Nesting strategies affect altitudinal distribution and habitat use in Alpine dung beetle communities

1. Dung beetles are key contributors to a suite of ecosystem services. Understanding the factors that dictate their distributions is a necessary step towards preventing negative impacts of biodiversity loss. 2. Alpine dung beetle communities were analysed along altitudinal gradients to assess how different components of the community, defined in terms of nesting strategy [dung‐ovipositing Aphodiidae (DOAs), soil‐ovipositing Aphodiidae (SOAs) and two paracoprid (PAR) groups, Geotrupidae and Scarabaeidae] and parameters relevant to dung removal rates (species richness, total biomass and functional diversity), are distributed, and to identify to which environmental factors they respond. 3. Species richness declined with altitude. There was no significant variation in functional diversity or total biomass in relation to altitude. There were significant variations when considered by nesting group: DOA species richness and biomass decreased, SOA biomass increased, and Geotrupidae biomass showed a non‐linear trend, as altitude increased. 4. Functional diversity and total species richness were positively related to vegetation cover. DOA species richness was highest in forest and scrub; SOA species richness was highest in grassland and PAR species richness was lowest in rocky areas. 5. Dung beetle species show different trends in species richness and biomass depending on nesting strategy. Management to promote the dung beetle community should include maintenance of a mosaic of habitat types. Given the likely importance of species richness and biomass to ecosystem functioning, and the complimentary effect of different dung beetle groups, such a strategy may protect and enhance the ecosystem services that Alpine dung beetles provide.     

Monitoring the ecosystem service provided by dung beetles offers benefits over commonly used biodiversity metrics and a traditional trapping method

Surveying terrestrial invertebrates often requires lethal techniques that can also kill non-target vertebrates. Removing the desirable components of biodiversity is at odds with the philosophy of ecological restoration and biodiversity conservation more generally. Moreover, commonly used metrics generated by such survey approaches (e.g. abundance and species richness) are only indirectly related to the ecosystem services (e.g. pollination) that are often of primary interest. We examined the relationship between rates of dung removal (a direct measure of an ecosystem service) and dung beetle abundance and species richness in a temperate region of New South Wales, Australia, and examined if dung removal in revegetated riparian areas of different ages were trending monotonically toward rates in areas with mature native vegetation. Pellets of pig manure and conventional traps were left at study sites for 48 h to examine the relationships between rates of dung removal and dung beetle abundance and richness. Regressions of abundance and richness with average percent dung removal were positive and significant, demonstrating the potential of the method as a non-lethal proxy. While the dung removal method cannot determine the species responsible, percentage dung removal was more time-efficient, costing 4 min per sample, while abundance and richness cost 13 and 17 min, respectively. Despite variability among replicates of the same habitat type, the trajectory across the restoration gradient showed an increase from sites recently revegetated toward those with mature woody vegetation. We interpreted these results as a positive response of dung beetle activity and an indication of recovery of this ecosystem service. We argue that responses that can be collected efficiently such as dung removal should be used if restorationists have limited resources for data collection and analysis; non-specialists are involved; knowledge of ecosystem function is required, and animal ethics constrain options.     

The effects of land use change on native dung beetle diversity and function in Australia's Wet Tropics

The impacts of land use change on biodiversity and ecosystem functions are variable, particularly in fragmented tropical rainforest systems with high diversity. Dung beetles (Scarabaeinae) are an ideal group to investigate the relationship between land use change, diversity and ecosystem function as they are easily surveyed, sensitive to habitat modification and perform many ecosystem functions. Although this relationship has been investigated for dung beetles in some tropical regions, there has been no study assessing how native dung beetles in Australia's tropical rainforests respond to deforestation, and what the corresponding consequences are for dung removal (a key ecosystem function fulfilled by dung beetles). In this study we investigated the relationship between dung beetle community attributes (determined through trapping) and function (using dung removal experiments that allowed different dung beetle functional groups to access the dung) in rainforest and cleared pasture in a tropical landscape in Australia's Wet Tropics. Species richness, abundance and biomass were higher in rainforest compared to adjacent pasture, and species composition between these land use types differed significantly. However, average body size and evenness in body size were higher in pasture than in rainforest. Dung removal was higher in rainforest than in pasture when both functional groups or tunnelers only could access the dung. Increased dung removal in the rainforest was explained by higher biodiversity and dominance of a small number of species with distinct body sizes, as dung removal was best predicted by the evenness in body size of the community. Our findings suggest that functional traits (including body size and dung relocation behaviour) present in a dung beetle community are key drivers of dung removal. Overall, our results show that deforestation has reduced native dung beetle diversity in Australian tropical landscapes, which negatively impacts on the capacity for dung removal by dung beetles in this region.     

Stronger dung removal in forests compared with grassland is driven by trait composition and biomass of dung beetles

1. Ecosystem processes depend on the biomass of the involved organisms, but their functional diversity may play an additional role. In particular, the exclusion of key functional groups through habitat disturbance may lead to the breakdown of ecosystem functions. Dung removal is an important process contributing to nutrient cycling and thus productivity in grazed ecosystems. 2. This study investigated the role of different functional groups of dung beetles in dung removal in different habitats within a wood-pasture in two different seasons. An experimental setting with 12 blocks and 108 dung pads was used to investigate short-term dung removal over 1 week of exposure. 3. Dung removal was most strongly affected by habitat type, with almost 40% lower levels in grassland than in adjacent forest and forest gaps. Of all assemblage characteristics, total biomass of tunneller species was the strongest predictor of dung removal, whereas functional diversity showed no significant effect. In accordance with the dung removal pattern at habitat type level, densities of large tunnellers were suppressed in grassland compared with forest. 4. It is concluded that dung removal is habitat-specific and large tunnellers play a disproportionate role in this important ecosystem function in temperate forests.     

Fire? They don't give a dung! The resilience of dung beetles to fire in a tropical savanna

1. Disturbance is a strong driver of community assembly and fire has long been recognised as one of the main disturbances of terrestrial ecosystems. This study tested the resilience of dung beetles to fire events in campos rupestres, which is a tropical savanna ecosystem that evolved under a frequent fire regime, by assessing the resistance and recovery of their communities. 2. Dung beetles were sampled before and after a fire event and the effect of fire on dung beetle richness, abundance, mean community biomass and composition was tested. The effects of time since last fire and fire frequency on the community were also tested. 3. No effect of fire occurrence, time since last fire and fire frequency on any community variable was found. 4. Some non‐mutually exclusive mechanisms promoting the resistance and recovery of dung beetles in campos rupestres could be acting in synergy. One potential mechanism is the mismatched seasonality between fire events and dung beetle occurrence, as fires occur during the dry season and dung beetles are present above ground during the rainy season. Furthermore, dung beetles are insects that remain buried during most of their lifetime, which could protect individuals from being burned. Another potential mechanism is the replacement of species in burned areas by the movement of individuals from unburned areas, attracted by resources and/or by metacommunity dynamics. 5. It is concluded that in this ‘fire‐dependent’ ecosystem, dung beetle communities are resilient to fire and seem not to be structured by this disturbance.     

Linking functional group richness and ecosystem functions of dung beetles: an experimental quantification

Dung beetles form an insect group that fulfils important functions in terrestrial ecosystems throughout the world. These include nutrient cycling through dung removal, soil bioturbation, plant growth, secondary seed dispersal and parasite control. We conducted field experiments at two sites in the northern hemisphere temperate region in which dung removal and secondary seed dispersal were assessed. Dung beetles were classified in three functional groups, depending on their size and dung manipulation method: dwellers, large and small tunnelers. Other soil inhabiting fauna were included as a fourth functional group. Dung removal and seed dispersal by each individual functional group and combinations thereof were estimated in exclusion experiments using different dung types. Dwellers were the most diverse and abundant group, but tunnelers were dominant in terms of biomass. All dung beetle functional groups had a clear preference for fresh dung. The ecosystem services in dung removal and secondary seed dispersal provided by dung beetles were significant and differed between functional groups. Although in absolute numbers more dwellers were found, large tunnelers were disproportionally important for dung burial and seed removal. In the absence of dung beetles, other soil inhabiting fauna, such as earthworms, partly took over the dung decomposing role of dung beetles while most dung was processed when all native functional groups were present. Our results, therefore, emphasize the need to conserve functionally complete dung ecosystems to maintain full ecosystem functioning.     

Forest edges have high conservation value for bird communities in mosaic landscapes

A major conservation challenge in mosaic landscapes is to understand how trait‐specific responses to habitat edges affect bird communities, including potential cascading effects on bird functions providing ecosystem services to forests, such as pest control. Here, we examined how bird species richness, abundance and community composition varied from interior forest habitats and their edges into adjacent open habitats, within a multi‐regional sampling scheme. We further analyzed variations in Conservation Value Index (CVI), Community Specialization Index (CSI) and functional traits across the forest‐edge‐open habitat gradient. Bird species richness, total abundance and CVI were significantly higher at forest edges while CSI peaked at interior open habitats, i.e., furthest from forest edge. In addition, there were important variations in trait‐ and species‐specific responses to forest edges among bird communities. Positive responses to forest edges were found for several forest bird species with unfavorable conservation status. These species were in general insectivores, understorey gleaners, cavity nesters and long‐distance migrants, all traits that displayed higher abundance at forest edges than in forest interiors or adjacent open habitats. Furthermore, consistently with predictions, negative edge effects were recorded in some forest specialist birds and in most open‐habitat birds, showing increasing densities from edges to interior habitats. We thus suggest that increasing landscape‐scale habitat complexity would be beneficial to declining species living in mosaic landscapes combining small woodlands and open habitats. Edge effects between forests and adjacent open habitats may also favor bird functional guilds providing valuable ecosystem services to forests in longstanding fragmented landscapes.     

Environmental consequences of deltamethrin residues in cattle feces in an African agricultural landscape

Pyrethroid insecticides are widely used to control ectoparasites of livestock, particularly ticks and biting flies. Their use in African livestock systems is increasing, driven by the need to increase productivity and local food security. However, insecticide residues present in the dung after treatment are toxic to dung‐inhabiting insects. In a semiarid agricultural habitat in Botswana, dung beetle adult mortality, brood ball production, and larval survival were compared between untreated cattle dung and cattle dung spiked with deltamethrin, to give concentrations of 0.01, 0.1, 0.5, or 1 ppm. Cattle dung‐baited pitfall traps were used to measure repellent effects of deltamethrin in dung on Scarabaeidae. Dung decomposition rate was also examined. There was significantly increased mortality of adult dung beetles colonizing pats that contained deltamethrin compared to insecticide‐free pats. Brood ball production was significantly reduced at concentrations of 1 ppm; larval survival was significantly reduced in dung containing 0.1 ppm deltamethrin and above. There was no difference in the number of Scarabaeidae attracted to dung containing any of the deltamethrin concentrations. Dung decomposition was significantly reduced even at the lowest concentration (0.01 ppm) compared to insecticide‐free dung. The widespread use of deltamethrin in African agricultural ecosystems is a significant cause for concern; sustained use is likely to damage dung beetle populations and their provision of environmentally and economically important ecosystem services. Contaminated dung buried by paracoprid (tunneling) beetles may retain insecticidal effects, with impacts on developing larvae below ground. Lethal and sublethal effects on entire dung beetle (Scarabaeidae) communities could impair ecosystem function in agricultural landscapes.     

Do riparian reserves support dung beetle biodiversity and ecosystem services in oil palm-dominated tropical landscapes?

Agricultural expansion and intensification are major threats to global biodiversity, ecological functions, and ecosystem services. The rapid expansion of oil palm in forested tropical landscapes is of particular concern given their high biodiversity. Identifying management approaches that maintain native species and associated ecological processes within oil palm plantations is therefore a priority. Riparian reserves are strips of forest retained alongside rivers in cultivated areas, primarily for their positive hydrological impact. However, they can also support a range of forest‐dependent species or ecosystem services. We surveyed communities of dung beetles and measured dung removal activity in an oil palm‐dominated landscape in Sabah, Malaysian Borneo. The species richness, diversity, and functional group richness of dung beetles in riparian reserves were significantly higher than in oil palm, but lower than in adjacent logged forests. The community composition of the riparian reserves was more similar to logged forest than oil palm. Despite the pronounced differences in biodiversity, we did not find significant differences in dung removal rates among land uses. We also found no evidence that riparian reserves enhance dung removal rates within surrounding oil palm. These results contrast previous studies showing positive relationships between dung beetle species richness and dung removal in tropical forests. We found weak but significant positive relationships between riparian reserve width and dung beetle diversity, and between reserve vegetation complexity and dung beetle abundance, suggesting that these features may increase the conservation value of riparian reserves. Synthesis and applications: The similarity between riparian reserves and logged forest demonstrates that retaining riparian reserves increases biodiversity within oil palm landscapes. However, the lack of correlation between dung beetle community characteristics and dung removal highlights the need for further research into spatial variation in biodiversity–ecosystem function relationships and how the results of such studies are affected by methodological choices.     

Experimental evidence for the effects of dung beetle functional group richness and composition on ecosystem function in a tropical forest

Much of the literature on the relationship between species richness or functional group richness and measures of ecosystem function focuses on a restricted set of ecosystem function measures and taxonomic groups. Few such studies have been carried out under realistic levels of diversity in the field, particularly in high diversity ecosystems such as tropical forests. We used exclusion experiments to study the effects of dung beetle functional group richness and composition on two interlinked and functionally important ecological processes, dung removal and secondary seed dispersal, in evergreen tropical forest in Sabah, Malaysian Borneo. Overall, both dung and seed removal increased with dung beetle functional group richness. However, levels of ecosystem functioning were idiosyncratic depending on the identity of the functional groups present, indicating an important role for functional group composition. There was no evidence for interference or competition among functional groups. We found strong evidence for overyielding and transgressive overyielding, suggesting complementarity or facilitation among functional groups. Not all mixtures showed transgressive overyielding, so that complementarity was restricted to particular functional group combinations. Beetles in a single functional group (large nocturnal tunnellers) had a disproportionate influence on measures of ecosystem function: in their absence dung removal is reduced by approximately 75%. However, a full complement of functional groups is required to maximize ecosystem functioning. This study highlights the importance of both functional group identity and species composition in determining the ecosystem consequences of extinctions or altered patterns in the relative abundance of species.     

Early-stage ecological influences of population recovery of large mammals on dung beetle assemblages in heavy snow areas

Past conservation initiatives and rapidly decreasing human populations in modern Japan have contributed to population recoveries of Sika deer (Cervus nippon), wild boar (Sus scrofa), and Japanese macaque (Macaca fuscata) throughout the country. Ironically, however, these recoveries have not always received a favorable reception, because these mammals can also be agricultural pests. To open public debate on the recoveries, based on a thorough understanding of their multifaceted roles in sustaining the local ecosystem, we evaluated the initial stage ripple effects caused by the mammalian population recovery on the community assembly of dung beetles, which are keystone decomposer organisms in terrestrial ecosystems. For the evaluation, we conducted manipulative snapshot experiments, using camera and pitfall traps, for mammal and dung beetle assemblages, respectively, in four different mountain ranges within the heavy snow areas of northern Japan, where the recovery of three mammal populations was at an early stage. The current findings implied that, although the feces of every recovering mammal species could provide valuable resources for most beetles, the ripple effects from the mammal population recoveries were subject to hysteresis of the local ecosystem, i.e., catastrophic shifts in ecosystems originating from the historical background of regional mammal defaunation. In particular, the abundance of tunnelers that could benefit from positive ripple effects decreased with an increase in past disturbances, which resulted in emptier forests, i.e., an ecosystem with fewer large mammals. The findings suggested that recovering populations of large mammals do not always contribute to the restoration of the original dung beetle communities, at least initially.     

Edge effects influence the composition and density of reef residents on subtidal restored oyster reefs

Within estuarine and coastal ecosystems globally, extensive habitat degradation and loss threaten critical ecosystem functions and necessitate widescale restoration efforts. There is abundant evidence that ecological processes and species interactions can vary with habitat characteristics, which has important implications for the design and implementation of restoration efforts aimed at enhancing specific ecosystem functions and services. We conducted an experiment examining how habitat characteristics (presence; edge vs. interior) influence the communities of resident fish and mobile invertebrates on restored oyster (Crassostrea virginica) reefs. Similar to previous studies, we found that restored reefs altered community composition and augmented total abundance and biomass relative to unstructured sand habitat. Community composition and biomass also differed between the edge and interior of individual reefs as a result of species-specific patterns over small spatial scales. These patterns were only weakly linked to oyster density, suggesting that other factors that vary between edge and interior (e.g. predator access or species interactions) are likely more important for community structure on oyster reefs. Fine-scale information on resident species' use of oyster reefs will help facilitate restoration by allowing decision makers to optimize the amount of edge versus interior habitat. To improve the prediction of faunal use and benefits from habitat restoration, we recommend investigations into the mechanisms shaping edge and interior preferences on oyster reefs.     

Down‐sizing of dung beetle assemblages over the last 53 000 years is consistent with a dominant effect of megafauna losses

The ongoing down‐sizing of the global mammal communities is assumed to have subsequent effects on mutualistic species communities. Dung beetles co‐evolved with large‐sized animals since millennia and depend on the megafauna feces of an appropriate size. Mammal community down‐sizing as a result of past and ongoing megafauna losses is therefore likely to result in a down‐sizing of dung beetle communities. However, empirical evidence for this co‐down‐sizing is lacking especially on larger spatial scales and over extended periods of time. Here, we show a significant down‐sizing of European dung beetle assemblages over the last ~53 000 years by relating Quaternary fossil records with trait information on body size of beetles. This significant down‐sizing of dung beetle communities was thereby not linear, but characterized by a weak decrease until the early Holocene but a strong acceleration in the recent pre‐history, from 6–7000 years BP onwards. This acceleration of down‐sizing coincides with the completion of the Quaternary megafauna extinction and the start of major shifts in human agricultural land‐use. In contrast, assemblage mean body size of non‐coprophagous scarabids as well as ground beetles – two groups of beetles with no or weak relations to megafauna – was observed to increase towards the present with an acceleration of body size increase coinciding with the onset of late‐glacial warming (14 200 years BP). In summary, the observed late‐Quaternary down‐sizing of European dung beetle communities is consistent with an effect of pre‐historic megafauna losses, and not with the coincident general warming. Ongoing down‐sizing of mammal communities is therefore likely to result in further down‐sizing of dung beetle assemblages, with potential effects on their important role for nutrient cycling and secondary seed dispersal in natural and extensive agro‐ecosystems. Future nature management initiatives could halt or even reverse this functional diversity loss via effective protection or restoration of megafauna communities.     

Dung Beetle Community and Functions along a Habitat-Disturbance Gradient in the Amazon: A Rapid Assessment of Ecological Functions Associated to Biodiversity

Although there is increasing interest in the effects of habitat disturbance on community attributes and the potential consequences for ecosystem functioning, objective approaches linking biodiversity loss to functional loss are uncommon. The objectives of this study were to implement simultaneous assessment of community attributes (richness, abundance and biomass, each calculated for total-beetle assemblages as well as small- and large-beetle assemblages) and three ecological functions of dung beetles (dung removal, soil perturbation and secondary seed dispersal), to compare the effects of habitat disturbance on both sets of response variables, and their relations. We studied dung beetle community attributes and functions in five land-use systems representing a disturbance gradient in the Brazilian Amazon: primary forest, secondary forest, agroforestry, agriculture and pasture. All response variables were affected negatively by the intensification of habitat disturbance regimes, but community attributes and ecological functions did not follow the same pattern of decline. A hierarchical partitioning analysis showed that, although all community attributes had a significant effect on the three ecological functions (except the abundance of small beetles on all three ecological functions and the biomass of small beetles on secondary dispersal of large seed mimics), species richness and abundance of large beetles were the community attributes with the highest explanatory value. Our results show the importance of measuring ecological function empirically instead of deducing it from community metrics.     

Pastoral Practices to Reverse Shrub Encroachment of Sub-Alpine Grasslands: Dung Beetles (Coleoptera,Scarabaeoidea) Respond More Quickly Than Vegetation

In recent decades, pastoral abandonment has produced profound ecological changes in the Alps. In particular, the reduction in grazing has led to extensive shrub encroachment of semi-natural grasslands, which may represent a threat to open habitat biodiversity. To reverse shrub encroachment, we assessed short-term effects of two different pastoral practices on vegetation and dung beetles (Coleoptera, Scarabaeoidea). Strategic placement of mineral mix supplements (MMS) and arrangement of temporary night camp areas (TNCA) for cattle were carried out during summer 2011 in the Val Troncea Natural Park, north-western Italian Alps. In 2012, one year after treatment, a reduction in shrub cover and an increase in bare ground cover around MMS sites was detected. A more intense effect was detected within TNCA through increases in forage pastoral value, and in the cover and height of the herbaceous layer. Immediately after treatment, changes in dung beetle diversity (total abundance, species richness, Shannon diversity, taxonomic and functional diversity) showed a limited disturbance effect caused by high cattle density. In contrast, dung beetle diversity significantly increased one year later both at MMS and TNCA sites, with a stronger effect within TNCA. Multivariate Regression Trees and associated Indicator Value analyses showed that some ecologically relevant dung beetle species preferred areas deprived of shrub vegetation. Our main conclusions are: i) TNCA are more effective than MMS in terms of changes to vegetation and dung beetles, ii) dung beetles respond more quickly than vegetation to pastoral practices, and iii) the main driver of the rapid response by dung beetles is the removal of shrubs. The resulting increase in dung beetle abundance and diversity, which are largely responsible for grassland ecosystem functioning, may have a positive effect on meso-eutrophic grassland restoration. Shrub encroachment in the Alps may therefore be reversed, and restoration of grassland enhanced, by using appropriate pastoral practices.     

Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration

I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.     

Use of habitat resources by scarab dung beetles in an savanna

In the Queen Elizabeth National Park, Uganda, we compared the scarab beetle assemblages in the dung of three wild ungulates (African buffalo, a ruminant foregut fermenter; hippopotamus, nonruminant foregut fermenter; and warthog, nonruminant hindgut fermenter). Dung was collected from two sandy-clay soils with different percentage of coarse sand. We aimed at investigating habitat resource selection by dung beetle species within a savanna natural contest with abundant and diverse food availability. Analyses were performed to detect differences for dung beetle assemblages in abundance, diversity, functional groups. Species richness in the three dung types and in the two soil types was similar. However, warthog dung and sandy-rich soil appeared the preferred habitat resources, in terms of abundance and biomass, while hippopotamus dung hosted the lowest values for these parameters. The analysis of functional groups revealed that slow-burying tunnellers held the major role, both in terms of abundance and biomass, and were mainly found in warthog dung.     

优势种去除对崇明东滩盐沼湿地生态系统的影响

盐沼生态系统环境梯度明显,物种组成较简单,是研究生物多样性与生态系统功能关系的理想对象。本研究以崇明东滩盐沼湿地为研究区域,研究优势种去除对植物群落结构以及底栖动物群落的影响。结果表明:(1)去除处理仅对植物群落分株密度有极显著效应(P0.01)。去除组和对照组物种组成差异随时间增加而减小,处理效应逐渐减弱。(2)去除组底栖动物密度均低于对照组,但差异不显著。(3)盐沼植物群落特征与底栖动物群落有密切关系,植物密度、冠层高度与底栖动物密度相关性极显著。去除优势种后,植物群落分株密度升高,群落内剩余物种占比有所上升,次优势种对群落的补偿效应具有较大贡献;而底栖动物群落密度下降,其生物量和多样性指数的变化趋势与密度并不一致。上述结果表明生物多样性变化影响了盐沼湿地生态系统植物群落和底栖动物群落结构,进而可能影响物质循环和能量流动过程。     

Dung beetle assemblages on tropical land‐bridge islands: small island effect and vulnerable species

Aim Using dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae) in a tropical land‐bridge island system, we test for the small island effect (SIE) in the species–area relationship and evaluate its effects on species richness and community composition. We also examine the determinants of species richness across island size and investigate the traits of dung beetle species in relation to their local extinction vulnerability following forest fragmentation. Location Lake Kenyir, a hydroelectric reservoir in north‐eastern Peninsular Malaysia. Methods We sampled dung beetles using human dung baited pitfall traps on 24 land‐bridge islands and three mainland sites. We used regression tree analyses to test for the SIE, as well as species traits related to local rarity, as an indication of extinction vulnerability. We employed generalized linear models (GLMs) to examine determinants for species richness at different scales and compared the results with those from conventional linear and breakpoint regressions. Community analyses included non‐metric multidimensional scaling, partial Mantel tests, nestedness analysis and abundance spectra. Results Regression tree analysis revealed an area threshold at 35.8 ha indicating an SIE. Tree basal area was the most important predictor of species richness on small islands (<35.8 ha). Results from GLMs supported these findings, with isolation and edge index also being important for small islands. The SIE also manifested in patterns of dung beetle community composition where communities on small islands (<35.8 ha) departed from those on the mainland and larger islands, and were highly variable with no significant nestedness, probably as a result of unexpected species occurrences on several small islands. The communities exhibited a low degree of spatial autocorrelation, suggesting that dispersal limitation plays a part in structuring dung beetle assemblages. Species with lower baseline density and an inability to forage on the forest edge were found to be rarer among sites and hence more prone to local extinction. Main conclusions We highlight the stochastic nature of dung beetle community composition on small islands and argue that this results in reduced ecosystem functionality. A better understanding of the minimum fragment size required for retaining functional ecological communities will be important for effective conservation management and the maintenance of tropical forest ecosystem stability.