Spatial patterns of species and plant traits in response to 20 years of grazing exclusion in subalpine grassland communities

Question: Does long‐term grazing exclusion affect spatial patterns of canopy height, plant species and traits in subalpine grassland communities? Are spatial patterns of species and traits similarly affected by grazing exclusion? Are changes in spatial patterns of species associated with changes in species abundances? Location: Subalpine grasslands, Vercors and Oisans Mountains, Alps (France). Methods: Spatial sampling of vegetation and measurements of plant traits were carried out within nine pairs of grazed and ungrazed 10 m × 10‐m plots in three species‐rich communities with different productivities. We estimated within‐plot spatial patterns of canopy height, species and aggregated trait values by measuring the extent (or patch size) and intensity of spatial dependence with Moran's I. Abundance‐weighted averages for species patch size and intensity of spatial dependence were calculated across all species per plot and across species per life form. Such measures derived from analysis of spatial dependence were considered spatial traits. Results: Response of spatial patterns to grazing exclusion was only detected in patch size, whereas intensity of spatial dependence was not affected. Changes in spatial patterns were community‐dependent because spatial traits based on patch size of canopy height and species increased following grazing exclusion only in the less productive community. Unexpectedly, changes in spatial patterns of species did not support changes in spatial patterns of trait values. Abundances and patch sizes of several life forms were significantly affected by grazing exclusion. However, at the scale investigated, changes in abundance of life forms did not correspond to changes in their spatial patterns and vice versa. Conclusion: In species‐rich communities, grazing alters spatial spread of species (i.e. patch size) rather than intra‐specific aggregation (i.e. intensity of spatial dependence). Results revealed possible mechanisms of species spatial reorganisation that are independent of abundance variation. Therefore, it is important to consider changes in spatial patterns in addition to changes in mean values of vegetation features when assessing impacts of grazing management.     

Plant patch structure modifies parasitoid assemblage richness of a specialist herbivore

1. The spatial structure of plant patches has been shown to affect host–parasitoid interactions, but its influence on parasitoid diversity remains largely ignored. Here we tested the prediction that parasitoid species richness of the specialist leafminer Liriomyza commelinae increases in larger and less isolated patches of its host plant Commelina erecta. We also explored whether parasitoid abundance and body size affected the occurrence of parasitoid species in local assemblages. 2. A total of 893 naturally established C. erecta patches were sampled on 18 sites around Córdoba city (Argentina). Also, two experiments were performed by creating patches differing in the number of plants and the distance from a parasitoid source. For these tests, plants were infected with the miner in the laboratory prior to placement in the field. 3. Plant patch size, independently of host abundance, positively affected the number of parasitoid species in both survey observations and experimental data. However, plant patch isolation did not influence parasitoid species richness. 4. The probability of finding rare parasitoid species increased with patch size, whereas occupation of isolated patches was independent of dispersal abilities (body size) of parasitoid species. 5. Overall, the results highlight the importance of considering spatial aspects such as the size of plant patches in the study of parasitoid communities.     

Patch size influences perceived patch quality for colonising Culex mosquitoes

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Scale and the guild functional response: Density-dependent predation varies with plot size

The predator functional response is an important mechanism determining the persistence of prey species; however, little is known about the effects of spatial scale on the functional response. We used a manipulative field experiment to quantify the effects of plot size on the guild functional response on the clam Mercenaria mercenaria, replicating the experiment in the summer in Chesapeake Bay, Virginia, and in the spring and fall in Indian River Lagoon, Florida, to examine the effects of predator and alternative prey abundance. In Virginia, the predation rate increased with both patch size and predator density, and was described by a modified sigmoid Type III functional response model that incorporated the effects of patch size. In Florida in the spring, the predator functional response was a Type III and did not vary with plot size, but in the fall it was a linear Type I at small plot sizes, and a Type III at a larger plot size. We hypothesize that the difference is primarily driven by changes in predator abundance and species between sites. In showing that the functional response can vary with plot size and season, our results indicate that small-scale experiments do not always scale up spatially or temporally. We suggest that the predictive power of such experiments may be limited by the complexity of the food web.     

Spatial metrics effect of forest fragmentation on forest bird abundance and site occupancy probability: the influence of patch size and isolation

The persistence of species taxa within fragmented habitats is dependent on the source–sink metapopulation processes, and forest patch size and isolation are key factors. Unveiling species–patch area and/or species–patch isolation relationships may help provide crucial information for species and landscape management. In this study, relationship between forest patch size and isolation with abundance and occupancy probability of forest-dependent birds was investigated. This study was based within a coastal landscape that faces deleterious human activities such as clearing for agriculture. The study aimed to answer the question of whether the size and extent of isolation of forest patches influence abundance and/or occupancy probability of forest-specialist and generalist birds. Two bird species, namely Tiny Greenbul Phyllastrephus debilis subsp. rabai and Yellow-bellied Greenbul Chlorocichla flaviventris, were used as models. Birds were surveyed using distance sampling methods, and spatial metrics were measured from satellite imagery. Focal forest size and distance between forest patches were the most influential metrics whereby abundance and occupancy probabilities increased with increasing patch size, but were negatively influenced by increasing gaps between patches. These findings provide evidence of the existence of patch size/ isolation–occupancy relationships characterised by higher occupancy rate of large patches and distance-dependent dispersal, which decreased with increasing gaps between patches. Controlling deleterious human activities that reduce forest size should be a priority for the long-term conservation of forest-dependent birds.     

Does the type of matrix matter? A quantitative review of the evidence

It has been increasingly recognized that the type of matrix surrounding habitat patches can affect biodiversity in landscapes, but there were only qualitative reviews of the subject focused on particular taxonomic groups. We present a quantitative review of studies from 1985 to 2008 that compared effects of different matrix types on individuals, populations and communities. We compiled 104 studies, most on animals, covering a broad range of landscape types and spatial scales. Most studies were empirical, focused on individuals and communities, and evaluated abundance/richness in the patch as the dependent variable. The type of matrix surrounding habitat patches influenced the studied parameters in 95% of the studies, but such effects were overall smaller compared to patch size or isolation effects. Matrix type effects were strongly species-specific, with different species responding differently to matrix type in 96% of studies comparing species or group of species. In 88% of studies, matrix types more similar in structure to the patch had higher quality for the studied organisms from the point of view of functional connectivity. Overall, the type of matrix is important, but patch size and isolation are the main determinants of ecological parameters in landscapes. Matrix quality generally increases with increasing structural similarity with habitat patches, a pattern that could be used as a general guideline for management of the matrix in fragmented landscapes.     

Impacts of forest fragmentation on species composition and forest structure in the temperate landscape of southern Chile

Aim Few studies have explicitly examined the influence of spatial attributes of forest fragments when examining the impacts of fragmentation on woody species. The aim of this study was to assess the diverse impacts of fragmentation on forest habitats by integrating landscape‐level and species‐level approaches. Location The investigation was undertaken in temperate rain forests located in southern Chile. This ecosystem is characterized by high endemism and by intensive recent changes in land use. Method Measures of diversity, richness, species composition, forest structure and anthropogenic disturbances were related to spatial attributes of the landscape (size, shape, connectivity, isolation and interior forest area) of forest fragments using generalized linear models. A total of 63 sampling plots distributed in 51 forest fragments with different spatial attributes were sampled. Results Patch size was the most important attribute influencing different measures of species composition, stand structure and anthropogenic disturbances. The abundance of tree and shrub species associated with interior and edge habitats was significantly related to variation in patch size. Basal area, a measure of forest structure, significantly declined with decreasing patch size, suggesting that fragmentation is affecting successional processes in the remaining forests. Small patches also displayed a greater number of stumps, animal trails and cow pats, and lower values of canopy cover as a result of selective logging and livestock grazing in relatively accessible fragments. However, tree richness and β‐diversity of tree species were not significantly related to fragmentation. Main conclusions This study demonstrates that progressive fragmentation by logging and clearance is associated with dramatic changes in the structure and composition of the temperate forests in southern Chile. If this fragmentation process continues, the ability of the remnant forests to maintain their original biodiversity and ecological processes will be significantly reduced.     

Experimental effects of habitat fragmentation on old-field canopy insects: community, guild and species responses

We examined the effects of habitat fragmentation on the species distributions, guild membership, and community structure of old-field insects using a fine-scale experimental approach. A continuous 1-ha goldenrod field was fragmented into four treatments that varied in both patch size and degree of isolation. Each treatment was replicated four times and arranged in a Latin square design. Canopy insects in fragmented patches were sampled with sweep nets during early and late summer 1995. The species richness of insects was significantly lower in fragmented than in unfragmented treatments during July, but was similar among treatments in September. Overall community abundance showed no treatment effect during either month. We also found significant row and column effects, suggesting there was spatial heterogeneity in species richness and abundance apart from treatment effects. Differences in species richness during July were primarily due to the loss of rare species in highly fragmented plots. Overall abundance was less responsive to community change because deletions of rare species in fragmented areas were not detected in abundance analyses. Four feeding guilds showed different responses to fragmentation: the species richness of sucking herbivores and the abundance of parasitoids were significantly reduced by fragmentation but predators and chewing herbivores were largely unaffected. Analyses of a subset of individual species within guilds suggest that the greater effects of fragmentation on sucking herbivores and parasitoids may be due to the degree of habitat specificity of guild members. The effects of small-scale habitat fragmentation were therefore detectable at the level of community, guild, and individual species. Changes in species richness, guild structure and species distributions were likely due to differential effects of habitat alteration on individual movements and patch selection rather than dispersal or demographic change. Nonetheless, the selective loss of rare species, differential guild effects and changes in species occupancy that we found in this small-scale experiment are also factors that are likely to operate in fragmented habitats over broader spatial scales. Received: 11 May 1998 / Accepted: 27 September 1998     

Disturbance and the community structure of stream invertebrates: patch-specific effects and the role of refugia

1. We have previously shown that the impact of spates on stream invertebrates may differ among patches separated by distances of metres or less. Here we analyse the species-specific flood responses of larval chironomids and adult and near mature copepods living in different patch types. Four patch types (with eight replicates of each) were compared: the sandy mid-channel, fine sediments around dams, coarse sediments around dams, and dam debris. Additionally, since some fine sediment patches had been shown previously to act as flow refugia while others did not, we also examined species-specific responses in refugium vs. non-refugium fine sediment patches. Detrended correspondence analysis was used to test for changes in assemblage structure (species composition and relative abundance). 2. Species richness was not altered in a predictable manner by floods; the least stable patch types (mid-channel and coarse patches) did not necessarily show reduced species richness during the spate. 3. As indicated by the spread of DCA ordination scores, there was generally a high degree of overlap in the species composition among the four patch types. Nevertheless, copepod species composition and relative abundance were more similar among patch types during the spate than pre-spate. Spates may induce a re-distribution of copepod species among the patch types. Chironomid species composition and relative abundance were no more similar among patch types during the spate than pre- or post-spate. 4. For both chironomids and copepods, species composition and relative abundance (as assessed by DCA ordination scores) in refugium patches changed more in response to the spate than in the non-refugium patches. An influx of individuals from just a few species for each group was responsible for the change in assemblage structure. Thus, despite the fact that our past work has shown that refugia may confer enhanced resistance and resilience of copepod and chironomid assemblages in terms of total faunal abundances, the present work suggests that resistance and resilience of the species composition of the community apparently are no greater in refugium patches than in non-refugium patches.     

Species- and community-level responses to habitat spatial changes in fragmented rainforests: assessing compensatory dynamics in amphibians and reptiles

The rapid loss and degradation of tropical forests threatens the maintenance of biodiversity across different spatial scales. Nevertheless, the extirpation and population decline of some disturbance-sensitive species may be compensated for by colonization and proliferation of disturbance-adapted species, thus allowing distributions of community-level attributes (e.g., abundance and diversity) to be preserved in human-modified tropical landscapes. To test this poorly assessed hypothesis we evaluated species- and community-level responses of amphibians and reptiles to differences in forest patch (patch size, shape, and distance to water bodies) and landscape metrics (old-growth forest cover, degree of fragmentation, and matrix composition) in the fragmented Lacandona rainforest, Mexico. We found that the abundance of several amphibian and reptile species was strongly associated with forest patch and landscape attributes, being particularly higher in larger patches surrounded by a greater forest cover. Such changes at the species level generated notable changes in reptile communities. In particular, the abundance, diversity, and evenness of reptile communities were strongly related to patch size, patch shape, and matrix composition. Yet, because of compensatory dynamics in amphibians, this group showed weak responses at the community level. Despite such compensatory dynamics, our results indicate that forest loss at the patch and landscape levels represents the main threat to both amphibians and reptiles, thus indicating that to preserve herpetological communities in this biodiversity hotspot, conservation initiatives should be focused on preventing further deforestation.     

Determinants of distribution and abundance in the clouded apollo butterfly: a landscape ecological approach

Recent studies on the determinants of distribution and abundance of animals at landscape level have emphasized the usefulness of the metapopulation approach, in which patch area and habitat connectivity have often proved to explain satisfactorily existing patch occupancy patterns. A different approach is needed to study the common situation in which suitable habitat is difficult to determine or does not occur in well‐defined habitat patches. We applied a landscape ecological approach to study the determinants of distribution and abundance of the threatened clouded apollo Parnassius mnemosyne butterfly within an area of 6 km² of agricultural landscape in south‐western Finland. The relative role of 24 environmental variables potentially affecting the distribution and abundance of the butterfly was studied using a spatial grid system with 2408 grid squares of 0.25 ha, of which 349 were occupied by the clouded apollo. Both the probability of butterfly presence and abundance in a 0.25 ha square increased with the presence of the larval host plant Corydalis solida the cover of semi‐natural grassland, the amount of solar radiation and spalial autocorrelation in butterfly occurrence. Additionally, butterfly abundance increased with overall mean patch size and decreased with maximum slope angle and wind speed. Two advantages of the employment of a spatial grid system included the avoidance of a subjective definition of suitable habitat patches and an evaluation of the relative significance of different components of habitat quality at the same time with habitat availability and connectivity. The large variation in habitat quality was influenced by the abundance of the larval host plant and adult nectar sources but also by climatological. topographical and structural factors. The application of a spatial grid system as used here has potential for a wide use in studies on landscape‐level distribution and abundance patterns in species with complex habitat requirements and habitat availability patterns.     

Internal Habitat Quality Determines the Effects of Fragmentation on Austral Forest Climbing and Epiphytic Angiosperms

Habitat fragmentation has become one of the major threats to biodiversity worldwide, particularly in the case of forests, which have suffered enormous losses during the past decades. We analyzed how changes in patch configuration and habitat quality derived from the fragmentation of austral temperate rainforests affect the distribution of six species of forest-dwelling climbing and epiphytic angiosperms. Epiphyte and vine abundance is primarily affected by the internal characteristics of patches (such as tree size, the presence of logging gaps or the proximity to patch edges) rather than patch and landscape features (such as patch size, shape or connectivity). These responses were intimately related to species-specific characteristics such as drought- or shade-tolerance. Our study therefore suggests that plant responses to fragmentation are contingent on both the species'' ecology and the specific pathways through which the study area is being fragmented, (i.e. extensive logging that shaped the boundaries of current forest patches plus recent, unregulated logging that creates gaps within patches). Management practices in fragmented landscapes should therefore consider habitat quality within patches together with other spatial attributes at landscape or patch scales.     

Biotic filtering and mass effects in small shrub patches: is arthropod community structure predictable based on the quality of the vegetation?

1. Community assembly is affected by four processes: dispersal, filtering effects (selection), ecological drift and evolution. The role of filtering relative to dispersal and drift should decline with patch size, hampering possibilities to predict which organisms will be observed within small‐sized patches. However, vegetation structure is known to have a marked impact on species assemblages, and plant quality may act as a biotic filter. This challenges the assumption of unpredictable species assemblages in small‐sized vegetation patches. 2. Using 32 stands of five shrub species in south‐west Finland, this study investigated whether biotic filtering effects caused by patch‐forming plants are strong enough to overcome the mixing of mobile arthropod assemblages across small patches. 3. Stochastic variation did not hide the signals of biotic filtering and dispersal in the small shrub patches. Habitat richness around the patches explained a three times larger share of variation in the species composition than did the identity of the patch‐forming plant, but it had less effect on the abundance of arthropods. A radius of 50–100 m around a patch explained the species composition best. 4. Abundance patterns varied between the feeding guilds; the patch‐forming shrub influenced the abundances of detritivores and leaf‐feeding herbivores, whereas the abundances of flower‐visiting herbivores appeared to track the flowering phenology of the plants. Shrub identity had little effect on omnivores or predators. Predator abundances were correlated with the abundance of potential prey. 5. The results of this study suggest that community composition within a vegetation patch may be predictable even if dispersal overrides local filtering effects, as suggested by the mass‐effects paradigm.     

Habitat amount hypothesis and passive sampling explain mammal species composition in Amazonian river islands

Nested structures of species assemblages have been frequently associated with patch size and isolation, leading to the conclusion that colonization–extinction dynamics drives nestedness. The ‘passive sampling’ model states that the regional abundance of species randomly determines their occurrence in patches. The ‘habitat amount hypothesis’ also challenges patch size and isolation effects, arguing that they occur because of a ‘sample area effect’. Here, we (a) ask whether the structure of the mammal assemblages of fluvial islands shows a nested pattern, (b) test whether species’ regional abundance predicts species’ occurrence on islands, and (c) ask whether habitat amount in the landscape and matrix resistance to biological flow predict the islands’ species composition. We quantified nestedness and tested its significance using null models. We used a regression model to analyze whether a species’ relative regional abundance predicts its incidence on islands. We accessed islands’ species composition by an NMDS ordination and used multiple regression to evaluate how species composition responds to habitat amount and matrix resistance. The degree of nestedness did not differ from that expected by the passive sampling hypothesis. Likewise, species’ regional abundance predicted its occurrence on islands. Habitat amount successfully predicted the species composition on islands, whereas matrix resistance did not. We suggest the application of habitat amount hypothesis for predicting species composition in other patchy systems. Although the island biogeography perspective has dominated the literature, we suggest that the passive sampling perspective is more appropriate for explaining the assemblages’ structure in this and other non‐equilibrium patch systems. Abstract in Portuguese is available with online material.     

Consequences for a host–parasitoid interaction of host-plant aggregation, isolation, and phenology

Abstract.  1. Spatial habitat structure can influence the likelihood of patch colonisation by dispersing individuals, and this likelihood may differ according to trophic position, potentially leading to a refuge from parasitism for hosts.
2. Whether habitat patch size, isolation, and host-plant heterogeneity differentially affected host and parasitoid abundance, and parasitism rates was tested using a tri-trophic thistle–herbivore–parasitoid system.
3.  Cirsium palustre thistles ( n = 240) were transplanted in 24 blocks replicated in two sites, creating a range of habitat patch sizes at increasing distance from a pre-existing source population. Plant architecture and phenological stage were measured for each plant and the numbers of the herbivore Tephritis conura and parasitoid Pteromalus elevatus recorded.
4. Mean herbivore numbers per plant increased with host-plant density per patch, but parasitoid numbers and parasitism rates were unaffected. Patch distance from the source population did not influence insect abundance or parasitism rates. Parasitoid abundance was positively correlated with host insect number, and parasitism rates were negatively density dependent. Host-plant phenological stage was positively correlated with herbivore and parasitoid abundance, and parasitism rates at both patch and host-plant scales.
5. The differential response between herbivore and parasitoid to host-plant density did not lead to a spatial refuge but may have contributed to the observed parasitism rates being negatively density dependent. Heterogeneity in patch quality, mediated by variation in host-plant phenology, was more important than spatial habitat structure for both the herbivore and parasitoid populations, and for parasitism rates.     

Competition versus facilitation: conspecific effects on pollinator visitation and seed set in the iris Lapeirousia oreogena

Ecological interactions between conspecific plants can range from facilitative to competitive depending on the spacing and abundance of individuals. Competitive interactions are expected when plants flower en‐masse and availability of pollinators is limited. We tested this prediction using Lapeirousia oreogena, a mass‐flowering South African iris that is pollinated by a single species of long‐proboscid fly. Controlled hand‐pollination experiments showed that L. oreogena is self‐compatible, but reliant on pollinator visits for seed set. Seed production per flower showed a significant negative relationship with patch size (and the correlated measure of number of individuals per patch), but was not affected by flower density or distance to neighbouring patches. There was a tendency for fly abundance to increase with patch size, but the rate of visits to individual flowers by flies was not affected by patch size. Seed set of hand‐pollinated flowers did not differ for plants in and out of dense patches, indicating that the large differences in seed set among patches were likely to reflect pollinator visits, rather than the genetic or physiological capacity of plants to produce seeds. The reduced fecundity of L. oreogena in large patches with a greater numbers of flowers is consistent with the idea that plants with highly specialized pollination systems can experience intra‐specific competition for pollination.     

The role of size and dominance in the feeding behaviour of coexisting hummingbirds

Interspecific competition can strongly influence community structure and limit the distribution and abundance of species. One of the main factors that determine hummingbird community structure is competition for food. The temporal and spatial distribution of nectar has a strong impact on hummingbird assemblages, shaping foraging niches according to hummingbird dominance and foraging strategy. We investigated whether body size and the degree of aggressive dominance influence feeding behaviour of hummingbirds in a temperate forest in northwestern Mexico (El Palmito, Mexico) when winter migrant hummingbirds are present in the community. First, we determined the dominance status of hummingbirds and evaluated the relationship between dominance and body mass, wing disc loading and migratory status. Secondly, we determined how hummingbird species used plant species differently. Thirdly, we examined whether the most dominant hummingbird species defended floral patches with more energy and/or with a larger number of flowers. At each flower patch, hummingbird species, number of hummingbird interactions, feeding time and number of flowers present were recorded. The total number of calories available within each floral patch was also determined. Our results demonstrate that the dominance hierarchy of 13 hummingbird species (migratory and resident) was correlated with body size but not wing disc loading, and that members of the hummingbird community showed a clear separation in resource use (by plant species). Hummingbirds at the top of the dominance hierarchy defended and fed on the best flower patches, defined by the quantity of calories available. Hence, the feeding behaviour of hummingbirds at El Palmito depends on the abundance of plant species used by hummingbirds and on the amount of energy available from each flower patch. Thus, hummingbird body size, aggressive dominance and defence of quality flower patches determines niche partitioning among species.     

Patch Size and Isolation Predict Plant Species Density in a Naturally Fragmented Forest

Studies of the effects of patch size and isolation on plant species density have yielded contrasting results. However, much of the available evidence comes from relatively recent anthropogenic forest fragments which have not reached equilibrium between extinction and immigration. This is a critical issue because the theory clearly states that only when equilibrium has been reached can the number of species be accurately predicted by habitat size and isolation. Therefore, species density could be better predicted by patch size and isolation in an ecosystem that has been fragmented for a very long time. We tested whether patch area, isolation and other spatial variables explain variation among forest patches in plant species density in an ecosystem where the forest has been naturally fragmented for long periods of time on a geological scale. Our main predictions were that plant species density will be positively correlated with patch size, and negatively correlated with isolation (distance to the nearest patch, connectivity, and distance to the continuous forest). We surveyed the vascular flora (except lianas and epiphytes) of 19 forest patches using five belt transects (50×4 m each) per patch (area sampled per patch = 0.1 ha). As predicted, plant species density was positively associated (logarithmically) with patch size and negatively associated (linearly) with patch isolation (distance to the nearest patch). Other spatial variables such as patch elevation and perimeter, did not explain among-patch variability in plant species density. The power of patch area and isolation as predictors of plant species density was moderate (together they explain 43% of the variation), however, a larger sample size may improve the explanatory power of these variables. Patch size and isolation may be suitable predictors of long-term plant species density in terrestrial ecosystems that are naturally and anthropogenically fragmented.     

What saplings can tell us about forest expansion over natural grasslands

Questions: 1. Do the species composition, richness and diversity of sapling communities vary significantly in differently sized patches? 2. Do forest patches of different sizes differ in woody plant colonization patterns? Location: São Francisco de Paula, Rio Grande do Sul, Brazil, 29°28'S,50°13'W. Methods: Three woody vegetation types, differing in structural development (patch size) and recovering for 10 years from cattle and burning disturbances, were sampled on grassland. We analysed the composition and complexity of the woody sapling communities, through relative abundance, richness and diversity patterns. We also evaluated recruitment status (residents vs. colonizers) of species in communities occurring in different forest patch size classes. Results : 1. There is a compositional gradient in sapling communities strongly associated with forest patch area. 2. Richness and diversity are positively correlated to patch area, but only in poorly structured patches; large patches present richness and diversity values similar to small patches. 3. Resident to colonizer abundance ratio increases from nurse plants to large patches. The species number proportion between residents and colonizers is similar in small and large patches and did not differ between these patch types. 4. Large patches presented a high number of exclusive species, while nurse plants and small patches did not. Conclusions: Woody plant communities in Araucaria forest patches are associated with patch structure development. Richness and diversity patterns are linked to patch colonization patterns. Generalist species colonize the understorey of nurse plants and small patches; resident species cannot recruit many new individuals. In large patches, sapling recruitment by resident adults precludes the immigration of new species into the patches, limiting richness and diversity.     

Landscape structure and the genetic effects of a population collapse

Both landscape structure and population size fluctuations influence population genetics. While independent effects of these factors on genetic patterns and processes are well studied, a key challenge is to understand their interaction, as populations are simultaneously exposed to habitat fragmentation and climatic changes that increase variability in population size. In a population network of an alpine butterfly, abundance declined 60–100% in 2003 because of low over-winter survival. Across the network, mean microsatellite genetic diversity did not change. However, patch connectivity and local severity of the collapse interacted to determine allelic richness change within populations, indicating that patch connectivity can mediate genetic response to a demographic collapse. The collapse strongly affected spatial genetic structure, leading to a breakdown of isolation-by-distance and loss of landscape genetic pattern. Our study reveals important interactions between landscape structure and temporal demographic variability on the genetic diversity and genetic differentiation of populations. Projected future changes to both landscape and climate may lead to loss of genetic variability from the studied populations, and selection acting on adaptive variation will likely occur within the context of an increasing influence of genetic drift.