Tradeoffs, competition, and coexistence in eastern deciduous forest ant communities

Ecologists have long sought to explain the coexistence of multiple potentially competing species in local assemblages. This is especially challenging in species-rich assemblages in which interspecific competition is intense, as it often is in ant assemblages. As a result, a suite of mechanisms has been proposed to explain coexistence among potentially competing ant species: the dominance–discovery tradeoff, the dominance–thermal tolerance tradeoff, spatial segregation, temperature-based niche partitioning, and temporal niche partitioning. Through a series of observations and experiments, we examined a deciduous forest ant assemblage in eastern North America for the signature of each of these coexistence mechanisms. We failed to detect evidence for any of the commonly suggested mechanisms of coexistence, with one notable exception: ant species appear to temporally partition foraging times such that behaviourally dominant species foraged more intensely at night, while foraging by subdominant species peaked during the day. Our work, though focused on a single assemblage, indicates that many of the commonly cited mechanisms of coexistence may not be general to all ant assemblages. However, temporal segregation may play a role in promoting coexistence among ant species in at least some ecosystems, as it does in many other organisms.     

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.     

The discovery-dominance trade-off is the exception, rather than the rule

1. Interspecific trade-offs are thought to facilitate coexistence between species at small spatial scales. The discovery-dominance trade-off, analogous to a competition-colonisation trade-off, is considered an important structuring mechanism in ant ecology. A trade-off between species' ability to discover food resources and to dominate them may explain how so many species apparently dependent on similar resources can coexist. 2. The discovery-dominance trade-off is thought to be broken by invasive species in enemy-free space or territorial species whose activity is fuelled by domination of carbohydrate resources. It may also be mediated by factors such as temperature and habitat structure. 3. We investigate the generality and form of the discovery-dominance relationship in an experiment using habitats of contrasting complexity across three continents. In addition, to assess how widespread the discovery-dominance trade-off is, we conducted a systematic review combining all empirical studies (published and from our experiment). 4. From our own fieldwork and meta-analyses of available studies, we find surprisingly little empirical support for the trade-off, with results indicating that mean effect sizes were either not significantly different from 0 or significantly positive. The trade-off was only detected in studies with parasitoids present. Additionally, experimental data from simple and complex habitats within each continent suggest that simple habitats may facilitate both food resource discovery and dominance. 5. We conclude that the discovery-dominance trade-off is the exception, rather than the rule. Instead, these abilities were commonly correlated. Real food resources provide many axes along which partitioning may occur, and discovery-dominance trade-offs are not a prerequisite for coexistence.     

Elevational gradients in phylogenetic structure of ant communities reveal the interplay of biotic and abiotic constraints on diversity

A central focus of ecology and biogeography is to determine the factors that govern spatial variation in biodiversity. Here, we examined patterns of ant diversity along climatic gradients in three temperate montane systems: Great Smoky Mountains National Park (USA), Chiricahua Mountains (USA), and Vorarlberg (Austria). To identify the factors which potentially shape these elevational diversity gradients, we analyzed patterns of community phylogenetic structure (i.e. the evolutionary relationships among species coexisting in local communities). We found that species at low‐elevation sites tended to be evenly dispersed across phylogeny, suggesting that these communities are structured by interspecific competition. In contrast, species occurring at high‐elevation sites tended to be more closely related than expected by chance, implying that these communities are structured primarily by environmental filtering caused by low temperatures. Taken together, the results of our study highlight the potential role of niche constraints, environmental temperature, and competition in shaping broad‐scale diversity gradients. We conclude that phylogenetic structure indeed accounts for some variation in species density, yet it does not entirely explain why temperature and species density are correlated.     

The role of competition by dominants and temperature in the foraging of subordinate species in Mediterranean ant communities

In this paper we test the influence of temperature and interference competition by dominant species on the foraging of subordinate species in Mediterranean ant communities. We have analyzed the changes in resource use by subordinate species in plots with different abundances of dominant ants, and in different periods of the day and the year, i.e., at different temperatures. The expected effects of competition by dominant species on foraging of subordinates were only detected for two species in the number of baits occupied per day, and for one species in the number of foragers at pitfall traps. In all three cases, subordinate species were less represented at baits or in traps in plots with a high density of dominants than in plots with a medium or low density of dominants. The number of workers per bait, and the foraging efficiency of subordinate species did not differ in plots differing in dominant abundance. Daily activity rhythms and curves of temperature versus foraging activity of subordinate species were also similar in plots with different abundance of dominant species, indicating no effect of dominants on the foraging times of subordinates. Instead, temperature had a considerable effect on the foraging of subordinate species. A significant relationship was found between maximum daily temperature and several variables related to foraging (the number of foragers at pitfall traps, the number of baits occupied per day, and the number of workers per bait) of a number subordinate species, both in summer and autumn. These results suggest that the foraging of subordinate ant species in open Mediterranean habitats is influenced more by temperature than by competition of dominants, although an effect of dominants on subordinates has been shown in a few cases. In ant communities living in these severe and variable environments, thermal tolerance reduces the importance of competition, and the mutual exclusion usually found between dominant and subordinate species appears to be the result of physiological specialization to different temperature ranges. Received: 8 May 1998 / Accepted: 30 July 1998     

Correlates of ecological dominance within Pheidole ants (Hymenoptera: Formicidae)

1. In any group of organisms, one can almost invariably find some species that are ecologically dominant (i.e. disproportionately more abundant and widespread), whereas others are comparatively less prevalent. Understanding of the causes of variation in ecological dominance has been elusive, particularly given that dominant and subordinate species often lack obvious features that could predict their abundance in nature. 2. In this study, physiological, behavioural, morphological, and phylogenetic information is integrated in an effort to understand the mechanisms underlying ecological dominance in ants using the hyperdiverse ant genus Pheidole (Formicidae: Myrmicinae) as a model system. Field estimates of the relative abundance of 10 Pheidole species were compared with potential correlates, which included behavioural (walking velocity), physiological (tolerance to high and low temperatures and desiccation), and morphological traits (body size and degree of dimorphism in the worker caste). A molecular phylogeny of the tested species was also generated to account for potential confounding effects of phylogenetic non‐independence. 3. Dominant Pheidole species were characterised by higher environmental tolerance with respect to temperature and humidity, as well as faster walking speeds. On the other hand, no morphological correlates of ecological dominance were detected. Interestingly, subordinate species showed no evidence of trade‐off in performance, being both more fragile to environmental challenges and slower in their walking speeds. 4. These results provide important insights into the mechanisms involved in local species coexistence in Pheidole.     

Critical thermal limits in Mediterranean ant species: trade-off between mortality risk and foraging performance

1. In Mediterranean ant communities, a close relationship has been found between activity rhythm in the period of maximum activity and position in the dominance hierarchy: subordinate species are active during the day, when conditions are more severe, while dominants are active during the afternoon and the night.
2. Results obtained in this study confirmed that the species foraging at higher temperatures were closer to their critical thermal limits than the species foraging at lower temperatures.
3. This enabled two extreme strategies of foraging in relation to temperature to be distinguished: (1) heat-intolerant ant species behaved as risk-averse species, foraging at temperatures very far from their critical thermal limits; and (2) heat-tolerant ant species behaved as risk-prone species, foraging very near their critical thermal limits and running a high heat mortality risk.
4. Heat-tolerant species benefited from this strategy by having better foraging performance at high temperatures.
5. This wide range of thermal niches may be one reason why Mediterranean ant faunas are so diverse in the face of limited diversity in vegetation and habitat structure: the daily range of temperature may be sufficiently great to meet the requirement both of heat-adapted and cold-adapted species as well as a spectrum of intermediate forms.     

Not enough niches: non‐equilibrial processes promoting species coexistence in diverse ant communities

Abstract Explanations for species coexistence in ant communities have traditionally focused on niche partitioning, particularly relating to differences in diet, foraging times and nesting requirements. Although niche separation is undoubtedly important, it seems insufficient to account for the high levels of local species richness that are commonly observed. This paper explores alternative explanations for ant species coexistence, focusing on factors that prevent competitive exclusion in diverse ant communities experiencing high levels of behavioural dominance, such as characteristically occurs in Australia. Very high species densities require two conditions to be met: first, a large number of species must successfully establish; and second, there must be a high rate of species persistence once established. In this context I advance five propositions based around three sets of arguments. First, ant sociality and modularity confers a high level of persistence once colonies are established, so that species coexistence is determined to a significant extent by processes operating at the establishment phase, rather than just by interactions between established colonies. Second, competitive outcomes are highly conditioned by environmental variation, which severely limits competitive exclusion. Finally, dominant species are highly patchy in space, and cannot comprehensively monopolize resources, such that there will usually be room for low densities of subordinate species. These propositions have relevance to neutral theories of community ecology, and to understanding intercontinental differences in local species richness.     

Species interactions and thermal constraints on ant community structure

Patterns of species occurrence and abundance are influenced by abiotic factors and biotic interactions, but these factors are difficult to disentangle without experimental manipulations. In this study, we used observational and experimental approaches to investigate the role of temperature and interspecific competition in controlling the structure of ground‐foraging ant communities in forests of the Siskiyou Mountains of southwestern Oregon. To assess the potential role of competition, we first used null model analyses to ask whether species partition temporal and/or spatial environments. To understand how thermal tolerances influence the structure of communities, we conducted a laboratory experiment to estimate the maximum thermal tolerance of workers and a field experiment in which we added shaded microhabitats and monitored the response of foragers. Finally, to evaluate the roles of temperature and interspecific competition in the field, we simultaneously manipulated shading and the presence of a dominant competitor (Formica moki). The foraging activity of species broadly overlapped during the diurnal range of temperatures. Species co‐occurrence patterns varied across the diurnal temperature range: species were spatially segregated at bait stations at low temperatures, but co‐occurred randomly at high temperatures. The decreased abundance of the co‐occurring thermophilic Temnothorax nevadensis in shaded plots was a direct effect of shading and not an indirect effect of competitive interactions. Thermal tolerance predicted the response of ant species to the shading experiment: species with the lowest tolerances to high temperatures showed the greatest increase in abundance in the shaded plots. Moreover, species with more similar thermal tolerance values segregated more frequently on baits than did species that differed in their thermal tolerances. Collectively, our results suggest that thermal tolerances of ants may mediate competitive effects in habitats that experience strong diurnal temperature fluctuations.     

Ant communities in taiga clearcuts: habitat effects and species interactions

We surveyed the structure of ant communities in young taiga forests by pitfall trapping in southern Finland The sampling sites were clearcut and planted with conifers 14-20 yr before the sampling The results indicated that the structure of the ant communities was largely determined by the top competitors the territorial species of the wood-ant group (Formica aquilonia and F lugubris) in the older, and the aggressive slavemaking ant ( F sanguinea )in the younger clearcuts Species interactions resulted in district spatial distributions of individual species depending on the competitive status of the species concerned Competition and slavemaking were the most important factors on larger spatial scales The spatial scale of competitive structuring was determined by the territory and colony sizes of the top competitors On a finer scale, variability in moisture and tree-canopy shading seemed to have enhanced coexistence of some competing submissive species by alleviating the effects of nest-site comp)edition and slavemaking Competition between the wood ants and the slavemaking ant affected indirectly the distribution and abundance of the species subject to slavemaking. F fusca and F lemam Similarly, the top competitors presumably affected the distributions of other interacting subordinate species indirectly through differential competitive effects on them Overall, species interactions seemed to have induced considerable determinism in ant-community succession in young forests     

Density‐dependent survival varies with species life‐history strategy in a tropical forest

Species coexistence in diverse communities likely results from multiple interacting factors. Mechanisms such as conspecific negative density dependence (CNDD) and varying life‐history strategies related to resource partitioning are known to influence plant fitness, and thereby community composition and diversity. However, we have little understanding of how these mechanisms interact and how they vary across life stages. Here, we document the interaction between CNDD and life‐history strategy, based on growth‐mortality trade‐offs, from seedling to adult tree for 47 species in a tropical forest. Species’ life‐history strategies remained consistent across stages: fast‐growing species had higher mortality than slow‐growing species at all stages. In contrast, mean CNDD was strongest at early life stages (i.e. seedling, sapling). Fast‐growing species tended to suffer greater CNDD than slow‐growing species at several, but not all life stages. Overall, our results demonstrate that coexistence mechanisms interact across multiple life stages to shape diverse tree communities.     

Ants on a mountain: spatial, environmental and habitat associations along an altitudinal transect in a centre of endemism

Mountains are biodiversity hotspots and provide spatially compressed versions of regional and continental variation. They might be the most cost effective way to measure the environmental associations of regional biotic communities and their response to global climate change. We investigated spatial variation in epigeal ant diversity along a north–south elevational transect over the Soutpansberg Mountain in South Africa, to see to what extent these patterns can be related to spatial (regional) and environmental (local) variables and how restricted taxa are to altitudinal zones and vegetation types. A total of 40,294 ants, comprising 78 species were caught. Ant richness peaked at the lowest elevation of the southern aspect but had a hump-shaped pattern along the northern slope. Species richness, abundance and assemblage structure were associated with temperature and the proportion of bare ground. Local environment and spatially structured environmental variables comprised more than two-thirds of the variation explained in species richness, abundance and assemblage structure, while space alone (regional processes) was responsible for <10%. Species on the northern aspect were more specific to particular vegetation types, whereas the southern aspect’s species were more generalist. Lower elevation species’ distributions were more restricted. The significance of temperature as an explanatory variable of ant diversity across the mountain could provide a predictive surrogate for future changes. The effect of CO₂-induced bush encroachment on the southern aspect could have indirect impacts complicating prediction, but ant species on the northern aspect should move uphill at a rate proportional to their thermal tolerance and the regional increases in temperature. Two species are identified that might be at risk of local extinction.     

祁连山国家公园青海片区蚂蚁物种多样性研究

本研究采用样地调查法研究了祁连山国家公园青海片区的蚂蚁物种多样性.在祁连山国家公园青海片区记录蚂蚁2亚科、6属、13种,发现1个中国新记录种:沃尔切胸蚁Temnthorax volgensis(Ruzsky),发现7个青海新记录种.光亮黑蚁 Formica candida Smith、满凹头蚁 Formica manc...     

Competition trade-offs in the organisation of a Mediterranean ant assemblage

Abstract.  1. The organisation of an ant assemblage inhabiting an olive orchard in central Italy was analysed and patterns of dominance among ant species were described in order to assess (i) the relationship between thermal dependency and degree of behavioural dominance, and (ii) the relationship between dominance and discovery ability.
2. Activity patterns of the most abundant species on trees were examined in a sample of 120 trees during spring and summer. The degree of behavioural dominance and the ability of different species to discover new food sources were assessed using tuna baiting on a subset of 80 trees.
3. Different ant species showed contrasting patterns of activity. Some species (such as Lasius lasioides , Camponotus lateralis , and Camponotus piceus ) were most active during the warmer part of the day, while others restricted their activity to the cooler hours ( Camponotus aethiops and Plagiolepis pygmaea ). Some species (such as Crematogaster scutellaris ) were active irrespective of the time of day.
4. No clear relationship was observed between temperature of maximal activity and degree of behavioural dominance. There was, however, a positive relationship between behavioural dominance and thermal range of activity. A positive relationship between dominance and ability to find resources, with the most behaviourally dominant and aggressive species being most efficient in finding food items, was also observed.
5. The results support the idea that the temperature–dominance relationship is much more complex in Mediterranean-type habitats than in other ecosystems. Of particular interest is the positive dominance–discovery relationship. This finding contrasts with previous investigations, which reported a negative relationship between dominance and discovery ability and suggested that this pattern plays a role in promoting the coexistence of species in ant communities.     

Designed and naturalised sward response to management: I. Patterns of herbage production

Pastures in the Appalachian region of the United States comprise a mix of grasses, legumes and forbs that tend to differ in productivity within and among years. A high degree of spatial variability in hill‐land pasture creates microsite conditions that influence botanical composition of pasture. The variation in sward composition presents logistical challenges to livestock producers who rely on a dependable supply of herbage mass and nutritive value to meet production goals. Our objective was to determine if forage communities sown for specific functions, for example, superior dry matter productivity, resource patch exploitation or targeted seasonal production, adapted to changing growing conditions within and among years. Productivity of communities differed among years reflecting the cumulative influences of time, ontogenetic and environmental variations. Maximum productivity was influenced by the specific forage community and less so by simple clipping and fertiliser management. Naturalised swards clipped to emulate hay management tended to have sustained herbage productivity but lower nutritive value when compared to sown communities. Rankings of dry matter productivity of communities were similar for each year where bioactive composition, high productivity and warm season tended to produce the most, and stoloniferous‐rhizomatous and naturalised pasture the least. Regardless of initial sward composition, effective number of species as an index of diversity increased when frequently clipped swards were not fertilised, and when infrequently clipped swards were fertilised. Dry matter production patterns were not influenced by the effective number of species in any forage community suggesting that key species sustained productivity with volunteer species making lesser contribution to total productivity. The species composition of forage plant communities appears to be more important than clipping or fertiliser management practices as a means to sustain forage productivity.     

红火蚁入侵强度对本地蚂蚁群落物种的发现-支配权衡的影响

【目的】发现-支配权衡(discovery-dominance trade off)在物种间的竞争起着重要作用。本研究旨在了解不同强度的红火蚁Solenopsis invicta入侵对本地蚂蚁群落物种的发现-支配权衡的影响,从而探讨红火蚁与本地蚂蚁的竞争机制。【方法】于2017年10-11月,在云南省楚雄州牟定县的一滇橄榄种植基地利用诱饵法和陷阱法结合的方式引诱和收集蚂蚁,观察并记录常见蚂蚁种类的发现能力、招募能力、支配能力、最早发现诱饵的时间段及个体数。【结果】红火蚁的入侵显著影响了本地蚂蚁群落物种的发现-支配权衡。强入侵区(5.6个活动蚁巢/100 m²)蚂蚁群落中物种的相对发现能力和相对支配能力不存在相关性;弱入侵区(0.14个活动蚁巢/100 m²)蚂蚁群落中两者存在较弱的负相关性,但不显著;对照区(无蚁巢)中两者存在显著负相关性(强入侵区:r=-0.01,P=0.97;弱入侵区:r=-0.60,P=0.21;对照区:r=-0.81,P=0.04)。强入侵区的红火蚁种群普遍在5 min之内就能发现诱饵,而弱入侵区的种群需要10～60 ...     

Dietary and Temporal Niche Differentiation in Tropical Ants—Can They Explain Local Ant Coexistence?

How species with similar ecological requirements avoid competitive exclusion remains contentious, especially in the species‐rich tropics. Niche differentiation has been proposed as a major mechanism for species coexistence. However, different niche dimensions must be studied simultaneously to assess their combined effects on diversity and composition of a community. In most terrestrial ecosystems, ants are among the most abundant and ubiquitous animals. Since they display direct, aggressive competition and often competitively displace subordinate species from resources, niche differentiation may be especially relevant among ants. We studied temporal and trophic niche differentiation in a ground ant community in a forest fragment in French Guiana. Different baits were presented during day and night to assess the temporal and dietary niches of the local species. They represented natural food resources such as sugars, carrion, excrements, seeds, and live prey. In addition, pitfalls provided a background measure of ant diversity. The communities attracted to the different baits significantly differed from each other, and even less attractive baits yielded additional species. We detected species specialized on living grasshoppers, sucrose, seeds, or dead insects. Community‐level differences between day and night were larger than those between baits, and many species were temporally specialized. In contrast to commonness, foraging efficiency of species was correlated to food specialization. We conclude that many ant species occupy different temporal or dietary niches. However, for many generalized species, the dietary, and temporal niche differentiation brought forward through our sampling effort, cannot alone explain their coexistence.     

Predicting future coexistence in a North American ant community

Global climate change will remodel ecological communities worldwide. However, as a consequence of biotic interactions, communities may respond to climate change in idiosyncratic ways. This makes predictive models that incorporate biotic interactions necessary. We show how such models can be constructed based on empirical studies in combination with predictions or assumptions regarding the abiotic consequences of climate change. Specifically, we consider a well‐studied ant community in North America. First, we use historical data to parameterize a basic model for species coexistence. Using this model, we determine the importance of various factors, including thermal niches, food discovery rates, and food removal rates, to historical species coexistence. We then extend the model to predict how the community will restructure in response to several climate‐related changes, such as increased temperature, shifts in species phenology, and altered resource availability. Interestingly, our mechanistic model suggests that increased temperature and shifts in species phenology can have contrasting effects. Nevertheless, for almost all scenarios considered, we find that the most subordinate ant species suffers most as a result of climate change. More generally, our analysis shows that community composition can respond to climate warming in nonintuitive ways. For example, in the context of a community, it is not necessarily the most heat‐sensitive species that are most at risk. Our results demonstrate how models that account for niche partitioning and interspecific trade‐offs among species can be used to predict the likely idiosyncratic responses of local communities to climate change.     

Energy gradients and the geographic distribution of local ant diversity

Geographical diversity gradients, even among local communities, can ultimately arise from geographical differences in speciation and extinction rates. We evaluated three models—energy-speciation, energy-abundance, and area—that predict how geographic trends in net diversification rates generate trends in diversity. We sampled 96 litter ant communities from four provinces: Australia, Madagascar, North America, and South America. The energy-speciation hypothesis best predicted ant species richness by accurately predicting the slope of the temperature diversity curve, and accounting for most of the variation in diversity. The communities showed a strong latitudinal gradient in species richness as well as inter-province differences in diversity. The former vanished in the temperature-diversity residuals, suggesting that the latitudinal gradient arises primarily from higher diversification rates in the tropics. However, inter-province differences in diversity persisted in those residuals—South American communities remained more diverse than those in North America and Australia even after the effects of temperature were removed.