Stephen F. Matter

Landscape Ecology of Mammals: Relationships between Density and Patch Size

A much discussed issue in landscape ecology is how processes that operate within spatially subdivided subpopulations scale-up to create a larger, landscape-level dynamic. A first step in answering this question is to ask to what degree subpopulations within a landscape vary in performance. Here we test the null hypothesis that densities of mammalian populations are constant over patches of varied size, i.e., that performance, as estimated via density, does not covary with patch area. Using a composite database from published studies, we found that densities of 20 of 32 species did not vary with patch area, while five showed increasing and seven decreasing density-area relationships. Studies reporting significant density-area relationships tended to include a greater number of patches of a greater range of sizes than those that reported no relationship, suggesting that statistical power may be an issue. Landscapes comprised of smaller, less-isolated patches tended to have negative density-area relationships and landscapes with larger, more isolated patches tended to have positive density-area relationships. Our results suggest that no consistent density-area relationship operates over all systems of patches. Instead, the patterns appear to be scale-dependent : frequent movement of individuals in the process of selecting habitats (patches) over smaller-scaled landscapes produced negative density-area relationships; movement of individuals among more isolated patches appeared to involve larger- and longer-scale population processes involving colonization and extinction and positive density-area relationships. Despite the fact that patches represent a central focus in landscape ecology, they appear to be a construct of human convenience rather than biological entities with a set number and kind of processes.

Among- and within-patch components of genetic diversity respond at different rates to habitat fragmentation: an empirical demonstration

Habitat fragmentation is a ubiquitous by-product of human activities that can alter the genetic structure of natural populations, with potentially deleterious effects on population persistence and evolutionary potential. When habitat fragmentation results in the subdivision of a population, random genetic drift then leads to the erosion of genetic diversity from within the resulting subpopulations and greater genetic divergence among them. Theoretical and simulation analyses predict that these two main genetic effects of fragmentation, greater differentiation among resulting subpopulations and reduced genetic diversity within them, will proceed at very different rates. Despite important implications for the interpretation of genetic data from fragmented populations, empirical evidence for this phenomenon has been lacking. In this analysis, we carry out an empirical study in populations of an alpine meadow-dwelling butterfly, which have become fragmented by increasing forest cover over five decades. We show that genetic differentiation among subpopulations (GST) is most highly correlated with contemporary forest cover, while genetic diversity within subpopulations (expected heterozygosity) is better correlated with the spatial pattern of forest cover 40 years in the past. Thus, where habitat fragmentation has occurred in recent decades, genetic differentiation among subpopulations can be near equilibrium while contemporary measures of within subpopulation diversity may substantially overestimate the equilibrium values that will eventually be attained.

An experimental examination of the effects of habitat quality on the dispersal and local abundance of the butterfly Parnassius smintheus

Abstract 1. Nectar flower abundance was manipulated through flower removal, and sex ratio was manipulated by moving individual butterflies within a series of nine alpine meadows. The movement and abundance of the butterfly Parnassius smintheus in the meadows were monitored using mark–release–recapture methods.

Interpatch movement of the red milkweed beetle, Tetraopes tetraophthalmus : individual responses to patch size and isolation

Individual movement patterns and the effects of host plant patch size and isolation on patch occupancy were examined for red milkweed beetles, Tetraopes tetraophthalmus, residing in a heterogeneous landscape. Male beetles were found to move both more often and farther between host plant patches than female beetles, and this difference affected the patterns of patch occupancy observed. Overall, unoccupied milkweed patches were smaller and more isolated than patches occupied by beetles. Patches uninhabited by females tended to be more isolated, but not necessarily smaller, than patches with female beetles, indicating that females may be affected more by patch isolation than patch size. Presence of male beetles on patches showed a stronger response to patch size than to patch isolation. Differences in movement between males and females illustrate the need for demographically based dispersal data. Comparisons of Tetraopes interpatch movement patterns between landscapes composed of patches of different size revealed that landscapes with overall smaller patches may have greater rates of interpatch movement.

Population density and area: the role of between- and within-patch processes

Abstract The relationship between population density and the size of host plant patches was investigated for the red milkweed beetle Tetraopestetraophthalmus inhabiting unmanipulated patches of Asclepias syriaca. The resource concentration hypothesis proposes that density-area patterns, specifically that of increasing herbivore density with patch size, are primarily a function of movement between host plant patches. This research investigated the degree to which movement accounted for density-area patterns. Poisson regression analysis of beetle abundance versus milkweed patch size revealed that beetle density tended to increase with patch size. The pattern of density and patch size resulted from local reproduction and residence time. The density of emerging beetles tended to increase with patch size while emigration rates were unrelated to patch size. Immigration rates were constant with patch size for male beetles, and decreased with patch size for female beetles. Net flux of beetles (immigration – emigration) did not vary with patch size for male beetles and decreased with patch size for female beetles. Comparisons are made between this system and previously studied systems where movement plays a significant role in forming density area patterns. Additionally, several hypotheses are presented which may account for greater in situ recruitment and residence time in large patches.

Edge avoidance and movement of the butterfly Parnassius smintheus in matrix and non-matrix habitat

We experimentally examined edge effects and movement patterns of the butterfly Parnassius smintheus in two habitat types, its preferred meadow habitat, and intervening forest matrix habitat. We followed the movement of 46 butterflies released at either 5 or 20m from a forest edge in either forest or meadow habitat. In contrast to theoretical predictions, we found that butterflies flew less frequently, shorter distances, and at lower rates in matrix habitat than they did in meadow habitat. Distance from the edge had little effect on these aspects of movement. Flight was strongly influenced by light levels with butterflies flying more readily at higher light levels. Light levels were higher in meadows than in forest explaining much of the difference in movement patterns. Turning angles showed that butterflies flying in meadow habitat avoided forest edges and that this effect extended nearly 25 m into meadows. Analysis of net displacement from the forest edge reinforced this result and showed that there may be attraction to the meadow for butterflies flying within forest.

Reproductive asynchrony in natural butterfly populations and its consequences for female matelessness.

1. Reproductive asynchrony, where individuals in a population are short-lived relative to the population-level reproductive period, has been identified recently as a theoretical mechanism of the Allee effect that could operate in diverse plant and insect species. The degree to which this effect impinges on the growth potential of natural populations is not yet well understood. 2. Building on previous models of reproductive timing, we develop a general framework that allows a detailed, quantitative examination of the reproductive potential lost to asynchrony in small natural populations. 3. Our framework includes a range of biologically plausible submodels that allow details of mating biology of different species to be incorporated into the basic reproductive timing model. 4. We tailor the parameter estimation methods of the full model (basic model plus mating biology submodels) to take full advantage of data from detailed field studies of two species of Parnassius butterflies whose mating status may be assessed easily in the field. 5. We demonstrate that for both species, a substantial portion of the female population (6.5-18.6%) is expected to die unmated. These analyses provide the first direct, quantitative evidence of female reproductive failure due to asynchrony in small natural populations, and suggest that reproductive asynchrony exerts a strong and largely unappreciated influence on the population dynamics of these butterflies and other species with similarly asynchronous reproductive phenology.

MIGRATION AND SURVIVAL OF PARNASSIUS SMINTHEUS: DETECTING EFFECTS OF HABITAT FOR INDIVIDUAL BUTTERFLIES

We examined the migration and survival of the butterfly Parnassius smintheus in a heterogeneous landscape consisting of 21 habitat patches imbedded in a matrix of meadow and forest habitat. We modified an existing mark–release–recapture model to account for multiple habitat types and fit the model to data for 839 and 873 individuals in two separate years. Migration was infrequent with only 24 and 27 observed movements between patches in each year. Daily within-patch survival was moderate (>0.90) and did not vary greatly with patch isolation. Estimated mortality during migration was low, but increased markedly for isolated populations. Despite the limited data, the model showed that forest matrix habitat reduced migration distance to a greater degree than did meadow habitat, indicating that the effective isolation of populations depends on both the habitat type and the distance between populations. This result concurs with previous investigations of these data, demonstrating the utility of the model even w...

Controlled experiments of habitat fragmentation: a simple computer simulation and a test using small mammals

Habitat fragmentation involves a reduction in the effective area available to a population and the imposition of hard patch edges. Studies seeking to measure effects of habitat fragmentation have compared populations in fragments of different size to estimate and area effect but few have examined the effect of converting open populations to closed ones (an effect of edges). To do so requires a shift in spatial scope-from comparison of individual fragments to that of fragmented versus unfragmented landscapes. Here we note that large-scale, “controlled” studies of habitat fragmentation have rarely been performed and are needed. In making our case we develop a simple computer simulation model based on how individual animals with home ranges are affected by the imposition of habitat edges, and use it to predict population-level responses to habitat fragmentation. We then compare predictions of the model with results from a field experiment on Peromyscus and Microtus. Our model treats the case where home ranges/territories fall entirely within or partially overlap with that of sample areas in continuous landscapes, but are restricted to areas within habitat fragments in impacted landscapes. Results of the simulations demonstrate that the imposition of hard edges can produce different population abundances for similar-sized areas in continuous and fragmented landscapes. This edge effect is disproportionately greater in small than large fragments and for species with larger than smaller home ranges. These predictions were generally supported by our field experiment. We argue that large-scale studies of habitat fragmentation are sorely needed, and that control-experiment contrasts of fragmented and unfragmented microlandscapes provide a logical starting point.

Synchrony, extinction, and dynamics of spatially segregated, heterogeneous populations

Most theoretical assessments of synchrony assume that populations are identical in their ability to produce and absorb dispersing individuals. Relaxing this assumption, I assessed how the local population growth rate, the proportion of individuals dispersing between populations, and different patterns in the immigration rate and carrying capacity with habitat patch size affect synchrony in local population dynamics and metapopulation extinction. All factors influenced both synchrony and extinction, and there were strong interactions between the factors. Local population growth rates had the largest effect on synchrony, followed by the pattern of carrying capacity with patch size and the proportion dispersing. The pattern of immigration with patch size had the smallest effect on synchrony. Metapopulation extinction was largely affected by the population growth rate, with the other factors having smaller but similar effects. Highly synchronous dynamics tended to be associated with greater metapopulation extinction within regions of chaotic local population growth; however, across the range of population growth investigated, high amounts of synchrony did not necessitate extinction. Differences among populations in the ability to absorb individuals mediated by interactions between growth rate, dispersal, patch size, and carrying capacity can substantially affect synchrony and extinction. Not considering how these factors interact may lead to erroneous conclusions regarding synchrony and extinction. Under certain situations, heterogeneity among local populations can promote metapopulation stability despite highly chaotic local dynamics.