Alejandro G. Farji-Brener

Putting plant resistance traits on the map: a test of the idea that plants are better defended at lower latitudes

• It has long been believed that plant species from the tropics have higher levels of traits associated with resistance to herbivores than do species from higher latitudes. A meta-analysis recently showed that the published literature does not support this theory. However, the idea has never been tested using data gathered with consistent methods from a wide range of latitudes. • We quantified the relationship between latitude and a broad range of chemical and physical traits across 301 species from 75 sites world-wide. • Six putative resistance traits, including tannins, the concentration of lipids (an indicator of oils, waxes and resins), and leaf toughness were greater in high-latitude species. Six traits, including cyanide production and the presence of spines, were unrelated to latitude. Only ash content (an indicator of inorganic substances such as calcium oxalates and phytoliths) and the properties of species with delayed greening were higher in the tropics. • Our results do not support the hypothesis that tropical plants have higher levels of resistance traits than do plants from higher latitudes. If anything, plants have higher resistance toward the poles. The greater resistance traits of high-latitude species might be explained by the greater cost of losing a given amount of leaf tissue in low-productivity environments.

Leaf-cutting ant nests near roads increase fitness of exotic plant species in natural protected areas

Understanding the mechanisms that promote the invasion of natural protected areas by exotic plants is a central concern for ecology. We demonstrated that nests of the leaf-cutting ant, Acromyrmex lobicornis, near roadsides promote the abundance, growth and reproduction of two exotic plant species, Carduus nutans and Onopordum acanthium, in a national park in northern Patagonia, Argentina and determine the mechanisms that produce these effects. Refuse dumps (RDs) from ant nests have a higher nutrient content than nearby non-nest soils (NNSs); foliar nutrient content and their 15N isotopic signature strongly suggest that plants reach and use these nutrients. Both species of exotic plants in RDs were 50–600% more abundant; seedlings had 100–1000% more foliar area and root and leaf biomass; and adult plants produced 100–300% more seeds than nearby NNS plants. Plants can thus gain access to and benefit from the nutrient content of ant RD, supporting the hypotheses that enhanced resource availability promotes exotic plant performance that could increase the likelihood of biological invasions. The two exotics produce an estimated of 8 385 000 more seeds ha−1 in areas with ant nests compared with areas without; this exceptional increase in seed production represents a potential threat to nearby non-invaded communities. We propose several management strategies to mitigate this threat. Removal efforts of exotics should be focused on ant RDs, where plants are denser and represent a higher source of propagules.

Seedling recruitment in a semi-arid Patagonian steppe: Facilitative effects of refuse dumps of leaf-cutting ants

Abstract Question: What is the influence of refuse dumps of leaf-cutting ants on seedling recruitment under contrasting moisture conditions in a semi-arid steppe? Location: Northwestern Patagonia, Argentina. Methods: In a greenhouse experiment, we monitored seedling recruitment in soil samples from refuse dumps of nests of the leaf-cutting ant Acromyrmex lobicornis and non-nest sites, under contrasting moisture conditions simulating wet and dry growing seasons. Results: The mean number of seedling species and individuals were higher in wet than in dry plots, and higher in refuse dump plots than in non-nest soil plots. The positive effect of refuse dumps on seedling recruitment was greater under low moisture conditions. Both the accumulation of discarded seeds by leaf-cutting ants and the passive trapping of blowing-seeds seems not explain the increased number of seeds in refuse dumps. Conversely, refuse dumps have higher water retention capacity and nutrient content than adjacent non-nest soils, allowing the recruitment of a greater number of species and individual seedlings. Conclusions: Nests of A. lobicornis may play an important role in plant recruitment in the study area, allowing a greater number of seedlings and species to be present, hence resulting in a more diverse community. Moreover, leaf-cutting ant nests may function as nurse elements, generating safe sites that enhance the performance of neighbouring seedlings mainly during the driest, stressful periods. Nomenclature: Correa (1969–1998).

Effect of fire on ground beetles and ant assemblages along an environmental gradient in NW Patagonia: Does habitat type matter?

ABSTRACT The response of beetle and ant assemblages to fire (2–5 y old) and the dependence of that response on habitat type were analyzed. Using pitfall traps, beetles (Coleoptera) and ants (Formicidae) were sampled in replicated forest, scrub, and steppe areas including paired unburnt and burnt plots. A total of 176 species of beetles (8245 individuals) and 22 species of ants (115,056 individuals) were captured. Most beetle families (65%) were captured in the forest, while most ant species were most frequently captured in the steppe (45%). Beetle abundance was the same in unburnt and burnt plots in the three habitats, but ant abundance was higher in burnt than in unburnt plots. Fire usually decreased species richness of both taxa, but the strength of this effect depended on the habitat type. Coleoptera richness was lower in burnt than in unburnt plots in the forest and steppe but was the same in unburnt and burnt scrub plots. Ant richness was lower in burnt forest and scrub plots and was similar between burnt and unburnt steppe plots. For both taxa, species composition of the forest assemblages was different between unburnt and burnt forest plots (difference almost significant for ants) but not between unburnt and burnt steppe plots. Beetle species composition in the scrub was different between unburnt and burnt plots, but ant species composition was the same. Our results support the idea that the consequences of fire on native beetle and ant assemblages depend on the habitat type but also on the taxonomic group under analysis. Beetles and ants showed differences in their habitat preference and in the way that they use the habitats (e.g., preference for vegetated or bare soil patches) along the environmental gradient. These differences explain why beetles were in general more severely affected than ants, and why fire differentially affected both taxa in the scrub (habitat of intermediate complexity). A good knowledge of the habitat utilization by different groups at the local scale and a regional perspective (e.g., habitat preference along an environmental gradient) are necessary to fully understand the effect of disturbances on native arthropod assemblages.

Waste Management in the Leaf-Cutting Ant Acromyrmex lobicornis: Division of Labour, Aggressive Behaviour, and Location of External Refuse Dumps

In leaf-cutting ants, the handling of waste materials from the fungus culture increases the risk of infection. Consequently, ants should manage their waste in a way that minimizes the spread of diseases. We investigated whether in Acromyrmex lobicornis, waste-worker ants (a) also perform roles in foraging or mound maintenance, (b) are morphologically different than other ant workers, and (c) are aggressively discriminated by other worker ants from the same colony. In addition, we investigated whether the location of external waste piles minimizes the probability that wastes spread to the ant nest. In the field, we (a) marked with different colours waste-workers, foragers and mound-workers and monitored whether these ants interchanged their tasks; (b) measured head width, head length, hind femur length and total length of waste-workers; foragers and mound-workers; (c) forced field encounters between waste-workers and foragers, and (d) measured the cardinal orientation of the waste piles in relation to the colony mound. Waste-worker ants did not perform other function outside the nest; neither foragers nor mound-workers managed the waste. Moreover, waste-workers were smaller than foragers and mound-workers, and were attacked if they tried to enter their nest using foraging entrances. The location of external refuse dumps also appears to reduce contamination risks. Waste piles always were down-slope, and often followed the prevailing wind direction. The importance of behaviours such as the division of labour, aggressions against waste-workers and nest compartmentalization (i.e., the orientation of external waste piles) to minimize the spread of pathogens is discussed.

The importance of treefall gaps as foraging sites for leaf-cutting ants depends on forest age

Variation in local abundance of herbivores is influenced by several abiotic and biotic factors, but it may often be determined by the local availability of plant resources (Quinn et al. 1997). Leaf-cutting ants, common defoliators of plants in the Neotropics, are more abundant in early successional forests than in old-growth forest (Jaffe & Vilela 1989, Vasconcelos & Cherrett 1995). A recent study showed that the high density of mature leaf-cutting ant nests in early successional forests was mainly determined by the availability of pioneer plant species, largely preferred by the leafcutters (Farji-Brener 2001). The dominance of pioneer species in early successional forests decreases foraging costs for leaf-cutting ant colonies, and thus these habitats can support a high density of mature colonies. On the other hand, the effective defence mechanisms of shade-tolerant species, which dominate primary habitats, could explain the low density of mature leaf-cutting ant colonies in old-growth forests (palatable forage hypothesis, see Farji-Brener 2001). According to this hypothesis, it is proposed that leaf-cutting ant colonies in old-growth forests should concentrate their foraging effort in treefall gaps, in which sun and pioneer plant species (preferred by leafcutters), are readily available (Farji-Brener 2001, Nichols-Orians 1991 a , b ; Wetterer 1994). However, this prediction of the palatable forage hypothesis has not yet been explicitly documented.

Ecological engineering by a native leaf-cutting ant increases the performance of exotic plant species

Numerous mechanisms are proposed to explain why exotic plants successfully invade natural communities. However, the positive effects of native engineers on exotic plant species have received less consideration. We tested whether the nutrient-rich soil patches created by a native ecological engineer (refuse dumps from the leaf-cutting ant Acromyrmex lobicornis) increase the performance of exotic more than native plants. In a greenhouse experiment, individuals from several native and exotic species were planted in pots with refuse dumps (RDs) and non-nest soils (NNSs). Total plant biomass and foliar nutrient content were measured at the end of the experiment. We also estimated the cover of exotic and native plant species in external RDs from 54 field ant nests and adjacent areas. Greenhouse plants showed more biomass and foliar nutrient content in RDs than in NNS pots. Nevertheless, differences in the final mean biomass among RD and NNS plants were especially great in exotics. Accordingly, the cover of exotic plants was higher in field RDs than in adjacent, non-nest soils. Our results demonstrated that plants can benefit from the enhanced nutrient content of ant RDs, and that A. lobicornis acts as an ecosystem engineer, creating a substrate that especially increases the performance of exotics. This supports the fluctuating resource hypothesis as a mechanism to promote biological invasions, and illustrates how this hypothesis may operate in nature. Since ant nests and exotic plants are more common in disturbed than in pristine environments, the role of ant nests in promoting biological invasions might be of particular interest. Proposals including the use of engineer species to restore disturbed habitats should be planned with caution because of their potential role in promoting invasions.

Competition for food between the exotic wasp Vespula germanica and the native ant assemblage of NW Patagonia: evidence of biotic resistance?

The success of a biological invasion may depend on the interactions between the invader and the native biota. However, little experimental evidence demonstrates whether local species can successfully compete with exotics. We experimentally determined the existence of competition for food between the exotic wasp Vespula germanica, one of the most recent Patagonian invaders, and the native ant assemblage. Both wasps and ants are generalist predators and scavengers, sharing habitat and food resources. We selected 30 sites within scrubland habitats where both ants and wasps were present. At each site, we placed containers with protein baits under three treatments: wasp exclusion, ant exclusion, and control (i.e., free access for wasps and ants). Ant exclusion increased the number of wasps (with regard to a control), but wasp exclusion did not affect ant abundance. This result suggests that native ants affect the foraging activity of exotic wasps but not vice versa. Aggressive behaviors and worker aggregation may explain the competitive advantage of ants. Ants bite wasp legs and massively aggregate on food sources, physically limiting the landing of wasps on baits. If the outcome of interactions at baits reported here influence wasp population-level parameters, this competitive interaction could be one of the factors explaining the low abundance of this exotic wasp in NW Patagonia in comparison with other invaded regions.

A meta-analysis of leaf-cutting ant nest effects on soil fertility and plant performance

1. Leaf‐cutting ants (LCAs) are considered as one of the most important agents of soil disturbances that affect vegetation patterns, but these assertions are based on isolated studies or anecdotal data. In this study, meta‐analysis techniques were used to quantitatively analyse the generality of these effects and determine some of their sources of variation.

To be or not to be faithful: flexible fidelity to foraging trails in the leaf-cutting ant Acromyrmex lobicornis

1. Ants using trails to forage have to select between two alternative routes at bifurcations, using two, potentially conflicting, sources of information to make their decision: individual experience to return to a previous successful foraging site (i.e. fidelity) and ant traffic. In the field, we investigated which of these two types of information individuals of the leaf‐cutting ant Acromyrmex lobicornis Emery use to decide which foraging route to take.