Regularity of species richness relationships to patch size and shape

This study aims to assess the degree of regularity in the effect of patch size and patch shape on plant species richness across a macroscale region, and to evaluate the implications for nature conservation.
Our study area covers south-eastern Norway and contains 16 agricultural landscapes with 2162 patches.
To analyse regularity a local linear mixed model (LLMM) was applied. This procedure estimates the richness trends due to shared effects of size and shape, and simultaneously provides the landscape-specific random effect. The latter is a direct estimate of the degree of irregularity between the landscapes, conditioned on specific values of size and shape.
The results show a positive interaction between the shape and size of patches, which is repeated for all landscapes. The shape of the patches produces more regular patterns in species richness than the size of patches. This we attribute to effects of dispersal and distance to neighbouring patches of different environmentally conditioned species pools. Large and complex patches have shorter average distance to neighbouring patches (of different types) than large simple-shaped (circular) patches have. We attribute the higher species richness of the former, given a similar area, to a higher number of species dispersed from the outside into the more complex plot. For small patches, however, the distance to the edge is short relative to normal dispersal distances, for patches of all shapes. This explains why the positive effect of shape complexity on species richness is stronger for large patches. This interpretation is supported by a strong spatial correlation conditioned on the most complex patches.
Theories of dynamics in biodiversity in patchy landscapes must consider shape as a regulator at the same level as size, and both shape and size of patches should be simultaneously taken into account for management planning.     

The decoupling of abundance and species richness in lizard communities

1. Patterns of species richness often correlate strongly with measures of energy. The more individuals hypothesis (MIH) proposes that this relationship is facilitated by greater resources supporting larger populations, which are less likely to become extinct. Hence, the MIH predicts that community abundance and species richness will be positively related. 2. Recently, Buckley & Jetz (2010, Journal of Animal Ecology, 79, 358-365) documented a decoupling of community abundance and species richness in lizard communities in south-west United States, such that richer communities did not contain more individuals. They predicted, as a consequence of the mechanisms driving the decoupling, a more even distribution of species abundances in species-rich communities, evidenced by a positive relationship between species evenness and species richness. 3. We found a similar decoupling of the relationship between abundance and species richness for lizard communities in semi-arid south-eastern Australia. However, we note that a positive relationship between evenness and richness is expected because of the nature of the indices used. We illustrate this mathematically and empirically using data from both sets of lizard communities. When we used a measure of evenness, which is robust to species richness, there was no relationship between evenness and richness in either data set. 4. For lizard communities in both Australia and the United States, species dominance decreased as species richness increased. Further, with the iterative removal of the first, second and third most dominant species from each community, the relationship between abundance and species richness became increasingly more positive. 5. Our data support the contention that species richness in lizard communities is not directly related to the number of individuals an environment can support. We propose an alternative hypothesis regarding how the decoupling of abundance and richness is accommodated; namely, an inverse relationship between species dominance and species richness, possibly because of ecological release.     

Bird communities in rainforest fragments: guild responses to habitat variables in Tabasco,Mexico

The effects of habitat loss and fragmentation in tropical forests are difficult to separate, as they usually occur concurrently. In the state park La Sierra, in Tabasco, Mexico, the rainforest is being cleared for pasture, and fragments are being used by local inhabitants. This study examined the response of bird feeding guilds to habitat characteristics, including human disturbance, in five fragments of different sizes (1 ∼ 4,500 ha, 2 ∼ 150 ha, and 2 ∼ 80 ha). Using point count observations, 125 species were recorded and were grouped into 11 feeding guilds. As expected, the largest fragment had higher species richness and abundances than the smaller fragments. However, five habitat features differed significantly among fragment sizes, including tree density, the number of tree stumps and the number of trails. Thus the larger fragment was also less disturbed. Fragment size alone was significant only for scavenger species richness, and for the abundance of bark gleaning insectivores and insectivore/nectarivores. Raptors were more diverse and abundant in the large fragment and less disturbed sites. Arboreal frugivores and bark or foliage gleaning insectivores, depended on higher trees and less disturbed sites. A better understanding of the mechanisms that affect persistence is essential for the planning of conservation actions.     

Patch- and landscape-scale effects on howler monkey distribution and abundance in rainforest fragments

Variation at both the patch and landscape scale is known to influence the distribution and abundance of arboreal monkeys in rainforest fragments. However, few studies have examined the factors associated with these different scales of focus simultaneously. Using stepwise logistic-regression and generalized linear models (GLMs), howler monkey Alouatta palliata distribution and abundance were examined as a function of patch quality (fragment area, shape, tree DBH and canopy height) and landscape connectivity (isolation, total forest area, fragment and road abundance, corridor abundance and length) in 119 rainforest fragments in northern Chiapas, Mexico. The positive correlation observed between monkey distribution (presence/absence) and both fragment area and abundance may be explained by increased resources within larger fragments and those whose proximity allows greater exploration by monkeys. In contrast, GLM analysis indicated that monkey abundance in inhabited fragments was positively correlated with corridor abundance, canopy height and fragment area. These relationships could be explained by greater reproductive investments by monkeys in forest fragments whose size, degree of perturbation and degree of connectivity with other fragments suggest greater overall resource availability. Future studies should explicitly include a multi-scale approach to understanding the factors affecting patterns of monkey distribution and abundance, particularly as this relates to measures of and interactions between patch connectivity and resource availability.     

Response of predators to prey abundance: separating the effects of prey density and patch size

We tested the relative and combined effects of prey density and patch size on the functional response (number of attacks per unit time and duration of attacks) of a predatory reef fish (Cheilodactylus nigripes (Richardson)) to their invertebrate prey. Fish attacked prey at a greater rate and for longer time in large than small patches of prey, but large patches had naturally greater densities of prey. We isolated the effects of patch size and prey density by reducing the density of prey in larger patches to equal that of small patches; thereby controlling for prey density. We found that the intensity at which fish attacked prey (combination of attack rate and duration) was primarily a response to prey density rather than the size of patch they occupied. However, there was evidence that fish spent more time foraging in larger than smaller patches independent of prey density; presumably because of the greater total number of prey available. These experimental observations suggest that fish can distinguish between different notions of prey abundance in ways that enhance their rate of consumption. Although fish may feed in a density dependent manner, a critical issue is whether their rate of consumption outstrips the rate of increase in prey abundance to cause density dependent mortality of prey.     

Species richness in soil bacterial communities: a proposed approach to overcome sample size bias

Estimates of species richness based on 16S rRNA gene clone libraries are increasingly utilized to gauge the level of bacterial diversity within various ecosystems. However, previous studies have indicated that regardless of the utilized approach, species richness estimates obtained are dependent on the size of the analyzed clone libraries. We here propose an approach to overcome sample size bias in species richness estimates in complex microbial communities. Parametric (Maximum likelihood-based and rarefaction curve-based) and non-parametric approaches were used to estimate species richness in a library of 13,001 near full-length 16S rRNA clones derived from soil, as well as in multiple subsets of the original library. Species richness estimates obtained increased with the increase in library size. To obtain a sample size-unbiased estimate of species richness, we calculated the theoretical clone library sizes required to encounter the estimated species richness at various clone library sizes, used curve fitting to determine the theoretical clone library size required to encounter the "true" species richness, and subsequently determined the corresponding sample size-unbiased species richness value. Using this approach, sample size-unbiased estimates of 17,230, 15,571, and 33,912 were obtained for the ML-based, rarefaction curve-based, and ACE-1 estimators, respectively, compared to bias-uncorrected values of 15,009, 11,913, and 20,909.     

Habitat selection determines abundance, richness and species composition of beetles in aquatic communities

Distribution and abundance patterns at the community and metacommunity scale can result from two distinct mechanisms. Random dispersal followed by non-random, site-specific mortality (species sorting) is the dominant paradigm in community ecology, while habitat selection provides an alternative, largely unexplored, mechanism with different demographic consequences. Rather than differential mortality, habitat selection involves redistribution of individuals among habitat patches based on perceived rather than realized fitness, with perceptions driven by past selection. In particular, habitat preferences based on species composition can create distinct patterns of positive and negative covariance among species, generating more complex linkages among communities than with random dispersal models. In our experiments, the mere presence of predatory fishes, in the absence of any mortality, reduced abundance and species richness of aquatic beetles by up to 80% in comparison with the results from fishless controls. Beetle species' shared habitat preferences generated distinct patterns of species richness, species composition and total abundance, matching large-scale field patterns previously ascribed to random dispersal and differential mortality. Our results indicate that landscape-level patterns of distribution and species diversity can be driven to a large extent by habitat selection behaviour, a critical, but largely overlooked, mechanism of community and metacommunity assembly.     

Amphibian and reptile biogeographic regions of Northern Eurasia,mapped separately

Distributions of amphibian and reptile faunas were separately delimited on a 1:20 000 000 vegetation map of Northern Eurasia divided into 245 10-degree-longitudinal segments of native subzone within the USSR borders as of 1990. All reptile and amphibian species recorded in every segment were listed, and the Jaccard indices were calculated, and the similarity matrix was studied with cluster analysis. Hierarchic classifications were made: the amphibian one consisting of 3 faunistic regions, divided into 4 subregions, 7 biologic provinces, and 23 districts. The reptile classification includes 4 faunistic regions, 7 subregions, 18 provinces, and 14 districts. The reptile classification has 1.5 times more provincial and district subdivisions than amphibian one. Environmental factors correlating with faunistic nonuniformity were revealed. Our amphibian and reptile schemes are 1.9 and 3.5 times more informative than those proposed earlier and account for 75 and 91% of variance in the faunal similarity coefficient of specific areas, respectively (multiple correlation coefficients 0.87 and 0.95). Environmental factors can explain 84 and 93% of faunistic nonuniformity (correlation coefficients 0.95 and 0.96).     

Environmental determinants of amphibian and reptile species richness in China

Understanding the factors that regulate geographical variation in species richness has been one of the fundamental questions in ecology for decades, but our knowledge of the cause of geographical variation in species richness remains poor. This is particularly true for herpetofaunas (including amphibians and reptiles). Here, using correlation and regression analyses, we examine the relationship of herpetofaunal species richness in 245 localities across China with 30 environmental factors, which include nearly all major environmental factors that are considered to explain broad-scale species richness gradients in such theories as ambient energy, water–energy dynamics, productivity, habitat heterogeneity, and climatic stability. We found that the species richness of amphibians and reptiles is moderately to strongly correlated with most of the environmental variables examined, and that the best fit models, which include explanatory variables of temperature, precipitation, net primary productivity, minimum elevation, and range in elevation, explain ca 70% the variance in species richness for both amphibians and reptiles after accounting for sample area. Although water and temperature are important explanatory variables to both amphibians and reptiles, water variables explain more variance in amphibian species richness than in reptile species richness whereas temperature variables explain more variance in reptile species richness than in amphibian species richness, which is consistent with different physiological requirements of the two groups of organisms.     

Understory bird communities in Amazonian rainforest fragments: species turnover through 25 years post-isolation in recovering landscapes

Inferences about species loss following habitat conversion are typically drawn from short-term surveys, which cannot reconstruct long-term temporal dynamics of extinction and colonization. A long-term view can be critical, however, to determine the stability of communities within fragments. Likewise, landscape dynamics must be considered, as second growth structure and overall forest cover contribute to processes in fragments. Here we examine bird communities in 11 Amazonian rainforest fragments of 1-100 ha, beginning before the fragments were isolated in the 1980s, and continuing through 2007. Using a method that accounts for imperfect detection, we estimated extinction and colonization based on standardized mist-net surveys within discreet time intervals (1-2 preisolation samples and 4-5 post-isolation samples). Between preisolation and 2007, all fragments lost species in an area-dependent fashion, with loss of as few as <10% of preisolation species from 100-ha fragments, but up to 70% in 1-ha fragments. Analysis of individual time intervals revealed that the 2007 result was not due to gradual species loss beginning at isolation; both extinction and colonization occurred in every time interval. In the last two samples, 2000 and 2007, extinction and colonization were approximately balanced. Further, 97 of 101 species netted before isolation were detected in at least one fragment in 2007. Although a small subset of species is extremely vulnerable to fragmentation, and predictably goes extinct in fragments, developing second growth in the matrix around fragments encourages recolonization in our landscapes. Species richness in these fragments now reflects local turnover, not long-term attrition of species. We expect that similar processes could be operating in other fragmented systems that show unexpectedly low extinction.     

Anodopetalum biglandulosum: Growth form and abundance in Tasmanian rainforest

Abstract Anodopetalum biglandulosum grows in a variety of habits in Tasmanian rainforest. Forty-eight stands were classified to produce a typology of growth forms for the species. The species has a fundamentally deterministic growth form and exhibits a variety of responses to canopy disruption. The responses are related to community floristics, forest structure and the disturbance regime. The roles of canopy proliferation and clonal spread were investigated for their ecological significance. In some forests A. biglandulosum may dominate the regrowth after tree-fall, resulting in a late successional shift in the forest structure.     

The relationship between seed size and abundance in plant communities: model predictions and observed patterns

Recent studies have suggested that seed size and plant abundance in communities are associated. However, inconsistent patterns have emerged from these studies, with varying mechanisms proposed to explain emergent relationships. We employ a theoretical framework, based on key theory lineages of vegetation dynamics and species coexistence, to examine relationships between species abundance and seed size. From these theory lineages, we identified four models and their predictions: the Seed size/number trade‐off model (SSNTM), the Succession model (SM), the Spatial competition model (SCM), and the Lottery model (LM). We then explored empirical evidence from ten diverse plant communities for seed size and abundance patterns, and related these patterns to model predictions. The SSNTM predicts a negative correlation between seed size and abundance. The SM predicts either a negative, positive or no correlation dependent on time since disturbance, while the SCM and LM make no predictions for a relationship between seed size and abundance. We found no evidence for consistent relationships between seed size and abundance across the ten communities. There were no consistent differences in seed size and abundance relationships between communities dominated by annuals compared to perennials. In three of the ten communities a significant positive seed size and abundance correlation emerged, which falsified the SSNTM as an important determinant of abundance structure in these communities. For sites in coastal woodland, the relationships between seed size and abundance were consistent with the predictions of the SM (although generally not significant), with fire being the disturbance. We suggest that the significant positive seed size and abundance correlations found may be driven by the association between large seeds and large growth forms, as large growth forms tend to be dominant. It seems likely that patterns of seed size and abundance in communities are determined by a complex interaction between environmental factors and correlations of plant attributes that determine a species’ strategy.     

Determinants of woody species richness in Scot pine and beech forests: climate, forest patch size and forest structure

We analysed patterns of woody species richness in Pinus sylvestris and Fagus sylvatica forests in Catalonia (NE Spain) from forestry inventory databank in relation to climate and landscape structure. Both types of forests are found within the same climatic range, although they have been managed following somewhat different goals. Overall, woody species richness significantly increased when conditions get closer to the Mediterranean ones, with milder temperatures. Differences between the two types of forests arose when comparing the relationship between richness and forest patch size. Woody species richness increased in pine forests with patch size, while the opposite trend was observed in beech forests. This pattern is explained by the different behaviour of structural canopy properties, since leaf area index and canopy cover showed a steeper increase with increasing forest patch size in Fagus forests than in Pinus ones. Accordingly, richness decreased with canopy cover in Fagus plots, but not in Pinus ones. We suggest that these differences would be related to management history, which may have enhanced the preservation of beech stands in larger forest landscape units.     

Bat species richness and abundance in tropical rain forest fragments and in agricultural habitats at Los Tuxtlas, Mexico

Faced with the rapid and extensive conversion of tropical rain forests to pasture lands and agricultural fields and with the need to preserve the remaining mammalian fauna, it is imperative to determine how the different species that form the mammalian community have responded to the anthropogenic alterations of their natural habitats To provide data in this direction, we sampled bats m 45 forest islands, m 20 agricultural habitats representing five types of vegetation (cocoa, coffee, mixed, citrus and allspice), in four live-fence sites and in four pasture sites at Los Tuxtlas, Veracruz, Mexico Sampling effort resulted in the capture of 2587 bats representing 35 species In forest habitats we detected 32 species We did not capture any bats at the four pasture sites, but the at the other agricultural habitats studied, we captured 38% of the bats and 77% of the species recorded Thirty-four percent of the species recorded were present at the live-fence habitats Isolating distance was an important variable influencing species richness in forests and in agricultural habitats Only 10% of the species recorded occurred m all the habitats studied, but 77% of the species occurred m a habitat other than ram forest Recaptures of bats indicated inter habitat movements in the fragmented landscape We discuss the conservation value for the bat fauna of agricultural islands of vegetation as elements reducing isolating distances among forest fragments     

Effects of herbivory and patch size on tree seedling survivorship in a fog-dependent coastal rainforest in semiarid Chile

The landscape (matrix) surrounding habitat fragments critically affects the biodiversity of those fragments due to biotic interchange and physical effects. However, to date, there have been only a limited number of studies on plant–animal interactions in fragmented landscapes, particularly on how tree seedling herbivory is affected by fragmentation. We have examined this question in a fog-dependent mosaic of rainforest fragments located on coastal mountaintops of semiarid Chile (30°S), where the effects of the surrounding semiarid matrix and forest patch size (0.1–22 ha) on tree seedling survival were simultaneously addressed. The rainforest is strongly dominated by the endemic evergreen tree species Aextoxicon punctatum (Olivillo, approx. 80% of basal area). To assess the magnitudes and causes of Olivillo seedling mortality, we set up a field experiment where 512 tree seedlings of known age were transplanted into four forest fragments of different sizes in four 1.5 × 3-m plots per patch; one-half of each plot was fenced off with chicken wire to exclude small mammals. The plots were monitored for 22 months. Overall, 50% of the plants died during the experiment. The exclusion of small mammals from the plots increased seedling survival by 25%, with the effect being greater in smaller patches where matrix-dwelling herbivores are more abundant. This experiment highlights the important role of the surrounding matrix in affecting the persistence of trees in forest fragments. Because herbivores from the matrix cause greater tree seedling mortality in small patches, their effects must be taken into account in forest conservation–restoration plans.     

A test of species–energy theory: patch occupancy and colony size in tropical rainforest litter‐nesting ants

Species–energy theory can account for spatial variation in the abundance and community composition of animals, though the mechanisms of species–energy theory are under contention. We evaluated three competing mechanisms at the local spatial scale by conducting an in vivo light manipulation over supplemental ant nests placed in the leaf litter of a Costa Rican tropical rainforest. We found that the light environment did not alter the 10% rate of occupation of the supplemental nests, but light did alter the size of colonies and the genus‐level composition of the community. Light levels in the foraging range were positively associated with colony sizes of all ants, whereas light levels directly on the nest site were predictive of the occurrence of ant genera. Colonies of specialized predators, dacetine ants, were larger in more shaded foraging environments, and the functional group of generalized myrmicines exhibited an opposite pattern, with smaller‐sized colonies in response to shading. Responses of twig‐dwelling ants to the light environment were most consistent with the metabolic cost hypothesis as a mechanism of species–energy theory. We found mixed support for the thermal energy availability hypothesis, and scant support for the chemical energy hypothesis, as the litter depth, a measure of prey density, was not predictive of ant responses. In summary, at the local scale, we found patterns in colony size and life history are governed by light‐dependent mechanisms.     

Enrichment planting to improve habitat quality and conservation value of tropical rainforest fragments

Many areas of tropical rainforest have been fragmented and the habitat quality of fragments is often poor. For example, on Borneo, many forest fragments are highly degraded by repeated logging of Dipterocarpaceae trees prior to fragmentation, and we examined the viability of enrichment planting as a potential management tool to enhance the conservation value of these forest fragments. We planted seedlings of three dipterocarp species with contrasting light demands and tolerances (Parashorea malaanonan (light demander), Dryobalanops lanceolata (intermediate), Hopea nervosa (shade tolerant)) in eight forest fragment sites (3–3529 ha), and compared seedling performance with four sites in continuous forest. Eighteen months after planting, survival rates of seedlings were equally high in fragment sites (mean survival = 63 %), and in continuous forest sites (mean survival = 68 %). By contrast, seedling growth and herbivory rates were considerably higher in fragments (by 60 % for growth and 45 % for herbivory) associated with higher light environments in degraded forest fragments compared with continuous forest sites. Among the three study species, H. nervosa seedlings had the highest survival rates overall, and P. malaanonan seedlings generally grew fastest and suffered highest herbivory rates. There were no interactions between species performance and the effects of fragment site area, forest structure or soil characteristics of sites suggesting that the three species responded similarly to fragmentation effects. High survival of planted seedlings implies that enrichment planting would be a successful forest management strategy to improve forest quality, and hence conservation value, of fragments.     

Effects of patch size and position above the substratum during early succession of subtidal soft-bottom communities

Early macrobenthos succession in small, disturbed patches on subtidal soft bottoms is facilitated by the arrival of post-larval colonizers, in particular by active and passive dispersers along the seafloor or through the water column. Using a field experiment at two contrasting sites (protected vs. exposed to wave action), we evaluated the role of (a) active and passive dispersal through the water column and (b) the influence of small-scale spatial variability during succession of subtidal macrobenthic communities in northern Chile. Containers of two sizes (surface area: small—0.12?m² and large—0.28?m²) at two positions above the natural substratum (height: low—3?cm and high—26?cm) were filled with defaunated sediment, installed at two sandy sublittoral sites (7–9?m water depth) and sampled after 7, 15, 30, 60 and 90?days, together with the natural bottom sediment. The experiment took place during austral fall (from late March to early July 2010), when both larval and post-larval stages are abundant. At the exposed site, early succession was driven by similar proportions of active and passive dispersers. A sequence from early, late and reference communities was also evident, but container position and size affected the proportional abundance of dispersal types. At the protected site, the successional process started with abundant colonization of active dispersers, but toward the end of the experiment, the proportion of swimmer/crawlers increased, thus resembling the dispersal types found in the natural community. At this site, the position above the sediment affected the proportional abundance of dispersal types, but patch size had no effect. This study highlights that macrobenthic post-larvae can reach at least 26?cm high above the bottom (actively or passively, depending on site exposure), thus playing an important role during early succession of sublittoral soft bottoms. The active or passive use of the sediment–water interphase may also play an important role in the connectivity of benthic populations and in the recovery after large-scale disturbances of sublittoral habitats.     

Heathland management effects on carabid beetle communities: the relationship between bare ground patch size and carabid biodiversity

Carabid beetles were sampled by pitfall trapping on Brentmoor Heath, Surrey, UK during the summers of 2009 and 2010 to determine the effects of bare ground patch size on beetle abundance, richness and diversity. Four patch sizes were investigated: 1, 4, 25 and 100?m² as well as the adjacent mown and unmown areas. A range of environmental parameters relating to soil characteristics, stones and nearby vegetation were measured at each patch and control habitat in order to distinguish the effect of patch size. Results show that bare ground is a valuable habitat for carabids, but that the response of their abundance, richness and diversity to bare ground patch size depends on a large number of environmental variables on and around the patches. When all variables are taken into account, smaller patches appear to benefit carabid abundance, richness and diversity. In the presence of Molinia caerulea at the patch edges, however, larger patches were more beneficial. Given this dependence on environmental variables, the perfect patch size for conservation of biodiversity is likely to be site specific and the best approach may be to use a variety of patch sizes at a range of successional stages.     

Mammal density and patterns of ectoparasite species richness and abundance

Patterns of species richness, prevalence and abundance of ectoparasites have rarely been investigated at both the levels of populations and species of hosts. Here, we investigated the effects in changes in small mammal density on species richness, abundance and prevalence of ectoparasitic fleas. The comparative analyses were conducted for different small mammal species and among several populations during a long-term survey. We tested the hypothesis that an increase in host density should be linked with an increase in parasite species richness both among host species and among populations within host species, as predicted by epidemiological models. We also used host species density data from literature. We found that host density has a major influence on the species richness of ectoparasite communities of small mammals among host populations. We found no relationship between data of host density from the literature and parasite species richness. In contrast with epidemiological hypotheses, we found no relationships between abundance, or prevalence, and host density, either among host species or among host populations. Moreover, a decrease in abundance of fleas in relation with an increase in host density was observed for two mammal species (Apodemus agrarius and A. flavicollis). The decrease or the lack of increase in flea abundance in relation with an increase in host density suggests anti-parasitic behavioural activities such as grooming.