Benjamin D. Hoffmann

Impact of an introduced ant on native rain forest invertebrates: Pheidole megacephala in monsoonal Australia

Pheidole megacephala is an exotic ant species that has severely affected native invertebrate biodiversity throughout the tropics. Its impacts have been documented extensively in relatively depauperate invertebrate communities, but not in species-rich habitats such as tropical rain forests. Here we describe the local distribution of P. megacephala and its impacts on native invertebrate assemblages in and around a rain forest patch at Howard Springs, in Australias monsoonal tropics. P. megacephala was found to be confined to a single area of approximately 25 ha, with its distribution centered on drainage lines and the rain forest. Significant but weak correlations were found between its abundance and vegetative canopy cover (positive) and distance from the rain forest (negative). In the most heavily infested area within the rain forest, the abundance of P. megacephala was 37–110 times that of total native ant abundance found within uninfested plots, as measured by pitfall traps. The abundance and richness of native ants and other invertebrates were significantly reduced in litter samples, pitfall catches and foliage beats where P. megacephala was present, inversely relative to the abundance of P. megacephala. Only two individuals of a single native ant species were found within the most infested plot, with native ant richness being reduced to about half in the least infested plot. The most persistent functional groups of native ants in infested plots were Cryptic species, which forage primarily within soil and leaf litter, and Opportunists, which exhibit highly generalised foraging behaviour. The highest abundance of P. megacephala corresponded with a 42–85% decrease in the abundance of other native invertebrates. Insect larvae were totally absent from foliage beats collected at the most heavily infested plot. P. megacephala was found overall to be expanding its range, averaging 12 m range expansion in the dry season and contracting 7 m in the wet season. It is able to spread into surrounding savanna habitats by occupying relatively sheltered microsites, such as beneath logs and at the bases of trees. However, it is unlikely to attain high population densities in open savanna habitats because of its relative intolerance of desiccation, and the prevalence of behaviourally dominant native ant species. Howard Springs is currently the only rain forest patch in monsoonal Australia known to be infested by P. megacephala, but clearly this ant is a serious potential threat to the regions rain forest invertebrate fauna.

Climate change, sea-level rise, and conservation: keeping island biodiversity afloat.

Island conservation programs have been spectacularly successful over the past five decades, yet they generally do not account for impacts of climate change. Here, we argue that the full spectrum of climate change, especially sea-level rise and loss of suitable climatic conditions, should be rapidly integrated into island biodiversity research and management.

Introduced and Invasive Species in Novel Rangeland Ecosystems: Friends or Foes?

Abstract Globally, new combinations of introduced and native plant and animal species have changed rangelands into novel ecosystems. Whereas many rangeland stakeholders (people who use or have an interest in rangelands) view intentional species introductions to improve forage and control erosion as beneficial, others focus on unintended costs, such as increased fire risk, loss of rangeland biodiversity, and threats to conservation efforts, specifically in nature reserves and parks. These conflicting views challenge all rangeland stakeholders, especially those making decisions on how best to manage novel ecosystems. To formulate a conceptual framework for decision making, we examined a wide range of novel ecosystems, created by intentional and unintentional introductions of nonnative species and land-use–facilitated spread of native ones. This framework simply divides decision making into two types: 1) straightforward–certain, and 2) complex–uncertain. We argue that management decisions to retain novel ecosystems are certain when goods and services provided by the system far outweigh the costs of restoration, for example in the case of intensively managed Cenchrus pastures. Decisions to return novel ecosystems to natural systems are also certain when the value of the system is low and restoration is easy and inexpensive as in the case of biocontrol of Opuntia infestations. In contrast, decisions whether to retain or restore novel ecosystems become complex and uncertain in cases where benefits are low and costs of control are high as, for example, in the case of stopping the expansion of Prosopis and Juniperus into semiarid rangelands. Decisions to retain or restore novel ecosystems are also complex and uncertain when, for example, nonnative Eucalyptus trees expand along natural streams, negatively affecting biodiversity, but also providing timber and honey. When decision making is complex and uncertain, we suggest that rangeland managers utilize cost–benefit analyses and hold stakeholder workshops to resolve conflicts. Resumen Mundialmente, nuevas combinaciones de plantas introducidas e inducidas y especies de animales han cambiado los pastizales a nuevos ecosistemas. Mientras que muchos de los interesados en los pastizales (personas que usan o tienen interés en los pastizales) ven un beneficio en la introducción de especies para el mejoramiento de la producción de forraje y control de la erosión, otros se interesan en los costos no planeados tales como el aumento en el riesgo de fuego, pérdida de biodiversidad en los pastizales y amenazas en los esfuerzos de conservación especialmente en reservas naturales y parques. Estos puntos de vista conflictivos son retos para todos los interesados en los pastizales, especialmente para la toma de decisiones en cómo manejar mejor los ecosistemas nuevos. Para formular un modelo conceptual para toma de decisiones, examinamos un amplio rango de ecosistemas nuevos, creados de manera intencional y no intencional de especies no nativas y el uso de tierras que facilitan la expansión de especies nativas. Este modelo simplemente divide la toma de decisiones en dos tipos: 1) francamente–seguro y 2) complejo–no seguro. Discutimos que las decisiones de manejo para mantener ecosistemas nuevos son seguras cuando los bienes y servicios proporcionados por el sistema sobrepasan por mucho el costo de restauración, por ejemplo en el caso de las praderas intensivas de Cenchrus. Las decisiones para devolver ecosistemas nuevos a sistemas naturales son también seguras cuando el valor del sistema es bajo y la restauración es fácil y barata como en el caso del control biológico de las infestaciones de Opuntia. En contraste, las decisiones ya sea de mantener o recuperar ecosistemas nuevos se complican y son inciertas en casos donde los beneficios son bajos y los costos altos, por ejemplo en el caso de detener la expansión del Prosopis y Juniperus en los pastizales semiáridos. También las decisiones para mantener o renovar un ecosistema nuevo son difíciles e inciertas cuando por ejemplo, especies no nativas como el Eucalipto se extienden sobre arroyos naturales afectando negativamente la biodiversidad pero también proveyendo madera y miel. Cuando el proceso de toma de decisiones es complejo e incierto sugerimos que los manejadores de pastizales usen el análisis de costo beneficio y talleres entre los interesados para resolver conflictos.

Climate mediates the effects of disturbance on ant assemblage structure

Many studies have focused on the impacts of climate change on biological assemblages, yet little is known about how climate interacts with other major anthropogenic influences on biodiversity, such as habitat disturbance. Using a unique global database of 1128 local ant assemblages, we examined whether climate mediates the effects of habitat disturbance on assemblage structure at a global scale. Species richness and evenness were associated positively with temperature, and negatively with disturbance. However, the interaction among temperature, precipitation and disturbance shaped species richness and evenness. The effect was manifested through a failure of species richness to increase substantially with temperature in transformed habitats at low precipitation. At low precipitation levels, evenness increased with temperature in undisturbed sites, peaked at medium temperatures in disturbed sites and remained low in transformed sites. In warmer climates with lower rainfall, the effects of increasing disturbance on species richness and evenness were akin to decreases in temperature of up to 9°C. Anthropogenic disturbance and ongoing climate change may interact in complicated ways to shape the structure of assemblages, with hot, arid environments likely to be at greatest risk.

Molecular phylogeny of Indo-Pacific carpenter ants (Hymenoptera: Formicidae, Camponotus) reveals waves of dispersal and colonization from diverse source areas

Ants that resemble Camponotus maculatus (Fabricius, 1782) present an opportunity to test the hypothesis that the origin of the Pacific island fauna was primarily New Guinea, the Philippines, and the Indo‐Malay archipelago (collectively known as Malesia). We sequenced two mitochondrial and four nuclear markers from 146 specimens from Pacific islands, Australia, and Malesia. We also added 211 specimens representing a larger worldwide sample and performed a series of phylogenetic analyses and ancestral area reconstructions. Results indicate that the Pacific members of this group comprise several robust clades that have distinctly different biogeographical histories, and they suggest an important role for Australia as a source of Pacific colonizations. Malesian areas were recovered mostly in derived positions, and one lineage appears to be Neotropical. Phylogenetic hypotheses indicate that the orange, pan‐Pacific form commonly identified as C. chloroticus Emery 1897 actually consists of two distantly related lineages. Also, the lineage on Hawaiʻi, which has been called C. variegatus (Smith, 1858), appears to be closely related to C. tortuganus Emery, 1895 in Florida and other lineages in the New World. In Micronesia and Polynesia the C. chloroticus‐like species support predictions of the taxon‐cycle hypothesis and could be candidates for human‐mediated dispersal.

Quantification of Supercolonial Traits in the Yellow Crazy Ant, Anoplolepis gracilipes

Abstract Supercoloniality is a social structure displayed by many invasive ant species, but there has been surprisingly little research quantifying the extent to which individual species display traits underlying such social organisation. This study quantifies three traits for the yellow crazy ant, Anoplolepis gracilipes Smith (Hymenoptera: Formicidae): little or no aggression between workers from different nests; the exchange of workers among nests; and resource exchange among nests, as well as supercolony structure arising from patterns of distribution and density of detections. Supercolonies displayed a structural continuum from being small ( < 10 ha) and “aggregated” with great continuity among detections through to being large (>10,000 ha) and “diffuse” with little continuity among detections. Smaller supercolonies had greater ant densities than larger supercolonies. In laboratory trials, no aggression was observed between workers from different nests sourced from different supercolonies, and paired nests merged within 24 hours. Workers lacked nest fidelity by rapidly populating artificial nests containing alien queens. The daily worker turnover rate per nest was estimated to be below 20%. Resources were readily moved among nests, with a resource being detected up to 13 m away from a source within 24 hours, and as far as 32 m after four days. The rate and distance of resource movement increased with increasing worker and nest density. This research has demonstrated that A. gracilipes displays supercoloniality equivalent to that of the well-studied Argentine ant Linepithema humile . Quantification of these traits is required for other supercolonial species to improve our understanding of this social strategy, especially for invasive ants to aid in understanding factors that promote invasion success and to improve management.

The end of Invasion Biology: intellectual debate does not equate to nonsensical science

Valéry et al. recently proposed to end the field of Invasion Biology on the grounds that it is based on an inadequate definition of the concept of biological invasion and that, as exotic species, native species should also be called invasive whenever they outbreak. We argue, on the contrary, that the sudden demographic dominance of native species cannot be termed invasion. Moreover, we claim that the suggestion of ending a fruitful and useful discipline because it does not conform to a subjective definition or because it still encompasses some debatable ideas and unresolved questions is both irrelevant and excessive. We believe that the thousands of researchers working in this discipline do not perform nonsensical science, and that their efforts to understand and limit biological invasions are compatible with debating on the key concept of that field.

A global database of ant species abundances

What forces structure ecological assemblages? A key limitation to general insights about assemblage structure is the availability of data that are collected at a small spatial grain (local assemblages) and a large spatial extent (global coverage). Here, we present published and unpublished data from 51 ,388 ant abundance and occurrence records of more than 2,693 species and 7,953 morphospecies from local assemblages collected at 4,212 locations around the world. Ants were selected because they are diverse and abundant globally, comprise a large fraction of animal biomass in most terrestrial communities, and are key contributors to a range of ecosystem functions. Data were collected between 1949 and 2014, and include, for each geo-referenced sampling site, both the identity of the ants collected and details of sampling design, habitat type, and degree of disturbance. The aim of compiling this data set was to provide comprehensive species abundance data in order to test relationships between assemblage structure and environmental and biogeographic factors. Data were collected using a variety of standardized methods, such as pitfall and Winkler traps, and will be valuable for studies investigating large-scale forces structuring local assemblages. Understanding such relationships is particularly critical under current rates of global change. We encourage authors holding additional data on systematically collected ant assemblages, especially those in dry and cold, and remote areas, to contact us and contribute their data to this growing data set.

The conundrum of the yellow crazy ant (Anoplolepis gracilipes) reproductive mode: no evidence for dependent lineage genetic caste determination

Asexual reproduction and hybridisation are often found among highly invasive plants and marine invertebrates. Recently, it has been suggested that clonality may enhance the success of invasive ants. In contrast, obligate hybridisation (dependent lineage genetic caste determination or DL GCD in ants) may decrease the chances of population persistence if one lineage is less prevalent than the other (asymmetry in lineage ratio). Genetic data available for the invasive yellow crazy ant (Anoplolepis gracilipes) suggest that it has an unconventional mode of reproduction that may involve asexual reproduction by workers or queens, or a form of genetic caste determination. Here, we investigated whether A. gracilipes reproduction involved DL GCD. The potential for worker reproduction was also assessed. We used microsatellite markers to assess the population structure of A. gracilipes workers, males, queens and sperm in queen spermathecae, from field collections in Arnhem Land. We found that a single queen lineage is present in Arnhem Land. The presence of a single lineage of queens discounts the possibility of DL GCD. Population structure separated queens and workers into different lineages, suggesting that these castes are determined genetically in A. gracilipes, or the mode of reproduction differs between workers and queens. Evidence for worker reproduction was weak. We conclude that the reproductive mode of A. gracilipes does not involve DL GCD. The resolution of the reproductive mode of A. gracilipes is complicated by a high prevalence of diploid males. The determination of the A. gracilipes reproductive mode remains a fascinating research question, and its resolution will improve our understanding of the contribution of the reproductive system to invasion success.

Genetic diversity is positively associated with fine‐scale momentary abundance of an invasive ant

Many introduced species become invasive despite genetic bottlenecks that should, in theory, decrease the chances of invasion success. By contrast, population genetic bottlenecks have been hypothesized to increase the invasion success of unicolonial ants by increasing the genetic similarity between descendent populations, thus promoting co-operation. We investigated these alternate hypotheses in the unicolonial yellow crazy ant, Anoplolepis gracilipes, which has invaded Arnhem Land in Australias Northern Territory. We used momentary abundance as a surrogate measure of invasion success, and investigated the relationship between A. gracilipes genetic diversity and its abundance, and the effect of its abundance on species diversity and community structure. We also investigated whether selected habitat characteristics contributed to differences in A. gracilipes abundance, for which we found no evidence. Our results revealed a significant positive association between A. gracilipes genetic diversity and abundance. Invaded communities were less diverse and differed in structure from uninvaded communities, and these effects were stronger as A. gracilipes abundance increased. These results contradict the hypothesis that genetic bottlenecks may promote unicoloniality. However, our A. gracilipes study population has diverged since its introduction, which may have obscured evidence of the bottleneck that would likely have occurred on arrival. The relative importance of genetic diversity to invasion success may be context dependent, and the role of genetic diversity may be more obvious in the absence of highly favorable novel ecological conditions.