Short-Term Temporal Patterns in Herbivore Beetle Assemblages in Polyculture Neotropical Forest Plantations.

Although insect herbivorous communities in tropical forests are known to exhibit strong seasonality, few studies have systematically assessed temporal patterns of variation in community structure and plant-herbivore interactions in early successional arboreal communities. We assessed seasonal and interannual variation of the diversity and composition of herbivorous beetles and the tree-herbivore network in a recently established polyculture forest plantation, during the dry and the rainy seasons of 2012 and of 2013. Species richness was similar between years, while the ecological diversity was higher in 2012. Comparing seasons, no differences were found in 2012, whereas in 2013, the species richness and ecological diversity were higher during the dry season. The species composition differed radically across years and seasons. Moreover, a quantitative nested pattern was consistently found across both temporal scales, more influenced by species densities. We found temporal changes in the species strength, whereas connectance and interaction evenness remained stable. Rapid temporal changes in the structural complexity of recently established polyculture plantations and the availability and quality of the trophic resources they offer may act as drivers of beetle diversity patterns, promoting rapid variation in herbivore composition and some interacting attributes. Nonetheless, network structure, connectance, and interaction evenness remained similar, suggesting that reorganizations in the distribution of species may determine the maintenance of the patterns of interaction. Further work assessing long-term temporal dynamics of herbivore beetle assemblages are needed to more robustly relate diversity and interaction patterns to biotic and abiotic factors and their implications in management programs.

Relative contribution of ecological and biological attributes in the fine-grain structure of ant-plant networks.

BACKGROUND: Ecological communities of interacting species analyzed as complex networks have shown that species dependence on their counterparts is more complex than expected at random. As for other potentially mutualistic interactions, ant-plant networks mediated by extrafloral nectar show a nested (asymmetric) structure with a core of generalist species dominating the interaction pattern. Proposed factors structuring ecological networks include encounter probability (e.g., species abundances and habitat heterogeneity), behavior, phylogeny, and body size. While the importance of underlying factors that influence the structure of ant-plant networks have been separately explored, the simultaneous contribution of several biological and ecological attributes inherent to the species, guild or habitat level has not been addressed. METHODS: For a tropical seasonal site we recorded (in 48 censuses) the frequency of pairwise ant-plant interactions mediated by extrafloral nectaries (EFN) on different habitats and studied the resultant network structure. We addressed for the first time the role of mechanistic versus neutral determinants at the 'fine-grain' structure (pairwise interactions) of ant-plant networks. We explore the simultaneous contribution of several attributes of plant and ant species (i.e., EFN abundance and distribution, ant head length, behavioral dominance and invasive status), and habitat attributes (i.e., vegetation structure) in prevailing interactions as well as in overall network topology (community). RESULTS: Our studied network was highly-nested and non-modular, with core species having high species strengths (higher strength values for ants than plants) and low specialization. Plants had higher dependences on ants than vice versa. We found that habitat heterogeneity in vegetation structure (open vs. shaded habitats) was the main factor explaining network and fine-grain structure, with no evidence of neutral (abundance) effects. DISCUSSION: Core ant species are relevant to most plants species at the network showing adaptations to nectar consumption and deterrent behavior. Thus larger ants interact with more plant species which, together with higher dependence of plants on ants, suggests potential biotic defense at a community scale. In our study site, heterogeneity in the ant-plant interactions among habitats is so prevalent that it emerges at community-level structural properties. High frequency of morphologically diverse and temporarily-active EFNs in all habitats suggests the relevance and seasonality of plant biotic defense provided by ants. The robust survey of ecological interactions and their biological/ecological correlates that we addressed provides insight of the interplay between adaptive-value traits and neutral effects in ecological networks.

Unravelling Darwin's entangled bank: architecture and robustness of mutualistic networks with multiple interaction types.

Trying to unravel Darwin's entangled bank further, we describe the architecture of a network involving multiple forms of mutualism (pollination by animals, seed dispersal by birds and plant protection by ants) and evaluate whether this multi-network shows evidence of a structure that promotes robustness. We found that species differed strongly in their contributions to the organization of the multi-interaction network, and that only a few species contributed to the structuring of these patterns. Moreover, we observed that the multi-interaction networks did not enhance community robustness compared with each of the three independent mutualistic networks when analysed across a range of simulated scenarios of species extinction. By simulating the removal of highly interacting species, we observed that, overall, these species enhance network nestedness and robustness, but decrease modularity. We discuss how the organization of interlinked mutualistic networks may be essential for the maintenance of ecological communities, and therefore the long-term ecological and evolutionary dynamics of interactive, species-rich communities. We suggest that conserving these keystone mutualists and their interactions is crucial to the persistence of species-rich mutualistic assemblages, mainly because they support other species and shape the network organization.

Trait-mediated indirect interactions of ant shape on the attack of caterpillars and fruits.

Mainly owing to their high diversity and abundance, ants are formidable as predators and defenders of foliage. Consequently, ants can exclude both invertebrate and vertebrate activity on plants via direct and indirect interactions as already shown in many previous studies. Here we present empirical evidence that objects resembling ant shape on dummy caterpillars were able to repel visually oriented predators. Moreover, we also show that rubber ants on dummy fruits can repel potential fruit dispersers. Our results have direct implications on the ecological and evolutionary dynamics of interactions in ant-based systems, as ant presence could affect the fitness of its partners. In short, our study highlights the importance of visual cues in interspecific interactions and opens a new way to study the effects of ant presence to test ecological and evolutionary hypotheses.

Influence of plant resistance traits in selectiveness and species strength in a tropical plant-herbivore network.

PREMISE OF THE STUDY: Plant-herbivore networks are highly specialized in their interactions, yet they are highly variable with regard to the relative importance of specific host species for herbivores. How host species traits determine specialization and species strength in this antagonistic network is still an unanswered question that we addressed in this study. METHODS: We assessed plant cover and antiherbivore resistance traits to assess the extent to which they accounted for the variation in specialization and strength of interactions among species in a plant-herbivore network. We studied a tropical antagonistic network including a diverse herbivore-host plant assemblages in different habitat types and climatic seasons, including host plants with different life histories. KEY RESULTS: Particular combinations of leaf toughness, trichome density, and phenolic compounds influenced herbivore specialization and host species strength, but with a significant spatiotemporal variation among plant life histories. Conversely, plant-herbivore network parameters were not influenced by plant cover. CONCLUSIONS: Our study highlights the importance of species-specific resistance traits of plants to understand the ecological and evolutionary consequences of plant-herbivore interaction networks. The novelty of our research lies in the use of a trait-based approach to understand the variation observed in diverse plant-herbivore networks.

Association patterns in saproxylic insect networks in three Iberian Mediterranean woodlands and their resistance to microhabitat loss.

The assessment of the relationship between species diversity, species interactions and environmental characteristics is indispensable for understanding network architecture and ecological distribution in complex networks. Saproxylic insect communities inhabiting tree hollow microhabitats within Mediterranean woodlands are highly dependent on woodland configuration and on microhabitat supply they harbor, so can be studied under the network analysis perspective. We assessed the differences in interacting patterns according to woodland site, and analysed the importance of functional species in modelling network architecture. We then evaluated their implications for saproxylic assemblages' persistence, through simulations of three possible scenarios of loss of tree hollow microhabitat. Tree hollow-saproxylic insect networks per woodland site presented a significant nested pattern. Those woodlands with higher complexity of tree individuals and tree hollow microhabitats also housed higher species/interactions diversity and complexity of saproxylic networks, and exhibited a higher degree of nestedness, suggesting that a higher woodland complexity positively influences saproxylic diversity and interaction complexity, thus determining higher degree of nestedness. Moreover, the number of insects acting as key interconnectors (nodes falling into the core region, using core/periphery tests) was similar among woodland sites, but the species identity varied on each. Such differences in insect core composition among woodland sites suggest the functional role they depict at woodland scale. Tree hollows acting as core corresponded with large tree hollows near the ground and simultaneously housing various breeding microsites, whereas core insects were species mediating relevant ecological interactions within saproxylic communities, e.g. predation, competitive or facilitation interactions. Differences in network patterns and tree hollow characteristics among woodland sites clearly defined different sensitivity to microhabitat loss, and higher saproxylic diversity and woodland complexity showed positive relation with robustness. These results highlight that woodland complexity goes hand in hand with biotic and ecological complexity of saproxylic networks, and together exhibited positive effects on network robustness.

Seasonal dynamics of the flower head infestation of Smallanthus maculatus by two nonfrugivorous tephritids.

Seasonal dynamics of the capitula infested by Dictyotrypeta sp. and Rhynencina spilogaster (Steyskal) (Diptera: Tephritidae) was evaluated throughout the flowering cycle of their host plant the sunflower, Smallanthus maculatus (Cavanilles) Robinson (Asterales: Asteraceae). In central Veracruz, Mexico, along 16 consecutive weeks, a total of 1,017 mature capitula were collected, recording the presence and abundance of immature stages (larvae and pupae) and their related parasitoids. Both fly species were present throughout the entire season, with overall infestation of 51.5% of the capitula examined. However, Dictyotrypeta sp. infested 11.3%, representing about one-fifth of them, and R. spilogaster was most abundant infesting four times as many capitula (42.9%), whereas both species were found together in only 2.6% of the capitula examined. Based on the temporal occurrence of larvae and pupae into flower heads as well as their associated parasitoids and times of emergence, Dictyotrypeta sp. had two yearly generations, and it seems that the second generation could enter a seasonal diapause; in contrast, R. spilogaster was a univoltine species that entered diapause that lasted until the next year.

Evaluating the spatio-temporal factors that structure network parameters of plant-herbivore interactions.

Despite the dynamic nature of ecological interactions, most studies on species networks offer static representations of their structure, constraining our understanding of the ecological mechanisms involved in their spatio-temporal stability. This is the first study to evaluate plant-herbivore interaction networks on a small spatio-temporal scale. Specifically, we simultaneously assessed the effect of host plant availability, habitat complexity and seasonality on the structure of plant-herbivore networks in a coastal tropical ecosystem. Our results revealed that changes in the host plant community resulting from seasonality and habitat structure are reflected not only in the herbivore community, but also in the emergent properties (network parameters) of the plant-herbivore interaction network such as connectance, selectiveness and modularity. Habitat conditions and periods that are most stressful favored the presence of less selective and susceptible herbivore species, resulting in increased connectance within networks. In contrast, the high degree of selectivennes (i.e. interaction specialization) and modularity of the networks under less stressful conditions was promoted by the diversification in resource use by herbivores. By analyzing networks at a small spatio-temporal scale we identified the ecological factors structuring this network such as habitat complexity and seasonality. Our research offers new evidence on the role of abiotic and biotic factors in the variation of the properties of species interaction networks.

Individual-based ant-plant networks: diurnal-nocturnal structure and species-area relationship.

Despite the importance and increasing knowledge of ecological networks, sampling effort and intrapopulation variation has been widely overlooked. Using continuous daily sampling of ants visiting three plant species in the Brazilian Neotropical savanna, we evaluated for the first time the topological structure over 24 h and species-area relationships (based on the number of extrafloral nectaries available) in individual-based ant-plant networks. We observed that diurnal and nocturnal ant-plant networks exhibited the same pattern of interactions: a nested and non-modular pattern and an average level of network specialization. Despite the high similarity in the ants' composition between the two collection periods, ant species found in the central core of highly interacting species totally changed between diurnal and nocturnal sampling for all plant species. In other words, this "night-turnover" suggests that the ecological dynamics of these ant-plant interactions can be temporally partitioned (day and night) at a small spatial scale. Thus, it is possible that in some cases processes shaping mutualistic networks formed by protective ants and plants may be underestimated by diurnal sampling alone. Moreover, we did not observe any effect of the number of extrafloral nectaries on ant richness and their foraging on such plants in any of the studied ant-plant networks. We hypothesize that competitively superior ants could monopolize individual plants and allow the coexistence of only a few other ant species, however, other alternative hypotheses are also discussed. Thus, sampling period and species-area relationship produces basic information that increases our confidence in how individual-based ant-plant networks are structured, and the need to consider nocturnal records in ant-plant network sampling design so as to decrease inappropriate inferences.

Highly nested diets in intrapopulation monkey-resource food webs.

Recently several studies have focused on the structure of ecological networks to provide insights into ecological and coevolutionary dynamics of interacting species. However, rarely have the tools of ecological networks been used to understand how feeding relationships vary among individuals of the same population. Here we use 7 years of data and network analyzed to examine the intrapopulation diet variation in a group of howler monkeys (Alouatta palliata). We show that individual monkey-resource food webs are nested, but not modular and the stability of these patterns is independent of time of day or season. Our findings indicated that individuals do not forage randomly when compared to null models and that the diets of more selective monkeys represent subsets of the diets of other individuals. Moreover, there are no subgroups that eat a particular set of available plant species more frequently than other sets, suggesting that the spatial strategy of group foraging plays an important role in the feeding ecology of each group given that individuals of the same group tend to share similar resources while the group remains at a feeding site. Since the diets of more selective individuals are a subset of other monkeys, we suggest that more selective monkeys are able to outcompete others for preferred foods. Additionally, we did not observe differences in nutritional content or spatial abundance of more frequently eaten plant species when compared with less frequently eaten species, but in most cases, the more frequently eaten plant species were Ficus (Moraceae). This reinforces the important role that Ficus trees play in howler monkey feeding ecology, likely due to its year-round availability.

Long-term temporal variation in the organization of an ant-plant network.

BACKGROUND AND AIMS: Functional groups of species interact and coevolve in space and time, forming complex networks of interacting species. A long-term study of temporal variation of an ant-plant network is presented with the aims of: (1) depicting its structural changes over a 20-year period; (2) detailing temporal variation in network topology, as revealed by nestedness and modularity analysis and other parameters (i.e. connectance, niche overlap); and (3) identifying long-term turnover in taxonomic structure (i.e. switches in ant resource use or plant visitor assemblages according to taxa). METHODS: Fieldwork was carried out at La Mancha, Mexico, and ant-plant interactions were observed between 1989 and 1991, between 1998 and 2000, and between May 2010 and 2011. Occurrences of ants on extrafloral nectaries (EFNs) were recorded. The resulting ant-plant networks were constructed from qualitative presence-absence data determined by a species-species matrix defined by the frequency of occurrence of each pairwise ant-plant interaction. KEY RESULTS: Network variation across time was stable and a persistent nested structure may have contributed to the maintenance of resilient and species-rich communities. Modularity was lower than expected, especially in the most recent networks, indicating that the community exhibited high overlap among interacting species (e.g. few species were hubs in the more recent network, being partly responsible for the nested pattern). Structurally, the connections created among modules by super-generalists gave cohesion to subsets of species that otherwise would remain unconnected. This may have allowed an increasing cascade-effect of evolutionary events among modules. Mutualistic ant-plant interactions were structured 20 years ago mainly by the subdominant nectarivorous ant species Camponotus planatus and Crematogaster brevispinosa, which monopolized the best extrafloral nectar resources and out-competed other species with broader feeding habits. Through time, these ants, which are still present, lost their position as network hubs and diminished in their importance in structuring the network; simultaneously, plants gained in importance. CONCLUSIONS: The long-term network analysis reveals a decrease in attended plant species richness, a notable increase in plant species participation from 1990 to 2010 (sustained by less plant taxonomic similarity in the older 1990 network), an increase in the number of ant species and a diminishing dominance of super-generalist ants. The structure of the community has remained highly nested and connected with low modularity, suggesting overall a more participative, homogeneous, cohesive interaction network. Although previous studies have suggested that interactions between ants and EFN-bearing plants are susceptible to seasonality, abiotic factors and perturbation, this cohesive structure appears to be the key for biodiversity and community maintenance.

Nectar secretion on fern fronds associated with lower levels of herbivore damage: field experiments with a widespread epiphyte of Mexican cloud forest remnants.

BACKGROUND AND AIMS: The oldest group of plants in which nectar secretions have been observed are the Polypodiopsida (ferns sensu lato). Nectaries have been reported in a dozen extant genera. The function of these nectaries has been investigated in several fern species, and in some circumstances has been demonstrated to have an antiherbivore role, attracting and maintaining biotic defence (ants and/or other predatory arthropods). This study documents foliar nectaries in Pleopeltis crassinervata, a widespread Central American epiphyte growing on a variety of trees in cloud forest areas of Veracruz, Mexico. This is a new record for this genus and species. METHODS: As previous experimental work on epiphytic species of Polypodium has demonstrated a protective role of ants for developing fronds, we conducted similar experiments (using nylon nail polish to cover nectaries rather than excluding ants with bands of sticky resin as in earlier work). The fronds of Pl. crassinervata developed over 6 weeks, at which time damage was assessed. The experiment was simultaneously conducted on a sympatric species lacking nectaries, Polypodium furfuraceum. Herbivore placement experiments were conducted with large and small caterpillars on both of these ferns. KEY RESULTS: Fronds with nectaries covered suffered greater damage from herbivores over the course of their development, compared with fronds that had uncovered nectaries functioning normally. The parallel experiment on Po. furfuraceum showed no difference between manipulated and control fronds. Six species of ants (Brachymyrmex minutus, Crematogaster formosa, Paratrechina longicornis, Solenopsis geminata, S. picea and Wasmannia auropunctata) were observed visiting nectaries of Pl. crassinervata; most were effective in removing herbivore larvae placed on the fronds. CONCLUSIONS: The long evolutionary history of ferns may explain why some previous studies of fern nectaries have shown little or no benefit to ferns from nectary visitors, as any coevolved herbivores are those resistant to ant defence. The results suggest that ants protect Pl. crassinervata fronds against herbivory. The presence of nectaries, and the relationship with ants, may contribute to this fern's widespread occurrence and persistence in the face of disturbance, though many other factors also play a role. Ant defence may be more likely to benefit a widespread species of disturbed habitats that encounters a wide range of non-adapted herbivores.

Host use and resource sharing by fruit/seed-infesting insects on Schoepfia schreberi (Olacaceae).

The interactions between the fruit fly Anastrepha spatulata Stone (Diptera: Tephritidae) and two species of moths, the gelechiid Coleotechnites sp. and an unidentified tortricid species, were examined on their effects on seed production in terms of their exploitation within fruits of Schoepfia schreberi J.F. Gmel (Santalales: Olacaceae). The study was carried out in three experimental sites during 3 yr. Under conditions of abundant fruit, A. spatulata was the dominant exploiter at the population level, as shown by its ability to infest the largest number of fruits of the three herbivores and substantially displace its moth competitors. In a separate experiment, when resource-partitioning species were excluded, A. spatulata infested twice as many fruits as the two moth species (44.3%). Field observations examined the ability of A. spatulata to locate suitable fruits. We found that, the variation in fruit characters influenced fruit suitability in relation to its size (4.1261 ± 0.0272 mm [mean ± SE]) and weight (0.0618 ± 0.0005 mg [mean ± SE]). Uninfested fruits produce viable seed but the interaction of all species (exploiting for the fruits) led to limited seed formation.

Breaking down complex Saproxylic communities: understanding sub-networks structure and implications to network robustness.

Saproxylic insect communities inhabiting tree hollow microhabitats correspond with large food webs which simultaneously are constituted by multiple types of plant-animal and animal-animal interactions, according to the use of trophic resources (wood- and insect-dependent sub-networks), or to trophic habits or interaction types (xylophagous, saprophagous, xylomycetophagous, predators and commensals). We quantitatively assessed which properties of specialised networks were present in a complex networks involving different interacting types such as saproxylic community, and how they can be organised in trophic food webs. The architecture, interacting patterns and food web composition were evaluated along sub-networks, analysing their implications to network robustness from random and directed extinction simulations. A structure of large and cohesive modules with weakly connected nodes was observed throughout saproxylic sub-networks, composing the main food webs constituting this community. Insect-dependent sub-networks were more modular than wood-dependent sub-networks. Wood-dependent sub-networks presented higher species degree, connectance, links, linkage density, interaction strength, and were less specialised and more aggregated than insect-dependent sub-networks. These attributes defined high network robustness in wood-dependent sub-networks. Finally, our results emphasise the relevance of modularity, differences among interacting types and interrelations among them in modelling the structure of saproxylic communities and in determining their stability.

Changes of a mutualistic network over time: reanalysis over a 10-year period.

We analyzed the structure of a multispecific network of interacting ants and plants bearing extrafloral nectaries recorded in 1990 and again in 2000 in La Mancha, Veracruz, Mexico. We assessed the replicability of the number of interactions found among species and also whether there had been changes in the network structure associated with appearance of new ant and plant species during that 10-year period. Our results show that the nested topology of the network was similar between sampling dates, group dissimilarity increased, mean number of interactions for ant species increased, the frequency distribution of standardized degrees reached higher values for plant species, more ant species and fewer plant species constituted the core of the more recent network, and the presence of new ant and plant species increased while their contribution to nestedness remained the same. Generalist species (i.e., those with the most links or interactions) appeared to maintain the stability of the network because the new species incorporated into the communities were linked to this core of generalists. Camponotus planatus was the most extreme generalist ant species (the one with the most links) in both networks, followed by four other ant species; but other species changed either their position along the continuum of generalists relative to specialists or their presence or absence within the network. Even though new species moved into the area during the decade between the surveys, the overall network structure remained unmodified.

Noninvasive collection of fresh hairs from free-ranging howler monkeys for DNA extraction.

The use of noninvasive collected samples as source of DNA in studies of wild primate populations has increased in recent years. Fresh-plucked hairs represent an important source of DNA, with relatively high quality and concentration. In this study, we describe a low-cost noninvasive technique for collecting fresh-plucked hairs used to obtain DNA samples from free-ranging black howler monkey populations (Alouatta pigra). We designed and manufactured darts made of wooden dowels, with the anterior part smeared with glue, which were projected with blowpipes to trap howler monkey hairs. All of the materials to make the darts are inexpensive and are available locally. We collected 89 samples from 76 individuals residing in 15 troops, and the total number of hairs obtained was 754. We found no differences in the number of hairs collected among sex-age classes or among localities but the percentage of darts recovered with sample varied among localities. Preliminary results indicate that over 96% of samples yielded DNA suitable for polymerase chain reaction-based microsatellite marker analysis. The technique proved successful for collecting fresh-plucked hairs of free-ranging black howler monkeys without any trauma to the animals and can be easily adapted to obtain samples from other wild primate and mammal species.

Predicted and verified distributions of Ateles geoffroyi and Alouatta palliata in Oaxaca, Mexico.

Primate conservation requires a better knowledge of the distributions and statuses of populations in both large areas of habitat and in areas for which we currently have no information. We focused on spider monkeys (Ateles geoffroyi) and howler monkeys (Alouatta palliata) in the state of Oaxaca, Mexico. This Mexican state has protected large tracts of forest, and has historical records for both primates, although very little is known about them. To update our knowledge of the distributions of these primates and identify potential areas in which they are present, we modeled their geographic distributions by characterizing their ecological niches using the genetic algorithm for rule-set production (GARP), performed interviews and carried out field surveys. The predicted distributions, surveys and interviews indicate that the distributions of these primates are restricted to northeastern Oaxaca. The results suggest that spider monkeys occupy a wider area and elevational range than howler monkeys. Throughout that range there is a wide variety of suitable habitats for these primates. Most of the sites where monkeys were recorded in the field are not officially protected and there was evidence of hunting and habitat destruction. It is important to improve protection, economic alternatives and environmental education as we move towards an integral solution for the conservation of these species. Validation of the GARP model was done for A. geoffroyi, since we had obtained enough field data for this species; this validation indicated that the predicted distribution of the species was statistically better than expected by chance. Hence, ecological niche modeling is a useful approach when performing an initial assessment to identify distribution patterns, detecting suitable areas for future exploration, and for conservation planning. Our findings provide an improved basis for primate conservation and productive fieldwork in Oaxaca.

Interaction intimacy affects structure and coevolutionary dynamics in mutualistic networks.

The structure of mutualistic networks provides clues to processes shaping biodiversity [1-10]. Among them, interaction intimacy, the degree of biological association between partners, leads to differences in specialization patterns [4, 11] and might affect network organization [12]. Here, we investigated potential consequences of interaction intimacy for the structure and coevolution of mutualistic networks. From observed processes of selection on mutualistic interactions, it is expected that symbiotic interactions (high-interaction intimacy) will form species-poor networks characterized by compartmentalization [12, 13], whereas nonsymbiotic interactions (low intimacy) will lead to species-rich, nested networks in which there is a core of generalists and specialists often interact with generalists [3, 5, 7, 12, 14]. We demonstrated an association between interaction intimacy and structure in 19 ant-plant mutualistic networks. Through numerical simulations, we found that network structure of different forms of mutualism affects evolutionary change in distinct ways. Change in one species affects primarily one mutualistic partner in symbiotic interactions but might affect multiple partners in nonsymbiotic interactions. We hypothesize that coevolution in symbiotic interactions is characterized by frequent reciprocal changes between few partners, but coevolution in nonsymbiotic networks might show rare bursts of changes in which many species respond to evolutionary changes in a single species.

Asymmetries in specialization in ant-plant mutualistic networks.

Mutualistic networks involving plants and their pollinators or frugivores have been shown recently to exhibit a particular asymmetrical organization of interactions among species called nestedness: a core of reciprocal generalists accompanied by specialist species that interact almost exclusively with generalists. This structure contrasts with compartmentalized assemblage structures that have been verified in antagonistic food webs. Here we evaluated whether nestedness is a property of another type of mutualism-the interactions between ants and extrafloral nectary-bearing plants--and whether species richness may lead to differences in degree of nestedness among biological communities. We investigated network structure in four communities in Mexico. Nested patterns in ant-plant networks were very similar to those previously reported for pollination and frugivore systems, indicating that this form of asymmetry in specialization is a common feature of mutualisms between free-living species, but not always present in species-poor systems. Other ecological factors also appeared to contribute to the nested asymmetry in specialization, because some assemblages showed more extreme asymmetry than others even when species richness was held constant. Our results support a promising approach for the development of multispecies coevolutionary theory, leading to the idea that specialization may coevolve in different but simple ways in antagonistic and mutualistic assemblages.