Underlying and proximate drivers of biodiversity changes in Mesoamerican biosphere reserves.

Protected areas are of paramount relevance to conserving wildlife and ecosystem contributions to people. Yet, their conservation success is increasingly threatened by human activities including habitat loss, climate change, pollution, and species overexploitation. Thus, understanding the underlying and proximate drivers of anthropogenic threats is urgently needed to improve protected areas' effectiveness, especially in the biodiversity-rich tropics. We addressed this issue by analyzing expert-provided data on long-term biodiversity change (last three decades) over 14 biosphere reserves from the Mesoamerican Biodiversity Hotspot. Using multivariate analyses and structural equation modeling, we tested the influence of major socioeconomic drivers (demographic, economic, and political factors), spatial indicators of human activities (agriculture expansion and road extension), and forest landscape modifications (forest loss and isolation) as drivers of biodiversity change. We uncovered a significant proliferation of disturbance-tolerant guilds and the loss or decline of disturbance-sensitive guilds within reserves causing a "winner and loser" species replacement over time. Guild change was directly related to forest spatial changes promoted by the expansion of agriculture and roads within reserves. High human population density and low nonfarming occupation were identified as the main underlying drivers of biodiversity change. Our findings suggest that to mitigate anthropogenic threats to biodiversity within biosphere reserves, fostering human population well-being via sustainable, nonfarming livelihood opportunities around reserves is imperative.

Linking Anthropogenic Landscape Perturbation to Herbivory and Pathogen Leaf Damage in Tropical Tree Communities.

Anthropogenic disturbance of tropical humid forests leads to habitat loss, biodiversity decline, landscape fragmentation, altered nutrient cycling and carbon sequestration, soil erosion, pest/pathogen outbreaks, among others. Nevertheless, the impact of these alterations in multitrophic interactions, including host-pathogen and vector-pathogen dynamics, is still not well understood in wild plants. This study aimed to provide insights into the main drivers for the incidence of herbivory and plant pathogen damage, specifically, into how vegetation traits at the local and landscape scale modulate such interactions. For this purpose, in the tropical forest of Calakmul (Campeche, Mexico), we characterised the foliar damage caused by herbivores and pathogens in woody vegetation of 13 sampling sites representing a gradient of forest disturbance and fragmentation in an anthropogenic landscape from well preserved to highly disturbed and fragmented areas. We also evaluated how the incidence of such damage was modulated by the vegetation and landscape attributes. We found that the incidence of damage caused by larger, mobile, generalist herbivores, was more sensitive to changes in landscape configuration, while the incidence of damage caused by small and specialised herbivores with low dispersal capacity was more influenced by vegetation and landscape composition. In relation to pathogen symptoms, the herbivore-induced foliar damage seems to be the main factor related to their incidence, indicating the enormous importance of herbivorous insects in the modulation of disease dynamics across tropical vegetation, as they could be acting as vectors and/or facilitating the entry of pathogens by breaking the foliar tissue and the plant defensive barriers. The incidence of pathogen damage also responded to vegetation structure and landscape configuration; the incidence of anthracnose, black spot, and chlorosis, for example, were favoured in sites surrounded by smaller patches and a higher edge density, as well as those with a greater aggregation of semi-evergreen forest patches. Fungal pathogens were shown to be an important cause of foliar damage for many woody species. Our results indicate that an increasing transformation and fragmentation of the tropical forest of southern Mexico could reduce the degree of specialisation in plant-herbivore interactions and enhance the proliferation of generalist herbivores (chewers and scrapers) and of mobile leaf suckers, and consequently, the proliferation of some symptoms associated with fungal pathogens such as fungus black spots and anthracnose. The symptoms associated with viral and bacterial diseases and to nutrient deficiency, such as chlorosis, could also increase in the vegetation in fragmented landscapes with important consequences in the health and productivity of wild and cultivated plant species. This is a pioneering study evaluating the effect of disturbances on multitrophic interactions, offering key insights on the main drivers of the changes in herbivory interactions and incidence of plant pathogens in tropical forests.

The Assembly of Tropical Dry Forest Tree Communities in Anthropogenic Landscapes: The Role of Chemical Defenses.

The effect of anthropogenic disturbance on plant community traits and tradeoffs remains poorly explored in tropical forests. In this study, we aimed to identify tradeoffs between defense and other plant functions related to growth processes in order to detect potential aboveground and edaphic environmental conditions modulating traits variation on plant communities, and to find potential assembly rules underlying species coexistence in secondary (SEF) and old-growth forests (OGF). We measured the foliar content of defense phytochemicals and leaf traits related to fundamental functions on 77 species found in SEF and OGF sites in the Jalisco dry forest ecoregion, Mexico, and we explored (1) the trait-trait and trait-habitat associations, (2) the intra and interspecies trait variation, and (3) the traits-environment associations. We found that phytochemical content was associated with high leaf density and leaf fresh mass, resulting in leaves resistant to drought and high radiation, with chemical and physical defenses against herbivore/pathogen attack. The phytochemicals and chlorophyll concentrations were negatively related, matching the predictions of the Protein Competition Model. The phylogenetic signal in functional traits, suggests that abundant clades share the ability to resist the harsh biotic and abiotic conditions and face similar tradeoffs between productive and defensive functions. Environmental filters could modulate the enhanced expression of defensive phytochemicals in SEF, while, in OGFs, we found a stronger filtering effect driving community assembly. This could allow for the coexistence of different defensive strategies in OGFs, where a greater species richness could dilute the prevalence of pathogens/herbivores. Consequently, anthropogenic disturbance could alter TDF ecosystem properties/services and functioning.

Homogenization of terrestrial mammals in fragmented rainforests: the loss of species turnover and its landscape drivers.

Understanding the factors and mechanisms shaping differences in species composition across space and time (ß-diversity) in human-modified landscapes has key ecological and applied implications. This topic is, however, challenging because landscape disturbance can promote either decreases (biotic homogenization) or increases (biotic differentiation) in ß-diversity. We assessed temporal differences in intersite ß-diversity of medium-bodied and large-bodied mammals in the fragmented Lacandona rainforest, Mexico. We hypothesized that, given the relatively short history of land-use changes in the region, and the gain and loss of some species caused by landscape spatial changes, ß-diversity would increase through time, especially its nestedness component. We estimated ß-diversity between 24 forest sites (22 forest patches and two continuous forest sites) in 2011 and 2017 to assess whether ß-diversity is decreasing or increasing in the region, and calculated its turnover and nestedness components to understand the mechanisms responsible for changes in ß-diversity, separately assessing mammal groups with different body mass, feeding guild, and habitat specialization. We then related such temporal changes in ß-diversity to temporal changes in five landscape variables (forest cover, matrix openness, number of patches, edge density and interpatch distance) to identify the landscape drivers of ß-diversity. In contrast with our expectations, ß-diversity decreased over time, suggesting an ongoing biotic homogenization process. This pattern was mostly driven by a decrease in species turnover in all mammal groups, especially in landscapes with decreasing forest cover and increasing forested matrices. Although the nestedness component showed a three-fold increase through time, species turnover was 22 and six times higher than nestedness in 2011 and 2017, respectively. The decreased turnover appears to be driven by an increase in dispersal (i.e., spillover) of native species among patches. The prevalence of species turnover over nestedness indicates that different forest sites have a fairly distinct subset of species (i.e., high complementarity in species composition). Therefore, conserving all remaining forest patches and increasing forest cover is of utmost importance to effectively maintain ß-diversity and conserve the total diversity (γ) of mammal assemblages in this Mesoamerican biodiversity hotspot.

Drivers potentially influencing host-bat fly interactions in anthropogenic neotropical landscapes at different spatial scales.

The anthropogenic modification of natural landscapes, and the consequent changes in the environmental conditions and resources availability at multiple spatial scales can affect complex species interactions involving key-stone species such as bat-parasite interactions. In this study, we aimed to identify the drivers potentially influencing host-bat fly interactions at different spatial scales (at the host, vegetation stand and landscape level), in a tropical anthropogenic landscape. For this purpose, we mist-netted phyllostomid and moormopid bats and collected the bat flies (streblids) parasitizing them in 10 sites representing secondary and old growth forest. In general, the variation in fly communities largely mirrored the variation in bat communities as a result of the high level of specialization characterizing host-bat fly interaction networks. Nevertheless, we observed that: (1) bats roosting dynamics can shape bat-streblid interactions, modulating parasite prevalence and the intensity of infestation; (2) a degraded matrix could favor crowding and consequently the exchange of ectoparasites among bat species, lessening the level of specialization of the interaction networks and promoting novel interactions; and (3) bat-fly interaction can also be shaped by the dilution effect, as a decrease in bat diversity could be associated with a potential increase in the dissemination and prevalence of streblids.

Structure and diversity of phyllostomid bat assemblages on riparian corridors in a human-dominated tropical landscape.

Tropical forests around the world have been lost, mainly because of agricultural activities. Linear elements like riparian vegetation in fragmented tropical landscapes help maintain the native flora and fauna. Information about the role of riparian corridors as a reservoir of bat species, however, is scanty. We assessed the value of riparian corridors on the conservation of phyllostomid bat assemblage in an agricultural landscape of southern Mexico. For 2 years (2011-2013), mist-netting at ground level was carried out twice during the dry season (December to May) and twice during the wet season (June to November) in different habitats: (1) riparian corridors in mature forest, (2) riparian corridors in pasture, (3) continuous forest away from riparian vegetation, and (4) open pastures. Each habitat was replicated three times. To determine the influence of vegetation structure on bat assemblages, all trees (≥10 cm dbh) were sampled in all habitats. Overall, 1752 individuals belonging to 28 species of Phyllostomidae were captured with Sternodermatinae being the most rich and abundant subfamily. Riparian corridors in mature forest and pastures had the greatest species richness and shared 65% of all species. Open pastures had the lowest richness and abundance of bats with no Phyllostominae species recorded. Six of the 18 species recorded could be considered as habitat indicators. There was a positive relationship between bat species composition and tree basal area. Our findings suggest that contrary to our expectations, bats with generalist habits and naturally abundant could be useful detector taxa of habitat modification, rather than bats strongly associated with undisturbed forest. Also in human-dominated landscapes, the maintenance of habitat elements such as large trees in riparian corridors can serve as reservoirs for bat species, especially for those that are strongly associated with undisturbed forest.

Influence of matrix type on tree community assemblages along tropical dry forest edges.

UNLABELLED: • PREMISE OF THE STUDY: Anthropogenic habitat edges have strong negative consequences for the functioning of tropical ecosystems. However, edge effects on tropical dry forest tree communities have been barely documented.• METHODS: In Chamela, Mexico, we investigated the phylogenetic composition and structure of tree assemblages (≥5 cm dbh) along edges abutting different matrices: (1) disturbed vegetation with cattle, (2) pastures with cattle and, (3) pastures without cattle. Additionally, we sampled preserved forest interiors.• KEY RESULTS: All edge types exhibited similar tree density, basal area and diversity to interior forests, but differed in species composition. A nonmetric multidimensional scaling ordination showed that the presence of cattle influenced species composition more strongly than the vegetation structure of the matrix; tree assemblages abutting matrices with cattle had lower scores in the ordination. The phylogenetic composition of tree assemblages followed the same pattern. The principal plant families and genera were associated according to disturbance regimes as follows: pastures and disturbed vegetation (1) with cattle and (2) without cattle, and (3) pastures without cattle and interior forests. All habitats showed random phylogenetic structures, suggesting that tree communities are assembled mainly by stochastic processes. Long-lived species persisting after edge creation could have important implications in the phylogenetic structure of tree assemblages.• CONCLUSIONS: Edge creation exerts a stronger influence on TDF vegetation pathways than previously documented, leading to new ecological communities. Phylogenetic analysis may, however, be needed to detect such changes.

Phyllostomid bat occurrence in successional stages of neotropical dry forests.

Tropical dry forests (TDFs) are highly endangered tropical ecosystems being replaced by a complex mosaic of patches of different successional stages, agricultural fields and pasturelands. In this context, it is urgent to understand how taxa playing critical ecosystem roles respond to habitat modification. Because Phyllostomid bats provide important ecosystem services (e.g. facilitate gene flow among plant populations and promote forest regeneration), in this study we aimed to identify potential patterns on their response to TDF transformation in sites representing four different successional stages (initial, early, intermediate and late) in three Neotropical regions: México, Venezuela and Brazil. We evaluated bat occurrence at the species, ensemble (abundance) and assemblage level (species richness and composition, guild composition). We also evaluated how bat occurrence was modulated by the marked seasonality of TDFs. In general, we found high seasonal and regional specificities in phyllostomid occurrence, driven by specificities at species and guild levels. For example, highest frugivore abundance occurred in the early stage of the moistest TDF, while highest nectarivore abundance occurred in the same stage of the driest TDF. The high regional specificity of phyllostomid responses could arise from: (1) the distinctive environmental conditions of each region, (2) the specific behavior and ecological requirements of the regional bat species, (3) the composition, structure and phenological patterns of plant assemblages in the different stages, and (4) the regional landscape composition and configuration. We conclude that, in tropical seasonal environments, it is imperative to perform long-term studies considering seasonal variations in environmental conditions and plant phenology, as well as the role of landscape attributes. This approach will allow us to identify potential patterns in bat responses to habitat modification, which constitute an invaluable tool for not only bat biodiversity conservation but also for the conservation of the key ecological processes they provide.

Functional regeneration and spectral reflectance of trees during succession in a highly diverse tropical dry forest ecosystem.

PREMISE OF THE STUDY: The function of most ecosystems has been altered by human activities. To asses the recovery of plant communities, we must evaluate the recovery of plant functional traits. The seasonally dry tropical forest (SDTF), a highly threatened ecosystem, is assumed to recover relatively quickly from disturbance, but an integrated evaluation of recovery in floristic, structural, and functional terms has not been performed. In this study we aimed to (a) compare SDTF plant functional, floristic, and structural change along succession; (b) identify tree functional groups; and (c) explore the spectral properties of different successional stages. METHODS: Across a SDTF successional gradient, we evaluated the change of species composition, vegetation structure, and leaf spectral reflectance and functional traits (related to water use, light acquisition, nutrient conservation, and CO(2) acquisition) of 25 abundant tree species. KEY RESULTS: A complete recovery of SDTF takes longer than the time period inferred from floristic or structural data. Plant functional traits changed along succession from those that maximize photoprotection and heat dissipation in early succession, where temperature is an environmental constraint, to those that enhance light acquisition in late succession, where light may be limiting. A spectral indicator of plant photosynthetic performance (photochemical reflectance index) discriminated between early and late succession. This constitutes a foundation for further exploration of remote sensing technologies for studying tropical succession. CONCLUSIONS: A functional approach should be incorporated as a regular descriptor of forest succession because it provides a richer understanding of vegetation dynamics than is offered by either the floristic or structural approach alone.

Local and landscape factors determining occurrence of phyllostomid bats in tropical secondary forests.

Neotropical forests are being increasingly replaced by a mosaic of patches of different successional stages, agricultural fields and pasture lands. Consequently, the identification of factors shaping the performance of taxa in anthropogenic landscapes is gaining importance, especially for taxa playing critical roles in ecosystem functioning. As phyllostomid bats provide important ecological services through seed dispersal, pollination and control of animal populations, in this study we assessed the relationships between phyllostomid occurrence and the variation in local and landscape level habitat attributes caused by disturbance. We mist-netted phyllostomids in 12 sites representing 4 successional stages of a tropical dry forest (initial, early, intermediate and late). We also quantitatively characterized the habitat attributes at the local (vegetation structure complexity) and the landscape level (forest cover, area and diversity of patches). Two focal scales were considered for landscape characterization: 500 and 1000 m. During 142 sampling nights, we captured 606 individuals representing 15 species and 4 broad guilds. Variation in phyllostomid assemblages, ensembles and populations was associated with variation in local and landscape habitat attributes, and this association was scale-dependent. Specifically, we found a marked guild-specific response, where the abundance of nectarivores tended to be negatively associated with the mean area of dry forest patches, while the abundance of frugivores was positively associated with the percentage of riparian forest. These results are explained by the prevalence of chiropterophilic species in the dry forest and of chiropterochorous species in the riparian forest. Our results indicate that different vegetation classes, as well as a multi-spatial scale approach must be considered for evaluating bat response to variation in landscape attributes. Moreover, for the long-term conservation of phyllostomids in anthropogenic landscapes, we must realize that the management of the habitat at the landscape level is as important as the conservation of particular forest fragments.