Size matters: nest colonization patterns for twig‐nesting ants

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Strong influences of a dominant,ground‐nesting ant on recruitment,and establishment of ant colonies and communities

Many factors drive the organization of communities including environmental factors, dispersal abilities, and competition. In particular, ant communities have high levels of interspecific competition and dominance that may affect community assembly processes. We used a combination of surveys and nest supplementation experiments to examine effects of a dominant ground‐nesting ant (Pheidole synanthropica) on (1) arboreal twig‐nesting, (2) ground‐foraging, and (3) coffee‐foraging ant communities in coffee agroecosystems. We surveyed these communities in high‐ and low‐density areas of P. synanthropica over 2 years. To test for effects on twig ant recruitment, we placed artificial nesting resources on coffee plants in areas with and without P. synanthropica. The first sampling period revealed differences in ant species composition on the ground, in coffee plants, and artificial nests between high‐ and low‐density sites of P. synanthropica. High‐density sites also had significantly lower recruitment of twig ants and had species‐specific effects on twig ant species. Prior to the second survey period, abundance of P. synanthropica declined in the high‐density sites, such that P. synanthropica densities no longer differed. Subsequent sampling revealed no difference in total recruitment of twig ants to artificial nests between treatments. Likewise, surveys of ground and coffee ants no longer showed significant differences in community composition. The results from the first experimental period, followed by survey results after the decline in P. synanthropica densities suggest that dominant ants can drive community assembly via both recruitment and establishment of colonies within the community.     

Changes in Species Richness,Abundance, and Composition of Arboreal Twig‐nesting Ants Along an Elevational Gradient in Coffee Landscapes

The distribution, diversity, and assembly of tropical insects have long intrigued ecologists, and for tropical ants, can be affected by competitive interactions, microhabitat requirements, dispersal, and availability and diversity of nesting sites. Arboreal twig‐nesting ants are limited by the number of hollow twigs available, especially in intensive agricultural systems. Ant diversity and abundance may shift along elevation gradients, but no studies have examined if the proportion of occupied twigs or richness of arboreal twig‐nesting ants vary with elevation. In coffee agroecosystems, there are over 40 species of arboreal twig‐nesting ants. We examined communities of twig‐nesting ants in coffee plants along an elevational gradient to answer the following questions: (1) Do species richness and colony abundance decline with elevation or show a mid‐elevation peak? (2) Does community composition change with elevation? (3) Is elevation an important predictor of change in ant abundance, richness, and relative abundance of common species? We surveyed 42 10 × 10 m plots in 2013 from 450 to1550 m elevation across a coffee landscape in Chiapas, Mexico. We sampled a total of 2211 hollow coffee twigs, 77.1 percent of which were occupied by one of 28 species of ants. Pseudomyrmex simplex was more abundant in lower elevations, whereas Pseudomyrmex ejectus dominated in high elevations. Species richness and the percent of occupied hollow twigs both peaked at mid‐elevations (800–1050 m). In sum, we found that species richness, abundance, and composition of arboreal twig‐nesting ants shift with elevation. These findings may provide important insights for understanding ant communities in coffee agroecosystems.     

Influences on the structure of suburban ant (Hymenoptera: Formicidae) communities and the abundance of Tapinoma sessile

Urbanization can alter the organization of ant communities and affect populations of urban pest ants. In this study, we sampled ant communities in urban and suburban yards to understand the habitat factors that shape these communities and influence the abundance of a common pest species, Tapinoma sessile (Say). We used pitfall traps to sample ant communities and a combination of pitfall traps and baiting to collect T. sessile at 24 sites in Knoxville, TN. In total, we collected 46 ant species. Ant species richness ranged from seven to 24 species per yard. Ant species richness tended to be lowest near houses, whereas T. sessile abundance was highest near houses. The best predictors of ant species richness in yards were canopy cover and presence of leaf litter: ant species richness peaked at mid-levels of canopy cover and was negatively correlated with the presence of leaf litter. Tapinoma sessile abundance increased with presence of logs, boards, or landscaping timbers and leaf litter in yards. Our results indicate that ant communities and the abundance of particular pest species in these urban and suburban landscapes are shaped by many of the same factors that structure ant communities in less anthropogenically disturbed environments.     

Impact of Flooding on the Species Richness, Density and Composition of Amazonian Litter-Nesting Ants

Litter-nesting ants are diverse and abundant in tropical forests, but the factors structuring their communities are poorly known. Here we present results of the first study to examine the impact of natural variation in flooding on a highly diverse (21 genera, 77 species) litter-nesting ant community in a primary Amazonian forest. Fifty-six 3 × 3 m plots experiencing strong variation in flooding and twenty-eight 3 × 3 m terra firme plots were exhaustively searched for litter-nesting ants to determine patterns of density, species richness and species composition. In each plot, flooding, litter depth, twig availability, canopy cover, plant density, percent soil nitrogen, carbon, and phosphorus were measured. Degree of flooding, measured as flood frequency and flood interval, had the strongest impact on ant density in flooded forest. Flooding caused a linear decrease in ant abundance, potentially due to a reduction of suitable nesting sites. However, its influence on species richness varied: low-disturbance habitat had species richness equal to terra firme forest after adjusting for differences in density. The composition of ant genera and species varied among flood categories; some groups known to contain specialist predators were particularly intolerant to flooding. Hypoponera STD10 appeared to be well-adapted to highly flooded habitat. Although flooding did not appear to increase species richness or abundance at the habitat scale, low-flooding habitat contained a mixture of species found in the significantly distinct ant communities of terra firme and highly flooded habitat.
     

Ants on the Move: Resource Limitation of a Litter‐nesting Ant Community in Costa Rica1

The leaf litter of tropical wet forests is replete with itinerant ant nests. Nest movement may help ants evade the constraints of stress and disturbance and increase access to resources. I studied how nest relocation and environmental factors may explain the density, size, and growth of leaf litter ant nests. I decoupled the relationships among litter depth, food abundance, and nest availability in a 4‐mo manipulation of food and leaf litter in a community of litter‐nesting ants in a lowland wet forest in Costa Rica. Over 4 mo, 290 1 m² treatment and control plots were sampled without replacement. Nest densities doubled in response to food supplementation, but did not decrease in response to litter removal or stress (from litter trampling). The supplementation of food increased the utilization of less favored nesting materials. In response to food supplementation and litter trampling, arboreal ants established nests in the litter, and growth rates of the most common ants (Pheidole spp.) increased. Colony growth was independent of colony size and growth rates of the most abundant ants. In general, I conclude that litter‐nesting ant density is driven primarily by food limitation, that nest relocation behavior significantly affects access to resource and the demographic structure of this community, and that nest fission may be a method to break the growth–reproduction trade‐off.     

The effects of two abundant ant species on soil nutrients and seedling recruitment in Brazilian Atlantic Forest

Ants can influence soil fertility and the spatial distribution of seeds, with possible effects on seedling recruitment. The ant species Pachycondyla striata Fr. Smith, 1858 and Odontomachus chelifer (Latreille, 1802) co-occur in many forest areas in the Neotropics. We assessed soil fertility and seed bank structure in soil samples close and distant (control) from ant nests in forest fragments. We also assessed the richness and abundance of seedlings on nests and control sites. In soil samples from ant nests, the concentration of phosphorus and potassium were respectively 55.6% and 36% higher than in control sites. Aluminium was 11–15% lower in soil samples from ant nests. In the greenhouse, soils from ant nests had higher plant abundance and species richness, but the same species composition in comparison with control sites. Although more plants emerged from soil samples of O. chelifer nests, in the field, the density and richness of seedlings were similar for the two ant species studied. Seedlings in the nest sites were, on average, 1.8 times more abundant and 1.6 times richer in species than in control sites. Our results showed that ant species can play a key role in seedling recruitment in forest fragments, where other animals with equivalent and positive effects, such as mammals, are missing.     

A test of species–energy theory: patch occupancy and colony size in tropical rainforest litter‐nesting ants

Species–energy theory can account for spatial variation in the abundance and community composition of animals, though the mechanisms of species–energy theory are under contention. We evaluated three competing mechanisms at the local spatial scale by conducting an in vivo light manipulation over supplemental ant nests placed in the leaf litter of a Costa Rican tropical rainforest. We found that the light environment did not alter the 10% rate of occupation of the supplemental nests, but light did alter the size of colonies and the genus‐level composition of the community. Light levels in the foraging range were positively associated with colony sizes of all ants, whereas light levels directly on the nest site were predictive of the occurrence of ant genera. Colonies of specialized predators, dacetine ants, were larger in more shaded foraging environments, and the functional group of generalized myrmicines exhibited an opposite pattern, with smaller‐sized colonies in response to shading. Responses of twig‐dwelling ants to the light environment were most consistent with the metabolic cost hypothesis as a mechanism of species–energy theory. We found mixed support for the thermal energy availability hypothesis, and scant support for the chemical energy hypothesis, as the litter depth, a measure of prey density, was not predictive of ant responses. In summary, at the local scale, we found patterns in colony size and life history are governed by light‐dependent mechanisms.     

戈壁生态系统蚁穴微生境对大型土壤动物多样性的影响

收获蚁蚁穴是戈壁生态系统中重要的微生境, 它通过汇集凋落物和改善土壤环境强烈影响动植物的分布及多样性。鉴于此, 本文选择戈壁荒漠收获蚁(Messor desertus)蚁穴为研究对象, 于2020年5月、6月和10月利用陷阱法调查了蚁穴及毗邻裸地大型土壤动物的种类组成及数量变化, 并分析其影响要素。结果表明: (1) 10月, 蚁穴和裸地间大型土壤动物群落组成存在显著差异, 而5月和6月二者间相差较小, 10月(62.9%)蚁穴和裸地大型土壤动物的平均相异性高于5月(34.8%)和6月(39.3%); (2) 5月, 蚁穴大型土壤动物均匀度指数显著低于裸地, 6月, 蚁穴大型土壤动物活动密度和类群丰富度均显著高于裸地, 10月, 蚁穴大型土壤动物类群丰富度和多样性指数均显著高于裸地; (3)荒漠收获蚁蚁穴显著提高了其他食性土壤动物的活动密度及类群丰富度, 还增强了捕食性和非捕食性土壤动物的相互作用关系并改变了荒漠收获蚁与蚁穴大型土壤动物类群的种间相互作用关系; (4) pRDA排序结果表明, 土壤电导率、全氮和粉粒含量是影响蚁穴和裸地大型土壤动物分布的主要土壤因子。总之, 戈壁生态系统荒漠收获蚁蚁穴微生境提高了大型土壤动物多样性, 改变了大型土壤动物类群间的营养和非营养关系, 这会影响大型土壤动物的营养结构及其功能。     

Arboreal Ant Abundance and Leaf Miner Damage in Coffee Agroecosystems in Mexico

Shaded coffee agroecosystems traditionally have few pest problems potentially due to higher abundance and diversity of predators of herbivores. However, with coffee intensification (e.g., shade tree removal or pruning), some pest problems increase. For example, coffee leaf miner outbreaks have been linked to more intensive management and increased use of agrochemicals. Parasitic wasps control the coffee leaf miner, but few studies have examined the role of predators, such as ants, that are abundant and diverse in coffee plantations. Here, we examine linkages between arboreal ant communities and coffee leaf miner incidence in a coffee plantation in Mexico. We examined relationships between incidence and severity of leaf miner attack and: (1) variation in canopy cover, tree density, tree diversity, and relative abundance of Inga spp. shade trees; (2) presence of Azteca instabilis, an arboreal canopy dominant ant; and (3) the number of arboreal twig‐nesting ant species and nests in coffee plants. Differences in vegetation characteristics in study plots did not correlate with leaf miner damage perhaps because environmental factors act on pest populations at a larger spatial scale. Further, presence of A. instabilis did not influence presence or severity of leaf miner damage. The proportion of leaves with leaf miner damage was significantly lower where abundance of twig‐nesting ants was higher but not where twig‐nesting ant richness was higher. These results indicate that abundance of twig‐nesting ants in shaded coffee plantations may contribute to maintenance of low leaf miner populations and that ants provide important ecosystem services in coffee agroecosystems.     

Local and landscape drivers of biodiversity of four groups of ants in coffee landscapes

Agriculture of varying management intensity dominates fragmented tropical areas and differentially impacts organisms across and within taxa. We examined impacts of local and landscape characteristics on four groups of ants in an agricultural landscape in Chiapas, Mexico comprised of forest fragments and coffee agroecosystems varying in habitat quality. We sampled ground ants found in leaf litter and rotten logs and arboreal ants found in hollow coffee twigs and on tree trunks. Then using vegetation and agrochemical indices and conditional inference trees, we examined the relative importance of local (e.g. vegetation, elevation, agrochemical) and landscape variables (e.g. distance to and amount of nearby forest and rustic coffee) for predicting richness and abundance of ants. Leaf litter ant abundance increased with vegetation complexity; richness and abundance of ants from rotten logs, twig-nests, and tree trunks were not affected by vegetation complexity. Agrochemical use did not affect species richness or abundance of any ant group. Several local factors (including humus mass, degree of decay of logs, number of hollow twigs, tree circumference, and absence of fertilizers) were significant positive predictors of abundance and richness of some ant groups. Two landscape factors (forest within 200 m, and distance from forest) predicted richness and abundance of twig-nesting and leaf litter ants. Thus, different ant groups were influenced by different characteristics of agricultural landscapes, but all responded primarily to local characteristics. Given that ants provide ecosystem services (e.g. pest control) in coffee farms, understanding ant responses to local and landscape characteristics will likely inform farm management decisions.     

A canopy dominant ant affects twig‐nesting ant assembly in coffee agroecosystems

One commonly studied driver of community assembly is the effect of dominant species on subordinate species. Dominant species may impact community assembly during competitive sorting, or recruitment. For ants, important and abundant species in the tropics, several factors may drive community assembly including competition, dispersal, priority effects, and environmental conditions. Although competition is a hallmark of ant ecology, few have examined the influence of patchily distributed dominant ants on other ant species and diversity, especially at the recruitment stage. Here, I consider the impacts of a canopy dominant ant species, Azteca instabilis, and changes in vegetation on twig‐nesting ant colony founding and ant community assembly in a coffee agroecosystem. I added artificial nests to coffee plants in areas with and without A. instabilis four times over a year, and then examined the occupation rate and identity of species colonizing nests. I also examined vegetation characteristics of sites where nests were added. The presence of A. instabilis on coffee plants drastically lowered colonization rates, but nest occupation increased with tree density, and with decreasing proportion of Inga spp. trees in the canopy. The presence of A. instabilis limited the number of nests occupied by six of the ten most common species; most rare species, however, were not affected by A. instabilis presence. Richness of colonizing ants in areas with A. instabilis was lower, but these effects did not significantly affect richness across broader scales. Despite large effects on individual species, species composition did not differ greatly in areas with and without A. instabilis, but some vegetation characteristics (basal area and tree richness) were predictive of ant composition. These results suggest that A. instabilis strongly affects founding events especially for common twig‐nesting species and that both vegetation and influences from this dominant species affect community assembly of twig‐nesting ants at the local scale.     

Evaluating community effects of a Keystone Ant,Azteca sericeasur,on Inga micheliana leaf litter decomposition in a shaded coffee agro-ecosystem

Our research examined the effect of Azteca sericeasur, a keystone arboreal ant, on the decomposition of leaf litter of the shade tree, Inga micheliana, in coffee agro-ecosystems. This interaction is important in understanding spatial heterogeneity in decomposition. We hypothesized that A. sericeasur could affect leaf litter decomposition by excluding other ants, which could release decomposers, like collembolans, from predation pressure. Determining the relative strengths of these interactions can illuminate the importance of A. sericeasur in decomposition and nutrient cycling processes. We assessed the ant and arthropod communities surrounding 10 pairs of trees, where each pair included one shade tree with an established A. sericeasur nest. Tuna baits were used in conjunction with pitfall traps to assess the ant and arthropod community, and litterbags with I. micheliana leaf litter were used to assess rates of decomposition. The species richness of ants did not change in proximity to A. sericeasur nests, though the ant communities were distinct. Abundance of Collembola and community composition of other invertebrates did not change in the presence of A. sericeasur nests, and there were no differences in leaf litter decomposition rates. This contradicts past studies that suggest A. sericeasur reduces ant species richness in its territory. We suggest that other ants may avoid A. sericeasur by moving within and beneath the leaf litter. Our results indicate that there is no net effect of A. sericeasur on leaf litter decomposition.     

Impact of plant cover on the cavity‐nesting ant Temnothorax crassispinus

1. Plant communities influence the availability of important resources for ants, such as nest sites and food, as well as environmental conditions. Thus, plants affect the abundance and distribution of ants. 2. In a field experiment, the influence of plant cover on the settlement of nest sites and per‐capita productivity of sexual individuals by the ant Temnothorax crassispinus was analysed. In July 2014, in five areas with patches of alien balsam Impatiens parviflora, and another five of native balsam I. noli‐tangere, transects composed of artificial nests were established; the nest sites were situated inside patches of balsams, and outside of them. Four hundred and forty artificial nests were used. One year later, the nests were collected. 3. Colonies of the ants three times more often inhabited nest sites outside the patches of both balsams. Besides, colonies with queens were more frequently found in nest sites located away from balsams. The per‐capita productivity of sexual individuals was higher in nests collected in patches of balsam, and the colonies from patches of alien balsam produced a more female‐biased sex ratio. 4. In terms of the impact on the ant, no clear differences were found between the alien balsam and the native one. The most important factor affecting the fitness of ants in areas dominated by balsams is the presence of herbaceous plant cover rather than whether the plant is alien or native.     

Leaf‐cutting ants as ecosystem engineers: topsoil and litter perturbations around Atta cephalotes nests reduce nutrient availability

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Oil palm expansion into rain forest greatly reduces ant biodiversity in canopy,epiphytes and leaf-litter

Oil palm cultivation is expanding rapidly into many of the world's most biodiverse tropical regions. One of the most functionally important and ecologically dominant animal groups in these environments is the ants. Here, we quantify the overall impacts of clear-felling lowland dipterocarp rainforest and conversion into oil palm plantation on ant diversity. At study sites in Sabah, Malaysia we collected ants from three microhabitats: 1 – the canopy, 2 – bird's nest ferns (Asplenium nidus complex, a common epiphyte in forest and oil palm), and 3 – leaf litter. We also measured temperature, humidity and light at collection sites to assess their impacts on ant community composition. Total ant species richness decreased from 309 to 110 (?64%) between forest and oil palm plantation. However, this impact was not the same across all microhabitats, with bird's nest ferns maintaining almost the same number of ant species in oil palm compared to forest (forest-oil palm, ferns: 36–35 (3% loss), canopy: 120–58 (52% loss), leaf litter: 216–56 (74% loss)). Relative abundance distributions remained the same for fern-dwelling ants, but became less even for oil palm ants in both the canopy and the leaf litter. These differences may be due in part to the ability of bird's nest ferns to provide a stable microclimate in hot, dry plantations. We also found that non-native ant species were more abundant in oil palm than in forest, and few forest ant species survived in plantations in any of the microhabitats. Only 59 of the 309 forest species persisted in oil palm plantations, corresponding to an 81% loss of forest species resulting from habitat conversion. Although oil palm supports many more ant species than has been previously reported, converting forest into plantation still leads to a dramatic reduction in species richness. The maintenance of forested areas is therefore vital for the conservation of ant biodiversity.     

云南橡胶林和天然次生林枯落物层蚂蚁物种多样性、群落结构差异及指示种

橡胶树Hevea brasiliensis是云南省重要的经济林木, 但对其生态服务功能尚存在争议。本研究以天然次生林为对照, 使用Winkler法对橡胶林枯落物层蚂蚁进行初步研究, 探讨橡胶林枯落物层蚂蚁的生态状况。于2012年10月和2013年4月采用Winkler袋法调查了云南省绿春县大黑山乡橡胶林和牛孔乡天然次生林枯落物层蚂蚁群落的物种多样性、 群落结构差异及指示种。结果表明： 橡胶林枯落物层蚂蚁多度（转换后）、 物种丰富度S和ACE值显著低于无干扰的天然次生林（P<0.05）; 蚂蚁多度（转换后）显著低于有干扰的天然次生林（P<0.05）, 而物种丰富度S和ACE值差异不显著。橡胶林枯落物层蚂蚁群落结构与两种天然次生林都不相似（F=3.93, df=12, P<0.01）。橡胶林中流浪种大头蚁属Pheidole的蚂蚁种类与天然次生林相比, 物种丰富度增加了100%。天然次生林枯落物层中蚂蚁指示种有3种, 分别为刘氏隆头蚁Strumigenys lewisi、 黄足厚结猛蚁Pachycondyla luteipes和女娲角腹蚁Recurvidris nuwa, 而橡胶林枯落物层中指示种仅为菱结大头蚁Pheidole nodus。枯落物层蚂蚁物种多样性与枯落物厚度呈显著正相关, 而枯落物盖度仅与蚂蚁多度（转换后）有相关性。结果说明, 橡胶林经过长期的经营管理, 生态环境趋于稳定, 对枯落物层蚂蚁群落具有一定的保护作用, 但与天然次生林相比, 蚂蚁多度（转换后）及群落结构仍显示出明显的不同。     

Litter Biomass and Nutrient Determinants of Ant Density, Nest Size, and Growth in a Costa Rican Tropical Wet Forest

The nutritional demands of animals vary by taxon. Across landscapes, communities of animals experience variability in the stoichiometry of carbon and nutrients within their resource base. Thus, we expect stoichiometry to contribute to the spatial variance in the demographic parameters of animal communities. Here, we measure how the composition of a litter-nesting tropical rainforest ant community is influenced by spatial variation in environmental stoichiometry relative to litter biomass, a known predictor of ant density. We found the density of ants and their nests were strongly related to litter biomass and carbon: phosphorus stoichiometry. The spatial variation in soil nutrients, which determines leaf litter stoichiometry, was an excellent predictor of nest size in the two most common genera of ants. We found a negative relationship between species' growth rate and local soil stocks of phosphorus. Overall, the density of litter-dwelling ants varied greatly across this tropical forest landscape and environmental stoichiometry can account for limits on ant density independent of the biomass of the leaf litter resource base.     

Spatial patterns of granivorous ant nest abundance and nest site selection in agricultural landscapes of Central Spain

Summary Patterns of abundance and site selection of granivorous ant nests were investigated in extensive cereal croplands of Central Spain. Nest densities and distributions were measured in two consecutive summers (1988 and 1989), together with habitat physiognomy and seed availability. Nest site selection patterns were analysed at two spatial scales (landscape and microhabitat) with respect to habitat physiognomy. Results indicate a very constant and predictable pattern of both nest abundance and nest site selection. Granivorous ant nests were most abundant in shrublands, and shrubby microsites were selected for nest placement. Croplands, and microsites with high covers of bare ground and litter, were avoided. These patterns were consistent between years despite a 1.7-fold increase in shrubland nest densities, that was attributed to the exceptionally dry winter between nest censuses. I suggest that winter survivorship of ant nests in the unploughed landscape units, and periodic ploughing in croplands, may be the main factors constraining granivorous ant densities in the landscape studied.     

Limitation of nesting resources for ants in Colombian forests and coffee plantations

Abstract.  1. This study examines limitation of nesting resources for leaf-litter and twig-nesting ants as a mechanism of diversity loss across an intensification gradient of coffee production in Colombia. Twelve farms were selected and classified into four management types: forest, polygeneric shade coffee, monogeneric shade coffee, and sun coffee (unshaded coffee monocultures).
2. At each of the farms, four treatment subplots were established at the corners of each of 10 25 m² plots: (i) twig augmentation (adding 10 empty bamboo twigs); (ii) litter augmentation (tripling existing litter profile); (iii) twig and litter augmentation; and (iv) no manipulation control, for a total of 480 subplots. A twig addition experiment was also performed on coffee bushes.
3. The results showed significantly more ant colonies in the forest and monogeneric shade coffee litter augmentation plots after 4 months. Litter-nesting ant species richness was higher in all three shade systems than in the sun coffee. The identities of ants nesting on coffee bushes were different from those in the soil level litter. Fewer species nested in bamboo twigs placed in litter in the most intensive systems.
4. More ants nested in the resource addition treatments, and more ant species were found in forested habitats; however, a single mechanism cannot explain the observed patterns. It was concluded that a combination of bottom-up and top-down effects might lead to the loss of associated fauna with the intensification of these agroecosystems.