Top-down response to spatial variation in productivity and bottom-up response to temporal variation in productivity in a long-term study of desert ants.

Under the Ecosystem Exploitation Hypothesis ecosystem productivity predicts trophic complexity, but it is unclear if spatial and temporal drivers of productivity have similar impacts. Long-term studies are necessary to capture temporal impacts on trophic structure in variable ecosystems such as deserts. We sampled ants and measured plant resources in the Simpson Desert, central Australia over a 22-year period, during which rainfall varied 10-fold. We sampled dune swales (higher nutrient) and crests (lower nutrient) to account for spatial variation in productivity. We asked how temporal and spatial variation in productivity affects the abundance of ant trophic guilds. Precipitation increased vegetation cover, with the difference more pronounced on dune crests; seeding and flowering also increased with precipitation. Generalist activity increased over time, irrespective of productivity. Predators were more active in more productive (swale) habitat, i.e. spatial impacts of productivity were greatest at the highest trophic level. By contrast, herbivores (seed harvesters and sugar feeders) increased with long-term rainfall; seed harvesters also increased as seeding increased. Temporal impacts of productivity were therefore greatest for low trophic levels. Whether productivity variation leads to top-down or bottom-up structured ecosystems thus depends on the scale and dimension (spatial or temporal) of productivity.

Long-term responses of desert ant assemblages to climate.

Productivity is a key driver of ecosystem structure and function, so long-term studies are critical to understanding ecosystems with high temporal variation in productivity. In some deserts, productivity, driven by moisture availability, varies immensely over time (rainfall) and space (landscape factors). At high productivity, species richness is expected to be driven in opposing directions by abundance (More Individuals Hypothesis - MIH) and competition. While studies investigating the impacts of spatial variation in productivity on community structure are common, the impacts of temporal variability on productivity are poorly understood. We tested how well rainfall predicted the activity, species numbers and assemblage composition of ants and if responses were moderated by landscape position. We also asked whether the number of species (richness per sampling unit and estimated species richness) responded directly to rainfall or was moderated by ant activity or competition from dominant ants. Over a 22-year period, when annual rainfall fluctuated between 79 mm and 570 mm, we sampled ants using pitfall traps in paired dune and swale habitats in the Simpson Desert, Australia. We used climate records over this period to model changes in ant assemblages. Activity of dominant ants responded primarily to long-term rainfall, increasing exponentially, while subordinate ants responded to short-term weather and time. Consistent with the MIH, the number of ant species was best predicted by activity, particularly that of subordinate ants. Activity of dominant ants had a declining positive effect on numbers of species. Landscape position strongly predicted species composition, while long-term rainfall determined composition at genus level but not species level. Over time, species composition fluctuated, but several genera consistently increased in activity. Productivity moderators such as long-term rainfall and landscape position are key drivers of ant activity and composition in the study ecosystem, acting indirectly on numbers of species. Numbers of species were explained largely by ant activity, making a strong case for the MIH, but not competition. Longer periods of low rainfall may indirectly reduce species richness in desert ecosystems. However, a trend to increasing richness over time may indicate that conservation management can ameliorate this impact.

Dominance-diversity relationships in ant communities differ with invasion.

The relationship between levels of dominance and species richness is highly contentious, especially in ant communities. The dominance-impoverishment rule states that high levels of dominance only occur in species-poor communities, but there appear to be many cases of high levels of dominance in highly diverse communities. The extent to which dominant species limit local richness through competitive exclusion remains unclear, but such exclusion appears more apparent for non-native rather than native dominant species. Here we perform the first global analysis of the relationship between behavioral dominance and species richness. We used data from 1,293 local assemblages of ground-dwelling ants distributed across five continents to document the generality of the dominance-impoverishment rule, and to identify the biotic and abiotic conditions under which it does and does not apply. We found that the behavioral dominance-diversity relationship varies greatly, and depends on whether dominant species are native or non-native, whether dominance is considered as occurrence or relative abundance, and on variation in mean annual temperature. There were declines in diversity with increasing dominance in invaded communities, but diversity increased with increasing dominance in native communities. These patterns occur along the global temperature gradient. However, positive and negative relationships are strongest in the hottest sites. We also found that climate regulates the degree of behavioral dominance, but differently from how it shapes species richness. Our findings imply that, despite strong competitive interactions among ants, competitive exclusion is not a major driver of local richness in native ant communities. Although the dominance-impoverishment rule applies to invaded communities, we propose an alternative dominance-diversification rule for native communities.

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.