Effects of the Argentine ant venom on terrestrial amphibians.

Invasive species have major impacts on biodiversity and are one of the primary causes of amphibian decline and extinction. Unlike other top ant invaders that negatively affect larger fauna via chemical defensive compounds, the Argentine ant (Linepithema humile) does not have a functional sting. Nonetheless, it deploys defensive compounds against competitors and adversaries. We estimated levels of ant aggression toward 3 native terrestrial amphibians by challenging juveniles in field ant trails and in lab ant foraging arenas. We measured the composition and quantities of toxin in L. humile by analyzing pygidial glands and whole-body contents. We examined the mechanisms of toxicity in juvenile amphibians by quantifying the toxin in amphibian tissues, searching for histological damages, and calculating toxic doses for each amphibian species. To determine the potential scope of the threat to amphibians, we used global databases to estimate the number, ranges, and conservation status of terrestrial amphibian species with ranges that overlap those of L. humile. Juvenile amphibians co-occurring spatially and temporally with L. humile die when they encounter L. humile on an ant trail. In the lab, when a juvenile amphibian came in contact with L. humile the ants reacted quickly to spray pygidial-gland venom onto the juveniles. Iridomyrmecin was the toxic compound in the spray. Following absorption, it accumulated in brain, kidney, and liver tissue. Toxic dose for amphibian was species dependent. Worldwide, an estimated 817 terrestrial amphibian species overlap in range with L. humile, and 6.2% of them are classified as threatened. Our findings highlight the high potential of L. humile venom to negatively affect amphibian juveniles and provide a basis for exploring the largely overlooked impacts this ant has in its wide invasive range.

Efectos del Veneno de la Hormiga Argentina sobre los Anfibios Terrestres Resumen Las especies invasoras tienen un impacto importante sobre la biodiversidad y son una de las causas principales del declive y extinción de los anfibios. A diferencia de otras hormigas super-invasoras que afectan negativamente a animales más grandes por medio de compuestos químicos de defensa, la hormiga argentina (Linepithema humile) no tiene unaguijón funcional. Sin embargo, esta hormiga despliega compuestos defensivos contra sus competidores y adversarios. Estimamos los niveles de agresión de las hormigas hacia tres anfibios terrestres nativos exponiendo a los anfibios juveniles en pistas de hormigas en el campo y en las arenas de forrajeo de las hormigas en el laboratorio. Medimos la composición y las cantidades de toxina que presenta L. humile por medio del análisis de las glándulas pigidiales y el contenido en el cuerpo completo. Examinamos los mecanismos de la toxicidad en los anfibios juveniles cuantificando la toxina en el tejido del anfibio, buscando daños histológicos y calculando las dosis tóxicas para cada especie de anfibio. Para determinar el alcance potencial de la amenaza para los anfibios usamos bases de datos mundiales para estimar el número, distribución y estado de conservación de las especies terrestres de anfibios con distribuciones que se solapan con la de L. humile. Los anfibios juveniles que co-ocurren temporal y espacialmente con L. humile mueren al encontrarse con esta especie de hormiga en sus pistas. En el laboratorio, cuando un anfibio juvenil entró en contacto con L. humile, las hormigas reaccionaron rápidamente rociando a estos juveniles con veneno proveniente de las glándulas pigidiales. La iridomyrmecina fue el compuesto tóxico que encontramos en las glándulas pigidiales. Después de ser absorbida por la piel del anfibio, se acumuló en el cerebro, los riñones y el hígado. La dosis tóxica para los anfibios depende de la especie. A nivel mundial, se estima que 817 especies de anfibios terrestres tienen una distribución que se solapa con la de L. humile, y el 6.2% de estas especies se encuentran clasificadas como amenazadas. Nuestros hallazgos resaltan el potencial alto del veneno de L. humile para tener efectos negativos sobre los anfibios juveniles y también proporcionan una base para la exploración de los impactos de esta hormiga en su amplio rango invasivo, los cuales generalmente son ignorados.

Does social thermal regulation constrain individual thermal tolerance in an ant species?

In ants, social thermal regulation is the collective maintenance of a nest temperature that is optimal for individual colony members. In the thermophilic ant Aphaenogaster iberica, two key behaviours regulate nest temperature: seasonal nest relocation and variable nest depth. Outside the nest, foragers must adapt their activity to avoid temperatures that exceed their thermal limits. It has been suggested that social thermal regulation constrains physiological and morphological thermal adaptations at the individual level. We tested this hypothesis by examining the foraging rhythms of six populations of A. iberica, which were found at different elevations (from 100 to 2,000 m) in the Sierra Nevada mountain range of southern Spain. We tested the thermal resistance of individuals from these populations under controlled conditions. Janzen's climatic variability hypothesis (CVH) states that greater climatic variability should select for organisms with broader temperature tolerances. We found that the A. iberica population at 1,300 m experienced the most extreme temperatures and that ants from this population had the highest heat tolerance (LT50 = 57.55°C). These results support CVH's validity at microclimatic scales, such as the one represented by the elevational gradient in this study. Aphaenogaster iberica maintains colony food intake levels across different elevations and mean daily temperatures by shifting its rhythm of activity. This efficient colony-level thermal regulation and the significant differences in individual heat tolerance that we observed among the populations suggest that behaviourally controlled thermal regulation does not constrain individual physiological adaptations for coping with extreme temperatures.

En hormigas, la termorregulación social es el mantenimiento colectivo de la temperatura del nido óptima para los individuos de la colonia. En la hormiga termófila Aphaenogaster iberica, hay dos comportamientos clave que regulan la temperatura del nido: la reubicación estacional y la profundidad variable del nido. Fuera del nido, las obreras recolectoras deben adaptar su actividad para evitar las temperaturas que excedan sus límites térmicos. Se ha sugerido que la termorregulación social limita las adaptaciones térmicas a nivel individual, fisiológicas y morfológicas. Examinamos esta hipótesis, estudiando los ritmos de actividad de recolección de alimento en seis poblaciones de A. iberica, a distinta altitud, desde 100 m a 2,000 m, en las montañas de Sierra Nevada, en el sur de España. Y analizamos la resistencia térmica de los individuos de estas poblaciones, en condiciones controladas. La Hipótesis de la Variabilidad Climática de Janzen (CVH) postula que una mayor variabilidad climática selecciona organismos con tolerancias térmicas más amplias. Encontramos que la población de 1,300 m era la que presentaba la mayor variabilidad climática, y que las hormigas de esta población tiene la mayor resistencia térmica individual (LT50 = 57.55°C), lo que confirma la validez de la CVH a una escala microclimática en el gradiente altitudinal estudiado. Encontramos que A. iberica puede compensar por la disminución de la temperatura media que acompaña al incremento en elevación. Las hormigas pueden cambiar sus ritmos de actividad sin afectar la entrada de alimento en la colonia, que tampoco se ve afectada por la elevación o la temperatura media diaria. A pesar de esta eficiente termorregulación a nivel de colonia, las diferencias estadísticamente significativas entre poblaciones observadas en la tolerancia térmica individual sugieren que la termorregulación controlada comportalmente no limita las adaptaciones fisiológicas individuales para enfrentarse a temperaturas extremas.

Evidence for locally adaptive metabolic rates among ant populations along an elevational gradient.

As global temperatures rise, the mechanistic links between temperature, physiology and behaviour will increasingly define predictions of ecological change. However, for many taxa, we currently lack consensus about how thermal performance traits vary within and across populations, and whether and how locally adaptive trait plasticity can buffer warming effects. The metabolic cold adaptation hypothesis posits that cold environments (e.g. high elevations and latitudes) select for high metabolic rates (MR), even after controlling for body size differences, and that this enables high activity levels when an organism is near its cold lower thermal limits. Steep MR reaction norms are further predicted at cold temperatures to enable rapid behavioural activation with rising temperatures needed to exploit brief thermal windows suitable for performing eco-evolutionary tasks. We tested these predictions by performing common garden experiments comparing thermal reaction norms of MR (from 15 to 32°C) and behaviour (from 10 to 40°C) across populations of the ant Aphaenogaster iberica sampled from a 2 km elevation gradient in the Sierra Nevada Mountains of southern Spain. As predicted, high-elevation ants had higher MR and steeper MR-temperature reaction norms. However, higher rates of energy use did not yield the predicted benefits of steeper activity-level reaction norms. The evidence for locally adaptive metabolic physiology only became apparent at intermediate temperatures, highlighting the importance of testing thermal performance hypotheses across thermal gradients, rather than focusing only on performance at thermal limits (i.e. critical thermal values). The partial support for the metabolic cold adaptation hypothesis highlights that while organisms likely show a wealth of unexplored metabolic temperature plasticity, the physiological mechanisms and eco-evolutionary trade-offs underlying such local adaptation remain obscure.

Origin and distribution of desert ants across the Gibraltar Straits.

The creation of geographic barriers has long been suspected to contribute to the formation of new species. We investigated the phylogeography of desert ants in the western Mediterranean basin in order to elucidate their mode of diversification. These insects which have a low dispersal capacity are recently becoming important model systems in evolutionary studies. We conducted an extensive sampling of species belonging to the Cataglyphis albicans group in the Iberian Peninsula (IP) and the northern Morocco (North Africa; NA). We then combined genetic, chemical and morphological analyses. The results suggest the existence of at least three and five clades in the IP and NA, respectively, whose delineation partially encompass current taxonomic classification. The three Iberian clades are monophyletic, but their origin in NA is uncertain (79% and 22% for Bayesian and Maximum Likelihood support, respectively). The estimation of divergence time suggests that a speciation process was initiated after the last reopening of the Gibraltar Straits ≈5.33 Ma. In the IP, the clades are parapatric and their formation may have been triggered by the fragmentation of a large population during the Pleistocene due to extended periods of glaciation. This scenario is supported by demographic analyses pointing at a recent expansion of Iberian populations that contrasts with the progressive contraction of the NA clades. Niche modeling reveals that this area, governed by favorable climatic conditions for desert ants, has recently increased in the IP and decreased in NA. Altogether, our data points at geoclimatic events as major determinants of species formation in desert ants, reinforcing the role of allopatric speciation.

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.

Queen Control or Queen Signal in Ants: What Remains of the Controversy 25 Years After Keller and Nonacs' Seminal Paper?

Ant queen pheromones (QPs) have long been known to affect colony functioning. In many species, QPs affect important reproductive functions such as diploid larvae sexualization and egg-laying by workers, unmated queens (gynes), or other queens. Until the 1990s, these effects were generally viewed to be the result of queen manipulation through the use of coercive or dishonest signals. However, in their seminal 1993 paper, Keller and Nonacs challenged this idea, suggesting that QPs had evolved as honest signals that informed workers and other colony members of the queen's presence and reproductive state. This paper has greatly influenced the study of ant QPs and inspired numerous attempts to identify fertility-related compounds and test their physiological and behavioral effects. In the present article, we review the literature on ant QPs in various contexts and pay special attention to the role of cuticular hydrocarbons (CHCs). Although the controversy generated by Keller and Nonacs' (Anim Behav 45:787-794, 1993) paper is currently less intensively debated, there is still no clear evidence which allows the rejection of the queen control hypothesis in favor of the queen signal hypothesis. We argue that important questions remain regarding the mode of action of QPs, and their targets which may help understanding their evolution.

Social Life in Arid Environments: The Case Study of Cataglyphis Ants.

Unlike most desert-dwelling animals, Cataglyphis ants do not attempt to escape the heat; rather, they apply their impressive heat tolerance to avoid competitors and predators. This thermally defined niche has promoted a range of adaptations both at the individual and colony levels. We have also recently discovered that within the genus Cataglyphis there are incredibly diverse social systems, modes of reproduction, and dispersal, prompting the tantalizing question of whether social diversity may also be a consequence of the harsh environment within which we find these charismatic ants. Here we review recent advances regarding the physiological, behavioral, life-history, colony, and ecological characteristics of Cataglyphis and consider perspectives on future research that will build our understanding of organic adaptive responses to desertification.

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.

Is phenotypic plasticity a key mechanism for responding to thermal stress in ants?

Unlike natural selection, phenotypic plasticity allows organisms to respond quickly to changing environmental conditions. However, plasticity may not always be adaptive. In insects, body size and other morphological measurements have been shown to decrease as temperature increases. This relationship may lead to a physiological conflict in ants, where larger body size and longer legs often confer better thermal resistance. Here, we tested the effect of developmental temperature (20, 24, 28 or 32 °C) on adult thermal resistance in the thermophilic ant species Aphaenogaster senilis. We found that no larval development occurred at 20 °C. However, at higher temperatures, developmental speed increased as expected and smaller adults were produced. In thermal resistance tests, we found that ants reared at 28 and 32 °C had half-lethal temperatures that were 2 °C higher than those of ants reared at 24 °C. Thus, although ants reared at higher temperatures were smaller in size, they were nonetheless more thermoresistant. These results show that A. senilis can exploit phenotypic plasticity to quickly adjust its thermal resistance to local conditions and that this process is independent of morphological adaptations. This mechanism may be particularly relevant given current rapid climate warming.

Native predators living in invaded areas: responses of terrestrial amphibian species to an Argentine ant invasion.

Predator-prey interactions play a key role in the success and impacts of invasive species. However, the effects of invasive preys on native predators have been poorly studied. Here, we first reviewed hypotheses describing potential relationships between native predators and invasive preys. Second, we examined how an invasive prey, the Argentine ant (Linepithema humile), affected a native terrestrial amphibian community. In the field, we looked at the structure of the amphibian community in invaded versus uninvaded areas and characterized amphibian trophic ecology. The amphibian community sampled seemed to show a species-dependent response in abundance to invasion: adults of the natterjack toad (Bufo calamita), the species demonstrating the highest degree of ant specialization, were less abundant in invaded areas. Although available ant biomass was significantly greater in invaded than in uninvaded areas (only Argentine ants occurred in the former), amphibians consumed relatively fewer ants in invaded areas. In the lab, we quantified amphibian consumption of Argentine ants versus native ants and assessed whether consumption patterns could have been influenced by prior exposure to the invader. The lab experiments corroborated the field results: amphibians preferred native ants over Argentine ants, and prior exposure did not influence consumption. Differences in preference explained why amphibians consumed fewer Argentine ants in spite of their greater relative availability; they might also explain why the most ant-specialized amphibians seemed to avoid invaded areas. Our results suggest the importance to account for predator feeding capacities and dietary ranges to understand the effects of invasive species at higher trophic levels.

Social coercion of larval development in an ant species.

Ants provide one of the best examples of the division of labor in animal societies. While the queens reproduce, workers generally refrain from laying eggs and dedicate themselves exclusively to domestic tasks. In many species, the small diploid larvae are bipotent and can develop either into workers or queens depending mostly on environmental cues. This generates a conflicting situation between the adults that tend to rear a majority of larvae into workers and the larvae whose individual interest may be to develop into reproductive queens. We tested the social regulation of larval caste fate in the fission-performing ant Aphaenogaster senilis. We first observed interactions between resident workers and queen- and worker-destined larvae in presence/absence of the queen. The results show that workers tend to specifically eliminate queen-destined larvae when the queen is present but not when she is absent or imprisoned in a small cage allowing for volatile pheromone exchanges. In addition, we found that the presence of already developed queen-destined larvae does not inhibit the development of younger still bipotent larvae into queens. Finally, we analyzed the cuticular hydrocarbon profiles of queen- and worker-destined larvae and found no significant quantitative or qualitative difference. Interestingly, the total amount of hydrocarbons on both larval castes is extremely low, which lends credence on the chemical insignificance hypothesis of larval ants. Overall, our results suggest that workers control larval development and police larvae that would develop into queens instead of workers. Such policing behavior is similar in many aspects to what is known of worker policing among adults.

A multidimensional functional trait analysis of resource exploitation in European ants.

The major factors explaining ecological variation in plants have been widely discussed over the last decade thanks to numerous studies that have examined the covariation that exists between pairs of traits. However, multivariate relationships among traits remain poorly characterized in animals. In this study, we aimed to identify the main multivariate trait dimensions that explain variance in important functional traits related to resource exploitation in ants. To this end, we created a large ant trait database. This database includes information on 11 traits that are important in ant resource exploitation; data were obtained for 150 European species found in different biomes. First, we examined the pairwise correlations between the traits included in the database. Second, we used multivariate analyses to identify potential trait dimensions. Our study shows that, to a great extent, resource exploitation strategies align along two main trait dimensions. The first dimension emerged in both the overall and group-specific analyses, where it accounted for the same pairwise trait correlations. The second dimension was more variable, as species were grouped by levels of taxonomy, habitat, and climate. These two dimensions included most of the significant pairwise trait correlations, thus highlighting that complementarity, but also redundancy, exists among different pairs of traits. The first dimension was associated with behavioral dominance: dominance was associated with large colony size, presence of multiple nests per colony, worker polymorphism, and a collective foraging 'strategy. The second dimension was associated with resource partitioning along dietary and microhabitat lines: it ranged from species that consume liquid foods, engage in group foraging, and mainly nest in the vegetation to species that consume insects and seeds, engage in individual foraging, and demonstrate strictly diurnal activity. Our findings establish a proficient ecological trait-based animal research that minimizes the number of traits to be measured while maximizing the number of relevant trait dimensions. Overall, resource exploitation in animals might be framed by behavioral dominance, foraging strategy, diet, and nesting habitat; the position of animal species within this trait space could provide relevant information about their distribution and abundance, for today as well as under future global change scenarios.

Recent speciation and secondary contact in endemic ants.

Gene flow is the main force opposing divergent selection, and its effects are greater in populations in close proximity. Thus, complete reproductive isolation between parapatric populations is not expected, particularly in the absence of ecological adaptation and sharp environmental differences. Here, we explore the biogeographical patterns of an endemic ant species, Cataglyphis floricola, for which two colour morphs (black and bicolour) coexist in parapatry throughout continuous sandy habitat in southern Spain. Discriminant analyses of six biometric measurements of male genitalia and 27 cuticular hydrocarbons reveal high differentiation between morphs. Furthermore, the low number of shared alleles derived from nuclear markers (microsatellites) between the morphs at their contact zone suggests the absence of recent gene flow. Mitochondrial DNA (COI) phylogenetic analysis and median-joining networks show that the black morph is basal to the bicolour morph, with unique haplotypes recovered for each morph. Mismatch distribution analysis and Bayesian skyline plots suggest that they are undergoing different demographic changes, with the bicolour and black morphs at demographic equilibrium and expansion, respectively. Thus, our results show complete reproductive isolation between the two colour morphs as evidenced from genetic, chemical and morphological data. We suggest that these divergence events could be explained by historical vicariance during the Pleistocene, in which reproductive traits experienced strong divergent selection between the morphs initiating or culminating speciation.

Ant functional responses along environmental gradients.

Understanding species distributions and diversity gradients is a central challenge in ecology and requires prior knowledge of the functional traits mediating species' survival under particular environmental conditions. While the functional ecology of plants has been reasonably well explored, much less is known about that of animals. Ants are among the most diverse, abundant and ecologically significant organisms on earth, and they perform a great variety of ecological functions. In this study, we analyse how the functional species traits present in ant communities vary along broad gradients in climate, productivity and vegetation type in the south-western Mediterranean. To this end, we compiled one of the largest animal databases to date: it contains information on 211 local ant communities (including eight climate variables, productivity, and vegetation type) and 124 ant species, for which 10 functional traits are described. We used traits that characterize different dimensions of the ant functional niche with respect to morphology, life history and behaviour at both individual and colony level. We calculated two complementary functional trait community indices ('trait average' and 'trait dissimilarity') for each trait, and we analysed how they varied along the three different gradients using generalized least squares models that accounted for spatial autocorrelation. Our results show that productivity, vegetation type and, to a lesser extent, each climate variable per se might play an important role in shaping the occurrence of functional species traits in ant communities. Among the climate variables, temperature and precipitation seasonality had a much higher influence on functional responses than their mean values, whose effects were almost lacking. Our results suggest that strong relationships might exist between the abiotic environment and the distribution of functional traits among south-western Mediterranean ant communities. This finding indicates that functional traits may modulate the responses of ant species to the environment. Since these traits act as the link between species distributions and the environment, they could potentially be used to predict community changes under future global change scenarios.

Ecological diversification preceded geographical expansion during the evolutionary radiation of Cataglyphis desert ants.

Biological diversity often arises as organisms adapt to new ecological conditions (i.e., ecological opportunities) or colonize suitable areas (i.e., spatial opportunities). Cases of geographical expansion followed by local ecological divergence are well described; they result in clades comprising ecologically heterogeneous subclades. Here, we show that the desert ant genus Cataglyphis likely originated in open grassland habitats in the Middle East ∼18 million years ago and became a taxon of diverse species specializing in prey of different masses. The genus then colonized the Mediterranean Basin around 9 million years ago. The result was the rapid accumulation of species, and the appearance of local assemblages containing species from different lineages that still displayed ancestral foraging specialties. These findings highlight that, in Cataglyphis, ecological diversification preceded geographical expansion, resulting in a clade composed of ecologically homogeneous subclades.

Surface lipids of queen-laid eggs do not regulate queen production in a fission-performing ant.

In animal societies, most collective and individual decision making depends on the presence of reproductive individuals. The efficient transmission of information among reproductive and non-reproductive individuals is therefore a determinant of colony organization. In social insects, the presence of a queen modulates multiple colonial activities. In many species, it negatively affects worker reproduction and the development of diploid larvae into future queens. The queen mostly signals her presence through pheromone emission, but the means by which these chemicals are distributed in the colony are still unclear. In several ant species, queen-laid eggs are the vehicle of the queen signal. The aim of this study was to investigate whether queen-laid eggs of the ant Aphaenogaster senilis possess queen-specific cuticular hydrocarbons and/or Dufour or poison gland compounds, and whether the presence of eggs inhibited larval development into queens. Our results show that the queen- and worker-laid eggs shared cuticular and Dufour hydrocarbons with the adults; however, their poison gland compounds were not similar. Queen-laid eggs had more dimethylalkanes and possessed a queen-specific mixture of cuticular hydrocarbons composed of 3,11 + 3,9 + 3,7-dimethylnonacosane, in higher proportions than did worker-laid eggs. Even though the queen-laid eggs were biochemically similar to the queen, their addition to experimentally queenless groups did not prevent the development of new queens. More studies are needed on the means by which queen ant pheromones are transmitted in the colony, and how these mechanisms correlates with life history traits.

Long-term recovery of Mediterranean ant and bee communities after fire in southern Spain.

Wildfires play a determinant role in the composition and structure of animal communities, especially for groups closely associated with the vegetation and soil, such as bees or ants. The effects of fire on animal communities depend on the functional traits of each group. Here, we assessed the impacts of fire and time since fire on the taxonomic and functional responses of ant and bee communities. We sampled 35 pine forests in Andalusia (southern Spain) that had experienced fire in the past (0 to 41 years ago). Specifically, we explored whether a) fire increased taxonomic and functional diversity and changed community composition in communities in the short term and b) fire influence (increase or decrease) on ant communities would be dependent on time since fire. We found that ant and bee taxonomic richness increased regardless of time since fire. Different approaches gave the same result, such as taxonomic diversity indexes (ant abundance, ant richness and ant Shannon diversity index), the changes in species richness in ant and bee communities, as well as the higher number of ant and bee species prone to the burned habitat than to the unburned habitat, using the Ihabitat Index. Besides environmental variables (such as the effects of different Pinus species or elevation), time since fire changed the taxonomic composition of ant communities and the functional composition of bee communities. Moreover, six of the 13 ant functional traits explored differed between burned and unburned areas, with the degree of difference declining as time since fire increased. For example, burned areas contained ant communities with more ground-nesting species and strictly diurnal species, functional traits that are characteristic of open areas. In contrast, other traits persisted in burned areas over the long term, notably a higher degree of worker polymorphism and species monogyny. Our study shows how much short- and long-term effects of fire on ant and bee communities differ; while richness increases in the long-term, some functional traits are also filtered in the short-term. We suggest that fire in Mediterranean coniferous ecosystems could have a positive effect on these groups and should not be overlooked.

Distinctive life traits and distribution along environmental gradients of dominant and subordinate Mediterranean ant species.

For most animal and plant species, life traits strongly affect their species-specific role, function or position within ecological communities. Previous studies on ant communities have mostly focused on the role of dominant species and the outcome of interspecific interactions. However, life traits of ant species have seldom been considered within a community framework. This study (1) analyses life traits related to ecological and behavioural characteristics of dominant and subordinate ant species from 13 sites distributed throughout the Iberian Peninsula, (2) determines how similar the ant species are within each of the two levels of the dominance hierarchy, and (3) establishes the distribution patterns of these different groups of species along environmental gradients. Our results showed that the differences between dominants and subordinates fall into two main categories: resource exploitation and thermal tolerance. Dominant species have more populated colonies and defend food resources more fiercely than subordinates, but they display low tolerance to high temperatures. We have identified different assemblages of species included within each of these two levels in the dominance hierarchy. The distribution of these assemblages varied along the environmental gradient, shifting from dominant Dolichoderinae and cryptic species in moist areas, to dominant Myrmicinae and hot climate specialists mainly in open and hot sites. We have been able to identify a set of life traits of the most common Iberian ant species that has enabled us to characterise groups of dominant and subordinate species. Although certain common features within the groups of both dominants and subordinates always emerge, other different features allow for differentiating subgroups within each of these groups. These different traits allow the different subgroups coping with particular conditions across environmental gradients.

Substrate temperature constrains recruitment and trail following behavior in ants.

In many ant species, foragers use pheromones to communicate the location of resources to nestmates. Mass-recruiting species deposit long-lasting anonymous chemical trails, while group-recruiting species use temporary chemical trails. We studied how high temperature influenced the foraging behavior of a mass-recruiting species (Tapinoma nigerrimum) and a group-recruiting species (Aphaenogaster senilis) through pheromone decay. First, under controlled laboratory conditions, we examined the effect of temperature on the trail pheromone of both species. A substrate, simulating soil, marked with gaster extract was heated for 10 min. at 25°, 35°, 45°, or 55 °C and offered to workers in a choice test. Heating gaster extract reduced the trail following behavior of the mass-recruiters significantly more than that of the group-recruiters. Second, analyses of the chemicals present on the substrate indicated that most T. nigerrimum gaster secretions vanished at 25 °C, and only iridodials persisted up to 55 °C. By contrast, A. senilis secretions were less volatile and resisted better to elevated temperatures to some extent. However, at 55 °C, the only chemicals that persisted were nonadecene and nonadecane. Overall, our results suggest that the foraging behavior of the group-recruiting species A. senilis is less affected by pheromone evaporation than that of the mass-recruiting species T. nigerrimum. This group-recruiting species might, thus, be particularly adapted to environments with fluctuating temperatures.

Temperature or competition: Which has more influence on Mediterranean ant communities?

Temperature and competition are two of the main factors determining ant community assemblages. Temperature may allow species to forage more or less efficiently throughout the day (in accordance with the maximum activity temperature of each species). Competition can be observed and quantified from species replacements occurring during resource exploitation. We studied the interspecific competitive interactions of ant communities from the Doñana Biological Reserve (southern Spain). Ants were sampled from pitfall traps and baits in three habitats with contrasted vegetation physiognomy (savin forest, pine forest, and dry scrubland). We measured the temperature during the competitive interactions between species and created a thermal competition index (TCI) to assess the relative contribution of temperature and numerical dominance to the competitive outcomes. Temperature had unequal effects on ant activity in each type of habitat, and modulated competitive interactions. The TCI showed that a species' success during pair interactions (replacements at baits) was driven by the proportion of workers between the two competing species and by the species-specific effect of temperature (how advantageous the temperature change is for each species during bait replacement). During competitive interactions, the effect of temperature (higher values of TCI) and numeric supremacy (higher worker proportion) gave higher success probabilities. Interspecific competitive relationships in these Mediterranean ant communities are habitat dependent and greatly influenced by temperature.