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.     

Vegetation connectivity increases ant activity and potential for ant‐provided biocontrol services in a tropical agroforest

In natural and managed systems, connections between trees are important structural resources for arboreal ant communities with ecosystem‐level effects. However, ongoing agricultural intensification in agroforestry systems, which reduces shade trees and connectivity between trees and crop plants, may hinder ant recruitment rates to resources and pest control services provided by ants. We examined whether increasing connectivity between coffee plants and shade trees in coffee plantations increases ant activity and enhances biological control of the coffee berry borer, the most devastating insect pest of coffee. Further, we examined whether artificial connections buffer against the loss of vegetation connectivity in coffee plants located at larger distances from the nesting tree. We used string to connect Inga micheliana shade trees containing Azteca sericeasur ant nests to coffee plants to compare ant activity before and after placement of the strings, and measured borer removal by ants on coffee plants with and without strings. Ant activity significantly increased after the addition of strings on connected plants, but not on control plants. Borer removal by ants was also three times higher on connected plants after string placement. Greater distance from the nesting tree negatively influenced ant activity on control coffee plants, but not on connected plants, suggesting that connections between coffee plants and nest trees could potentially compensate for the negative effects that larger distances pose on ant activity. Our study shows that favoring connectivity at the local scale, by artificially adding connections, promotes ant activity and may increase pest removal in agroecosystems. Abstract in Spanish is available with online material.     

Associations between invasive eucalyptus psyllids and arthropod litter communities under tree canopies in southern California

Honeydew‐producing psyllids are an important pest of eucalyptus (Myrtaceae) in California, USA, and may influence surrounding litter arthropod communities. In particular, the introduced Australian psyllids Glycaspis brimblecombei Moore and Eucalyptolyma maideni Froggatt (both Hemiptera: Psyllidae) may facilitate the prevalence of invasive ant species. We examined ground‐dwelling arthropod communities under eucalyptus trees infested by psyllids. We used a model comparison approach to examine the association of psyllid infestation, ant abundance, and environmental factors with ground arthropod abundance and richness. We found a significant positive association between ant activity on eucalyptus trees and psyllid abundance. Higher psyllid abundance and higher Argentine ant abundance were associated with increased arthropod richness. Irrigation was also associated with increased arthropod richness and abundance. Regardless of location collected, arthropod communities collected in pitfall traps under trees with high psyllid abundance had high similarity to arthropod communities under trees with high ant activity. Abundance of isopods was positively associated with both ant and psyllid abundance. Other arthropod groups differed in their association with ants and psyllids. Argentine ants may exacerbate pest impacts and may also decrease the effectiveness of biological control programs for eucalyptus lerp psyllids.     

Size matters: nest colonization patterns for twig‐nesting ants

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Nutrient transfer supports a beneficial relationship between the canopy ant,Azteca trigona,and its host tree

1. Energy fluxes between ants and plants have been a focal point for documenting mutualistic behaviour. Plants can provide resources to ants through the production of extrafloral nectaries. In exchange, ants can fertilise plants through their nutrient‐ and microbe‐rich refuse. 2. Here, we test a potential facultative mutualism between the carton‐nesting canopy ant, Azteca trigona, and their host trees. Through observational and experimental approaches, this study documents how nutrient transfer provides a basis for this beneficial ant–plant relationship. 3. In a greenhouse experiment, fertilisation with sterilised refuse (i.e. nutrients only) increased seedling growth three‐fold, while the refuse with its natural microbial community increased growth 11‐fold. 4. Total root density was doubled in refuse piles compared with the surrounding area in situ. On average, refuse provides host trees and the surrounding plant community with access to a > 800% increase in N, P and K relative to leaf litter. 5. Azteca trigona preferentially nests in trees with extrafloral nectaries and on large, longer‐lived tree species. 6. Given the nutrient‐poor nature of the Neotropics, host trees probably experience significant benefits from refuse fertilisation. Conversely, A. trigona benefit from long‐term stable structural support for nests and access to nutrient‐rich extrafloral nectaries. Without clear costs to either A. trigona or host trees, it is proposed that these positive interactions are preliminary evidence of a facultative mutualism.     

Does Argentine ant invasion affect prey availability for foliage-gleaning birds?

Food availability during the breeding season plays a critical role in reproductive success of insectivorous birds. Given that the invasive Argentine ant (Linepithema humile) is known to alter arthropod communities, we predicted that its invasion may affect the availability of food resources for coexisting foliage-gleaning birds. With this aim we studied, for 3 years, foliage arthropods occurring on cork oaks (Quercus suber) and tree heaths (Erica arborea) in invaded and non-invaded secondary forests of the northeastern Iberian Peninsula. Our results show that Argentine ants interact with arboreal foliage arthropods in a different manner than the native ants they displace do. The invasive ant impacted the arthropod community by reducing order diversity and ant species richness and by causing extirpation of most native ant species. Arthropod availability for foliage gleaners’ nestlings diminished in invaded cork oaks, mainly responding to the abundance and biomass depletion of caterpillars. Results suggest that the reproduction of canopy-foraging foliage-gleaning species that mostly rely on caterpillars to feed their young could be compromised by the Argentine ant invasion. Thus, the Argentine ant could be promoting bottom-up effects in the trophic web through its effects on the availability of arthropod preys for insectivorous birds.     

Contribution of cocoa plantations to the conservation of native ants (Insecta: Hymenoptera: Formicidae) with a special emphasis on the Atlantic Forest fauna of southern Bahia,Brazil

By maintaining a forest-like structure, shaded cocoa plantations contribute to the conservation of ants that usually live in the soil, leaf litter or canopy of tropical forests. Here we synthesize the available information on the diversity and community structure of ants in shaded cocoa plantations in the Atlantic forest region of Brazil, compare ant assemblages in cocoa agroforests with forests and other forms of agriculture, and discuss how these shaded plantations contribute to the conservation of the ants in the Atlantic Forest region. We also discuss ants of economical importance and of special interest, including Camponotus, Dolichoderus, Gnamptogenys, Pachycondyla, Pseudomyrmex and other litter dwelling genera. We discuss the situation of the tramp ant Wasmannia auropunctata in the Bahian cocoa-producing region where it is considered as native, and that of the two cryptobiotic genera Thaumatomyrmex and Typhlomyrmex, as well as that of proven and possible endangered army ant and Ponerini species. A total of 192 ant species from four strata were found in extensive sampling of a cocoa plantation with a relatively simple shade canopy (comprised primarily of Erythrina). Species richness in the cocoa plantations corresponded roughly to that of low diversity native forests, and species composition of cocoa plantations was most similar to native habitats (forest and mangroves) while ant composition in other agricultural habitats was most similar to that of urban areas. Although occurrences of Wasmannia auropunctata were similar in cocoa plantations and forests, abundance of Thaumatomyrmex and Typhlomyrmex, generally thought to be rare ants, was relatively high in cocoa plantations. These results, from cocoa plantations with relatively simple shade, demonstrate the importance of cocoa for ant conservation in the Atlantic forest region of Brazil. It is likely that cocoa plantations with a greater number of vegetation strata and higher tree species richness (such as traditional cabruca plantations) provide even more important habitat for ants generally and for ant species of conservation concern.     

Influence of season on leaf litter ant (Hymenoptera: Formicidae) diversity in the locality of Minko'o (Southern,Cameroon)

Seasonality is known to influence ant activity in many tropical rain forests in the world such as South America and Africa. We surveyed ant fauna in the leaf litter in the locality of Minko'o. The work aimed to evaluate the effect of seasonal variation on the diversity and composition of litter ants. Ants were sampled from November 2015 to June 2017, using four sampling methods: visual capture, bait, pitfall trap and extraction. Species richness, Shannon diversity index and analysis of similarities were used to characterise diversity of ant communities between seasons. We collected 306 ant species, shared out between 56 genera and 11 subfamilies. Subfamilies Myrmicinae, Ponerinae, Dolichoderinae, Formicinae, Dorylinae, Cerapachyinae and Pseudomyrmecinae occurred in all the seasons. Species richness was highest in major dry season with 243 species followed by minor rainy season with 188, major rainy season with 177 species and finally minor dry season that recorded the lowest with 155 species. Kruskal–Wallis test showed that ant species richness did not differ between seasons (p > 0.05). Species diversity index indicated that diversity was the highest during minor dry season (H′ = 4.24), followed by the major dry season (H′ = 4.23), minor rainy season (H′ = 4.21) and lowest during major rainy season (H′ = 4.06). Eight most frequents ants have been recorded: Axinidris sp.1, Camponotus flavomarginatus, Monomorium guineense, Myrmicaria opaciventris, Odontomachus troglodytes, Carebara perpusilla, Paltothyreus tarsatus and Pheidole megacephala. Assessment of the seasonal effect on diversity reveals that dry season is richer and more diverse than rainy season and the season significantly influence the diversity of litter ants.     

The influence of nest availability on the abundance and diversity of twig-dwelling ants in a Papua New Guinea forest

Tropical ant communities are frequently diverse, but highly patchy in nature. The availability of suitable nest sites may be a regulating force in structuring litter ant communities. Our aim was to examine ant resource utilization in naturally occurring twigs, and to modify the availability of these resources in order to quantify the influence of nest availability on ant communities in a Papua New Guinean forest. First, we compared ant communities that assemble in artificial twigs (drilled, wooden dowels), naturally occurring twigs, and the leaf litter. A total of 55 ant species were captured: 33 from the leaf litter, 29 from naturally occurring twigs, and only 12 from artificial nests. Significantly different communities formed in each of the three nest types. Second, we examined how the density of natural or artificial nest material influenced the ant abundance and species richness. Plots had between 5 and 96 potential nest sites. An average of only 11.2% of these twigs was colonized. Both species richness and the total abundance of adult ants were significantly positively correlated with increasing naturally occurring twig density. Conversely, increasing the availability of artificial nests from 5 to 20 per plot had no significant effect on the proportion of artificial nests colonized, species richness, or the colony size. We observed that ant species richness and abundance increased with natural twig density, at least for naturally occurring communities. But why so many twigs remain vacant and available for ant colonization remains unknown. Other biotic and abiotic factors likely influence the use of nesting habitat in these ant communities.     

Life in leaf litter: novel insights into community dynamics of bacteria and fungi during litter decomposition

Microorganisms play a crucial role in the biological decomposition of plant litter in terrestrial ecosystems. Due to the permanently changing litter quality during decomposition, studies of both fungi and bacteria at a fine taxonomic resolution are required during the whole process. Here we investigated microbial community succession in decomposing leaf litter of temperate beech forest using pyrotag sequencing of the bacterial 16S and the fungal internal transcribed spacer (ITS) rRNA genes. Our results reveal that both communities underwent rapid changes. Proteobacteria, Actinobacteria and Bacteroidetes dominated over the entire study period, but their taxonomic composition and abundances changed markedly among sampling dates. The fungal community also changed dynamically as decomposition progressed, with ascomycete fungi being increasingly replaced by basidiomycetes. We found a consistent and highly significant correlation between bacterial richness and fungal richness (R = 0.76, P < 0.001) and community structure (R_Mantel = 0.85, P < 0.001), providing evidence of coupled dynamics in the fungal and bacterial communities. A network analysis highlighted nonrandom co‐occurrences among bacterial and fungal taxa as well as a shift in the cross‐kingdom co‐occurrence pattern of their communities from the early to the later stages of decomposition. During this process, macronutrients, micronutrients, C:N ratio and pH were significantly correlated with the fungal and bacterial communities, while bacterial richness positively correlated with three hydrolytic enzymes important for C, N and P acquisition. Overall, we provide evidence that the complex litter decay is the result of a dynamic cross‐kingdom functional succession.     

Response of the ant community to attributes of fragments and vegetation in a northeastern Atlantic Rain Forest area, Brazil

The objective of this study was to determine the effects of forest fragmentation on ant richness in a landscape of Atlantic Forest in Northeast Brazil. More specifically, the ant richness was related to the attributes of fragments (area and distance from the fragment central point to the edge), landscape (forest cover surrounding the fragments), and tree community (plant density, richness, and percentage of shade tolerant species). The surveys were carried out in 19 fragments located in Alagoas State from October 2007 to March 2008. Samples were collected through a 300 m transect established in the center of each fragment, where 30 1-m2 leaf litter samples were collected at 10 m intervals. A total of 146 ant species was collected, which belonged to 42 genera, 24 tribes and nine subfamilies. The attributes of fragments and landscape did not influence ant richness. On the other hand, tree density explained ca. 23% of ant richness. In relation to functional groups, both density and richness of trees explained the richness of general myrmicines (the whole model explained ca. 42% of the variation in this group) and percentage of shade tolerant trees explained the richness of specialist predator ants (30% for the whole model). These results indicate that ant fauna is more influenced by vegetation integrity than by fragment size, distance to edge or forest cover surrounding fragments.     

It is about time: genetic variation in the timing of leaf‐litter inputs influences aquatic ecosystems

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Drought negatively affects communities on a foundation tree: growth rings predict diversity

Understanding how communities respond to extreme climatic events is important for predicting the impact of climate change on biodiversity. The plant vigor and stress hypotheses provide a theoretical framework for understanding how arthropods respond to stress, but are rarely tested at the community level. Following a record drought, we compared the communities of arthropods on pinyon pine (Pinus edulis) that exhibited a gradient in physical traits related to environmental stress (e.g., growth rate, branch dieback, and needle retention). Six patterns emerged that show how one of the predicted outcomes of climate change in the southwestern USA (i.e., increased drought severity) alters the communities of a foundation tree species. In accordance with the plant vigor hypothesis, increasing tree stress was correlated with an eight to tenfold decline in arthropod species richness and abundance. Trees that were more similar in their level of stress had more similar arthropod communities. Both foliage quantity and quality contributed to arthropod community structure. Individual species and feeding groups differed in their responses to plant stress, but most were negatively affected. Arthropod richness (r ² = 0.48) and abundance (r ² = 0.48) on individual trees were positively correlated with the tree’s radial growth during drought. This relationship suggests that tree ring analysis may be used as a predictor of arthropod diversity, which is similar to findings with ectomycorrhizal fungi. A contrast of our findings on arthropod abundance with published data on colonization by mutualistic fungi on the same trees demonstrates that at low stress these two communities respond differently, but at high stress both are negatively affected. These results suggest that the effect of extreme climatic events such as drought on foundation tree species are likely to decrease multi-trophic diversity and shift arthropod community composition, which in turn could cascade to affect other associated taxa.     

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.     

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.     

Group hunting in a ponerine ant,Leptogenys nitida Smith

Field observations on the emigration and foraging behaviours of the southern African ponerine ant, Leptogenys nitida, were undertaken at Mtunzini, Natal, South Africa. These colonies have a single ergatoid queen and 200–1000 workers. The nest sites are found in the leaf litter and these nests are moved frequently over distances ranging from 0.5 to 5 m. Leptogenys nitida is a diurnal predator of arthropods dwelling in the leaf litter. Up to 500 workers participate in each foraging trail, and are not led by definite scouts. Ants form clear trunk trails and fan out at various intervals to search for prey. The prey is searched for and retrieved cooperatively. From laboratory tests it was determined that ants will follow pygidial gland extracts, with the poison gland extract eliciting a limited response. The type of army ant behaviour observed in L. nitida seems to be different to that observed in other ponerine ants.     

Plant palatability to leaf‐cutter ants (Atta laevigata) and litter decomposability in a Neotropical woodland savanna

Although leaf‐cutter ants have been recognized as the dominant herbivore in many Neotropical ecosystems, their role in nutrient cycling remains poorly understood. Here we evaluated the relationship between plant palatability to leaf‐cutter ants and litter decomposability. Our rationale was that if preference and decomposability are related, and if ant consumption changes the abundance of litter with different quality, then ant herbivory could affect litter decomposition by affecting the quality of litter entering the soil. The study was conducted in a woodland savanna (cerrado denso) area in Minas Gerais, Brazil. We compared the decomposition rate of litter produced by trees whose fresh leaves have different degrees of palatability to the leaf‐cutter ant Atta laevigata. Our experiments did not indicate the existence of a significant relationship between leaf palatability to A. laevigata and leaf‐litter decomposability. Although the litter mixture composed of highly palatable plant species showed, initially, a faster decay rate than the mixture of poorly palatable species, this difference was no longer visible after about 6 months. Results were consistent regardless of whether litter invertebrates were excluded or not from litter bags. Similarly, experiments comparing the decomposition rate of litter from pairs of related plant species also showed no association between plant palatability and decomposition. Decomposition rate of the more palatable species was faster, slower or similar to that of the less palatable species depending upon the particular pair of species being compared. We suggest that the traits that mostly influence the decomposition rate of litter produced by cerrado trees may not be the same as those that influence plant palatability to leaf‐cutter ants. Atta laevigata select leaves of different species based – at least in part – on their nitrogen content, but N content was a poor predictor of the decomposition rates of the species we studied.     

The presence of aggressive ants is associated with fewer insect visits to and altered microbe communities in coffee flowers

The process of dispersal can shape ecological communities, but its influence is thought to be small compared to the effects of environmental variation or direct species interactions, particularly for microbial communities. Ants can influence movement patterns of insects and the microbes they vector, potentially affecting microbial establishment on plants, including in agroecosystems. Here, we examine how the presence of aggressive ants, which can influence floral visitation by bees and other pollinators, shapes the community composition of bacteria and fungi on coffee flowers in farms that differ in shade management intensity. We hypothesized that the presence of aggressive ants should reduce the frequency and diversity of floral visitors. Finally, we predicted that the effects of ants should be stronger in the low-shade farm, which has a less diverse community of floral visitors. We sampled microbial communities from nectar and pistils of coffee flowers near and far from nests of the aggressive ant Azteca sericeasur across two farms that vary in shade management and diversity of floral visitors. Bacterial and fungal community composition was characterized using Illumina sequencing of the 16S and ITS regions of the rRNA gene. Consistent with our expectation, Azteca presence was associated with a decrease in the number and diversity of visitors, visit duration and number of flowers visited. Azteca presence influenced microbial communities, but effects differed between farms. Azteca nests were associated with higher bacterial diversity in both farms, but the difference between flowers on trees with and without Azteca was greater in the high-shade farm. Azteca nests were associated with higher fungal diversity in the high-shade farm, but not the low-shade farm. In addition, the presence of ants was strongly associated with species composition of fungi and bacteria in flowers, but differentiation between ant and no-ant communities was greater in the low-shade farm. Specific operational taxonomic units (OTUs) were differentially associated with the presence of ants. We conclude that indirect interactions that influence dispersal may have large effects on microbial community composition, particularly in ephemeral microbial communities.     

Leaf-cutting ant preferences for five native tropical plantation tree species growing under different light conditions

To examine the susceptibility of five Costa Rican tree species to leaf-cutter ants (Atta cephalotes L.: Formicidae, Attini), young and mature leaves from trees that were grown in a plantation under full sun and partial shade were offered to six leaf-cutter colonies located in full sun and six in patial shade. In addition to offering leaf disks to the ants, we offered large pieces of leaves to assess the effect of cutting leaves on food choices. Leaf-cutters responded differently to each plant species, preferring Virola koschyni Warburg (Myristicaceae) and Hyeronima alchorneoides Allemao (Euphorbiaceae) over Stryphnodendrum microstachyum Poeppig & Endlicher (Mimosoideae), Pentaclethra macroloba Willdenow (Fabaceae) and Vochysia ferruginea Martius (Vochysiaceae). In agreement with previous studies, interspecific differences among the tree species in water and saponin content appeared to account for the observed ant preferences among the five tree species: leaf-cutter ants preferred leaves and disks with more water and less saponins and the five tree species varied significantly in these traits. An observed positive correlation between preference and phenolic/nitrogen ratio appears to be spurious, and is instead due to a negative correlation between water content and nitrogen content. For the first time, Atta nest location has been shown to affect consumption: nests located in partial shade removed more leaf material than those from the sun plots. However, nest location had no effect on preference ranking of the tree species tested. Pieces of leaves from all the tree species grown in partial shade were significantly more removed than those trees grown in full sun. Leaf age, toughness, leaf specific weights, ant activity, and colony were not correlated with food choices. Because of the observed preferences, the five tree species should not be considered as equal candidates for plantation purposes. However, complete characterization of the candidate status of tree species for plantations in the neotropics must include information on the ability of such species to tolerate pest attacks in addition to their natural defenses to attack.