Sequencing stepping stones: a raster-based GIS model for routing a connectivity corridor through a fragmented landscape

Landscape structure, habitat fragmentation, and the ecology of insects

Biodiversity is severely threatened by habitat destruction. As a consequence of habitat destruction, the remaining habitat becomes more fragmented. This results in time-lagged population extirpations in remaining fragments when these are too small to support populations in the long term. If these time-lagged effects are ignored, the long-term impacts of habitat loss and fragmentation will be underestimated. We quantified the magnitude of time-lagged effects of habitat fragmentation for 157 nonvolant terrestrial mammal species in Madagascar, one of the biodiversity hotspots with the highest rates of habitat loss and fragmentation. We refined species' geographic ranges based on habitat preferences and elevation limits and then estimated which habitat fragments were too small to support a population for at least 100 years given stochastic population fluctuations. We also evaluated whether time-lagged effects would change the threat status of species according to the International Union for the Conservation of Nature (IUCN) Red List assessment framework. We used allometric relationships to obtain the population parameters required to simulate the population dynamics of each species, and we quantified the consequences of uncertainty in these parameter estimates by repeating the analyses with a range of plausible parameter values. Based on the median outcomes, we found that for 34 species (22% of the 157 species) at least 10% of their current habitat contained unviable populations. Eight species (5%) had a higher threat status when accounting for time-lagged effects. Based on 0.95-quantile values, following a precautionary principle, for 108 species (69%) at least 10% of their habitat contained unviable populations, and 51 species (32%) had a higher threat status. Our results highlight the need to preserve continuous habitat and improve connectivity between habitat fragments. Moreover, our findings may help to identify species for which time-lagged effects are most severe and which may thus benefit the most from conservation actions.

Studies examining the effects of anthropogenic habitat fragmentation on both neutral and adaptive genetic variability are still scarce. We compared tadpole fitness-related traits (viz. survival probability and body size) among populations of the common frog (Rana temporaria) from fragmented (F) and continuous (C) habitats that differed significantly in population sizes (C > F) and genetic diversity (C > F) in neutral genetic markers. Using data from common garden experiments, we found a significant positive relationship between the mean values of the fitness related traits and the amount of microsatellite variation in a given population. While genetic differentiation in neutral marker loci (F(ST)) tended to be more pronounced in the fragmented than in the continuous habitat, genetic differentiation in quantitative traits (Q(ST)) exceeded that in neutral marker traits in the continuous habitat (i.e. Q(ST) > F(ST)), but not in the fragmented habitat (i.e. Q(ST) approximately F(ST)). These results suggest that the impact of random genetic drift relative to natural selection was higher in the fragmented landscape where populations were small, and had lower genetic diversity and fitness as compared to populations in the more continuous landscape. The findings highlight the potential importance of habitat fragmentation in impairing future adaptive potential of natural populations.

It is important to understand the relative effects of landscape habitat loss, habitat fragmentation, and matrix quality on biodiversity, so that potential management options can be appropriately ranked. However, their effects and relative importance may change with the size of the landscape considered because the multiple (and potentially conflicting) ecological processes that are influenced by landscape structure occur at different spatial scales (e.g. dispersal, predation, foraging). We estimated the relative effects of habitat loss, habitat fragmentation, and matrix quality (measured as the amount of forest, the proportion of forest area contained in large core forests, and the density of roads respectively) on fragmentation‐sensitive forest birds in southern Ontario, Canada using a range of landscape sizes (0.8–310 km 2 ). We used three complementary statistical approaches to estimate relative effects of these correlated landscape factors – 1) multiple regression, 2) information theoretic (AIC) estimates of the most parsimonious model, and 3) multi‐model inference to average effects across all supported models. We controlled for spatial autocorrelation, local habitat, roadside sampling bias, time of day, season, habitat heterogeneity, and the interaction between the effects of habitat amount and fragmentation. We found that relative effects of habitat amount and fragmentation were scale dependent; habitat amount had a consistently positive effect that was consistent over more than two orders of magnitude in landscape area (~1–300 km 2 ). In contrast, the effects of habitat fragmentation depended on the size of the landscape considered. Indeed, for veery Catharus fuscescens , habitat fragmentation had positive effects at one scale and negative effects at another. The effects of matrix quality were generally weak and changed little with scale. For the number of fragmentation sensitive species and the presence of veery, habitat amount was most important in large landscapes and habitat fragmentation in small landscapes but for the presence of ovenbird Seiurus aurocapilla , habitat amount was most important at all scales.

I reviewed and reconciled predictions of four models on the effect of habitat fragmentation on the population extinction threshold, and I compared these predictions to results from empirical studies. All four models predict that habitat fragmentation can, under some conditions, increase the extinction threshold such that, in more fragmented landscapes, more habitat is required for population persistence. However, empirical studies have shown both positive and negative effects of habitat fragmentation on population abundance and distribution with about equal frequency, suggesting that the models lack some important process(es). The two colonization–extinction (CE) models predict that fragmentation can increase the extinction threshold by up to 60–80%; i.e., the amount of habitat required for persistence can shift from <5% of the landscape to >80% of the landscape, with a shift from completely clumped to completely fragmented habitat. The other two models (birth–immigration–death–emigration, or BIDE models) predict much smaller potential effects of fragmentation on the extinction threshold, of no more than a 10–20% shift in the amount of habitat required for persistence. This difference has important implications for conservation. If fragmentation can have a large effect on the extinction threshold, then alteration of habitat pattern (independent of habitat amount) can be an effective tool for conservation. On the other hand, if the effects of fragmentation on the extinction threshold are small, then this is a limited option. I suggest that the difference in model predictions results from differences in the mechanisms by which the models produce the extinction threshold. In the CE models, the threshold occurs by an assumed reduction in colonization rate with decreasing habitat amount. In the BIDE models, loss of habitat is assumed to increase the proportion of the population that spends time in the matrix, where reproduction is not possible and the mortality rate is assumed to be higher (than in breeding habitat). Habitat loss therefore decreases the overall reproduction rate and increases the overall mortality rate on the landscape. I hypothesize that this imposes a constraint on the potential for habitat fragmentation to mitigate effects of habitat loss in BIDE models. To date, empirical studies of the independent effects of habitat loss and fragmentation suggest that habitat loss has a much larger effect than habitat fragmentation on the distribution and abundance of birds, supporting the BIDE model prediction, at least for this taxon.

Effects of small-scale habitat fragmentation, habitat corridors and mainland dispersal on soil decomposer organisms

In fragmented landscapes the relationship between the probability of occurrence of single species and the amount of suitable habitat is usually not proportional, with a threshold habitat level below which the population becomes extinct. Ecological theory predicts that, although the reduction in species’ occurrence probabilities (and eventually the extinction threshold) is a direct consequence of habitat loss, habitat fragmentation might reduce species’ occurrence probabilities and affect the location of this threshold by reducing its predicted occurrence to lower levels of habitat amount. However, little is known about the validity of this extinction threshold hypothesis. Here, we performed analyses on the relationships between the probability of occurrence of eight tree species and the availability of forest habitat for two different empirical scenarios of low and moderate to high fragmentation. We partitioned the effects of habitat amount versus fragmentation by using two metrics: (1) the percentage of forest cover, and (2) the proportion of this percentage occurring in the largest forest patch. We find that, although decreasing forest cover had negative effects on the occurrence of tree species irrespective of fragmentation levels, forest fragmentation significantly modified the response pattern in six tree species, although only one species confirmed the extinction threshold hypothesis, which we interpret as a consequence of high degree of forest specialism and low dispersal ability. For most species, fragmentation either had positive effects or did not affect significantly the species’ probability of occurrence. This indicates that the effects of habitat fragmentation on tree species are weak relative to the effects of habitat amount, which is the main determinant of the reduction in species’ occurrence probabilities, and eventually species extinction, in fragmented landscapes.

Fragmentation of natural habitats generally has negative effects on the reproductive success of many plant species; however, little is known about epiphytic plants. We assessed the impact of forest fragmentation on plant–pollinator interactions and female reproductive success in two epiphytic Tillandsia species with contrasting life history strategies (polycarpic and monocarpic) in Chamela, Jalisco, Mexico, over three consecutive years. Hummingbirds were the major pollinators of both species and pollinator visitation rates were similar between habitat conditions. In contrast, the composition and frequency of floral visitors significantly varied between habitat conditions in polycarpic and self-incompatible T. intermedia but not in monocarpic self-compatible T. makoyana. There were no differences between continuous and fragmented habitats in fruit set in either species, but T. makoyana had a lower seed set in fragmented than in continuous forests. In contrast, T. intermedia had similar seed set in both forest conditions. These results indicate that pollinators were effective under both fragmented and continuous habitats, possibly because the major pollinators are hummingbird species capable of moving across open spaces and human-modified habitats. However, the lower seed set of T. makoyana under fragmented conditions suggests that the amount and quality of pollen deposited onto stigmas may differ between habitat conditions. Alternatively, changes in resource availability may also cause reductions in seed production in fragmented habitats. This study adds to the limited information on the effects of habitat fragmentation on the reproductive success of epiphytic plants, showing that even related congeneric species may exhibit different responses to human disturbance. Plant reproductive systems, along with changes in pollinator communities associated with habitat fragmentation, may have yet undocumented consequences on gene flow, levels of inbreeding and progeny quality of dry forest tillandsias.

Habitat loss and fragmentation threaten the long-term viability of innumerable species of plants and animals. At the same time, habitat fragmentation may impose strong natural selection and lead to evolution of life histories with possible consequences for demographic dynamics. The Baltic populations of the Glanville fritillary butterfly (Melitaea cinxia) inhabit regions with highly fragmented habitat (networks of small dry meadows) as well as regions with extensive continuous habitat (calcareous alvar grasslands). Here, we report the results of common garden studies on butterflies originating from two highly fragmented landscapes (FL) in Finland and Sweden and from two continuous landscapes (CL) in Sweden and Estonia, conducted in a large outdoor cage (32 by 26 m) and in the laboratory. We investigated a comprehensive set of 51 life-history traits, including measures of larval growth and development, flight performance, and adult reproductive behavior. Seventeen of the 51 traits showed a significant difference between fragmented versus CL. Most notably, the growth rate of postdiapause larvae and several measures of flight capacity, including flight metabolic rate, were higher in butterflies from fragmented than CL. Females from CL had shorter intervals between consecutive egg clutches and somewhat higher life-time egg production, but shorter longevity, than females from FL. These results are likely to reflect the constant opportunities for oviposition in females living in continuous habitats, while the more dispersive females from FL allocate more resources to dispersal capacity at the cost of egg maturation rate. This study supports theoretical predictions about small population sizes and high rate of population turnover in fragmented habitats selecting for increased rate of dispersal, but the results also indicate that many other life-history traits apart from dispersal are affected by the degree of habitat fragmentation.

The effects of habitat fragmentation on plant–animal interactions may emerge at different spatial scales, depending on the species-specific perception response of the interacting animals. Furthermore, changes in habitat cover and configuration commonly occur simultaneously, hampering efforts to understand and mitigate the impact of fragmentation on these biotic interactions. In order to account for the relative influence of habitat loss and fragmentation on plant–animal interactions, we quantified habitat structure in sixteen sectors (nested circular areas of 100 and 200 m radii) in four different localities (four sectors per locality) across the Cantabrian Range in NW Spain. In the center of each 100 m radius sector, we measured the magnitude of two ecologically opposite (mutualistic vs antagonistic) interactions in individual holly trees Ilex aquifolium which strongly determine the regeneration process in this plant species: frugivory by birds and seed predation by rodents. We found that habitat fragmentation, though not habitat loss, affected the magnitude of both plant–animal interactions. However, these effects were conditioned by the strong differences in spatial heterogeneity in habitat structure between localities. In fact, the effect of habitat fragmentation on both plant–animal interactions disappeared when the locality in which sectors were sited was taken into account. This study highlights that 1) habitat spatial configuration, far from being a negligible component of habitat structure, is in fact able to influence key ecological processes such as plant–animal interactions, and 2) the potential spatial and structural complexity of localities makes a regional approach (i.e. that involving many localities) indispensable in the quest for comprehensive understanding of the effects of habitat structure on biodiversity in real-world fragmented landscapes.

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.

Habitat loss and fragmentation has long been considered the primary cause for biodiversity loss and ecosystem degradation worldwide, and is a key research topic in landscape ecology (Wu 2013). Habitat fragmentation often refers to the reduction of continuous tracts of habitat to smaller, spatially distinct remnant patches, and habitat loss typically occurs concurrently with habitat fragmentation (Collinge 2009). Although some habitats are naturally patchy in terms of abiotic and biotic conditions (Wu and Loucks 1995), human actions have profoundly fragmented landscapes across the word (Haddad et al. 2015), altering the quality and connectivity of habitats. Therefore, understanding the causes and consequences of habitat fragmentation is critical to preserving biodiversity and ecosystem functioning. From May 4th to 10th, 2015, an International Workshop on Habitat Fragmentation and Biodiversity Conservation, held at the Thousand Island Lake, Zhejiang, China, discussed threats to biodiversity in fragmented landscapes and how fragmentation research can identify and help mitigate these threats. To meet these challenges, the Workshop had three goals. The first was to synthesize key findings in fragmentation science. Second was to identify important remaining research questions concerning the relationships between habitat fragmentation, biodiversity, and ecosystem functioning at local, regional, and global scales. Finally, we examined the unique roles of field-based fragmentation experiments in addressing these questions. The Workshop’s findings are relevant to the broader ecological community, and we present them here to stimulate research that will advance landscape ecology and conservation biology.

Simulated effects of habitat loss and fragmentation on a solitary mustelid predator

Crested Tit Parus cristatus young from first broods dispersed 1 week later if they were born in small isolated pine plots (‘habitat fragments’) compared with individuals in a large pine forest (‘continuous habitat’). This delay in dispersal was caused by an extended period between fledging and dispersal. In second broods, the delay was even longer due to the interbrood interval being 9 days longer in habitat fragments. As nestlings in habitat fragments had a lower body‐mass, and age at dispersal was negatively correlated with nestling body‐mass within each nest, the postponed dispersal from fragments might be explained partly by a lower body‐mass. Alternatively, postponed dispersal from fragments could result from a barrier effect caused by reluctance to cross inhospitable habitat.Immigration by young from first broods into habitat fragments was delayed by approximately 3 weeks, and proportionally more second brood emigrants were recovered in this type of habitat. These results are in agreement with the hypothesis that fragments are second‐choice habitat. Early immigrants into continuous habitat had a higher probability of settlement in winter flocks compared with late ones, independent of condition or age. Therefore, Crested Tits born in habitat fragments probably have a lower chance of settling in first‐choice habitat.

The effects of habitat destruction and fragmentation on survival rate and population size were tested in an experimental model system consisting of seven enclosed populations of root voles, Microtus oeconomus. The study comprised two consecutive parts. First, the effect of habitat fragmentation was investigated by comparing survival in three populations inhabiting continuous habitat with that in for populations inhabiting fragmented habitat. Second, the continuous habitats were experimentally fragmented by destroying 50% of the area inhabited by the populations, producing the same configuration as in the fragmented habitats. These extensive habitat manipulations were expected to affect survival rates due to changes in the spatiosocial structuring of the populations. Throughout the study, populations were monitored by an intensive live—trapping program. Cormack—Jolly—Sebver methodology was used for statistical modelling and estimation of survival rates. The seven experimental populations differed considerably in population dynamics: some grew, while others remained remarkably stable throughout the experiment. In contrast to expectations, neith population size nor survival rates seemed to be affected by the experimental treatments. survival rate seemed to play a minor role in determining population size. Emigration rates, which also seemed to have little direct demographic importance, were considerably higher in fragmented than in continuous habitats during the first (pre—destruction) part of the study. Surprisingly, this difference persisted after habitat destruction, when all populations experienced the same habitat configuration. We conclude that survival rates in these root vole populations were not dependent on major alterations of spatiosocial structure, possibly owing to adaptation to natural disturbance regimes.