The roles of landscape context, niche breadth, and range boundaries in predicting species responses to habitat alteration

Extant species in human‐dominated landscapes differ in their sensitivity to habitat loss and fragmentation, although extinctions induced by environmental alteration reduce variation and result in a surviving subset of species with some degree of ‘resistance’. Here, we test the degree to which variable responses to habitat alteration are (1) essentially an inherent property of a taxon subject to constraints imposed by its geographical range, as suggested by Swihart et al. (2003), (2) a function of the landscape in which a species occurs, or (3) a function of spatial trends occurring on large scales. We used data collected on 33 vertebrate species during 2001–04 across the upper Wabash River basin, Indiana, in 35 square ‘landscapes’, each 23 km² in size. Six species of forest rodent, six species of grassland rodents, seven species of bats, eight species of aquatic turtles, and six species of amphibians were sampled at 504, 212, 590, 228, and 625 patches, respectively. The fraction of patches of primary habitat (e.g. forests for tree squirrels, wetlands for aquatic turtles) occupied by a target species was used as a response variable. On a basin‐wide scale, 47% of variation in proportional occupancy among species could be explained by taxon‐specific variables; occupancy rates were related positively to niche breadth and negatively to the proximity of a geographical range boundary. After controlling for species effects, landscape‐level occupancy rates varied significantly for 16 of 33 species, with variation partitioned among landscape variables alone (mean = 11% of variation), spatial trend variables alone (26%), and both variable sets jointly (8%). Among landscape variables, percentage forest cover positively affected occupancy rates of three bat species and a tree squirrel. Variation in occupancy rates among landscapes was consistent with large‐scale spatial trends for 13 species. Our findings demonstrate the general importance of niche breadth as a predictor of species responses to habitat alteration and highlight the importance of viewing the effects of habitat loss and fragmentation at multiple spatial scales.     

Changes in habitat specificity of species at their climatic range boundaries

Species are thought to have more restricted niches towards their range boundaries, although this has rarely been quantified systematically. We analysed transect data for 41 butterfly species along climatic gradients within Britain and show that 71% of species have broader niches at sites with milder winters. Shifts in habitat associations are considerable across most species' ranges; averaged across all 41 species, we estimate that if 26% of individuals were associated with the favoured habitat on the species' warmest transect, then 70% of individuals would be confined to this habitat on the species' coldest transect. Species with more southerly distributions in Britain showed the greatest changes in their habitat associations. We conclude that geographic variation in realized niche breadth is common and relatively large, especially near range boundaries, and should be taken into account in conserving species under changing climates.     

Impacts of climate warming and habitat loss on extinctions at species' low-latitude range boundaries

Polewards expansions of species' distributions have been attributed to climate warming, but evidence for climate‐driven local extinctions at warm (low latitude/elevation) boundaries is equivocal. We surveyed the four species of butterflies that reach their southern limits in Britain. We visited 421 sites where the species had been recorded previously to determine whether recent extinctions were primarily due to climate or habitat changes. Coenonympha tullia had become extinct at 52% of study sites and all losses were associated with habitat degradation. Aricia artaxerxes was extinct from 50% of sites, with approximately one‐third to half of extinctions associated with climate‐related factors and the remainder with habitat loss. For Erebia aethiops (extinct from 24% of sites), approximately a quarter of the extinctions were associated with habitat and three‐quarters with climate. For Erebia epiphron, extinctions (37% of sites) were attributed mainly to climate with almost no habitat effects. For the three species affected by climate, range boundaries retracted 70–100 km northwards (A. artaxerxes, E. aethiops) and 130–150 m uphill (E. epiphron) in the sample of sites analysed. These shifts are consistent with estimated latitudinal and elevational temperature shifts of 88 km northwards and 98 m uphill over the 19‐year study period. These results suggest that the southern/warm range margins of some species are as sensitive to climate change as are northern/cool margins. Our data indicate that climate warming has been of comparable importance to habitat loss in driving local extinctions of northern species over the past few decades; future climate warming is likely to jeopardize the long‐term survival of many northern and mountain species.     

Modelling the effect of habitat fragmentation on range expansion in a butterfly

There is an increasing need for conservation programmes to make quantitative predictions of biodiversity responses to changed environments. Such predictions will be particularly important to promote species recovery in fragmented landscapes, and to understand and facilitate distribution responses to climate change. Here, we model expansion rates of a test species (a rare butterfly, Hesperia comma) in five landscapes over 18 years (generations), using a metapopulation model (the incidence function model). Expansion rates increased with the area, quality and proximity of habitat patches available for colonization, with predicted expansion rates closely matching observed rates in test landscapes. Habitat fragmentation constrained expansion, but in a predictable way, suggesting that it will prove feasible both to understand variation in expansion rates and to develop conservation programmes to increase rates of range expansion in such species.     

Climate change and human colonization triggered habitat loss and fragmentation in Madagascar

The relative effect of past climate fluctuations and anthropogenic activities on current biome distribution is subject to increasing attention, notably in biodiversity hot spots. In Madagascar, where humans arrived in the last ~4 to 5,000 years, the exact causes of the demise of large vertebrates that cohabited with humans are yet unclear. The prevailing narrative holds that Madagascar was covered with forest before human arrival and that the expansion of grasslands was the result of human‐driven deforestation. However, recent studies have shown that vegetation and fauna structure substantially fluctuated during the Holocene. Here, we study the Holocene history of habitat fragmentation in the north of Madagascar using a population genetics approach. To do so, we infer the demographic history of two northern Madagascar neighbouring, congeneric and critically endangered forest dwelling lemur species—Propithecus tattersalli and Propithecus perrieri—using population genetic analyses. Our results highlight the necessity to consider population structure and changes in connectivity in demographic history inferences. We show that both species underwent demographic fluctuations which most likely occurred after the mid‐Holocene transition. While mid‐Holocene climate change probably triggered major demographic changes in the two lemur species range and connectivity, human settlements that expanded over the last four millennia in northern Madagascar likely played a role in the loss and fragmentation of the forest cover.     

苜蓿草地生境丧失与破碎化对昆虫物种丧失与群落重建的影响

生境破碎化包括生境丧失与破碎化两个相对独立的过程,为探讨这两个过程各自对生物多样性的影响,本文利用苜蓿草地实验模型系统(EMS)构建了36个小区研究不同生境丧失与破碎化对昆虫群落及不同类群的影响,包括18个破碎化小区与18个连续小区,破碎化小区全部采用1 m×1 m(H=1)破碎,连续小区苜蓿连片(H=0),生境丧失采...     

How fire interacts with habitat loss and fragmentation

Biodiversity faces many threats and these can interact to produce outcomes that may not be predicted by considering their effects in isolation. Habitat loss and fragmentation (hereafter ‘fragmentation’) and altered fire regimes are important threats to biodiversity, but their interactions have not been systematically evaluated across the globe. In this comprehensive synthesis, including 162 papers which provided 274 cases, we offer a framework for understanding how fire interacts with fragmentation. Fire and fragmentation interact in three main ways: (i) fire influences fragmentation (59% of 274 cases), where fire either destroys and fragments habitat or creates and connects habitat; (ii) fragmentation influences fire (25% of cases) where, after habitat is reduced in area and fragmented, fire in the landscape is subsequently altered because people suppress or ignite fires, or there is increased edge flammability or increased obstruction to fire spread; and (iii) where the two do not influence each other, but fire interacts with fragmentation to affect responses like species richness, abundance and extinction risk (16% of cases). Where fire and fragmentation do influence each other, feedback loops are possible that can lead to ecosystem conversion (e.g. forest to grassland). This is a well-documented threat in the tropics but with potential also to be important elsewhere. Fire interacts with fragmentation through scale-specific mechanisms: fire creates edges and drives edge effects; fire alters patch quality; and fire alters landscape-scale connectivity. We found only 12 cases in which studies reported the four essential strata for testing a full interaction, which were fragmented and unfragmented landscapes that both span contrasting fire histories, such as recently burnt and long unburnt vegetation. Simulation and empirical studies show that fire and fragmentation can interact synergistically, multiplicatively, antagonistically or additively. These cases highlight a key reason why understanding interactions is so important: when fire and fragmentation act together they can cause local extinctions, even when their separate effects are neutral. Whether fire–fragmentation interactions benefit or disadvantage species is often determined by the species' preferred successional stage. Adding fire to landscapes generally benefits early-successional plant and animal species, whereas it is detrimental to late-successional species. However, when fire interacts with fragmentation, the direction of effect of fire on a species could be reversed from the effect expected by successional preferences. Adding fire to fragmented landscapes can be detrimental for species that would normally co-exist with fire, because species may no longer be able to disperse to their preferred successional stage. Further, animals may be attracted to particular successional stages leading to unexpected responses to fragmentation, such as higher abundance in more isolated unburnt patches. Growing human populations and increasing resource consumption suggest that fragmentation trends will worsen over coming years. Combined with increasing alteration of fire regimes due to climate change and human-caused ignitions, interactions of fire with fragmentation are likely to become more common. Our new framework paves the way for developing a better understanding of how fire interacts with fragmentation, and for conserving biodiversity in the face of these emerging challenges.     

The relationship between range size and niche breadth: a test using five species of Gammarus (Amphipoda)

Data for five closely related species of gammarid crustaceans are used to examine interspecific relationships between the breadth of fundamental tolerance or capacity and geographical range size. Gammarus duebeni is, almost without exception, the most tolerant species and that with the best physiological performance. Although there is some limited variation, the remaining species can be ranked broadly in the sequence G. zaddachi  > G. salinus  >  G. oceanicus > G. locusta . The wide tolerance and high performance of G. duebeni is associated with the occupation of a wider range of environmental 'types' than any other of the species. In terms of geographical range size, the species can be ranked from most to least widespread in the sequence G. oceanicus  > G. duebeni  >  G. zaddachi  >  G. salinus  >  G. locusta . This provides little support for Brown's hypothesis, or the argument that the more widely distributed species within a taxonomic assemblage also tend to have the widest fundamental niches. However, if marine ( G. oceanicus and G. locusta ) and estuarine ( G. duebeni , G. zaddachi , G. salinus) species are considered separately, then in each case the species with the largest geographical range is also the most tolerant/best performer. In this sense, the jack-of-all-trades is the master-of-all, rather than the master-of-none.     

The geometry of habitat fragmentation: Effects of species distribution patterns on extinction risk due to habitat conversion

Land‐use changes, which cause loss, degradation, and fragmentation of natural habitats, are important anthropogenic drivers of biodiversity change. However, there is an ongoing debate about how fragmentation per se affects biodiversity in a given amount of habitat. Here, we illustrate why it is important to distinguish two different aspects of fragmentation to resolve this debate: (a) geometric fragmentation effects, which exclusively arise from the spatial distributions of species and habitat fragments, and (b) demographic fragmentation effects due to reduced fragment sizes, and/or changes in fragment isolation, edge effects, or species interactions. While most empirical studies are primarily interested in quantifying demographic fragmentation effects, geometric effects are typically invoked as post hoc explanations of biodiversity responses to fragmentation per se. Here, we present an approach to quantify geometric fragmentation effects on species survival and extinction probabilities. We illustrate this approach using spatial simulations where we systematically varied the initial abundances and distribution patterns (i.e., random, aggregated, or regular) of species as well as habitat amount and fragmentation per se. As expected, we found no geometric fragmentation effects when species were randomly distributed. However, when species were aggregated, we found positive effects of fragmentation per se on survival probability for a large range of scenarios. For regular species distributions, we found weakly negative geometric effects. These findings are independent of the ecological mechanisms which generate nonrandom species distributions. Our study helps to reconcile seemingly contradictory results of previous fragmentation studies. Since intraspecific aggregation is a ubiquitous pattern in nature, our findings imply widespread positive geometric fragmentation effects. This expectation is supported by many studies that find positive effects of fragmentation per se on species occurrences and diversity after controlling for habitat amount. We outline how to disentangle geometric and demographic fragmentation effects, which is critical for predicting the response of biodiversity to landscape change.     

Climate envelope modelling reveals intraspecific relationships among flowering phenology, niche breadth and potential range size in Arabidopsis thaliana

Species often harbour large amounts of phenotypic variation in ecologically important traits, and some of this variation is genetically based. Understanding how this genetic variation is spatially structured can help to understand species' ecological tolerances and range limits. We modelled the climate envelopes of Arabidopsis thaliana genotypes, ranging from early- to late-flowering, as a function of several climatic variables. We found that genotypes with contrasting alleles at individual flowering time loci differed significantly in potential range size and niche breadth. We also found that later flowering genotypes had more restricted range potentials and narrower niche breadths than earlier flowering genotypes, indicating that local selection on flowering can constrain or enhance the ability of populations to colonise other areas. Our study demonstrates how climate envelope models that incorporate ecologically important genetic variation can provide insights into the macroecology of a species, which is important to understand its responses to changing environments.     

Home range size scales to habitat amount and increasing fragmentation in a mobile woodland specialist

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Regional occupancy in unicellular eukaryotes: a reflection of niche breadth,habitat availability or size‐related dispersal capacity?

1. The distribution patterns of unicellular and multicellular organisms have recently been shown to differ profoundly, with the former probably being mostly cosmopolitan, whereas the latter are mostly restricted to certain regions. However, the within‐region distribution patterns of these two organism groups may be rather similar. 2. We predicted that the degree of regional occupancy in unicellular eukaryotes would be related to niche characteristics, dispersal ability and size, as has been found previously for multicellular organisms. The niche characteristics we considered were niche position, that measures marginality in species habitat distribution, and niche breadth, that measures amplitude in species habitat distribution. Niche characteristics were determined using Outlying Mean Index (OMI) analysis. 3. We found that the regional occupancy in our model group of unicellular eukaryotes, stream diatoms, was primarily a reflection of the niche position of a species or, more generally, habitat availability. Thus, non‐marginal species (i.e. species that occupied common habitat conditions across the region) tended to be more widely distributed than marginal species (i.e. species that were restricted to a limited range of rare habitat conditions). This finding was further supported by the general linear model, with niche position, niche breadth, maximum size and attachment mode as explanatory variables: niche position was by far the most important variable accounting for variability in regional occupancy, with significant amounts of additional variation related to niche breadth and maximum size of diatoms. 4. Thus, the degree of regional occupancy among unicellular eukaryotes may be primarily governed by habitat availability, supporting former findings for multicellular organisms.     

Of niches and distributions: range size increases with niche breadth both globally and regionally but regional estimates poorly relate to global estimates

The relationship between species’ niche breadth (i.e. the range of environmental conditions under which a species can persist) and range size (i.e. the extent of its spatial distribution) has mostly been tested within geographically restricted areas but rarely at the global extent. Here, we not only tested the relationship between range size (derived from species’ distribution data) and niche breadth (derived from species’ distribution and co‐occurrence data) of 1255 plant species at the regional extent of the European Alps, but also at the global extent and across both spatial scales for a subset of 180 species. Using correlation analyses, linear models and variation partitioning, we found that species’ realized niche breadth estimated at the regional level is a weak predictor of species’ global niche breadth and range size. Against our expectations, distribution‐derived niche breadth was a better predictor for species’ range size than the co‐occurrence‐based estimate, which should, theoretically, account for more than the climatically determined niche dimensions. Our findings highlight that studies focusing on the niche breadth vs range size relationship must explicitly consider spatial mismatches that might have confounded and diminished previously reported relationships.     

Distribution, abundance and niche breadth of birds: scale matters

We used local habitat niche breadth, local abundance and body size of non-passerine afrotropical birds in Tsavo East National Park (Kenya) to predict species distributional ranges in Kenya and across Africa. Univariate analysis revealed a significant positive correlation between local abundance and distribution only on the scale of Kenya. Performing a multiple regression analysis, local abundance, local habitat niche breadth and body size explained a significant part of the variance in bird distribution, again only on the Kenyan scale. From these results, we speculate that on continental scales distributions may be more influenced by macroclimatic conditions and historical factors, whereas distributions on regional scales are predominantly influenced by ecological factors.     

Effects of habitat loss, habitat fragmentation, and isolation on the density, species richness, and distribution of ladybeetles in manipulated alfalfa landscapes

Abstract.  1. Habitat loss and fragmentation are the main causes of changes in the distribution and abundance of organisms, and are usually considered to negatively affect the abundance and species richness of organisms in a landscape. Nevertheless, habitat loss and fragmentation have often been confused, and the reported negative effects may only be the result of habitat loss alone, with habitat fragmentation having nil or even positive effects on abundance and species richness.
2. Manipulated alfalfa micro-landscapes and coccinellids (Coleoptera: Coccinellidae) are used to test the effects habitat loss (0% or 84%), fragmentation (4 or 16 fragments), and isolation (2 or 6 m between fragments) on the density, species richness, and distribution of native and exotic species of coccinellids.
3. Generally, when considering only the individuals in the remaining fragments, habitat loss had variable effects while habitat fragmentation had a positive effect on the density of two species of coccinellids and on species richness, but did not affect two other species. Isolation usually had no effect. When individuals in the whole landscape were considered, negative effects of habitat loss became apparent for most species, but the positive effects of fragmentation remained only for one species.
4. Native and exotic species of coccinellids did not segregate in the different landscapes, and strong positive associations were found most often in landscapes with higher fragmentation and isolation.
5. The opposing effects of habitat loss and fragmentation may result in a nil global effect; therefore it is important to separate their effects when studying populations in fragmented landscapes.     

Reconstruction of the historical range alters niche estimates in an endangered rodent

Defining historical baselines is critical for species conservation. Under the niche reduction hypothesis, species in decline may be restricted disproportionately from parts of their environmental niche. This bias likely has important implications for modeling species’ distributions if only contemporary occurrences (i.e. post‐range reduction) are used, because suitable habitat will be classified as unsuitable. Unfortunately, robust historical occurrence data is rarely available for sensitive species. In this study, we documented historical locations of the endangered, keystone giant kangaroo rat Dipodomys ingens by examining aerial imagery for burrow mounds. These burrow mounds are readily identifiable and distinguishable from other soil disturbances. We found giant kangaroo rat burrows well outside the currently accepted estimate of their historical distribution. Following the niche reduction hypothesis, we found that giant kangaroo rats have been extirpated from the flattest, hottest, driest parts of their range due to agricultural conversion. This reduction in their realized niche led to significant changes between historical and contemporary models of their distribution. We found that giant kangaroo rats may have occupied up to 56% more habitat historically than currently believed. Our results provide new guidance for managers working on restoration and habitat protection for this ecosystem engineer. This study highlights the critical importance of modeling historical distributions using the entire environmental niche once occupied by species of conservation need.     

Possible effects of habitat fragmentation and climate change on the range of forest plant species

Global circulation models predict an increase in mean annual temperature between 2.1 and 4.6 °C by 2080 in the northern temperate zone. The associated changes in the ratio of extinctions and colonizations at the boundaries of species ranges are expected to result in northward range shifts for a lot of species. However, net species colonization at northern boundary ranges, necessary for a northward shift and for range conservation, may be hampered because of habitat fragmentation. We report the results of two forest plant colonization studies in two fragmented landscapes in central Belgium. Almost all forest plant species (85%) had an extremely low success of colonizing spatially segregated new suitable forest habitats after c . 40 years. In a landscape with higher forest connectivity, colonization success was higher but still insufficient to ensure large-scale colonization. Under the hypothesis of net extinction at southern range boundaries, forest plant species dispersal limitation will prevent net colonization at northern range boundaries required for range conservation.     

Dispersal capacity and diet breadth modify the response of wild bees to habitat loss

Habitat loss poses a major threat to biodiversity, and species-specific extinction risks are inextricably linked to life-history characteristics. This relationship is still poorly documented for many functionally important taxa, and at larger continental scales. With data from five replicated field studies from three countries, we examined how species richness of wild bees varies with habitat patch size. We hypothesized that the form of this relationship is affected by body size, degree of host plant specialization and sociality. Across all species, we found a positive species–area slope (z = 0.19), and species traits modified this relationship. Large-bodied generalists had a lower z value than small generalists. Contrary to predictions, small specialists had similar or slightly lower z value compared with large specialists, and small generalists also tended to be more strongly affected by habitat loss as compared with small specialists. Social bees were negatively affected by habitat loss (z = 0.11) irrespective of body size. We conclude that habitat loss leads to clear shifts in the species composition of wild bee communities.