Species' borders: a unifying theme in ecology

Biologists have long been fascinated by species' borders, and with good reason. Understanding the ecological and evolutionary dynamics of species' borders may prove to be the key that unlocks new understanding across a wide range of biological phenomena. After all, geographic range limits are a point of entry into understanding the ecological niche and threshold responses to environmental change. Elucidating patterns of gene flow to, and returning from, peripheral populations can provide important insights into the nature of adaptation, speciation and coevolution. Species' borders form natural laboratories for the study of the spatial structure of species interactions. Comparative studies from the center to the margin of species' ranges allow us to explore species' demographic responses along gradients of increasing environmental stress. Range dynamics further permit investigation into invasion dynamics and represent bellwethers for a changing climate. This set of papers explores ecological and evolutionary dynamics of species' borders from diverse empirical and theoretical perspectives.     

Oceanic dispersal barriers in a holoplanktonic gastropod

Pteropods, a group of holoplanktonic gastropods, are regarded as bioindicators of the effects of ocean acidification on open ocean ecosystems, because their thin aragonitic shells are susceptible to dissolution. While there have been recent efforts to address their capacity for physiological acclimation, it is also important to gain predictive understanding of their ability to adapt to future ocean conditions. However, little is known about the levels of genetic variation and large‐scale population structuring of pteropods, key characteristics enabling local adaptation. We examined the spatial distribution of genetic diversity in the mitochondrial cytochrome c oxidase I (COI) and nuclear 28S gene fragments, as well as shell shape variation, across a latitudinal transect in the Atlantic Ocean (35°N–36°S) for the pteropod Limacina bulimoides. We observed high levels of genetic variability (COI π = 0.034, 28S π = 0.0021) and strong spatial structuring (COI Φ_ST = 0.230, 28S Φ_ST = 0.255) across this transect. Based on the congruence of mitochondrial and nuclear differentiation, as well as differences in shell shape, we identified a primary dispersal barrier in the southern Atlantic subtropical gyre (15–18°S). This barrier is maintained despite the presence of expatriates, a gyral current system, and in the absence of any distinct oceanographic gradients in this region, suggesting that reproductive isolation between these populations must be strong. A secondary dispersal barrier supported only by 28S pairwise Φ_ST comparisons was identified in the equatorial upwelling region (between 15°N and 4°S), which is concordant with barriers observed in other zooplankton species. Both oceanic dispersal barriers were congruent with regions of low abundance reported for a similar basin‐scale transect that was sampled 2 years later. Our finding supports the hypothesis that low abundance indicates areas of suboptimal habitat that result in barriers to gene flow in widely distributed zooplankton species. Such species may in fact consist of several populations or (sub)species that are adapted to local environmental conditions, limiting their potential for adaptive responses to ocean changes. Future analyses of genome‐wide diversity in pteropods could provide further insight into the strength, formation and maintenance of oceanic dispersal barriers.     

Density‐dependent dispersal and the formation of range borders.

Knowledge about the mechanisms of range formation is crucial for scientifically based species conservation strategies in the face of ongoing global climate change. In recent years an increasing amount of studies have focused on the influences of density‐dependent dispersal on demographic and biogeographical patterns. However, it still remains unclear, to what extent and in what ways this strategy would affect the range formation of species. In order to fill this gap, we present a study using individual‐based simulations of a species with discrete generations living along a dispersal mortality gradient. We compare the evolution of range sizes for species following density‐dependent and density‐independent emigration. Furthermore we assess the influence of environmental stochasticity and Allee effects on range formation, as both processes are known to play an important role for dispersal evolution. We find that density‐dependent dispersal always results in much wider ranges than unconditional dispersal. Increasing environmental stochasticity, a predicted consequence of climate change, can remarkably expand the ranges of species living in such connectivity gradients if dispersal decisions are based on local population density. A strong Allee effect causes range contraction for both strategies, but the effect is considerably less dramatic under density‐dependent compared to density‐independent emigration. We strongly recommend accounting for these findings in future attempts to model species’ range shifts due to climate change.     

Rapid divergence of mussel populations despite incomplete barriers to dispersal

Striking genetic structure among marine populations at small spatial scales is becoming evident with extensive molecular studies. Such observations suggest isolation at small scales may play an important role in forming patterns of genetic diversity within species. Isolation‐by‐distance, isolation‐by‐environment and historical priority effects are umbrella terms for a suite of processes that underlie genetic structure, but their relative importance at different spatial and temporal scales remains elusive. Here, we use marine lakes in Indonesia to assess genetic structure and assess the relative roles of the processes in shaping genetic differentiation in populations of a bivalve mussel (Brachidontes sp.). Marine lakes are landlocked waterbodies of similar age (6,000–10,000 years), but with heterogeneous environments and varying degrees of connection to the sea. Using a population genomic approach (double‐digest restriction‐site‐associated DNA sequencing), we show strong genetic structuring across populations (range F_ST: 0.07–0.24) and find limited gene flow through admixture plots. At large spatial scales (>1,400 km), a clear isolation‐by‐distance pattern was detected. At smaller spatial scales (<200 km), this pattern is maintained, but accompanied by an association of genetic divergence with degree of connection. We hypothesize that (incomplete) dispersal barriers can cause initial isolation, allowing priority effects to give the numerical advantage necessary to initiate strong genetic structure. Priority effects may be strengthened by local adaptation, which the data may corroborate by showing a high correlation between mussel genotypes and temperature. Our study indicates an often‐neglected role of (evolution‐mediated) priority effects in shaping population divergence.     

Climate and habitat barriers to dispersal in the highly mobile grey wolf

We reanalysed published data to evaluate whether climate and habitat are barriers to dispersal in one of the most mobile and widely distributed mammals, the grey wolf (Canis lupus). Distance-based redundancy analysis (dbRDA) was used to examine the amount of variation in genetic distances that could be explained by an array of environmental factors, including geographical distance. Patterns in genetic variation were also examined using MDS plots among populations and relationships between genetic structure and individual environmental variables were further explored using the BIOENV procedure. We found that, contrary to a previous report, a pattern of isolation with distance is evident on a continental scale in the North American wolf population. This pattern is apparently related to climate and habitat. Specifically, vegetation types appear to play a role in the genetic dissimilarities among populations. When we controlled for the effect of spatial variation, climate was still associated with genetic distance. Further, partitioning of geographical distances into latitudinal and longitudinal axes revealed that the east-west gradient had the strongest relationship with genetic distance. We suggest two possible mechanisms by which environmental conditions may influence the dispersal decisions made by wolves.     

Habitat suitability,corridors and dispersal barriers for large carnivores in Poland

Carnivores are often particularly sensitive to landscape fragmentation. Ecological corridors may help to connect local populations, ensuring gene flow and retaining viable meta-populations. We aimed to establish habitat suitability models for two large carnivores in Poland, the grey wolf Canis lupus Linnaeus, 1758 and the Eurasian lynx Lynx lynx Linnaeus, 1758, based on ecological niche factor analysis (ENFA). Secondly, we calculated least cost paths (LCPs) based on cost values obtained from ENFA. Thirdly, we determined structures that might act as barriers, thus diminishing the value of the corridor unless appropriate conservation measures are taken. We compared some of the results with actual dispersal data of four lynx in eastern Poland. Results indicate that both species are highly marginalised. Less habitat that is currently available in Poland is suitable for lynx than for wolves. We determined a total of 76 LCPs. Comparison of these theoretical corridors with actual dispersal routes suggests that the traits of calculated LCPs are mostly within the range of those of real routes. We highlight a variety of features that might act as barriers, such as major roads (including planned highways), urbanized areas, and large un-forested areas. We give suggestions where concerted conservation efforts (eg wildlife passages) might be particularly well-directed.     

Detection of barriers to dispersal is masked by long lifespans and large population sizes

Population genetic analyses of species inhabiting fragmented landscapes are essential tools for conservation. Occasionally, analyses of fragmented populations find no evidence of isolation, even though a barrier to dispersal is apparent. In some cases, not enough time may have passed to observe divergence due to genetic drift, a problem particularly relevant for long‐lived species with overlapping generations. Failing to consider this quality during population structure analyses could result in incorrect conclusions about the impact of fragmentation on the species. We designed a model to explore how lifespan and population size influence perceived population structure of isolated populations over time. This iterative model tracked how simulated populations of variable lifespan and population size were affected by drift alone, using a freshwater mussel, Quadrula quadrula (mapleleaf), as a model system. In addition to exhibiting dramatic lifespan variability among species, mussels are also highly imperiled and exhibit fragmentation by dams throughout the range of many species. Results indicated that, unless population size was small (<50 individuals) or lifespan short (<22 years), observing genetic divergence among populations was unlikely. Even if wild populations are isolated, observing population structure in long‐lived mussels from modern damming practices is unlikely because it takes longer for population structure to develop in these species than most North American dams have existed. Larger population sizes and longer lifespans increase the time needed for significant divergence to occur. This study helps illuminate the factors that influence genetic responses by populations to isolation and provides a useful model for conservation‐oriented research.     

Landscape and oceanic barriers shape dispersal and population structure in the island nematode Pristionchus pacificus

In this study evolutionary host plant patterns at ranks from order to species were analysed using spatial evolutionary and ecological vicariance analysis (SEEVA), based on a multigene phylogeny of 45 ascomycete fungal species. The objective was to understand speciation events and host associations in Ophiognomonia (Gnomoniaceae). Species of this genus are perithecial fungi that occur as endophytes, pathogens, and latent saprobes on plants in the families of Betulaceae, Fagaceae, Juglandaceae, Lauraceae, Malvaceae, Platanaceae, Rosaceae, Salicaceae, and Sapindaceae. A second objective was to determine whether speciation events are influenced by host conservatism, host specialization, or host switching at different taxonomic host ranks. Host differences between sister clades were interpreted using the divergence index (D) from the SEEVA analysis, ranging from 0 for no divergence to 1 for maximum possible divergence. Several fungal subclades showed clear patterns of host order/family conservatism (D = 1.00) for hosts in Betulaceae, Fagaceae, Juglandaceae, and Rosaceae. Clear trends of host specialization at host genus and species ranks (D = 1.00) were suggested within these host families. Independent host jumps were observed for two species at the family rank and three at the order rank. As a result of this study, host specificity and specialization is hypothesized as a mechanism that can strongly contribute to speciation patterns in fungal pathogens. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 1–16.     

Cryptic barriers to dispersal within a lake allow genetic differentiation of Eurasian perch

Gene flow between coexisting or nearby populations normally prevents genetic divergence and local adaptation. Despite this, there are an increasing number of reports of sympatric sister taxa, indicating potential divergence and speciation in the face of gene flow. A large number of such reported cases involve lake-dwelling fish, which are expected to run into few physical barriers to dispersal within their aquatic habitat. However, such cases may not necessarily reflect sympatric speciation if cryptic dispersal barriers are common in lakes and other aquatic systems. In this study, we examined genetic differentiation in perch (Perca fluviatilis L.) from nine locations in a single, small lake (24 km(2)), using microsatellites. We detected significant genetic differentiation in all but two pairwise comparisons. These patterns were not consistent with divergence by distance or the existence of kin groups. Instead, they suggest that cryptic barriers to dispersal exist within the lake, allowing small-scale genetic divergence. Such an observation suggests that allopatric (or parapatric) divergence may be possible, even in small, apparently homogenous environments such as lakes. This has important consequences for how we currently view evidence from nature for sympatric speciation.     

Amazon's vulnerability to climate change heightened by deforestation and man‐made dispersal barriers

Species migrations in response to climate change have already been observed in many taxonomic groups worldwide. However, it remains uncertain if species will be able to keep pace with future climate change. Keeping pace will be especially challenging for tropical lowland rainforests due to their high velocities of climate change combined with high rates of deforestation, which may eliminate potential climate analogs and/or increase the effective distances between analogs by blocking species movements. Here, we calculate the distances between current and future climate analogs under various climate change and deforestation scenarios. Under even the most sanguine of climate change models (IPSL_CM4, A1b emissions scenario), we find that the median distance between areas in the Amazon rainforest and their closest future (2050) climate analog as predicted based on just temperature changes alone is nearly 300 km. If we include precipitation, the median distance increases by over 50% to >475 km. Since deforestation is generally concentrated in the hottest and driest portions of the Amazon, we predict that the habitat loss will have little direct impact on distances between climate analogs. If, however, deforested areas also act as a barrier to species movements, nearly 30% or 55% of the Amazon will effectively have no climate analogs anywhere in tropical South America under projections of reduced or Business‐As‐Usual deforestation, respectively. These ‘disappearing climates’ will be concentrated primarily in the southeastern Amazon. Consequently, we predict that several Amazonian ecoregions will have no areas with future climate analogs, greatly increasing the vulnerability of any populations or species specialized on these conditions. These results highlight the importance of including multiple climatic factors and human land‐use in predicting the effects of climate change, as well as the daunting challenges that Amazonian diversity faces in the near future.     

种子醇溶蛋白提取及检测条件探索

以大麦、小麦、玉米、高粱和苏丹草的种子为材料提取种子的醇溶蛋白,分析了不同提取剂及不同固液比[种子粉末样品质量(g)与提取剂的体积(mL)的比例]对种子醇溶蛋白提取效果的影响,并对SDS-PAGE检测醇溶蛋白中的不同胶浓度、厚度以及样品上样量等的影响进行了研究.结果表明,60%的正丙醇、乙二醇、异丙醇和叔丁醇分别是小麦、大麦、玉米以及高粱和苏丹草的最佳提取剂,将1∶6比例提取的种子醇溶蛋白以15μL上样,0.5mm厚度的15%分离胶电泳可以得到清晰的电泳检测图谱.     

Wetlands as barriers: effects of vegetated waterways on downstream dispersal of zebra mussels

1. Stream flow is a major vector for zebra mussel spread among inland lakes. Veligers have been found tens to hundreds of km from upstream source lakes in unvegetated stream and river systems. It has been suggested, however, that the downstream transport of zebra mussels is restricted by wetland ecosystems. We hypothesized that vegetated waterways, (i.e. wetland streams) would hinder the downstream dispersal of zebra mussels in connected inland lake systems. 2. Veliger abundance, recruitment and adult mussels were surveyed in four lake‐wetland systems in southeastern Michigan, U.S.A. from May to August 2006. Sampling was conducted downstream of the lakes invaded by zebra mussels, beginning at the upstream edge of aquatic vegetation and continuing downstream through the wetland streams. 3. Veliger abundance decreased rapidly in vegetated waterways compared to previously reported rates of decrease in non‐vegetated streams. Veligers were rarely found more than 1 km downstream from where vegetation began. Newly recruited individuals and adults were extremely rare beyond open water in the wetland systems. 4. Densely vegetated aquatic ecosystems limit the dispersal of zebra mussels downstream from invaded sources. Natural, remediated and constructed wetlands may therefore serve as a protective barrier to help prevent the spread of zebra mussels and other aquatic invasive species to other lakes and ecosystems.     

Landscape genetics of an endangered salt marsh endemic: Identifying population continuity and barriers to dispersal

Preserving the genetic diversity of endangered species is fundamental to their conservation and requires an understanding of genetic structure. In turn, identification of landscape features that impede gene flow can facilitate management to mitigate such obstacles and help with identifying isolated populations. We conducted a landscape genetic study of the endangered salt marsh harvest mouse (Reithrodontomys raviventris), a species endemic to the coastal marshes of the San Francisco Estuary of California. We collected and genotyped?>?500 samples from across the marshes of Suisun Bay which contain the largest remaining tracts of habitat for the species. Cluster analyses and a population tree identified three geographically discrete populations. Next, we conducted landscape genetic analyses at two scales (the entire study area and across the Northern Marshes) where we tested 65 univariate models of landscape features and used the best supported to test multivariable analyses. Our analysis of the entire study area indicated that open water and elevation (>?2 m) constrained gene flow. Analysis of the Northern Marshes, where low elevation marsh habitat is more continuous, indicated that geographic distance was the only significant predictor of genetic distance at this scale. The identification of a large, connected population across Northern Marshes achieves a number of recovery targets for this stronghold of the species. The identification of landscape features that act as barriers to dispersal enables the identification of isolated and vulnerable populations more broadly across the species range, thus aiding conservation prioritization.

     

Landscape genetics of fishers (Martes pennanti) in the Northeast: dispersal barriers and historical influences

Habitat fragmentation and overtrapping are thought to have resulted in severe population declines for fisher (Martes pennanti) across the northeastern United States, and by the end of the 1930s only 3 remnant populations remained. Subsequent trapping cessation, extensive reintroduction programs, and natural recolonization have helped fishers to reclaim much of their historical range. The degree to which these processes have impacted genetic structure in this species, however, remains unknown. We used 11 microsatellites from tissue samples (n = 432) of fishers to characterize contemporary population structure in light of historical population structure and thus to determine the relative influence of anthropogenic disturbances and natural landscape features in shaping genetic structure of the contemporary population. Our results indicated that 3 well-differentiated contemporary populations are present that correspond well with what would be expected based on their reported history. A course barrier to dispersal appears in the western portion of the study area associated with several lakes including Lake George and Great Sacandaga Lake. Large-scale reintroduction efforts and natural recolonizations have largely had predictable impacts on population structure. An important exception is the substantial impact of the reintroduction of fishers to Vermont.