Movement trajectories and habitat partitioning of small mammals in logged and unlogged rain forests on Borneo

1. Non-volant animals in tropical rain forests differ in their ability to exploit the habitat above the forest floor and also in their response to habitat variability. It is predicted that specific movement trajectories are determined both by intrinsic factors such as ecological specialization, morphology and body size and by structural features of the surrounding habitat such as undergrowth and availability of supportive structures. 2. We applied spool-and-line tracking in order to describe movement trajectories and habitat segregation of eight species of small mammals from an assemblage of Muridae, Tupaiidae and Sciuridae in the rain forest of Borneo where we followed a total of 13,525 m path. We also analysed specific changes in the movement patterns of the small mammals in relation to habitat stratification between logged and unlogged forests. Variables related to climbing activity of the tracked species as well as the supportive structures of the vegetation and undergrowth density were measured along their tracks. 3. Movement patterns of the small mammals differed significantly between species. Most similarities were found in congeneric species that converged strongly in body size and morphology. All species were affected in their movement patterns by the altered forest structure in logged forests with most differences found in Leopoldamys sabanus. However, the large proportions of short step lengths found in all species for both forest types and similar path tortuosity suggest that the main movement strategies of the small mammals were not influenced by logging but comprised generally a response to the heterogeneous habitat as opposed to random movement strategies predicted for homogeneous environments. 4. Overall shifts in microhabitat use showed no coherent trend among species. Multivariate (principal component) analysis revealed contrasting trends for convergent species, in particular for Maxomys rajah and M. surifer as well as for Tupaia longipes and T. tana, suggesting that each species was uniquely affected in its movement trajectories by a multiple set of environmental and intrinsic features.     

Fine‐scale habitat preferences and habitat partitioning by three mycophagous mammals in tropical wet sclerophyll forest,north‐eastern Australia

Abstract Fine‐scale habitat preferences of three co‐occurring mycophagous mammals were examined in a tropical wet sclerophyll forest community in north‐eastern Australia. Two of the three mammal species responded to fine‐scale variation in vegetation and landform around individual trap locations. At a broad scale, the northern bettong (Bettongia tropica), an endangered marsupial endemic to the Australian wet tropics region, showed a preference for ridges over mid‐slopes and gullies, irrespective of forest type. In contrast, the northern brown bandicoot (Isoodon macrourus), a widespread marsupial, displayed a preference for Eucalyptus woodland over adjacent Allocasuarina forest, irrespective of topographic category. The giant white‐tailed rat (Uromys caudimaculatus), a rodent endemic to the wet tropics, showed no particular preference for either forest type or topographic category. A multiple regression model of mammal capture success against three principal habitat gradients constructed from 21 habitat variables using principal component analysis indicated strong species‐specific preferences for fine‐scale vegetation assemblages. Bettongs preferred areas of Eucalyptus woodland with sparse ground cover, low densities of certain grass species, high density of tree stems and few pig diggings. Bandicoots, in contrast, favoured areas in both forest types with dense ground cover, fewer tree stems and greater numbers of pig diggings; that is, characteristics least favoured by bettongs. The striking differences in fine‐scale habitat preferences of these two mammals of similar body size and broad habitat requirements suggest a high degree of fine‐scale habitat partitioning. White‐tailed rats did not show preference for any of the habitat gradients examined.     

Beta‐diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly

Site‐to‐site variation in species composition (β‐diversity) generally increases from low‐ to high‐diversity regions. Although biogeographical differences in community assembly mechanisms may explain this pattern, random sampling effects can create this pattern through differences in regional species pools. Here, we compared assembly mechanisms between spatially extensive networks of temperate and tropical forest plots with highly divergent species pools (46 vs. 607 species). After controlling for sampling effects, β‐diversity of woody plants was similar and higher than expected by chance in both forests, reflecting strong intraspecific aggregation. However, different mechanisms appeared to explain aggregation in the two forests. In the temperate forest, aggregation reflected stronger environmental correlations, suggesting an important role for species‐sorting (e.g. environmental filtering) processes, whereas in the tropics, aggregation reflected stronger spatial correlations, more likely reflecting dispersal limitation. We suggest that biogeographical differences in the relative importance of different community assembly mechanisms contribute to these striking gradients in global biodiversity.     

Non‐interacting impacts of fertilization and habitat area on plant diversity via contrasting assembly mechanisms

Aim

The local‐ and regional‐based forms of anthropogenic change reducing grassland diversity are generally identified, but these scale‐dependent processes tend to co‐occur with unclear interactive effects. Here, we explicitly test how common local and regional perturbations simultaneously affect plant alpha and beta diversity in a multiyear community assembly experiment using fragments of grassland habitat of various sizes. We hypothesized that local disturbances and decreasing patch size would interact, suppressing local diversity while homogenizing composition among patches.

Location

North America.

Methods

We conducted a three‐year grassland assembly experiment, factorially manipulating local perturbation (nitrogen addition and mowing) and patch area for 36 patches over 13 ha. We quantified the individual and interactive effects of these local and regional factors on plant alpha and beta diversity within (quadrat scale) and among patches (patch scale). We also used a null model approach to disentangle between stochastic‐ and niche‐based assembly mechanisms.

Results

We detected a gradient of assembly outcomes driven by two non‐interacting factors—the effects of N fertilization on alpha (negative) and beta (positive) diversity regardless of spatial scale and the scale‐dependant effect of increasing patch size on alpha (positive) and beta (positive) diversity. These effects unfolded over time, with the constraints on richness and composition shifting from dispersal‐based during the first sampling year to perturbation‐and size‐based factors at year two and three. Fertilization effects were driven by a mixture of deterministic (i.e., selection at the species level) and stochastic (i.e., random extinctions) processes resulting in a decline in local richness but an increase in spatial heterogeneity in species composition. Area appeared to influence alpha diversity mainly via stochastic “sampling effect”—larger patches represented a larger sample of the regional pool. Niche‐based processes, however, led to convergence in beta diversity among smaller patches driving a positive overall effect of area on beta diversity.

Main conclusion

Our results illustrate how diversity regulation in contemporary grasslands can be simultaneously shaped by local and regional factors acting additively but via contrasting assembly mechanisms that operate at different spatial and temporal scales.

     

Species turnover (β‐diversity) in ectomycorrhizal fungi linked to uptake capacity

Ectomycorrhizal (EcM) fungal communities may be shaped by both deterministic and stochastic processes, potentially influencing ecosystem development and function. We evaluated community assembly processes for EcM fungi of Pseudotsuga menziesii among 12 sites up to 400 km apart in southwest British Columbia (Canada) by investigating species turnover (β‐diversity) in relation to soil nitrogen (N) availability and physical distance. We then examined functional traits for an N‐related niche by quantifying net fluxes of , and protons on excised root tips from three contrasting sites using a microelectrode ion flux measurement system. EcM fungal communities were well aligned with soil N availability and pH, with no effect of site proximity (distance–decay curve) on species assemblages. Species turnover was significant (β_1/2 = 1.48) along soil N gradients, with many more Tomentella species on high N than low N soils, in contrast to Cortinarius species. Ammonium uptake was greatest in the spring on the medium and rich sites and averaged over 190 nmol/m²/s for Tomentella species. The lowest uptake rates of were by nonmycorrhizal roots of axenically grown seedlings (10 nmol/m²/s), followed by Cortinarius species (60 nmol/m²/s). EcM roots from all sites displayed only marginal uptake of nitrate (8.3 nmol/m²/s). These results suggest uptake capacity is an important functional trait influencing the assembly of EcM fungal communities. The diversity of EcM fungal species across the region arguably provides critical belowground adaptations to organic and inorganic N supply that are integral to temperate rainforest ecology.     

Common ancestry or environmental trait filters: cross‐continental comparisons of trait–habitat relationships in tropical anuran amphibian assemblages

Aim To investigate whether trait–habitat relations in biological communities converge across three global regions. The goal is to assess the role of habitat templets in shaping trait assemblages when different assembly mechanisms are operating and to test whether trait–habitat relations reflect a common evolutionary history or environmental trait filters. Location Guiana Shield, South America; Upper Guinea Forest Block, West Africa; Borneo rain forests, Southeast Asia. Methods We compared large anuran amphibian data sets at both the regional and cross‐continental scale. We applied a combination of three‐table ordinations (RLQ) and permutation model‐based multivariate fourth‐corner statistics to test for trait–habitat relationships at both scales and used phylogenetic comparative methods to quantify phylogenetic signal in traits that enter these analyses. Results Despite the existence of significant trait–habitat links and congruent trait patterns, we did not find evidence for the existence of a universal trait–habitat relationship at the assemblage level and no clear sign for cross‐continental convergence of trait–habitat relations. Patterns rather varied between continents. Despite the fact that a number of traits were conserved across phylogenies, the phylogenetic signal varied between regions. Trait–habitat relations therefore not only reflect a common evolutionary history, but also more recently operating environmental trait filters that ultimately determine the trait composition in regional assemblages. Main conclusions Integrating trait–habitat links into analyses of biological assemblages can enhance the predictive power and general application of species assembly rules in community and macroecology, particularly when phylogenetic comparative methods are simultaneously applied. However, in order to predict trait composition based on habitat templets, trait–habitat links cannot be assumed to be universal but rather have to be individually established in different regions prior to model building. Only then can direct trait‐based approaches be useful tools for predicting fundamental community patterns.     

Testing and interpreting the shared space‐environment fraction in variation partitioning analyses of ecological data

Variation partitioning analyses combined with spatial predictors (Moran's eigenvector maps, MEM) are commonly used in ecology to test the fractions of species abundance variation purely explained by environment and space. However, while these pure fractions can be tested using a classical residuals permutation procedure, no specific method has been developed to test the shared space‐environment fraction (SSEF). Yet, the SSEF is expected to encompass a major driver of community assembly, that is, an induced spatial dependence effect (ISD; i.e. the reflection of a spatially structured habitat filter on a species distribution). A reliable test of this fraction is therefore crucial to properly test the presence of an ISD on ecological data. To bridge the gap, we propose to test the SSEF through spatially‐constrained null models: torus‐translations, and Moran spectral randomisations. We investigated the type I error rate and statistical power of our method based on two real environmental datasets and simulations of tree distributions. Ten types of tree distribution displaying contrasted aggregation properties were simulated, and their abundances were sampled in 153 regularly‐distributed 20 × 20 m quadrats. The SSEF was tested for 1000 simulated tree distributions either unrelated to the environment, or filtered by environmental variables displaying contrasting spatial structures. The method proposed provided a correct type I error rate (< 0.05). The statistical power was high (> 0.9) when abundances were filtered by an environmental variable structured at broad scale. However, the spatial resolution allowed by the sampling design limited the power of the method when using a fine‐scale filtering variable. This highlighted that an ISD can be properly detected providing that the spatial pattern of the filtering process is correctly captured by the sampling design of the study. An R function to apply the SSEF testing method is provided and detailed in a tutorial.     

Intraspecific trait variation increases species diversity in a trait‐based grassland model

Intraspecific trait variation (ITV) is thought to play a significant role in community assembly, but the magnitude and direction of its influence are not well understood. Although it may be critical to better explain population persistence, species interactions, and therefore biodiversity patterns, manipulating ITV in experiments is challenging. We therefore incorporated ITV into a trait‐ and individual‐based model of grassland community assembly by adding variation to the plants’ functional traits, which then drive life‐history tradeoffs. Varying the amount of ITV in the simulation, we examine its influence on pairwise‐coexistence and then on the species diversity in communities of different initial sizes. We find that ITV increases the ability of the weakest species to invade most, but that this effect does not scale to the community level, where the primary effect of ITV is to increase the persistence and abundance of the competitively‐average species. Diversity of the initial community is also of critical importance in determining ITV's efficacy; above a threshold of interspecific diversity, ITV does not increase diversity further. For communities below this threshold, ITV mainly helps to increase diversity in those communities that would otherwise be low‐diversity. These findings suggest that ITV actively maintains diversity by helping the species on the margins of persistence, but mostly in habitats of relatively low alpha and beta diversity.     

Co‐occurrence patterns in a diverse arboreal ant community are explained more by competition than habitat requirements

A major goal of community ecology is to identify the patterns of species associations and the processes that shape them. Arboreal ants are extremely diverse and abundant, making them an interesting and valuable group for tackling this issue. Numerous studies have used observational data of species co‐occurrence patterns to infer underlying assembly processes, but the complexity of these communities has resulted in few solid conclusions. This study takes advantage of an observational dataset that is unusually well‐structured with respect to habitat attributes (tree species, tree sizes, and vegetation structure), to disentangle different factors influencing community organization. In particular, this study assesses the potential role of interspecific competition and habitat selection on the distribution patterns of an arboreal ant community by incorporating habitat attributes into the co‐occurrence analyses. These findings are then contrasted against species traits, to explore functional explanations for the identified community patterns. We ran a suite of null models, first accounting only for the species incidence in the community and later incorporating habitat attributes in the null models. We performed analyses with all the species in the community and then with only the most common species using both a matrix‐level approach and a pairwise‐level approach. The co‐occurrence patterns did not differ from randomness in the matrix‐level approach accounting for all ant species in the community. However, a segregated pattern was detected for the most common ant species. Moreover, with the pairwise approach, we found a significant number of negative and positive pairs of species associations. Most of the segregated associations appear to be explained by competitive interactions between species, not habitat affiliations. This was supported by comparisons of species traits for significantly associated pairs. These results suggest that competition is the most important influence on the distribution patterns of arboreal ants within the focal community. Habitat attributes, in contrast, showed no significant influence on the matrix‐wide results and affected only a few associations. In addition, the segregated pairs shared more biological characteristic in common than the aggregated and random ones.     

Larval habitat for the avian malaria vector Culex quinquefasciatus (Diptera: Culicidae) in altered mid‐elevation mesic‐dry forests in Hawai'i

Effective management of avian malaria (Plasmodium relictum) in Hawai'i's endemic honeycreepers (Drepanidinae) requires the identification and subsequent reduction or treatment of larval habitat for the mosquito vector, Culex quinquefasciatus (Diptera: Culicidae). We conducted ground surveys, treehole surveys, and helicopter aerial surveys from 2001–2003 to identify all potential larval mosquito habitat within two 100+ ha mesic‐dry forest study sites in Hawai'i Volcanoes National Park, Hawai'i; ‘Ainahou Ranch and Mauna Loa Strip Road. At ‘Ainahou Ranch, anthropogenic sites (43%) were more likely to contain mosquitoes than naturally occurring (8%) sites. Larvae of Cx. quinquefasciatus were predominately found in anthropogenic sites while Aedes albopictus larvae occurred less frequently in both anthropogenic sites and naturally‐occurring sites. Additionally, moderate‐size (≈20–22,000 liters) anthropogenic potential larval habitat had >50% probability of mosquito presence compared to larger‐ and smaller‐volume habitat (<50%). Less than 20% of trees surveyed at ‘Ainahou Ranch had treeholes and few mosquito larvae were detected. Aerial surveys at ‘Ainahou Ranch detected 56% (95% CI: 42–68%) of the potential larval habitat identified in ground surveys. At Mauna Loa Strip Road, Cx. quinquefasciatus larvae were only found in the rock holes of small intermittent stream drainages that made up 20% (5 of 25) of the total potential larval habitat. The volume of the potential larval habitat did not influence the probability of mosquito occurrence at Mauna Loa Strip Road. Our results suggest that Cx. quinquefasciatus abundance, and subsequently avian malaria, may be controlled by larval habitat reduction in the mesic‐dry landscapes of Hawai'i where anthropogenic sources predominate.     

Using presence‐only modelling to predict Asian elephant habitat use in a tropical forest landscape: implications for conservation

Aim Asian elephants, Elephas maximus, are threatened throughout their range by a combination of logging, large scale forest conversion and conflict with humans. We investigate which environmental factors, both biotic and abiotic, constrain the current distribution of elephants. A spatially explicit habitat model is constructed to find core areas for conservation and to assess current threats. Location Ulu Masen Ecosystem in the province of Nanggroe Aceh Darussalam on the island of Sumatra, Indonesia. Methods A stratified survey was conducted at 12 sites (300 transects) to establish the presence of elephants. Presence records formed the basis to model potential habitat use. Ecological niche factor analysis (ENFA) is used to describe their niche and to identify key factors shaping elephant distribution. An initial niche model was constructed to describe elephant niche structure, and a second model focused on identifying core areas only. To assess the threat of habitat encroachment, overlap between the elephants’ optimal niche and the occurrence of forest encroachment is computed. Results Elephants were recorded throughout the study area from sea level to 1600 m a.s.l. The results show that the elephant niche and consequently habitat use markedly deviates from the available environment. Elephant presence was positively related to forest cover and vegetation productivity, and elephants were largely confined to valleys. A spatially explicit model showed that elephants mainly utilize forest edges. Forest encroachment occurs throughout the elephants range and was found within 80% of the elephants’ ecological niche. Main conclusions In contrast to general opinion, elephant distribution proved to be weakly constrained by altitude, possibly because of movement routes running through mountainous areas. Elephants were often found to occupy habitat patches in and near human‐dominated areas. This pattern is believed to reflect the displacement of elephants from their former habitat.     

On‐farm habitat restoration counters biotic homogenization in intensively managed agriculture

To slow the rate of global species loss, it is imperative to understand how to restore and maintain native biodiversity in agricultural landscapes. Currently, agriculture is associated with lower spatial heterogeneity and turnover in community composition (β‐diversity). While some techniques are known to enhance α‐diversity, it is unclear whether habitat restoration can re‐establish β‐diversity. Using a long‐term pollinator dataset, comprising ～9,800 specimens collected from the intensively managed agricultural landscape of the Central Valley of California, we show that on‐farm habitat restoration in the form of native plant ‘hedgerows’, when replicated across a landscape, can boost β‐diversity by approximately 14% relative to unrestored field margins, to levels similar to some natural communities. Hedgerows restore β‐diversity by promoting the assembly of phenotypically diverse communities. Intensively managed agriculture imposes a strong ecological filter that negatively affects several important dimensions of community trait diversity, distribution, and uniqueness. However, by helping to restore phenotypically diverse pollinator communities, small‐scale restorations such as hedgerows provide a valuable tool for conserving biodiversity and promoting ecosystem services.     

Species–area curve and distance–decay relationships indicate habitat thresholds of ectomycorrhizal fungi in an old‐growth Pseudotsuga menziesii landscape

Aim

Ectomycorrhizal fungi (EMF) are a diverse and essential biota of forests that are vulnerable to species loss through reductions in late‐seral habitat. We examined how the spatial ecology of this biota, particularly distance–decay and species–area relationships, could better inform habitat thresholds for EMF conservation planning.

Location

Southeast Vancouver Island near Victoria, British Columbia, Canada.

Methods

Using a stratified sampling design, 11 plots (0.15 ha in size) were established at 0.05–17.5 km apart across 2,800 ha of mesic old‐growth Pseudotsuga menziesii var. menziesii and Tsuga heterophylla forests. EMF communities were compiled through molecular analysis of root tips and sporocarps.

Results

The EMF community was comprised of many Cortinarius, Piloderma, Russula and Tricholoma species typical of mesotrophic habitat. A total of 238 EMF species were observed, of which 86 species were detected only once. The ratio of average species richness per plot (84 taxa) to total richness was low at 0.35, and inherent stochasticity of the EMF community was estimated to be 31% community dissimilarity for species incidence. Distance decay of EMF communities was nonlinear, with an estimated slope break at 2.6 km, followed by a largely unchanging trend in β‐diversity. Accumulated species–area curves were fitted best by the cumulative Weibull sigmoid model, and the asymptote (367 species) at approx. 50 ha was consistent with nonparametric estimates of γ‐diversity (342–362 spp.).

Main conclusions

Old‐growth forests host an impressive amount of EMF diversity, and many of the Ramaria, Inocybe and Russula species are likely to be endemic to the Pacific Northwest. Both niche‐ and neutral‐based processes influenced EMF community composition, resulting in a minimum threshold of 50 ha (1.8% of the sample area) for capturing γ‐diversity. These spatial patterns will help design and evaluate conservation efforts, such as retention forestry, to sustain fully diverse EMF communities over managed landscapes.