Agriculturally altered vegetation, especially oil-palm plantations, is rapidly increasing in Southeast Asia. Low species diversity is associated with this commodity, but data on anuran diversity in oil-palm plantations are lacking. We investigated how anuran biological diversity differs between forest and oil-palm plantation, and whether observed differences in biological diversity of these areas is linked to specific environmental factors. We hypothesized that biological diversity is lower in plantations and that plantations support a larger proportion of disturbance-tolerant species than forest. We compared species richness, abundance, and community composition between plantation and forest areas and between site types within plantation and forest (forest stream vs. plantation stream, forest riparian vs. plantation riparian, forest terrestrial vs. plantation terrestrial). Not all measures of biological diversity differed between oil-palm plantations and secondary forest sites. Anuran community composition, however, differed greatly between forest and plantation, and communities of anurans in plantations contained species that prosper in disturbed areas. Although plantations supported large numbers of breeding anurans, we concluded the community consisted of common species that were of little conservation concern (commonly found species include Fejervarya limnocharis, Microhyla heymonsi, and Hylarana erythrea). We believe that with a number of management interventions, oil-palm plantations can provide habitat for species that dwell in secondary forests.
The diversity and abundance of macroinvertebrate shredders were investigated in 52 forested streams (local scale) from nine catchments (regional scale) covering a large area of peninsular Malaysia. A total of 10,642 individuals of aquatic macroinvertebrates were collected, of which 18.22% were shredders. Biodiversity of shredders was described by alpha (αaverage), beta (β) and gamma diversity (γ) measures. We found high diversity and abundance of shredders in all catchments, represented by 1,939 individuals (range 6-115 and average per site of 37.29±3.48 SE) from 31 taxa with 2-13 taxa per site (αaverage=6.98±0.33 SE) and 10-15 taxa per catchment (γ=13.33±0.55 SE). At the local scale, water temperature, stream width, depth and altitude were correlated significantly with diversity (Adj-R2=0.205). Meanwhile, dissolved oxygen, stream velocity, water temperature, stream width and altitude were correlated to shredder abundance (Adj-R2=0.242). At regional scale, however, water temperature was correlated negatively with β and γ diversity (r2=0.161 and 0.237, respectively) as well as abundance of shredders (r2=0.235). Canopy cover was correlated positively with β diversity (r2=0.378) and abundance (r2=0.266), meanwhile altitude was correlated positively with β (quadratic: r2=0.175), γ diversity (quadratic: r2=0.848) as well as abundance (quadratic: r2=0.299). The present study is considered as the first report describing the biodiversity and abundance of shredders in forested headwater streams across a large spatial scale in peninsular Malaysia. We concluded that water temperature has a negative effect while altitude showed a positive relationship with diversity and abundance of shredders. However, it was difficult to detect an influence of canopy cover on shredder diversity.
The exceptionally high species richness of arthropods in tropical rainforests hinges on the complexity of the forest itself: that is, on features such as the high plant diversity, the layered nature of the canopy and the abundance and the diversity of epiphytes and litter. We here report on one important, but almost completely neglected, piece of this complex jigsaw-the intricate network of rhizomorph-forming fungi that ramify through the vegetation of the lower canopy and intercept falling leaf litter. We show that this litter-trapping network is abundant and intercepts substantial amounts of litter (257.3 kg ha(-1)): this exceeds the amount of material recorded in any other rainforest litter-trapping system. Experimental removal of this fungal network resulted in a dramatic reduction in both the abundance (decreased by 70.2 ± 4.1%) and morphospecies richness (decreased by 57.4 ± 5.1%) of arthropods. Since the lower canopy levels can contain the highest densities of arthropods, the proportion of the rainforest fauna dependent on the fungal networks is likely to be substantial. Fungal litter-trapping systems are therefore a crucial component of habitat complexity, providing a vital resource that contributes significantly to rainforest biodiversity.
Selective logging is one of the major drivers of tropical forest degradation, causing important shifts in species composition. Whether such changes modify interactions between species and the networks in which they are embedded remain fundamental questions to assess the 'health' and ecosystem functionality of logged forests. We focus on interactions between lianas and their tree hosts within primary and selectively logged forests in the biodiversity hotspot of Malaysian Borneo. We found that lianas were more abundant, had higher species richness, and different species compositions in logged than in primary forests. Logged forests showed heavier liana loads disparately affecting slow-growing tree species, which could exacerbate the loss of timber value and carbon storage already associated with logging. Moreover, simulation scenarios of host tree local species loss indicated that logging might decrease the robustness of liana-tree interaction networks if heavily infested trees (i.e. the most connected ones) were more likely to disappear. This effect is partially mitigated in the short term by the colonization of host trees by a greater diversity of liana species within logged forests, yet this might not compensate for the loss of preferred tree hosts in the long term. As a consequence, species interaction networks may show a lagged response to disturbance, which may trigger sudden collapses in species richness and ecosystem function in response to additional disturbances, representing a new type of 'extinction debt'.
Selective logging is the most prevalent land-use change in the tropics. Despite the resulting degradation of forest structure, selectively logged forests still harbor a substantial amount of biodiversity leading to suggestions that their protection is the next best alternative to conserving primary, old-growth forests. Restoring carbon stocks under Reducing Emissions from Deforestation and Forest Degradation (REDD+) schemes is a potential method for obtaining funding to protect logged forests, via enrichment planting and liberation cutting of vines. This study investigates the impacts of restoring logged forests in Borneo on avian phylogenetic diversity, the total evolutionary history shared across all species within a community, and on functional diversity, with important implications for the protection of evolutionarily unique species and the provision of many ecosystem services. Overall and understorey avifaunal communities were studied using point count and mist netting surveys, respectively. Restoration caused a significant loss in phylogenetic diversity and MPD (mean pairwise distance) leaving an overall bird community of less total evolutionary history and more closely related species compared to unlogged forests, while the understorey bird community had MNTD (mean nearest taxon distance) that returned toward the lower levels found in a primary forest, indicating more closely related species pairs. The overall bird community experienced a significant loss of functional strategies and species with more specialized traits in restored forests compared to that of unlogged forests, which led to functional clustering in the community. Restoration also led to a reduction in functional richness and thus niches occupied in the understorey bird community compared to unlogged forests. While there are additional benefits of restoration for forest regeneration, carbon sequestration, future timber harvests, and potentially reduced threat of forest conversion, this must be weighed against the apparent loss of phylogenetic and functional diversity from unlogged forest levels, making the biodiversity-friendliness of carbon sequestration schemes questionable under future REDD+ agreements. To reduce perverse biodiversity outcomes, it is important to focus restoration only on the most degraded areas or at reduced intensity where breaks between regimes are incorporated.
Selective logging is the dominant form of human disturbance in tropical forests, driving changes in the abundance of vertebrate and invertebrate populations relative to undisturbed old-growth forests. A key unresolved question is understanding which physiological mechanisms underlie different responses of species and functional groups to selective logging. Regulation of oxidative status is thought to be one major physiological mechanism underlying the capability of species to cope with environmental changes. Using a correlational cross-sectional approach, we compared a number of oxidative status markers among 15 understorey bird species in unlogged and selectively logged forest in Borneo in relation to their feeding guild. We then tested how variation of markers between forest types was associated with that in population abundance. Birds living in logged forests had a higher activity of the antioxidant enzyme superoxide dismutase and a different regulation of the glutathione cycle compared to conspecific birds in unlogged forest. However, neither oxidative damage nor oxidized glutathione differed between forest types. We also found that omnivores and insectivores differed significantly in all markers related to the key cellular antioxidant glutathione irrespective of the forest type. Species with higher levels of certain antioxidant markers in a given type of forest were less abundant in that forest type compared to the other. Our results suggest that there was little long-term effect of logging (last logging rotation occurred ~15 years prior to the study) on the oxidative status of understorey bird species. However, it is unclear if this was owing to plasticity or evolutionary change. Our correlative results also point to a potential negative association between some antioxidants and population abundance irrespective of the forest type.
The distribution patterns of tropical ectomycorrhizal (ECM) fungi along altitudinal gradients remain largely unknown. Furthermore, despite being an iconic site for biodiversity research, virtually nothing is known about the diversity and spatial patterns of fungi on Mt Kinabalu and neighbouring mountain ranges. We carried out deep DNA sequencing of soil samples collected between 425 and 4000 m above sea level to compare richness and community composition of ECM fungi among altitudinal forest types in Borneo. In addition, we tested whether the observed patterns are driven by habitat or by geometric effect of overlapping ranges of species (mid-domain effect). Community composition of ECM fungi was strongly correlated with elevation. In most genera, richness peaked in the mid-elevation montane forest zone, with the exception of tomentelloid fungi, which showed monotonal decrease in richness with increasing altitude. Richness in lower-mid- and mid-elevations was significantly greater than predicted under the mid-domain effect model. We provide the first insight into the composition of ECM fungal communities and their strong altitudinal turnover in Borneo. The high richness and restricted distribution of many ECM fungi in the montane forests suggest that mid-elevation peak richness is primarily driven by environmental characteristics of this habitat and not by the mid-domain effect.
Functional diversity is an integrative approach to better understand biodiversity across space and time. In the present study, we investigated the spatiotemporal patterns (i.e., elevation and season) and environmental determinants of anuran functional diversity on Tianping Mountain, northwest Hunan, China. Specifically, 10 transects were established from low (300 m a.s.l.) to high (1 492 m a.s.l.) elevations, and anuran communities were sampled in spring, early summer, midsummer, and autumn in 2017. Four functional diversity indices were computed for each transect in each season using ecomorphological functional traits. Our results demonstrated that these indices had contrasting responses to increasing elevations. However, they did not differ significantly among seasons in terms of temporal patterns. Interestingly, the unique spatiotemporal functional diversity patterns were impacted by distinct environmental variables, such as leaf litter cover, water temperature, number of trees, and water conductivity.
Both niche and stochastic dispersal processes structure the extraordinary diversity of tropical plants, but determining their relative contributions has proven challenging. We address this question using airborne imaging spectroscopy to estimate canopy β-diversity for an extensive region of a Bornean rainforest and challenge these data with models incorporating niches and dispersal. We show that remotely sensed and field-derived estimates of pairwise dissimilarity in community composition are closely matched, proving the applicability of imaging spectroscopy to provide β-diversity data for entire landscapes of over 1000 ha containing contrasting forest types. Our model reproduces the empirical data well and shows that the ecological processes maintaining tropical forest diversity are scale dependent. Patterns of β-diversity are shaped by stochastic dispersal processes acting locally whilst environmental processes act over a wider range of scales.
A total of 29 female black flies were captured by a hand net as they swarmed around humans in Tam Dao National Park, Vinh Phuc Province, Vietnam. They included one species of the subgenus Gomphostilbia (Simulium (Gomphostilbia) asakoae Takaoka & Davies) and five species of the subgenus Simulium, of which one species is described as Simulium (Simulium) vietnamense sp. nov. and the other four species (S. (S.) chungi Takaoka & Huang, S. (S.) grossifilum Takaoka & Davies, S. (S.) maenoi Takaoka & Choochote, and S. (S.) rufibasis Brunetti) are newly recorded from Vietnam.
Diversity responses to land-use change are poorly understood at local scales, hindering our ability to make forecasts and management recommendations at scales which are of practical relevance. A key barrier in this has been the underappreciation of grain-dependent diversity responses and the role that β-diversity (variation in community composition across space) plays in this. Decisions about the most effective spatial arrangement of conservation set-aside, for example high conservation value areas, have also neglected β-diversity, despite its role in determining the complementarity of sites. We examined local-scale mammalian species richness and β-diversity across old-growth forest, logged forest, and oil palm plantations in Borneo, using intensive camera- and live-trapping. For the first time, we were able to investigate diversity responses, as well as β-diversity, at multiple spatial grains, and across the whole terrestrial mammal community (large and small mammals); β-diversity was quantified by comparing observed β-diversity with that obtained under a null model, in order to control for sampling effects, and we refer to this as the β-diversity signal. Community responses to land use were grain dependent, with large mammals showing reduced richness in logged forest compared to old-growth forest at the grain of individual sampling points, but no change at the overall land-use level. Responses varied with species group, however, with small mammals increasing in richness at all grains in logged forest compared to old-growth forest. Both species groups were significantly depauperate in oil palm. Large mammal communities in old-growth forest became more heterogeneous at coarser spatial grains and small mammal communities became more homogeneous, while this pattern was reversed in logged forest. Both groups, however, showed a significant β-diversity signal at the finest grain in logged forest, likely due to logging-induced environmental heterogeneity. The β-diversity signal in oil palm was weak, but heterogeneity at the coarsest spatial grain was still evident, likely due to variation in landscape forest cover. Our findings suggest that the most effective spatial arrangement of set-aside will involve trade-offs between conserving large and small mammals. Greater consideration in the conservation and management of tropical landscapes needs to be given to β-diversity at a range of spatial grains.
Understanding how anthropogenic disturbance influences patterns of community composition and the reinforcing interactive processes that structure communities is important to mitigate threats to biodiversity. Competition is considered a primary reinforcing process, yet little is known concerning disturbance effects on competitive interaction networks. We examined how differences in ant community composition between undisturbed and disturbed Bornean rainforest, is potentially reflected by changes in competitive interactions over a food resource. Comparing 10 primary forest sites to 10 in selectively-logged forest, we found higher genus richness and diversity in the primary forest, with 18.5% and 13.0% of genera endemic to primary and logged respectively. From 180 hours of filming bait cards, we assessed ant-ant interactions, finding that despite considered aggression over food sources, the majority of ant interactions were neutral. Proportion of competitive interactions at bait cards did not differ between forest type, however, the rate and per capita number of competitive interactions was significantly lower in logged forest. Furthermore, the majority of genera showed large changes in aggression-score with often inverse relationships to their occupancy rank. This provides evidence of a shuffled competitive network, and these unexpected changes in aggressive relationships could be considered a type of competitive network re-wiring after disturbance.
Corals and their associated fauna are extremely diverse in tropical waters and form major reefs. In the high-latitude temperate zone, corals living near their distribution limit are considered marginal communities because they are particularly extremely sensitive to environmental and climatic changes. In this study, we examined the diversity and host usage of coral-associated barnacles on Jeju Island, Korea, the northern coral distribution limit in the East China Sea. In this study, only three coral-associated barnacles-from two genera in two subfamilies-were collected. The Pyrgomatinid barnacles Cantellius arcuatus and Cantellius cf. euspinulosum were found only on the corals Montipora millepora and Alveopora japonica, respectively. The Megatrematinid barnacle Pyrgomina oulastreae, relatively a generalist, was found on Psammocora spp. (both profundacella and albopicta) and Oulastrea crispata corals. The host usage of these three barnacles does not overlap. DNA barcode sequences of the C. arcuatus specimens collected in the present study matched those collected in Kochi in Japan, Taiwan, Malaysia and Papua New Guinea, suggesting that this species has a wide geographical distribution. C. arcuatus covers a wider host range in Taiwan waters, inhabiting Montipora spp. and Porites spp., which suggests that the host specificity of coral-associated barnacles varies with host availability. C. cf. euspinulosum probably has a very narrow distribution and host usage. The sequences of C. cf. euspinulosum on Jeju Island do not match those of any known sequences of Cantellius barnacles in the Indo-Pacific region. P. oulastreae probably prefers cold water because it has been reported in temperate regions. Coral-associated barnacles in marginal communities have considerably lower diversity than their subtropical and tropical counterparts. When host availability is limited, marginal coral-associated barnacles exhibit higher host specificity than those in subtropical and tropical reef systems.
One of the most urgent contemporary tasks for taxonomists and evolutionary biologists is to estimate the number of species on earth. Recording alpha diversity is crucial for protecting biodiversity, especially in areas of elevated species richness, which coincide geographically with increased anthropogenic environmental pressures - the world's so-called biodiversity hotspots. Although the distribution of Puddle frogs of the genus Occidozyga in South and Southeast Asia includes five biodiversity hotspots, the available data on phylogeny, species diversity, and biogeography are surprisingly patchy. Samples analyzed in this study were collected throughout Southeast Asia, with a primary focus on Sundaland and the Philippines. A mitochondrial gene region comprising ~ 2000 bp of 12S and 16S rRNA with intervening tRNA Valine and three nuclear loci (BDNF, NTF3, POMC) were analyzed to obtain a robust, time-calibrated phylogenetic hypothesis. We found a surprisingly high level of genetic diversity within Occidozyga, based on uncorrected p-distance values corroborated by species delimitation analyses. This extensive genetic diversity revealed 29 evolutionary lineages, defined by the > 5% uncorrected p-distance criterion for the 16S rRNA gene, suggesting that species diversity in this clade of phenotypically homogeneous forms probably has been underestimated. The comparison with results of other anuran groups leads to the assumption that anuran species diversity could still be substantially underestimated in Southeast Asia in general. Many genetically divergent lineages of frogs are phenotypically similar, indicating a tendency towards extensive morphological conservatism. We present a biogeographic reconstruction of the colonization of Sundaland and nearby islands which, together with our temporal framework, suggests that lineage diversification centered on the landmasses of the northern Sunda Shelf. This remarkably genetically structured group of amphibians could represent an exceptional case for future studies of geographical structure and diversification in a widespread anuran clade spanning some of the most pronounced geographical barriers on the planet (e.g., Wallace's Line). Studies considering gene flow, morphology, ecological and bioacoustic data are needed to answer these questions and to test whether observed diversity of Puddle frog lineages warrants taxonomic recognition.
To propose proper conservation measures and to elucidate coexistence mechanisms of sympatric carnivore species, we assessed temporal activity patterns of the sympatric carnivore species using 37,379 photos collected for more than 3 years at three study sites in Borneo. We categorized activity patterns of nine carnivore species (one bear, three civets, two felids, one skunk, one mustelid, one linsang) by calculating the photo-capturing proportions at each time period (day, night, twilight). We then evaluated temporal activity overlaps by calculating the overlap coefficients. We identified six nocturnal (three civets, one felid, one skunk, one linsang), two diurnal (one felid, one mustelid), and one cathemeral (bear) species. Temporal activity overlaps were high among the nocturnal species. The two felid species possessing morphological and ecological similarities exhibited clear temporal niche segregation, but the three civet species with similar morphology and ecology did not. Broad dietary breadth may compensate for the high temporal niche overlaps among the nocturnal species. Despite the high species richness of Bornean carnivores, almost half are threatened with extinction. By comparing individual radio-tracking and our data, we propose that a long-term study of at least 2 or 3 years is necessary to understand animals' temporal activity patterns, especially for sun bears and civets, by camera-trapping and to establish effective protection measures.
The distributions of amphibians, birds and mammals have underpinned global and local conservation priorities, and have been fundamental to our understanding of the determinants of global biodiversity. In contrast, the global distributions of reptiles, representing a third of terrestrial vertebrate diversity, have been unavailable. This prevented the incorporation of reptiles into conservation planning and biased our understanding of the underlying processes governing global vertebrate biodiversity. Here, we present and analyse the global distribution of 10,064 reptile species (99% of extant terrestrial species). We show that richness patterns of the other three tetrapod classes are good spatial surrogates for species richness of all reptiles combined and of snakes, but characterize diversity patterns of lizards and turtles poorly. Hotspots of total and endemic lizard richness overlap very little with those of other taxa. Moreover, existing protected areas, sites of biodiversity significance and global conservation schemes represent birds and mammals better than reptiles. We show that additional conservation actions are needed to effectively protect reptiles, particularly lizards and turtles. Adding reptile knowledge to a global complementarity conservation priority scheme identifies many locations that consequently become important. Notably, investing resources in some of the world's arid, grassland and savannah habitats might be necessary to represent all terrestrial vertebrates efficiently.
Most palm oil currently available in global markets is sourced from certified large-scale plantations. Comparatively little is sourced from (typically uncertified) smallholders. We argue that sourcing sustainable palm oil should not be determined by commercial certification alone and that the certification process should be revisited. There are so-far unrecognized benefits of sourcing palm oil from smallholders that should be considered if genuine biodiversity conservation is to be a foundation of 'environmentally sustainable' palm oil production. Despite a lack of certification, smallholder production is often more biodiversity-friendly than certified production from large-scale plantations. Sourcing palm oil from smallholders also alleviates poverty among rural farmers, promoting better conservation outcomes. Yet, certification schemes - the current measure of 'sustainability' - are financially accessible only for large-scale plantations that operate as profit-driven monocultures. Industrial palm oil is expanding rapidly in regions with weak environmental laws and enforcement. This warrants the development of an alternative certification scheme for smallholders. Greater attention should be directed to deforestation-free palm oil production in smallholdings, where production is less likely to cause large scale biodiversity loss. These small-scale farmlands in which palm oil is mixed with other crops should be considered by retailers and consumers who are interested in promoting sustainable palm oil production. Simultaneously, plantation companies should be required to make their existing production landscapes more compatible with enhanced biodiversity conservation.
This study explored the conservation values of communally reserved forests (CRFs), which local indigenous communities deliberately preserve within their area of shifting cultivation. In the current landscape of rural Borneo, CRFs are the only option for conservation because other forested areas have already been logged or transformed into plantations. By analyzing their alpha and beta diversity, we investigated how these forests can contribute to restore regional biodiversity. Although CRFs were fragmented and some had been disturbed in the past, their tree species diversity was high and equivalent to that of primary forests. The species composition of intact forests and forests disturbed in the past did not differ clearly, which indicates that past logging was not intensive. All CRFs contained unique and endangered species, which are on the IUCN Red List, Sarawak protected plants, or both. On the other hand, the forest size structure differed between disturbed and intact CRFs, with the disturbed CRFs consisting of relatively smaller trees. Although the beta diversity among CRFs was also high, we found a high contribution of species replacement (turnover), but not of richness difference, in the total beta diversity. This suggests that all CRFs have a conservation value for restoring the overall regional biodiversity. Therefore, for maintaining the regional species diversity and endangered species, it would be suitable to design a conservation target into all CRFs.