A comparison of the responses of two tropical specialist herbivores to host plant patch size

Summary The effects of host plant patch size on the abundances of two specialist herbivores (the chrysomelid beetle, Acalymma innubum and the pentatomid bug, Piezosternum subulatum) were investigated in a natural forest community in the Virgin Islands. Abundances were compared early and late in the season in different sized patches of the cucurbit host plant (Cayaponia americana) growing in open habitat (with no surrounding plant community) and forest habitat (with diverse surrounding plant community). For both herbivore species, adult abundances per patch were positively correlated with patch leaf area, but there was a significant patch size effect (i.e., correlation between herbivore density per unit plant and patch leaf area) only for beetles in the forest habitat. Both herbivore species were significantly affected by surrounding plant diversity, but in opposite ways: beetles were more abundant in open patches whereas bugs were more abundant in forest patches. Relationships between abundance and patch size in open and forest patches changed through the season for both herbivore species. These changing abundance patterns are discussed with respect to (1) increases in the diversity of the plant community surrounding host plant patches, and (2) differences in herbivore movement patterns.     

Landscape-scale habitat fragmentation is positively related to biodiversity,despite patch-scale ecosystem decay

Positive effects of habitat patch size on biodiversity are often extrapolated to infer negative effects of habitat fragmentation on biodiversity at landscape scales. However, such cross-scale extrapolations typically fail. A recent, landmark, patch-scale analysis (Chase et al., 2020, Nature 584, 238–243) demonstrates positive patch size effects on biodiversity, that is, ‘ecosystem decay’ in small patches. Other authors have already extrapolated this result to infer negative fragmentation effects, that is, higher biodiversity in a few large than many small patches of the same cumulative habitat area. We test whether this extrapolation is valid. We find that landscape-scale patterns are opposite to their analogous patch-scale patterns: for sets of patches with equal total habitat area, species richness and evenness decrease with increasing mean size of the patches comprising that area, even when considering only species of conservation concern. Preserving small habitat patches will, therefore, be key to sustain biodiversity amidst ongoing environmental crises.     

Distribution and colonisation ability of three parasitoids and their herbivorous host in a fragmented landscape

Habitat fragmentation can disrupt communities of interacting species even if only some of the species are directly affected by fragmentation. For instance, if parasitoids disperse less well than their herbivorous hosts, habitat fragmentation may lead to higher herbivory in isolated plant patches due to the absence of the third trophic level. Community-level studies suggest that parasitoids tend to have limited dispersal abilities, on the order of tens of metres, much smaller than that of their hosts, while species-oriented studies document dispersal by parasitoids on the scale of kilometres. In this study the distribution patterns of three parasitoid species with different life histories and their moth host, Hadena bicruris, a specialist herbivore of Silene latifolia, were compared in a large-scale network of natural fragmented plant patches along the rivers Rhine and Waal in the Netherlands. We examined how patch size and isolation affect the presence of each species. Additionally, experimental plots were used to study the colonisation abilities of the species at different distances from source populations.In the natural plant patches the presence of the herbivore and two of the parasitoids, the gregarious specialist Microplitis tristis and the gregarious generalist Bracon variator were not affected by patch isolation at the scale of the study, while the solitary specialist Eurylabus tristis was. In contrast to the herbivore, the presence of all parasitoid species declined with plant patch size. The colonisation experiment confirmed that the herbivore and M. tristis are good dispersers, able to travel at least 2 km within a season. B. variator showed intermediate colonisation ability and E. tristis showed very limited colonisation ability at this spatial scale. Characteristics of parasitoid species that may contribute to differences in their dispersal abilities are discussed.     

Soil nutrient status determines how elephant utilize trees and shape environments

1. Elucidation of the mechanism determining the spatial scale of patch selection by herbivores has been complicated by the way in which resource availability at a specific scale is measured and by vigilance behaviour of the herbivores themselves. To reduce these complications, we studied patch selection by an animal with negligible predation risk, the African elephant. 2. We introduce the concept of nutrient load as the product of patch size, number of patches and local patch nutrient concentration. Nutrient load provides a novel spatially explicit expression of the total available nutrients a herbivore can select from. 3. We hypothesized that elephant would select nutrient-rich patches, based on the nutrient load per 2500 m(2) down to the individual plant scale, and that this selection will depend on the nitrogen and phosphorous contents of plants. 4. We predicted that elephant would cause more adverse impact to trees of lower value to them in order to reach plant parts with higher nutrient concentrations such as bark and root. However, elephant should maintain nutrient-rich trees by inducing coppicing of trees through re-utilization of leaves. 5. Elephant patch selection was measured in a homogenous tree species stand by manipulating the spatial distribution of soil nutrients in a large field experiment using NPK fertilizer. 6. Elephant were able to select nutrient-rich patches and utilized Colophospermum mopane trees inside these patches more than outside, at scales ranging from 2500 down to 100 m(2) . 7. Although both nitrogen and phosphorus contents of leaves from C. mopane trees were higher in fertilized and selected patches, patch choice correlated most strongly with nitrogen content. As predicted, stripping of leaves occurred more in nutrient-rich patches, while adverse impact such as uprooting of trees occurred more in nutrient-poor areas. 8. Our results emphasize the necessity of including scale-dependent selectivity in foraging studies and how elephant foraging behaviour can be used as indicators of change in the availability of nutrients.     

Mixed effects of habitat fragmentation on species richness and community structure in a microarthropod microecosystem

Abstract.  1. Theory is unclear about the optimal degree of isolation of habitat fragments where the aim is to maximise species richness. In a field-based microecosystem of Collembola and predatory and non-predatory mites, moss patches of the same total area were fragmented to varying degrees. The habitat was left for several months to allow the communities to approach a new state of equilibrium.
2. The species richness (in particular of predatory mites) of a given area of habitat was greater when it was part of a large mainland area than part of an island, in agreement with theory.
3. Conversely, species richness and abundance were largely unaffected by fragmentation of a fixed area of island habitat. In this case, it is suggested here that the advantages of several small patches (e.g. reduced impact of environmental stochasticity, wider range of habitats overall) were equally balanced by the advantages of a single large patch (e.g. reduced effect of demographic stochasticity, wider range of habitats within a single patch, reduced edge effect), or that both effects were small.
4. The shapes of rank–abundance curves were similar among the levels of fragmentation of a fixed area of island habitat, implying that fragmentation had little impact on community structure. Conversely, the species composition of non-predatory mites varied weakly, but significantly, by fragmentation.     

The distinct effects of habitat fragmentation on population size

We sought to understand how the separation of habitats into spatially isolated fragments influences the abundance of organisms. Using a simple, deterministic model of population growth, we compared analytically exact solutions predicting abundance of consumers in two isolated patches with abundance of consumers in a single large patch where the carrying capacity of the large patch is the sum of the carrying capacities of the isolated ones. For the deterministic model, the effect of fragmentation was to slow the rate of population growth in the fragmented habitat relative to the intact one. We also analyzed a stochastic version of the model to examine the effect of fragmentation on population abundance when resources vary randomly in time. For the stochastic model, the effect of fragmentation was to reduce population abundance. We proved in closed-form, that for a non-equilibrium population exhibiting logistic population growth, fragmentation will reduce population size even when the total carrying capacity is not affected by fragmentation. We provide a theoretical basis for the prediction that habitat fragmentation amplifies the effect of habitat loss on the abundance of mobile organisms.     

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.     

Fragmentation in calcareous grasslands: species specialization matters

Habitat fragmentation resulting from anthropogenic land-use change may negatively affect both biodiversity and ecosystem structure and function. However, susceptibility to fragmentation varies between species and may be influenced by for instance specialization, functional traits and trophic level. We examined how total and specialist species richness, species composition and functional trait composition at two trophic levels (vascular plants and sap-feeding hoppers) vary with habitat fragmentation (patch size and connectivity) in dry calcareous grasslands in southeast Norway. We found that fragmentation affected plant and hopper species composition both totally and of habitat specialists, but with a net species loss only for the specialists, indicating greater susceptibility of specialized species. Reductions in patch size and increasing isolation negatively affected plant specialists with different sets of traits, effectively reducing the number of species with trait combinations suitable to persist in small and isolated patches. Fragmentation influenced trait composition of the total hopper community, but not of habitat specialists. A lesser degree of habitat association could explain why hoppers, despite belonging to a higher tropic level, seemed to be less susceptible to fragmentation than plants. Nonetheless, our study shows that habitat fragmentation affects both species richness, species composition and trait composition of plants and hoppers, indicating that fragmentation leads not only to a loss of species, but also alters dominance hierarchies and the functionality of grassland communities.     

Combined effects of patch size and plant nutritional quality on local densities of insect herbivores

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.     

Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance

Seagrasses form temporally dynamic, fragmented subtidal landscapes in which both large- and small-scale habitat structure may influence faunal survival and abundance. We compared the relative influences of seagrass (Zostera marina L.) habitat fragmentation (patch size and isolation) and structural complexity (shoot density) on juvenile blue crab (Callinectes sapidus Rathbun) survival and density in a Chesapeake Bay seagrass meadow. We tethered crabs to measure relative survival, suction sampled for crabs to measure density, and took seagrass cores to measure shoot density in patches spanning six orders of magnitude (ca. 0.25-30,000 m²) both before (June) and after (September) seasonally predictable decreases in seagrass structural complexity and increases in seagrass fragmentation. We also determined if juvenile blue crab density and seagrass shoot density varied between the edge and the interior of patches. In June, juvenile blue crab survival was not linearly related to seagrass patch size or to shoot density, but was significantly lower in patches separated by large expanses of unvegetated sediment (isolated patches) than in patches separated by <1 m of unvegetated sediment (connected patches). In September, crab survival was inversely correlated with seagrass shoot density. This inverse correlation was likely due to density-dependent predation by juvenile conspecifics (i.e. cannibalism); juvenile blue crab density increased with seagrass shoot density, was inversely correlated with crab survival, and was greater in September than in June. Shoot density effects on predator behavior and on conspecific density also likely caused crab survival to be lower in isolated patches than in connected patches in June. Isolated patches were either large (patch area >3000 m²) or very small (<1 m²). Large isolated patches had the lowest shoot densities, which may have allowed predators to easily find tethered crabs. Very small isolated patches had the highest shoot densities and consequently a high abundance of predators (=juvenile conspecifics). Though shoot density did not differ between the edge and the interior of patches, crabs were more abundant in the interior of patches than at the edge. These results indicate that seagrass fragmentation does not have an overriding influence on juvenile blue crab survival and density, and that crab cannibalism and seasonal changes in landscape structure may influence relationships between crab survival and seagrass habitat structure. Habitat fragmentation, structural complexity, faunal density, and time all must be incorporated into future studies on faunal survival in seagrass landscapes.     

How predator incursions affect critical patch size: the role of the functional response

Understanding the impact of habitat edges provides a key to deciphering how community dynamics change as functions of habitat structure and spatial scale. Motivated by studies of predation on bird nests in forest fragments and other cases of "cross-boundary subsidies," we present results from a partial differential equation model in which a patch-resident prey species suffers incidental mortality from a generalist predator species residing in the surrounding matrix habitat. We demonstrate that predator intrusions have the potential to induce critical patch size effects for the prey species, even when the prey's dynamics would otherwise preclude such effects. We also demonstrate that the existence of critical patch size effects depends on the functional response of the predator, with Lotka-Volterra and Type II functional responses generating the effect (but not Type III). We conclude by discussing how predator-induced critical patch size effects can influence opportunities for regionwide persistence of the prey by altering the fraction and spatial distribution of meaningful patches within a metapopulation.     

Insular patterns of calicioid lichens in a boreal old-growth forest-wetland mosaic

Fragmentation of the forested landscape poses a threat to many aspects of biodiversity associated with old-growth forests Studies of the effects of forest fragmentation are often complicated by the variation in composition and age of patches and the matrix This study used a system of isolated stands where patch age and composition were similar and the matrix variability negligible The patches were composed of old-growth Picea abies stands of varying size and shape in a wetland matrix The study organisms were epiphytic crustose calicioid lichens (also known as Caliciales), many of which are very substrate-specific and restricted to old-growth stands The aim of the study was to measure the effect of patch size, patch isolation, habitat and substrate quality on the species riochness and composition of epiphytic calicioids Twenty-four patches ranging from 0 4 to 15 9 ha in size were studied All species of calicioid lichens were registered in 0 1 ha plots in each patch Isolation was measured as the percentage of available habitat within 400 m of a patch Twenty-two species were found with an average of 9 48 ± 0 26 (SE) species per patch and 292 ± 0 18 (SE) species per tree Species richness at patch level correlated with stand structure, primarily tree density, while number of species per tree (reflecting population size) was strongly correlated with island size and several stand variables There was no effect of isolation on species richness Species composition was influenced by both substrate variables and patch size The species composition on the islands showed a significant nestedness, i e species composition on species-poor islands constituted a non-random subset of the species composition on species-rich islands We propose that the explanation for the strong relationship between species richness at tree level and stand size is an edge effect which implies that unaffected interior areas only occur on large islands The different microclimate of the patch edge enables only the hardiest species to establish large populations there whilst shade and moisture demanding species are restricted to the interiors of larger islands     

Inter‐patch movement in an experimental system: the effects of life history and the environment

An important process for the persistence of populations subjected to habitat loss and fragmentation is the dispersal of individuals between habitat patches. Dispersal involves emigration from a habitat patch, movement between patches through the surrounding landscape, and immigration into a new suitable habitat patch. Both landscape and physical condition of the disperser are known to influence dispersal ability, although disentangling these effects can often be difficult in the wild. In one of the first studies of its kind, we used an invertebrate model system to investigate how dispersal success is affected by the interaction between the habitat condition, as determined by food availability, and life history characteristics (which are also influenced by food availability). Dispersal of juvenile and adult mites (male and female) from either high food or low food natal patches were tested separately in connected three patch systems where the intervening habitat patches were suitable (food supplied) or unsuitable (no food supplied). We found that dispersal success was reduced when low food habitat patches were coupled to colonising patches via unsuitable intervening patches. Larger body size was shown to be a good predictor of dispersal success, particularly when the intervening landscape is unsuitable. Our results suggest that there is an interaction between habitat fragmentation and habitat suitability in determining dispersal success: if patches degrade in suitability and this affects the ability to disperse successfully then the effective connectance across landscapes may be lowered. Understanding these consequences will be important in informing our understanding of how species, and the communities in which they are embedded, may potentially respond to habitat fragmentation.     

Distribution of birds in natural landscape mosaics of old-growth forests in northern Sweden: relations to habitat area and landscape context

We censused breeding birds for three years in natural landscape mosaics of virgin old-growth spruce forest and mire in a large protected forest area in northern Sweden Twenty forest patches, ranging from 0 2 to 17 8 ha in size, were selected in two matrix types, dominated by forest and mire, respectively Patches were very similar with regards to habitat features There was a strong effect of patch area on species richness, but no effect of matrix type Standardization of species richness by rarefaction revealed that small patches (<5 ha) had fewer and large patches (>10 ha) more species than expected Overall distribution of species across patches showed a highly significant nested pattern, indicating that a few habitat generalists occupy all size classes, whereas more demanding species avoid small patches regardless of landscape composition Individual species tended to be distributed evenly across patch classes and no significant edge effect in terms of density of birds was found Our results have bearings on actions to preserve avian diversity in northern boreal forests small patches (<5 ha) provide habitat only for habitat generalists, and therefore larger (>10 ha) patches should be preserved     

The effects of forest fragmentation on species richness on the Obudu Plateau,south‐eastern Nigeria

This study examines how fragmentation through changes in patch size and increasing distances between forest patches affects bird species richness. Numbers of bird species were higher in large forest patches and decrease with decreasing patch size. Bird species richness also decreased with increased isolation of forest patches. Suitable trees should be planted in the gaps between the forests that will create wildlife corridors to maintain connectivity.     

Does fragmentation of Urtica habitats affect phytophagous and predatory insects differentially?

Effects of habitat fragmentation on the insect community of stinging nettle (Urtica dioica L.) were studied, using 32 natural nettle patches of different area and degree of isolation in an agricultural landscape. Habitat fragmentation reduced the species richness of Heteroptera, Auchenorrhyncha, and Coleoptera, and the abundance of populations. Habitat isolation and area reduction did not affect all insect species equally. Monophagous herbivores had a higher probability of absence from small patches than all (monophagous and polyphagous) herbivore species, and the percentage of monophagous herbivores increased with habitat area. Abundance and population variability of species were negatively correlated and could both be used as a predictor of the percentage of occupied habitats. Species richness of herbivores correlated (positively) with habitat area, while species richness of predators correlated (negatively) with habitat isolation. In logistic regressions, the probability of absence of monophagous herbivores from habitat patches could only be explained by habitat area (in 4 out of 10 species) and predator absence probability only by habitat isolation (in 3 out of 14 species). Presumably because of the instability of higher-trophic-level populations and dispersal limitation, predators were more affected by habitat isolation than herbivores, while they did not differ from herbivore populations with respect to abundance or variability. Thus increasing habitat connectivity in the agricultural landscape should primarily promote predator populations. Received: 4 February 1998 / Accepted: 4 May 1998     

Support for the habitat amount hypothesis from a global synthesis of species density studies

Decades of research suggest that species richness depends on spatial characteristics of habitat patches, especially their size and isolation. In contrast, the habitat amount hypothesis predicts that (1) species richness in plots of fixed size (species density) is more strongly and positively related to the amount of habitat around the plot than to patch size or isolation; (2) habitat amount better predicts species density than patch size and isolation combined, (3) there is no effect of habitat fragmentation per se on species density and (4) patch size and isolation effects do not become stronger with declining habitat amount. Data on eight taxonomic groups from 35 studies around the world support these predictions. Conserving species density requires minimising habitat loss, irrespective of the configuration of the patches in which that habitat is contained.     

Community structure and demography of small-mammal populations in insular montane forests in southern India

bstract Small mammals were studied in insular montane forest patches in the Upper Nilgiris in southern India from February 1994 to September 1996. Nine patches were selected at two sites, one with a single large 600-ha patch, the other with several small patches ranging in size from 0.2 to 60 ha. The population characteristics and community structure of small mammals were studied in relation to patch size and habitat structure within the patches. The two most abundant species were Rattus rattus (2–36 individuals/ha) and Suncus montanus (0–11 individuals/ha), while the abundance of seven other species recorded were very low. The population characteristics of R. rattus examined were density, biomass, proportions of juveniles, sub-adults and adults, mean weight of all animals, mean weight of adults, sex ratio and persistence. The proportion of adults, sub-adults, mean weight of animals, and mean weight of adult males were correlated with patch size. Persistence of sub-adult females in the large patch was especially high. The density and biomass of other small-mammal species were also studied. Two indices of diversity, species richness and proportion of R.␣rattus were compared as measures of community structure. Seven habitat characteristics were measured; of these, canopy cover, canopy height and tree density were correlated with the size of the patch. Density and biomass of species other than R. rattus and proportion of R. rattus were correlated with canopy height. Density and biomass of species other than R. rattus were highest in smaller patches. While the population characteristics of R. rattus may be affected by patch size, the density of rare species may be influenced by factors related to lower canopy height. Migration between patches may be an important factor in maintaining populations in these patches. Received: 5 January 1998 / Accepted: 23 March 1998     

Fragment size affects plant herbivory via predator loss

Fragmentation and resultant changes in patch size are predicted to alter species diversity and community composition, yet the consequences of these differences for species interactions are poorly understood. Theory predicts that predators are more sensitive to fragmentation than their prey, resulting in greater predator loss in small patches. Predator loss, in turn, is predicted to 1) increase herbivory rates overall, and 2) cause herbivores to shift feeding from plants that act as refugia to those that are preferred forage. We tested these predictions in an old‐field community using two experiments. The first was a large‐scale experiment that included hundreds of arthropod species in fragments of various sizes, and used goldenrod and switchgrass to assess herbivory. Our second experiment manipulated densities of a focal predator species and a focal prey species to determine if changes in densities, rather than other characteristics of fragments, were sufficient to cause the trends observed in the first experiment. We found that predator densities declined in small fragments whereas herbivore densities showed the opposite trend. Total herbivory mirrored herbivore densities by increasing in small patches, and this mean increase was driven by large increases in goldenrod herbivory but declines in switchgrass herbivory. Experimental manipulation of densities confirmed that herbivores preferentially feed on goldenrod, and that predators depress herbivory on goldenrod but have a negligible effect on switchgrass. Our results suggest that fragmentation alters trophic interactions by causing declines in predator densities and increases in herbivore densities, but that feeding preferences of herbivores may generate unequal impacts among plant species.     

The role of patch area and habitat diversity in explaining native plant species richness in disturbed suburban forest patches in northern Belgium

Relatively easy measurable patch characteristics (especially habitat diversity measures) have proven to be valuable indicators of forest plant species richness in forest fragments of relatively undisturbed areas. Urban and suburban forest patches, however, are characterized by a specific landscape ecological context implying that specific processes may influence ecosystem functioning and hence that other abiotic indicators for plant diversity are more appropriate. We studied the relation between functional ecological plant species groups and suburban forest patch characteristics such as patch area, habitat diversity and isolation. Some components of species richness were related to the isolation of the patches. In contrast to previous similar large-scale fragmentation studies in more rural areas, further results stressed the overwhelming importance of patch area relative to habitat variables in determining species richness. This suggests (1) the occurrence of density-dependent species extinction processes in small forest patches; or (2) the existence of external deterministic factors which put a major constraint on species richness in small patches. We tend to support the latter hypothesis and propose forest disturbance and associated black cherry (Prunus serotina Ehrh.) invasion as such a possible external factor. Small forest patches may be more sensitive to disturbance and biological invasion due to various reasons. Hence large forest patches are to be preferred for plant conservation in the suburban area.