Long-term spatiotemporal variation in density of a tropical folivore: responses to a complex disturbance regime.

The Anthropocene is a time of unprecedented and accelerating rates of environmental change that includes press (e.g., climate change) and pulse disturbances (e.g., cyclonic storms, land use change) that interact to affect spatiotemporal dynamics in the density, distribution, and biodiversity of organisms. We leverage three decades of spatially explicit data on the density of a tropical folivore (Lamponius portoricensis [Insecta, Phasmida]) in a hurricane-mediated ecosystem (montane rainforest of Puerto Rico), along with associated environmental attributes, to disentangle the effects of interacting disturbances at multiple spatial scales. Spatiotemporal variation in density at a small spatial scale is affected by disturbance-related characteristics (hurricane severity, time after most recent major hurricane, ambient temperature, and understory temperature), legacies of previous land use, and understory habitat structure. Nonetheless, only a small proportion of spatiotemporal variation in density was related to those characteristics. In contrast, the majority of interannual variation in mean density at a larger scale was related to disturbance characteristics and understory habitat structure. These factors combine to affect a weak and declining trend in the density of L. portoricensis over time. The low resistance of L. portoricensis to Hurricane Hugo, as compared to Hurricanes Georges and Maria, likely arose because a drought followed Hurricane Hugo. The disturbance regime of the region is predicted to include increases in ambient temperatures, frequency of high-intensity storms, and frequency of droughts. Such trends may combine to threaten the conservation status of L. portoricensis, and other species with which it shares similar life history characteristics.

Effects of Host Species Identity and Diet on the Biodiversity of the Microbiota in Puerto Rican Bats.

Microbiota perform vital functions for their mammalian hosts, making them potential drivers of host evolution. Understanding effects of environmental factors and host characteristics on the composition and biodiversity of the microbiota may provide novel insights into the origin and maintenance of these symbiotic relationships. Our goals were to (1) characterize biodiversity of oral and rectal microbiota in bats from Puerto Rico; and (2) determine the effects of geographic location and host characteristics on that biodiversity. We collected bats and their microbiota from three sites, and used four metrics (species richness, Shannon diversity, Camargo evenness, Berger-Parker dominance) to characterize biodiversity. We quantified the relative importance of site, host sex, host species-identity, and host foraging-guild on biodiversity of the microbiota. Microbe biodiversity was highly variable among conspecifics. Geographical location exhibited consistent effects, whereas host sex did not. Within each host guild, host species exhibited consistent differences in biodiversity of oral microbiota and of rectal microbiota. Oral microbe biodiversity was indistinguishable between guilds, whereas rectal microbe biodiversity was significantly greater in carnivores than in herbivores. The high intraspecific and spatial variation in microbe biodiversity necessitate a relatively large number of samples to statistically isolate the effects of environmental or host characteristics on the microbiota. Species-specific biodiversity of oral microbiota suggests these communities are structured by direct interactions with the host immune system via epithelial receptors. In contrast, the number of microbial taxa that a host gut supports may be driven by host diet-diversity or composition.

A canonical metacommunity structure over 3 decades: ecologically consistent but spatially dynamic patterns in a hurricane-prone montane forest.

The Anthropocene is a time of rapid change induced by human activities, including pulse and press disturbances that affect the species composition of local communities and connectivity among them, giving rise to spatiotemporal dynamics at multiple scales. We evaluate effects of global warming and repeated intense hurricanes on gastropod metacommunities in montane tropical rainforests of Puerto Rico for each of 28 consecutive years. Specifically, we quantified metacommunity structure each year; assessed effects of global warming, hurricane-induced disturbance, and secondary succession on interannual variation in metacommunity structure; and evaluated legacies of previous land use on metacommunity structure. Gastropods were sampled annually during a 28-year period characterized by disturbance and succession associated with 3 major hurricanes (Hurricanes Hugo, Georges, and Maria). For each year, we evaluated coherence (the extent to which the environmental distributions of species are uninterrupted along a common latent environmental gradient), species range turnover, and species range boundary clumping; and conducted co-occurrence analyses for each pair of species. We used generalized linear mixed-effects model to evaluate long-term responses of the metacommunity to aspects of global warming and disturbance. Metacommunity structure was remarkably stable, with consistent patterns of species co-occurrence. Disturbance, warming, and successional stage had little effect on metacommunity structure. Despite great temporal variation in environmental conditions, groups of species tracked their niche through space and time to maintain the same general structure. Consequently, metacommunity structure was highly resistant and resilient to multiple disturbances, even those that greatly altered forest structure.

Arthropods are not declining but are responsive to disturbance in the Luquillo Experimental Forest, Puerto Rico.

A number of recent studies have documented long-term declines in abundances of important arthropod groups, primarily in Europe and North America. These declines are generally attributed to habitat loss, but a recent study [B.C. Lister, A. Garcia, Proc. Natl. Acad. Sci. USA 115, E10397-E10406 (2018)] from the Luquillo Experimental Forest (LEF) in Puerto Rico attributed declines to global warming. We analyze arthropod data from the LEF to evaluate long-term trends within the context of hurricane-induced disturbance, secondary succession, and temporal variation in temperature. Our analyses demonstrate that responses to hurricane-induced disturbance and ensuing succession were the primary factors that affected total canopy arthropod abundances on host trees, as well as walkingstick abundance on understory shrubs. Ambient and understory temperatures played secondary roles for particular arthropod species, but populations were just as likely to increase as they were to decrease in abundance with increasing temperature. The LEF is a hurricane-mediated system, with major hurricanes effecting changes in temperature that are larger than those induced thus far by global climate change. To persist, arthropods in the LEF must contend with the considerable variation in abiotic conditions associated with repeated, large-scale, and increasingly frequent pulse disturbances. Consequently, they are likely to be well-adapted to the effects of climate change, at least over the short term. Total abundance of canopy arthropods after Hurricane Maria has risen to levels comparable to the peak after Hurricane Hugo. Although the abundances of some taxa have declined over the 29-y period, others have increased, reflecting species turnover in response to disturbance and secondary succession.

Multiple dimensions of bat biodiversity along an extensive tropical elevational gradient.

Research concerning spatial dynamics of biodiversity generally has been limited to considerations of the taxonomic dimension, which is insensitive to interspecific variation in ecological or evolutionary characteristics that play important roles in species assembly and provide linkages to ecosystem services. Consequently, the assumption that the taxonomic dimension is a good surrogate for other dimensions remains unconfirmed. We assessed variation in taxonomic (species richness) as well as phylogenetic and functional (Rao's quadratic entropy, a measurement of dispersion) dimensions of bat biodiversity along an elevational gradient in the Manu Biosphere Reserve of Peru. Phylogenetic dispersion was based on relatedness of species derived from a mammalian supertree. Functional dispersion was estimated separately for each of six functional components that reflect particular niche axes (e.g. diet, foraging strategy, body size) and for all functional components combined. Species richness declined nonlinearly with elevation, whereas phylogenetic dispersion and functional dispersion based on all functional components were not significantly associated with elevation (orthogonal polynomial regression). Moreover, considerable heterogeneity in the form of elevational relationships existed among functional components. After accounting for variation in species richness, dispersion of phylogenetic, diet and foraging strategy attributes were significantly greater than expected at high elevations, whereas dispersion of body size was significantly less than expected at high elevations. Species richness was a poor surrogate for phylogenetic or functional dispersion. Functional dispersion based on multiple components obscured patterns detected by particular components and hindered identification of mechanistic explanations for elevational variation in biodiversity. Variation in phylogenetic dispersion effectively captured the composite variation represented by all functional components, suggesting a phylogenetic signal in functional attributes. Mechanisms that give rise to variation in richness do not fully account for variation in phylogenetic or functional characteristics of assemblages. Greater than expected phylogenetic, diet and foraging strategy dispersion at high elevations were associated with the loss of phylogenetically or functionally redundant species, suggesting that increasing interspecific competition with decreasing productivity resulted in competitive exclusion. In contrast, low dispersion of size attributes at high elevations suggests the importance of abiotic filtering that favours small-sized species that can more easily enter torpor.

Long-term dynamics of tropical walking sticks in response to multiple large-scale and intense disturbances.

Understanding the effects of disturbance and secondary succession on spatio-temporal patterns in the abundance of species is stymied by a lack of long-term demographic data, especially in response to infrequent and high intensity disturbances, such as hurricanes. Moreover, resistance and resilience to hurricane-induced disturbance may be mediated by legacies of previous land use, although such interactive effects are poorly understood, especially in tropical environments. We address these central issues in disturbance ecology by analyzing an extensive dataset, spanning the impacts of Hurricanes Hugo and Georges, on the abundance of a Neotropical walking stick, Lamponius portoricensis, in tabonuco rainforest of Puerto Rico during the wet and dry seasons from 1991 to 2007. By synthesizing data from two proximate sites in tabonuco forest, we show that resistance to Hurricane Hugo (97% reduction in abundance) was much less than resistance to Hurricane Georges (21% reduction in abundance). Based on a powerful statistical approach (generalized linear mixed-effects models with Poisson error terms), we documented that the temporal trajectories of abundance during secondary succession (i.e., patterns of resilience) differed between hurricanes and among historical land use categories, but that the effects of hurricanes and land use histories were independent of each other. These complex results likely arise because of differences in the intensities of the two hurricanes with respect to microclimatic effects (temperature and moisture) in the forest understory, as well as to time-lags in the response of L. portoricensis to changes in the abundance and distribution of preferred food plants (Piper) in post-hurricane environments.

Elements of metacommunity structure of Paraguayan bats: multiple gradients require analysis of multiple ordination axes.

Techniques to evaluate elements of metacommunity structure (EMS; coherence, species turnover and range boundary clumping) have been available for several years. Such approaches are capable of determining which idealized pattern of species distribution best describes distributions in a metacommunity. Nonetheless, this approach rarely is employed and such aspects of metacommunity structure remain poorly understood. We expanded an extant method to better investigate metacommunity structure for systems that respond to multiple environmental gradients. We used data obtained from 26 sites throughout Paraguay as a model system to demonstrate application of this methodology. Using presence-absence data for bats, we evaluated coherence, species turnover and boundary clumping to distinguish among six idealized patterns of species distribution. Analyses were conducted for all bats as well as for each of three feeding ensembles (aerial insectivores, frugivores and molossid insectivores). For each group of bats, analyses were conducted separately for primary and secondary axes of ordination as defined by reciprocal averaging. The Paraguayan bat metacommunity evinced Clementsian distributions for primary and secondary ordination axes. Patterns of species distribution for aerial insectivores were dependent on ordination axis, showing Gleasonian distributions when ordinated according to the primary axis and Clementsian distributions when ordinated according to the secondary axis. Distribution patterns for frugivores and molossid insectivores were best described as random. Analysis of metacommunities using multiple ordination axes can provide a more complete picture of environmental variables that mold patterns of species distribution. Moreover, analysis of EMS along defined gradients (e.g., latitude, elevation and depth) or based on alternative ordination techniques may complement insights based on reciprocal averaging because the fundamental questions addressed in analyses are contingent on the ordination technique that is employed.

Geographical ecology of Paraguayan bats: spatial integration and metacommunity structure of interacting assemblages.

We examined the relative contributions of regional spatial characteristics and local environmental conditions in determining Paraguayan bat species composition. We used a suite of full and partial redundancy analyses to estimate four additive partitions of variance in bat species composition: (a) unexplained variation, (b) that explained purely by spatial characteristics, (c) that explained purely by local environmental conditions and (d) that explained jointly by space and environment. The spatial component to bat species composition was greater than the environmental component and both pure spatial and pure environmental characteristics accounted for significant amounts of variation in bat species composition. Results from variance decomposition suggest that the mass effects model describes metacommunity structure of Paraguayan bats better than species sorting or neutral models. Such mass effects may potentially be general for bats and could explain the inability of purely local factors to fully account for bat community organization. Mass effects also have substantial conservation implications because rescue effects may enhance the persistence of mobile species in fragmented landscapes with relatively few protected sites.