite loci. We first used spatially explicit and nonspatial Bayesian clustering algorithms to partition the sample into discrete clusters and evaluate hypotheses of 'isolation by barriers' (IBB). We further characterized the relationships between genetic distance and geographical ('isolation by distance', IBD) and ecological distances ('isolation by resistance', IBR) obtained from optimized landscape models. Using a reciprocal causal modelling approach, we competed the IBD, IBR and IBB hypotheses with each other to unravel factors driving population genetic structure. Additionally, we further assessed spatially explicit indices of genetic diversity using sGD across potentially overlapping genetic neighbourhoods that matched the inferred population structure. Our results revealed a complex spatial genetic cline that appears to be driven jointly by IBD and partial barriers to gene flow (IBB) associated with poor habitat and interspecific competition. Habitat loss and fragmentation, in synergy with past overharvesting and possible interspecific competition with sympatric stone marten (Martes foina), are likely the main factors responsible for the spatial genetic structure we observed. These results emphasize the need for a more thorough evaluation of discrete and clinal hypotheses governing gene flow in landscape genetic studies, and the potential influence of different limiting factors affecting genetic structure at different spatial scales.

Landscape effects on gene flow for a climate-sensitive montane species, the American pika.

Climate change is arguably the greatest challenge to conservation of our time. Most vulnerability assessments rely on past and current species distributions to predict future persistence but ignore species' abilities to disperse through landscapes, which may be particularly important in fragmented habitats and crucial for long-term persistence in changing environments. Landscape genetic approaches explore the interactions between landscape features and gene flow and can clarify how organisms move among suitable habitats, but have suffered from methodological uncertainties. We used a landscape genetic approach to determine how landscape and climate-related features influence gene flow for American pikas (Ochotona princeps) in Crater Lake National Park. Pikas are heat intolerant and restricted to cool microclimates; thus, range contractions have been predicted as climate changes. We evaluated the correlation between landscape variables and genetic distance using partial Mantel tests in a causal modelling framework, and used spatially explicit simulations to evaluate methods of model optimization including a novel approach based on relative support and reciprocal causal modelling. We found that gene flow was primarily restricted by topographic relief, water and west-facing aspects, suggesting that physical restrictions related to small body size and mode of locomotion, as well as exposure to relatively high temperatures, limit pika dispersal in this alpine habitat. Our model optimization successfully identified landscape features influencing resistance in the simulated data for this landscape, but underestimated the magnitude of resistance. This is the first landscape genetic study to address the fundamental question of what limits dispersal and gene flow in the American pika.

Spatiotemporal variation in resource selection: insights from the American marten (Martes americana).

Behavioral and genetic adaptations to spatiotemporal variation in habitat conditions allow species to maximize their biogeographic range and persist over time in dynamic environments. An understanding of these local adaptations can be used to guide management and conservation of populations over broad extents encompassing diverse habitats. This understanding is often achieved by identifying covariates related to species' occurrence in multiple independent studies conducted in relevant habitats and seasons. However, synthesis across studies is made difficult by differences in the model covariates evaluated and analytical frameworks employed. Furthermore, inferences may be confounded by spatiotemporal variation in which habitat attributes are limiting to the species' ecological requirements. In this study, we sought to quantify spatiotemporal variation in resource selection by the American marten (Martes americana) in forest ecosystems of the Pacific Northwest, USA. We developed resource selection functions for both summer and winter based on occurrence data collected in mesic and xeric forest habitats. Use of a consistent analytical framework facilitated comparisons. Habitat attributes predicting marten occurrence differed strongly between the two study areas, but not between seasons. Moreover, the spatial scale over which covariates were calculated greatly influenced their predictive power. In the mesic environment, marten resource selection was strongly tied to riparian habitats, whereas in the xeric environment, marten responded primarily to canopy cover and forest fragmentation. These differences in covariates associated with marten occurrence reflect differences in which factors were limiting to marten ecology in each study area, as well as local adaptations to habitat variability. Our results highlight the benefit of controlled meta-replication studies in which analyses of multiple study areas and seasons at varying spatial scales are integrated into a single framework.