The Leptogenys Roger, 1861 (Formicidae, Ponerinae) of Hong Kong SAR with additional records from Guangdong, China.

Leptogenys is the most diverse genus of the ant subfamily Ponerinae and is widely distributed across the world's tropical and subtropical regions. More than 40 species are known from the Oriental realm displaying a wide range of ecologies, although their life history traits remain poorly understood, and new species are frequently discovered. Here, a faunal review of the genus from Hong Kong SAR, southern China is provided. A total of nine species are recorded, with one new species, Leptogenysgrohli Hamer, Lee & Guénard, sp. nov. described. Ecological and biogeographic information, including new information on reproductive modes for two species are provided with the ergatoids of L.binghamii Forel, 1900 and L.rufidaZhou et al., 2012 described. Additional records for five of these species within the neighbouring province of Guangdong are also provided. Finally, an illustrated key to species known from Hong Kong is presented, as well as notes on each species' distribution, ecology, and behaviour. An updated provincial distributional checklist of the Leptogenys species of Mainland China and Taiwan is also supplied.

Generalized mutualisms promote range expansion in both plant and ant partners.

Mutualism improves organismal fitness, but strong dependence on another species can also limit a species' ability to thrive in a new range if its partner is absent. We assembled a large, global dataset on mutualistic traits and species ranges to investigate how multiple plant-animal and plant-microbe mutualisms affect the spread of legumes and ants to novel ranges. We found that generalized mutualisms increase the likelihood that a species establishes and thrives beyond its native range, whereas specialized mutualisms either do not affect or reduce non-native spread. This pattern held in both legumes and ants, indicating that specificity between mutualistic partners is a key determinant of ecological success in a new habitat. Our global analysis shows that mutualism plays an important, if often overlooked, role in plant and insect invasions.

A checklist of Nylanderia (Hymenoptera: Formicidae: Formicinae) from Hong Kong and Macao SARs, with an illustrated identification key for species in Southeast China and Taiwan.

Species of the ant genus Nylanderia constitute some of the most common ground dwelling Formicinae in tropical and subtropical areas. The genus includes numerous species introduced into new regions, especially within urban or disturbed environments. Here, we review the Nylanderia species found within Hong Kong and Macao, which are both highly urbanized regions and of critical importance to Southern China's policy-makers and non-governmental actors concerned with the regional spread of non-native species. A total of 11 species and 2 morphospecies are recorded in Hong Kong and 6 species and 2 morphospecies are recorded in Macao. We also provide a list of new records for the genus in Hong Kong and Macao, along with taxonomic accounts for all the species recorded. Additionally, the present work provides an illustrated identification key for the species of Nylanderia occurring in Southeast Mainland China and Taiwan. We briefly discuss the possible biogeographical implications of our findings, providing additional support to the exotic status of N. bourbonica in continental Southeast Asia. Lastly, we give our perspective on the ramifications of local inaccessibility to type specimens to current taxonomic practices.

The role of body mass in limiting post heat-coma recovery ability in terrestrial ectotherms.

Under global warming, animal species show shrinking body size responses, cascading deep changes in community structure and ecosystem functions. Although the exact physiological mechanisms behind this phenomenon remain unsolved, smaller individuals may benefit from warming climate more than larger ones. Heat-coma, a physiological state with severe consequences on locomotion ability, is often considered as an "ecological death" scenario under which individuals are unable to escape and exposed to predation, further heat injury, and other hazards. Species are expected to increasingly encounter heat-coma temperature thresholds under warming climate, and body size may be an important trait for thermoregulation in particular for ectotherms. The relationship between heat-coma and shrinking body size remains, however, unclear. Yet, recovery after short-term heat-coma is possible, but little is known about its importance in thermal adaptation and how organismal size correlates with post heat-coma recovery. Here, using ants as a model system, we firstly examined the fate of heat-comatose individuals under field conditions to quantify the ecological benefits of post heat-coma recovery. Then, we quantified ants' recovery ability after heat-coma using a dynamic thermal assay in the laboratory and asked if thermal resilience varies between species with different body mass. Our results confirm that heat-coma represents an inherent ecological death where individuals failed to recover from coma suffer strong predation pressure. Additionally, following phylogenetic signals inclusion, organisms with small mass were more likely to recover, supporting the temperature-size rule in thermal adaptation and recent studies showing a decrease in body size composition of ectotherm community under warmer climatic conditions. Body size as a fundamental trait in ecology thus affects ectotherm survival under thermal stress, which may drive species body size adaptations and community composition under future warming scenarios.

The impact of land use on non-native species incidence and number in local assemblages worldwide.

While the regional distribution of non-native species is increasingly well documented for some taxa, global analyses of non-native species in local assemblages are still missing. Here, we use a worldwide collection of assemblages from five taxa - ants, birds, mammals, spiders and vascular plants - to assess whether the incidence, frequency and proportions of naturalised non-native species depend on type and intensity of land use. In plants, assemblages of primary vegetation are least invaded. In the other taxa, primary vegetation is among the least invaded land-use types, but one or several other types have equally low levels of occurrence, frequency and proportions of non-native species. High land use intensity is associated with higher non-native incidence and frequency in primary vegetation, while intensity effects are inconsistent for other land-use types. These findings highlight the potential dual role of unused primary vegetation in preserving native biodiversity and in conferring resistance against biological invasions.

The global spread and invasion capacities of alien ants.

Many alien species are neither cultivated nor traded but spread unintentionally, and their global movements, capacities to invade ecosystems, and susceptibility to detection by biosecurity measures are poorly known.1,2,3,4 We addressed these key knowledge gaps for ants, a ubiquitous group of stowaway and contaminant organisms that include some of the world's most damaging invasive species.5,6,7,8,9,10 We assembled a dataset of over 146,000 occurrence records to comprehensively map the human-mediated spread of 520 alien ant species across 525 regions globally. From descriptions of the environments in which species were collected within individual regions-such as in imported cargoes, buildings, and outdoor settings-we determined whether different barriers to invasion had been overcome11 and classified alien ant species under three levels of invasion capacity corresponding to increasing biosecurity threat. We found that alien species of different invasion capacities had different sources and sinks globally. For instance, although the diversity of indoor-confined species peaked in the Palearctic realm, that of species able to establish outdoors peaked in the Nearctic and Oceanian realms, and these mainly originated from the Neotropical and Oriental realms. We also found that border interceptions worldwide missed two-thirds of alien species with naturalization capacity, many associated with litter and soil. Our study documents the vast spread of alien ants globally while highlighting avenues for more targeted biosecurity responses, such as prioritizing the screening of imports from regions that are diversity hotspots for species of high invasion capacity and improving the detection of cryptic alien invertebrates dwelling in substrates.

Species richness of bat flies and their associations with host bats in a subtropical East Asian region.

BACKGROUND: Understanding the interactions between bat flies and host bats offer us fundamental insights into the coevolutionary and ecological processes in host-parasite relationships. Here, we investigated the identities, host specificity, and patterns of host association of bat flies in a subtropical region in East Asia, which is an understudied region for bat fly research. METHODS: We used both morphological characteristics and DNA barcoding to identify the bat fly species found on 11 cavernicolous bat species from five bat families inhabiting Hong Kong. We first determined the phylogenetic relationships among bat fly species. Then, we elucidated the patterns of bat-bat fly associations and calculated the host specificity of each bat fly species. Furthermore, we assembled the mitogenomes of three bat fly species from two families (Nycteribiidae and Streblidae) to contribute to the limited bat fly genetic resources available. RESULTS: We examined 641 individuals of bat flies and found 20 species, of which many appeared to be new to science. Species of Nycteribiidae included five Nycteribia spp., three Penicillidia spp., two Phthiridium spp., one Basilia sp., and one species from a hitherto unknown genus, whereas Streblidae included Brachytarsina amboinensis, three Raymondia spp., and four additional Brachytarsina spp. Our bat-bat fly association network shows that certain closely related bat flies within Nycteribiidae and Streblidae only parasitized host bat species that are phylogenetically more closely related. For example, congenerics of Raymondia only parasitized hosts in Rhinolophus and Hipposideros, which are in two closely related families in Rhinolophoidea, but not other distantly related co-roosting species. A wide spectrum of host specificity of these bat fly species was also revealed, with some bat fly species being strictly monoxenous, e.g. nycteribiid Nycteribia sp. A, Phthiridium sp. A, and streblid Raymondia sp. A, while streblid B. amboinensis is polyxenous. CONCLUSIONS: The bat fly diversity and specificity uncovered in this study have shed light on the complex bat-bat fly ecology in the region, but more bat-parasite association studies are still needed in East Asian regions like China as a huge number of unknown species likely exists. We highly recommend the use of DNA barcoding to support morphological identification to reveal accurate host-ectoparasite relationships for future studies.

GABI-I: The global ant biodiversity informatics-island database.

Island systems are known to harbor disproportionate amounts of geographically restricted biodiversity and to experience high rates of species loss, and they ultimately represent critical systems with significant conservation values. However, knowledge of the biodiversity value of insular systems remains highly fragmented and incomplete for many groups of organisms, especially insects. This gap limits our understanding of their global significance for biodiversity and inhibits prioritization for future exploration and conservation efforts. Here, we developed a new database to present current knowledge on ants on 2678 islands globally, based on nominal species, and provide information on their native or exotic status. In total, this database contains 7010 ant species (44.6% of the known global ant fauna) that have been recorded on islands globally from 449,232 records. In addition, this database identifies 108 large islands (area > 200 km2 ) that have received no ant sampling efforts globally. This new data set provides the most comprehensive understanding of ant diversity and composition on islands globally, opening up new opportunities to address questions on a multitude of research questions and fields related to biogeography, ecology, and evolution. This data set also provides a roadmap for future exploration and conservation actions in connection with ants on islands as well as overall ant diversity, with updates available as new records and taxonomic updates are published. There are no copyright restrictions on this database and users should cite this data paper in publications when using the data.

Ant communities in disturbed subtropical landscapes: is climate more important than stochastic processes?

Functional trait-based approaches have provided advances in the understanding of community assembly rules. Broad generalisations remain, however, limited due to the idiosyncratic nature of taxa and ecosystems, especially in tropical regions. We use fine scale resolution (30 m grid) environmental variables and community surveys from nearly 100 secondary tropical forest sites to study niche-based or neutral assembly mechanisms in ground dwelling ants. This provides a unique opportunity for understanding fine scale drivers of taxonomic, functional and phylogenetic diversity in a region characterized by large topographic and climatic differences on a relatively small geographic scale. Precipitation emerged as the most consistent environmental correlate, in shaping taxonomic, phylogenetic and functional aspects of the communities. Functional diversity was weakly associated with topography and temperature related variables. The fourth corner model revealed that femur, scape and mandible length were key traits in response to precipitation, and that communities showed a functional homogenization towards shorter appendages at wetter sites. Our results suggest that neutral and deterministic assembly processes act in concert to shape the taxonomic, phylogenetic and functional aspects of leaf litter ant assemblages. The use of multiple complementary metrics and approaches along environmental gradients are powerful to reveal the subtilities of assembly processes and provide insight into the ways future communities might change.

The abundance, biomass, and distribution of ants on Earth.

Knowledge on the distribution and abundance of organisms is fundamental to understanding their roles within ecosystems and their ecological importance for other taxa. Such knowledge is currently lacking for insects, which have long been regarded as the "little things that run the world". Even for ubiquitous insects, such as ants, which are of tremendous ecological significance, there is currently neither a reliable estimate of their total number on Earth nor of their abundance in particular biomes or habitats. We compile data on ground-dwelling and arboreal ants to obtain an empirical estimate of global ant abundance. Our analysis is based on 489 studies, spanning all continents, major biomes, and habitats. We conservatively estimate total abundance of ground-dwelling ants at over 3 × 1015 and estimate the number of all ants on Earth to be almost 20 × 1015 individuals. The latter corresponds to a biomass of ∼12 megatons of dry carbon. This exceeds the combined biomass of wild birds and mammals and is equivalent to ∼20% of human biomass. Abundances of ground-dwelling ants are strongly concentrated in tropical and subtropical regions but vary substantially across habitats. The density of leaf-litter ants is highest in forests, while the numbers of actively ground-foraging ants are highest in arid regions. This study highlights the central role ants play in terrestrial ecosystems but also major ecological and geographic gaps in our current knowledge. Our results provide a crucial baseline for exploring environmental drivers of ant-abundance patterns and for tracking the responses of insects to environmental change.

The global distribution of known and undiscovered ant biodiversity.

Invertebrates constitute the majority of animal species and are critical for ecosystem functioning and services. Nonetheless, global invertebrate biodiversity patterns and their congruences with vertebrates remain largely unknown. We resolve the first high-resolution (~20-km) global diversity map for a major invertebrate clade, ants, using biodiversity informatics, range modeling, and machine learning to synthesize existing knowledge and predict the distribution of undiscovered diversity. We find that ants and different vertebrate groups have distinct features in their patterns of richness and rarity, underscoring the need to consider a diversity of taxa in conservation. However, despite their phylogenetic and physiological divergence, ant distributions are not highly anomalous relative to variation among vertebrate clades. Furthermore, our models predict that rarity centers largely overlap (78%), suggesting that general forces shape endemism patterns across taxa. This raises confidence that conservation of areas important for small-ranged vertebrates will benefit invertebrates while providing a "treasure map" to guide future discovery.

Trait-mediated competition drives an ant invasion and alters functional diversity.

The assumption that differences in species' traits reflect their different niches has long influenced how ecologists infer processes from assemblage patterns. For instance, many assess the importance of environmental filtering versus classical limiting-similarity competition in driving biological invasions by examining whether invaders' traits are similar or dissimilar to those of residents, respectively. However, mounting evidence suggests that hierarchical differences between species' trait values can distinguish their competitive abilities (e.g. for the same resource) instead of their niches. Whether such trait-mediated hierarchical competition explains invasions and structures assemblages is less explored. We integrate morphological, dietary, physiological and behavioural trait analyses to test whether environmental filtering, limiting-similarity competition or hierarchical competition explain invasions by fire ants on ant assemblages. We detect both competition mechanisms; invasion success is not only explained by limiting similarity in body size and thermal tolerance (presumably allowing the invader to exploit different niches from residents), but also by the invader's superior position in trait hierarchies reflecting competition for common trophic resources. We find that the two mechanisms generate complex assemblage-level functional diversity patterns-overdispersion in some traits, clustering in others-suggesting their effects are likely missed by analyses restricted to a few traits and composite trait diversity measures.

Testing the reliability and ecological implications of ramping rates in the measurement of Critical Thermal maximum.

Critical Thermal maximum (CTmax) is often used to characterize the upper thermal limits of organisms and represents a key trait for evaluating the fitness of ectotherms. The lack of standardization in CTmax assays has, however, introduced methodological problems in its measurement, which can lead to questionable estimates of species' upper thermal limits. Focusing on ants, which are model organisms for research on thermal ecology, we aim to obtain a reliable ramping rate that will yield the most rigorous measures of CTmax for the most species. After identifying three commonly used ramping rates (i.e., 0.2, 0.5 and 1.0°C min-1) in the literature, we experimentally determine their effects on the CTmax values of 27 species measured using dynamic assays. Next, we use static assays to evaluate the accuracy of these values in function of the time of exposure. Finally, we use field observations of species' foraging activities across a wide range of ground temperatures to identify the most biologically relevant CTmax values and to develop a standardized method. Our results demonstrate that the use of a 1°C min-1 ramping rate in dynamic assays yields the most reliable CTmax values for comparing ant species' upper thermal limits, which are further validated in static assays and field observations. We further illustrate how methodological biases in physiological trait measurements can affect subsequent analyses and conclusions on community comparisons between strata and habitats, and the detection of phylogenetic signal (Pagel's λ and Bloomberg's K). Overall, our study presents a methodological framework for identifying a reliable and standardized ramping rate to measure CTmax in ants, which can be applied to other ectotherms. Particular attention should be given to CTmax values obtained with less suitable ramping rates, and the potential biases they may introduce to trait-based research on global warming and habitat conversion, as well as inferences about phylogenetic conservatism.

Warm and arid regions of the world are hotspots of superorganism complexity.

Biologists have long been fascinated by the processes that give rise to phenotypic complexity of organisms, yet whether there exist geographical hotspots of phenotypic complexity remains poorly explored. Phenotypic complexity can be readily observed in ant colonies, which are superorganisms with morphologically differentiated queen and worker castes analogous to the germline and soma of multicellular organisms. Several ant species have evolved 'worker polymorphism', where workers in a single colony show quantifiable differences in size and head-to-body scaling. Here, we use 256 754 occurrence points from 8990 ant species to investigate the geography of worker polymorphism. We show that arid regions of the world are the hotspots of superorganism complexity. Tropical savannahs and deserts, which are typically species-poor relative to tropical or even temperate forests, harbour the highest densities of polymorphic ants. We discuss the possible adaptive advantages that worker polymorphism provides in arid environments. Our work may provide a window into the environmental conditions that promote the emergence of highly complex phenotypes.

Diversity begets diversity: Low resource heterogeneity reduces the diversity of nut-nesting ants in rubber plantations.

One of the most general patterns in ecology is the positive relationship between environmental heterogeneity and local diversity. On the one hand, increased resource heterogeneity provides more resources for diverse consumers in the community. On the other hand, increased structural heterogeneity creates variation in the environment's physical structure, thus allowing the coexistence of diverse species with different environmental requirements. Here, we examined the relative importance of resource and structural heterogeneity in determining the taxonomic, functional, and phylogenetic diversity of nut-nesting ants in natural rainforest and rubber plantation. The species richness of nut-nesting ants was 70% higher in rainforest than in rubber plantation. The clustered functional and phylogenetic structure in rubber plantation suggested a strong effect of environmental filtering in shaping ant functional and phylogenetic structure. Nesting heterogeneity (nut diversity) was the major factor explaining variation in taxonomic, functional, and phylogenetic diversity, suggesting that resource heterogeneity plays a major role in shaping the biodiversity patterns of nut-nesting ants. Overall, these results indicate that decreased resource diversity following the conversion of rainforest to rubber plantation can drive biodiversity loss in nut-nesting ants, through its effect on reducing both ant species, functional, and phylogenetic diversity. The decline in species richness and functional and phylogenetic diversity in the local ant community might have major effects on ecosystem functioning.

ATLANTIC ANTS: a data set of ants in Atlantic Forests of South America.

Ants, an ecologically successful and numerically dominant group of animals, play key ecological roles as soil engineers, predators, nutrient recyclers, and regulators of plant growth and reproduction in most terrestrial ecosystems. Further, ants are widely used as bioindicators of the ecological impact of land use. We gathered information of ant species in the Atlantic Forest of South America. The ATLANTIC ANTS data set, which is part of the ATLANTIC SERIES data papers, is a compilation of ant records from collections (18,713 records), unpublished data (29,651 records), and published sources (106,910 records; 1,059 references), including papers, theses, dissertations, and book chapters published from 1886 to 2020. In total, the data set contains 153,818 ant records from 7,636 study locations in the Atlantic Forest, representing 10 subfamilies, 99 genera, 1,114 ant species identified with updated taxonomic certainty, and 2,235 morphospecies codes. Our data set reflects the heterogeneity in ant records, which include ants sampled at the beginning of the taxonomic history of myrmecology (the 19th and 20th centuries) and more recent ant surveys designed to address specific questions in ecology and biology. The data set can be used by researchers to develop strategies to deal with different macroecological and region-wide questions, focusing on assemblages, species occurrences, and distribution patterns. Furthermore, the data can be used to assess the consequences of changes in land use in the Atlantic Forest on different ecological processes. No copyright restrictions apply to the use of this data set, but we request that authors cite this data paper when using these data in publications or teaching events.

Mangroves are an overlooked hotspot of insect diversity despite low plant diversity.

BACKGROUND: The world's fast disappearing mangrove forests have low plant diversity and are often assumed to also have a species-poor insect fauna. We here compare the tropical arthropod fauna across a freshwater swamp and six different forest types (rain-, swamp, dry-coastal, urban, freshwater swamp, mangroves) based on 140,000 barcoded specimens belonging to ca. 8500 species. RESULTS: We find that the globally imperiled habitat "mangroves" is an overlooked hotspot for insect diversity. Our study reveals a species-rich mangrove insect fauna (>3000 species in Singapore alone) that is distinct (>50% of species are mangrove-specific) and has high species turnover across Southeast and East Asia. For most habitats, plant diversity is a good predictor of insect diversity, but mangroves are an exception and compensate for a comparatively low number of phytophagous and fungivorous insect species by supporting an unusually rich community of predators whose larvae feed in the productive mudflats. For the remaining tropical habitats, the insect communities have diversity patterns that are largely congruent across guilds. CONCLUSIONS: The discovery of such a sizeable and distinct insect fauna in a globally threatened habitat underlines how little is known about global insect biodiversity. We here show how such knowledge gaps can be closed quickly with new cost-effective NGS barcoding techniques.

A new subterranean species and an updated checklist of Strumigenys (Hymenoptera, Formicidae) from Macao SAR, China, with a key to species of the Greater Bay Area.

In the past few decades, sampling of leaf litter with Winkler extractors revealed how abundant and ubiquitous ants from the genus Strumigenys are. It is now known that this genus has the third greatest number of species within the Formicidae family. However, very few subterranean species are known, which may be due to the current under-sampling of the soil stratum. Here, a holistic sampling approach, including the use of subterranean traps, was employed in Macao SAR, China. Subterranean traps allowed the collection of a new cryptic ant species: Strumigenys subterranea Brassard, Leong & Guénard, sp. nov. Moreover, extensive sampling of the leaf litter in secondary forests provided four new species records for the genus. The list of Macanese Strumigenys is here updated, elevating the known diversity in Macao to nine species. Furthermore, to facilitate further research on ants of the Guangdong-Hong Kong-Macao Greater Bay Area, a key to the 29 Strumigenys species known from the region is provided. Lastly, recommendations for sampling methods to assess ant biodiversity underground are discussed. In conclusion, this study highlights the importance of using extensive sampling methods, and suggests that relatively small patches of secondary forests within cities can hold a surprisingly high diversity as well as some undescribed species.

Ants of the Hengduan Mountains: a new altitudinal survey and updated checklist for Yunnan Province highlight an understudied insect biodiversity hotspot.

China's Hengduan Mountain region has been considered one of the most diverse regions in the northern hemisphere. Its stunning topography with many deep valleys and impassable mountain barriers has promoted an astonishing diversification in many groups of organisms including plants, birds, mammals, and amphibians. However, the insect biodiversity in this region is still poorly known. Here, the first checklist of ant species from the Southern Hengduan Mountain region is presented, generated by sampling ant diversity using a wide array of collection methods, including Winkler leaf litter extraction, vegetation beating, and hand collection. 130 species/morphospecies from nine subfamilies and 49 genera were identified. Among them, 17 species from 13 genera represent new records for Yunnan province, and eight species are newly recorded for China. Moreover, we believe 41 novel morphospecies (31% of the total collected taxa) will prove to be new to science. These results highlight the rich ant fauna of this region and strongly support its status as a biodiversity hotspot. The current ant species checklist for the whole of Yunnan Province was updated by recording 550 named species from 99 genera. Taken together, our results suggest that the Yunnan ant fauna still remains under-sampled, and future sampling will likely yield many more species, among them many undescribed ones.

The Ants (Hymenoptera, Formicidae) of Sri Lanka: a taxonomic research summary and updated checklist.

An updated checklist of the ants (Hymenoptera: Formicidae) of Sri Lanka is presented. These include representatives of eleven of the 17 known extant subfamilies with 341 valid ant species in 79 genera. Lioponera longitarsus Mayr, 1879 is reported as a new species country record for Sri Lanka. Notes about type localities, depositories, and relevant references to each species record are given. Accounts of the dubious and some undetermined species from Sri Lanka are also provided. 82 species (24%) are endemic whereas 18 species that are non-native to Sri Lanka are recorded. The list provides a synthesis of the regional taxonomical work carried out to date and will serve as a baseline for future studies on the ant fauna of this biodiversity hotspot.