Peter M. Jørgensen

Beta‐diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly

Site-to-site variation in species composition (β-diversity) generally increases from low- to high-diversity regions. Although biogeographical differences in community assembly mechanisms may explain this pattern, random sampling effects can create this pattern through differences in regional species pools. Here, we compared assembly mechanisms between spatially extensive networks of temperate and tropical forest plots with highly divergent species pools (46 vs. 607 species). After controlling for sampling effects, β-diversity of woody plants was similar and higher than expected by chance in both forests, reflecting strong intraspecific aggregation. However, different mechanisms appeared to explain aggregation in the two forests. In the temperate forest, aggregation reflected stronger environmental correlations, suggesting an important role for species-sorting (e.g. environmental filtering) processes, whereas in the tropics, aggregation reflected stronger spatial correlations, more likely reflecting dispersal limitation. We suggest that biogeographical differences in the relative importance of different community assembly mechanisms contribute to these striking gradients in global biodiversity.

Functional trait space and the latitudinal diversity gradient

Significance We present a conceptual framework for testing theories for the latitudinal gradient of species richness in terms of variation in functional diversity at the alpha, beta, and gamma scales. We compared ecological community theory with large-scale observational data of tree species richness and functional diversity. We found that the patterns of functional trait diversity are not consistent with any one theory of species diversity. These conflicting results indicate that none of the broad classes of biodiversity theory considered here is alone able to explain the latitudinal gradient of species diversity in terms of functional trait space. The processes causing the latitudinal gradient in species richness remain elusive. Ecological theories for the origin of biodiversity gradients, such as competitive exclusion, neutral dynamics, and environmental filtering, make predictions for how functional diversity should vary at the alpha (within local assemblages), beta (among assemblages), and gamma (regional pool) scales. We test these predictions by quantifying hypervolumes constructed from functional traits representing major axes of plant strategy variation (specific leaf area, plant height, and seed mass) in tree assemblages spanning the temperate and tropical New World. Alpha-scale trait volume decreases with absolute latitude and is often lower than sampling expectation, consistent with environmental filtering theory. Beta-scale overlap decays with geographic distance fastest in the temperate zone, again consistent with environmental filtering theory. In contrast, gamma-scale trait space shows a hump-shaped relationship with absolute latitude, consistent with no theory. Furthermore, the overall temperate trait hypervolume was larger than the overall tropical hypervolume, indicating that the temperate zone permits a wider range of trait combinations or that niche packing is stronger in the tropical zone. Although there are limitations in the data, our analyses suggest that multiple processes have shaped trait diversity in trees, reflecting no consistent support for any one theory.

Habitat area and climate stability determine geographical variation in plant species range sizes

Despite being a fundamental aspect of biodiversity, little is known about what controls species range sizes. This is especially the case for hyperdiverse organisms such as plants. We use the largest botanical data set assembled to date to quantify geographical variation in range size for ∼ 85 000 plant species across the New World. We assess prominent hypothesised range-size controls, finding that plant range sizes are codetermined by habitat area and long- and short-term climate stability. Strong short- and long-term climate instability in large parts of North America, including past glaciations, are associated with broad-ranged species. In contrast, small habitat areas and a stable climate characterise areas with high concentrations of small-ranged species in the Andes, Central America and the Brazilian Atlantic Rainforest region. The joint roles of area and climate stability strengthen concerns over the potential effects of future climate change and habitat loss on biodiversity.

Shifts in trait means and variances in North American tree assemblages: species richness patterns are loosely related to the functional space

One of the key hypothesized drivers of gradients in species richness is environmental filtering, where environmental stress limits which species from a larger species pool gain membership in a local community owing to their traits. Whereas most studies focus on small-scale variation in functional traits along environmental gradient, the effect of large-scale environmental filtering is less well understood. Furthermore, it has been rarely tested whether the factors that constrain the niche space limit the total number of coexisting species. We assessed the role of environmental filtering in shaping tree assemblages across North America north of Mexico by testing the hypothesis that colder, drier, or seasonal environments (stressful conditions for most plants) constrain tree trait diversity and thereby limit species richness. We assessed geographic patterns in trait filtering and their relationships to species richness pattern using a comprehensive set of tree range maps. We focused on four key plant functional traits reflecting major life history axes (maximum height, specific leaf area, seed mass, and wood density) and four climatic variables (annual mean and seasonality of temperature and precipitation). We tested for significant spatial shifts in trait means and variances using a null model approach. While we found significant shifts in mean species’ trait values at most grid cells, trait variances at most grid cells did not deviate from the null expectation. Measures of environmental harshness (cold, dry, seasonal climates) and lower species richness were weakly associated with a reduction in variance of seed mass and specific leaf area. The pattern in variance of height and wood density was, however, opposite. These findings do not support the hypothesis that more stressful conditions universally limit species and trait diversity in North America. Environmental filtering does, however, structure assemblage composition, by selecting for certain optimum trait values under a given set of conditions.

Spatial patterns of above-ground structure, biomass and composition in a network of six Andean elevation transects

Background: The Amazon to Andes transition zone provides large expanses of relatively pristine forest wilderness across environmental gradients. Such elevational gradients are an excellent natural laboratory for establishing long-term interactions between forest ecosystems and environmental parameters, which is valuable for understanding ecosystem responses to environmental change. Aims: This study presents data on elevational trends of forest structure (biomass, basal area, height, stem density), species richness, and composition from six elevational transects in the Andes. Methods: We analysed the spatial patterns of forest structure, above-ground biomass and composition from 76 permanent plots, ranging from lowland Amazonian rain forest to high-elevation cloud forests in Ecuador, Peru, and Bolivia. Results: Forest above-ground woody biomass stocks ranged from 247 Mg ha−1 (Peru, 210 m) to 86 Mg ha−1 (Peru, 3450 m), with significantly decreasing trends of tree height and biomass and an increasing trend of stem density with increasing elevation. We observed an increase in forest richness at three taxonomic levels at mid-elevation, followed by a decrease in richness within the cloud immersion zone. Conclusions: The transects show an increase in stem density, a decline in tree height and above-ground coarse wood biomass and a hump-shaped trend in species richness with increasing elevation. These results suggest that environmental change could lead to significant shifts in the properties of these ecosystems over time.

Limited sampling hampers “big data” estimation of species richness in a tropical biodiversity hotspot

Macro-scale species richness studies often use museum specimens as their main source of information. However, such datasets are often strongly biased due to variation in sampling effort in space and time. These biases may strongly affect diversity estimates and may, thereby, obstruct solid inference on the underlying diversity drivers, as well as mislead conservation prioritization. In recent years, this has resulted in an increased focus on developing methods to correct for sampling bias. In this study, we use sample-size-correcting methods to examine patterns of tropical plant diversity in Ecuador, one of the most species-rich and climatically heterogeneous biodiversity hotspots. Species richness estimates were calculated based on 205,735 georeferenced specimens of 15,788 species using the Margalef diversity index, the Chao estimator, the second-order Jackknife and Bootstrapping resampling methods, and Hill numbers and rarefaction. Species richness was heavily correlated with sampling effort, and only rarefaction was able to remove this effect, and we recommend this method for estimation of species richness with “big data” collections.

New Insights into the Evolution of Passiflora subgenus Decaloba (Passifloraceae): Phylogenetic Relationships and Morphological Synapomorphies

Abstract Phylogenetic relationships of Passiflora subgenus Decaloba were examined using 148 taxa and four molecular markers: nuclear nrITS, ncpGS, cp trnL-F, and ndhF. Relationships of subgenus Decaloba to the other four Passiflora subgenera (Astrophea, Deidamioides, Tetrapathea, and Passiflora) are investigated, as are relationships among the eight supersections within subgenus Decaloba. Results indicate that subgenus Deidamioides is not monophyletic. Subgenus Astrophea + subgenus Deidamioides (section Tryphostemmatoides) together form the most basally branching lineage in the genus, followed by a clade comprised of subgenus Passiflora + subgenus Deidamioides (sections Tetrastylis, Polyanthea, and Deidamioides). Passiflora obovata (subgenus Deidamioides section Mayapathanthus) is resolved as part of subgenus Decaloba. The Old World subgenus Tetrapathea is supported as sister to subgenus Decaloba. Subgenus Decaloba is monophyletic and contains seven major lineages that generally correspond to currently recognized supersections. Within subgenus Decaloba, supersection Pterosperma is most basally branching, followed by supersection Hahniopathanthus + P. obovata. The New World species Passiflora multiflora, the type of supersection Multiflora, is resolved as sister to a monophyletic Old World supersection Disemma. The remainder of the former supersection Multiflora is paraphyletic with respect to supersection Auriculata. Supersections Cieca, Bryonioides, and Decaloba are monophyletic. Within supersection Decaloba, two main clades are resolved: 1) section Xerogona + section Decaloba pro parte and 2) the remainder of section Decaloba. The molecular phylogeny supports a New World origin for Passiflora, with two independent radiations to the Old World. Morphological synapomorphies are discussed for the major clades, documenting a pattern of remarkable evolutionary lability in several notable characters.

The discovery of the Amazonian tree flora with an updated checklist of all known tree taxa.

Amazonia is the most biodiverse rainforest on Earth, and the debate over how many tree species grow there remains contentious. Here we provide a checklist of all tree species collected to date, and describe spatial and temporal trends in data accumulation. We report 530,025 unique collections of trees in Amazonia, dating between 1707 and 2015, for a total of 11,676 species in 1225 genera and 140 families. These figures support recent estimates of 16,000 total Amazonian tree species based on ecological plot data from the Amazonian Tree Diversity Network. Botanical collection in Amazonia is characterized by three major peaks, centred around 1840, 1920, and 1980, which are associated with flora projects and the establishment of inventory plots. Most collections were made in the 20th century. The number of collections has increased exponentially, but shows a slowdown in the last two decades. We find that a species’ range size is a better predictor of the number of times it has been collected than the species’ estimated basin-wide population size. Finding, describing, and documenting the distribution of the remaining species will require coordinated efforts at under-collected sites.

Seasonal drought limits tree species across the Neotropics

Within the tropics, the species richness of tree communities is strongly and positively associated with precipitation. Previous research has suggested that this macroecological pattern is driven by the negative effect of water-stress on the physiological processes of most tree species. This process implies that the range limits of taxa are defined by their ability to occur under dry conditions, and thus in terms of species distributions it predicts a nested pattern of taxa distribution from wet to dry areas. However, this ‘dry-tolerance’ hypothesis has yet to be adequately tested at large spatial and taxonomic scales. Here, using a dataset of 531 inventory plots of closed canopy forest distributed across the Western Neotropics we investigated how precipitation, evaluated both as mean annual precipitation and as the maximum climatological water deficit, influences the distribution of tropical tree species, genera and families. We find that the distributions of tree taxa are indeed nested along precipitation gradients in the western Neotropics. Taxa tolerant to seasonal drought are disproportionally widespread across the precipitation gradient, with most reaching even the wettest climates sampled; however, most taxa analysed are restricted to wet areas. Our results suggest that the ‘dry tolerance’ hypothesis has broad applicability in the worlds most species-rich forests. In addition, the large number of species restricted to wetter conditions strongly indicates that an increased frequency of drought could severely threaten biodiversity in this region. Overall, this study establishes a baseline for exploring how tropical forest tree composition may change in response to current and future environmental changes in this region.

Linking environmental filtering and disequilibrium to biogeography with a community climate framework

We present a framework to measure the strength of environmental filtering and disequilibrium of the species composition of a local community across time, relative to past, current, and future climates. We demonstrate the framework by measuring the impact of climate change on New World forests, integrating data for climate niches of more than 14000 species, community composition of 471 New World forest plots, and observed climate across the most recent glacial-interglacial interval. We show that a majority of communities have species compositions that are strongly filtered and are more in equilibrium with current climate than random samples from the regional pool. Variation in the level of current community disequilibrium can be predicted from Last Glacial Maximum climate and will increase with near-future climate change.