Inter-specific aggression generates ant mosaics in canopies of primary tropical rainforest

The ant mosaic is a concept of the non-random spatial distribution of individual ant species in trees built upon the assumption of interspecific behavioural associations. However, colony identity and environmental variance may also play a role in species distribution. Here we assess the presence of ant mosaics in a primary forest ecosystem and whether they are structured by species' aggressive behaviours or by habitat filtering. We sampled arboreal ants from vertically stratified baits exposed in 225 canopy trees in a 9-ha plot of primary lowland forest in Papua New Guinea, the largest forest area surveyed to detect ant mosaics. We performed behavioural tests on conspecific ants from adjacent trees to determine the territories of individual colonies. We explored the environmental effects on the ant communities using information on the plot vegetation structure and topography. Furthermore, we created a novel statistical method to test for the community non-random spatial structure across the plot via spatial randomisation of individual colony territories. Finally, we linked spatial segregation among the four most common species to experimentally assessed rates of interspecies aggression. The ant communities comprised 57 species of highly variable abundance and vertical stratification. Ant community composition was spatially dependent, but it was not affected by tree species composition or canopy connectivity. Only local elevation had a significant but rather small effect. Individual colony territories ranged from one tree to 0.7 ha. Species were significantly over-dispersed, with their territory overlap significantly reduced. The level of aggression between pairs of the four most common species was positively correlated with their spatial segregation. Our study demonstrates the presence of ant mosaics in tropical pristine forest, which are maintained by interspecific aggression rather than habitat filtering, with vegetation structure having a rather small and indirect effect, probably linked to microclimate variability.     

Extreme life history plasticity and the evolution of invasive characteristics in a native ant

Disturbance resulting from urbanization is a leading cause of biotic homogenization worldwide. Native species are replaced with widespread non-native species and ants are among the world’s most notorious invaders. To date, all documented cases of ant invasions involve exotic introduced species that are spread around the world by human-mediated dispersal. I investigated the effect of urbanization on the evolution of invasive characteristics in a native ant species, the odorous house ant, Tapinoma sessile (Say). Colony social structure, life history traits, and the spatial pattern of nest distribution were compared by sampling T. sessile across a gradient of three distinct habitats: natural, semi-natural, and urban. Results demonstrate a remarkable transition in colony social and spatial structure and life history traits between natural and urban environments. In natural habitats, T. sessile colonies are comprised of small, monogyne (single queen), and monodomous (single nest) colonies. In urban areas, T. sessile often exhibit extreme polygyny and polydomy, form large supercolonies, and become a dominant pest. Results also suggest that urban T. sessile colonies may have a negative impact on native ant abundance and diversity. In the natural environment T. sessile coexisted with a wide array of other ant species, while very few ant species were present in the urban environment invaded by T. sessile. Habitat degradation and urbanization can lead to extreme changes in social and spatial colony structure and life history traits in a native ant species and can promote the evolution of invasive characteristics such as polygyny, polydomy, and supercolonial colony structure.     

Spatial structure of litter-dwelling ant distribution in a subtropical dry forest

Understanding the spatial patterns of species distribution is essential to characterize the structure of communities, to optimize species inventories and to evaluate the impact of biotic and abiotic variables. Here we describe the spatial structure of the distribution of leaf litter ant species, and of biotic factors that could explain it, in a subtropical semi-deciduous forest of the Argentinian Chaco, characterized by a dense understorey of shrubs and terrestrial bromeliads. Environmental variables (leaf litter quantity and ground bromeliad density) were measured and ants were collected in 1 m² quadrats distributed along two 200 m transects at intervals of 1.25 m. Overall 87 species were collected. Sixteen positive associations and a single negative association were observed between the 11 most frequent species taken pair-wise. Our results suggest that the spatial distribution of leaf litter ants was determined at two different scales. At a small scale (period below 10 m) a periodic spatial structure, likely due to intraspecific competition, produced a succession of peaks of abundance separated by gaps. At a larger scale (period around 50 m), periodically distributed environmental factors induced aggregates of colonies of species responding positively to these factors. A high quantity of leaf litter and, to a lesser extent, a high density of ground bromeliads promoted a high density and a high species richness of ants. Numerically dominant ants being generally positively associated, interspecific competition was apparently weak. All ant species whose abundance was correlated with an environmental factor were not completely spatially structured by it. This suggests that some other factors, such as intraspecific competition, may have counter-effects. Received 14 March 2005; revised 26 April 2005; accepted 10 May 2005.     

Partitioning of resources amongst seed-harvesters in an ant community in semi-arid Australia

Resource partitioning among five species of seed-harvesting ants in a semi-arid saltbush and adjacent grassland habitat of southwestern New South Wales is analysed. While habitat differences between species were only slight, the utilization of two resource dimensions, food type and time available for activity by them, varied markedly in response to environmental changes. The degree to which species overlapped in their use of these resources also fluctuated as conditions varied. Thus, both the manner and extent to which the harvester ant species interact is rapidly and continually changing in response to environmental fluctuations. The results suggest that, at most times, there is very little pressure due to exploitative competition within the harvester ant community.     

Structure and succession of ant assemblages in a north European sand dune area

Ant assemblages of a successional dune area (Tvärminne, south Finland) were analysed at three levels: the distribution of species among habitats, the distribution of colonies within habitats and the spatio-temporal distribution of individuals. The distribution of ant species among habitat patches representing different stages of succession is correlated with the composition of the epigeic fauna, the number and condition of dead twigs on the ground and the vegetation architecture. The composition of ant assemblages seems relatively independent of the plant species composition, and there are uncoordinated successional steps between the vegetation and ant communities.
In the early successional stages interference competition is weak and insignificant, as indicated by the random spatial arrangement of colonies, the absence of postcompetitive niche segregation and the low encounter rates of ant workers on the ground and at baits. In the aggressive behavioural hierarchy, Formica cinerea is a submissive species as compared with either Formica sanguinea or Lasius alienus.     

An Ant Mosaic Revisited: Dominant Ant Species Disassemble Arboreal Ant Communities but Co-Occur Randomly

The spatial distributions of many tropical arboreal ant species are often arranged in a mosaic such that dominant species have mutually exclusive distributions among trees. These dominant species can also mediate the structure of the rest of the arboreal ant community. Little attention has been paid to how diet might shape the effects of dominant species on one another and the rest of the ant community. Here, we take advantage of new information on the diets of many tropical arboreal ant species to examine the intra- and inter-guild effects of dominant species on the spatial distribution of one another and the rest of the tropical arboreal ant community in a cocoa farm in Bahia, Brazil. Using null model analyses, we found that all ant species, regardless of dominance status or guild membership, co-occur much less than expected by chance. Surprisingly, the suite of five dominant species showed random co-occurrence patterns, suggesting that interspecific competition did not shape their distribution among cocoa trees. Across all species, there was no evidence that competition shaped co-occurrence patterns within guilds. Co-occurrence patterns of subordinant species were random on trees with dominant species, but highly nonrandom on trees without dominant species, suggesting that dominant species disassemble tropical arboreal ant communities. Taken together, our results highlight the often complex nature of interactions that structure species-rich tropical arboreal ant assemblages.     

Species richness and structure of ant communities in a dynamic archipelago: effects of island area and age

Aim To assess how ant species richness and structure of ant communities are influenced by island age (disturbance history) in a dynamic archipelago. Location Cabra Corral dam, Salta Province, north‐west Argentina (25°08′ S, 65°20′ W). Methods Ant species richness on remaining fragments (islands) of a flooded forest was determined, as well as island area, isolation and age. Simple linear regressions were performed to assess relationships between ant species richness and those insular variables. Furthermore, a stepwise multiple linear regression analysis was conducted in order to determine the relative influence of each insular variable on ant species richness. Islands were categorized in two age classes (old and young) and co‐occurrence analyses were applied within each class to evaluate changes in community structure because of interspecific competition. Results Simple regression analyses indicated a moderate, positive effect of island area on ant species richness. Weak, marginally non‐significant relationships were found between ant species richness and both island isolation and island age, showing the tendency for there to be a decrease in ant species richness with island isolation and that ant species richness might be higher in old islands. The multiple regression analysis indicated that island isolation and age had no significant effects on the number of ant species, island area being the only independent variable retained in the analysis. On the contrary, whereas a random pattern of species co‐occurrence was found on young islands, ant communities in old islands showed a significantly negative pattern of species co‐occurrence, suggesting that the effect of competition on community structure was stronger on older islands than on younger islands. Main conclusions Island area was the most important variable explaining ant species richness on the islands of Cabra Corral dam. However, both island isolation and island age (or disturbance history) might also contribute to shape the observed community patterns. The present study also shows that island age significantly affects the strength with which interspecific interactions structure ant communities on islands.     

Genetic population structure, queen supersedure and social polymorphism in a social Hymenoptera

In social insects, the emergence of multiple queening is linked to changes in a suite of traits such as the reproductive life span of queens, mating patterns and population structure. We investigated queen turnover, colony longevity, spatial distribution patterns and genetic differentiation in a population of the socially polymorphic ant Formica fusca. Genetic differentiation between the social forms was absent, and mating patterns were similar in the two forms. The spatial distribution of single- and multi-queen colonies indicated an absence of colony reproduction by budding in both colony types. However, the rate of queen supersedure was high in multi-queen colonies and absent in single-queen ones. The social structure of colonies remained stable across years, but colony mortality did not differ between the two social forms. These results imply that differences between social types may appear and persist also in sympatry, and that these differences may occur in some traits, but not others, despite the presence of homogenizing gene flow.     

Ant mosaics in a tropical rainforest in Australia and elsewhere: A critical review

Abstract The concept of ‘ant mosaics’ has been established to describe the structure of arboreal ant communities in plantations and other relatively simple forest systems. It is essentially built upon the existence of negative and positive associations between ant species plus the concept of dominance hierarchies. Whether this concept can be applied to ant communities in more complex mature tropical rain forests has been questioned by some authors. Here we demonstrate that some previous attempts to prove or disprove the existence of such ant mosaics sampled by knockdown insecticide canopy fogging in near pristine tropical forests may have been thwarted by poor statistical power and too coarse spatial resolution, and the conclusions may be highly dependent on ant species and forest stratum selected for the study. Moreover, the presence or absence of ant mosaics may be driven by the density of suitable resources. We use an intensively studied ant community in the lowland rainforests of North‐East Queensland, Australia to outline processes that may lead to ant mosaic patterns, reasoning that competition for highly predictable resources in space and time such as honeydew and nectar is a fundamental process to maintain the mosaic structure. Honeydew and nectar sources, particularly their amino acids, are of crucial importance for nourishment of arboreal ant species. We use canopy fogging data from the same site in Australia and from two mature rainforests in South‐East Asia to compare spatial avoidance and co‐occurrence patterns implied by ant mosaics. Significant negative and positive associations were found among the three most abundant ant species in each dataset. Several problems with such spatial analyses are discussed, and we suggest that studies of ant mosaics in complex rainforest communities would benefit from a more focused approach on patterns of resource distribution and their differential utilisation by ants.     

Ant assemblage structure on cocoa trees in smallholder farms in the Centre Region of Cameroon

We investigated the ant community structure in cocoa farms in the Centre Region of Cameroon. Ants were collected on the cocoa trees during the years 2006 and 2007 using chemical knock‐down. We tested the hypothesis of the existence of deterministic factor in the structuration of ant mosaic using C‐Score; we assessed the relationship between the numerical dominant and subdominant ant species using Spearman correlation test and discussed on the influence of vegetation structure and farm management on the ant community structure. A total of 53 ant species belonging to 20 genera and five subfamilies were identified from a set of 51,525 workers collected. C‐score analysis supported the hypothesis that ant community were structured by competition. Negative relationships were found between dominant ant species. Farming practices which were mainly pruning, chemical treatment and habitat structure appeared to influence the ecological status and distribution of dominant ant species.     

Ectoparasite community structure on three closely related seabird hosts: a multiscale approach combining ecological and genetic data

Parasite communities can be structured at different spatial scales depending on the level of organization of the hosts; hence, examining this structure should be a multiscale process. We investigated ectoparasite community structure in three closely related seabird hosts, the Mediterranean Cory's shearwater Calonectris diomedea diomedea , the Atlantic Cory's shearwater C. d. borealis and the Cape Verde shearwater C. edwardsii . This community was composed of three lice ( Halipeurus abnormis , Austromenopon echinatum and Saemundssonia peusi) and one flea species ( Xenopsylla gratiosa ), and was considered at the infra-, component and regional community levels. We examined temporal and spatial structuring of the infracommunities, the influence of host aggregation and body condition on the component community, and the effect of genetic and geographic connectivity among host populations on the regional community. Ectoparasite infracommunities showed substantial species overlaps in temporal patterns of abundance, but species were spatially segregated within the host body. Within component communities, all ectoparasite species showed an aggregated distribution in abundance. However, aggregation patterns and their relationships with the spatial distribution of hosts within the breeding colony differed among ectoparasite species, mainly reflecting ecological differences between fleas and lice. At the regional scale, similarity in ectoparasite communities correlated with geographic distances among host colonies, but not with genetic distances. This result suggests differences in climate and habitat characteristics among host localities as a major determinant of regional communities, rather than host connectivity. Taken together, our results highlight the importance of the geographic distribution of host breeding colonies and the spatial segregation within the host body as key factors in determining ectoparasite community structure in Calonectris shearwaters.     

热带森林不同演替阶段蚂蚁巢穴的分布特征及其影响因素

为探明热带森林蚂蚁巢穴的分布特征及其影响因素, 采用样方法研究了西双版纳不同演替阶段热带森林定居巢穴蚂蚁的种类及其巢穴的密度、盖度和空间分布特征, 并分析了土壤理化环境与蚂蚁种类总数、巢穴密度及盖度的相关性。结果表明, 不同演替阶段热带森林蚂蚁种类总数、巢穴的密度及盖度大小顺序为: 小果野芭蕉 (Musa acuminata)群落>白背桐(Mallotus paniculatus)群落>思茅崖豆(Mellettia leptobotrya)群落, 并且热带森林的演替类型显著影响蚂蚁种类总数及巢穴密度, 而对巢穴盖度的影响未达到显著水平; 蚂蚁种类总数、巢穴密度与土壤总有机碳和水解氮显著正相关, 与土壤容重和土壤含水率显著负相关, 但所选择的土壤理化指标与巢穴盖度的相关性均未达到显著水平; 蚂蚁巢穴的空间分布呈随机分布格局。我们的数据表明, 不同演替阶段热带森林所形成的植被类型及土壤环境状况共同影响定居的蚂蚁种类总数与筑巢密度。     

Spatial Distribution of Dominant Arboreal Ants in a Malagasy Coastal Rainforest: Gaps and Presence of an Invasive Species

We conducted a survey along three belt transects located at increasing distances from the coast to determine whether a non-random arboreal ant assemblage, such as an ant mosaic, exists in the rainforest on the Masoala Peninsula, Madagascar. In most tropical rainforests, very populous colonies of territorially dominant arboreal ant species defend absolute territories distributed in a mosaic pattern. Among the 29 ant species recorded, only nine had colonies large enough to be considered potentially territorially dominant; the remaining species had smaller colonies and were considered non-dominant. Nevertheless, the null-model analyses used to examine the spatial structure of their assemblages did not reveal the existence of an ant mosaic. Inland, up to 44% of the trees were devoid of dominant arboreal ants, something not reported in other studies. While two Crematogaster species were not associated with one another, Brachymyrmex cordemoyi was positively associated with Technomyrmex albipes, which is considered an invasive species—a non-indigenous species that has an adverse ecological effect on the habitats it invades. The latter two species and Crematogaster ranavalonae were mutually exclusive. On the other hand, all of the trees in the coastal transect and at least 4 km of coast were occupied by T. albipes, and were interconnected by columns of workers. Technomyrmex albipes workers collected from different trees did not attack each other during confrontation tests, indicating that this species has formed a supercolony along the coast. Yet interspecific aggressiveness did occur between T. albipes and Crematogaster ranavalonae, a native species which is likely territorially dominant based on our intraspecific confrontation tests. These results suggest that the Masoala rainforest is threatened by a potential invasion by T. albipes, and that the penetration of this species further inland might be facilitated by the low density of native, territorially dominant arboreal ants normally able to limit its progression.     

The Regulation of Ant Colony Foraging Activity without Spatial Information

Many dynamical networks, such as the ones that produce the collective behavior of social insects, operate without any central control, instead arising from local interactions among individuals. A well-studied example is the formation of recruitment trails in ant colonies, but many ant species do not use pheromone trails. We present a model of the regulation of foraging by harvester ant (Pogonomyrmex barbatus) colonies. This species forages for scattered seeds that one ant can retrieve on its own, so there is no need for spatial information such as pheromone trails that lead ants to specific locations. Previous work shows that colony foraging activity, the rate at which ants go out to search individually for seeds, is regulated in response to current food availability throughout the colony's foraging area. Ants use the rate of brief antennal contacts inside the nest between foragers returning with food and outgoing foragers available to leave the nest on the next foraging trip. Here we present a feedback-based algorithm that captures the main features of data from field experiments in which the rate of returning foragers was manipulated. The algorithm draws on our finding that the distribution of intervals between successive ants returning to the nest is a Poisson process. We fitted the parameter that estimates the effect of each returning forager on the rate at which outgoing foragers leave the nest. We found that correlations between observed rates of returning foragers and simulated rates of outgoing foragers, using our model, were similar to those in the data. Our simple stochastic model shows how the regulation of ant colony foraging can operate without spatial information, describing a process at the level of individual ants that predicts the overall foraging activity of the colony.     

Positive interactions between desert granivores: localized facilitation of harvester ants by kangaroo rats

Facilitation, when one species enhances the environment or performance of another species, can be highly localized in space. While facilitation in plant communities has been intensely studied, the role of facilitation in shaping animal communities is less well understood. In the Chihuahuan Desert, both kangaroo rats and harvester ants depend on the abundant seeds of annual plants. Kangaroo rats, however, are hypothesized to facilitate harvester ants through soil disturbance and selective seed predation rather than competing with them. I used a spatially explicit approach to examine whether a positive or negative interaction exists between banner-tailed kangaroo rat (Dipodomys spectabilis) mounds and rough harvester ant (Pogonomyrmex rugosus) colonies. The presence of a scale-dependent interaction between mounds and colonies was tested by comparing fitted spatial point process models with and without interspecific effects. Also, the effect of proximity to a mound on colony mortality and spatial patterns of surviving colonies was examined. The spatial pattern of kangaroo rat mounds and harvester ant colonies was consistent with a positive interspecific interaction at small scales (<10 m). Mortality risk of vulnerable, recently founded harvester ant colonies was lower when located close to a kangaroo rat mound and proximity to a mound partly predicted the spatial pattern of surviving colonies. My findings support localized facilitation of harvester ants by kangaroo rats, likely mediated through ecosystem engineering and foraging effects on plant cover and composition. The scale-dependent effect of kangaroo rats on abiotic and biotic factors appears to result in greater founding and survivorship of young colonies near mounds. These results suggest that soil disturbance and foraging by rodents can have subtle impacts on the distribution and demography of other species.     

蚂蚁群落与栖境关系研究进展及新趋势

生物群落与栖境的关系是生态学研究的核心之一,蚂蚁群落由于在陆地生态系统中的生物量、分布以及具备的生态功能的重要性,是研究这种关系的理想对象。在查阅大量文献的基础上,简述了蚂蚁物种多样性与栖境关系研究现状。介绍了蚂蚁功能群划分以及在不同尺度上与栖境关系的应用研究,评述了功能群应用的限制。阐明了功能特征的定义以及基于形态特征和营养级方面的蚂蚁群落功能特征与栖境的研究,并对功能特征的研究趋势进行了展望。     

Microsite and litter cover effects on seed banks vary with seed size and dispersal mechanisms: implications for revegetation of degraded saline land

Seed movements and fates are important for restoration as these determine spatial patterns of recruitment and ultimately shape plant communities. This article examines litter cover and microsite effects on seed availability at a saline site revegetated with Eucalyptus sargentii tree rows interplanted with 5?C6 rows of saltbush (Atriplex spp.). As litter accumulation decreases with increasing distance from tree rows, soil seed banks were compared between paired bare and litter-covered zones within three microsites; tree row, saltbush row closest to tree row and saltbush mid-row (middle row of saltbush between tree rows). Germinable seed banks of the four most abundant species with contrasting seed sizes and dispersal mechanisms were assessed to test the hypotheses that: (i) microsites with litter cover contain higher seed densities than bare areas, but that (ii) microsite and litter effects will vary depending on seed size and dispersal mechanisms. Overall, litter cover increased seed densities, however, litter effects varied with seed size, with no effect on small-seeded species and litter increasing densities of large-seeded species. Seed bank composition also differed between tree and shrub microsites due to differences in seed morphology and dispersal mechanisms. Water-dispersed species were unaffected by microsite but densities of wind-dispersed species, including Atriplex spp., were higher in saltbush microsites. Densities of wind-dispersed species also differed between the two saltbush microsites despite similar litter cover. Future plantings should consider row spacing and orientation, as well as the dimensions of seeding mounds and associated neighbouring depressions, to maximize litter and seed-trapping by microsites.     

Ecosystem engineering and predation: the multi-trophic impact of two ant species

1. Ants are ubiquitous ecosystem engineers and generalist predators and are able to affect ecological communities via both pathways. They are likely to influence any other terrestrial arthropod group either directly or indirectly caused by their high abundance and territoriality. 2. We studied the impact of two ant species common in Central Europe, Myrmica rubra and Lasius niger, on an arthropod community. Colony presence and density of these two ant species were manipulated in a field experiment from the start of ant activity in spring to late summer. 3. The experiment revealed a positive influence of the presence of one ant colony on densities of decomposers, herbivores and parasitoids. However, in the case of herbivores and parasitoids, this effect was reversed in the presence of two colonies. 4. Generally, effects of the two ant species were similar with the exception of their effect on Braconidae parasitoid densities that responded positively to one colony of M. rubra but not of L. niger. 5. Spider density was not affected by ant colony manipulation, but species richness of spiders responded positively to ant presence. This effect was independent of ant colony density, but where two colonies were present, spider richness was significantly greater in plots with two M. rubra colonies than in plots with one colony of each ant species. 6. To test whether the positive ecosystem engineering effects were purely caused by modified properties of the soil, we added in an additional experiment (i) the soil from ant nests (without ants) or (ii) unmodified soil or (iii) ant nests (including ants) to experimental plots. Ant nest soil on its own did not have a significant impact on densities of decomposers, herbivores or predators, which were significantly, and positively, affected by the addition of an intact nest. 7. The results suggest an important role of both ant species in the grassland food web, strongly affecting the densities of decomposers, herbivores and higher trophic levels. We discuss how the relative impact via bottom-up and top-down effects of ants depends on nest density, with a relatively greater top-down predatory impact at higher densities.     

Experimental confirmation of competition between two dominant species of Iridomyrmex (Hymenoptera: Formicidae)

The well-defined ant succession following disturbance of coastal heathland in eastern Australia includes community replacement triggered by one dominant species of Iridomyrmex (species C) being replaced by another (species A). This paper investigates the consequences of experimentally reducing the abundance of species A by injecting petrol into the nest entrances of colonies. The response of species C was monitored at 2 and 10 months after removal and shows highly significant increases in the number of pitfall traps occupied and the area of territory maintained by this species. It is confirmed that interspecific competition plays an important role in maintaining the observed mosaic pattern of ant distribution and is also important in ant succession. The mechanism of competition is interference or, more exactly, territorial competition. The species appear to actively defend territorial space by patrolling territory boundaries that are breached when the defenders are eliminated.