Movement activity and habitat use of Carabus ullrichii (Coleoptera: Carabidae): The forest edge as a mating site?

Some carabid species are not restricted to a single habitat only, but use various types of habitats. In these species, relatively little is known about the utilization of occupied habitats and factors affecting their movement within these habitats. In this study, we focus on the movement activity of ubiquitous Carabus ullrichii during its reproductive period at the border of two types of habitats, a meadow and a forest. We tracked 21 adult individuals using radio telemetry and recorded in total 1,687 position fixes. Movement activity was associated with the type of habitat and specific environmental conditions such as time of the day and air temperature. Both sexes activated preferably at dusk and during the night, although males were most active at temperatures around 15°C, while females showed no preference for temperature. Males were able to walk as fast as females, but they were more associated with forest edge. We assume that the inner edge of the forest could be used as a mating site and after mating males stay there and wait for new females with which to mate, while fertilized females disperse into the surroundings. They moved further into the closed forest where they were likely looking for oviposition sites and food resources to support reproduction. Exclusively females were recorded to visit the meadow at a greater distance from the forest edge and their movements there were almost always direct.     

Genetic Consequences of Habitat Fragmentation in Black-and-Gold Howler (<Emphasis Type="BoldItalic">Alouatta caraya</Emphasis>) Populations from Northern Argentina

Human-induced habitat fragmentation might seriously affect behavioural patterns and the survival of species whose ecological requirements strongly depend on specific environmental conditions. We compared the genetic structure and dispersal patterns of 2 populations of Alouatta caraya (Plathyrrhini, Atelidae) to understand how habitat reduction and fragmentation affect gene flow in this species. We sampled individuals from 7 groups living in continuous forest (CF, n = 46, 22 males and 24 females), and 11 groups that inhabit a fragmented forest (FF, n = 50, 24 males and 26 females). FST values based on 11 microsatellite loci showed a recent genetic differentiation among groups in the FF. In contrast, the CF showed no differentiation among groups. Further, FST values between sexes, as well as kinship relationships, also exhibited differences between habitats. In the CF, both males and females disperse, leading to nondifferentiated groups composed of adults that are not close relatives. Conversely, in the FF, some groups are differentiated, males disperse more than females, and groups are composed of closely related adult females. Our results suggest that habitat fragmentation modifies the dispersal patterns of black-and-gold howlers. These differences between habitats may reflect a reduced gene flow, providing genetic evidence that suggests that habitat fragmentation severely limits the howler’s ability to disperse. An increasing level of isolation due to uncontrolled deforestation may cause similar loss of genetic diversity on other arboreal primates, and nonprimates that depend on forest continuity to disperse, reducing their abilities to cope with environmental changes.     

Effects of landscape matrix on population connectivity of an arboreal mammal,Petaurus breviceps

Ongoing habitat loss and fragmentation is considered a threat to biodiversity as it can create small, isolated populations that are at increased risk of extinction. Tree‐dependent species are predicted to be highly sensitive to forest and woodland loss and fragmentation, but few studies have tested the influence of different types of landscape matrix on gene flow and population structure of arboreal species. Here, we examine the effects of landscape matrix on population structure of the sugar glider (Petaurus breviceps) in a fragmented landscape in southeastern South Australia. We collected 250 individuals across 12 native Eucalyptus forest remnants surrounded by cleared agricultural land or exotic Pinus radiata plantations and a large continuous eucalypt forest. Fifteen microsatellite loci were genotyped and analyzed to infer levels of population differentiation and dispersal. Genetic differentiation among most forest patches was evident. We found evidence for female philopatry and restricted dispersal distances for females relative to males, suggesting there is male‐biased dispersal. Among the environmental variables, spatial variables including geographic location, minimum distance to neighboring patch, and degree of isolation were the most important in explaining genetic variation. The permeability of a cleared agricultural matrix to dispersing gliders was significantly higher than that of a pine matrix, with the gliders dispersing shorter distances across the latter. Our results added to previous findings for other species of restricted dispersal and connectivity due to habitat fragmentation in the same region, providing valuable information for the development of strategies to improve the connectivity of populations in the future.     

No evidence for divergence in male harmfulness or female resistance in response to changes in the opportunity for dispersal

The outcome of sexual conflict can depend on the social environment, as males respond to changes in the inclusive fitness payoffs of harmfulness and harm females less when they compete with familiar relatives. Theoretical models also predict that if limited male dispersal predictably enhances local relatedness while maintaining global competition, kin selection can produce evolutionary divergences in male harmfulness among populations. Experimental tests of these predictions, however, are rare. We assessed rates of dispersal in female and male seed beetles Callosobruchus maculatus, a model species for studies of sexual conflict, in an experimental setting. Females dispersed significantly more often than males, but dispersing males travelled just as far as dispersing females. Next, we used experimental evolution to test whether limiting dispersal allowed the action of kin selection to affect divergence in male harmfulness and female resistance. Populations of C. maculatus were evolved for 20 and 25 generations under one of three dispersal regimens: completely free dispersal, limited dispersal and no dispersal. There was no divergence among treatments in female reproductive tract scarring, ejaculate size, mating behaviour, fitness of experimental females mated to stock males or fitness of stock females mated to experimental males. We suggest that this is likely due to insufficient strength of kin selection rather than a lack of genetic variation or time for selection. Limited dispersal alone is therefore not sufficient for kin selection to reduce male harmfulness in this species, consistent with general predictions that limited dispersal will only allow kin selection if local relatedness is independent of the intensity of competition among kin.     

Does sex matter? Gender‐specific responses to forest fragmentation in Neotropical bats

Understanding the consequences of habitat modification on wildlife communities is central to the development of conservation strategies. However, albeit male and female individuals of numerous species are known to exhibit differences in habitat use, sex‐specific responses to habitat modification remain little explored. Here, we used a landscape‐scale fragmentation experiment to assess, separately for males and females, the effects of fragmentation on the abundance of Carollia perspicillata and Rhinophylla pumilio, two widespread Neotropical frugivorous bats. We predicted that sex‐specific responses would arise from higher energetic requirements from pregnancy and lactation in females. Analyses were conducted independently for each season, and we further investigated the joint responses to local and landscape‐scale metrics of habitat quality, composition, and configuration. Although males and females responded similarly to a fragmentation gradient composed by continuous forest, fragment interiors, edges, and matrix habitats, we found marked differences between sexes in habitat use for at least one of the seasons. Whereas the sex ratio varied little in continuous forest and fragment interiors, females were found to be more abundant than males in edge and matrix habitats. This difference was more prominent in the dry season, the reproductive season of both species. For both species, abundance responses to local‐ and landscape‐scale predictors differed between sexes and again, differences were more pronounced in the dry season. The results suggest considerable sex‐mediated responses to forest disruption and degradation in tropical bats and complement our understanding of the impacts of fragmentation on tropical forest vertebrate communities.     

Habitat use and dispersal characteristic by <Emphasis Type="Italic">Stethophyma grossum</Emphasis>: the role of habitat isolation and stable habitat conditions towards low dispersal

Species in a highly fragmented environment, such as the intensively used agricultural landscapes of Europe, are expected to be in danger of extinction. We hypothesize according to Kisdi’s theory (Am Nat 159:579–596, 2002) that species in fragmented landscapes with isolated habitats in general tend to possess low dispersal. In order to verify this hypothesis we studied the movement patterns of Stethophyma grossum, a hygrophilous species of wetlands, by mark–release–recapture techniques in a landscape with scattered suitable habitats over 3 years. The study focused on the major population in this landscape (site #1) as dispersal behaviour was assumed to be greatest. Actually, marked individuals of S. grossum were never found in any further suitable habitats in close vicinity to site #1. Despite that the peatland meadow of study site #1 was all over covered with homogenous vegetation only 6% (1.8 ha) of the whole area (30 ha) were occupied by S. grossum. The mean recapture rate over 3 years amounted to 39% with no significant differences between males and females. Both covered little distances within their mean range size of 1.8 ha; the median distances were 36.91 m for males and 26.65 m for females. We confirm the hypothesis that sub-populations of species in longstanding naturally isolated habitats, which habitat conditions have been stable; evolved low dispersal with little movements which are routine movements to find mating partners or food.     

Whip it good! Geographic consistency in male songs and variability in female songs of the duetting eastern whipbird Psophodes olivaceus

Geographic variation in male bird songs has been studied extensively, but there have been few investigations of geographic variation in female songs or sex differences in patterns of geographic variation. We compared patterns of variation in male and female songs of eastern whipbirds Psophodes olivaceus by analyzing recordings from 16 populations across the species’ geographic range in eastern Australia. We found remarkably different patterns of geographic variation between the sexes. Female eastern whipbird songs are easily categorized into discrete song types. Song types are shared between nearby females, but female songs show pronounced differences at a continental scale. In contrast, male eastern whipbird songs show high consistency throughout the species’ geographic range. All recorded males share the ability to transpose the frequency of the introductory whistle and most recorded males share the ability to vary the direction of the slope of the terminal whip crack. For eight of nine measured variables, female songs show significantly higher levels of variation than male songs. We discuss whether sex differences in dispersal, song learning strategies, and song function may explain these sex differences in patterns of song variation. We suggest that eastern whipbirds have experienced a decoupling of male and female song learning strategies and that the songs of each sex have responded to different selective pressures in the context of countersinging interactions. We speculate that consistency in male songs throughout the geographic range of eastern whipbirds may arise through female preference for males that perform large bandwidth whip cracks.     

Variation in fine‐scale genetic structure and local dispersal patterns between peripheral populations of a South American passerine bird

The distribution of suitable habitat influences natal and breeding dispersal at small spatial scales, resulting in strong microgeographic genetic structure. Although environmental variation can promote interpopulation differences in dispersal behavior and local spatial patterns, the effects of distinct ecological conditions on within‐species variation in dispersal strategies and in fine‐scale genetic structure remain poorly understood. We studied local dispersal and fine‐scale genetic structure in the thorn‐tailed rayadito (Aphrastura spinicauda), a South American bird that breeds along a wide latitudinal gradient. We combine capture‐mark‐recapture data from eight breeding seasons and molecular genetics to compare two peripheral populations with contrasting environments in Chile: Navarino Island, a continuous and low density habitat, and Fray Jorge National Park, a fragmented, densely populated and more stressful environment. Natal dispersal showed no sex bias in Navarino but was female‐biased in the more dense population in Fray Jorge. In the latter, male movements were restricted, and some birds seemed to skip breeding in their first year, suggesting habitat saturation. Breeding dispersal was limited in both populations, with males being more philopatric than females. Spatial genetic autocorrelation analyzes using 13 polymorphic microsatellite loci confirmed the observed dispersal patterns: a fine‐scale genetic structure was only detectable for males in Fray Jorge for distances up to 450 m. Furthermore, two‐dimensional autocorrelation analyzes and estimates of genetic relatedness indicated that related males tended to be spatially clustered in this population. Our study shows evidence for context‐dependent variation in natal dispersal and corresponding local genetic structure in peripheral populations of this bird. It seems likely that the costs of dispersal are higher in the fragmented and higher density environment in Fray Jorge, particularly for males. The observed differences in microgeographic genetic structure for rayaditos might reflect the genetic consequences of population‐specific responses to contrasting environmental pressures near the range limits of its distribution.     

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.     

Mark–recapture on large spatial scale reveals long distance dispersal in the Marsh Fritillary,Euphydryas aurinia

1. Long distance dispersal (LDD), or movements far beyond the occupied habitat borders, maintains the integrity of metapopulations in fragmented landscapes. Recent studies on butterflies increasingly reveal that LDD exists even in species that were long regarded as sedentary. Mark–recapture (MR) studies covering larger study areas typically reveal movements among distant colonies. 2. We studied dispersal of the EU‐protected, regionally endangered Euphydryas aurinia Rottemburg butterfly in the Czech Republic, using two complementary MR approaches. The single system study was carried out for eight seasons within 30 habitat patches covering 28 ha. The multiple populations study was carried out for a single season, but covering almost all Czech colonies of the species (82 colonies, 110 distinct patches, total area 324 ha within ca 1500 km²). 3. Single system mean lifetime movements were consistently higher for males, but slopes of dispersal kernel power functions were shallower for females, implying that higher proportions of females crossed distances of several kilometres. 4. The multiple populations study allowed detection of 51 lifetime movements exceeding 5 km (41 males, 10 females) and 14 movements exceeding 10 km (13 males, 1 female). Both mean lifetime movements and slopes of the dispersal kernels varied among systems, with no consistent pattern between sexes. All Czech Republic populations are within 0.1% movement probability of both sexes, whereas 1% movement probability delimits three separate management units. 5. Dispersal predictions from local data underestimate total mobility, warning against the use of local MR data for extrapolating long‐distance movements. Local dispersal data, however, remain useful for analysing finer details of insect mobility.     

Habitat loss and fragmentation reduce effective gene flow by disrupting seed dispersal in a neotropical palm

Habitat loss and fragmentation often reduce gene flow and genetic diversity in plants by disrupting the movement of pollen and seed. However, direct comparisons of the contributions of pollen vs. seed dispersal to genetic variation in fragmented landscapes are lacking. To address this knowledge gap, we partitioned the genetic diversity contributed by male gametes from pollen sources and female gametes from seed sources within established seedlings of the palm Oenocarpus bataua in forest fragments and continuous forest in northwest Ecuador. This approach allowed us to quantify the separate contributions of each of these two dispersal processes to genetic variation. Compared to continuous forest, fragments had stronger spatial genetic structure, especially among female gametes, and reduced effective population sizes. We found that within and among fragments, allelic diversity was lower and genetic structure higher for female gametes than for male gametes. Moreover, female gametic allelic diversity in fragments decreased with decreasing surrounding forest cover, while male gametic allelic diversity did not. These results indicate that limited seed dispersal within and among fragments restricts genetic diversity and strengthens genetic structure in this system. Although pollen movement may also be impacted by habitat loss and fragmentation, it nonetheless serves to promote gene flow and diversity within and among fragments. Pollen and seed dispersal play distinctive roles in determining patterns of genetic variation in fragmented landscapes, and maintaining the integrity of both dispersal processes will be critical to managing and conserving genetic variation in the face of continuing habitat loss and fragmentation in tropical landscapes.     

Successful despite poor flight performance: range expansion is associated with enhanced exploratory behaviour and fast development

Anthropogenic interference forces species to respond to changing environmental conditions. One possible response is dispersal and concomitant range shifts, allowing individuals to escape unfavourable conditions or to track the shifting climate niche. Range expansions depend on both dispersal capacity and the ability to establish populations beyond the former range. We here compare well‐established core populations with recently established edge populations in the currently northward expanding butterfly Lycaena tityrus. Edge populations were characterized by shorter development times and smaller size, a higher sensitivity to high temperature and an enhanced exploratory behaviour. The differences between core and edge populations found suggest adaptation to local climates and an enhanced dispersal ability in edge populations. In particular, enhanced exploratory behaviour may be advantageous in all steps of the dispersal process and may have facilitated the current range expansion. This study describes differences associated with a current range expansion, knowledge which might be useful for a better understanding of species responses to environmental change. We further report on variation between males and females in morphology and flight behaviour, with males showing a longer flight endurance and more pronounced exploratory behaviour than females.     

Dispersal of four fritillary butterflies within identical landscape

Both species-specific traits and landscape configuration, such as area and connectivity of habitat patches plus the character of uninhabitable matrix, affect animal movements in fragmented landscapes. Difficulties with disentangling species-specific and landscape effects have obscured comparisons among species, hindering the understanding of dispersal in metapopulations. To circumvent this complication, we performed a mark–recapture study of four related nymphalid butterflies within identical landscape and in single season. The studied species were three Melitaeinae checkerspots (Euphydryas aurinia, Melitaea athalia, Melitaea diamina) and one Argynnini fritillary (Brenthis ino). Applying the Virtual Migration model revealed that (1) except for mortality within habitat, model parameters differed from those found for the studied species elsewhere; (2) the three Melitaeinae species were more akin in movement parameters than the Argynnini representative (i.e., B. ino); (3) within Melitaeinae, differences between sexes were more prominent than differences among species; (4) Melitaeinae males left natal patches more readily than females, while the opposite applied to B. ino; (5) males of M. diamina and both sexes of B. ino exhibited highest values of dispersal mortality; (6) except for females of M. diamina and both sexes of B. ino, immigration and emigration scaled with area in females but not in males. Finding (1) demonstrates that geometry of habitat network affects mobility considerably and that transferring dispersal parameters across systems is unwarranted. Still, (2–6) demonstrate that within identical networks, related species follow similar dispersal patterns, suggesting that conservation scenarios suitable for a well-studied model species would suite related species as well.     

Dispersal mood revealed by shifts from routine to direct flights in the meadow brown butterfly Maniola jurtina

A comprehensive mechanistic approach to dispersal requires the translation of the whole mobility register of the target organism into movement rules that could subsequently be used to model its displacements. According to the optimality paradigm, this procedure implies a cost–benefit analysis of mobility patterns taking into account not only movements, but also their external context and the internal state of the moving individuals. Using this framework, we detected a ‘dispersal mood’ in some individuals of the meadow brown butterfly Maniola jurtina. These adopted a direct flight strategy, which was topologically different from the previously documented foray search strategy. Those individuals that used the direct flight strategy moved straighter as soon as they left the habitat and avoided heading back to their patch of origin, which is the best inter‐patch search strategy when dispersal risks and costs are high. The direct flight strategy was conditional to sex: females used it twice as much as males. We suggest that this sex bias was due to female investment in offspring, which is maximized by male avoidance and spatial bet hedging. Inter‐patch dispersal of gravid females is crucial for the persistence of M. jurtina populations in spatially and temporally unpredictable environments.     

Niche separation in space and time between two sympatric sister species—a case of ecological pleiotropy

We investigate the niche separation in space and time between the Palearctic sister species Leptidea sinapis and L. reali (Lepidoptera, Pieridae) in central Sweden. Using field sampling, we show that L. reali is a habitat specialist confined to meadows, whereas L. sinapis is a habitat generalist also inhabiting forests. This difference in habitat utilization was corroborated by experimental release of laboratory-reared L. sinapis and L. reali in two adjacent forest and meadow habitats during their natural flight period; virtually all recaptured L. reali that were released in the forest were later caught in the meadow, whereas L. sinapis shifted equally often from meadow to forest as in the opposite direction. In the field, both species fly in May–June, but L. reali appears on average a week earlier in spring and has a substantial second generation in July, whereas L. sinapis is practically univoltine. When overwintered pupae were incubated under identical conditions in the laboratory, females did, however, not differ in phenology, and L. sinapis males actually emerged earlier than L. reali males. When larvae were reared at 23°C on the host plant Lotus corniculatus at a range of daylengths, both species produced a substantial proportion of directly developing individuals at an 18.5 h daylength or longer. When reared at 23°C and a 22 h daylength, L. reali showed an overall higher propensity to develop directly than L. sinapis on plant species originating from both the meadow and the forest habitat. Both Leptidea species showed a lower propensity to enter direct development on forest associated plants than on meadow associated plants. Hence, we suggest that the difference in phenology and voltinism between L. sinapis and L. reali is largely the result of environmentally implemented ecological pleiotropic effects caused by the between-species difference in habitat preference.     

Mating Behavior Drives Seed Dispersal by the Long‐wattled Umbrellabird Cephalopterus penduliger

Frugivores exhibit considerable variation in the seed dispersal services they provide. Understanding what drives these differences is a key goal for ecologists because of the central role seed dispersal plays in shaping ecological and genetic diversity in plant populations. The lek‐mating system of the Long‐wattled Umbrellabird (Cephalopterus penduliger) provides a powerful lens to examine how mating behavior may impact seed dispersal outcomes. As in all lek‐breeding species, male Umbrellabirds congregate in traditional sites (leks) to display, whereas females are solitary and visit leks only rarely. This study demonstrates how differences in mating behavior between the sexes drive distinctive seed movement and deposition patterns by male vs. female Umbrellabirds. Using radio tracking and gut retention trials, we documented divergent movement patterns between the sexes that are directly attributable to mating behavior differences. These movement differences led males to disperse seeds long distances from source trees and to deposit the majority of seeds they ingested within the lek; females dispersed seeds shorter distances and more evenly across the landscape. We empirically confirmed that the density of dispersed seeds was higher in leks than in control areas outside the lek, yet found no evidence that this higher density of seeds in leks reduced probability of seedling establishment. This research not only provides a mechanistic explanation for long dispersal distances and high levels of genetic diversity previously reported for seeds in Umbrellabird leks, but also highlights the importance of explicitly considering behavior in studies of animal‐mediated seed dispersal.     

Effects of meadow suitability on female behaviour in the alpine butterfly Parnassius smintheus

Abstract  1. Many butterfly populations persist in networks of naturally fragmented habitat patches. Movement and reproductive decisions made by adult females are critical to the persistence of these populations because colonisation of extinct habitat patches in the network requires emigration of fecund adult females from their natal meadow and their subsequent establishment in the extinct patch.
2. Movement and oviposition behaviours of mated Parnassius smintheus females released in suitable meadows (a good- and a poor-quality meadow) and an unsuitable meadow were compared, to determine whether adult females consider meadow suitability for their offspring despite frequent oviposition events off the larval host plant.
3. Bootstrap and correlated random walk analyses of female step lengths and turn angles demonstrated that females flew more randomly in the unsuitable meadow than in the suitable meadows. Although females tended to turn the sharpest angle between landing sites in the good-quality meadow, and fly the smallest distance between landing sites and displace the smallest distance from the release site in the suitable meadows, no significant differences were detected in turn angle, step length, and dispersal rates between suitable and unsuitable meadows.
4. Results from female flight observations and a caged oviposition study suggest that females lay significantly more eggs in suitable habitat than in unsuitable habitat despite not ovipositing on the host plant, and support the above findings.
5. Movement and oviposition behaviours of adult female P. smintheus promote their retention within meadows that can support their offspring.     

Individual and Temporal Variation in Habitat Association of an Alien Carnivore at Its Invasion Front

Gathering information on how invasive species utilize the habitat is important, in order to better aim actions to reduce their negative impact. We studied habitat use and selection of 55 GPS-marked raccoon dogs (30 males, 25 females) at their invasion front in Northern Sweden, with particular focus on differences between males and females, between movement states, and between seasons and times of the day. Daily movement pattern was used to classify GPS-locations into dispersing and settled. We focused on both anthropogenic and natural landscape characteristics. Since we did not have any a priori knowledge about the spatial scale of raccoon dog habitat selection, we first assessed how landscape characteristics of random points changed with distance from the GPS-location they were paired to. Because changes in habitat use became less pronounced at approximately 5 km for all variables, we focused on habitat use at two spatial scales: fine (500 m) and coarse (5 km). Habitat selection was strongest at the coarse scale, and reflected the results found for habitat use. Raccoon dogs selected agricultural areas and wetlands, lower altitudes, and shallow slopes, and avoided forests, open natural areas, and areas close to water and roads. There were no differences in habitat selection between males and females, or between movement states. This lack of sexual segregation increases the probability of encountering potential mates during dispersal, and therefore the likelihood for reproduction in new areas. The seasonal and diurnal pattern of habitat use may provide guidance for where and when to aim management efforts.     

Divergence in Female Duetting Signals in the Enchenopa binotata Species Complex of Treehoppers (Hemiptera: Membracidae)

Sexual communication often involves signal exchanges between the sexes, or duetting, in which mate choice is expressed through response signals. With both sexes acting as signalers and receivers, variation in the signals of males and females may be important for mate choice, reproductive isolation, and divergence. In the Enchenopa binotata species complex – a case study of sympatric speciation in which vibrational duetting may have an important role – male signals are species‐specific, females choose among males on the basis of signal traits that reflect species and individual differences, and female preferences have exerted divergent selection on male signals. Here, we describe variation in female signals in the E. binotata species complex. We report substantial species differences in the spectral and temporal features of female signals, and in their timing relative to male signals. These differences were similar in range to differences in male signals in the E. binotata complex. We consider processes that might contribute to divergence in female signals, and suggest that signal evolution in the E. binotata complex may be influenced by mate choice in both sexes.     

The effects of artificial and natural barriers on the movement of small mammals in Banff National Park, Canada

Barriers that dissect continuous habitat, such as roads, initiate the process of habitat fragmentation and can filter or eliminate animal movement through otherwise pristine areas. A severe road barrier appears to exist in Banff National Park where the Trans‐Canada Highway (TCH) is known to impede the movement of some animals, but the effects of this barrier relative to natural barriers and continuous habitat have not been assessed. To compare animal movement in response to these three contexts, we translocated three species of murid rodents (meadow voles, Microtus pennsylvanicus, deer mice, Peromyscus maniculatus, and red‐backed voles, Clethrionomys gapperi) across the TCH (an artificial barrier), a forested strip in the median of the TCH (a natural barrier), in the adjacent highway verge (continuous grassy) and in nearby forest (continuous forested). We coated murids with fluorescent dye, released them at standardized distances, and followed the resulting trails to obtain detailed information on movement paths. Overall, individuals were 20% less successful crossing the TCH than natural (forested) barriers and 10% less successful crossing natural barriers than continuous habitat, but exhibited marked variation among species. Path tortuosity was negatively related to return success; more convoluted paths were associated with lower success across all treatments. Deer mice returned to their home‐ranges more often than the other two species, perhaps because of their generalist habitat preferences, nocturnal activities, or larger home ranges. In all treatments, the fractal dimensions of movement paths were very qualitatively similar to those observed for return success and tortuosity, indicating that the responses of these species were not influenced by spatial scale. Together, our results suggest that (a) murid species differ in the movement limitation imposed by both natural and artificial barriers, perhaps as a consequence of differences in habitat preferences and activity patterns, and (b) murid responses to a road barrier are only moderately different from responses to the natural barrier of a forest edge in the context of translocation experiments.