Shifting dispersal modes at an expanding species’ range margin

While it is generally recognized that noncontiguous (long‐distance) dispersal of small numbers of individuals is important for range expansion over large geographic areas, it is often assumed that colonization on more local scales proceeds by population expansion and diffusion dispersal (larger numbers of individuals colonizing adjacent sites). There are few empirical studies of dispersal modes at the front of expanding ranges, and very little information is available on dispersal dynamics at smaller geographic scales where we expect contiguous (diffusion) dispersal to be prevalent. We used highly polymorphic genetic markers to characterize dispersal modes at a local geographic scale for populations at the edge of the range of a newly invasive grass species (Brachypodium sylvaticum) that is undergoing rapid range expansion in the Pacific Northwest of North America. Comparisons of Bayesian clustering of populations, patterns of genetic diversity, and gametic disequilibrium indicate that new populations are colonized ahead of the invasion front by noncontiguous dispersal from source populations, with admixture occurring as populations age. This pattern of noncontiguous colonization was maintained even at a local scale. Absence of evidence for dispersal among adjacent pioneer sites at the edge of the expanding range of this species suggests that pioneer populations undergo an establishment phase during which they do not contribute emigrants for colonization of neighbouring sites. Our data indicate that dispersal modes change as the invasion matures: initial colonization processes appear to be dominated by noncontiguous dispersal from only a few sources, while contiguous dispersal may play a greater role once populations become established.     

Consequences of large-scale processes for the conservation of bird populations

1.  Detailed studies of population ecology are usually carried out in relatively restricted areas in which emigration and immigration play a role. We used a modelling approach to explore the population consequences of such dispersal and applied ideas from our simulations to the conservation of wild birds.
2.  Our spatial model incorporates empirically derived variation in breeding output between habitats, density dependence and dispersal. The outputs indicate that dispersal can have considerable consequences for population abundance and distribution. The abundance of a species within a patch can be markedly affected by the surrounding habitat matrix.
3.  Dispersal between habitats may result in lower population densities at the edge of good quality habitat blocks and could partially explain why some species are restricted to large habitat fragments.
4.  Habitat deterioration may not only lead to population declines within that habitat but also in adjacent habitats of good quality. This may confound studies attempting to diagnose population declines.
5.  Although mobile species have the advantages of colonizing sites within metapopulations, dispersal into poorer quality territories may markedly reduce total populations.
6.  There are two main approaches to conservation: one is to concentrate on establishing and maintaining protected areas, while the other involves conservation of the wider countryside. If dispersal is an important process then protecting only isolated areas may be insufficient to maintain the populations within them.     

Allee effects may slow the spread of parasites in a coastal marine ecosystem

Allee effects are thought to mediate the dynamics of population colonization, particularly for invasive species. However, Allee effects acting on parasites have rarely been considered in the analogous process of infectious disease establishment and spread. We studied the colonization of uninfected wild juvenile Pacific salmon populations by ectoparasitic salmon lice (Lepeophtheirus salmonis) over a 4-year period. In a data set of 68,376 fish, we observed 85 occurrences of precopular pair formation among 1,259 preadult female and 613 adult male lice. The probability of pair formation was dependent on the local abundance of lice, but this mate limitation is likely offset somewhat by mate-searching dispersal of males among host fish. A mathematical model of macroparasite population dynamics that incorporates the empirical results suggests a high likelihood of a demographic Allee effect, which can cause the colonizing parasite populations to die out. These results may provide the first empirical evidence for Allee effects in a macroparasite. Furthermore, the data give a rare detailed view of Allee effects in colonization dynamics and suggest that Allee effects may dampen the spread of parasites in a coastal marine ecosystem.     

Invasion ecology of wild pigs (<Emphasis Type="Italic">Sus scrofa</Emphasis>) in Florida,USA: the role of humans in the expansion and colonization of an invasive wild ungulate

Wild pigs (Sus scrofa) are the most widely distributed invasive wild ungulate in the United States, yet the factors that influence wild pig dispersal and colonization at the regional level are poorly understood. Our objective was to use a population genetic approach to describe patterns of dispersal and colonization among populations to gain a greater understanding of the invasion process contributing to the expansion of this species. We used 52 microsatellite loci to produce individual genotypes for 482 swine sampled at 39 locations between 2014 and 2016. Our data revealed the existence of genetically distinct subpopulations (F _ST  = 0.1170, p < 0.05). We found evidence of both fine-scale subdivision among the sampling locations, as well as evidence of long term genetic isolation. Several locations exhibited significant admixture (interbreeding) suggesting frequent mixing of individuals among locations; up to 14% of animals were immigrants from other populations. This pattern of admixture suggested successive rounds of human-assisted translocation and subsequent expansion across Florida. We also found evidence of genetically distinct populations that were isolated from nearby populations, suggesting recent introduction by humans. In addition, proximity to wild pig holding facilities was associated with higher migration rates and admixture, likely due to the escape or release of animals. Taken together, these results suggest that human-assisted movement plays a major role in the ecology and rapid population growth of wild pigs in Florida.     

Protected areas for kiwi in mainland forests of New Zealand: how large should they be?

This paper examines, theoretically, how dispersal affects the viability of brown kiwi populations in protected areas of different size. Brown kiwi are threatened by introduced mammalian predators in mainland forests and are likely to persist only in managed forests where predators are controlled. In each protected area, the kiwi population will function as a net source, with an outflow of juveniles into the adjoining forest and minimal backflow into the reserve. Computer simulations show the minimum area of forest required for population viability increases non-linearly as the mean dispersal distance of juveniles increases. Preliminary measurements of the mean dispersal distance of brown kiwi in the wild suggest kiwi populations are unlikely to be viable in protected areas of less than 10 000 ha. Our estimate of the forest area requirement for viable populations of brown kiwi agrees with those derived by earlier workers using biogeographic techniques.     

Mitochondrial haplotype distribution,seed dispersal and patterns of postglacial expansion of ponderosa pine

Maternally inherited mtDNA in a secondary contact zone of ponderosa pine revealed a cline less than 10 km wide - much narrower than previously described. A survey of 76 populations gave no evidence either of intermixing or of a mosaic contact zone. Such sharp contact zones are consistent with diffusive range expansion, rather than long distance colonization. However, evidence for long distance seed dispersal events was found in two populations where haplotypes were observed far from their main area of occurrence. The results suggest a small number of long distance colonists with diffusive dispersal from these centres.     

Colonization in metapopulations: a review of theory and observations

In metapopulation dynamics turnover of populations in isolated patches may be frequent. Regional survival of a species in such a system with frequent extinctions hinges on its colonization ability. Colonization is more than just dispersal; when a propagule reaches a new patch it faces higher extinction probabilities than does an established population. Extinction models as well as empirical data suggest that a large propagule with a potential for rapid increase in a varying environment, or with a low mortality rate in an environment perceived as constant, has a higher probability of successful colonization. Large variation in population size when it is still small increases the risk of failure. Factors introducing such variation are demographic stochasticity and environmental variation. It is hard to single out demographic traits that ensure good colonizing ability, since colonization can be achieved in many different ways, but generalists and species with self-fertilization seem to be superior.     

Neutral processes forming large clones during colonization of new areas

In species reproducing both sexually and asexually clones are often more common in recently established populations. Earlier studies have suggested that this pattern arises due to natural selection favouring generally or locally successful genotypes in new environments. Alternatively, as we show here, this pattern may result from neutral processes during species’ range expansions. We model a dioecious species expanding into a new area in which all individuals are capable of both sexual and asexual reproduction, and all individuals have equal survival rates and dispersal distances. Even under conditions that favour sexual recruitment in the long run, colonization starts with an asexual wave. After colonization is completed, a sexual wave erodes clonal dominance. If individuals reproduce more than one season, and with only local dispersal, a few large clones typically dominate for thousands of reproductive seasons. Adding occasional long‐distance dispersal, more dominant clones emerge, but they persist for a shorter period of time. The general mechanism involved is simple: edge effects at the expansion front favour asexual (uniparental) recruitment where potential mates are rare. Specifically, our model shows that neutral processes (with respect to genotype fitness) during the population expansion, such as random dispersal and demographic stochasticity, produce genotype patterns that differ from the patterns arising in a selection model. The comparison with empirical data from a post‐glacially established seaweed species (Fucus radicans) shows that in this case, a neutral mechanism is strongly supported.     

秦岭大熊猫种群扩散格局及研究方法

种群扩散格局是研究种群扩散规律和机制的关键信息,也是制定物种保护对策的重要基础。大型动物种群扩散格局研究方法为扩散生态学研究的薄弱领域,并制约扩散生态学的发展。以秦岭大熊猫为研究对象,根据2000年以来的种群调查数据,基于大熊猫领域的特性,利用GIS的扩展区分析功能和景观分析方法研究了大熊猫种群分布区及动态;基于聚集的特性,利用GIS的核密度分析功能对大熊猫种群多度和聚集状况及空间变化进行了分析。发现2012年的秦岭大熊猫种群分布区较2000年增加5.5%(即15307.8hm~2),高密度种群聚集区从2处变成1处,种群聚集程度进一步增加、聚集格局的完整性大大提升,尤以中密度聚集区增长最显著,种群格局呈明显的分布区扩张、聚集度增加的态势。表明基于物种的生物学特性,立足于种群分布和多度格局变化,通过长期调查和监测可以有效掌握物种的种群扩散格局;大型动物可根据其生物学特性探索可行的方法与量化种群扩散的参数来研究其扩散格局,从而促进大型动物种群扩散研究的开展。     

Genetic structure of the cyclic fossorial water vole (Arvicola terrestris): landscape and demographic influences

Genetic structure can be strongly affected by landscape features and variation through time and space of demographic parameters such as population size and migration rate. The fossorial water vole (Arvicola terrestris) is a cyclic species characterized by large demographic fluctuations over short periods of time. The outbreaks do not occur everywhere at the same time but spread as a wave at a regional scale. This leads to a pattern of large areas (i.e. some hundreds of km2), each with different vole abundances, at any given time. Here, we describe the abundance and genetic structures in populations of the fossorial water vole. We use the data to try to understand how landscape and demographic features act to shape the genetic structure. The spatial variability of vole abundance was assessed from surface indices, collected in spring 2002 (April) in eastern central France. Genetic variability was analysed using eight microsatellite loci at 23 localities sampled between October 2001 and April 2002. We found some congruence between abundance and genetic structures. At a regional scale, the genetic disruptions were associated with both sharp relief and transition between an area of low abundance and another of high abundance. At a local scale, we observed a variation of the isolation-by-distance pattern according to the abundance level of vole populations. From these results we suggest that the dispersal pattern in cyclic rodent populations varies throughout the demographic cycle.     

Genetic analysis of complex demographic scenarios: spatially expanding populations of the cane toad, Bufo marinus

Inferring the spatial expansion dynamics of invading species from molecular data is notoriously difficult due to the complexity of the processes involved. For these demographic scenarios, genetic data obtained from highly variable markers may be profitably combined with specific sampling schemes and information from other sources using a Bayesian approach. The geographic range of the introduced toad Bufo marinus is still expanding in eastern and northern Australia, in each case from isolates established around 1960. A large amount of demographic and historical information is available on both expansion areas. In each area, samples were collected along a transect representing populations of different ages and genotyped at 10 microsatellite loci. Five demographic models of expansion, differing in the dispersal pattern for migrants and founders and in the number of founders, were considered. Because the demographic history is complex, we used an approximate Bayesian method, based on a rejection-regression algorithm, to formally test the relative likelihoods of the five models of expansion and to infer demographic parameters. A stepwise migration-foundation model with founder events was statistically better supported than other four models in both expansion areas. Posterior distributions supported different dynamics of expansion in the studied areas. Populations in the eastern expansion area have a lower stable effective population size and have been founded by a smaller number of individuals than those in the northern expansion area. Once demographically stabilized, populations exchange a substantial number of effective migrants per generation in both expansion areas, and such exchanges are larger in northern than in eastern Australia. The effective number of migrants appears to be considerably lower than that of founders in both expansion areas. We found our inferences to be relatively robust to various assumptions on marker, demographic, and historical features. The method presented here is the only robust, model-based method available so far, which allows inferring complex population dynamics over a short time scale. It also provides the basis for investigating the interplay between population dynamics, drift, and selection in invasive species.     

Measuring ungulate tolerance to human with flight distance: a reliable visitor management tool?

Tourist-induced behavioural changes in large vertebrates are of concern for protected area management as they trigger a trade-off: large vertebrates attract visitors, but induced behavioral changes can reduce animal fitness and cause animals to avoid tourist-frequented zones. Behavioural response of animals to tourists is often studied to ensure informed management decision-making, a task frequently supported by flight distance analysis. In this context, guanaco (Lama guanicoe) response to tourists was studied in a protected area using two complementary methods: flight distance and sighting frequency analyses. Flight response analyses show that guanacos develop a considerable tolerance to vehicles and pedestrians in tourist areas, a reaction that extends approximately 500 m around visited areas. Such analyses thus point to (i) few areas being underused as a consequence of human presence and (ii) a low risk of tame animals being poached outside the park, all potentially leading to the conclusion that tourist visits are sustainable. However, guanaco sighting frequencies during 107 fieldwork days along 3 years show a significant reduction in sightings on days with higher Park visitor numbers. Moreover we present a formal procedure for the definition of a threshold for this to happen (247 visitors/day in our case). This suggests the potential risk of negative effects on guanaco population and a lower probability of guanaco sightings if Park visitor numbers rise beyond current figures. Results allow to conclude that assessment of human disturbance to flagship species in protected areas requires further methods in addition to flight distance.     

Effects of Culling on Mesopredator Population Dynamics

Anthropogenic changes in land use and the extirpation of apex predators have facilitated explosive growth of mesopredator populations. Consequently, many species have been subjected to extensive control throughout portions of their range due to their integral role as generalist predators and reservoirs of zoonotic disease. Yet, few studies have monitored the effects of landscape composition or configuration on the demographic or behavioral response of mesopredators to population manipulation. During 2007 we removed 382 raccoons (Procyon lotor) from 30 forest patches throughout a fragmented agricultural ecosystem to test hypotheses regarding the effects of habitat isolation on population recovery and role of range expansion and dispersal in patch colonization of mesopredators in heterogeneous landscapes. Patches were allowed to recolonize naturally and demographic restructuring of patches was monitored from 2008–2010 using mark-recapture. An additional 25 control patches were monitored as a baseline measure of demography. After 3 years only 40% of experimental patches had returned to pre-removal densities. This stagnant recovery was driven by low colonization rates of females, resulting in little to no within-patch recruitment. Colonizing raccoons were predominantly young males, suggesting that dispersal, rather than range expansion, was the primary mechanism driving population recovery. Contrary to our prediction, neither landscape connectivity nor measured local habitat attributes influenced colonization rates, likely due to the high dispersal capability of raccoons and limited role of range expansion in patch colonization. Although culling is commonly used to control local populations of many mesopredators, we demonstrate that such practices create severe disruptions in population demography that may be counterproductive to disease management in fragmented landscapes due to an influx of dispersing males into depopulated areas. However, given the slow repopulation rates observed in our study, localized depopulation may be effective at reducing negative ecological impacts of mesopredators in fragmented landscapes at limited spatial and temporal scales.     

Colonization of Languedoc vineyards by phytoseiid mites (Acari: Phytoseiidae): influence of wind and crop environment

A natural increase of phytoseiid mite populations (Kampimodromus aberrans, Typhlodromus pyri and Phytoseius plumifer) was observed in vineyards in Languedoc, Burgundy and Corsica under integrated pest management strategies. The aim of the present study was to characterize the mechanisms of this colonization in space and time in Languedoc. The abundance of phytoseiid mites in the vegetation close to three grape fields was determined twice a year (May and July). Aerial (funnels with water) and soil (felt strip) traps were placed in and around grape fields, in order to assess the colonization potential provided by aerial dispersal and ambulatory locomotion. The populations of phytoseiid mites in the crops were studied twice a month in order to gain information on the make up of the dispersal populations. The species K. aberrans was found in largest quantities in the traps, in the natural vegetation and in the crops. Predatory mite dispersal occurred essentially by aerial dispersal and was dependent on the wind intensity and wind direction. Identical sex ratios were observed in migrant populations and in populations present in the grape fields, woody areas and hedges. A large proportion of immatures was found to move by aerial dispersal. The colonization potential (rapidity, intensity and regularity) was directly associated with the abundance of the phytoseiids and the proximity of natural vegetation. A deep, dense and tall woody area containing suitable host plants for predatory mites constituted the most stable source of phytoseiid mites. Natural colonization of vineyards provides considerable phytoseiid mite potential that could be managed in an agricultural landscape.     

The feeding habits of wolves in relation to large prey availability in northern Italy

We investigated wolf feeding habits in relation to the abundance of wild and domestic ungulates to test the hypothesis that large prey are preferred and that their abundance affects the use of other food categories and diet breadth. We determined diet composition by scat analysis from December 1987 to December 1992. The research was carried out in three study areas located in northern Italy and characterised by marked differences in wild and domestic ungulate abundance. In study area A (low wild and domestic ungulate availability) fruits, livestock, other vertebrates and wild ungulates made up the bulk of the diet (71% in volume). In area B (high availability of livestock) wolf diet was mainly based on sheep and wild boars (80% in volume). In study area C (high availability of wild ungulates) wild ungulates were the main food of wolves (90% in volume). Significant differences were found among study areas in the mean percentage volume of all food categories and in particular for wild ungulates, livestock, other vertebrates and fruits (p < 0.0001 in all cases). Diet breadth decreased in areas with high availability of large wild and domestic herbivores. The use of livestock species was lower where there was high abundance, richness and diversity of the wild ungulate guild. Selection for wild ungulate species was partially affected by their abundance: however other factors as prey social behaviour, adaptability to the habitat (for introduced species), and body size could have an important role in species selection by wolves. In particular in area C wild boars were selected for, roe and red deers avoided, and fallow deers and mouflons used as available. Livestock species were used in relation to their abundance and accessibility, in particular sheep were selected for and cattle avoided; but if calves bom in the pastures were considered as the only available cattle, they were selected for and sheep were used as available. Large and in particular wild herbivores were found to be of great importance for the wolf population maintenance in northern Italy, one of the most important recovery areas of Mediterranean wolves.     

Deer-mediated expansion of a rare plant species

Numerous plant colonizations have been putatively attributed to deer, based on plant species traits, fur brushing or dung analyses. But, in woodlands, direct links between the expansion of zoochorous plant species and ungulate presence have seldom been reported. Based on coupled floristic and browsing surveys, repeated in time, we analysed the causes of the spatio-temporal progression of the epizoochorous species Cynoglossum germanicum over 30 years in a network covering an 11000 ha forested area in north-eastern France. In this area, deer populations reached a peak in the 1970s, then were reduced in order to meet forest management requirements. Although initially rare and protected locally, C. germanicum has displayed an unexpected fast colonization rate during the last few decades but only in the northern part of the forest, which previously had the highest animal populations. Absent in the initial 1976 survey, C. germanicum occurred in 8% of the plots in 1981, then 46% in 2006. Logistic regression models revealed that the probability of occurrence of C. germanicum in 2006 increased not only with light indicator values, in accordance with its ecological requirements, but also with past deer browsing pressure. This result provides direct evidence of long-lasting impacts of deer populations on plant species distribution. Combining two complementary traits, animal transport and herbivory avoidance, C. germanicum benefited from epizoochorous dispersal and, once established, was protected from deer browsing by the presence of toxic proteins in its tissues. Due to the triggering role of ungulates, this species switched from the status of rare to that of colonizer within only a few decades.     

Mitochondrial differentiation in a polymorphic land snail: evidence for Pleistocene survival within the boundaries of permafrost

The genetic differentiation of populations having colonized formerly unsuitable habitats after the Pleistocene glaciations depends to a great extent on the speed of expansion. Slow dispersers maintain their refugial diversity whereas fast dispersal leads to a reduction of diversity in the newly colonized areas. During the Pleistocene, almost the entire current range of the land snail Arianta arbustorum has repeatedly been covered with ice or been subjected to permafrost. Owing to the low potential for dispersal of land snails, slow (re)colonization of the wide range from southern refugia can be excluded. Alternatively, fast, passive dispersal from southern refugia or survival in and expansion from multiple refugia within the area subjected to permafrost may account for the current distribution. To distinguish between these scenarios we reconstructed a phylogeography based on the sequences of a fragment of the cytochrome oxidase I from 133 individuals collected at 45 localities and analysed the molecular variance. Seventy-five haplotypes were found that diverged on average at 7.52% of positions. This high degree of diversity suggests that A. arbustorum is an old species in which the population structure, isolation and the hermaphroditic nature have reduced the probability of lineage extinction. The genetic structure was highly significant with the highest variance partition found among regions. Geographic distance and mitochondrial differentiation were not congruent. Lineages had overlapping ranges. The clear genetic differentiation and the patchy pattern of haplotype distribution suggest that colonization of formerly unsuitable habitats was mainly achieved from multiple populations from within the permafrost area.     

High haplotype diversity with fine-scale structure in a recently established population of an endangered orchid

New populations of endangered plants may establish outside of protected areas if a suitable habitat becomes available. However, it is unclear whether such populations are genetically uniform, as a result of a founder effect, or whether they contain genetic variation resulting from continuous gene flow from source populations, and subsequent recruitment after establishment. We addressed this by examining haplotype variation in the endangered orchid, Anacamptis robusta, which has formed a new population outside of its protected area within the past 20 years. To assess population growth, the number of A. robusta flowering plants were counted every year for 22 years in both the new population and populations within the reserve. Haplotype diversity and fine-scale structure were examined with spatial autocorrelation analysis, both in the new population and representative populations from the protected area. The number of flowering individuals in the new population increased from 9 to 2,277 between 2003 and 2012, whereas within the reserve flowering population sizes varied between years. Seventeen of 23 haplotypes detected for A. robusta were detected in the new population, with seven of these occurring more frequently in the new population than reserve populations. In the new population, there was strong fine-scale spatial structure of haplotypes, similar to patterns found in large populations from the protected area, suggesting multiple colonization events and subsequent local recruitment. This highlights that ongoing demographic and genetic monitoring of plant populations is vital to improve our understanding of population colonization and the conservation of narrow endemics.     

Genetic diversity and spatial genetic structure of African wild dogs (<Emphasis Type="Italic">Lycaon pictus</Emphasis>) in the Greater Limpopo transfrontier conservation area

The Greater Limpopo Transfrontier Conservation Area (GLTFCA) is one of the last refuges for the endangered African wild dog and hosts roughly one-tenth of the global population. Wild dogs in this area are currently threatened by human encroachment, habitat fragmentation and scarcity of suitable connecting habitat between protected areas. We derived genetic data from mitochondrial and nuclear markers to test the following hypotheses: (i) demographic declines in wild dogs have caused a loss of genetic variation, and (ii) Zimbabwean and South African populations in the GLTFCA have diverged due to the effects of isolation and genetic drift. Genetic patterns among five populations, taken with comparisons to known information, illustrate that allelic richness and heterozygosity have been lost over time, presumably due to effects of inbreeding and genetic drift. Genetic structuring has occurred due to low dispersal rates, which was most apparent between Kruger National Park and the Zimbabwean Lowveld. Immediate strategies to improve gene flow should focus on increasing the quality of habitat corridors between reserves in the GLTFCA and securing higher wild dog survival rates in unprotected areas, with human-mediated translocation only undertaken as a last resort.     

Long-distance dispersal in red foxes <Emphasis Type="Italic">Vulpes vulpes</Emphasis> revealed by GPS tracking

Dispersal is a fundamental process that facilitates population and range expansion by providing a mechanism for colonization and metapopulation linkages. Yet quantifying the dispersal process, particularly long-distance dispersal events, has been inherently difficult due to technological and observational limitations. Additionally, dispersal distance calculated as the straight-line distance between initiation and settlement fails to account for the actual movement path of the animal during dispersal. Here, we highlight six long-distance dispersal events, representing some of the longest dispersal distances recorded for red foxes. Cumulative dispersal movements ranged from 132 to 1036 km and occurred within both sexes (1 female, 5 males). With one exception, dispersal events ranged from 7 to 22 days and tended to be directed north-northwest. Importantly, cumulative movements were up to five times longer than straight-line distances, with two foxes traveling an additional 114 and 256 km before returning to, and settling in, areas previously encountered during dispersal. This suggests a role of habitat assessment and homing behavior during dispersal and indicates that the capacity and potential for dispersal are not limiting factors to either sex in a red fox population. Dispersal capacity should thus be considered regarding transboundary management and disease control of red fox populations.