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.     

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.     

Size matters: nest colonization patterns for twig‐nesting ants

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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.     

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.     

Territory defense by the ant Azteca trigona: maintenance of an arboreal ant mosaic

Mosaics of exclusive foraging territories, produced by intra-and interspecific competition, are commonly reported from arboreal ant communities throughout the tropics and appear to represent a recurring feature of community organization. This paper documents an ant mosaic within mangrove forests of Panama and examines the behavioral mechanisms by which one of the common species, Azteca trigona, maintains its territories. Most of the mangrove canopy is occupied by mutually exclusive territories of the ants A. trigona, A. velox, A. instabilis, and Crematogaster brevispinosa. When foraging workers of A. trigona detect workers of these territorial species, they organize an alarm recruitment response using pheromonal and tactile displays. Nestmates are attracted over short distances by an alarm pheromone originating in the pygidial gland and over longer distances by a trail pheromone produced by the Pavan's gland. Recruits are simultaneously alerted by a tactile display. No evidence was found for chemical marking of the territory. Major workers are proportionally more abundant at territory borders than on foraging trails in the interior of the colony. The mechanisms of territory defense in A. trigona are remarkably similar to those of ecologically analogous ants in the Old World tropics.     

Experiments with artificial nests provide evidence for ant community stratification and nest site limitation in a tropical forest

Ants are dominant in tropical forests and many species nest in hollow cavities. The manner in which species are vertically stratified in these complex habitats is not known, with lack of nest sites being proposed to limit ant populations. Here, we assess ant community stratification and nest site limitation in a lowland rainforest in New Guinea using experimental addition of artificial bamboo nests of two cavity sizes (small: ~12 mm large: ~32 mm diameter) placed at ground level, in the understorey, and in the canopy. We also conducted a pilot experiment to test the utility of nest translocation. Nests were checked for occupancy after 10 weeks and half of the occupied nests were then translocated between forest plots, while keeping same vertical position. Occupancy of small nests was much higher in the understorey and canopy than at ground level (~75% vs. ~25%). Translocation was successful, as a majority of nests was inhabited by the same species before and after translocation and there was no impact of translocation to a different plot compared to the control, except for a reduction in colony size at ground level. Our experiment demonstrates a vertical stratification in community composition of ants nesting in hollow dead cavities and shows that these ants are more nest site limited in the higher strata than at ground level. Use of small artificial cavities has great potential for future experimental studies, especially for those focused on arboreal ants, as occupancy is high and translocation does not negatively affect their colony size. Abstract in Tok Pisin is available with online material.     

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.     

Biotic and abiotic determinants of the formation of ant mosaics in primary Neotropical rainforests

1. Ants are widespread in tropical rainforests, including in the canopy where territorially dominant arboreal species represent the main part of the arthropod biomass. 2. By mapping the territories of dominant arboreal ant species and using a null model analysis and a pairwise approach this study was able to show the presence of an ant mosaic on the upper canopy of a primary Neotropical rainforest (c. 1 ha sampled; 157 tall trees from 28 families). Although Neotropical rainforest canopies are frequently irregular, with tree crowns at different heights breaking the continuity of the territories of dominant ants, the latter are preserved via underground galleries or trails laid on the ground. 3. The distribution of the trees influences the structure of the ant mosaic, something related to the attractiveness of tree taxa for certain arboreal ant species rather than others. 4. Small‐scale natural disturbances, most likely strong winds in the area studied (presence of canopy gaps), play a role by favouring the presence of two ant species typical of secondary formations: Camponotus femoratus and Crematogaster levior, which live in parabiosis (i.e. share territories and nests but lodge in different cavities) and build conspicuous ant gardens. In addition, pioneer Cecropia myrmecophytic trees were recorded.     

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.     

Escape to the high canopy—thermal deficiency causes niche expansion in a forest‐floor ant

The small formicoxenine ant Temnothorax saxonicus was known from about 40 localities in Central Europe nesting in anorganic substrates on floor of xerothermous forests whereas investigations of 198 tree canopies in 19 forest sites of the same region provided no indication for arboreal nesting or foraging. We present the first evidence for canopy‐nesting populations of T. saxonicus on old Quercus trees in 3 sites having maximum calibrated topsoil temperatures of 17.9 ± 0.3 °C which were significantly (P < 0.007) lower than 22.8 ± 2.0 °C measured in 5 sites with ground‐nesting populations. The thermal deficit on forest floor inhibits brood development in ground nests and caused a moving to canopy were maximum calibrated temperatures of the, now wooden, substrates are at least 26.1 °C for the whole canopy and 30.8 °C in more sun‐exposed spots. T. saxonicus competed here successfully with the obligatory canopy ants T. affinis and T. corticalis. The distributional data of this rope‐climbing study support former results that highest nest densities of small arboreal ants occur in temperate climate over the entire canopy mantle of single trees situated in open land or in park‐like environments but occur in the top of the canopy in tree stands with high degree of canopy closure.     

Stage‐dependent responses to emergent habitat heterogeneity: consequences for a predatory insect population in a coffee agroecosystem

Interactions among members of biological communities can create spatial patterns that effectively generate habitat heterogeneity for other members in the community, and this heterogeneity might be crucial for their persistence. For example, stage‐dependent vulnerability of a predatory lady beetle to aggression of the ant, Azteca instabilis, creates two habitat types that are utilized differently by the immature and adult life stages of the beetle. Due to a mutualistic association between A. instabilis and the hemipteran Coccus viridis – which is A. orbigera main prey in the area – only plants around ant nests have high C. viridis populations. Here, we report on a series of surveys at three different scales aimed at detecting how the presence and clustered distribution of ant nests affect the distribution of the different life stages of this predatory lady beetle in a coffee farm in Chiapas, Mexico. Both beetle adults and larvae were more abundant in areas with ant nests, but adults were restricted to the peripheries of highest ant activity and outside the reach of coffee bushes containing the highest densities of lady beetle larvae. The abundance of adult beetles located around trees with ants increased with the size of the ant nest clusters but the relationship is not significant for larvae. Thus, we suggest that A. orbigera undergoes an ontogenetic niche shift, not through shifting prey species, but through stage‐specific vulnerability differences against a competitor that renders areas of abundant prey populations inaccessible for adults but not for larvae. Together with evidence presented elsewhere, this study shows how an important predator is not only dependent on the existence of two qualitatively distinct habitat types, but also on the spatial distribution of these habitats. We suggest that this dependency arises due to the different responses that the predator's life stages have to this emergent spatial pattern.     

Predaceous ants, beach replenishment, and nest placement by sea turtles

Ants known for attacking and killing hatchling birds and reptiles include the red imported fire ant (Solenopsis invicta Buren), tropical fire ant [Solenopsis geminata (Fabr.)], and little fire ant [Wasmannia auropunctata (Roger)]. We tested whether sea turtle nest placement influenced exposure to predaceous ants. In 2000 and 2001, we surveyed ants along a Florida beach where green turtles (Chelonia mydas L.), leatherbacks (Dermochelys coriacea Vandelli), and loggerheads (Caretta caretta L.) nest. Part of the beach was artificially replenished between our two surveys. As a result, mean beach width experienced by nesting turtles differed greatly between the two nesting seasons. We surveyed 1,548 sea turtle nests (2000: 909 nests; 2001: 639 nests) and found 22 ant species. S. invicta was by far the most common species (on 431 nests); S. geminata and W. auropunctata were uncommon (on 3 and 16 nests, respectively). In 2000, 62.5% of nests had ants present (35.9% with S. invicta), but in 2001, only 30.5% of the nests had ants present (16.4% with S. invicta). Turtle nests closer to dune vegetation had significantly greater exposure to ants. Differences in ant presence on turtle nests between years and among turtle species were closely related to differences in nest placement relative to dune vegetation. Beach replenishment significantly lowered exposure of nests to ants because on the wider beaches turtles nested farther from the dune vegetation. Selective pressures on nesting sea turtles are altered both by the presence of predaceous ants and the practice of beach replenishment.     

Impacts of Argentine ants on avian nesting success

Biological invasions can have severe and widespread impacts on ecological communities. A few species of ants have become particularly damaging invaders but quantitative data of their impacts on many taxa is still lacking. We provide experimental evidence using artificial nests baited with quail eggs that the invasive Argentine ant (Linepithema humile) can be a significant avian nest predator – Argentine ants recruited to more nests and in higher abundance than the native ant species they displace. However, at a site invaded by Argentine ants, we monitored over 400 nests of a ground-nesting species, the Dark-eyed Junco (Junco hyemalis), and found that less than 2% of nests failed as a result of Argentine ant predation/infestation. A review of the literature also suggests that Argentine ants may not be a serious threat to bird nests relative to other predators or parasites. However, invasive ants with the capability of overwhelming prey though stinging (specifically the red-imported fire ant, Solenopsis invicta), may have a higher impact on avian nesting success. Received 14 January 2005; revised 28 April 2005; accepted 12 May 2005.     

Effects of canopy connectivity on the arboreal ant community in coffee shade trees

Canopy connectivity influences foraging, movement, and competition in arboreal ant communities. Understanding how canopy connectivity affects arboreal ant communities could inform the development of management practices that maximize services from known biocontrol agents. We experimentally manipulated connectivity between the crowns of large shade trees to investigate the effects of canopy connectivity on arboreal ant species richness and composition in a coffee agroecosystem. A linear mixed-effects analysis showed that the number of species observed at baits set in tree crowns increased significantly after the crowns had been connected with nylon ropes. Crowns that were connected increased in similarity of ant species composition, particularly between adjacent connected crowns. Connectivity may increase the number of species present in tree crowns by allowing ants to move and forage in the canopy while bypassing trunks with more aggressive, territorial species such as Azteca sericeasur. Because twig-nesting species in the upper canopy have been shown to act as biocontrol agents of herbivores, an increase in species richness in tree crowns could have positive implications for agricultural pest-control services.     

Temporal and spatial patterns of foraging in two syntopic ant species,Manica yessensis and Formica lemani,on Mount Fuji,Japan

Spatiotemporal characteristics of terrestrial foraging were studied in two ant species, Manica yessensis and Formica lemani, in a volcanic desert on the southeast slope of Mount Fuji, Gotenba, Japan. Both ants are common in this habitat, and they construct underground nests in this dry area with sparse vegetation. Nests of M. yessensis have multiple nest-openings on the surface, whereas nests of F. lemani have very few openings, but their nesting and foraging areas overlap completely. A “mark-and-observe” method applied to M. yessensis demonstrated that worker ants of this species move between openings more than 3 m away. A study plot (6 m × 12 m quadrat) was set up, in which all nest-openings of both species were mapped. Day-long observations on numbers of foragers in this plot revealed that foraging M. yessensis are active in morning and evening, while F. lemani continues foraging all day, but both species cease activity at night. Associations between locations of foragers and nest-openings differed significantly between the two species, that is, surface foraging of M. yessensis workers was largely confined to the vicinity of their nest-openings, whereas foragers of F. lemani travelled far from their nest-openings. The function of multiple nest-openings in M. yessensis is discussed.     

Nesting preferences of the threatened wood ant <Emphasis Type="Italic">Formica exsecta</Emphasis> (Hymenoptera: Formicidae); implications for conservation in Scotland

Wood ants are a dominant and ecologically important component of northern coniferous forests with interactions at many trophic levels. Each species exhibits specific habitat preferences which need to be understood if conservation measures are to be successful. In Britain, the rare narrow-headed ant Formica exsecta has disappeared from much of its former range and is now largely restricted to the highlands of Scotland where it is found in open canopy woodland and along forest edges. Nest locale of one small and vulnerable population at the edge of its current range, were compared with those of random locations within the same habitat and with actual nests within the stronghold of the Abernethy-Glenmore complex in Strathspey. After data exploration with Redundancy Analysis, stepwise multiple regression was used to create a model which best estimated the variance in nest location using a parsimonious selection of vegetation and environmental variables. The input variables included were light, soil moisture, altitude, tree stature and distribution, vegetation structure and composition, and ground characteristics. F. exsecta clearly exhibited preferences for the position of nest mounds in relation to light, vegetation and tree cover. Forest location was also important in determining which variables nests were affected by. This study highlights the importance of maintaining a dynamic mosaic of different-aged woodland enabling early successional species such as F. exsecta, with suitable areas to move to as conditions change and allowing the co-existence of all wood ant species.     

A native ant armed to limit the spread of the Argentine ant

Argentine ants (Linepithema humile) usually actively displace native ants through a combination of rapid recruitment, high numerical dominance and intense aggressive fights. However, in some cases, native ants can offer a strong resistance. In Corsica, a French Mediterranean island, local resistance by the dominant Tapinoma nigerrimum has been proposed as a factor limiting Argentine ant invasion. With the aim of evaluating the abilities of T. nigerrimum in interference and exploitative competition, this study tested in the laboratory the aggressive interactions between this native dominant ant and the invasive Argentine ant. We used four different assays between L. humile and T. nigerrimum: (1) worker dyadic interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a competition for space and food. This study confirms the ability of Argentine ants to compete with native species, by initiating more fights, using cooperation and simultaneously deploying physical and chemical defenses. However, despite Argentine ant fighting capabilities, T. nigerrimum was more efficient in both interference and exploitative competition. Its superiority was obvious in the space and food competition assays, where T. nigerrimum dominated food in 100% of the replicates after 1 h and invaded Argentine ant nests while the reverse was never observed. The death feigning behavior exhibited by Argentine ant workers also suggests the native ant’s superiority. Our study thus demonstrates that T. nigerrimum can offer strong competition and so may be able to limit the spread of Argentine ants in Corsica. This confirms that interspecific competition from ecologically dominant native species can affect the invasion success of invaders, notably by decreasing the likelihood of incipient colony establishment and survival.     

Competitive dominance in the organisation of Mediterranean ant communities

1. Competition by dominant species is thought to be key to structuring ant communities. However, recent findings suggest that the effect of dominant species on communities is less pronounced than previously assumed. 2. The aim of the present study was to identify the role of dominant ants in the organisation of Mediterranean communities, particularly the role of competition in invaded and uninvaded communities. The effects on ant assemblages of two dominant ants, the invasive Argentine ant and the native ant, Tapinoma nigerrimum Nylander, were assessed. 3. The abundances of both dominant ants were significantly correlated with a decrease in native ant richness at traps. However, only the invasive ant was associated with a reduction in diversity and abundance of other ant species at site scale. In the presence of T. nigerrimum, species co‐occurrence patterns were segregated or random. Community structure in both the dominant‐free and the Argentine ant sites showed random patterns of species co‐occurrence. 4. The present findings indicate that dominant ants regulate small‐scale diversity by competition. However, at the broader scale of the assemblage, T. nigerrimum may only affect species distribution, having no apparent effect on community composition. Moreover, we find no evidence that inter‐specific competition shapes species distribution in coastal Mediterranean communities free of dominant ants. 5. These results show that dominant species may affect ant assemblages but that the nature and the intensity of such effects are species and scale dependent. This confirms the hypothesis that competitive dominance may be only one of a range of factors that structure ant communities.     

Aggressive interactions between the introduced Argentine ant, Linepithema humile and the native odorous house ant, Tapinoma sessile

The Argentine ant (Linepithema humile) is an invasive species that disrupts the balance of natural ecosystems by displacing indigenous ant species throughout its introduced range. The mechanisms by which Argentine ants effectively compete against native ant species have been previously addressed in field studies that centered on interference and exploitation competition at baits and mainly examined the colony-level performance of Argentine ants. Detailed behavioral observations explaining the basis for the strong competitive ability of L. humile are comparatively rare. To gain a better understanding of the mechanisms by which Argentine ants displace native ants we examined the aggressive interactions between the Argentine ants and the odorous house ant, Tapinoma sessile in four different aggression assays: (1) worker dyad interactions, (2) symmetrical group interactions, (3) intruder introductions into an established resident colony, and (4) a resource competition assay which focused on competition for food and nesting space. Our results demonstrate a clear disparity between worker-level and colony-level fighting ability of Argentine ants and provide behavioral evidence to explain the superior interference ability of Argentine ants in group assays. Argentine ants experienced mixed success in fighting against odorous house ants in dyad interactions, but gradually gained a numerical advantage in symmetrical group interactions by active cooperation among nestmates. Results of the resource competition assay indicate that Argentine ants recruit rapidly, numerically dominate food and nesting sites, and aggressively displace T. sessile from baits. Taken together, the results of these assays allow us to pinpoint the behavioral mechanisms responsible for the remarkable competitive ability of Argentine ants.