Patterns in the structure of grassland butterfly communities along a gradient of human disturbance: further analysis based on the generalist/specialist concept

Kitahara and Fujii, in 1994, analyzed the butterfly communities along a gradient of human disturbance by applying the generalist/specialist concept. Butterfly species were classified into generalist or specialist species based on their voltinism (seasonal time dimension) and potential larval resource breadth (food dimension). The community structure and species composition showed the systematic changes along the gradient. To verify the generality of those trends, we monitored five grassland butterfly communities with varying degrees of human disturbance twice a month during 1985 by the line transect method at the foot of Mt. Fuji, central Japan, and analyzed their structure in a manner similar to that employed by Kitahara and Fujii. Most results were consistent with the patterns recognized by Kitahara and Fujii. The route (community) order based on increasing human disturbance was strongly and negatively correlated with butterfly species richness but with neither butterfly species diversity (H′) nor evenness (J′). Also, the degree of human disturbance was significantly and negatively correlated with the number of specialist species, but not with that of generalists, in a community. Butterfly species richness was more strongly correlated with the number of specialist species than with that of generalists. Our analyses also showed that the generalist species were distributed more widely over the communities than were the specialists. However, in contrast to the trend revealed by Kitahara and Fujii, there was no significant difference in the population densities and in the spatial population variability between the two species groups. As a whole, our analyses confirmed the consistency of most community patterns detected by Kitahara and Fujii. The causes of the inconsistencies in some patterns were thought to be mainly the present habitat conditions with a relatively short growing season at high altitudes. Received: October 19, 1999 / Accepted: June 5, 2000     

The influence of matrix and edges on species richness patterns of ground beetles (Coleoptera: Carabidae) in habitat islands

Aim The aim of this study was to analyse whether, and how, the inclusion of habitat specialists and edge‐preferring species modifies the species–area relationship predictions of the island biogeography theory for an insect group (ground beetles, Coloptera: Carabidae) living in natural fragments. Species–habitat island area relationships applied to terrestrial habitat islands can be distorted by the indiscriminate inclusion of all species occurring in the fragments. Matrices surrounding terrestrial habitat fragments can provide colonists that do not necessarily distinguish the fragment from the matrix and can survive and reproduce there. Edge‐preferring species can further distort the expected relationship, as smaller fragments have larger edge:core ratios. Location Nineteen forest fragments were studied in the Bereg Plain, Hungary, and SW Ukraine. This area contains natural forest patches, mainly of oak and hornbeam, and supports a mountain entomofauna. Methods Ground beetles (Carabidae) present in the 19 forest patches were categorized into generalists, forest specialists and edge‐preferring species. We analysed the relationship between species richness and fragment area using species richness in the different categories. Results The assemblages contained a high share of generalist species (species that occur also in the surrounding matrix). Forest patch size and the number of generalist species showed a marginally significant negative relationship, indicating that generalist species were more important in smaller patches. Forest specialist species richness was correlated positively with patch area. Edge‐preferring species were shown to influence the species–area relationship: the number of edge‐preferring species increased with the edge:area ratio. Main conclusions Both generalist and edge‐preferring species can considerably distort the species–area relationship. Island biogeography theory can be applied to habitat islands only if the habitat islands are defined correctly from the viewpoint of the target species.     

Bird community patterns in response to the island features of urban woodlots in eastern China

Many studies have demonstrated the changes in the spatial patterns of plant and animal communities with respect to habitat fragmentation. Insular communities tend to exhibit some special patterns in connection with the characteristics of island habitats. In this paper, the relationships between richness, assemblage, and abundance of bird communities with respect to island features were analyzed in 20 urban woodlots in Hangzhou, China. Field investigations of bird communities, using the line transect method, were conducted from January to December, 1997. Each woodlot was surveyed 16 times during the year. Results indicated that bird richness was higher, per unit area, in the smaller woodlots than the larger ones, and overall bird density decreased with the increase in the size of woodlot. However, the evenness of species abundance increased with the area, and small woodlots were usually dominated by higher density species and large woodlots by medium density species. Most species occurring in the small woodlots also occurred in larger woodlots. Also, bird communities among urban woodlots showed a nestedness pattern in assemblage. These patterns implied that the main impacts of woodland habitat fragmentation are: (1) species are constricted and thus species number will increase at a given sample size; (2) as surface area decreases, the proportion of forest edge species as to interior species will increase; (3) community abundance will therefore increase per unit area but most individuals will be from a few dominant species; and (4) overall species diversity will decrease at a habitat level as well as at a region level. These patterns of community in response to the island features were therefore summarized as “island effects in community”. The underlying processes of such observations were also examined in this paper. Woodlot area, edge ratio, isolation, and habitat nestedness were considered as the important factors forming the island effects in community. High heterogeneity between habitats usually contributed most to the maintenance of regional biodiversity, especially in urban woodlots. __________ Translated from Acta Ecologica Sinica, 2005, 25(4): 657–663 [译自: 生态学报, 2005, 25(4): 657–663]     

How does landscape context contribute to effects of habitat fragmentation on diversity and population density of butterflies?

Aim Studies on habitat fragmentation of insect communities mostly ignore the impact of the surrounding landscape matrix and treat all species equally. In our study, on habitat fragmentation and the importance of landscape context, we expected that habitat specialists are more affected by area and isolation, and habitat generalists more by landscape context. Location and methods The study was conducted in the vicinity of the city of Göttingen in Germany in the year 2000. We analysed butterfly communities by transect counts on thirty‐two calcareous grasslands differing in size (0.03–5.14 ha), isolation index (2100–86,000/edge‐to‐edge distance 55–1894 m), and landscape diversity (Shannon–Wiener: 0.09–1.56), which is correlated to percentage grassland in the landscape. Results A total of 15,185 butterfly specimens belonging to fifty‐four species are recorded. In multiple regression analysis, the number of habitat specialist (n = 20) and habitat generalist (n = 34) butterfly species increased with habitat area, but z‐values (slopes) of the species–area relationships for specialists (z = 0.399) were significantly steeper compared with generalists (z = 0.096). Generalists, but not specialists, showed a marginally significant increase with landscape diversity. Effects of landscape diversity were scale‐dependent and significant only at the smallest scale (landscape context within a 250 m radius around the habitat). Habitat isolation was not related to specialist and generalist species numbers. In multiple regression analysis the density of specialists increased significantly with habitat area, whereas generalist density increased only marginally. Habitat isolation and landscape diversity did not show any effects. Main conclusions Habitat area was the most important predictor of butterfly community structure and influenced habitat specialists more than habitat generalists. In contrast to our expectations, habitat isolation had no effect as most butterflies could cope with the degree of isolation in our study region. Landscape diversity appeared to be important for generalist butterflies only.     

Biodiversity and community structure of temperate butterfly species within a gradient of human disturbance: An analysis based on the concept of generalist vs. Specialist strategies

We monitored nine butterfly communities with varying degrees of human disturbance by conducting a census twice a month during 1980 by the line transect method in and around Tsukuba City, central Japan. We analyzed the biodiversity and community structures using the generalist/specialist concept. The site (community) order based on decreasing human disturbance was positively correlated with butterfly species diversity (H′), species richness (the total number of species), and the number of specialist species in a community, but not with the number of generalist species. The number of generalist species was rather constant, irrespective of the degree of human disturbance. Thus, both the butterfly species diversity and species richness were more dependent on the specialists than the generalists. Our analyses also showed that the generalist species were distributed widely over the communities, and they maintained high population densities, resulting in high rank status in abundance in a community, with more spatial variation in density per species. Specialist species showed the opposite trends. These results demonstrate that the generalist/specialist concept is a powerful tool applicable to analyse the biodiversity and structure of natural communities.     

Area mediated shifts in bird community composition: a study on a fragmented Mediterranean grassland

The effects of habitat fragmentation on birds have often been studied in forest specialist species. Here we aimed at comparing the response of open habitat birds within a range of habitat specialization. The study area was a Mediterranean pseudo-steppe, designated as important for conservation yet fragmented by tree encroachment. We defined bird species dependency on steppe-like habitat by a correspondence analysis, allowing us to distinguish between specialists, generalists and scrubland species. We studied species abundance in relation to fragment area, testing whether species representation in fragments differed from those in continuous habitat. This analysis showed a contrasted response to fragment size between “open habitat” specialist species and generalist ones. Open habitat species were under-represented in the smallest fragments, while generalist were over-represented in small fragments in comparison to their distribution in continuous habitats. We discuss how these results can be linked to species habitat requirements. We find that scrubland species seem to be favoured by encroachment of woody vegetation, as they are able to explore and use the wooded matrix; however specialist species are restricted to open patches and are sensitive to a reduction in patch size. This allows us to predict how different species can exhibit a different sensitivity to habitat fragmentation.     

Large and least isolated fragments preserve habitat specialist spiders best in dry sandy grasslands in Hungary

The role of fragment size, isolation and habitat diversity in the conservation of spider assemblages living in fragmented landscape were studied in dry sandy grasslands (East Hungary, Nyírség). Spiders were collected using pitfall traps at eight dry grassland fragments from 2001 to 2009 from March to October fortnightly. We tested the rules of island biogeography, which suggest that the species richness increases with the size and decreases with the isolation of fragments. The habitat diversity is an important factor for species richness, since large areas usually have more habitats; therefore, the number of species may be higher in these areas. During the 9-year study period, altogether 10,544 individuals belonging to 106 species were collected. Contradicting the classical theory, we found a significant negative relationship between the total number of spider species and the grassland size, while the ratio of sandy grassland specialist spider species increased with fragment size. The relationship between the ratio of generalist species and the fragment size was not significant. The overall species richness and the isolation of studied grasslands did not show a significant relationship. The ratio of sandy grassland specialist species decreased, while the ratio of generalist species increased with the increasing of isolation. The habitat diversity did not show any effect on spider species richness. We concluded that to conserve the habitat specialist species it is recommended to preserve the large and least isolated grassland fragments, furthermore to increase the size of small fragments with the restoration of the adjacent croplands.     

19th century woodland structure controls stand-scale epiphyte diversity in present-day Scotland

Axioms developed from island biogeography theory (i.e. species–area relationships, effects of fragmentation and isolation) are central to the development of conservation strategy. Within this context, the 'extinction debt' hypothesis brings into question an often assumed relationship between species richness and present-day spatial habitat structure (i.e. extent, fragmentation), suggesting instead that the richness and composition of biological communities may lag behind spatial changes in habitat. We examined evidence for an extinction debt among epiphytic lichens, a highly diverse biological group of significant conservation concern. Using sites in Scotland, we compared epiphyte species richness in smaller-scale habitat units (aspen stands) to larger-scale woodland structure (extent and fragmentation) measured at two spatial scales (1 km² and 4 km²) and for two timeframes, modern (1990s to 2000s) and historic (1860s to 1880s). Species richness was positively related to woodland extent and negatively related to woodland fragmentation; however, richness was explained better by historic woodland structure at a 1-km² scale, than by modern woodland structure. The results indicate: (1) a coupling of stand-scale epiphyte assembly and dynamics of the wider woodland ecosystem, and (2) a significant lag in the response of epiphyte species richness to habitat spatial structure. However, the effect of spatial habitat structure is different between species groups with contrasting traits. The effect of decreasing woodland extent on epiphyte richness is generally more severe for microlichens (comprising a greater number of rare and specialist species) than the more generalist macrolichens.     

Does habitat loss affect the communities of plants and insects equally in plant–pollinator interactions? Preliminary findings

Habitat loss is a major threat to biodiversity and ecosystem function. As habitats are lost, one factor affecting their community structures is the niche-width demand of species, which ranges from specialist to generalist. This study focused on specialist and generalist species in plant–pollinator interactions and tested the hypothesis that plant and pollinator communities become more generalized as habitat loss increases. The study was made in seven selected sites in southern Ontario, Canada, at the level of landscape that is characterized by distributed forests within intensively managed agricultural fields. We quantified both the degree of habitat loss and the degree of specialization/generalization for each of the plant and insect communities using a sampling method of hexagonal transects. Regression analysis indicated a significant relationship between the increase of habitat loss and the shift to generalization in insect, but not in plant, communities. Our results suggest that, in plant–pollinator interactions, insect communities are more sensitive and/or quicker than plant communities to respond to the effects of habitat loss.     

Determinants of species richness in generalist and specialist Mediterranean butterflies: the negative synergistic forces of climate and habitat change

Although it is well established that butterfly richness is affected by climate and human factors (e.g. habitat disturbance and degradation) at different spatial scales, the drivers behind these changes vary greatly according to the geographical region and the ecology of the species concerned. It is essential that this variation be understood if trends in diversity are to be predicted with any degree of confidence under a scenario of global change. Here we examine patterns of butterfly species richness among groups differing in degree of habitat specialization, diet breadth and mobility in the north‐west Mediterranean Basin, a European hotspot for this taxon. We analyze a large number of butterfly communities and take into consideration the main potential drivers, that include climatic, geographic and resource variables, landscape structure and human environmental impact at different spatial scales. Our study shows that both climatic and anthropogenic factors play an important role in determining butterfly species richness in the north‐west Mediterranean Basin, but that their relative impact differs between specialist and generalist groups. At lower altitudes, water availability, a product of the interplay between temperature and rainfall, and negative effects of temperature appear as the most determinant factors. Maximum diversity was observed at mid‐altitudes, which reveals the importance from a conservation point of view of Mediterranean mountain ranges. The results suggest serious population declines in specialist species restricted to mountain areas as a result of climate warming in combination with habitat loss caused by the abandonment of grazing and mowing. They also suggest negative trends for generalist species due to an increase in aridity in combination with an increase in intensification of human land use in lowland areas. Such synergies are expected to lead to rapid declines in Mediterranean butterfly populations in the coming years, thereby posing a severe threat for the conservation of European biodiversity.     

Landscape composition and habitat area affects butterfly species richness in semi-natural grasslands

During the last 50 years, the distribution and abundance of many European butterfly species associated with semi-natural grasslands have declined. This may be the result of deteriorating habitat quality, but habitat loss, resulting in decreasing area and increasing isolation of remaining habitat, is also predicted to result in reduced species richness. To investigate the effects of habitat loss on species richness, we surveyed butterflies in semi-natural grasslands of similar quality and structure, but situated in landscapes of different habitat composition. Using spatially explicit habitat data, we selected one large (6–10 ha) and one small (0.5–2 ha) grassland site (pasture) in each of 24 non-overlapping 28.2 km² landscapes belonging to three categories differing in the proportion of the area that consisted of semi-natural grasslands. After controlling for local habitat quality, species richness was higher in grassland sites situated in landscapes consisting of a high proportion of grasslands. Species richness was also higher in larger grassland sites, and this effect was more pronounced for sedentary than for mobile species. However, the number of species for a given area did not differ between large and small grasslands. There was also a significant relationship between butterfly species richness and habitat quality in the form of vegetation height and abundance of flowers. In contrast, butterfly density was not related to landscape composition or grassland size. When species respond differently to habitat area or landscape composition this leads to effects on community structure, and nestedness analysis showed that depauperate communities were subsets of richer ones. Both grassland area and landscape composition may have contributed to this pattern, implying that small habitat fragments and landscapes with low proportions of habitat are both likely to mainly contain common generalist species. Based on these results, conservation efforts should aim at preserving landscapes with high proportions of the focal habitat.     

Island biogeography of bats in Baja California, Mexico: patterns of bat species richness in a near-shore archipelago

Aim We studied the relationship between the size and isolation of islands and bat species richness in a near‐shore archipelago to determine whether communities of vagile mammals conform to predictions of island biogeography theory. We compared patterns of species richness in two subarchipelagos to determine whether area per se or differences in habitat diversity explain variations in bat species richness. Location Islands in the Gulf of California and adjacent coastal habitats on the Baja California peninsula in northwest Mexico. Methods Presence–absence surveys for bats were conducted on 32 islands in the Gulf of California using acoustic and mist‐net surveys. We sampled for bats in coastal habitats of four regions of the Baja peninsula to characterize the source pool of potential colonizing species. We fitted a semi‐log model of species richness and multiple linear regression and used Akaike information criterion model selection to assess the possible influence of log₁₀ area, isolation, and island group (two subarchipelagos) on the species richness of bats. We compared the species richness of bats on islands with greater vegetation densities in the southern gulf (n = 20) with that on drier islands with less vegetation in the northern gulf (n = 12) to investigate the relationship between habitat diversity and the species richness of bats. Results Twelve species of bats were detected on islands in the Gulf of California, and 15 species were detected in coastal habitats on the Baja peninsula. Bat species richness was related to both area and isolation of islands, and was higher in the southern subarchipelago, which has denser vegetation. Log₁₀ area was positively related to bat species richness, which increased by one species for every 5.4‐fold increase in island area. On average, richness declined by one species per 6.25 km increase in isolation from the Baja peninsula. Main conclusions Our results demonstrate that patterns of bat species richness in a near‐shore archipelago are consistent with patterns predicted by the equilibrium theory of island biogeography. Despite their vagility, bats may be more sensitive to moderate levels of isolation than previously expected in near‐shore archipelagos. Differences in vegetation and habitat xericity appear to be associated with richness of bat communities in this desert ecosystem. Although observed patterns of species richness were consistent with those predicted by the equilibrium theory, similar relationships between species richness and size and isolation of islands may arise from patch‐use decision making by individuals (optimal foraging strategies).     

Factors determining mammal species richness on habitat islands and isolates: habitat diversity, disturbance, species interactions and guild assembly rules

• 1 For over three decades the equilibrium theory of island biogeography has galvanized studies in ecological biogeography. Studies of oceanic islands and of natural habitat islands share some similarities to continental studies, particularly in developed regions where habitat fragmentation results from many land uses. Increasingly, remnant habitat is in the form of isolates created by the clearing and destruction of natural areas. Future evolution of a theory to predict patterns of species abundance may well come from the application of island biogeography to habitat fragments or isolates.
• 2 In this paper we consider four factors other than area and isolation that influence the number and type of mammal species coexisting in one place: habitat diversity, habitat disturbance, species interactions and guild assembly rules. In all examples our data derive from mainland habitat, fragmented to differing degrees, with different levels of isolation.
• 3 Habitat diversity is seen to be a good predictor of species richness. Increased levels of disturbance produce a relatively greater decrease in species richness on smaller than on larger isolates. Species interactions in the recolonization of highly disturbed sites, such as regenerating mined sites, is analogous to island colonization. Species replacement sequences in secondary successions indicate not just how many, but which species are included. Lastly, the complement of species established on islands, or in insular habitats, may be governed by guild assembly rules. These contributions may assist in taking a renewed theory into the new millennium.

     

Conservation implications of competition between generalist and specialist rodents in Mediterranean afforested landscape

Density dependent habitat selection at the community level is regarded as a major determinant of biodiversity at the local scale, and data on these processes and how they are affected by human activities is highly applicable to conservation. By studying the competitive relationships between a specialist and a generalist we can acquire valuable insights about how different environmental elements determine species abundance and distribution and consequently biodiversity. Here we describe a study of density dependent processes that determine the community structure of two rodents: a specialist—the broad toothed mouse (Apodemus mystacinus), and a generalist—the common spiny mouse (Acomys cahirinus) in a Mediterranean maqui habitat, and how this structure is impacted by anthropogenic planting of pine stands. We carried out two field experiments: The first, based on open field trapping, looking at how rodent communities change with habitat structure. The second experiment was an enclosure study aimed at validating the habitat preferences and competitive relationship between the specialist and the generalist. We identified asymmetric competition relationships in which the specialist was dominant over the generalist. Competition intensity was lower in maqui with >10% oak cover, although both species abundances were high. Competition was found only during the limiting season (summer). Based on these findings we produced management recommendations to keep indigenous small mammals’ biodiversity high. Density dependent habitat selection processes play a central role in determining biodiversity, and understanding the mechanisms motivating these processes is needed if alterations in biodiversity in response to human disturbance are to be understood.     

Spatial and ecological population genetic structures within two island‐endemic Aeonium species of different niche width

The Crassulacean genus Aeonium is a well‐known example for plant species radiation on oceanic archipelagos. However, while allopatric speciation among islands is documented for this genus, the role of intra‐island speciation due to population divergence by topographical isolation or ecological heterogeneity has not yet been addressed. The aim of this study was to investigate intraspecific genetic structures and to identify spatial and ecological drivers of genetic population differentiation on the island scale. We analyzed inter simple sequence repeat variation within two island‐endemic Aeonium species of La Palma: one widespread generalist that covers a large variety of different habitat types (Ae. davidbramwellii) and one narrow ecological specialist (Ae. nobile), in order to assess evolutionary potentials on this island. Gene pool differentiation and genetic diversity patterns were associated with major landscape structures in both species, with phylogeographic implications. However, overall levels of genetic differentiation were low. For the generalist species, outlier loci detection and loci–environment correlation approaches indicated moderate signatures of divergent selection pressures linked to temperature and precipitation variables, while the specialist species missed such patterns. Our data point to incipient differentiation among populations, emphasizing that ecological heterogeneity and topographical structuring within the small scales of an island can foster evolutionary processes. Very likely, such processes have contributed to the radiation of Aeonium on the Canary Islands. There is also support for different evolutionary mechanisms between generalist and specialist species.     

Do asynchronies in extinction debt affect the structure of trophic networks? A case study of antagonistic butterfly larvae–plant networks

Habitat loss and fragmentation affect species richness in fragmented habitats and can lead to immediate or time‐delayed species extinctions. Asynchronies in extinction and extinction debt between interacting species may have severe effects on ecological networks. However, these effects remain largely unknown. We evaluated the effects of habitat patch and landscape changes on antagonistic butterfly larvae–plant trophic networks in Mediterranean grasslands in which previous studies had shown the existence of extinction debt in plants but not in butterflies. We sampled current species richness of habitat‐specialist and generalist butterflies and vascular plants in 26 grasslands. We assessed the direct effects of historical and current patch and landscape characteristics on species richness and on butterfly larvae–plant trophic network metrics and robustness. Although positive species‐ and interactions–area relationships were found in all networks, structure and robustness was only affected by patch and landscape changes in networks involving the subset of butterfly specialists. Larger patches had more species (butterflies and host plants) and interactions but also more compartments, which decreased network connectance but increased network stability. Moreover, most likely due to the rescue effect, patch connectivity increased host‐plant species (but not butterfly) richness and total links, and network robustness in specialist networks. On the other hand, patch area loss decreased robustness in specialist butterfly larvae–plant networks and made them more prone to collapse against host plant extinctions. Finally, in all butterfly larvae–plant networks we also detected a past patch and landscape effect on network asymmetry, which indicates that there were different extinction rates and extinction debts for butterflies and host plants. We conclude that asynchronies in extinction and extinction debt in butterfly–plant networks provoked by patch and landscape changes caused changes in species richness and network links in all networks, as well as changes in network structure and robustness in specialist networks.     

Vascular plant species richness in relation to habitat diversity and island area in the Finnish Archipelago

Aim The aim of this study is to explore the interrelationships between island area, species number and habitat diversity in two archipelago areas. Location The study areas, Brunskär and Getskär, are located in an archipelago in south‐western Finland. Methods The study areas, 82 islands in Brunskär and 78 in Getskär, were classified into nine habitat types based on land cover. In the Brunskär area, the flora (351 species) was surveyed separately for each individual habitat on the islands. In the Getskär area, the flora (302 species) was surveyed on a whole‐island basis. We used standard techniques to analyse the species–area relationship on a whole‐island and a habitat level. We also tested our data for the small island effect (SIE) using breakpoint and path analysis models. Results Species richness was significantly associated with both island area and habitat diversity. Vegetated area in particular, defined as island area with the rock habitat subtracted, proved to be a strong predictor of species richness. Species number had a greater association with island area multiplied by the number of habitats than with island area or habitat number separately. The tests for a SIE in the species–area relationship showed the existence of a SIE in one of the island groups. No SIE could be detected for the species–vegetated area relationship in either of the island groups. The strength of the species–area relationship differed considerably between the habitats. Main conclusions The general principles of island biogeography apply well to the 160 islands in this study. Vascular plant diversity for small islands is strongly influenced by physiographic factors. For the small islands with thin and varying soil cover, vegetated area was the most powerful predictor of species richness. The species–area curves of various habitats showed large variations, suggesting that the measurement of habitat areas and establishment of habitat‐based species lists are needed to better understand species richness on islands. We found some evidence of a SIE, but it is debatable whether this is a ‘true’ SIE or a soil cover/habitat characteristics feature.     

Habitat structure and predators: choices and consequences for rodent habitat specialists and generalists

Summary Both habitat structure and risk of predation are thought to influence rodent community composition in different habitats, but experiments on the degree to which these factors determine the use of habitat by rodents are lacking. I sought to discover (1) if cover density altered habitat choice and (2) if cover density affected the vulnerability to predators of two rodents, a habitat specialist and a habitat generalist. In laboratory experiments, the habitat specialist, the red-backed vole (Clethrionomys gapperi), preferred greater densities of both vertical (wall) and horizontal (ceiling) cover. The habitat generalist, the deer mouse (Peromyscus maniculatus), also preferred greater densities of vertical and horizontal cover, but its preferences were weaker and more inconsistent than those of C. gapperi In tests of vulnerability to domestic ferrets, C. gapperi were more vulnerable in arenas with less vertical cover, while P. maniculatus did not differ in vulnerability between the two vertical cover densities used. Vulnerability to predators in differing densities of horizontal cover was not tested because of the reduced differences in preference for this cover type between the rodent species. Risk of predation is one explanation for C. gapper's inherent preference for denser cover.     

Subsidized Island Biogeography Hypothesis: another new twist on an old theory

We present a new hypothesis for predicting and describing patterns of species diversity on small islands and habitat fragments. We have modified the traditional island biogeography equilibrium theory to incorporate the influence of spatial subsidies from the surrounding matrix, which vary among islands and habitat fragments, on species diversities. The modification indicates three possible directions for the effects of spatial subsidies on diversity, which depend on where the focal community falls on the hypothesized unimodal curve of the productivity–diversity relationship. The idea is novel because no recent syntheses of productivity–diversity–area relationships examine the role of allochthonous resources on recipient communities’ diversity patterns.     

Flying between Sky Islands: The Effect of Naturally Fragmented Habitat on Butterfly Population Structure

High elevation montane areas are called “sky islands” when they occur as a series of high mountains separated by lowland valleys. Different climatic conditions at high elevations makes sky islands a specialized type of habitat, rendering them naturally fragmented compared to more continuous habitat at lower elevations. Species in sky islands face unsuitable climate in the intervening valleys when moving from one montane area to another. The high elevation shola-grassland mosaic in the Western Ghats of southern India form one such sky island complex. The fragmented patches make this area ideal to study the effect of the spatial orientation of suitable habitat patches on population genetic structure of species found in these areas. Past studies have suggested that sky islands tend to have genetically structured populations, possibly due to reduced gene flow between montane areas. To test this hypothesis, we adopted the comparative approach. Using Amplified Fragment Length Polymorphisms, we compared population genetic structures of two closely related, similar sized butterfly species: Heteropsis oculus, a high elevation shola-grassland specialist restricted to the southern Western Ghats, and Mycalesis patnia, found more continuously distributed in lower elevations. In all analyses, as per expectation the sky island specialist H. oculus exhibited a greater degree of population genetic structure than M. patnia, implying a difference in geneflow. This difference in geneflow in turn appears to be due to the natural fragmentation of the sky island complexes. Detailed analysis of a subset of H. oculus samples from one sky island complex (the Anamalais) showed a surprising genetic break. A possible reason for this break could be unsuitable conditions of higher temperature and lower rainfall in the intervening valley region. Thus, sky island species are not only restricted by lack of habitat continuity between montane areas, but also by the nature of the intervening habitat.