Extrafloral‐nectaries and interspecific aggressiveness regulate day/night turnover of ant species foraging for nectar on Bionia coriacea

Plants bearing extrafloral nectaries (EFNs) vary the secretion of nectar between day and night, which creates turnover in the composition of interacting ant species. Daily variation in the composition of ant species foraging on vegetation is commonly observed, but its mechanisms are poorly understood. We evaluated the daily variation in nectar availability and interspecific aggressiveness between ants as possible regulatory mechanisms of the turnover in ant–plant interactions. We hypothesized that (i) plants would interact with more ant species during periods of higher secretion of nectar and that (ii) aggressive ant species would compete for nectar, creating a daily turnover of species collecting nectar. We tested this hypothesis by measuring the production of nectar during the day and night and by experimentally removing EFNs of Bionia coriacea (=Camptosema coriaceum) (Nees & Mart.) Benth. (Fabaceae: Faboideae) plants in a Brazilian savanna (Cerrado). We then compared the abundance and composition of ant species between those treatments and during the day. Our results indicate that more ant workers forage on plants during the day, when nectar was sugary, while more ant species forage at night, when aggressiveness between ant species was lower. We also detected a day/night turnover in ant species composition. Ant species foraging for nectar during the day were not the same at night, and this turnover did not occur on plants without EFNs. Both dominant ant species, diurnal Camponotus crassus (Hymenoptera: Formicidae) and nocturnal Camponotus rufipes (Hymenoptera: Formicidae), were the most aggressive species, attacking other ants in their specific periods of forage while also being very aggressive toward each other. However, this aggressiveness did not occur in the absence of nectar, which allowed non‐aggressive nocturnal ant species to forage only during the daytime, disrupting the turnover. We conclude that extrafloral‐nectar presence and interspecific aggressiveness between ants, along with other environmental factors, are important mechanisms creating turnovers in ants foraging on plants.     

Endemic predators, invasive prey and native diversity

Interactions between native diversity and invasive species can be more complex than is currently understood. Invasive ant species often substantially reduce diversity in the native ants diversity that act as natural control agents for pest insects. In Indonesia (on the island of Sulawesi), the third largest cacao producer worldwide, we show that a predatory endemic toad (Ingerophrynus celebensis) controls invasive ant (Anoplolepis gracilipes) abundance, and positively affects native ant diversity. We call this the invasive-naivety effect (an opposite of enemy release), whereby alien species may not harbour anti-predatory defences against a novel native predator. A positive effect of the toads on native ants may facilitate their predation on insect vectors of cacao diseases. Hence, toads may increase crop yield, but further research is needed on this aspect. Ironically, amphibians are globally the most threatened vertebrate class and are strongly impacted by the conversion of rainforest to cacao plantations in Sulawesi. It is, therefore, crucial to manage cacao plantations to maintain these endemic toads, as they may provide critical ecosystem services, such as invasion resistance and preservation of native insect diversity.     

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.     

Aphid‐tending ants on introduced fennel: can resources derived from non‐native plants alter the trophic position of higher‐order consumers?

1. Although plant invasions often reduce insect abundance and diversity, non‐native plants that support phytophagous insects can subsidise higher trophic levels via elevated herbivore abundance. 2. Here ant–aphid interactions on non‐native fennel on Santa Cruz Island, California are examined. Fennel hosts abundant, honeydew‐producing fennel aphids. The patchiness of fennel and the relative lack of honeydew‐producing insects on other plants at our study sites suggest that assimilation of fennel‐derived honeydew would increase the abundance and decrease the trophic position of the omnivorous, aphid‐tending Argentine ant. 3. To assess the strength of the ant–aphid interaction, a comparison of ant abundance on and adjacent to fennel prior to and 3 weeks after experimental aphid removal was performed. Compared with control plants with aphids, ants declined in abundance on and around fennel plants following aphid removal. At the habitat scale, pitfall traps in fennel‐dominated habitats captured more ants than in fennel‐free scrub habitats. 4. To determine if assimilation of aphid‐produced honeydew reduces the ant's trophic position, variation in δ¹⁵N values among ants, plants and other arthropods was analysed. Unexpectedly, δ¹⁵N values for ants in fennel‐dominated habitats were higher than those of arthropod predators from the same sites and also higher than those of ants from fennel‐free habitats. 5. Our results illustrate how introduced plants that support phytophagous insects appear to transfer energy to higher trophic levels via elevated herbivore abundance. Although assimilation of fennel‐derived honeydew did not appear to reduce consumer trophic position, spatial variation in alternative food resources might obscure contributions from honeydew.     

Intraspecific Food‐Robbing and Neighborhood Competition: Consequences for Anti‐Robber Vigilance and Colony Productivity

Most social animals have mechanisms to distinguish group members from outsiders, in part to prevent the exploitation of resources reserved for members of the group. Nevertheless, specialized thieves of the Neotropical ant, Ectatomma ruidum, also known as the ‘thieving ant’, regularly enter and steal resources from distinct, neighboring colonies. Here, we examine the mechanisms and consequences of thievery in a population of E. ruidum. We show that (1) individuals from nearby colonies were accepted more often than those from farther colonies; (2) rejection rates decreased as individuals interacted more with non‐nestmates from the same source colony; and (3) colonies that were experimentally treated to reduce thievery rates had improved productivity. The boost in productivity with thievery reduction was greater in low density populations than in high density populations. We conclude that, as in other species, thievery has negative fitness costs to E. ruidum. However, greater acceptance of neighbors than non‐neighbors and increased acceptance after habituation to non‐nestmates suggest a proximate explanation for the presence of thievery. Moreover, lower fitness costs of thievery at high nesting density, combined with observations of extraordinarily high densities of E. ruidum throughout its range, suggest there is little selection pressure among these ants to guard against thieves, thus providing an ultimate explanation why thievery persists among litter‐foraging ants.     

Plant killing by Neotropical acacia ants: ecology,decision‐making,and head morphology

Mutualistic species often associate with several partners that vary in the benefits provided. In some protective ant–plant mutualisms, ants vary in the extent at which they kill neighboring vegetation. Particularly, in acacia ants (Pseudomyrmex), the area around the host tree that ants keep free from vegetation (“clearings”) vary depending on the species. This study assessed whether interspecific variation in clearing size corresponds to workers biting on plant tissue of different thickness. As expected, workers from species making the largest clearings bit more often on thicker plant tissues than workers from species making smaller clearings. Because head shape affects mandible force, I also assessed whether pruning on thick tissue in mutualistic ant species or being a predator in non‐mutualistic species correlated with broader heads, which yield stronger mandible force. The species with the broader heads were non‐mutualistic predators or mutualistic pruners of thick tissues, which suggest that pruning neighboring vegetation in non‐predatory species demands force even when the ants do not kill prey with their mandibles. The findings reveal that clearing size variation in mutualistic ant partners of plants can also be observed at the level of individual decision‐making processes among workers, and suggest that head morphology could be a trait under selection in protective ant–plant mutualisms. Abstract in Spanish is available with online material.     

Foraging activity of leaf‐cutting ants changes light availability and plant assemblage in Atlantic forest

1. Leaf‐cutting ants (LCAs) have often been denoted as ecosystem engineers because of their multifarious effects on the vegetation, particularly via nest‐driven environmental changes. However, the non‐trophic impacts of LCAs on forest dynamics via foliage harvesting across sizeable foraging zones (so‐called associated ecosystem engineering) are still poorly investigated. 2. Here, light availability and sapling assemblages were assessed within foraging areas and ant‐free control zones of 16 Atta cephalotes colonies located in a large remnant of Atlantic forest in northeastern Brazil. 3. Canopy openness and total light transmission were 1.4 and 1.6 times higher in foraging zones than in control areas. In parallel, sapling density and species richness decreased constantly from control to foraging zone plots. Additionally, shade‐tolerant species exhibited reduced abundance across foraging zones. A non‐metric multidimensional scaling ordination based on taxonomic similarity primarily segregated foraging zone and control plots; foraging zone plots converged to be more similar to each other as well. Finally, some plant species emerged as indicators of LCA‐free zones. 4. These results suggest that LCA foraging activity in the forest canopy directly increases the light availability and indirectly affects the recruitment and the structure of local plant assemblages. 5. Such a biologically significant effect on the light environment and its cascades confirms LCAs as potent ecosystem engineers, particularly as a plant assembly force, which operates beyond the spatial reach of their well‐described nest effects.     

Coccinellid interactions mediated by vegetation heterogeneity

Environmental heterogeneity can have profound effects on agroecosystem function and it is important for improving ecosystem services such as biological control. Promoting system diversity via non‐crop plants is one method for increasing habitat heterogeneity within farmscapes. Non‐crop plants provide access to refuges and alternative food resources provide multiple benefits to enhance populations of arthropod predators. In this study, we examined the effects of small‐scale spatial structure on life‐stage specific interactions between the native coccinellid, Hippodamia convergensGuérin‐Méneville, and the exotic Harmonia axyridis (Pallas) (both Coleoptera: Coccinellidae), which overlap in spatial distribution in many crop systems. Squash [Cucurbita pepo L. (Cucurbitaceae)] and non‐crop mugwort [Artemisia vulgaris L. (Asteraceae)] plants with and without aphids were used as a model of spatial heterogeneity in micro‐ and mesocosm experiments. In response to factorial treatment combinations, we evaluated oviposition behavior, egg predation, larval survival, and larval predator‐prey and predator‐predator interactions. Adult H. convergens displayed higher foraging activity on aphids when exposed to complex habitats containing a non‐crop plant. In the presence of the exotic coccinellid, H. convergens preferred to deposit eggs on the non‐crop plant. Furthermore, a combination of spatial heterogeneity and prey availability reduced larval intraguild predation and cannibalism, and improved reproductive output of H. convergens by reducing intra‐ and interspecific egg predation. Our results provide evidence that life‐stage‐specific intraguild interactions are mediated by access to non‐crop plants. Thus, the introduction or maintenance of non‐crop plants has the potential to enhance coexistence of multiple natural enemies and improve top‐down control of pests.     

Ant mosaics occur in SE Asian oil palm plantation but not rain forest and are influenced by the presence of nest‐sites and non‐native species

Interaction networks within biotic communities can be dramatically altered by anthropogenic habitat modification. Ants, an important ecological group, often interact competitively to form mosaic‐like patterns in disturbed plantation habitats, in which dominant species form mutually exclusive territories. However, the existence of these ant mosaics in pristine forests is contentious. Here we assess the relative strengths of ant competitive interactions in oil palm plantation and primary rain forest in Sabah, Malaysia, using null models of species co‐occurrence. We use two metrics: the C‐score, which measures mean degree of overall co‐occurrence, and a novel metric, the C_var‐score, which measures the variance in degree of co‐occurrence. We also investigate the role of nest sites by collecting ants from canopy and leaf litter microhabitats, and from epiphytic ferns, an important nest site for canopy ants. Furthermore, we assess whether non‐native species, which were widespread in oil palm plantation (61 occurrences vs five in rain forest) are important in driving the formation of ant mosaics. We found no evidence for ant mosaics in any primary forest microhabitat. In oil palm plantation, segregation between species was pronounced in epiphytes, weak in the rest of the canopy and absent in leaf litter communities. Intriguingly, exclusion of non‐native ant species from analyses increased the degree of negative species co‐occurrence in all three microhabitats, with species segregation in the oil palm canopy becoming statistically significant. Our results suggest that invasion of plantation habitats by non‐native species does not drive increased species segregation in ant communities. Rather, high degrees of species segregation might relate to changes in the importance of canopy nest sites, with colonies competing more strongly for these in plantations. In primary forests, weaker nest‐site limitation and the highly complex, more vertically stratified, non‐uniform canopy could lead to random co‐occurrence between ant species at the scales studied here.     

Biodiversity responses to land‐use and restoration in a global biodiversity hotspot: Ant communities in Brazilian Cerrado

Given that land‐use change is the main cause of global biodiversity decline, there is widespread interest in adopting land‐use practices that maintain high levels of biodiversity, and in restoring degraded land that previously had high biodiversity value. In this study, we use ant taxonomic and functional diversity to examine the effects of different land uses (agriculture, pastoralism, silviculture and conservation) and restoration practices on Cerrado (Brazilian savanna) biodiversity. We also examine the extent to which ant diversity and composition can be explained by vegetation attributes that apply across the full land management spectrum. We surveyed vegetation attributes and ant communities in five replicate plots of each of 13 land‐use and restoration treatments, including two types of native vegetation as reference sites: cerrado sensu stricto and cerradão. Several land‐use and restoration treatments had comparable plot richness to that of the native reference habitats. Ant species and functional composition varied systematically among land‐use treatments following a gradient from open habitats such as agricultural fields to forested sites. Tree basal area and grass cover were the strongest predictors of ant species richness. Losses in ant diversity were higher in land‐use systems that transform vegetation structure. Among productive systems, therefore, uncleared pastures and old pine plantations had similar species composition to that occurring in cerrado sensu stricto. Restoration techniques currently applied to sites that were previously Cerrado have focused on returning tree cover, and have failed to restore ant communities typical of savanna. To improve restoration outcomes for Cerrado biodiversity, greater attention needs to be paid to the re‐establishment and maintenance of the grass layer, which requires frequent fire. At the broader scale, conservation planning in agricultural landscapes, should recognize the value of land‐use mosaics and the risks of homogenization.     

Similar yet different: differential response of a praying mantis to ant‐mimicking spiders

Batesian mimics typically dupe visual predators by resembling noxious or deadly model species. Ants are unpalatable and dangerous to many arthropod taxa, and are popular invertebrate models in mimicry studies. Ant mimicry by spiders, especially jumping spiders, has been studied and researchers have examined whether visual predators can distinguish between the ant model, spider mimic and spider non‐mimics. Tropical habitats harbour a diverse community of ants, their mimics and predators. In one such tripartite mimicry system, we investigated the response of an invertebrate visual predator, the ant‐mimicking praying mantis (Euantissa pulchra), to two related ant‐mimicking spider prey of the genus Myrmarachne, each closely mimicking its model ant species. We found that weaver ants (Oecophylla smaragdina) were much more aggressive than carpenter ants (Camponotus sericeus) towards the mantis. Additionally, mantids exhibited the same aversive response towards ants and their mimics. More importantly, mantids approached carpenter ant‐mimicking spiders significantly more than often that they approached weaver ant‐mimicking spiders. Thus, in this study, we show that an invertebrate predator, the praying mantis, can indeed discriminate between two closely related mimetic prey. The exact mechanism of the discrimination remains to be tested, but it is likely to depend on the level of mimetic accuracy by the spiders and on the aggressiveness of the ant model organism.     

Elevated temperatures alter an ant‐aphid mutualism

Ant‐hemipteran mutualisms are keystone interactions that can be variously affected by warming: these mutualisms can be strengthened or weakened, or the species can transition to new mutualist partners. We examined the effects of elevated temperatures on an ant‐aphid mutualism in the subalpine zone of the Rocky Mountains in Colorado, USA. In this system, inflorescences of the host plant, Ligusticum porteri Coult. & Rose (Apiaceae), are colonized by the ant‐tended aphid Aphis asclepiadis Fitch or less frequently by the non‐ant tended aphid Cavariella aegopodii (Scopoli) (both Hemiptera: Aphididae). Using an 8‐year observational study, we tested for two key mechanisms by which ant‐hemipteran mutualisms may be altered by climate change: shifts in species identity and phenological mismatch. Whereas the aphid species colonizing the host plant is not changing in response to year‐to‐year variation in temperature, we found evidence that a phenological mismatch between ants and aphids could occur. In warmer years, colonization of host plant inflorescences by ants is decreased, whereas for A. asclepiadis aphids, host plant colonization is mostly responsive to date of snowmelt. We also experimentally established A. asclepiadis colonies on replicate host plants at ambient and elevated temperatures. Ant abundance did not differ between aphid colonies at ambient vs. elevated temperatures, but ants were less likely to engage in tending behaviors on aphid colonies at elevated temperatures. Sugar composition of aphid honeydew was also altered by experimental warming. Despite reduced tending by ants, aphid colonies at elevated temperatures had fewer intraguild predators. Altogether, our results suggest that higher temperatures may disrupt this ant‐aphid mutualism through both phenological mismatch and by altering benefits exchanged in the interaction.     

Ant seed predation,pesticide applications and farmers' income from tropical multi‐cropping gardens

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Territorial ants depress plant growth through cascading non‐trophic effects in an alpine meadow

Understanding non‐trophic interactions is critical to mechanistically linking community structure and ecosystem functioning. Despite the widespread occurrence of territoriality across animal taxa and ecosystems, the cascading ecological consequences of non‐trophic interactions between territorial animals and intruders have been poorly studied. We experimentally investigated the non‐trophic interaction between territorial ants and members of a dung decomposer community (i.e. predatory arthropods, maggots and coprophagous beetles) in an alpine meadow. We further examined how this non‐trophic interaction cascaded to influence ecosystem properties including dung removal rate, soil nutrient status and aboveground plant biomass surrounding dung pats. Results indicated that territorial interference of ants on key decomposers cascaded to affect plant growth. Specifically, ants significantly decreased the abundance of coprophagous beetles at the time of their peak‐abundance and hence decreased dung removal rates and soil nitrogen concentrations, ultimately decreasing aboveground plant biomass. The strength of this non‐trophic cascading effect was comparable to those reported in studies addressing trophic cascades triggered by predator–prey interactions. Our findings suggest that the non‐trophic interactions and associated cascading effects stemming from territorial behavior should be incorporated into ecological network modeling and research addressing biodiversity–ecosystem functioning relationships.     

Argentine ants displace floral arthropods in a biodiversity hotspot

Argentine ant (Linepithema humile (Mayr)) invasions are often associated with the displacement of ground‐dwelling arthropods. Argentine ant invasions can also exert other effects on the community through interactions with plants and their associated arthropods. For example, carbohydrate resources (e.g. floral or extrafloral nectar) may influence foraging behaviour and interactions among ants and other arthropods. In South Africa's Cape Floristic Region, Argentine ants and some native ant species are attracted to the floral nectar of Leucospermum conocarpodendron Rourke (Proteaceae), a native tree that also has extrafloral nectaries (EFNs). Despite having relatively low abundance in pitfall traps, Argentine ants visited inflorescences more frequently and in higher abundance than the most frequently observed native ants, Camponotus spp., though neither native nor Argentine ant floral foraging was influenced by the EFNs. Non‐metric multidimensional scaling revealed significant dissimilarity in arthropod communities on inflorescences with Argentine ants compared to inflorescences with native or no ants, with Coleoptera, Diptera, Hymenoptera, Arachnida, Orthoptera, and Blattaria all being underrepresented in inflorescences with Argentine ants compared to ant‐excluded inflorescences. Native honeybees (Apis mellifera capensis Eschscholtz) spent 75% less time foraging on inflorescences with Argentine ants than on inflorescences without ants. Neither Argentine ant nor native ant visits to inflorescences had a detectable effect on seed set of Le. conocarpodendron. However, a pollen supplementation experiment revealed that like many other proteas, Le. conocarpodendron is not pollen‐limited. Flower predation was negatively associated with increased ant visit frequency to the inflorescences, but did not differ among inflorescences visited by native and Argentine ants. Displacement of arthropods appears to be a consistent consequence of Argentine ant invasions. The displacement of floral arthropods by Argentine ants may have far‐reaching consequences for this biodiversity hotspot and other regions that are rich in insect‐pollinated plants.     

Twig-Nesting Ants: The Hidden Predators of the Coffee Berry Borer in Chiapas,Mexico

Coffee is a globally important crop that is subject to numerous pest problems, many of which are partially controlled by predatory ants. Yet several studies have proposed that these ecosystem services may be reduced where agricultural systems are more intensively managed. Here we investigate the predatory ability of twig-nesting ants on the main pest of coffee, the coffee berry borer (Hypothenemus hampei) under different management systems in southwest Chiapas, Mexico. We conducted both laboratory and field experiments to examine which twig-nesting ant species, if any, can prey on free-living borers or can remove borers embedded in coffee fruits and whether the effects of the twig-nesting ant community differ with habitat type. Results indicate that several species of twig-nesting ants are effective predators of both free-living borers and those embedded in coffee fruits. In the lab, Pseudomyrmex ejectus, Pseudomyrmex simplex, and Pseudomyrmex PSW-53 effectively removed free-living and embedded borers. In the field, abundance, but not diversity, of twig-nesting ant colonies was influenced by shade management techniques, with the highest colony abundance present in the sites where shade trees were recently pruned. However, borer removal rates in the field were significant only in the shadiest site, but not in more intensively managed sites. This study provides evidence that twig-nesting ants can act as predators of the coffee berry borer and that the presence of twig-nesting ants may not be strongly linked to shade management intensity, as has been suggested for other arthropod predators of the borer.     

Foraging in highly dynamic environments: leaf‐cutting ants adjust foraging trail networks to pioneer plant availability

Major shifts in the availability of palatable plant resources are of key relevance to the ecology of leaf‐cutting ants in human‐modified landscapes. However, our knowledge is still limited regarding the ability of these ants to adjust their foraging strategy to dynamic environments. Here, we examine a set of forest stand attributes acting as modulating forces for the spatiotemporal architecture of foraging trail networks developed by Atta cephalotes L. (Hymenoptera: Formicidae: Attini). During a 12‐month period, we mapped the foraging systems of 12 colonies located in Atlantic forest patches with differing size, regeneration age, and abundance of pioneer plants, and examined the variation in five trail system attributes (number of trails, branching points, leaf sources, linear foraging distance, and trail complexity) in response to these patch‐related variables. Both the month‐to‐month differences (depicted in annual trail maps) and the steadily accumulating number of trails, trail‐branching points, leaf sources, and linear foraging distance illustrated the dynamic nature of spatial foraging and trail complexity. Most measures of trail architecture correlated positively with the number of pioneer trees across the secondary forest patches, but no effects from patch age and size were observed (except for number of leaf sources). Trail system complexity (measured as fractal dimension; Df index) varied from 1.114 to 1.277 along the 12 months through which ant foraging was monitored, with a marginal trend to increase with the abundance of pioneer stems. Our results suggest that some leaf‐cutting ant species are able to generate highly flexible trail networks (via fine‐tuned adjustment of foraging patterns), allowing them to profit from the continuous emergence/recruitment of palatable resources.     

Negative effects of ant foraging on spiders in Douglas-fir canopies

 Spiders and ants are potential competitors and mutual predators. Indirect evidence from previous research has suggested that ant foraging may significantly lower the abundance of arboreal spiders in young Douglas-fir plantations in western Oregon. This study tested the effect of foraging by ants, dominated by Camponotus spp., on spider assemblages in Douglas-fir canopies in a 5-month ant-exclusion experiment. The biomass of potential prey organisms on foliage, dominated by Psocoptera, increased significantly by 1.9- to 2.4-fold following ant exclusion. The removal of ants did not affect the abundance of flying arthropods in the vicinity of tree canopies as indicated by sticky trap catches. The abundance of hunting spiders, the majority being Salticidae, increased significantly by 1.5- to 1.8-fold in trees without ants in the late summer; neither the abundance of web-building spiders nor the average body size of hunting and web-building spiders were significantly affected by ant removal. Spider diversity and community structure did not differ significantly between control and ant-removal trees. The majority of prey captured by ants were Aphidoidea (48.1%) and Psocoptera (12.5%); spiders represented only 1.4% of the ants’ diet. About 40% of observed ants were tending Cinara spp. aphids. Our observations suggest that the lower abundance of hunting spiders in control canopies with ants may be due to interference competition with ants resulting from ant foraging and aphid-tending activities. Direct predation of spiders by ants appeared to be of minor importance in this study system. This study did not provide sufficient evidence for exploitative competition for prey between ants and spiders. Received: 21 February 1996 / Accepted: 14 August 1996     

The role of resource dispersion in promoting the co‐occurrence of dominant and subordinate ant species

Dominant competitors govern resource use in many communities, leading to predictions of local exclusion and lower species diversity where dominant species are abundant. However, subordinate and dominant species frequently co‐occur. One mechanism that could facilitate resource sharing and co‐occurrence of dominant and subordinate competitors is fine‐scale resource dispersion. Here, we distributed 6 g of a food resource into 1, 2, 8, 32 or 64 units in small 0.40 m² areas centred on nests of the dominant ant Monomorium sydneyense. We tested three hypotheses. First, we hypothesized that the species richness and abundance of foraging ants would increase with increasing resource dispersion. Accordingly, species richness doubled and total ant abundance was two orders of magnitude higher in high resource dispersion treatments. Secondly, we hypothesized that increasing resource dispersion would reduce competitive interactions such as resource turnover events and lower the probability of food resources being occupied. Substantial support for this hypothesis was observed. Finally, we tested the hypothesis that the foraging time of each species would be proportional to the relative abundance of each species solely in high resource dispersion treatments. Expected and observed foraging times were statistically similar for only the dominant ant M. sydneyense. The subdominant Pheidole rugosula increased its foraging time much more than was expected, while two subordinate ants showed no relationship between observed and expected times. Thus, while increasing resource dispersion significantly increased overall species richness, this increase in co‐occurrence did not correlate with a significant increase in foraging time for the two subordinate species. Rather, changes in resource dispersion appeared to benefit only the subdominant species. Inter‐site variation appeared more important for other subordinate species indetermining co‐occurrence and foraging time. Multiple mechanisms facilitate co‐occurrence and resource sharing in this community, and probably in most other communities.     

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.