Biogeography of Great Basin butterflies: revisiting patterns, paradigms, and climate change scenarios

We used comprehensive data on butterfly distributions from six mountain ranges in the Great Basin to explore three connected biogeographic issues. First, we examined species richness and occurrence patterns both within and among mountain ranges. Only one range had a significant relationship between species richness and area. Relationships between species richness and elevation varied among mountain ranges. Species richness decreased as elevation increased in one range, increased as elevation increased in three ranges, and was not correlated in two ranges. In each range, distributional patterns were nested, but less vagile species did not always exhibit greater nestedness. Second, we compared our work with similar studies of montane mammals. Results from both taxonomic groups suggest that it may be appropriate to modify existing general paradigms of the biogeography of montane faunas in the Great Basin. Third, we revisited and refined previous predictions of how butterfly assemblages in the Great Basin may respond to climate change. The effects of climate change on species richness of montane butterflies may vary considerably among mountain ranges. In several ranges, few if any species apparently would be lost. Neither local species composition nor the potential order of species extirpations appears to be generalizable among ranges.     

Differences in habitat quality explain nestedness in a land snail meta-community

We set up two alternative hypotheses on how environmental variables could foster nestedness; one of “nested habitats” and another of “nested habitat quality”. The former hypothesis refers to situations where the nestedness of species depends on a nestedness of discrete habitats. The latter considers situations where all species in an assemblage increase in abundance along the same environmental gradient, but differ in specialisation or tolerance. We tested whether litter‐dwelling land snails (terrestrial gastropods) in boreal riparian forest exhibited a nested community structure, whether such a pattern was related to differences in environmental variables among sites, and which of the two hypotheses that best could account for the found pattern. We sampled litter from 100 m² plots in 29 mature riparian forest sites along small streams in the boreal zone of Sweden. The number of snail species varied between 3 and 14 per site. Ranking the species‐by‐site matrix by PCA scores of the first ordination axis revealed a similarly significant nested pattern as when the matrix was sorted by number of species, showing that the species composition in this meta‐community can be properly described as nested. Several environmental variables, most notably pH index, were correlated with the first PCA axis. All but two species had positive eigenvectors in the PCA ordination and the abundance increased considerably along the gradient for most of the species implying that the hypothesis of “nested habitats” was rejected in favour of the “nested habitat quality” hypothesis. Analyses of nestedness have seldom been performed on equal sized plots, and our study shows the importance of understanding that variation in environmental variables among sites can result in nested communities. The conservation implications are different depending on which of our two hypotheses is supported; a conservation focus on species “hotspots” is more appropriate if the communities are nested because of “nested habitat quality”.     

Distributions of forest birds and butterflies in the Andaman islands, Bay of Bengal: nested patterns and processes

The distributional patterns of forest birds and butterflies in the Andaman islands, an oceanic chain located off SE Asia, were tested for nestedness. Both taxa were highly nested. Nestedness could be due to colonization or extinction processes, area or distance effects or nestedness of habitats. Nestedness in forest bird distributions were strongly influenced by area and habitat related factors. Habitats were significantly nested in all three island groups, however most strongly for the North Andamans. However forest bird distributions in the North Andamans, as indicated by row order in the packed matrix, was not correlated with habitat diversity, suggesting that habitat related factors alone cannot account for these patterns. Other causal influences could be passive sampling, where common and abundant species and habitats are more likely to have a widespread distribution than rare species and habitats. The nested subset pattern seen in two unrelated taxa suggests that the Andamans are extinction dominated and that the protection of forests on large islands is critical for the conservation of its biodiversity.     

Comparative influence of spatial scale on beta diversity within regional assemblages of birds and butterflies

Aim We examined whether variation in species composition of breeding birds and resident butterflies in the Great Basin of North America depended on sampling grain (the smallest resolvable unit of study) and on the relative proximity of sampling units across the landscape. We also compared patterns between the two taxonomic groups with reference to their life‐history characteristics. Location Data for our analyses were collected from 1996 to 2003 in three adjacent mountain ranges in the central Great Basin (Lander and Nye counties, Nevada, USA): the Shoshone Mountains, Toiyabe Range and Toquima Range. Methods Data on species composition for both taxonomic groups were collecting using standard inventory methods for birds and butterflies in temperate regions. Data were compiled at three sampling grains, sites (average 12 ha), canyons (average 74 ha) and mountain ranges. For each sampling grain in turn, we calculated similarity of species composition using the Jaccard index. First, we investigated whether mean similarity of species composition among the three ranges differed as a function of the grain size at which data were compiled. Secondly, we explored whether mean similarity of species composition was greater for canyons within the same mountain range than for canyons within different mountain ranges. Thirdly, we examined whether mean similarity of species composition at the site level was different for sites within the same canyon, sites within different canyons in the same mountain range, and sites within canyons in different mountain ranges. We used a Bayesian model to analyse these comparisons. Results For both taxonomic groups, mean similarity of species composition increased as the sampling grain increased. The effect of spatial grain was somewhat greater for birds than for butterflies, especially when the intermediate sampling grain was compared with the smallest sampling grain. Similarity of species composition of butterflies at each sampling grain was greater than similarity of species composition of birds at the same grain. Mean similarity of species composition of both birds and butterflies at the canyon level and site level was affected by relative proximity of sampling locations; beta diversity increased as the relative isolation of sampling locations increased. Main conclusions The sensitivity of beta diversity to sampling grain likely reflects the effect of local environmental heterogeneity: as sampling grain increases, biotic assemblages appear more homogeneous. Although breeding birds in our study system have larger home ranges than resident butterflies, birds may have more specialized resource requirements related to vegetation structure and composition, especially at small sampling scales. The degree of variation in species composition of both taxonomic groups suggests that spatially extensive sampling will be more effective for drawing inferences about regional patterns of species diversity than intensive sampling at relatively few, smaller sites.     

Can human disturbance promote nestedness? A case study with breeding birds in urban habitat fragments

Several factors, singly or in combination, have proven relevant in promoting nestedness in animal communities (area, isolation, habitat structure, etc.); however, little empirical evidence exists about the role of human disturbance. The goal of this paper was to assess whether human disturbance (pedestrians) may affect bird species composition in such a way as to generate a nested distributional pattern. The study was conducted in an urban fragmented landscape, the city of Madrid, where wooded parks were suitable fragments for many bird species, and had different levels of human visitation. At the community level, the distribution of species was significantly nested in two consecutive breeding seasons. Using two different procedures ("temperature" and "departures" methods) to analyse causality in nestedness, I found that pedestrian rate, fragment size and the diversity of stems were significantly correlated to the nested pattern. When analysed simultaneously, and controlling for their independent effects, these factors still accounted for nestedness. Pedestrian rate was the only factor significantly associated with changes in species composition between years. At the individual species level, 74 and 41% of species were significantly nested in relation to pedestrian rate in 1997 and 1998, respectively; however, these percentages were independent of foraging substrate and body size. Besides the classic area and habitat diversity effects, human disturbance can also promote nestedness: locally, by restraining the time and space of foraging and breeding opportunities, thus reducing fragment suitability, and regionally, by increasing extinction and decreasing colonization probabilities in highly disturbed fragments.     

Nestedness, biogeographic theory, and the design of nature reserves

I examine the relationship between nested distributional patterns and the degree to which several small reserves will contain more species than would a single reserve of equal total area (SLOSS). Nestedness is a common property of species distributions on real and habitat islands. However, there is considerable variation in nestedness among species distributions, some of which is related to the physical and biological background of the archipelagoes. Nestedness does not vary according to the taxonomic group examined (with the exception of aquatic invertebrates). Nestedness does vary between real and habitat islands (with aquatic invertebrates excluded), but not between oceanic and land-bridge islands. The more a biota is nested, the more likely it is that a single large reserve would preserve more species. However, nestedness is a rather poor predictor of SLOSS, as the vast majority of archipelagoes support a strategy of several small reserves, even though almost all of them are significantly nested. Nestedness says little about optimal reserve design and management, and appears to be a weak conservation tool. Received: 30 May 1995 / Accepted: 1 April 1997     

The influence of colonization in nested species subsets

Biotic communities inhabiting collections of insular habitat patches often exhibit compositional patterns described as nested subsets. In nested biotas, the assemblages of species in relatively depauperate sites comprise successive subsets of species in relatively richer sites. In theory, nestedness may result from selective extinction, selective colonization, or other mechanisms, such as nested habitats. Allopatric speciation is expected to reduce nestedness. Previous studies, based largely on comparisons between land-bridge and oceanic archipelagos, have emphasized the role of selective extinction. However, colonization could also be important in generating strong patterns of nestedness. We apply a recently published index of nestedness to more than 50 island biogeographic data sets, and examine the roles of colonization, extinction, endemism, and, to a limited extent, habitat variability on the degree on nestedness. Most data sets exhibit a significant degree of nestedness, although there is no general tendency for land-bridge biotas to appear more nested than oceanic ones. Endemic species are shown to generally reduce nestedness. Comparisons between groups of non-endemic species differing in overwater or inter-patch dispersal ability indicate that superior dispersers generally exhibit a greater degree of nestedness than poorer dispersers, a result opposite that expected if colonization were a less predictable process than extinction. These results suggest that frequent colonization is likely to enhance nestedness, thereby increasing the compositional overlap among insular biotas. The prevalence of selective extinction in natural communities remains in question. The importance of colonization in generating and maintaining nested subsets suggests that (1) minimum critical areas will be difficult to determine from patterns of species distributions on islands; (2) multiple conservation sites are likely to be required to preserve communities in subdivided landscapes; and (3) management of dispersal processes may be as important to preserving species and communities as is minimizing extinctions.     

Effects of site and species characteristics on nested patterns of species composition in sedge meadows

Biotas of both geographical islands and habitat islands are often nested subsets of the biotas of successively more species-rich islands within the same system. The life history characteristics of a species may determine how that species contributes to the general pattern of species nestedness. Here, I investigate the floras of 56 sedge meadow wetlands in northern Illinois (USA) in order to characterize the degree of nestedness in these communities, determine which individual plant species contribute to the nested pattern, and investigate species characteristics that might be related to nonrandom patterns of distribution in individual plant species. The entire assemblage of species at all sedge meadows was significantly nested. Species richness and area were significantly correlated, and the nested pattern was closely related to site area, suggesting that species drop out of the assemblage in a predictable order as site area decreases. Some individual species exhibited nonrandom distributions across the sites, occurring more often in large, species-rich sites. Large sites were more likely than smaller sites to contain conservative species, i.e., those typical of pristine natural habitat, whereas nonconservative species were distributed more randomly among sites. Nested patterns of distribution of conservative species with respect to site area may result from their high probability of extinction on small sites or from a tendency for required habitats to co-occur on the same large sites. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nestedness and niche‐based species loss in moorland plant communities

Communities in isolated habitat patches surrounded by inhospitable matrices often form a nested subset pattern. However, the underlying causal mechanisms and conservation implications of nestedness in regional communities remain controversial. The nested ranks of species in a nested species‐by‐site matrix may reflect a gradient of species vulnerability to extinction or of colonization ability. However, nestedness analysis has rarely been used to explore determinants of species rank; consequently, little is known of underpinning mechanisms. In this study, we examined nestedness in moorland plant communities widely interspersed within the subalpine zone of northern Japan. Moorland sites differed in area (1000–160 000 m²) and were naturally isolated from one another to various extents within an inhospitable forest matrix. We also determined whether site characteristics (physical and morphometric measures) and species characteristics (niche position and breadth, based on species’ traits) are related to nestedness. Moorland plant communities in the study area were significantly nested. The pH and moorland kernel density (proxy for spatial clustering of moorlands around the focal site) were the most important predictors of moorland site nested rank in a nestedness matrix. Niche breadths of species (measured as variation in leaf mass area and height) predicted species’ nested ranks. Selective environmental tolerances imposed by environmental harshness and selective extinction caused by declines in site carrying capacities probably account for the nested subset pattern in moorland plant communities. The nested rank of species in the nestedness matrix can therefore be translated into the potential order of species loss explainable by species niche breadths (based on variation in functional traits). Complementary understanding of the determinants of site ranking and species ranking in the nestedness matrix provides powerful insight into ecological processes underlying nestedness and into the ways by which communities are assembled or disassembled by such processes.     

Does variation in patch use among butterfly species contribute to nestedness at fine spatial scales?

A nested pattern occurs whenever the species observed in depauperate habitat patches are a subset of those found in more species‐rich patches. Ecologists have documented many instances of nestedness caused by population‐level processes such as colonization and extinction at biogeographic scales. However, few researchers have examined whether nestedness may exist at fine scales due to the ways in which individual organisms discriminate among potential habitat patches. In 1999, we experimentally fragmented an old‐field habitat into patches of varying size to test whether nestedness could exist on a fine spatial scale. Five treatments of differing patch size were replicated five times in a Latin square design by selectively mowing 15×15 m² plots within an old‐field (patch areas: 225, 180, 135, 90, and 45 m²). Specifically, we tested whether butterflies foraging within a network of patches differing in area conformed to a nested subset structure. We also classified species according to (1) their flight height while foraging (high or low), and (2) their adult habitat breadth (ubiquitous, general, or restricted) to determine whether nestedness could be explained by difference in species’ tendency to discriminate among patches differing in area.
We found significant evidence that a community of foraging Lepidoptera conformed to a nested subset structure based on the difference between the observed nestedness within the butterfly community and the nestedness obtained from randomly generated species presence/absence matrices. Poisson regression analyses demonstrated that high‐flying, habitat‐restricted species avoided the smallest patches (90 and 45 m²) in favor of larger remnants, whereas low‐flying, habitat generalists used all patch sizes. Thus, our study is one of the first to demonstrate that nestedness among species subsets can be observed at fine spatial scales (within a single 1.5 hectare field) and may be maintained by species behavioral differences: discriminating species (i.e. high‐flying, habitat restricted) avoided the smallest patches, and less discriminating species (i.e. low‐flying, ubiquitous) were distributed throughout the field without regard to patch size. Our results also suggest that nestedness should be viewed as yet another scalar pattern in ecology, generated by variation in patch use by individuals at fine‐scales as well as the more traditionally invoked processes of extinction and colonization of species at broad‐scales.     

Nested Coleoptera and Orthoptera on sand dunes in the Basin and Range Province of western North America

The sand dunes of the Basin and Range Province of western North America contain obligate and endemic species of Coleoptera and Orthoptera. These dune habitats represent islands on which the isolated insular faunas are in a state of relaxation. The calculated temperature metric used in this study reflects the relative measure of disorder, by which the degree of nestedness can be determined. Sixteen dunes in the Basin and Range Province are shown to comprise a nested subset of obligate Coleoptera and Orthoptera. These sixteen sand dunes remain nested even when the endemic species are excluded from the analysis. The absence of endemic species slightly decreased the calculated temperature of the island-dune archipelago. Therefore, endemic species present in the sand dunes do not significantly contribute to the high degree of nestedness of dune obligate Coleoptera and Orthoptera in the Basin and Range Province. The dunes can also be separated into five distinct sub-basins, two of which contain only one sand dune. These sub-basins are not significantly nested, but together define the nested structure of the Basin and Range Province.     

Butterflies as an indicator group of riparian ecosystem assessment

Riparian ecosystems play an important role in modulating a range of ecosystem processes that affect aquatic and terrestrial organisms. Butterflies are a major herbivore in terrestrial ecosystems and are also common in riparian ecosystems. Since butterflies use plants for larval food and adult nectar sources in riparian ecosystems, butterfly diversity can be utilized to evaluate riparian ecosystems. We compiled butterfly data from 33 sites in three riparian ecosystem types across the country and compared butterfly diversity in terms of number of species and quality index in relation to riparian environmental variables. Number of butterfly and plant species was not different among three riparian habitat types. Additionally, there was no significant ecological variable to distinguish the butterfly communities on three riparian habitats. Non-metric multi-dimensional scaling ordination showed that butterfly communities in three riparian ecosystem types differed from each other, and butterfly riparian quality index was the main variable for butterfly assemblages. Five indicator species for moor and another five species for riverine riparian ecosystems were identified. Three and one indicator species for moor and riparian ecosystems, respectively, were plant specialists, while 44 butterflies were general feeders, feeding on a wide range of hostplants in several habitats. These results suggest that butterfly species use actively riparian habitats for nectar and larval food, and the butterfly riparian quality index can be employed to track faunal change in riparian habitats, which are frequently threatened by disturbances such as water level and climate changes, and invasive species.     

Seasonality and nestedness of bird communities in urban parks in Valencia,Spain

Although nested species subset patterns and mechanisms promoting them have constituted the focus of a considerable research effort, little attention has been paid to the role of seasonality in generating or moderating these patterns. I conducted monthly censuses of 130 urban parks in the city of Valencia throughout an annual cycle to assess whether seasonality influenced nested patterns of bird species assemblages. Specifically I tested the hypothesis that an increase in the movement of birds among parks outside of the breeding season, may reduce nestedness of resident birds. Results obtained in this study showed that assemblages of resident bird species were nested during the breeding season but not outside it, thus giving some support to the hypothesis. However, a reduction in nestedness outside the breeding season was not explained by an increase in the occupancy of parks but with an increase in the number of absences from parks that had been inhabited during the breeding season, most likely due to the use of habitats outside parks by some species (finches mainly). Results suggest the importance of both selective extinction and colonization processes in contributing to the degree of nestedness during the breeding season. However, environmental stress in the form of human disturbance apparently had little effect on the distributions of individual species. Habitat nestedness did not appear to be a factor as parks of all sizes were similar in number and type of habitats. An assessment of seasonality in other landscapes is needed to further understand its general effects on nestedness.     

Insular biogeography of the montane butterfly faunas in the Great Basin: comparison with birds and mammals

Summary Butterfly species lists were assembled for 18 Great Basin mountain ranges for which distributional data on mammals and birds have been analysed previously by other workers. The ranges represent remnant islands of the boreal habitat that once was continuous across the Great Basin but is now restricted to higher elevations as a result of climatic change at the close of the Pleistocene. The effects of biogeographic factors (area, distance, elevation) and habitat diversity on butterfly species number were examined. The Great Basin boreal butterfly faunas were found to be depauperate overall relative that of the principal mainland source, the Rocky Mountains, and were found to have fewer species than predicted by the mainland species-area data. However, only a weak area effect, and no distance effect, was detected by bivariate and multivariate analysis. Furthermore, the habitat diversity score found to explain virtually all the variation in bird species number in the same ranges in previous studies is only marginally significantly correlated with butterflies. When the butterflies are subdivided according to their vagility, the relative differences in the species-area correlation and slope (z-value) between the vagility categories were consistent with those found previously for mammals and birds, and, as predicted by theory, less vagile taxa exhibit higher species-area correlations and z-values. Overall, differences in the insular biogeography of buttterflies and vertebrates seem to reflect fundamental ecological differences between the taxa.     

Ecological traits reveal functional nestedness of bird communities in habitat islands: a global survey

The widespread destruction and fragmentation of natural habitats around the world creates a strong incentive to understand how species and communities respond to such pressures. The vast majority of research into habitat fragmentation has focused solely on species presence or absence. However, analyses using innovative functional methodologies offer the prospect of providing new insights into the key questions surrounding community structure in fragmented systems. A key topic in fragmentation research is nestedness (i.e. the ordered composition of species assemblages involving a significant tendency for packing of the presence–absence matrix into a series of proper subsets). To date, nestedness analyses have been concerned solely with nestedness of species membership. Here, we capitalize on the publication of a recent nestedness index (traitNODF) in which the branch lengths of functional dendrograms are incorporated into the standard NODF nestedness index. Using bird community data from 18 forest‐habitat‐island studies, and measurements of eight continuous functional traits from over 1000 bird species, we conduct the first synthetic analysis of nestedness from a functional perspective (i.e. a nestedness analysis which incorporates how similar species are in terms of their ecological traits). We use two null models to test the significance of any observed functional nestedness, and investigate the role of habitat island area in driving functional nestedness. We also determine whether functional nestedness is driven primarily by species composition or by differences in species’ traits. We found that the majority (94%) of datasets were functionally nested by island area when a permutation null model was used, although only 11–22% of datasets were significantly functionally nested when a more conservative fixed‐fixed null model was used. Species composition was always the most important driver of functional nestedness, but the effect of differences in species traits was occasionally quite large. Our results isolate the importance of island area in driving functional nestedness where it does occur and show that habitat loss results in the ordered loss of functional traits. This analysis demonstrates the potential insights that may derive from testing for ordered patterns of functional diversity. Synthesis The widespread fragmentation of natural habitats around the world creates a strong incentive to understand how ecological communities respond to such pressures. A key topic in this research agenda is nestedness; however, to date, nestedness analyses have been concerned solely with species presence or absence. Using data from 18 bird‐habitat‐island studies we conduct the first synthetic analysis of nestedness from a functional perspective (i.e. a nestedness analysis which incorporates how similar species are in terms of their ecological traits). Our findings suggest that many bird‐habitat island communities are significantly functionally nested, although our results were sensitive to the null model used. Our study demonstrates the benefits of testing for ordered patterns of functional diversity.     

Testing multiple assembly rule models in avian communities on islands of an inundated lake,Zhejiang Province,China

Aim A fundamental question in community ecology is whether general assembly rules determine the structure of natural communities. Although many types of assembly rules have been described, including Diamond’s assembly rules, constant body‐size ratios, favoured states, and nestedness, few studies have tested multiple assembly rule models simultaneously. Therefore, little is known about the relative importance of potential underlying factors such as interspecific competition, inter‐guild competition, selective extinction and habitat nestedness in structuring community composition. Here, we test the above four assembly rule models and examine the causal basis for the observed patterns using bird data collected on islands of an inundated lake. Location Thousand Island Lake, China. Methods  We collected data on presence–absence matrices, body size and functional groups for bird assemblages on 42 islands from 2007 to 2009. To test the above four assembly rule models, we used null model analyses to compare observed species co‐occurrence patterns, body‐size distributions and functional group distributions with randomly generated assemblages. To ensure that the results were not biased by the inclusion of species with extremely different ecologies, we conducted separate analyses for the entire assemblage and for various subset matrices classified according to foraging guilds. Results The bird assemblages did not support predictions by several competitively structured assembly rule models, including Diamond’s assembly rules, constant body‐size ratios, and favoured states. In contrast, bird assemblages were highly significantly nested and were apparently shaped by extinction processes mediated through area effects and habitat nestedness. The nestedness of bird assemblages was not a result of passive sampling or selective colonization. These results were very consistent, regardless of whether the entire assemblage or the subset matrices were analysed. Main conclusions Our results suggest that bird assemblages were shaped by extinction processes mediated through area effects and habitat nestedness, rather than by interspecific or inter‐guild competition. From a conservation point of view, our results indicate that we should protect both the largest islands with the most species‐rich communities and habitat‐rich islands in order to maximize the number of species preserved.     

Nestedness of butterfly assemblages in the Zhoushan Archipelago,China: area effects,life-history traits and conservation implications

The nested subset pattern (nestedness) of faunal assemblages has been a research focus in the fields of island biogeography and conservation biology in recent decades. However, relatively few studies have described nestedness in butterfly assemblages in oceanic archipelago systems. Moreover, previous studies often quantified nestedness using inappropriate nestedness metrics and random fill algorithms with high Type I errors. The aims of this study are to examine the existence of nestedness and underlying causal mechanisms of butterfly assemblages in the Zhoushan Archipelago, China. We used the line-transect method to determine butterfly occupancy and abundance on 42 study islands from July to August 2014. We obtained butterfly life-history traits (wingspan, body weight and minimum area requirement) by field work and island geographical features (area and isolation) from the literature. We used the recently developed metric WNODF to estimate nestedness. Partial Spearman rank correlation was used to evaluate the associations of nestedness and island geographical features as well as butterfly life-history traits related to species extinction risk and colonization ability. The butterfly assemblages were significantly nested. Island area and minimum area requirement of butterflies were significantly correlated with nestedness after controlling for other independent variables. In contrast, the nestedness of butterflies did not appear to result from passive sampling or selective colonization. However, multi-year studies are needed to confirm that target effects are not muddling these results. Our results indicate that selective extinction may be the main driver of nestedness of butterfly assemblages in our study system. From a conservation viewpoint, we should protect both large islands and species with large area requirement to maximize the number of species preserved.     

Ecological niche modelling of montane mammals in the Great Basin, North America: examining past and present connectivity of species across basins and ranges

Aim The goal of this study was to determine the extent of suitable habitats across the basins and ranges of the Great Basin for 13 montane mammals in the present and during the Last Glacial Maximum (LGM). For all these mammal species, we test whether: (1) more suitable habitat was available in basin areas during the LGM; (2) suitable habitat shifted upwards in elevation between the LGM and the present; (3) more ranges have suitable habitat than are currently occupied; and (4) these species are currently restricted to suitable habitats at higher‐elevation range areas. We also examine whether and how much distributional response varies among these montane mammal species. Location The Great Basin of western North America. Methods We re‐examine the past and present distributions of 13 Great Basin montane mammals using ecological niche modelling techniques that utilize now widely available species occurrence data and new, fine‐scale past climatological GIS layers in the present and at the LGM. These methods provide an objective, repeatable means for visual comparison of past and present modelled distributions for species examined in previous biogeographical studies. Results Our results indicate greater areal and lower elevational suitable habitat in the LGM than at present for nearly all montane mammals, and that there is more suitable habitat at present than is currently occupied. Our results also show that lowland areas provide suitable dispersal routes between ranges for most of the montane mammals both at the LGM and at present. However, three of the 13 species have little to no predicted suitable habitat in the LGM near currently occupied ranges, in contrast to the pattern for the other 10. For these species, the model results support more recent long‐distance colonization. Main conclusions Our finding of suitable lowland dispersal routes in the present for most species supports and greatly extends similar findings from single‐species studies. Our results also provide a visually striking confirmation that changes in species distribution and colonization histories of Great Basin montane mammals vary in a fashion related to the tolerances and requirements of each of these species; this has previously been hypothesized but not rigorously tested for multiple montane mammals in the region.     

Nested bird and micro-habitat assemblages in a peatland archipelago

Biotic assemblages of insular habitats are nested when poor assemblages are subsets of richer ones. Nestedness of species assemblages is frequent and may result from selective extinction or frequent colonization in insular habitats. It may also be created by a nested distribution of habitats among islands or by sampling bias. We sampled 67 isolated peatlands (7–843 ha) in southern Quebec, Canada, to measure nestedness of bird species assemblages among peatlands and assess the habitat nestedness hypothesis. Species and microhabitat assemblages were both strongly nested among peatlands. Whether sites were ranked by species richness, microhabitat richness or peatland area had no effect on nestedness. However, microhabitat nestedness was significantly reduced when sites were sorted by area rather than by microhabitat richness. As expected, if bird-microhabitat associations are responsible for the nested pattern of distribution, we found a positive correlation between the contributions of bird species and microhabitats to individual site nestedness. Nevertheless, microhabitat assemblages were significantly less nested than bird species assemblages, possibly because of frequent recolonization by birds or uneven sampling among sites. Received: 12 June 1998 / Accepted: 20 September 1998     

Ecosystem engineers maintain a rare species of butterfly and increase plant diversity

We evaluated whether ecosystem engineers can accomplish two conservation goals simultaneously: (1) indirectly maintain populations of an endangered animal through habitat modification and (2) increase riparian plant diversity. We tested for effects of a prominent ecosystem engineer, the beaver Castor canadensis, on populations of St. Francis' satyr butterfly Neonympha mitchellii francisci and plant species richness and composition. We performed our test by surveying riparian vegetation communities in all stages of beaver‐influenced wetland succession. We found that beavers created wetland habitats that supported plant species not found elsewhere in riparian zones and increased plant species diversity across the landscape by creating a novel combination of patch types. Our results confirmed what others have found about engineering effects on plant diversity, but these results further demonstrated a case where ecosystem engineers indirectly maintain populations of rare animals by modifying the composition and diversity of plant communities within wetlands. Our research demonstrates how an ecosystem engineer can influence habitat availability and composition of plant communities important for an endangered insect, and maintain overall plant species diversity by increasing habitat heterogeneity.