Juliano Sarmento Cabral

Late Quaternary climate change shapes island biodiversity

Island biogeographical models consider islands either as geologically static with biodiversity resulting from ecologically neutral immigration–extinction dynamics, or as geologically dynamic with biodiversity resulting from immigration–speciation–extinction dynamics influenced by changes in island characteristics over millions of years. Present climate and spatial arrangement of islands, however, are rather exceptional compared to most of the Late Quaternary, which is characterized by recurrent cooler and drier glacial periods. These climatic oscillations over short geological timescales strongly affected sea levels and caused massive changes in island area, isolation and connectivity, orders of magnitude faster than the geological processes of island formation, subsidence and erosion considered in island theory. Consequences of these oscillations for present biodiversity remain unassessed. Here we analyse the effects of present and Last Glacial Maximum (LGM) island area, isolation, elevation and climate on key components of angiosperm diversity on islands worldwide. We find that post-LGM changes in island characteristics, especially in area, have left a strong imprint on present diversity of endemic species. Specifically, the number and proportion of endemic species today is significantly higher on islands that were larger during the LGM. Native species richness, in turn, is mostly determined by present island characteristics. We conclude that an appreciation of Late Quaternary environmental change is essential to understand patterns of island endemism and its underlying evolutionary dynamics.

Oceanic island biogeography through the lens of the general dynamic model: Assessment and prospect

The general dynamic model of oceanic island biogeography (GDM) has added a new dimension to theoretical island biogeography in recognizing that geological processes are key drivers of the evolutionary processes of diversification and extinction within remote islands. It provides a dynamic and essentially non‐equilibrium framework generating novel predictions for emergent diversity properties of oceanic islands and archipelagos. Its publication in 2008 coincided with, and spurred on, renewed attention to the dynamics of remote islands. We review progress, both in testing the GDMs predictions and in developing and enhancing ecological–evolutionary understanding of oceanic island systems through the lens of the GDM. In particular, we focus on four main themes: (i) macroecological tests using a space‐for‐time rationale; (ii) extensions of theory to islands following different patterns of ontogeny; (iii) the implications of GDM dynamics for lineage diversification and trait evolution; and (iv) the potential for downscaling GDM dynamics to local‐scale ecological patterns and processes within islands. We also consider the implications of the GDM for understanding patterns of non‐native species diversity. We demonstrate the vitality of the field of island biogeography by identifying a range of potentially productive lines for future research.

Mechanistic simulation models in macroecology and biogeography: state-of-art and prospects

Macroecology and biogeography are concerned with understanding biodiversity patterns across space and time. In the past, the two disciplines have addressed this question mainly with correlative approaches, despite frequent calls for more mechanistic explanations. Recent advances in computational power, theoretical understanding, and statistical tools are, however, currently facilitating the development of more system-oriented, mechanistic models. We review these models, identify different model types and theoretical frameworks, compare their processes and properties, and summarize emergent findings. We show that ecological (physiology, demographics, dispersal, biotic interactions) and evolutionary processes, as well as environmental and human-induced drivers, are increasingly modelled mechanistically; and that new insights into biodiversity dynamics emerge from these models. Yet, substantial challenges still lie ahead for this young research field. Among these, we identify scaling, calibration, validation, and balancing complexity as pressing issues. Moreover, particular process combinations are still understudied, and so far models tend to be developed for specific applications. Future work should aim at developing more flexible and modular models that not only allow different ecological theories to be expressed and contrasted, but which are also built for tight integration with all macroecological data sources. Moving the field towards such a ‘systems macroecology’ will test and improve our understanding of the causal pathways through which eco-evolutionary processes create diversity patterns across spatial and temporal scales. This article is protected by copyright. All rights reserved.

Biogeographic, climatic and spatial drivers differentially affect α, β and γ diversities on oceanic archipelagos

Island biogeographic studies traditionally treat single islands as units of analysis. This ignores the fact that most islands are spatially nested within archipelagos. Here, we took a fundamentally different approach and focused on entire archipelagos using species richness of vascular plants on 23 archipelagos worldwide and their 174 constituent islands. We assessed differential effects of biogeographic factors (area, isolation, age, elevation), current and past climate (temperature, precipitation, seasonality, climate change velocity) and intra-archipelagic spatial structure (archipelago area, number of islands, area range, connectivity, environmental volume, inter-island distance) on plant diversity. Species diversity of each archipelago (γ) was additively partitioned into α, β, nestedness and replacement β-components to investigate the relative importance of environmental and spatial drivers. Multiple regressions revealed strong effects of biogeography and climate on α and γ, whereas spatial factors, particularly number of islands, inter-island distance and area range, were key to explain β. Structural equation models additionally suggested that γ is predominantly determined by indirect abiotic effects via its components, particularly β. This highlights that β and the spatial arrangement of islands are essential to understand insular ecology and evolution. Our methodological framework can be applied more widely to other taxa and archipelago-like systems, allowing new insights into biodiversity origin and maintenance.

Functional leaf traits of vascular epiphytes: vertical trends within the forest, intra‐ and interspecific trait variability, and taxonomic signals

Summary Analysing functional traits along environmental gradients can improve our understanding of the mechanisms structuring plant communities. Within forests, vertical gradients in light intensity, temperature and humidity are often pronounced. Vascular epiphytes are particularly suitable for studying the influence of these vertical gradients on functional traits because they lack contact with the soil and thus individual plants are entirely exposed to different environmental conditions, from the dark and humid understorey to the sunny and dry outer canopy. In this study, we analysed multiple aspects of the trait-based ecology of vascular epiphytes: shifts in trait values with height above ground (as a proxy for vertical environmental gradients) at community and species level, the importance of intra- vs. interspecific trait variability, and trait differences among taxonomic groups. We assessed ten leaf traits for 1151 individuals belonging to 83 epiphyte species of all major taxonomic groups co-occurring in a Panamanian lowland forest. Community mean trait values of many leaf traits were strongly correlated with height and particularly specific leaf area and chlorophyll concentration showed nonlinear, negative trends. Intraspecific trait variability was pronounced and accounted for one-third of total observed trait variance. Intraspecific trait adjustments along the vertical gradient were common and seventy per cent of all species showed significant trait–height relationships. In addition, intraspecific trait variability was positively correlated with the vertical range occupied by species. We observed significant trait differences between major taxonomic groups (orchids, ferns, aroids, bromeliads). In ferns, for instance, leaf dry matter content was almost twofold higher than in the other taxonomic groups. This indicates that some leaf traits are taxonomically conserved. Our study demonstrates that vertical environmental gradients strongly influence functional traits of vascular epiphytes. In order to understand community composition along such gradients, it is central to study several aspects of trait-based ecology, including both community and intraspecific trends of multiple traits.

Branchfall as a Demographic Filter for Epiphyte Communities: Lessons from Forest Floor-Based Sampling

Local variation in the abundance and richness of vascular epiphytes is often attributed to environmental characteristics such as substrate and microclimate. Less is known, however, about the impacts of tree and branch turnover on epiphyte communities. To address this issue, we surveyed branches and epiphytes found on the forest floor in 96 transects in two forests (Atlantic rainforest in Brazil and Caribbean rainforest in Panama). In the Brazilian forest, we additionally distinguished between edge and core study sites. We quantified branch abundance, epiphyte abundance, richness and proportion of adults to investigate the trends of these variables over branch diameter. Branches <2 cm in diameter comprised >90% of all branches on the forest floor. Abundance and richness of fallen epiphytes per transect were highest in the Brazilian core transects and lowest in the Panamanian transects. The majority of epiphytes on the floor (c. 65%) were found attached to branches. At all three study sites, branch abundance and branch diameter were negatively correlated, whereas epiphyte abundance and richness per branch, as well as the proportion of adults were positively correlated with branch diameter. The relationship between branch diameter and absolute epiphyte abundance or richness differed between study sites, which might be explained by differences in forest structure and dynamics. In the Panamanian forest, epiphytes had been previously inventoried, allowing an evaluation of our surveying method by comparing canopy and forest floor samplings. Individuals found on the forest floor corresponded to 13% of all individuals on branches <10 cm in diameter (including crowns), with abundance, richness and composition trends on forest floor reflecting canopy trends. We argue that forest floor surveys provide useful floristic and, most notably, demographic information particularly on epiphytes occurring on the thinnest branches, which are least accessible. Here, branchfall acts as an important demographic filter structuring epiphyte communities.

Karyotype diversity and genome size variation in Neotropical Maxillariinae orchids

Orchidaceae is a widely distributed plant family with very diverse vegetative and floral morphology, and such variability is also reflected in their karyotypes. However, since only a low proportion of Orchidaceae has been analysed for chromosome data, greater diversity may await to be unveiled. Here we analyse both genome size (GS) and karyotype in two subtribes recently included in the broadened Maxillariinea to detect how much chromosome and GS variation there is in these groups and to evaluate which genome rearrangements are involved in the species evolution. To do so, the GS (14 species), the karyotype - based on chromosome number, heterochromatic banding and 5S and 45S rDNA localisation (18 species) - was characterised and analysed along with published data using phylogenetic approaches. The GS presented a high phylogenetic correlation and it was related to morphological groups in Bifrenaria (larger plants - higher GS). The two largest GS found among genera were caused by different mechanisms: polyploidy in Bifrenaria tyrianthina and accumulation of repetitive DNA in Scuticaria hadwenii. The chromosome number variability was caused mainly through descending dysploidy, and x=20 was estimated as the base chromosome number. Combining GS and karyotype data with molecular phylogeny, our data provide a more complete scenario of the karyotype evolution in Maxillariinae orchids, allowing us to suggest, besides dysploidy, that inversions and transposable elements as two mechanisms involved in the karyotype evolution. Such karyotype modifications could be associated with niche changes that occurred during species evolution.

Interactions between ecological, evolutionary, and environmental processes unveil complex dynamics of island biodiversity

Aims Understanding how biodiversity emerges and varies in space and time is central to ecology and biogeography. Multiple processes affect biodiversity at different scales and organizational levels, hence progress in understanding biodiversity dynamics requires the integration of these underlying processes. Here we present BioGEEM (BioGeographical Eco-Evolutionary Model), a spatially-explicit, process-based model that integrates all processes hypothesized to be relevant for biodiversity dynamics and that can be used to evaluate their relative roles. Location Hypothetical oceanic islands Methods The model is stochastic, grid-based, and integrates ecological (metabolic constraints, demography, dispersal, and competition), evolutionary (mutation and speciation), and environmental (geo-climatic dynamics) processes. Plants on oceanic islands served as model system. We used the full model to test hypotheses about emergent patterns at different spatio-temporal scales and organizational levels (populations, species, communities, and assemblages), switching off processes to assess the importance 1) of competition for realistic population and range dynamics; 2) metabolic constraints for endemism and community composition; 3) environmental dynamics and 4) speciation for biogeographical patterns. Results The full model generated multiple patterns matching empirical and theoretical expectations. For example, populations were largest on young, species-poor islands. Species, particularly endemics, were better able to fill their potential range on small, species-poor islands. Richness gradients peaked at mid-elevations. The proportion of endemics was highest on old, large, and isolated environments within the islands. Species and trait richness showed unimodal temporal trends. Switching off selected processes affected these patterns, and we found most of our hypotheses supported. Main conclusions Integrating ecological, evolutionary, and environmental processes is essential to simultaneously generate realistic spatio-temporal dynamics at population, species, community, and assemblage level. Finally, large-scale biodiversity dynamics emerged directly from biological processes which make this mechanistic model a valuable ‘virtual long-term field station’ to study the linkages between biogeography and ecology.

Integrating community assembly and biodiversity to better understand ecosystem function: the Community Assembly and the Functioning of Ecosystems (CAFE) approach

The research of a generation of ecologists was catalysed by the recognition that the number and identity of species in communities influences the functioning of ecosystems. The relationship between biodiversity and ecosystem functioning (BEF) is most often examined by controlling species richness and randomising community composition. In natural systems, biodiversity changes are often part of a bigger community assembly dynamic. Therefore, focusing on community assembly and the functioning of ecosystems (CAFE), by integrating both species richness and composition through species gains, losses and changes in abundance, will better reveal how community changes affect ecosystem function. We synthesise the BEF and CAFE perspectives using an ecological application of the Price equation, which partitions the contributions of richness and composition to function. Using empirical examples, we show how the CAFE approach reveals important contributions of composition to function. These examples show how changes in species richness and composition driven by environmental perturbations can work in concert or antagonistically to influence ecosystem function. Considering how communities change in an integrative fashion, rather than focusing on one axis of community structure at a time, will improve our ability to anticipate and predict changes in ecosystem function.

The geography of the Anthropocene differs between the land and the sea

Climate change and other anthropogenic drivers of biodiversity change are unequally distributed across the world. Despite the implications for biodiversity change, it is unknown if the geographic patterns of drivers differ between the terrestrial and marine realm. Using global datasets on human population density, land and resource exploitation, pollution, species invasions, and climate change, we found stronger positive correlations among drivers in the terrestrial than in the marine realm, leading to areas of especially intense human impact on land. Climate change tended to be negatively correlated with other drivers in the terrestrial realm whereas the opposite was true in the marine realm. We show that different regions of the world are exposed to distinct ‘anthropogenic threat complexes’, comprising suites of drivers of varying intensities. Our global analysis highlights the broad conservation priorities needed to mitigate the drivers shaping biodiversity responses to anthropogenic change.