Deliberately introduced dung beetles in Australia: 12 years of occurrence and abundance records from 2001 to 2022.

Since 1968, the Australian Dung Beetle Project has carried out field releases of 43 deliberately introduced dung beetle species for the biological control of livestock dung and dung-breeding pests. Of these, 23 species are known to have become established. For most of these species, sufficient time has elapsed for population expansion to fill the extent of their potential geographic range through both natural and human-assisted dispersal. Consequently, over the last 20 years, extensive efforts have been made to quantify the current distribution of these introduced dung beetles, as well as the seasonal and spatial variation in their activity levels. Much of these data and their associated metadata have remained unpublished, and they have not previously been synthesized into a cohesive dataset. Here, we collate and report data from the three largest dung beetle monitoring projects from 2001 to 2022. Together, these projects encompass data collected from across Australia, and include records for all 23 species of established dung beetles introduced for biocontrol purposes. In total, these data include 22,718 presence records and 213,538 absence records collected during 10,272 sampling events at 546 locations. Most presence records (97%) include abundance data. In total, 1,752,807 dung beetles were identified as part of these data. The distributional occurrence and abundance data can be used to explore questions such as factors influencing dung beetle species distributions, dung beetle biocontrol, and insect-mediated ecosystem services. These data are provided under a CC-BY-NC 4.0 license and users are encouraged to cite this data paper when using the data.

Global meta-analysis reveals overall higher nocturnal than diurnal activity in insect communities.

Insects sustain key ecosystem functions, but how their activity varies across the day-night cycle and the underlying drivers are poorly understood. Although entomologists generally expect that more insects are active at night, this notion has not been tested with empirical data at the global scale. Here, we assemble 331 quantitative comparisons of the abundances of insects between day and night periods from 78 studies worldwide and use multi-level meta-analytical models to show that insect activity is on average 31.4% (CI: -6.3%-84.3%) higher at night than in the day. We reveal diel preferences of major insect taxa, and observe higher nocturnal activity in aquatic taxa than in terrestrial ones, as well as in warmer environments. In a separate analysis of the small subset of studies quantifying diel patterns in taxonomic richness (31 comparisons from 13 studies), we detect preliminary evidence of higher nocturnal richness in tropical than temperate communities. The higher overall (but variable) nocturnal activity in insect communities underscores the need to address threats such as light pollution and climate warming that may disproportionately impact nocturnal insects.

Dung beetles increase plant growth: a meta-analysis.

The ecosystem services provided by dung beetles are well known and valued. Dung beetles bury dung for feeding and breeding, and it is generally thought that the process of burying dung increases nutrient uptake by plant roots, which promotes plant growth. Many studies have tested the effects of dung beetles on plant growth, but there has been no quantitative synthesis of these studies. Here we use a multi-level meta-analysis to estimate the average effect of dung beetles on plant growth and investigate factors that moderate this effect. We identified 28 publications that investigated dung beetle effects on plant growth. Of these, 24 contained the minimum quantitative data necessary to include in a meta-analysis. Overall, we found that dung beetles increased plant growth by 17%; the 95% CI for possible values for the true increase in plant growth that were most compatible with our data, given our statistical model, ranged from 1% to 35%. We found evidence that the dung beetle-plant growth relationship is influenced by the plant measurement type and the number of beetles accessing the dung. However, beetles did not increase plant growth in all quantitative trials, as individual effect sizes ranged from -72% to 806%, suggesting important context-dependence in the provision of ecosystem services.

Intraspecific leaf trait variation mediates edge effects on litter decomposition rate in fragmented forests.

There is strong trait dependence in species-level responses to environmental change and their cascading effects on ecosystem functioning. However, there is little understanding of whether intraspecific trait variation (ITV) can also be an important mechanism mediating environmental effects on ecosystem functioning. This is surprising, given that global change processes such as habitat fragmentation and the creation of forest edges drive strong trait shifts within species. On 20 islands in the Thousand Island Lake, China, we quantified intraspecific leaf trait shifts of a widely distributed shrub species, Vaccinium carlesii, in response to habitat fragmentation. Using a reciprocal transplant decomposition experiment between forest edge and interior on 11 islands with varying areas, we disentangled the relative effects of intraspecific leaf trait variation versus altered environmental conditions on leaf decomposition rates in forest fragments. We found strong intraspecific variation in leaf traits in response to edge effects, with a shift toward recalcitrant leaves with low specific leaf area and high leaf dry matter content from forest interior to the edge. Using structural equation modeling, we showed that such intraspecific leaf trait response to habitat fragmentation had translated into significant plant afterlife effects on leaf decomposition, leading to decreased leaf decomposition rates from the forest interior to the edge. Importantly, the effects of intraspecific leaf trait variation were additive to and stronger than the effects from local environmental changes due to edge effects and habitat loss. Our experiment provides the first quantitative study showing that intraspecific leaf trait response to edge effects is an important driver of the decrease in leaf decomposition rate in fragmented forests. By extending the trait-based response-effect framework toward the individual level, intraspecific variation in leaf economics traits can provide the missing functional link between environmental change and ecological processes. These findings suggest an important area for future research on incorporating ITV to understand and predict changes in ecosystem functioning in the context of global change.

Complex battlefields favor strong soldiers over large armies in social animal warfare.

In social animals, success can depend on the outcome of group battles. Theoretical models of warfare predict that group fighting ability is proportional to two key factors: the strength of each soldier in the group and group size. The relative importance of these factors is predicted to vary across environments [F. W. Lanchester, Aircraft in Warfare, the Dawn of the Fourth Arm (1916)]. Here, we provide an empirical validation of the theoretical prediction that open environments should favor superior numbers, whereas complex environments should favor stronger soldiers [R. N. Franks, L. W. Partridge, Anim. Behav. 45, 197-199 (1993)]. We first demonstrate this pattern using simulated battles between relatively strong and weak soldiers in a computer-driven algorithm. We then validate this result in real animals using an ant model system: In battles in which the number of strong native meat ant Iridomyrmex purpureus workers is constant while the number of weak non-native invasive Argentine ant Linepithema humile workers increases across treatments, fatalities of I. purpureus are lower in complex than in simple arenas. Our results provide controlled experimental evidence that investing in stronger soldiers is more effective in complex environments. This is a significant advance in the empirical study of nonhuman warfare and is important for understanding the competitive balance among native and non-native invasive ant species.

Forest edges increase pollinator network robustness to extinction with declining area.

Edge effects often exacerbate the negative effects of habitat loss on biodiversity. In forested ecosystems, however, many pollinators actually prefer open sunny conditions created by edge disturbances. We tested the hypothesis that forest edges have a positive buffering effect on plant-pollinator interaction networks in the face of declining forest area. In a fragmented land-bridge island system, we recorded ~20,000 plant-pollinator interactions on 41 islands over 3 yr. We show that plant richness and floral resources decline with decreasing forest area at both interior and edge sites, but edges maintain 10-fold higher pollinator abundance and richness regardless of area loss. Edge networks contain highly specialized species, with higher nestedness and lower modularity than interior networks, maintaining high robustness to extinction following area loss while forest interior networks collapse. Anthropogenic forest edges benefit community diversity and network robustness to extinction in the absence of natural gap-phase dynamics in small degraded forest remnants.

Disentangling biotic and abiotic drivers of intraspecific trait variation in woody plant seedlings at forest edges.

In fragmented forests, edge effects can drive intraspecific variation in seedling performance that influences forest regeneration and plant composition. However, few studies have attempted to disentangle the relative biotic and abiotic drivers of intraspecific variation in seedling performance. In this study, we carried out a seedling transplant experiment with a factorial experimental design on three land-bridge islands in the Thousand Island Lake, China, using four common native woody plant species. At different distances from the forest edge (2, 8, 32, 128 m), we transplanted four seedlings of each species into each of three cages: full-cage, for herbivore exclusion; half-cage, that allowed herbivore access but controlled for caging artifacts; and no-cage control. In the 576 cages, we recorded branch architecture, leaf traits, and seedling survival for each seedling before and after the experimental treatment. Overall, after one full growing season, edge-induced abiotic drivers and varied herbivory pressure led to intraspecific variation in seedling performance, including trade-offs in seedling architecture and resource-use strategies. However, responses varied across species with different life-history strategies and depended on the driver in question, such that the abiotic and biotic effects were additive across species, rather than interactive. Edge-induced abiotic variation modified seedling architecture of a shade-tolerant species, leading to more vertical rather than lateral growth at edges. Meanwhile, increased herbivory pressure resulted in a shift toward lower dry matter investment in leaves of a light-demanding species. Our results suggest that edge effects can drive rapid directional shifts in the performance and intraspecific traits of some woody plants from early ontogenetic stages, but most species in this study showed negligible phenotypic responses to edge effects. Moreover, species-specific responses suggest the importance of interspecific differences modulating the degree of trait plasticity, implying the need to incorporate individual-level responses when understanding the impact of forest fragmentation on plant communities.

Re-establishment of cavity-nesting bee and wasp communities along a reforestation gradient in southern Amazonia.

Global initiatives to reforest degraded areas have intensified in recent years, in an attempt to reverse the environmental impacts of habitat loss on species and ecosystem provided by them. However, the effectiveness of such reforestation initiatives in re-establishing biodiversity is still poorly understood. Here, we test how reforestation type and intensity applied to deforested areas affect the reestablishment of communities of cavity-nesting bees and wasps. We deployed experimental trap-nests along a reforestation gradient of increasing structural similarity to primary forest, after 18 years of reforestation. We found that reestablishment, in terms of abundance and richness of both bees and wasps, was greatest at an intermediate point along the reforestation gradient. However, these communities were highly dissimilar to primary forest, and recovery of intact insect community composition was only achieved when reforestation was more similar in structure to natural forests. This effect was more pronounced for bees than for wasps. Our findings suggest that along the reforestation gradient, services provided by wasps will be more easily recovered than those provided by bees. Our results have important implications for the challenges of restoring and maintaining species biodiversity as well as their associated ecosystem services.

A systematic revision of Draculoides (Schizomida: Hubbardiidae) of the Pilbara, Western Australia, Part I: the Western Pilbara.

The schizomid fauna of mainland Australia currently comprises 60 species within seven named genera, of which five are endemic to the continent: Attenuizomus Harvey, 2000, Brignolizomus Harvey, 2000, Draculoides Harvey, 1992, Julattenius Harvey, 1992, Notozomus Harvey, 2000. Most Australian schizomids have been described from eastern and northern Australia, but there is also a significant subterranean fauna that has been found in hypogean habitats in the semi-arid Pilbara region of Western Australia. The vast majority of these species can be assigned to the genus Draculoides and this study is the first in a proposed series to revise this highly diverse genus. We treat the species found in the western Pilbara region, which includes 13 new species and 13 previously named species, using morphological characters and multi-locus sequence data. We also incorporate a molecular "mini-barcode" approach for COI, 12S and ITS2 to diagnose the new species. The new species are named: Draculoides akashae Abrams and Harvey, n. sp., D. belalugosii Abrams and Harvey, n. sp., D. carmillae Abrams and Harvey, n. sp., D. christopherleei Abrams and Harvey, n. sp., D. claudiae Abrams and Harvey, n. sp., D. immortalis Abrams and Harvey, n. sp., D. karenbassettae Abrams and Harvey, n. sp., D. mckechnieorum Abrams and Harvey, n. sp., D. minae Abrams and Harvey, n. sp., D. noctigrassator Abrams and Harvey, n. sp., D. nosferatu Abrams and Harvey, n. sp., D. piscivultus Abrams and Harvey, n. sp. and D. warramboo Abrams and Harvey, n. sp. We also provide the first descriptions of males of D. anachoretus (Harvey, Berry, Edward and Humphreys, 2008) and D. gnophicola (Harvey, Berry, Edward and Humphreys, 2008). All of the new species are subterranean-dwelling, short-range endemic species that occur in regions subject to mining activities, rendering them of high conservation significance.

Multi-Species Phylogeography of Arid-Zone Sminthopsinae (Marsupialia: Dasyuridae) Reveals Evidence of Refugia and Population Expansion in Response to Quaternary Change.

Historical population contraction and expansion events associated with Pleistocene climate change are important drivers of intraspecific population structure in Australian arid-zone species. We compared phylogeographic patterns among arid-adapted Dasyuridae (Sminthopsis and Planigale) with close phylogenetic relationships and similar ecological roles to investigate the drivers of phylogeographic structuring and the importance of historical refugia. We generated haplotype networks for two mitochondrial (control region and cytochrome b) and one nuclear (omega-globin) gene from samples distributed across each species range. We used ΦST to test for a genetic population structure associated with the four Pilbara subregions, and we used expansion statistics and Bayesian coalescent skyline analysis to test for signals of historical population expansion and the timing of such events. Significant population structure associated with the Pilbara and subregions was detected in the mitochondrial data for most species, but not with the nuclear data. Evidence of population expansion was detected for all species, and it likely began during the mid-late Pleistocene. The timing of population expansion suggests that these species responded favorably to the increased availability of arid habitats during the mid-late Pleistocene, which is when previously patchy habitats became more widespread. We interpret our results to indicate that the Pilbara region could have acted as a refugium for small dasyurids.

FragSAD: A database of diversity and species abundance distributions from habitat fragments.

Habitat destruction is the single greatest anthropogenic threat to biodiversity. Decades of research on this issue have led to the accumulation of hundreds of data sets comparing species assemblages in larger, intact, habitats to smaller, more fragmented, habitats. Despite this, little synthesis or consensus has been achieved, primarily because of non-standardized sampling methodology and analyses of notoriously scale-dependent response variables (i.e., species richness). To be able to compare and contrast the results of habitat fragmentation on species' assemblages, it is necessary to have the underlying data on species abundances and sampling intensity, so that standardization can be achieved. To accomplish this, we systematically searched the literature for studies where abundances of species in assemblages (of any taxa) were sampled from many habitat patches that varied in size. From these, we extracted data from several studies, and contacted authors of studies where appropriate data were collected but not published, giving us 117 studies that compared species assemblages among habitat fragments that varied in area. Less than one-half (41) of studies came from tropical forests of Central and South America, but there were many studies from temperate forests and grasslands from all continents except Antarctica. Fifty-four of the studies were on invertebrates (mostly insects), but there were several studies on plants (15), birds (16), mammals (19), and reptiles and amphibians (13). We also collected qualitative information on the length of time since fragmentation. With data on total and relative abundances (and identities) of species, sampling effort, and affiliated meta-data about the study sites, these data can be used to more definitively test hypotheses about the role of habitat fragmentation in altering patterns of biodiversity. There are no copyright restrictions. Please cite this data paper and the associated Dryad data set if the data are used in publications.

Too hot to handle: Cenozoic aridification drives multiple independent incursions of Schizomida (Hubbardiidae) into hypogean environments.

The formation of the Australian arid zone, Australia's largest and youngest major biome, has been recognized as a major driver of rapid evolutionary radiations in terrestrial plants and animals. Here, we investigate the phylogenetic diversity and evolutionary history of subterranean short-tailed whip scorpions (Schizomida: Hubbardiidae), which are a significant faunal component of Western Australian hypogean ecosystems. We sequenced two mitochondrial (12S, COI) and three nuclear DNA markers (18S, 28S, ITS2) from ∼600 specimens, largely from the genera Draculoides and Paradraculoides, including 20 previously named species and an additional 56 newly identified operational taxonomic units (OTUs). Phylogenetic analyses revealed a large and rapid species radiation congruent with Cenozoic aridification of the continent, in addition to the identification of a new genus in Western Australia and the first epigean schizomid from the Pilbara. Here, we also synonymise Paradraculoides with Draculoides (new synonymy), due to paraphyly and a lack of reliable characters to define the two genera. Our results are consistent with multiple colonisations of the subterranean realm from epigean ancestors as their forest habitat fragmented and retracted, with ongoing fragmentation and diversification of lineages underground. These findings illustrate the remarkable diversity and high incidence of short-range endemism of Western Australia's subterranean fauna, which has important implications for identifying and managing short-range endemic subterranean fauna. They also highlight the advantages of including molecular data in subterranean fauna surveys as all specimens can be utilized, regardless of sex and life stage. Additionally, we have provided the first multi-gene phylogenetic framework for Australian schizomids, which will enable researchers and environmental consultants to identify new taxa or align them to existing lineages.

The scorpion-tailed orb-weaving spiders (Araneae, Araneidae, Arachnura) in Australia and New Zealand.

The scorpion-tailed orb-weaving spiders in the genus Arachnura Vinson, 1863 (Araneidae Clerck, 1757) are revised for Australia and New Zealand. Arachnura higginsii (L. Koch, 1872) only occurs in Australia and A. feredayi (L. Koch, 1872) only in New Zealand. A single female collected in south-eastern Queensland (Australia) is here tentatively identified as A. melanura Simon, 1867, but it is doubtful that this species has established in Australia. Two juveniles from northern Queensland do not conform to the diagnoses of any of the above species and are illustrated pending a more thorough revision of the genus in South-East Asia and the Pacific region. An unidentified female from Westport (New Zealand) does not conform to the diagnoses of A. feredayi and A. higginsii, but is not described due to its poor preservation status. Arachnura caudatella Roewer, 1942 (replacement name for Epeira caudata Bradley, 1876), originally described from Hall Sound (Papua New Guinea) and repeatedly catalogued for Australia, is considered a nomen dubium.

A global strategy to mitigate the environmental impact of China's ruminant consumption boom.

Rising demand for ruminant meat and dairy products in developing countries is expected to double anthropogenic greenhouse gas and ammonia emissions from livestock by 2050. Mitigation strategies are urgently needed to meet demand while minimizing environmental impacts. Here, we develop scenarios for mitigating emissions under local vs global supply policies using data from 308 livestock farms across mainland China, where emissions intensities are ~50% higher than those in developed nations. Intensification of domestic production and globalized expansion through increased trade result in reductions in global emissions by nearly 30% over a business-as-usual scenario, but at the expense of trading partners absorbing the associated negative externalities of environmental degradation. Only adoption of a mixed strategy combining global best-practice in sustainable intensification of domestic production, with increased green-source trading as a short-term coping strategy, can meet 2050 demand while minimizing the local and global environmental footprint of China's ruminant consumption boom.

Soil fertility shapes belowground food webs across a regional climate gradient.

Changes in soil fertility during pedogenesis affect the quantity and quality of resources entering the belowground subsystem. Climate governs pedogenesis, yet how climate modulates responses of soil food webs to soil ageing remains unexplored because of the paucity of appropriate model systems. We characterised soil food webs along each of four retrogressive soil chronosequences situated across a strong regional climate gradient to show that belowground communities are predominantly shaped by changes in fertility rather than climate. Basal consumers showed hump-shaped responses to soil ageing, which were propagated to higher-order consumers. There was a shift in dominance from bacterial to fungal energy channels with increasing soil age, while the root energy channel was most important in intermediate-aged soils. Our study highlights the overarching importance of soil fertility in regulating soil food webs, and indicates that belowground food webs will respond more strongly to shifts in soil resources than climate change.

Landscape context alters cost of living in honeybee metabolism and feeding.

Field metabolic rate (FMR) links the energy budget of an animal with the constraints of its ecosystem, but is particularly difficult to measure for small organisms. Landscape degradation exacerbates environmental adversity and reduces resource availability, imposing higher costs of living for many organisms. Here, we report a significant effect of landscape degradation on the FMR of free-flying Apis mellifera, estimated using 86Rb radio-isotopic turnover. We validated the relationship between 86Rb kb and metabolic rate for worker bees in the laboratory using flow-through respirometry. We then released radioisotopically enriched individuals into a natural woodland and a heavily degraded and deforested plantation. FMRs of worker bees in natural woodland vegetation were significantly higher than in a deforested landscape. Nectar consumption, estimated using 22Na radio-isotopic turnover, also differed significantly between natural and degraded landscapes. In the deforested landscape, we infer that the costs of foraging exceeded energetic availability, and honeybees instead foraged less and depended more on stored resources in the hive. If this is generally the case with increasing landscape degradation, this will have important implications for the provision of pollination services and the effectiveness and resilience of ecological restoration practice.

Non-random food-web assembly at habitat edges increases connectivity and functional redundancy.

Habitat fragmentation dramatically alters the spatial configuration of landscapes, with the creation of artificial edges affecting community structure and dynamics. Despite this, it is not known how the different food webs in adjacent habitats assemble at their boundaries. Here we demonstrate that the composition and structure of herbivore-parasitoid food webs across edges between native and plantation forests are not randomly assembled from those of the adjacent communities. Rather, elevated proportions of abundant, interaction-generalist parasitoid species at habitat edges allowed considerable interaction rewiring, which led to higher linkage density and less modular networks, with higher parasitoid functional redundancy. This was despite high overlap in host composition between edges and interiors. We also provide testable hypotheses for how food webs may assemble between habitats with lower species overlap. In an increasingly fragmented world, non-random assembly of food webs at edges may increasingly affect community dynamics at the landscape level.

Woody plant richness does not influence invertebrate community reassembly trajectories in a tree diversity experiment.

Understanding the relationship between plant diversity and diversity at higher trophic levels is important from both conservation and restoration perspectives. Although there is strong evidence for bottom-up maintenance of biodiversity, this is based largely on studies of simplified grassland systems. Recently, studies in the TreeDivNet global network of tree diversity experiments have begun to test whether these findings are generalizable to more complex ecosystems, such as woodlands. We monitored invertebrate community reassembly over 5 yr of experimental woodland restoration at the TreeDivNet Ridgefield site in southwest Australia, testing the effects of woody plant species richness and herb-layer manipulation on invertebrate community structure and ant species composition. From 2010 to 2014, we sampled ground-dwelling invertebrates using pitfall traps in herbicide vs. no-herbicide subplots nested within each of 10 woody plant treatments varying in richness from zero (bare controls) to eight species, which produced a total of 211, 235 invertebrates, including 98, 979 ants belonging to 74 species. In mixed model analyses, the presence of woody plants was an important driver of faunal community reassembly (relative to bare control plots), but faunal responses to woody plant treatment combinations were idiosyncratic and unrelated to woody plant richness across treatments. We also found that a herbicide-induced reduction in herbaceous plant cover and richness had a positive effect on ant richness and caused more rapid convergence of invertebrate community composition toward the composition of a woodland reference site. These findings show that woody plant richness did not have direct positive effects on the diversity and community reassembly trajectories of higher trophic levels in our woodland system. From a management perspective, this suggests that even low-diversity restoration or carbon sequestration plantings can potentially lead to faunal reassembly outcomes that are comparable to more complex re-planting designs.

Mutualists or parasites? Context-dependent influence of symbiotic fly larvae on carnivorous investment in the Albany pitcher plant.

Carnivorous plants allocate more resources to carnivorous structures under nutrient-limited conditions, and relative investment can also be influenced by animals (infauna) that live in association with these plants and feed on their prey. We investigated these effects within a population of the pitcher plant Cephalotus follicularis containing varying densities of larvae of the fly Badisis ambulans. For plants with a relatively high proportion of adult pitchers, increasing larval density was associated with lower relative leaf allocation to new pitcher buds. For plants with relatively few adult pitchers, however, there was greater relative leaf allocation to pitcher buds with increasing larval density. In a field experiment, there was no significant effect of experimental larval presence or absence on the change in carbon-to-nitrogen (C/N) ratio of plants. Although the direction of the correlation between B. ambulans larvae and relative investment in carnivorous and non-carnivorous structures depends on the relative number of mature structures, whether the larvae enhance or reduce nutrient stress under different conditions remains unclear. The change in C/N was, however, less variable for pitchers that contained larvae, suggesting a stabilizing effect. Eighteen of 52 experimental pitchers were damaged by an unknown species, causing the pitcher fluid to drain. These pitchers were significantly more likely to survive if they contained larvae. These results suggest that the relationship between infauna and host varies with the initial resource status and environmental context of the host plant.

Apparent competition drives community-wide parasitism rates and changes in host abundance across ecosystem boundaries.

Species have strong indirect effects on others, and predicting these effects is a central challenge in ecology. Prey species sharing an enemy (predator or parasitoid) can be linked by apparent competition, but it is unknown whether this process is strong enough to be a community-wide structuring mechanism that could be used to predict future states of diverse food webs. Whether species abundances are spatially coupled by enemy movement across different habitats is also untested. Here, using a field experiment, we show that predicted apparent competitive effects between species, mediated via shared parasitoids, can significantly explain future parasitism rates and herbivore abundances. These predictions are successful even across edges between natural and managed forests, following experimental reduction of herbivore densities by aerial spraying of insecticide over 20 hectares. This result shows that trophic indirect effects propagate across networks and habitats in important, predictable ways, with implications for landscape planning, invasion biology and biological control.