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     

Responses of biotic interactions of dominant and subordinate species to decadal warming and simulated rotational grazing in Tibetan alpine meadow

Warming increases competition among plant species in alpine communities by ameliorating harsh environmental conditions, such as low temperatures. Grazing, as the main human activity, may mitigate the effect of warming, as previously reported. However, it is critical to refine the effects of warming on biotic interactions among species, for example, by taking the competitive ability of species into consideration. Based on a 10-year warming and grazing experiment in a Tibetan alpine meadow, we evaluated interspecific biotic interactions of dominant and subordinate species, using the approach of interspecific spatial associations. Warming significantly increased competition between subordinate and dominant species as well as among subordinate species, but not among dominant species. Moreover, facilitation of dominant-subordinate species also increased under warming. Simulated rotational grazing had similar effects to warming, with increasing interspecific competition. Our results show that, when studying the effects of warming on biotic interactions among species, it is necessary to characterize different species pairs relative to their competitive ability, and that simulated rotational grazing does not mitigate the effects of warming in the long term. Our results also provide evidence that the spatial pattern of species is a critical mechanism in species coexistence.     

Social rank‐dependent expression of arginine vasotocin in distinct preoptic regions in male Oryzias latipes

This study examined arginine vasotocin (AVT) expression in the brains of dominant and subordinate male medaka Oryzias latipes after short‐ and long‐term competition. High AVT expression in distinct preoptic regions was found in dominants and subordinates within minutes of encountering each other. During long‐term competition, AVT expression remained high in dominants but not in subordinates.     

Dominance and species co-occurrence in highly diverse ant communities: a test of the interstitial hypothesis and discovery of a three-tiered competition cascade

The role of competitive exclusion is problematic in highly diverse ant communities where exceptional species richness occurs in the face of exceptionally high levels of behavioural dominance. A possible non-niche–based explanation is that the abundance of behaviourally dominant ants is highly patchy at fine spatial scales, and subordinate species act as insinuators by preferentially occupying these gaps—we refer to this as the interstitial hypothesis. To test this hypothesis, we examined fine-scale patterns of ant abundance and richness according to a three-tiered competition hierarchy (dominants, subdominants and subordinates) in an Australian tropical savanna using pitfall traps spaced at 2 m intervals. Despite the presence of gaps in the fine-scale abundance of individual species, the combined abundance of dominant ants (species of Iridomyrmex, Papyirus and Oecophylla) was relatively uniform. There was therefore little or no opportunity for subordinate species to preferentially occupy gaps in the foraging ranges of dominant species, and we found no relationship between the abundance of dominant ants and nondominant species richness at fine spatial scales. However, we found a negative relationship between subdominant and subordinate ants, a negative relationship between dominant and subdominant ants, and a positive relationship between dominant and subordinate ants. These results suggest that dominant species actually promote species richness by neutralizing the effects of subdominant species on subordinate species. Such indirect interactions have very close parallels with three-tiered trophic cascades in food webs, and we propose a “competition cascade” where the interactions are through a competition rather than trophic hierarchy.     

Effect of the Presence of Subordinates on Dominant Female Behaviour and Fitness in Hierarchies of the Dwarf Angelfish Centropyge bicolor

In many hierarchical animal societies, dominant individuals control group membership owing to their power to evict subordinates. In such groups, the presence of subordinates, and therefore group stability, is continually dependent on subordinates being tolerated by dominants. The dominant decision to tolerate or evict is, in turn, dependent on the costs and benefits to dominants of subordinate presence. We investigated the effect of subordinate presence on dominants in the female dominance hierarchy of the dwarf angelfish Centropyge bicolor, using both observations of natural groups and experimental removals of subordinates. We found that the presence of subordinates had no effect on dominant access to resources, as measured by dominant foraging rates and home range areas, nor on dominant fitness, as measured by growth rates and spawning frequencies. Our results suggest that the presence of subordinates has a neutral effect on the current fitness of dominants, so that dominants have no great incentive to evict subordinates. We discuss the possibility that tolerance of subordinates might be further explained by considering future fitness, as dominant females in these haremic protogynous angelfish stand to inherit the male position, whereupon subordinate females change from potential competition to useful mates.     

The devil is in the detail: Nonadditive and context‐dependent plant population responses to increasing temperature and precipitation

In climate change ecology, simplistic research approaches may yield unrealistically simplistic answers to often more complicated problems. In particular, the complexity of vegetation responses to global climate change begs a better understanding of the impacts of concomitant changes in several climatic drivers, how these impacts vary across different climatic contexts, and of the demographic processes underlying population changes. Using a replicated, factorial, whole‐community transplant experiment, we investigated regional variation in demographic responses of plant populations to increased temperature and/or precipitation. Across four perennial forb species and 12 sites, we found strong responses to both temperature and precipitation change. Changes in population growth rates were mainly due to changes in survival and clonality. In three of the four study species, the combined increase in temperature and precipitation reflected nonadditive, antagonistic interactions of the single climatic changes for population growth rate and survival, while the interactions were additive and synergistic for clonality. This disparity affects the persistence of genotypes, but also suggests that the mechanisms behind the responses of the vital rates differ. In addition, survival effects varied systematically with climatic context, with wetter and warmer + wetter transplants showing less positive or more negative responses at warmer sites. The detailed demographic approach yields important mechanistic insights into how concomitant changes in temperature and precipitation affect plants, which makes our results generalizable beyond the four study species. Our comprehensive study design illustrates the power of replicated field experiments in disentangling the complex relationships and patterns that govern climate change impacts across real‐world species and landscapes.     

Both facilitation and limiting similarity shape the species coexistence in dry alkali grasslands

Facilitation is an important driver of community assembly, and often overwhelms the effect of competition in stressed habitats. Thus, net effect of biotic interactions is often positive in stressed grasslands, where dominant species and litter can protect the subordinate species. Besides facilitation, niche partitioning can also support species coexistence leading to limiting similarity between subordinate species. Our aim was to provide a detailed analysis of fine-scale biotic interactions in stressed alkali grasslands. We supposed, that there are positive relationships between the main biomass fractions and species richness. We expected the expansion of trait ranges and the increase of trait dissimilarity with increasing biomass scores (total litter, green biomass of dominant species) and species richness. We studied the relationships between main biomass fractions, species richness, functional diversity and functional trait indices (ranges, weighted means and Rao indices). We used fine-scale biomass sampling in nine stands of dry alkali grasslands dominated by Festuca pseudovina. The detected relationships were always positive between the main biomass fractions (green biomass of dominant species, total litter and green biomass of subordinate species) and species richness. We found that the green biomass of dominant species and total litter increased ranges and dissimilarity of functional traits. Our results suggest that in dry alkali grasslands facilitation is crucial in shaping vegetation composition. The green biomass of dominant species and total litter increased the biomass production of subordinate species leading to overyielding. We found that mechanisms of facilitation and limiting similarity were jointly shaping the species coexistence in stressed grasslands, such as alkali grasslands.     

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.     

Range position and climate sensitivity: The structure of among‐population demographic responses to climatic variation

Species’ distributions will respond to climate change based on the relationship between local demographic processes and climate and how this relationship varies based on range position. A rarely tested demographic prediction is that populations at the extremes of a species’ climate envelope (e.g., populations in areas with the highest mean annual temperature) will be most sensitive to local shifts in climate (i.e., warming). We tested this prediction using a dynamic species distribution model linking demographic rates to variation in temperature and precipitation for wood frogs (Lithobates sylvaticus) in North America. Using long‐term monitoring data from 746 populations in 27 study areas, we determined how climatic variation affected population growth rates and how these relationships varied with respect to long‐term climate. Some models supported the predicted pattern, with negative effects of extreme summer temperatures in hotter areas and positive effects on recruitment for summer water availability in drier areas. We also found evidence of interacting temperature and precipitation influencing population size, such as extreme heat having less of a negative effect in wetter areas. Other results were contrary to predictions, such as positive effects of summer water availability in wetter parts of the range and positive responses to winter warming especially in milder areas. In general, we found wood frogs were more sensitive to changes in temperature or temperature interacting with precipitation than to changes in precipitation alone. Our results suggest that sensitivity to changes in climate cannot be predicted simply by knowing locations within the species’ climate envelope. Many climate processes did not affect population growth rates in the predicted direction based on range position. Processes such as species‐interactions, local adaptation, and interactions with the physical landscape likely affect the responses we observed. Our work highlights the need to measure demographic responses to changing climate.     

Interactions among salt marsh plants vary geographically but not latitudinally along the California coast

The strength of species interactions often varies geographically and locally with environmental conditions. Competitive interactions are predicted to be stronger in benign environments while facilitation is expected to be stronger in harsh ones. We tested these ideas with an aboveground neighbor removal experiment at six salt marshes along the California coast. We determined the effect of removals of either the dominant species, Salicornia pacifica, or the subordinate species on plant cover, aboveground biomass and community composition, as well as soil salinity and moisture. We found that S. pacifica consistently competed with the subordinate species and that the strength of competition varied among sites. In contrast with other studies showing that dominant species facilitate subordinates by moderating physical stress, here the subordinate species facilitated S. pacifica shortly after removal treatments were imposed, but the effect disappeared over time. Contrary to expectations based on patterns observed in east coast salt marshes, we did not see patterns in species interactions in relation to latitude, climate, or soil edaphic characteristics. Our results suggest that variation in interactions among salt marsh plants may be influenced by local‐scale site differences such as nutrients more than broad latitudinal gradients.     

Recruitment facilitation can promote coexistence and buffer population growth in metacommunities

Although positive species interactions are ubiquitous in nature, theory has generally focused on the role of negative interactions to explain patterns of species diversity. Here, we incorporate recruitment facilitation, a positive interaction prevalent in marine and terrestrial systems, into a metacommunity framework to assess how the interplay between colonisation, competition and facilitation mediates coexistence. We show that when subordinate species facilitate the recruitment of dominant species, multi-species metacommunities can persist stably even if the colonisation rate of the dominant species is greater than that of the subordinate species. In addition, recruitment facilitation can buffer population growth from changes in colonisation rates, and thus explain the paradoxical mismatch between patterns of abundance and recruitment in marine systems. Overall, our results demonstrate that recruitment facilitation can have profound effects on the assembly, dissolution and regulation of metacommunities by mediating the relative influence of local and regional processes on population abundance and species diversity.     

Arbuscular mycorrhizal fungi reduce the differences in competitiveness between dominant and subordinate plant species

In grassland communities, plants can be classified as dominants or subordinates according to their relative abundances, but the factors controlling such distributions remain unclear. Here, we test whether the presence of the arbuscular mycorrhizal (AM) fungus Glomus intraradices affects the competitiveness of two dominant (Taraxacum officinale and Agrostis capillaris) and two subordinate species (Prunella vulgaris and Achillea millefolium). Plants were grown in pots in the presence or absence of the fungus, in monoculture and in mixtures of both species groups with two and four species. In the absence of G. intraradices, dominants were clearly more competitive than subordinates. In inoculated pots, the fungus acted towards the parasitic end of the mutualism–parasitism continuum and had an overall negative effect on the growth of the plant species. However, the negative effects of the AM fungus were more pronounced on dominant species reducing the differences in competitiveness between dominant and subordinate species. The effects of G. intraradices varied with species composition highlighting the importance of plant community to mediate the effects of AM fungi. Dominant species were negatively affected from the AM fungus in mixtures, while subordinates grew identically with and without the fungus. Therefore, our findings predict that the plant dominance hierarchy may flatten out when dominant species are more reduced than subordinate species in an unfavourable AM fungal relationship (parasitism).     

Higher reproductive skew among birds than mammals in cooperatively breeding species

While competition for limited breeding positions is a common feature of group life, species vary widely in the extent to which reproduction is shared among females (‘reproductive skew’). In recent years, there has been considerable debate over the mechanisms that generate variation in reproductive skew, with most evidence suggesting that subordinates breed when dominants are unable to prevent them from doing so. Here, we suggest that viviparity reduces the ability of dominant females to control subordinate reproduction and that, as a result, dominant female birds are more able than their mammal counterparts to prevent subordinates from breeding. Empirical data support this assertion. This perspective may increase our understanding of how cooperative groups form and are stabilized in nature.     

Architectural and physiological mechanisms of reduced size inequality in CO2-enriched stands of common ragweed (Ambrosia artemisiifolia)

Testing whether and how subordinate individuals differ from dominants in the utilization of enriched CO₂ atmospheres is important for understanding future stand and community structure. We hypothesized that subordinate and dominant Ambrosia artemisiifolia L. (Asteraceae) (common ragweed) plants growing in dense stands would not equally acquire or utilize carbon gains from CO₂-enrichment, and that the resulting disproportionate growth gains to subordinates would reduce size inequalities in competing stands. We grew experimental stands of A. artemisiifolia in either ambient (360 μL L⁻¹) or twice ambient (720 μL L⁻¹) levels of atmospheric CO₂. We compared the relative growth, photosynthetic capacity, and architecture of subordinate and dominant plants in each treatment, and assessed size inequalities using the stand-level coefficient of variation (CV). In elevated CO₂, plants grew larger, but subordinate plants shifted more mass to upper stem allocation than dominants. Dominant plants demonstrated reduced leaf-level photosynthetic gains in elevated CO₂ compared with subordinate plants. Reduced CVs in plant size reflected smaller proportional growth gains by dominants over subordinates in elevated vs. ambient stands. We conclude that differences in the architectural and physiological responses of subordinate and dominant ragweed plants reduce competition and allow subordinate plants to catch up to dominants in elevated CO₂ conditions.     

Differential influence of native and introduced arbuscular mycorrhizal fungi on growth of dominant and subordinate plants

The impact of arbuscular mycorrhizal fungi (AMF) on plant ecosystems has been intensively reported. In this research, we explored the difference between native and introduced AMF in promoting the growth of dominant and subordinate plant species. In glasshouse experiments, dominants and subordinates from subtropical grasslands were colonized by native AMF or introduced AMF, Glomus versiforme. The biomass revealed that mycorrhizal dependencies (MD) on the native AMF of the dominants were much higher than those of the subordinates, while MD on the introduced AMF changed following the replacement of native AMF with introduced AMF. A close relationship between biomass promotion and increase in phosphorus uptake was observed, indicating the important role of AMF-enhanced nutrient acquisition by roots. Our results show that plant community structures are partly determined by MD on native AMF, and could be modified by introducing exogenous AMF species.     

A test of biotic interactions among two alpine plant species in Australia

This study examines the extent to which interactions among two common alpine/subalpine plant species and their neighbours at the Bogong High Plains in southern Australia are characterized by competition or facilitation. The two target species were Celmisia pugioniformis (Asteraceae) and Carex breviculmis (Cyperaceae). Biotic interactions were examined using vegetation removal manipulations over three growing seasons at five sites across the altitudinal range of tall alpine herbfield communities. Observations recorded growth and mortality. Results for C. pugioniformis clearly indicated facilitation as a dominant process across all sites and seasons. Plants that had their neighbours removed tended to perform worse than plants that had their neighbours left intact. Growth observations for Ca. breviculmis were less clear, but again suggested facilitation. Mortality was distinctly higher among Ca. breviculmis individuals that had their neighbours removed relative to those with neighbours left intact. Results collectively suggest the removal of neighbours acts to reduce growth and increase mortality in C. pugioniformis and Ca. breviculmis throughout the altitudinal range of tall alpine herbfields at the Bogong High Plains. Facilitative and competitive interactions need to be recognized in efforts aimed at mitigating climate change‐associated impacts on the ecology of alpine plant communities. The extent to which biotic interactions may exacerbate or buffer abiotic change is difficult to predict, emphasizing the need for ecological monitoring.     

Biotic interactions mediate the expansion of black mangrove (Avicennia germinans) into salt marshes under climate change

Many species are expanding their distributions to higher latitudes due to global warming. Understanding the mechanisms underlying these distribution shifts is critical for better understanding the impacts of climate changes. The climate envelope approach is widely used to model and predict species distribution shifts with changing climates. Biotic interactions between species, however, may also influence species distributions, and a better understanding of biotic interactions could improve predictions based solely on climate envelope models. Along the northern Gulf of Mexico coast, USA, subtropical black mangrove (Avicennia germinans) at the northern limit of its distribution grows sympatrically with temperate salt marsh plants in Florida, Louisiana, and Texas. In recent decades, freeze‐free winters have led to an expansion of black mangrove into salt marshes. We examined how biotic interactions between black mangrove and salt marsh vegetation along the Texas coast varied across (i) a latitudinal gradient (associated with a winter‐temperature gradient); (ii) the elevational gradient within each marsh (which creates different marsh habitats); and (iii) different life history stages of black mangroves (seedlings vs. juvenile trees). Each of these variables affected the strength or nature of biotic interactions between black mangrove and salt marsh vegetation: (i) Salt marsh vegetation facilitated black mangrove seedlings at their high‐latitude distribution limit, but inhibited black mangrove seedlings at lower latitudes; (ii) mangroves performed well at intermediate elevations, but grew and survived poorly in high‐ and low‐marsh habitats; and (iii) the effect of salt marsh vegetation on black mangroves switched from negative to neutral as black mangroves grew from seedlings into juvenile trees. These results indicate that the expansion of black mangroves is mediated by complex biotic interactions. A better understanding of the impacts of climate change on ecological communities requires incorporating context‐dependent biotic interactions into species range models.     

The Effect of Social Hierarchy on Captive Coyote (Canis latrans) Foraging Behavior

Foraging efficiency of individuals in pack forming species may be influenced by social dynamics within a pack. The effects of social hierarchy in particular may influence individual foraging behavior in canids, such as coyotes (Canis latrans). To examine the impact of social hierarchy on foraging behavior, we tested 16 captive coyotes in eight naturally established dominant–subordinate pairs, using the guesser–knower paradigm. We measured the efficiency of subordinate coyotes to relocate a food resource when alone and then allowed pairs to forage together, such that subordinates had prior knowledge of food location but dominants did not. To determine whether (1) subordinates used a direct or discursive strategy to obtain food in the presence of a dominant and (2) dominants used an exploitative or independent strategy to obtain food in the presence of a subordinate with previous knowledge, we measured their search efficiency (e.g., correct choice of area, feeder, and latency to correct feeder). Results showed subordinates learned to relocate food and increase efficiency when alone. In a social context, however, subordinate efficiency decreased. That is, subordinates approached the correct area, but searched more feeders before finding the correct one. Dominants initially used an independent search strategy but then quickly displaced the subordinate and monopolized the resource, reducing subordinate efficiency further. Despite continual displacement and reduction in efficiency, subordinates did not alter their foraging strategy over time. Our results suggest prior information can improve individual foraging advantage, but that social status strongly impacts individual foraging efficiency in social species such as coyotes.     

Allostatic load, social status and stress hormones: the costs of social status matter

Cooperation and social support are the major advantages of living in social groups. However, there are also disadvantages arising from social conflict and competition. Social conflicts may increase allostatic load, which is reflected in increased concentrations of glucocorticoids. We applied the emerging concept of allostasis to investigate the relation between social status and glucocorticoid concentrations. Animals in a society experience different levels of allostatic load and these differences may predict relative glucocorticoid concentrations of dominant and subordinate individuals. We reviewed the available data from free-ranging animals and generated, for each sex separately, phylogenetic independent contrasts of allostatic load and relative glucocorticoid concentrations. Our results suggest that the relative allostatic load of social status predicts whether dominants or subordinates express higher or lower concentrations of glucocorticoids. There was a significant correlation between allostatic load of dominance and relative glucocorticoid concentrations in both females and males. When allostatic load was higher in dominants than in subordinates, dominants expressed higher levels of glucocorticoids; when allostatic load was similar in dominants and subordinates, there were only minor differences in glucocorticoid concentrations; and when allostatic load was lower in dominants than in subordinates, subordinates expressed higher levels of glucocorticoids than dominants. To our knowledge, this is the first model that consistently explains rank differences in glucocorticoid concentrations of different species and sexes. The heuristic concept of allostasis thus provides a testable framework for future studies of how social status is reflected in glucocorticoid concentrations.