Niche partitioning of two Palearctic passerine migrants with Afrotropical residents in their West African winter quarters

In bird migration systems, the question of coexistence and competitionof migrants with residents in the nonbreeding season and theirrole in shaping the evolution of present avian communities ismuch debated. However, conclusions are often drawn in a speculativeway. In the Palearctic-African bird migration system, many studieshave addressed the question of coexistence of Palearctic breedingbirds with Afrotropical species in the former's wintering grounds.These studies have led to some generalizations concerning habitatselection and foraging ecology about the traits that might enablemigrants to coexist with residents. Migrants were thereforeassumed to forage in more open habitats, in more peripheralparts of the vegetation and with a higher foraging speed thanresidents. Furthermore, they were also assumed to be more flexiblein foraging behavior by using a wider range of foraging tactics,but some studies revealed contradicting results. We studiedthe ecology of Palearctic migrants in Comoé NationalPark, Ivory Coast, West Africa, during three successive wintersto explore the factors of niche partitioning between migrantsand residents and to test the hypothesis of whether there arecommon behavioral traits in migrants. Therefore, we comparedthe ecology of two Palearctic breeding species: pied flycatcher,Ficedula hypoleuca, and willow warbler, Phylloscopus trochilus,with resident members of the respective guilds. With respectto use of microhabitat, foraging speed, and intake rates, wecould not confirm the above-mentioned generalizations, showingthat care has to be taken into account when drawing conclusionsfrom few studies for a whole migration system on a huge continent.However, both migrating species were more flexible in foragingbehavior than were their Afrotropical counterparts. As thereare hints that this is also the case when migrants are comparedwith residents on their breeding grounds, we suggest that thisflexibility enables migrants to partition resources with residentsand, therefore, coexist with Afrotropical species. We discuss,however, whether this flexibility is an adaptation to migratorybehavior or a prerequisite for the evolution of migration. Therole that competition plays in present communities cannot besolved with a few observational studies because of the followingproblems. First, it is difficult to detect competition in thefield. Second, there are constraints of performing field experimentsthat have not previously been performed in Africa, and third,there are several possible hypothetical scenarios about therole of competition in shaping present communities, includingfactors that might have been important in the past and are thereforeimpossible to detect at present.     

SYSTEMATIC NOTES (Part I)

Declines in populations of Palearctic migrants wintering in the Sahel of Africa have been linked to the impacts of climate change and habitat degradation in the region. Despite this, there is an almost complete lack of data on the density and distribution of Palearctic migrants wintering in the Sahel and whether they have the same habitat requirements as similar, resident Afrotropical species. We measured the density of five species of Palearctic warblers (Sylviidae) and 10 species of Afrotropical gleaning passerines (Sylviidae, Nectariniidae, Malaconotidae and Ploceidae) at 16 sites in the Sahel of northern Nigeria between October and April during two winters. Two species of Afrotropical gleaner (Hippolais pallida and Ploceus luteolus) showed seasonal variation in abundance, but this variation was unlikely to have decreased Afrotropical densities sufficiently to change the degree of competition experienced by Palearctic migrants. This observation, combined with a positive correlation between abundances of Afrotropical and Palearctic species, suggests that these two groups occur together and have similar spatial and temporal habitat requirements, and therefore possibly similar responses to habitat degradation. Sylvia communis appears to be the principal species utilising the region during spring migration, presumably for fattening prior to the trans-Saharan crossing, and is thus perhaps the most vulnerable species to habitat loss in the region.     

Habitat selection and coexistence of migrants and Afrotropical residents

In this paper I discuss factors that influence the habitat choice of small migrant birds on their Afrotropical wintering grounds and the ecological isolation of migrants and residents. The main characteristic of migrants is the use of resources which are sporadic in space and time. The majority of migrants occur in seasonal savannas and open woodland, mostly using temporarily and locally abundant food sources generally unused by residents. Migrants are more eurytopic and exploit more open parts of the habitats than ecologically similar tropical species. In some guilds, the foraging speed and rate of migrants is higher and they use their wings more often in comparison to residents. There is weak evidence that niche shifts of residents are induced by the arrival of migrants. Overt interspecific interactions seem to be infrequent, except in a guild of ground-feeding birds which rely on a rather predictable but poor food supply. In this guild, residents, but not intra-African migrants, dominated Palaearctic migrants in aggressive encounters irrespective of body-size. In contrast to this, dominance was size-dependent among wintering Palaearctic migrants.     

Independent trophic behavior and breeding success of a resident flycatcher and a coexisting migratory congener

Historical biases towards the study of Palearctic and Nearctic bird migration systems has resulted in a dearth of information on the ecology of intratropical migrants, which likely also play important ecological roles within their communities. For instance, there is little information on the foraging ecology and breeding biology of co‐occurring migrants and residents within the intratropical migratory system. Thus, we used two congeneric flycatchers – the resident Plain‐crested Elaenia Elaenia cristata (Aves: Tyrannidae), and the migrant Lesser Elaenia E. chiriquensis (Tyrannidae) – to check whether either of them showed foraging niche plasticity mediating their coexistence and the effects of such syntopy on their breeding behavior and fitness. To do so, we used an ensemble of methods that included the evaluation of food resource phenology, foraging behavior observations, stable isotope ratio analysis in birds’ tissues, and nest monitoring. Our results confirmed that residents have foraging behavior plasticity but reveals its trophic behavior and breeding success is independent of the coexistence with migrants. Since such results depart from the predictions based on the current (and often Northern‐biased) theories of bird migration, we discuss alternative hypotheses explaining our findings, including the influence of physiology on diet and the behavioral responses of nest predators. Once there is a clear knowledge gap about the interspecific interactions between residents and intratropical migrants, our study represents a simple, yet important, step towards understanding the mechanisms underlying this system.     

Fine‐scale habitat use during the non‐breeding season suggests that winter habitat does not limit breeding populations of a declining long‐distance Palearctic migrant

For migrant birds, which habitats are suitable during the non‐breeding season influences habitat availability, population resilience to habitat loss, and ultimately survival. Consequently, habitat preferences during winter and whether habitat segregation according to age and sex occurs directly influences migration ecology, survival and breeding success. We tested the fine‐scale habitat preferences of a declining Palearctic migrant, the whinchat Saxicola rubetra, on its wintering grounds in west Africa. We explored the influence of habitat at the territory‐scale and whether dominance‐based habitat occupancy occurs by describing the variation in habitat characteristics across wintering territories, the degree of habitat change within territories held throughout winter, and whether habitat characteristics influenced territory size and space‐use within territories or differed with age and sex. Habitat characteristics varied substantially across territories and birds maintained the same territories even though habitat changed significantly throughout winter. We found no evidence of dominance‐based habitat occupancy; instead, territories were smaller if they contained more perching shrubs or maize crops, and areas with more perching shrubs were used more often within territories, likely because perches are important for foraging and territory defence. Our findings suggest that whinchats have non‐specialised habitat requirements within their wintering habitat of open savannah and farmland, and respond to habitat variation by adjusting territory size and space‐use within their territories instead of competing with conspecifics. Whinchats show a tolerance for human‐modified habitats and results support previous findings that some crop types may provide high‐quality wintering habitat by increasing perch density and foraging opportunities. By having non‐specialised requirements within broad winter habitat types, migrants are likely to be flexible to changing wintering conditions in Africa, both within and across winters, so possibly engendering some resilience to the rapid anthropogenic habitat degradation occurring throughout their wintering range.     

Does life-history variability in salmonids affect habitat use by juveniles? A comparison among streams open and closed to anadromy

1. Migratory and resident forms of salmonids coexist in many river systems. Although such coexistence is widespread, little is known about its ecological basis and no studies have compared the habitat use of premigratory juveniles and residents. 2. We employed a comparative approach to explore the differential habitat use of juvenile anadromous and resident brook trout. This required the investigation of habitat use in streams closed to anadromy, containing only resident brook trout Salvelinus fontinalis ('resident-only' streams) and streams open to anadromy, containing coexisting Atlantic salmon Salmo salar and anadromous and resident brook trout ('migrant-resident' streams). 3. We demonstrate that fast habitats (riffles) are occupied more frequently in streams with migratory brook trout relative to riffle habitats of streams with only resident brook trout. In contrast, occupation of slow current velocities (pools) was observed in both migrant-resident and resident-only streams as both stream types contain resident brook trout. The net effect is a wider distribution of occupied habitats (pool and riffles) in migrant-resident streams relative to resident-only streams, resulting in few, if any, unused habitats. 4. These results are consistent with previously reported bioenergetic, morphological and stable isotope differences observed between anadromous and resident brook trout. 5. Our findings suggest that a link exists between juvenile habitat use, metabolic costs and life-history strategies.     

Habitat use and densities of co‐existing migrant Willow Warblers Phylloscopus trochilus and resident eremomelas Eremomela spp. in Zimbabwe

Capsule Migrant Willow Warblers occupy more woodland types and occur at higher densities than ecologically‐similar resident Afrotropical warblers.

Aims To compare population densities of Willow Warblers and eremomelas in adjacent acacia, mopane and miombo woodlands, and assess the abundance of potential invertebrate prey in each habitat type, in order to investigate whether Palearctic migrants use more open habitats and are more flexible in habitat use than their Afrotropical counterparts in the same feeding guild.

Methods Using distance sampling we carried out four replicated sets of point counts in acacia woodland and three sets of counts in miombo and mopane between December 1999 and February 2000. We noted the tree species in which we saw warblers foraging and took beating‐tray samples of potential arthropod prey present on tree foliage in each of the three habitats.

Results Willow Warbler density in acacia woodland increased from 1.80 ± 0.54 (se) birds/ha in early December to 7.15 ± 1.41 birds/ha in late January after influxes of later arrivals. Densities of Willow Warblers in miombo and mopane were much lower (1.14 ± 0.28 and 0.38 ± 0.23 birds/ha, respectively) and did not show significant changes. Burnt‐necked Eremomelas averaged 0.74 ± 0.34 birds/ha in acacia woodland, and in miombo densities of Green‐capped and Yellow‐bellied Eremomelas were 0.23 ± 0.17 and 0.34 ± 0.26 birds/ha, respectively. Densities in mopane were too low to estimate reliably. Willow Warblers and Green‐capped Eremomelas showed some apparent preferences in tree species used for foraging but differences in tree use were not obviously related to the abundance of arthropod taxa present as potential prey.

Conclusion Willow Warblers occupied more habitats at greater density than similar Afrotropical warblers. They appear to favour acacia, but their settlement patterns and the reasons for disparities between densities of immigrants and residents are unclear.     

A preliminary assessment of the habitat selection of two Palaearctic migrant passerine species in West Africa

We investigated the habitat selected by two Palaearctic migrants (Pied Flycatcher, Ficedula hypoleuca, Willow Warbler, Phylloscopus trochilus) in a patchy landscape in Ivory Coast and compared it with the habitat selection of Afrotropical species in the same foraging guilds. Transect counts were used to test the hypothesis that migrants use more open and more seasonal habitats and have a broader use of habitats compared with resident species. Habitats compared were, in order of decreasing tree density, gallery forest, an isolated forest and bush/tree savanna. The isolated forest had the most pronounced seasonal changes (deciduous trees) and was the one with the most diverse vegetation structure. The habitat where both migrants were most frequent was the isolated forest, and thus occurred in the habitat with the most pronounced seasonal change. Diversity of habitats selected was highest in migrants but in the Pied Flycatcher this was possibly an artefact due to subdominant individuals being excluded from the preferred habitat by territorial birds. Potential competition for habitat with Afrotropical species was found to be low.     

Altitudinal migration by birds: a review of the literature and a comprehensive list of species

Altitudinal migration is the seasonal altitudinal movement of birds from breeding areas to non‐breeding or wintering areas at different elevations. Although this type of migration is widely reported, questions remain concerning the number of species that perform altitudinal migration, possible variation among different taxa and geographic locations in the extent of altitudinal migration, and the foraging guilds of altitudinal migrants. We conducted an extensive bibliographic survey and compiled a list of altitudinal migrant birds worldwide. We characterized species in terms of their foraging guilds because the spatial distribution of food resources along altitudinal gradients is often evoked as a driver of bird altitudinal migration. We identified 1238 species of altitudinal migrants, ~10% of the ~10,000 extant species of birds. We found a strong geographic bias in publications focusing on avian altitudinal migration toward the United States and Costa Rica, and a paucity of studies in megadiverse regions such as the Afrotropical and Indomalayan realms, and areas in the Neotropics other than Costa Rica. We also found that most species of altitudinal migrants were invertivores rather than frugivores or nectarivores. This general pattern held true for all zoogeographic realms except the Neotropics, where nectarivores and frugivores predominated among altitudinal migrants. The prevalence of invertivore birds among altitudinal migrants is not unexpected because this is the most common foraging guild among birds worldwide. Overall, we found no prevalence of any specific foraging guild among altitudinal migrants across zoogeographic regions. The results of studies to date suggest that altitudinal migration by birds may be driven by a number of factors, including access to increased food resources for breeding or molting, weather conditions, and mating and nesting opportunities. However, to better understand the mechanisms underlying altitudinal migration, broadening the geographic scope of studies is paramount, with additional study of altitudinal migration especially needed in the megadiverse tropical regions of sub‐Saharan Africa, Southeast Asia, and South America.     

Temperate migrants and resident bird species in Afro‐tropical savannahs show similar levels of ecological generalism

The specificity of an animal's habitat requirements will determine its ability to deal with anthropogenic climate and habitat change. Migratory birds are thought to be particularly vulnerable to such change, but theory predicts that they should be largely generalists. This prediction was tested with the aim of assessing whether migratory Palaearctic‐breeding birds wintering in the savannah biome of Africa are more or less generalist in their habitat use compared with taxonomically and ecologically similar Afro‐tropical resident species. The degree of specialization of these species groups to certain habitat characteristics was assessed and compared by calculating the relative occurrence of the species along habitat gradients, where wide occurrence indicates generalism and narrow occurrence indicates specialism. Palaearctic migrants as a group could not clearly be distinguished as generalists relative to Afro‐tropical residents with respect to habitat attributes. The only indication of greater flexibility in Palaearctic migrants was a significant tendency to use habitats over a wider latitudinal range. The results suggest that migrants are generalists, but not necessarily more generalist than taxonomically similar resident species that also occur over a wide range of habitat types within the savannah biome. The availability of specific habitat requirements on the wintering grounds in Africa is therefore unlikely to be a primary limiting factor for many Afro‐Palaearctic migratory bird species.     

Impacts of passage migrant songbirds on behaviour and habitat use of resident Sardinian Warblers Sylvia melanocephala in Gibraltar

Explanations for the coexistence of migratory and sedentary (resident) birds are much debated: some authors have suggested that migrants make use of surplus resources not exploited by residents, whereas others have recorded ecological shifts in residents during periods when migrants are present. These shifts have been ascribed to the presence of migrants but could equally reflect seasonal variation in the distribution of resources required by sedentary species. Here we assess how the abundance of migrants affects the territorial behaviour and habitat use of resident Sardinian Warblers Sylvia melanocephala at a stop‐over site on the Rock of Gibraltar. There was a significant positive relationship between the abundance of migrants and the distances moved, time spent calling and the species and functional diversity of plants used by Sardinian Warblers on a daily basis, suggesting that migrants have a significant impact on residents.     

Timing and location of reproduction in African waterfowl: an overview of >100 years of nest records

The timing and location of reproduction are fundamental elements of reproductive success for all organisms. Understanding why animals choose to reproduce at particular times and in particular places is also important for our understanding of other aspects of organismal ecology, such as their habitat requirements, movement strategies, and biogeography. Although breeding patterns in waterfowl are relatively well documented, most studies are from northern temperate regions and the influences of location and time of year on breeding in Afrotropical ducks (Anatidae) are poorly understood. We outline six alternative (but not mutually exclusive) hypotheses that might explain where and when Afrotropical ducks choose to breed. To explore these hypotheses, we assembled and analyzed a new database of c. 22,000 breeding records for 16 Afrotropical ducks and one introduced Palearctic species (the Mallard Anas platyrhynchos). The full database is available on line as an appendix to this article. We identified five distinct breeding strategies as well as two outliers. Peak breeding for 9 of 16 indigenous duck species occurs during the dry season. We found no evidence for spatial synchrony or spatial autocorrelation in breeding, suggesting a high level of flexibility in waterfowl responses to prevailing conditions in any given year. More intensive analyses of alternative hypotheses are needed, but our initial analysis suggests that the timing of breeding for the majority of Afrotropical ducks is driven by a combination of resource availability and predation risk.     

An assemblage of wintering waders in coastal Kenya: activity budget and habitat use

To contribute to the poorly known ecology of Palearctic immigrant waders in East Africa, we studied the foraging system (roosting and foraging behaviours, 24‐h activity rhythm, time‐activity budget and habitat use) of eleven wader species overwintering on the Kenyan coast. The waders congregated in communal roosts and completely overlapped in their foraging rhythms, which in this tidal environment were completely determined by tides, and which were independent of the dark/light cycle. All species, both tactile and visual foragers, fed both by day and by night. This study, along with similar findings for other areas of the Palaeotropics, shows that wader activity throughout the 24‐h day is essentially uniform, and is only slightly influenced by the dark/light cycle, but is modified by tidal levels. The total time devoted to feeding by each wader species was related to their body mass. Each species used a specific micro‐habitat for foraging.     

Foraging in space and time structure an African small mammal community

We used live-trapping and foraging to test for the effect of habitat selection and diet on structuring a community of six small mammals and one bird within the Soutpansberg, South Africa. We established grids that straddled adjacent habitats: woodland, rocky hillside, and grassland. Trapping and foraging were used to estimate abundance, habitat use, and species-specific foraging costs. The species with the highest abundance and foraging activity in a habitat, activity time, or food was considered the most efficient and presumed to have a competitive advantage. All species exhibited distinct patterns of spatial and temporal habitat preference which provided the main mechanism of coexistence, followed by diet selection. The study species were organized into three assemblages (α diversity): grassland, Rhabdomys pumilio, Dendromus melanotis, and Mus minutoides.; woodland, Aethomys ineptus and Micaelamys namaquensis; and rock-dwelling, M. namaquensis and Elephantulus myurus. Francolinus natalensis foraged in open rocky areas and under wooded islands within the grassland. Species organization across the habitats suggested that feeding opportunities are available within all habitats; however, distinct habitat preferences resulted from differing foraging aptitudes and efficiencies of the competing species. At Lajuma, species distribution and coexistence are promoted through distinct habitat preferences that were shaped by competition and species-specific foraging costs. The combination of trapping and foraging provided a mechanistic approach that integrates behavior into community ecology by ‘asking’ the animal to reveal its perspective of the environment. Using spatial and temporal foraging decisions—as behavioral indicators—enables us to guide our understanding for across-taxa species coexistence.     

Foraging Ecology Predicts Learning Performance in Insectivorous Bats

Bats are unusual among mammals in showing great ecological diversity even among closely related species and are thus well suited for studies of adaptation to the ecological background. Here we investigate whether behavioral flexibility and simple- and complex-rule learning performance can be predicted by foraging ecology. We predict faster learning and higher flexibility in animals hunting in more complex, variable environments than in animals hunting in more simple, stable environments. To test this hypothesis, we studied three closely related insectivorous European bat species of the genus Myotis that belong to three different functional groups based on foraging habitats: M. capaccinii, an open water forager, M. myotis, a passive listening gleaner, and M. emarginatus, a clutter specialist. We predicted that M. capaccinii would show the least flexibility and slowest learning reflecting its relatively unstructured foraging habitat and the stereotypy of its natural foraging behavior, while the other two species would show greater flexibility and more rapid learning reflecting the complexity of their natural foraging tasks. We used a purposefully unnatural and thus species-fair crawling maze to test simple- and complex-rule learning, flexibility and re-learning performance. We found that M. capaccinii learned a simple rule as fast as the other species, but was slower in complex rule learning and was less flexible in response to changes in reward location. We found no differences in re-learning ability among species. Our results corroborate the hypothesis that animals’ cognitive skills reflect the demands of their ecological niche.     

Patterns of bird migration phenology in South Africa suggest northern hemisphere climate as the most consistent driver of change

Current knowledge of phenological shifts in Palearctic bird migration is largely based on data collected on migrants at their breeding grounds; little is known about the phenology of these birds at their nonbreeding grounds, and even less about that of intra‐African migrants. Because climate change patterns are not uniform across the globe, we can expect regional disparities in bird phenological responses. It is also likely that they vary across species, as species show differences in the strength of affinities they have with particular habitats and environments. Here, we examine the arrival and departure of nine Palearctic and seven intra‐African migratory species in the central Highveld of South Africa, where the former spend their nonbreeding season and the latter their breeding season. Using novel analytical methods based on bird atlas data, we show phenological shifts in migration of five species – red‐backed shrike, spotted flycatcher, common sandpiper, white‐winged tern (Palearctic migrants), and diederik cuckoo (intra‐African migrant) – between two atlas periods: 1987–1991 and 2007–2012. During this time period, Palearctic migrants advanced their departure from their South African nonbreeding grounds. This trend was mainly driven by waterbirds. No consistent changes were observed for intra‐African migrants. Our results suggest that the most consistent drivers of migration phenological shifts act in the northern hemisphere, probably at the breeding grounds.     

Evolution of altitudinal migration in passerines is linked to diet

Bird migration is typically associated with a latitudinal movement from north to south and vice versa. However, many bird species migrate seasonally with an upslope or downslope movement in a process termed altitudinal migration. Globally, 830 of the 6,579 Passeriformes species are considered altitudinal migrants and this pattern has emerged multiple times across 77 families of this order. Recent work has indicated an association between altitudinal migration and diet, but none have looked at diet as a potential evolutionary driver. Here, we investigated potential evolutionary drivers of altitudinal migration in passerines around the world by using phylogenetic comparative methods. We tested for evolutionary associations between altitudinal migration and foraging guild and primary habitat preference in passerines species worldwide. Our results indicate that foraging guild is evolutionarily associated with altitudinal migration, but this relationship varies across zoogeographical regions. In the Nearctic, herbivorous and omnivorous species are associated with altitudinal migration, while only omnivorous species are associated with altitudinal migration in the Palearctic. Habitat was not strongly linked to the evolution of altitudinal migration. While our results point to diet as a potentially important driver of altitudinal migration, the evolution of this behavior is complex and certainly driven by multiple factors. Altitudinal migration varies in its use (for breeding or molting), within a species, population, and even at the individual level. As such, the evolution of altitudinal migration is likely driven by an ensemble of factors, but this study provides a beginning framework for understanding the evolution of this complex behavior.     

Passerine migration strategies and body mass variation along geographic sectors across East Africa, the Middle East and the Arabian Peninsula

Using the body mass estimates of 12 long-distance migrating Palearctic passerine species monitored at successive sites across the Eastern Africa flyway, we tested whether birds modulate their body mass according to specific seasonal demands across different geographic sectors. We compared body mass estimates across latitudinal distances and geographic sectors in Europe, the desert, Northeast Africa and East Africa. Our results show that, depending on the species and season considered, the average body mass increase or decrease is variable at and among different geographic sectors. By comparing the variation in body mass between different ecological sectors, we were able to show when and where migrants accumulate their migratory fuel reserves during migration.     

Movement ecology of Afrotropical birds: Functional traits provide complementary insights to species identity

Effects of anthropogenic activities on habitats and species communities and populations are complex and vary across species depending on their ecological traits. Movement ecology may provide important insights into species’ responses to habitat structures and quality. We investigated how movement behavior across a human‐modified landscape depends on species identity and species traits, with particular focus on habitat specialization, feeding guilds, and dispersal behavior. We tracked 34 individuals of nine Afrotropical bird species during three years in an anthropogenic riparian landscape of East Africa. We investigated whether species’ functional traits predicted their habitat use and movement behavior better than species’ identities. Our results indicate that habitat specialists mainly occur in dense riparian thickets, while habitat generalists do occur in agricultural land. Home‐ranges of omnivorous habitat generalists are larger than of frugivorous and insectivorous generalists and omnivorous and insectivorous specialists. Movement speed was highest in settlement areas for all species, with activity peaks during morning and afternoon for habitat specialists. Our results reveal that functional traits and species identity provide complementary insights into responses of organisms to habitat structures and habitat quality.     

Habitat selection as a mechanism of resource partitioning in two cryptic bat species Pipistrellus pipistrellus and Pipistrellus pygmaeus

Ecomorphological studies of bat communities often reveal the spatial and temporal coexistence of morphologically similar species, leading to suggestions that these communities are structured by non-deterministic processes. However, the diversification of echolocation call structure in bats allows for considerable morphological similarity while still permitting niche differentiation based on specialisation for prey type and habitat structure. The recent separation of a common Palaearctic bat, the pipistrelle, into Pipistrellus pipistrellus and P. pygmaeus, which are sympatrically distributed throughout their range, raises the question as to whether these two morphologically similar species partition resources in time and space.
To test the hypothesis that the coexistence of these cryptic species is facilitated by differential habitat use, 14 P . pipistrellus , and 12 P. pygmaeus were radio-tracked from adjacent maternity roosts, in northeast Scotland, from May to September 2002/2003. The two species showed distinct habitat partitioning with P. pygmaeus foraging predominantly in riparian woodland and over water, and P. pipistrellus foraging along woodland edges and short isolated tree lines. Inter-specific overlap in habitat use was low and consequently foraging ranges were segregated spatially.
The degree of habitat partitioning revealed in these species, which show considerable overlap in echolocation call parameters and functional morphology, suggests that morphological features, whilst useful in separating chiropteran species into coarse-grained foraging guilds, may not predict fine-grained ecological segregation.