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Habitat destruction, habitat restoration and eigenvector-eigenvalue relations.

According to metapopulation theory, the capacity of a habitat patch network to support the persistence of a species is measured by the metapopulation capacity of the patch network. Mathematically, metapopulation capacity is given by the leading eigenvalue lambda(M) of an appropriately constructed non-negative n x n matrix M, where n is the number of habitat patches. Both habitat destruction (in the sense of destruction of entire patches) and habitat deterioration (in the sense of partial destruction of patches) lower the metapopulation capacity of the patch network. The effect of gradual habitat deterioration is given by the derivative of lambda(M) with respect to patch attributes and may be straightforwardly evaluated by sensitivity analysis. In contrast, destruction of entire patches leads to a rank modification of matrix M, the effect of which on lambda(M) may be derived from eigenvector-eigenvalue relations. Eigenvector-eigenvalue relations have previously been analyzed only for symmetric matrices, which restricts their use in biological applications. In this paper I generalize some of the previous results by deriving eigenvector-eigenvalue relations for general non-symmetric matrices. In addition to the exact eigenvector-eigenvalue relations, I also derive eigenvalue perturbation formulae for rank-one modifications. These results lead to simple and intuitive approximation formulae, which may be used e.g. to assess the contribution of particular habitat patches to the metapopulation capacity of the landscape. The mathematical results presented are not restricted to the metapopulation context, but they should find a number of useful applications in biology, engineering and other applied sciences, where the removal (or addition) of matrix rows and columns often corresponds in a natural manner to decreasing (or increasing) the degrees of freedom of the focal system.

Animals↗

Habitat destruction, fragmentation, and disturbance promote invasion by habitat generalists in a multispecies metapopulation.

Species invasions are extremely common and are vastly outpacing the ability of resource agencies to address each invasion, one species at a time. Management actions that target the whole landscape or ecosystem may provide more cost-effective protection against the establishment of invasive species than a species-by-species approach. To explore what ecosystem-level actions might effectively reduce invasions, we developed a multispecies, multihabitat metapopulation model. We assume that species that successfully establish themselves outside their native range tend to be habitat generalists and that a tradeoff exists between competitive ability and habitat breadth, such that habitat specialists are competitively superior to habitat generalists. In this model, habitat destruction, fragmentation, and short-term disturbances all favor invasion by habitat generalists, despite the inferior competitive abilities of generalist species. Our model results illustrate that providing relatively undisturbed habitat and preventing further habitat degradation and fragmentation can provide a highly cost-effective defense against invasive species.

Competitive Behavior↗

Effects of habitat destruction and resource supplementation in a predator-prey metapopulation model.

We developed a mean field, metapopulation model to study the consequences of habitat destruction on a predator-prey interaction. The model complements and extends earlier work published by Bascompte and Solé (1998, J. theor. Biol.195, 383-393) in that it also permits use of alternative prey (i.e., resource supplementation) by predators. The current model is stable whenever coexistence occurs, whereas the earlier model is not stable over the entire domain of coexistence. More importantly, the current model permits an assessment of the effect of a generalist predator on the trophic interaction. Habitat destruction negatively affects the equilibrium fraction of patches occupied by predators, but the effect is most pronounced for specialists. The effect of habitat destruction on prey coexisting with predators is dependent on the ratio of extinction risk due to predation and prey colonization rate. When this ratio is less than unity, equilibrial prey occupancy of patches declines as habitat destruction increases. When the ratio exceeds one, equilibrial prey occupancy increases even as habitat destruction increases; i.e., prey "escape" from predation is facilitated by habitat loss. Resource supplementation reduces the threshold colonization rate of predators necessary for their regional persistence, and the benefit derived from resource supplementation increases in a nonlinear fashion as habitat destruction increases. We also compared the analytical results to those from a stochastic, spatially explicit simulation model. The simulation model was a discrete time analog of our analytical model, with one exception. Colonization was restricted locally in the simulation, whereas colonization was a global process in the analytical model. After correcting for differences between nominal and effective colonization rates, most of the main conclusions of the two types of models were similar. Some important differences did emerge, however, and we discuss these in relation to the need to develop fully spatially explicit analytical models. Finally, we comment on the implications of our results for community structure and for the conservation of prey species interacting with generalist predators.

Animals↗

Climate change and habitat destruction: a deadly anthropogenic cocktail.

Climate change and habitat destruction are two of the greatest threats to global biodiversity. Lattice models have been used to investigate how hypothetical species with different characteristics respond to habitat loss. The main result shows that a sharp threshold in habitat availability exists below which a species rapidly becomes extinct. Here, a similar modelling approach is taken to establish what determines how species respond to climate change. A similar threshold exists for the rate of climate change as has been observed for habitat loss-patch occupancy remains high up to a critical rate of climate change, beyond which species extinction becomes likely. Habitat specialists, especially those of relatively poor colonizing ability are least able to keep pace with climate change. The interaction between climate change and habitat loss might be disastrous. During climate change, the habitat threshold occurs sooner. Similarly, species suffer more from climate change in a fragmented habitat.

Adaptation, Physiological↗

Effects of habitat destruction in a prey-predator metapopulation model

A mean field, metapopulation model of a predator-prey interaction is developed in order to understand the consequences of habitate destruction at different trophic levels. Such a model allows is to explore different ecological scenarios (donor control vs. top-down control) by changing a single parameter. The response to habitat destruction is qualitatively the same for both predator and prey, although there are interesting differences linked to the trophic position. A similar decrease in the colonization rates affects the two species quite differently. Predators diminish faster than prey, and furthermore, the fraction of occupied sites decreases more sharply as colonization rates are lowered, i.e. there is a nonlinear relationship between regional abundance and colonization rate. There is a well-defined threshold in the colonization rate below which the predator becomes extinct. Thus, dispersal rate is critical for predicting the consequences of habitat destruction. Finally, these results are compared with the behavior of a spatially explicit simulation. The only difference between the analytical model and the simulation is that colonization is no longer a global phenomenon in the latter, but it is a local process, constrained to the nearest patches. The bulk of the results are similar to the mean field behavior, and we comment on some differences related with non-homogeneity and real space. Some general implications for conservation biology and biological control are outlined. Copyright 1998 Academic Press

Journal Article↗

Habitat Destruction and Competitive Coexistence in Spatially Explicit Models with Local Interactions.

We investigate the competition-colonization trade-off in a two-species competition model in various heterogeneous environments. The competitiveness of species is hierarchically ordered and the dynamics of the model are described by spatially local interactions of individuals. The size of the local neighborhood may depend on the species. The habitat consists of inhabitable and permanently destroyed sites. It was previously observed (Nee & May, 1992; Tilman et al., 1994, 1997) that destruction of habitat may aid the inferior species and cause the superior species to go extinct. These conclusions were based on an n-species model in which the spatial arrangement of destroyed sites was ignored. We examine the effects of different spatial arrangements of destroyed sites on survival and coexistence of the two species. We conclude that the spatial arrangement plays an important role and can influence the outcome qualitatively. The key quantity for predicting the outcome of habitat destruction on species survival is the relationship between the mean dispersal distance and the mean distance between inhabitable and destroyed sites. We contrast the hierarchical model with a version of the Lotka-Volterra model of interspecific competition and show that habitat destruction can alter the dominance relationship between species by reducing intraspecific competition.Copyright 1998 Academic Press

Journal Article↗

Habitat Deterioration, Habitat Destruction, and Metapopulation Persistence in a Heterogenous Landscape

Levins's unstructured metapopulation model predicts that the equilibrium fraction of empty habitat patches is a constant function of the fraction h of suitable patches in the landscape and that this constant equals the threshold value for metapopulation persistence. Levins's model thus suggests that the minimum amount of suitable habitat necessary for metapopulation persistence can be estimated from the fraction of empty patches at steady state. In this paper we construct several more realistic structured metapopulation models that include variation in patch quality and the rescue effect. These models predict both positive and negative correlations between the fractions of suitable patches and empty patches. The type of correlation depends in an intricate manner on the strength of the rescue effect and on the quality distribution of the patches to be destroyed. Empty patches can be considered as the resource limiting metapopulation growth. Our results demonstrate that the correlation between the fractions of suitable patches and empty patches is positive if and only if the average value of the resource decreases as the number of patches increases. Copyright 1997 Academic Press. Copyright 1997 Academic Press

Journal Article↗

Habitat destruction in mutualistic metacommunities.

We investigate a mutualistic metacommunity where the strength of the mutualistic interaction between species is measured by the extent to which the presence of one species on a patch either reduces the extinction rate of the others present on the same patch or increases their ability to colonize other patches. In both cases, a strong enough mutualism enables all species to persist at habitat densities where they would all be extinct in the absence of the interaction. However, a mutualistic interaction that enhances colonization enables the species to persist at lower habitat density than one that suppresses extinction. All species abruptly go extinct (catastrophe) when the habitat density is decreased infinitesimally below a critical value. A comparison of the mean field or spatially implicit case with unrestricted dispersal and colonization to all patches in the system with a spatially explicit case where dispersal is restricted to the immediate neighbours of the original patch leads to the intriguing conclusion that restricted dispersal can be favourable for species that have a beneficial effect on each other when habitat conditions are adverse. When the mutualistic interaction is strong enough, the extinction threshold or critical amount of habitat required for the persistence of all species is lower when the dispersal is locally restricted than when unrestricted ! The persistence advantage for all species created by the mutualistic interaction increases substantially with the number of species in the metacommunity, as does the advantage for restricted dispersal over global dispersal.

Adaptation, Physiological↗

Habitat destruction, environmental catastrophes, and metapopulation extinction.

The extinction process of fragmented populations, characterized by a small number of conspecifics inhabiting each patch, is heavily affected by natural and human disturbance. To evaluate the risk of extinction we consider a network of identical patches connected by passive or active dispersal and hosting a finite, discrete number of individuals. We discuss three types of disturbance affecting the metapopulation: permanent loss of habitat patches, erosion of existing patches, and random catastrophes that wipe out the entire population of a patch. Starting from an infinite-dimensional Markov model that fully accounts for demographic stochasticity, we reduce it to finite dimension via moment closure with negative-binomial approximation. The compact models obtained in this way account for the dynamics of the fraction of empty patches, the average number of individuals in occupied patches, and the variance of their distribution. After comparing the performance of these compact models with that of the infinite-dimensional model in the case of no disturbances, we then proceed to computing persistence-extinction boundaries as bifurcation lines of the compact models in the space of demographic and disturbance parameters. We consider bifurcations with respect to demographic and environmental parameters and contrast our results with those of previous theories. We find out that environmental catastrophes increase the risk of extinction for both frequent and infrequent dispersers, while the random loss of patches has a much larger influence on frequent dispersers. This influence can be counterbalanced by active dispersal. Local erosion of habitat fragments has a larger influence on infrequent than on frequent dispersers. We finally discuss the important synergistic effects of disturbances acting simultaneously.

Disasters↗

Habitat destruction in a simple predator-prey patch model: how predators enhance prey persistence and abundance.

We model a metapopulation of predator-prey patches using both spatially implicit or mean-field (MF) and spatially explicit (SE) approaches. We show that in the MF model there are parameter regimes for which prey cannot persist in the absence of predators, but can in their presence. In addition, there are parameter regimes for which prey may persist in isolation, but the presence of predators will increase prey patch density. Predators may thus enhance prey persistence and overall abundance. The key mechanism responsible for this effect is the occurrence of prey dispersal from patches that are occupied by both prey and predators. In addition, these patches should be either long-lived, such as that occurs when predators keep prey from overexploiting its local resource, or the presence of a predator on a patch should significantly enhance the prey dispersal out of that patch. In the SE approach these positive effects of predators on prey persistence and abundance occur for even larger parameter ranges than in the MF model. Prey dispersal from predator-prey patches may thus be important for persistence of both species as a community, independent of the modeling framework studied. Comparison of the MF and SE approaches shows that local dispersal constraints can have the edge over global dispersal for the persistence of the metapopulation in regimes where the two species have a beneficial effect on each other. In general, our model provides an example of feedback in multiple-species metapopulations that can make the implementation of conservation schemes based on single-species arguments very risky.

Animals↗

Challenges to introducing and managing disturbance regimes for Holocarpha macradenia, an endangered annual grassland forb.

Introducing rare plants to new sites for conservation to offset effects of habitat destruction requires detailed knowledge of habitat requirements, plant demography, and management needs. We conducted a factorial experiment replicated at three coastal prairie sites to test the effects of clipping frequency and litter accumulation on seed germination, seedling survival, reproduction, and seedling recruitment of introduced populations of the endangered, tall-stature, annual forb, Holocarpha macradenia (DC.) E. Greene. Clipping favored H. macradenia, primarily by enhancing seed germination and flower production. Litter accumulation had no effect on seed germination, even after 5 years of treatments. Seedling recruitment was highly site specific with large numbers of recruits recorded at only one of three sites. Although recruitment of seedlings was higher in clipped plots for 2-3 years, by 4-5 years after introduction very few seedlings survived to reproduction in any treatment. We attribute this result to a combination of poor habitat quality, small population size, and lack of a seed bank. We were unsuccessful in introducing this relatively well-studied species of concern to apparently suitable habitat at multiple sites in multiple years, which suggests that translocating rare plant populations to mitigate for habitat destruction is an expensive and highly uncertain endeavor.

Asteraceae↗

The competitive dynamics of metapopulations subject to the Allee-like effect.

It is well recognized that individuals of many species can benefit from the presence of conspecifics, a concept broadly referred to as the Allee effect. At the metapopulation level, there is an analogous but essentially different phenomenon called the Allee-like effect that leads to metapopulation extinction thresholds at low habitat occupancy. But so far not adequate attention has been paid to this phenomenon. In this paper, the Allee-like effect is introduced into a metapopulation model of one species and also that of a three-state two-species competitive system. Phase plane analysis is used to investigate the dynamics of these models. We demonstrate that the Allee-like effect alone could lead to multiple stable states in three-state two-species competitive systems at the metapopulation level, and the number of stable states decrease as the Allee-like effect becomes more severe. Severe Allee-like effects may make coexistence impossible and may even lead to the extinction of both species even if their initial habitat occupancies are high and suitable habitats are enough. It is especially noticeable that depending on their initial conditions one species may exclude the other one that subjects to a weaker Allee-like effect than the former, while the second species always excludes the first one when both species are assumed to be in the absence of the Allee-like effect. We also investigate the habitat destructive effect on the Allee-like system mentioned above. Research indicates that the existence of the Allee-like effect makes a metapopulation more susceptible to habitat destruction. All in all, the Allee-like effect is probably a destabilizing factor that, together with habitat destruction, would affect the continuous existence of species. These conclusions may have important implications for conservation and metacommunity organization.

China↗

Increased pollen flow counteracts fragmentation in a tropical dry forest: an example from Swietenia humilis Zuccarini.

Habitat destruction and the resultant fragmentation of the remaining forest are a common phenomenon in the tropics. Most investigations emphasize the potential dangers of fragmentation in isolating patches of forest and exposing populations to loss of species diversity through founder effects, genetic drift, inbreeding, and restricted gene flow. However, a limited number of studies have shown that gene flow may be extensive in tropical trees, suggesting that it may occur between forest fragments and also "isolated" remnant trees. There is an urgent need to quantify pollen flow within and between forest fragments to test the veracity of such views and determine the genetic value of such fragments for in situ conservation. Microsatellite markers are used to genotype individuals of Swietenia humilis from a highly fragmented forest mosaic to directly quantify pollen-mediated gene flow. Distances of pollen flow more than 10 times greater than previously reported were detected. Our results show that some tropical angiosperm tree species may be much more adaptable and resilient to habitat destruction and fragmentation than previously considered. The description of many remnant trees as isolated or "living dead" may be more a conditioning of human perception than a true reflection of their potential conservation value.

Ecology↗

Influence of volcanic activity on the population genetic structure of Hawaiian Tetragnatha spiders: fragmentation, rapid population growth and the potential for accelerated evolution.

Volcanic activity on the island of Hawaii results in a cyclical pattern of habitat destruction and fragmentation by lava, followed by habitat regeneration on newly formed substrates. While this pattern has been hypothesized to promote the diversification of Hawaiian lineages, there have been few attempts to link geological processes to measurable changes in population structure. We investigated the genetic structure of three species of Hawaiian spiders in forests fragmented by a 150-year-old lava flow on Mauna Loa Volcano, island of Hawaii: Tetragnatha quasimodo (forest and lava flow generalist), T. anuenue and T. brevignatha (forest specialists). To estimate fragmentation effects on population subdivision in each species, we examined variation in mitochondrial and nuclear genomes (DNA sequences and allozymes, respectively). Population subdivision was higher for forest specialists than for the generalist in fragments separated by lava. Patterns of mtDNA sequence evolution also revealed that forest specialists have undergone rapid expansion, while the generalist has experienced more gradual population growth. Results confirm that patterns of neutral genetic variation reflect patterns of volcanic activity in some Tetragnatha species. Our study further suggests that population subdivision and expansion can occur across small spatial and temporal scales, which may facilitate the rapid spread of new character states, leading to speciation as hypothesized by H. L. Carson 30 years ago.

Animals↗

Quantitative evaluation of the petal shape variation in Primula sieboldii caused by breeding process in the last 300 years.

Primula sieboldii: (E. Morren) has been a popular garden plant at least since the Edo period, about 300 years ago. We compared petal form between cultivars and wild populations in order to characterise the changes that have occurred during domestication. The comparison was made using EF-PCA analysis, which describes overall petal shape mathematically by transforming petal contour coordinates into elliptic Fourier descriptors; it subsequently summarises these descriptors by principal component analysis (PCA). Rearing cultivars in a common-garden experiment identified the PCs with a substantial genetic element. A clear heritable component was detected for the PCs characterising symmetrical variation in flower shape, but not the asymmetrical variation. Wild populations of this species have become endangered owing to habitat destruction by human activity, and many lowland floodplain habitats have been lost. Variation within the remaining wild populations was significantly lower than in the cultivars for PC1 (aspect ratio), PC3 (curvature of proximal and distal parts) and petal area; but not for PC2 (depth of head notch) and PC4 (position of the centre of gravity). The shifts in petal form from the wild populations to the cultivars parallel those seen in other crop-types following domestication, including an increase in size and diversity of forms: cultivars have shallower head notches, more fan-shaped petals and larger petals than do wild P. sieboldii.

Analysis of Variance↗

Quantifying remaining forested habitat within the historic distribution of the cotton-top tamarin (Saguinus oedipus) in Colombia: Implications for long-term conservation.

Landsat Thematic Mapper (TM) data were used to classify forested areas within the historic distribution of the endangered cotton-top tamarin (Saguinus oedipus) in Colombia. This species continues to be threatened by habitat destruction, and we observed a 31% decrease of the forested habitat within the tamarins' historic distribution in 1990-2000. It is estimated that since the establishment of some protected areas for cotton-top tamarins and other native Colombian wildlife (Parque Nacional Natural Paramillo, Santuario de Fauna y Flora Los Colorados, and Reserva Forestal de Montes de Maria), almost 43% of the original forested area has been lost. Given that the human population of Colombia increases annually by 1.6% [Patel, 2002], it is important to target specific areas for tamarin protection while creating mitigation strategies to compensate for economic growth. The results of this study provide valuable information to assist in the long-term development of effective conservation strategies for this endangered primate.

Animals↗

The past impact of livestock husbandry on dispersal of plant seeds in the landscape of Denmark.

The recent decline in species richness in (semi)-natural habitats in northern Europe has largely been attributable to habitat destruction, and to subsequent limitation in seed dispersal among fragments. However, some habitat types were probably split up already in the historical landscape, but the segregated parts were probably not isolated to the present degree. This paper seeks evidence for livestock as vectors for propagules at 3 spatial scales in the past cultural landscape. Three main scales at which livestock acted as seed dispersers are important: free movement in the landscape (1-10 km), driving animals to mast feeding or to manors (10-50 km), and the export of living animals (hundreds of km). The emerging picture is for most plant species a dramatically decreased chance of dispersal in the modern landscape. The consequence is probably decreasing species richness in (semi)-natural plant communities, such as pasture, meadow, and heathland.

Animal Husbandry↗