USING ISOLATION BY DISTANCE AND EFFECTIVE DENSITY TO ESTIMATE DISPERSAL SCALES IN ANEMONEFISH

Robust estimates of dispersal are critical for understanding population dynamics and local adaptation, as well as for successful spatial management. Genetic isolation by distance patterns hold clues to dispersal, but understanding these patterns quantitatively has been complicated by uncertainty in effective density. In this study, we genotyped populations of a coral reef fish (Amphiprion clarkii) at 13 microsatellite loci to uncover fine‐scale isolation by distance patterns in two replicate transects. Temporal changes in allele frequencies between generations suggested that effective densities in these populations are 4–21 adults/km. A separate estimate from census densities suggested that effective densities may be as high as 82–178 adults/km. Applying these effective densities with isolation by distance theory suggested that larval dispersal kernels in A. clarkii had a spread near 11 km (4–27 km). These kernels predicted low fractions of self‐recruitment in continuous habitats, but the same kernels were consistent with previously reported, high self‐recruitment fractions (40–60%) when realistic levels of habitat patchiness were considered. Our results suggested that ecologically relevant larval dispersal can be estimated with widely available genetic methods when effective density is measured carefully through cohort sampling and ecological censuses, and that self‐recruitment studies should be interpreted in light of habitat patchiness.     

Sibling resemblance in natal dispersal distance and direction in the Ortolan Bunting Emberiza hortulana

Natal dispersal distance and direction determine the likelihood that siblings will settle close together and hence the risk of inbreeding. Several studies have shown a sibling resemblance in dispersal distance, but few studies have analysed sibling resemblance in dispersal direction or the distance between siblings after dispersal at the landscape level. I studied the entire Norwegian population of Ortolan Buntings Emberiza hortulana, which is patchily distributed in an area covering c. 500 km². Males and females did not differ in dispersal distance (overall median 3.7 km), but directions were different. Natal dispersal distances and directions were positively related within sibling pairs, but comparisons with control individuals suggested that any effects were due to spatial effects of configuration of habitat patches in the study area. Brother–sister pairs (n = 16) were at least as similar as brother–brother pairs (n = 18). Distance between siblings after natal dispersal increased with dispersal distance, but even so, five of 35 sibling pairs settled < 1 km apart, despite dispersal distances of 8.3–9.9 km for two of these pairs. Including movements later in life, eight sibling pairs were < 1 km apart at some time (four pairs of brothers and four brother–sister pairs), and in one case a brother mated with its sister. Another case of mating between close relatives (father and daughter) involved short female natal dispersal. These data indicate that female‐biased natal dispersal and long‐distance dispersal may reduce, but do not exclude, the possibility of inbreeding.     

Reduced dispersal propensity in the wingless waterstrider Aquarius najas in a highly fragmented landscape

Dispersal behaviour of animals in fragmented habitats has generated intensive theoretical attention but empirical data on the evolution of dispersal are still relatively scarce. Theory predicts reduced dispersal propensity in small and isolated habitat patches. We tested these predictions in the waterstrider Aquarius najas, a wingless species with special habitat demands. Flightlessness constrains insect dispersal and as a stream specialist A. najas cannot survive on still water. Lakes therefore represent a dispersal barrier for this species. We measured dispersal propensity of the waterstrider A. najas which originated from ten fragmented populations. In the experiment, we transplanted laboratory-grown individuals to the field. We did not find differences between sex in dispersal propensity. However, we found that waterstriders that originated from small and isolated patches moved less than individuals from large and more continuous habitats. This suggests that the cost of dispersal over hostile surrounding habitats may be high. We conclude that a low likelihood of dispersal in A. najas is an adaptation to small and isolated stream habitats.     

INFLUENCE OF VEGETATION,TOPOGRAPHY, AND ROADS ON COUGAR MOVEMENT IN SOUTHERN CALIFORNIA

Abstract: Models of individual movement can help conserve wide-ranging carnivores on increasingly human-altered landscapes, and cannot be constructed solely by analyzing the daytime resting locations typically collected in carnivore studies. We examined the movements of 10 female and 7 male cougars (Puma concolor) at 15-min intervals during 44 nocturnal or diel periods of hunting or traveling in the Santa Ana Mountain Range of southern California, USA, between 1988 and 1992. Cougars tended to move in a meandering path (mean turning angle ∼54°), and distance moved (mean and mode ∼300 m) was not correlated with turning angle. Cougars used a broader range of habitats for nocturnal or diel movements than for previously described daybed locations for this same population. Riparian vegetation ranked highest in a compositional analysis of vegetation types selected during movement; grassland, woodland and urbanized sites were least selected. During periods of stasis (we presume many of these were stalking locations), patterns of selection were less marked. Cougars spent a disproportionate amount of time in highly ranked vegetation types, and traveled slowest through riparian habitats and fastest through human-dominated areas. Our results suggest that travel speed may provide an efficient index of habitat selection in concert with other types of analysis. Hunting or traveling individuals consistently used travel paths that were less rugged than their general surroundings. Traveling cougars avoided 2-lane paved roads, but dirt roads may have facilitated movement. Maintenance and restoration of corridors between large wildlands is essential to conserving cougars in southern California. Our results indicate that riparian vegetation, and other vegetation types that provide horizontal cover, are desirable features in such corridors, that dirt roads should not impede cougar use of corridors, and that corridors should lie along routes with relatively gentle topography. Our results suggest that cougars do not key in on highway-crossing structures in a way that creates a prey trap. Our empirical frequency distributions of distances and turning angles, along with cougar responses to vegetation, topography, and roads can help parameterize an individually-based movement model for cougars in human-altered landscapes.     

Multistage,Long-Range Natal Dispersal by a Global Positioning System-Collared Scandinavian Wolf

Abstract: We document a new record dispersal for wolves worldwide. The natal straight-line dispersal distance of a Global Positioning System-collared female wolf from the Scandinavian population was 1,092 km from southeast Norway to northeast Finland, with a multistage actual travel distance of >10,000 km. Natural gene flow to the isolated, inbred Scandinavian wolf population may occur if survival of dispersers is improved.     

Morphological variation associated with dispersal capacity in a tree‐killing bark beetle Dendroctonus ponderosae Hopkins

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Increases in Midwestern cougars despite harvest in a source population

Although cougars (Puma concolor) appear to be recolonizing the midwestern United States, there is concern that hunting in source populations (primarily the Black Hills, SD and WY, USA) may prevent cougars from dispersing eastward. We use carcass data of cougars (n =147 carcasses at known locations, of which 111 were of known sex) in the Midwest collected during 1990–2015 to quantify whether cougar hunting in the Black Hills affected cougar distribution and presence in the Midwest. We separated carcass data into 2 time periods: before hunting in the Black Hills (i.e., pre-hunt; 1990–2004) and after hunting (i.e., post-hunt; 2005–2015). We hypothesized that if hunting prevented dispersal into the Midwest, cougar distribution would be random and their presence less, relative to the pre-hunt period. We also were interested in sex ratios of carcasses over time, given the importance of that demographic metric to the potential establishment of viable populations. During the pre-hunt period, 25 carcasses were dispersed randomly in the Midwest. During the post-hunt period, we found nearly 4 times the number of carcasses in the Midwest (n = 86), carcasses were significantly clustered, and a greater percentage of carcasses were female (pre-hunt n = 6 [24%]; post-hunt n = 27 [31%]). Relative to the pre-hunt period, we observed a 460-km northward shift in the directional distribution of carcass locations during the post-hunt period. These findings do not support the idea that hunting in the Black Hills has prevented cougar presence from increasing in the Midwest. Alternatively, we suggest the potential for immigration from cougar populations farther to the west as an explanation for the increase in cougar presence (particularly females) confirmed after the initiation of cougar hunting in the Black Hills. © 2019 The Wildlife Society.     

Effects of conditionally expressed phenotypes and environment on amphibian dispersal in nature

Individuals vary greatly in the distance they disperse, and in doing so, strongly affect ecological and evolutionary processes. Dispersal, when viewed as a component of phenotype, can be affected independently or jointly by environment. However, among taxa with complex life cycles that occupy different habitats over ontogeny, the effects of environment on dispersal and the interaction between environment and phenotype remains poorly understood. Here, we conducted a field experiment to measure how dispersal distance was affected by phenotype, environment experienced before and after metamorphosis, and their interaction. We manipulated the environment encountered by a pond‐breeding salamander Ambystoma annulatum during the aquatic larval stage and again as dispersing terrestrial juveniles. After assaying juvenile phenotype (exploration behavior, body condition, and morphology), we then measured the initial distance dispersed by juveniles. The distance moved by dispersing salamanders was affected by attributes of both larval and juvenile habitat, with salamanders that encountered low quality habitat in either life stage moving the farthest. However, we did not find support for an interactive effect of phenotype and environment affecting the distance moved by dispersers. Interestingly, exploration behavior explained the distance moved by philopatric animals but not dispersing ones. Our findings indicate that the environment experienced before metamorphosis can affect juvenile dispersal behavior, and demonstrates the need to consider dispersal in species with complex life cycles to understand the coupling between local and regional population dynamics.     

What is the required minimum landscape size for dispersal studies?

Among small animals dispersal parameters are mainly obtained by traditional methods using population studies of marked individuals. Dispersal studies may underestimate the rate and distance of dispersal, and be biased because of aggregated habitat patches and a small study area. The probability of observing long distance dispersal events decreases with distance travelled by the organisms. In this study a new approach is presented to solve this methodological problem. An extensive mark-release-recapture programme was performed in an area of 81 km(2) in southern Sweden. To estimate the required size of the study area for adequate dispersal measures we examined the effect of study area size on dispersal distance using empirical data and a repeated subsampling procedure. In 2003 and 2004, two species of diurnal burnet moths (Zygaenidae) were studied to explore dispersal patterns. The longest confirmed dispersal distance was 5600 m and in total 100 dispersal events were found between habitat patches for the two species. The estimated dispersal distance was strongly affected by the size of the study area and the number of marked individuals. For areas less than 10 km(2) most of the dispersal events were undetected. Realistic estimates of dispersal distance require a study area of at least 50 km(2). To obtain adequate measures of dispersal, the marked population should be large, preferably over 500 recaptured individuals. This result was evident for the mean moved distance, mean dispersal distance and maximum dispersal distance. In general, traditional dispersal studies are performed in small study areas and based on few individuals and should therefore be interpreted with care. Adequate dispersal measures for insects obtained by radio-tracking and genetic estimates (gene flow) is still a challenge for the future.     

Forest edges and fire ants alter the seed shadow of an ant-dispersed plant

Exotic species invade fragmented, edge-rich habitats readily, yet the distinct impacts of habitat edges and invaders on native biota are rarely distinguished. Both appear detrimental to ant-dispersed plants such as bloodroot, Sanguinaria canadensis. Working in northeastern Georgia (USA), an area characterized by a rich ant-dispersed flora, fragmented forests, and invasions by the red imported fire ant, Solenopsis invicta , I monitored the interactions between ants and S. canadensis seeds in uninvaded forest interiors, uninvaded forest edges, invaded forest interiors, and invaded forest edges. I observed 95% of the seed dispersal events that occurred within the 60-min observation intervals. Seed collection rates were similar among all four (habitat × invasion) groups. The presence of invasive ants had a strong effect on seed dispersal distance: S. invicta collected most seeds in invaded sites, but was a poorer disperser than four of five native ant taxa. Habitat type (interior versus edge) had no effect on seed dispersal distance, but it had a strong effect on seed dispersal direction. Dispersal towards the edge was disproportionately rare in uninvaded forest edges, and ants in those habitats moved the average dispersed seed approximately 70 cm away from that edge. Dispersal direction was also skewed away from the edge in uninvaded forest interiors and invaded forest edges, albeit non-significantly. This biased dispersal may help explain the rarity of myrmecochorous plants in younger forests and edges, and their poor ability to disperse between fragments. This is the first demonstration that forest edges and S. invicta invasion influence seed dispersal destination and distance, respectively. These forces act independently.     

Sex, isolation and fidelity: unbiased long-distance dispersal in a terrestrial amphibian

Amphibians in general are considered poor dispersers and thus their dispersal curve should be dominated by short movements. Additionally, as male toads do not compete for females and sexual selection is by female choice, dispersal should be male-biased. Furthermore, since adults are site-loyal and polygynous, juveniles should move farther and faster than adults. We tested the hypotheses that dispersal would be limited and both sex- and age-biased in a population of Fowler's toads Bufo fowleri at Lake Erie, Ontario, Canada. Based on a mark-recapture study of 2816 toads, 1326 recaptured at least once, we found that although the toads did show high site fidelity, the dispersal curve was highly skewed with a significant "tail" where the maximum distance moved by an adult was 34 km. Dispersal was neither sex-biased nor age-biased despite clear theoretical predictions that dispersal should be biased towards males and juveniles. We conclude that the resource competition hypothesis of sex-biased dispersal does not predict dispersal tendencies as readily for amphibians as for mammals and birds. Toad dispersal only appears to be juvenile-biased because the juveniles are more abundant than the adults, not because they are the more active dispersers.     

Piping Plovers make decisions regarding dispersal based on personal and public information in a variable coastal ecosystem

ABSTRACT An increasing number of studies have revealed that public information influences dispersal decisions in a wide variety of species. However, few empirical studies have explored the effect of environmental predictability on the use of public information. We applied a model‐selection approach to data gathered from 1998 to 2006 in five eastern Canada provinces to determine the factors influencing breeding dispersal distances of Piping Plovers (Charadrius melodus melodus), a species that encounters substantial interannual changes in habitat quality. We examined the following hypotheses: (1) individuals that are unsuccessful breeders disperse greater distances than individuals that breed successfully and (2) dispersal distances of failed breeders are negatively correlated with the reproductive success of locally breeding conspecifics. We found that mean dispersal distance was greater for plovers that did not breed successfully the previous year (x′= 17.8 ± 27.4 [SD] km, N= 15) than for those that were successful (x′= 5.8 ± 26.4 km, N= 71). Failed breeders whose neighbors produced no fledged young dispersed over distances 34.8 times greater than those whose neighbors produced four fledglings. Our results show that public information is used in habitats whose environmental predictability is lower than predicted by theoretical studies. Thus, dispersal appears to be triggered by a combination of failure to hatch at least one egg, and the failure of neighboring pairs to produce fledglings. Individuals therefore appear to preferentially return to sites where breeding success is high. This may partly explain why Piping Plovers use only a fraction of the suitable breeding habitat within their geographic range. Consequently, although suitable habitat is present elsewhere within the breeding range, the loss of traditional nesting sites is likely to significantly impact productivity of this endangered species.     

Spatio‐temporal Genetic Structure of a Tropical Bee Species Suggests High Dispersal Over a Fragmented Landscape

Habitat destruction threatens biodiversity by reducing the amount of available resources and connectivity among geographic areas. For organisms living in fragmented habitats, population persistence may depend on dispersal, which maintains gene flow among fragments and can prevent inbreeding within them. It is centrally important to understand patterns of dispersal for bees living in fragmented areas given the importance of pollination systems and recently documented declines in bee populations. We used population and landscape genetic techniques to characterize patterns of dispersal over a large fragmented area in southern Costa Rica for the orchid bee species Euglossa championi. First, we estimated levels of genetic differentiation among forest fragments as ?_PT, an analog to the traditional summary statistic F_ST, as well as two statistics that may more adequately represent levels of differentiation, G’_ST and D_est. Second, we used a Bayesian approach to determine the number and composition of genetic groups in our sample. Third we investigated how genetic differentiation changes with distance. Fourth, we determined the extent to which deforested areas restrict dispersal. Finally, we estimated the extent to which there were temporal differences in allele frequencies within the same forest fragments. Within years we found low levels of differentiation even over 80 km, and no effect of land use type on level of genetic differentiation. However, we found significant genetic differentiation between years. Taken together our results suggest that there are high levels of gene flow over this geographic area, and that individuals show low site fidelity over time.     

Fine-scale population differentiation and gene flow in a terrestrial salamander (Plethodon cinereus) living in continuous habitat

Several recent studies have shown that amphibian populations may exhibit high genetic subdivision in areas with recent fragmentation and urban development. Less is known about the potential for genetic differentiation in continuous habitats. We studied genetic differentiation of red-backed salamanders (Plethodon cinereus) across a 2-km transect through continuous forest in Virginia, USA. Mark-recapture studies suggest very little dispersal for this species, whereas homing experiments and post-Pleistocene range expansion both suggest greater dispersal abilities. We used six microsatellite loci to examine genetic population structure and differentiation between eight subpopulations of red-backed salamanders at distances from 200 m to 2 km. We also used several methods to extrapolate dispersal frequencies and test for sex-biased dispersal. We found small, but detectable differentiation among populations, even at distances as small as 200 m. Differentiation was closely correlated with distance and both Mantel tests and assignment tests were consistent with an isolation-by-distance model for the population. Extrapolations of intergenerational variance in spatial position (sigma(2)<15 m(2)) and pair-wise dispersal frequencies (4 Nm < 25 for plots separated by 300 m) both suggest limited gene flow. Additionally, tests for sex-biased dispersal imply that dispersal frequency is similarly low for both sexes. We suggest that these low levels of gene flow and the infrequent dispersal observed in mark-recapture studies may be reconciled with homing ability and range expansion if dispersing animals rarely succeed in breeding in saturated habitats, if dispersal is flexible depending on the availability of habitat, or if dispersal frequency varies across the geographic range of red-backed salamanders.     

Spatial connectivity in an adult‐sedentary reef fish with extended pelagic larval phase

Understanding the spatial scale of demographic connectivity in marine reef fishes dispersing pelagic larvae is a challenging task because of the technical difficulties associated with tagging and monitoring the movements of progeny at early life stages. Several studies highlighted a strong importance of local retention with levels of dispersal of ecological significance restricted to short distances. To date little information is available in species where pelagic dispersal lasts for long periods of time. In this work, population structure and connectivity were studied in the grey triggerfish, Balistes capriscus. Grey triggerfish larvae and juveniles remain associated with floating Sargassum sp. beds for an estimated period of 4–7 months before settling on benthic habitats where they remain sedentary as adults. Analysis of genetic variation among populations along the continental shelf of the northern Gulf of Mexico and U.S. east coast, encompassing over 3,100 km of coastline, revealed homogeneous allele frequencies and a weak isolation‐by‐distance pattern. Moment and maximum‐likelihood estimates of dispersal parameters both indicated occurrence of large neighbourhoods with estimates of the dispersal distribution parameter σ of 914 and 780 km, respectively. Simulated distributions of dispersal distances using several distribution functions all featured substantial fractions of long‐distance dispersal events with the 90% percentiles of travel distance prior to settlement averaging 1,809 km. These results suggest a high dependency of local recruitment on the output of nonlocal spawning stocks located hundreds of kilometres away and a reduced role of local retention in this species.     

Ecological effects of fear: How spatiotemporal heterogeneity in predation risk influences mule deer access to forage in a sky‐island system

Forage availability and predation risk interact to affect habitat use of ungulates across many biomes. Within sky‐island habitats of the Mojave Desert, increased availability of diverse forage and cover may provide ungulates with unique opportunities to extend nutrient uptake and/or to mitigate predation risk. We addressed whether habitat use and foraging patterns of female mule deer (Odocoileus hemionus) responded to normalized difference vegetation index (NDVI), NDVI rate of change (green‐up), or the occurrence of cougars (Puma concolor). Female mule deer used available green‐up primarily in spring, although growing vegetation was available during other seasons. Mule deer and cougar shared similar habitat all year, and our models indicated cougars had a consistent, negative effect on mule deer access to growing vegetation, particularly in summer when cougar occurrence became concentrated at higher elevations. A seemingly late parturition date coincided with diminishing NDVI during the lactation period. Sky‐island populations, rarely studied, provide the opportunity to determine how mule deer respond to growing foliage along steep elevation and vegetation gradients when trapped with their predators and seasonally limited by aridity. Our findings indicate that fear of predation may restrict access to the forage resources found in sky islands.     

Pollen flow in Psiguria warscewiczii: a comparison of Heliconius butterflies and hummingbirds

Summary Pollen dispersal by male and female Heliconius butterflies and hermit hummingbirds was examined in a natural population of the tropical vine Psiguria warscewiczii. Fluorescent dyes were used to track flowers visited by individual pollinators. Mean distance of pollen dispersal was significantly higher by males than by female Heliconius. Butterflies dispersed pollen greater distances than did hummingbirds. There were no significant differences in dispersal distance among species of Heliconius. The number of flowers or plants to which dye was dispersed was greater for Heliconius than for hummingbirds.The home range behavior and site specificity exhibited by Heliconius, together with the preference by most Heliconius species in the study site for Psiguria led us to examine the relationship between the butterflies' home range and its daily foraging among Psiguria. A Heliconius mark-recapture program was conducted simultaneously with the dye program. This study showed that individual butterflies, for which extensive recaptures were made, visited an area of Psiguria plants on any one day that corresponded to the area in which those butterflies were caught or observed. The area over which a butterfly moved was termed its home range; within home ranges relative movement by butterflies was estimated as the mean distance between sequential captures. This estimate differed significantly among species of Heliconius; males of all species moving greater distances than females.     

Cougars are recolonizing the midwest: Analysis of cougar confirmations during 1990–2008

Although cougars (Puma concolor) were extirpated from much of midwestern North America around 1900, hard evidence of cougar presence has increased and populations have become established in the upper portions of the Midwest during the past 20 years. Recent occurrences of cougars in the Midwest are likely due to dispersal of subadult cougars into the region from established western populations, and may be indicative of further recolonization and range expansion. We compiled confirmed locations of cougars (i.e., via carcasses, tracks, photos, video, and DNA evidence) collected during 1990–2008 in 14 states and provinces of midwestern North America. We separated our study area into 2 regions (east and west), calculated number and types of confirmations, and assessed trends in confirmations during the study period. We recorded 178 cougar confirmations in the Midwest and the number of confirmations increased during the study period (r = 0.79, P ≤ 0.001). Confirmations by state or province ranged from 1 (Kansas, Michigan, and Ontario) to 67 (Nebraska). Carcasses were the most prevalent confirmation type (n = 56). Seventy-six percent of known-sex carcass confirmations were males, consistent with predominantly male-biased dispersal in cougars. More confirmations (P = 0.05) were recorded in the western region than the eastern region . Seventy-nine percent of cougar confirmations occurred within 50 km of highly suitable habitat (i.e., forest areas with steep terrain and low road and human densities). Given the number of cougar confirmations, the increasing frequency of occurrences, and that long-distance dispersal has been documented via radiocollared individuals, our research suggests that cougars are continuing to recolonize midwestern North America. © 2012 The Wildlife Society.