A Systematic Review on Opuntia (Cactaceae; Opuntioideae) Flower-Visiting Insects in the World with Emphasis on Mexico: Implications for Biodiversity Conservation.

Opuntia spp. are cacti with high ecological, economic and conservation interest in semiarid environments, particularly in Mexico. We conducted a systematic search of the existing peer-reviewed literature about the state of knowledge of pollination ecology on these plants. We documented the most studied Opuntia species worldwide with an emphasis on Mexico. We found that only 15% of Opuntia species described have been investigated so far, and studies were mainly focused on comprehension of the biology of a single species. Despite the economic and cultural importance of Opuntia, there is a significant lack of knowledge about the flower-visiting insects and their taxonomic identity. We provide a checklist of the insect species associated with Opuntia spp. Through a circular network, we visualize the complex Opuntia flower-visiting insect relationship, and we detected a set of key species constituting the generalist core of the networks constructed. Since pollination is crucial for crop production, a better understanding of ecological interactions would inform management measures to strengthen biodiversity and agriculture sustainability as well as productivity in arid and marginal lands. Further research on pollination ecology is needed to improve the conservation status of the insects associated with Opuntia species.

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.

Diversity Patterns of Tropical Epigeal Beetle Assemblages Associated with Monoculture and Polyculture Plantations with Big-Leaf Mahogany.

Although commercial forest plantations have experienced a major growth in the tropics over the past decades, little attention has been paid to their role in the conservation of epigeal arthropod communities. We studied diversity patterns of the epigeal beetle community in monoculture and polyculture forest plantations with big-leaf mahogany (Swietenia macrophylla). Likewise, we explored the existence of indicator species of each plantation type. Our findings highlight that each plantation type promotes multiple impacts on diversity patterns. We found that monocultures positively influenced overall beetle species richness and ecological diversity. When broken down by guild, both predator and decomposer species richness were similar between monoculture and polyculture, whereas for beetle diversity we found contrasting responses by guild: decomposer diversity was greater in monoculture whereas predator diversity was higher in polyculture. In addition, species composition differed between monoculture and polyculture, except for the predator guild. Species turnover was the main component explaining beta diversity patterns at all levels, indicating that each plantation type promotes biologically distinct epigeal assemblages. Few superabundant heliophile species dominated the beetle community structure; moreover, monocultures had a composition skewed towards heliophile species whereas polyculture favored umbrophile species. These patterns could be attributed to differences in habitat complexity between plot types, namely differences in tree cover. Additionally, indicator species only were identified in polycultures, reflecting their higher spatial complexity. Monoculture and polyculture plantations with big-leaf mahogany are complementary agroecosystems for preserving diverse epigeal beetle communities and should be considered valuable tools for conservation purposes in the tropics.

Diversity and deadwood-based interaction networks of saproxylic beetles in remnants of riparian cloud forest.

We studied the saproxylic beetle community inhabiting deadwood in remnants of riparian cloud forests in "La Antigua" basin, in central Veracruz (Mexico). We assessed the influence of deadwood features (tree species, trunk position, trunk diameter, trunk volume and decomposition stages) on saproxylic beetle diversity. In order to assess the stability of beetle species-deadwood interactions, we also analyzed the ecological networks structure. A total of 63 deadwood trunks, belonging to four tree species, were sampled by standardized hand-collection throughout well-preserved remnants of riparian cloud forest. We found that tree species and deadwood decay stage are the main drivers that determine the diversity and stability of saproxylic beetle species interactions. Our results indicate that Quercus corrugata is the main tree species in terms of maintaining the significantly highest saproxylic beetle diversity, but with no stable interactions (saproxylic beetle-deadwood). A nested network structure was detected for Clethra mexicana and Liquidambar styraciflua, with a pool of core (generalist) saproxylic beetle species. We observed that beetle diversity from the early and late deadwood stages comprises distinct assemblages and the four stages of decomposition showed a nested network structure. During deadwood succession, community composition and guilds changed among networks; the early successional stage had more specialized xylophagous beetles, while other guilds (mycophagous, saprophagous and zoophagous) arrive later and become the core species in the advanced stages of decomposition networks. Heliscus tropicus (Passalidae) is a key species constituting the core of all of the networks and could be considered an ecosystem engineer in cloud forests. By exploring links between saproxylic beetles and deadwood characteristics, we can further our understanding of species interaction in order to develop management strategies oriented towards the protection of species and their habitats in this threatened ecosystem.

Temporal Shifts and Cactus-Beetle Networks in an Intertropical Semiarid Zone in Mexico.

Cactus-dominated semiarid scrubland covers 40% of Mexican territory. This ecosystem is highly dynamic and undergoes drastic changes caused by seasonality. These temporal changes influence saprophagous insect communities associated with Cactaceae species. In this study, we analyzed the beetle community associated with decayed cactus species at the Barranca de Metztitlán Biosphere Reserve in central Mexico. We compared the diversity and composition of beetle communities in rainy and dry seasons; moreover, the network architecture of cactus-beetle interactions was examined. High dominance and abundance were detected in rainy assemblages, whereas the dry season had less abundance but more ecological diversity. A nested structure was found between individual cactus species and beetle species, as well as in an intrapopulation network between fragments of the columnar cacti Isolatocereus dumortieri (Scheidw.) Backeb. (Cactaceae), and beetle species for both seasons (rainy and dry). This finding shows more generalist than specialist beetle species inhabiting cactus species. Further research is still needed to understand whether the presence of these beetle species is determined by microhabitat conditions or the abundance of prey associated with decayed cacti. This is the first step in untangling the complex interactions among cactus-beetle species involved in the decomposition process of cacti in semiarid environments. This study provides evidence of temporal shifts in abundance and diversity patterns of these beetles associated with decayed cacti; furthermore, we did not detect an influence of seasonality on the structure of cactus-beetle interactions.

Populations and assemblages living on the edge: dung beetles responses to forests-pasture ecotones.

Edge effects alter insect biodiversity in several ways. However, we still have a limited understanding on simultaneous responses of ecological populations and assemblages to ecotones, especially in human modified landscapes. We analyze edge effects on dung beetle populations and assemblages between livestock pastures and native temperate forests (Juniperus and pine-oak forests (POFs)) to describe how species abundances and assemblage parameters respond to edge effects through gradients in forest-pasture ecotones. In Juniperus forest 13 species avoided the ecotones: six species showed greater abundance in forest interior and seven in pasturelands, while the other two species had a neutral response to the edge. In a different way, in POF we found five species avoiding the edge (four with greater abundance in pastures and only one in forest), two species had a neutral response, and two showed a unimodal pattern of abundance near to the edge. At the assemblage level edge effects are masked, as species richness, diversity, functional richness, functional evenness, and compositional incidence dissimilarity did not vary along forest-pasture ecotones. However, total abundance and functional divergence showed higher values in pastures in one of the two sampling localities. Also, assemblage similarity based on species' abundance showed a peak near to the edge in POF. We propose that conservation efforts in human-managed landscapes should focus on mitigating current and delayed edge effects. Ecotone management will be crucial in livestock dominated landscapes to conserve regional biodiversity and the environmental services carried out by dung beetles.

The role of sex and age in the architecture of intrapopulation howler monkey-plant networks in continuous and fragmented rain forests.

We evaluated the structure of intrapopulation howler monkey-plant interactions by focusing on the plant species consumed by different sex and age classes in continuous and fragmented forests in southern Mexico. For this we used network analysis to evaluate the impact of fragmentation on howler population traits and on resource availability and food choice. A total of 37 tree and liana species and seven plant items (bark, immature fruits, flowers, mature fruits, immature leaves, mature leaves and petioles) were consumed, but their relative consumption varied according to sex and age classes and habitat type. Overall, adult females consumed the greatest number of plant species and items while infants and juveniles the lowest. For both continuous and fragmented forests, we found a nested diet for howler monkey-plant networks: diets of more selective monkeys represent subsets of the diets of other individuals. Nestedness was likely due to the high selectivity of early life stages in specific food plants and items, which contrasts with the generalized foraging behaviour of adults. Information on the extent to which different plant species and primate populations depend on such interactions in different habitats will help to make accurate predictions about the potential impact of disturbances on plant-animal interaction networks.

The Multiple Impacts of Tropical Forest Fragmentation on Arthropod Biodiversity and on their Patterns of Interactions with Host Plants.

Tropical rain forest fragmentation affects biotic interactions in distinct ways. Little is known, however, about how fragmentation affects animal trophic guilds and their patterns of interactions with host plants. In this study, we analyzed changes in biotic interactions in forest fragments by using a multitrophic approach. For this, we classified arthropods associated with Heliconia aurantiaca herbs into broad trophic guilds (omnivores, herbivores and predators) and assessed the topological structure of intrapopulation plant-arthropod networks in fragments and continuous forests. Habitat type influenced arthropod species abundance, diversity and composition with greater abundance in fragments but greater diversity in continuous forest. According to trophic guilds, coleopteran herbivores were more abundant in continuous forest and overall omnivores in fragments. Continuous forest showed a greater diversity of interactions than fragments. Only in fragments, however, did the arthropod community associated with H aurantiaca show a nested structure, suggesting novel and/or opportunistic host-arthropod associations. Plants, omnivores and predators contributed more to nestedness than herbivores. Therefore, Heliconia-arthropod network properties do not appear to be maintained in fragments mainly caused by the decrease of herbivores. Our study contributes to the understanding of the impact of fragmentation on the structure and dynamics of multitrophic arthropod communities associated with a particular plant species of the highly biodiverse tropical forests. Nevertheless, further replication of study sites is needed to strengthen the conclusion that forest fragmentation negatively affects arthropod assemblages.