Local- and landscape-scale variables shape insect diversity in an urban biodiversity hot spot.

Local community structure is shaped by processes acting at local and landscape scales. The relative importance of drivers operating across different spatial scales is difficult to test without observations across regional or latitudinal gradients. Cities exhibit strong but predictable environmental gradients overlaying a mosaic of highly variable but repeated habitat types within a constrained area. Thus, cities present a unique opportunity to explore how both local and landscape factors influence local biotic communities. We used insect communities to examine the interactions among local environmental variables (such as temperature and relative humidity), local habitat characteristics (such as plant community composition), and broad-scale patterns of urbanization (including biophysical, human-built, and socioeconomic variables) on local insect abundance, species richness, and species composition in Los Angeles, a hot, dry, near-desert city. After accounting for seasonal trends, insect species richness and abundance were highest in drier and hotter sites, but the magnitude of local environmental effects varied with the degree of urbanization. In contrast, insect species composition was best predicted by broad-scale urbanization trends, with the more native communities occurring in less urbanized sites and more cosmopolitan insects occurring in highly urbanized sites. However, insect species richness and abundance were >30% higher and insect composition was similar across sites that hosted either native or drought-tolerant plants, regardless of the degree of urbanization. These results demonstrate that urban insect biodiversity is a product of interacting mechanisms working at both local and landscape scales. However, local-scale changes to urban habitats, such as cultivating plants that are adapted to the natural environment nearest the city, can positively impact urban biodiversity regardless of location.

Temperature accounts for the biodiversity of a hyperdiverse group of insects in urban Los Angeles.

The urban heat island effect is a worldwide phenomenon that has been linked to species distributions and abundances in cities. However, effects of urban heat on biotic communities are nearly impossible to disentangle from effects of land cover in most cases because hotter urban sites also have less vegetation and more impervious surfaces than cooler sites within cities. We sampled phorid flies, one of the largest, most biologically diverse families of true flies (Insecta: Diptera: Phoridae), at 30 sites distributed within the central Los Angeles Basin, where we found that temperature and the density of urban land cover are decoupled. Abundance, richness, and community composition of phorids inside urban Los Angeles were most parsimoniously accounted for by mean air temperature in the week preceding sampling. Sites with intermediate mean temperatures had more phorid fly individuals and higher richness. Communities were more even at urban sites with lower minimum temperatures and sites located further away from natural areas, suggesting that communities separated from natural source populations may be more homogenized. Species composition was best explained by minimum temperature. Inasmuch as warmer areas within cities can predict future effects of climate change, phorid fly communities are likely to shift nonlinearly under future climates in more natural areas. Exhaustive surveys of biotic communities within cities, such as the one we describe here, can provide baselines for determining the effects of urban and global climate warming as they intensify.

Distribution and dietary regulation of an associated facultative Rhizobiales-related bacterium in the omnivorous giant tropical ant, Paraponera clavata.

We document a facultative Bartonella-like Rhizobiales bacterium in the giant tropical ant, Paraponera clavata. In a lowland tropical rainforest in Costa Rica, 59 colonies were assayed for the prevalence of the Bartonella-like bacterium (BLB), 14 of which were positive. We addressed three questions: First, how does the prevalence of BLB within colonies vary with environmental conditions? Second, how does diet affect the prevalence of BLB in P. clavata? Third, how does the distribution of BLB among colonies reflect ambient differences in food resources and foraging habits? A variety of environmental variables that may be predictive of the presence of BLB were measured, and diet manipulations were conducted to test whether the prevalence of BLB responded to supplemental carbohydrate or prey. The ambient frequency of BLB is much higher in young secondary forests, but is nearly absent from older secondary forests. The prevalence of BLB inside field colonies increased over the duration of a 2-week carbohydrate supplementation; however, water and prey supplementation did not alter the prevalence of BLB. The diets of the colonies located in young secondary forest, compared to other habitats, have a diet richer in carbohydrates and lower in prey. The abundance of carbohydrate, or the relative lack of N, in a colony's diet influences the occurrence of the BLB microbe in P. clavata. As experimental diet manipulations can affect the facultative presence of an N-cycling microbe, a consistent diet shift in diet may facilitate the emergence of tighter symbioses.

Drivers of arthropod biodiversity in an urban ecosystem.

Our world is becoming increasingly urbanized with a growing human population concentrated around cities. The expansion of urban areas has important consequences for biodiversity, yet the abiotic drivers of biodiversity in urban ecosystems have not been well characterized for the most diverse group of animals on the planet, arthropods. Given their great diversity, comparatively small home ranges, and ability to disperse, arthropods make an excellent model for studying which factors can most accurately predict urban biodiversity. We assessed the effects of (i) topography (distance to natural areas and to ocean) (ii) abiotic factors (mean annual temperature and diurnal range), and (iii) anthropogenic drivers (land value and amount of impervious surface) on the occurrence of six arthropod groups represented in Malaise trap collections run by the BioSCAN project across the Greater Los Angeles Area. We found striking heterogeneity in responses to all factors both within and between taxonomic groups. Diurnal temperature range had a consistently negative effect on occupancy but this effect was only significant in Phoridae. Anthropogenic drivers had mixed though mostly insignificant effects, as some groups and species were most diverse in highly urbanized areas, while other groups showed suppressed diversity. Only Phoridae was significantly affected by land value, where most species were more likely to occur in areas with lower land value. Los Angeles can support high regional arthropod diversity, but spatial community composition is highly dependent on the taxonomic group.

To drink or grasp? How bullet ants (Paraponera clavata) differentiate between sugars and proteins in liquids.

Flexibility in behavior can increase the likelihood that a forager may respond optimally in a fluctuating environment. Nevertheless, physiological or neuronal constraints may result in suboptimal responses to stimuli. We observed foraging workers of the giant tropical ant (also referred to as the "bullet ant"), Paraponera clavata, as they reacted to liquid solutions with varying concentrations of sugar and protein. We show that when protein/sucrose concentration is high, many bullet ants will often try to grasp at the droplet, rather than gather it by drinking. Because P. clavata actively hunt for prey, fixed action patterns and rapid responses to protein may be adaptively important, regardless of the medium in which it is presented. We conclude that, in P. clavata, food-handling decisions are made in response to the nutrient content of the food rather than the texture of the food. Further, we suggest that colonies that maintain a mixture of individuals with consistent fixed or flexible behavioral responses to food-handling decisions may be better adapted to fluctuating environmental conditions, and we propose future studies that could address this.

The ecology of nest movement in social insects.

Social insect colonies are typically mobile entities, moving nests from one location to another throughout the life of a colony. The majority of social insect species-ants, bees, wasps, and termites-have likely adopted the habit of relocating nests periodically. The syndromes of nest relocation include legionary nomadism, unstable nesting, intrinsic nest relocation, and adventitious nest relocation. The emergence of nest movement is a functional response to a broad range of potential selective forces, including colony growth, competition, foraging efficiency, microclimate, nest deterioration, nest quality, parasitism, predation, and seasonality. Considering the great taxonomic and geographic distribution of nest movements, assumptions regarding the nesting biology of social insects should be reevaluated, including our understanding of population genetics, life-history evolution, and the role of competition in structuring communities.

More individuals but fewer species: testing the 'more individuals hypothesis' in a diverse tropical fauna.

A positive relationship between species richness and productivity is often observed in nature, but the causes remain contentious. One mechanism, the 'more individuals hypothesis' (MIH), predicts richness increases monotonically with density, as a function of resource flux. To test the MIH, we manipulated resource abundance in a community of tropical rainforest litter ants and measured richness and density responses. A unimodal relationship between richness and density most closely fitted the control and disturbance (resource removal) treatments in contrast to expectations of the MIH. Resource addition resulted in a monotonic increase in richness relative to density, a shift from the pattern in the control. In the disturbance treatment, richness was greater than in the control, opposite to expectations of the MIH. While large-scale correlations between ant diversity and net primary productivity or temperature are reconcilable with the MIH, key elements of the hypothesis are not supported.

Canopy and litter ant assemblages share similar climate-species density relationships.

Tropical forest canopies house most of the globe's diversity, yet little is known about global patterns and drivers of canopy diversity. Here, we present models of ant species density, using climate, abundance and habitat (i.e. canopy versus litter) as predictors. Ant species density is positively associated with temperature and precipitation, and negatively (or non-significantly) associated with two metrics of seasonality, precipitation seasonality and temperature range. Ant species density was significantly higher in canopy samples, but this difference disappeared once abundance was considered. Thus, apparent differences in species density between canopy and litter samples are probably owing to differences in abundance-diversity relationships, and not differences in climate-diversity relationships. Thus, it appears that canopy and litter ant assemblages share a common abundance-diversity relationship influenced by similar but not identical climatic drivers.

Climatic drivers of hemispheric asymmetry in global patterns of ant species richness.

Although many taxa show a latitudinal gradient in richness, the relationship between latitude and species richness is often asymmetrical between the northern and southern hemispheres. Here we examine the latitudinal pattern of species richness across 1003 local ant assemblages. We find latitudinal asymmetry, with southern hemisphere sites being more diverse than northern hemisphere sites. Most of this asymmetry could be explained statistically by differences in contemporary climate. Local ant species richness was positively associated with temperature, but negatively (although weakly) associated with temperature range and precipitation. After contemporary climate was accounted for, a modest difference in diversity between hemispheres persisted, suggesting that factors other than contemporary climate contributed to the hemispherical asymmetry. The most parsimonious explanation for this remaining asymmetry is that greater climate change since the Eocene in the northern than in the southern hemisphere has led to more extinctions in the northern hemisphere with consequent effects on local ant species richness.

Spurious and functional correlates of the isotopic composition of a generalist across a tropical rainforest landscape.

BACKGROUND: The isotopic composition of generalist consumers may be expected to vary in space as a consequence of spatial heterogeneity in isotope ratios, the abundance of resources, and competition. We aim to account for the spatial variation in the carbon and nitrogen isotopic composition of a generalized predatory species across a 500 ha. tropical rain forest landscape. We test competing models to account for relative influence of resources and competitors to the carbon and nitrogen isotopic enrichment of gypsy ants (Aphaenogaster araneoides), taking into account site-specific differences in baseline isotope ratios. RESULTS: We found that 75% of the variance in the fraction of 15N in the tissue of A. araneoides was accounted by one environmental parameter, the concentration of soil phosphorus. After taking into account landscape-scale variation in baseline resources, the most parsimonious model indicated that colony growth and leaf litter biomass accounted for nearly all of the variance in the delta15N discrimination factor, whereas the delta13C discrimination factor was most parsimoniously associated with colony size and the rate of leaf litter decomposition. There was no indication that competitor density or diversity accounted for spatial differences in the isotopic composition of gypsy ants. CONCLUSION: Across a 500 ha. landscape, soil phosphorus accounted for spatial variation in baseline nitrogen isotope ratios. The delta15N discrimination factor of a higher order consumer in this food web was structured by bottom-up influences - the quantity and decomposition rate of leaf litter. Stable isotope studies on the trophic biology of consumers may benefit from explicit spatial design to account for edaphic properties that alter the baseline at fine spatial grains.

Isotopic evidence for oligotrophication of terrestrial ecosystems.

Human societies depend on an Earth system that operates within a constrained range of nutrient availability, yet the recent trajectory of terrestrial nitrogen (N) availability is uncertain. Examining patterns of foliar N concentrations and isotope ratios (δ15N) from more than 43,000 samples acquired over 37 years, here we show that foliar N concentration declined by 9% and foliar δ15N declined by 0.6-1.6‰. Examining patterns across different climate spaces, foliar δ15N declined across the entire range of mean annual temperature and mean annual precipitation tested. These results suggest declines in N supply relative to plant demand at the global scale. In all, there are now multiple lines of evidence of declining N availability in many unfertilized terrestrial ecosystems, including declines in δ15N of tree rings and leaves from herbarium samples over the past 75-150 years. These patterns are consistent with the proposed consequences of elevated atmospheric carbon dioxide and longer growing seasons. These declines will limit future terrestrial carbon uptake and increase nutritional stress for herbivores.

Bringing ecology blogging into the scientific fold: measuring reach and impact of science community blogs.

The popularity of science blogging has increased in recent years, but the number of academic scientists who maintain regular blogs is limited. The role and impact of science communication blogs aimed at general audiences is often discussed, but the value of science community blogs aimed at the academic community has largely been overlooked. Here, we focus on our own experiences as bloggers to argue that science community blogs are valuable to the academic community. We use data from our own blogs (n = 7) to illustrate some of the factors influencing reach and impact of science community blogs. We then discuss the value of blogs as a standalone medium, where rapid communication of scholarly ideas, opinions and short observational notes can enhance scientific discourse, and discussion of personal experiences can provide indirect mentorship for junior researchers and scientists from underrepresented groups. Finally, we argue that science community blogs can be treated as a primary source and provide some key points to consider when citing blogs in peer-reviewed literature.

Non-native Ants Are Smaller than Related Native Ants.

I compare the sizes of non-native and native ants to evaluate how worker size may be related to the ability of a species to invade new habitats. I compare the size of 78 non-native ant species belonging to 26 genera with the size of native congeneric species; native ants are larger than non-native ants in 22 of 26 genera. Ants were sorted by genera into fighting and nonfighting groups, based on observations of interspecific interactions with other ant species. In all of the genera with monomorphic worker castes that fight during competition, the non-native species were smaller than the native species. The genera that engage in combat had a higher frequency of significantly smaller size in non-native ants. I selected Wasmannia auropunctata for further studies, to compare native and non-native populations. Specimens of W. auropunctata from non-native populations were smaller than conspecific counterparts from its native habitat. I consider hypotheses to explain why non-native ants are smaller in size than native ants, including the role of colony size in interspecific fights, changes in life history, the release from intraspecific fighting, and climate. The discovery that fighting non-natives are smaller than their closest native relatives may provide insight into the mechanisms for success of non-native species, as well as the role of worker size and colony size during interspecific competition.

Tradeoffs in the evolution of caste and body size in the hyperdiverse ant genus Pheidole.

The efficient investment of resources is often the route to ecological success, and the adaptability of resource investment may play a critical role in promoting biodiversity. The ants of the "hyperdiverse" genus Pheidole produce two discrete sterile castes, soldiers and minor workers. Within Pheidole, there is tremendous interspecific variation in proportion of soldiers. The causes and correlates of caste ratio variation among species of Pheidole remain enigmatic. Here we test whether a body size threshold model accounts for interspecific variation in caste ratio in Pheidole, such that species with larger body sizes produce relatively fewer soldiers within their colonies. We evaluated the caste ratio of 26 species of Pheidole and found that the body size of workers accounts for interspecific variation in the production of soldiers as we predicted. Twelve species sampled from one forest in Costa Rica yielded the same relationship as found in previously published data from many localities. We conclude that production of soldiers in the most species-rich group of ants is regulated by a body size threshold mechanism, and that the great variation in body size and caste ratio in Pheidole plays a role in niche divergence in this rapidly evolving taxon.