Ice age legacies in the geographical distribution of tree species richness in Europe

Aim This study uses a high‐resolution simulation of the Last Glacial Maximum (LGM) climate to assess: (1) whether LGM climate still affects the geographical species richness patterns in the European tree flora and (2) the relative importance of modern and LGM climate as controls of tree species richness in Europe. Location The parts of Europe that were unglaciated during the LGM. Methods Atlas data on the distributions of 55 tree species were linked with data on modern and LGM climate and climatic heterogeneity in a geographical information system with a 60‐km grid. Four measures of species richness were computed: total richness, and richness of the 18 most restricted species, 19 species of medium incidence (intermediate species) and 18 most widespread species. We used ordinary least‐squares regression and spatial autoregressive modelling to test and estimate the richness–climate relationships. Results LGM climate constituted the best single set of explanatory variables for richness of restricted species, while modern climate and climatic heterogeneity was best for total and widespread species richness and richness of intermediate species, respectively. The autoregressive model with all climatic predictors was supported for all richness measures using an information‐theoretic approach, albeit only weakly so for total species richness. Among the strongest relationships were increases in total and intermediate richness with climatic heterogeneity and in restricted richness with LGM growing‐degree‐days. Partial regression showed that climatic heterogeneity accounted for the largest unique variation fraction for intermediate richness, while LGM climate was particularly important for restricted richness. Main conclusions LGM climate appears to still affect geographical patterns of tree species richness in Europe, albeit the relative importance of modern and LGM climate depends on range size. Notably, LGM climate is a strong richness control for species with a restricted range, which appear to still be associated with their glacial refugia.     

Biogeographic anomalies in the species richness of Chilean forests: Incorporating evolution into a climatic – historic scenario

Broad‐scale richness gradients are closely associated with temperature and water availability. However, historical and evolutionary processes have also contributed to shape current diversity patterns. In this paper we focus on the potential influences of Pleistocene glaciation and phylogenetic niche conservatism (the tendency for traits to be maintained during diversification) on the tree diversity gradient in Chile, and we quantify its primary climatic correlates. Tree species richness is greatest at mid latitudes, particularly in the Andes and Coastal ranges, and decreases abruptly to the south and north. Regression tree analysis identified annual precipitation and annual temperature as the primary probable drivers of this gradient. Ice cover during the Last Glacial Maximum was also identified as an ‘important’ variable, but the contemporary and historical predictors are strongly collinear. Geographically weighted regression indicated that the relationships between richness and environmental variables vary regionally: the relationship between tree richness and precipitation is stronger in north‐central Chile, whereas tree richness and temperature are most strongly associated in south‐central Chile. By assigning each species the age of the family to which it belongs and averaging all species in each geographical unit, we also found that species from the oldest families are distributed mainly in mid to high latitudes and species from younger families are distributed mainly at lower latitudes. This pattern is closely associated with annual precipitation. Thus, the ecological component of tree richness follows contemporary climatic gradients of both energy and water, but the aridification of the Atacama Desert was an important driver over evolutionary time. The influence of recent Pleistocene glaciation remains unresolved but it cannot be discounted.     

Species richness of gall-forming insects in a tropical rain forest: correlations with plant diversity and soil fertility

We tested two hypotheses to explain changes in species richness ofgall-forming insects. The first hypothesis proposes that gall-forming insectspecies richness increases as more potential host–plant species areavailable. The second hypothesis implies that soil fertility affects plantcolonization by gall-forming insects. Seven sites, representing strongdifferences in vegetation and soil were chosen at the Lacandona tropical rainforest region, Chiapas, Mexico. Overall, we found 1522 individual plantsbelonging to 340 different plant species. From this, we found gall-forminginsects on 737 (43.9%) plants and on 74 (22%) of total plant species. We found asignificant negative correlation between gall-forming insect species richnessand species richness of plants, which does not support the hypothesis that plantspecies richness is an important factor in generating the radiation ofgall-forming insects. Using phosphorus as an indicator of soil fertility, wefound the lowest number of plants with gall-forming insects and the smallestgall-forming insect load per individual plant in the more fertile soil(alluvial). In contrast, the highest number of plants with galls and the highestgall-forming insect load per plant were found at a savanna-like vegetationsite, where the poorest soil was recorded. These results did not support thesoil fertility hypothesis in terms of species richness, but did with respect toabundance of plants with galls.     

Spatio‐temporal scaling of biodiversity and the species–time relationship in a stream fish assemblage

1. The increase of species richness with the area of the habitat sampled, that is the species–area relationship, and its temporal analogue, the species–time relationship (STR), are among the few general laws in ecology with strong conservation implications. However, these two scale‐dependent phenomena have rarely been considered together in biodiversity assessment, especially in freshwater systems. 2. We examined how the spatial scale of sampling influences STRs for a Central‐European stream fish assemblage (second‐order Bernecei stream, Hungary) using field survey data in two simulation‐based experiments. 3. In experiment one, we examined how increasing the number of channel units, such as riffles and pools (13 altogether), and the number of field surveys involved in the analyses (12 sampling occasions during 3 years), influence species richness. Complete nested curves were constructed to quantify how many species one observes in the community on average for a given number of sampling occasions at a given spatial scale. 4. In experiment two, we examined STRs for the Bernecei fish assemblage from a landscape perspective. Here, we evaluated a 10‐year reach level data set (2000–09) for the Bernecei stream and its recipient watercourse (third‐order Kemence stream) to complement results on experiment one and to explore the mechanisms behind the observed patterns in more detail. 5. Experiment one indicated the strong influence of the spatial scale of sampling on the accumulation of species richness, although time clearly had an additional effect. The simulation methodology advocated here helped to estimate the number of species in a diverse combination of spatial and temporal scale and, therefore, to determine how different scale combinations influence sampling sufficiency. 6. Experiment two revealed differences in STRs between the upstream (Bernecei) and downstream (Kemence) sites, with steeper curves for the downstream site. Equations of STR curves were within the range observed in other studies, predominantly from terrestrial systems. Assemblage composition data suggested that extinction–colonisation dynamics of rare, non‐resident (i.e. satellite) species influenced patterns in STRs. 7. Our results highlight that the determination of species richness can benefit from the joint consideration of spatial and temporal scales in biodiversity inventory surveys. Additionally, we reveal how our randomisation‐based methodology may help to quantify the scale dependency of diversity components (α, β, γ) in both space and time, which have critical importance in the applied context.     

Environment versus dispersal in the assembly of western Amazonian palm communities

Aim It is a central issue in ecology and biogeography to understand what governs community assembly and the maintenance of biodiversity in tropical rain forest ecosystems. A key question is the relative importance of environmental species sorting (niche assembly) and dispersal limitation (dispersal assembly), which we investigate using a large dataset from diverse palm communities. Location Lowland rain forest, western Amazon River Basin, Peru. Methods We inventoried palm communities, registering all palm individuals and recording environmental conditions in 149 transects of 5 m × 500 m. We used ordination, Mantel tests and indicator species analysis (ISA) to assess compositional patterns, species responses to geographical location and environmental factors. Mantel tests were used to assess the relative importance of geographical distance (as a proxy for dispersal limitation) and environmental differences as possible drivers of dissimilarity in palm species composition. We repeated the Mantel tests for subsets of species that differ in traits of likely importance for habitat specialization and dispersal (height and range size). Results We found a strong relationship between compositional dissimilarity and environmental distance and a weaker but also significant relationship between compositional dissimilarity and geographical distance. Consistent with expectations, relationships with environmental and geographical distance were stronger for understorey species than for canopy species. Geographical distance had a higher correlation with compositional dissimilarity for small‐ranged species compared with large‐ranged species, whereas the opposite was true for environmental distance. The main environmental correlates were inundation and soil nutrient levels. Main conclusions The assembly of palm communities in the western Amazon appears to be driven primarily by species sorting according to hydrology and soil, but with dispersal limitation also playing an important role. The importance of environmental characteristics and geographical distance varies depending on plant height and geographical range size in agreement with functional predictions, increasing our confidence in the inferred assembly mechanisms.     

Spatiotemporal patterns and dynamics of species richness and abundance of woody plant functional groups in a tropical forest landscape of Hainan Island, South China

Tropical forests are among the most species-diverse ecosystems on Earth. Their structures and ecological functions are complex to understand. Functional group is defined as a group of species that play similar roles in an ecosystem. The functional group approach has been regarded as an effective way of linking the compositions of complex ecosystems with their ecological functions. To understand the variation of functional groups in species-rich ecosystems after disturbance, the present study investigated the spatial pattern and temporal dynamics of woody plants in a typically fragmented natural forest landscape of Hainan Island in South China. The study area was classified into eight landscape types based on vegetation type, disturbance manner and the time of recovery. The woody plant species were aggregated into seven functional groups based on the growth form, successional status and plant size. The results gained from the present study showed that all functional groups, except for the emergent and canopy tree species, were present in all eight landscape types. Each landscape type had different numbers of dominant functional groups. There are similar species richness and stem abundance structure among functional groups between mid-successional clear cut lowland rainforest and old growth tropical coniferous forest. This similarity exists in selective logged lowland rainforest and old-growth lowland rainforest, as well as among landscape types of montane rainforest. The functional groups with the same successional status had similar patterns of species richness and stem abundance ratios among different landscape types. The variation patterns of functional groups along the successional stages in terms of species richness and stem abundance among the tropical lowland rainforest landscape types were more similar to each other than those in the tropical montane reinforest landscape types. This study provides further support for the competition-colonization tradeoff and successional niche theory as opposed to models of neutrality and ecological equivalence.     

The implications of Tertiary and Quaternary sea level rise events for avian distribution patterns in the lowlands of northern South America

Aim To assess the correspondence between current avian distributions in the lowlands of northern South America with respect to the hypothesized importance of sea level rise and other events over the past 15 million years on speciation. Location Tropical lowlands of north‐western South America. Methods To establish which bird taxa may have originated in each area of endemism, I examined the ranges of all bird species occurring in the Pacific and the Caribbean lowlands. To determine land and sea distribution during a sea level rise of 100 m in north‐western South America and eastern Central America, I traced the 100 m contour line from the Geoatlas of Georama and the Global 30‐Arc‐Second Elevation Data (GTOPO30) produced by the US Geological Survey. Results During a sea level rise of ～100 m, marine incursions would have occurred from the Caribbean Sea and the Pacific Ocean. Several areas of tropical forest and dry/arid vegetation would have been isolated, currently known as the Darién, Chocó, Nechí, Catatumbo, Tumbesian and Guajiran areas of animal and plant endemism. Main conclusions A large part of the high diversity of forest and nonforest birds and other animals and plants in the Pacific rain forest and the Caribbean woodland likely arose as the result of sea level rises, dispersal and other geological and climatic events.     

The more‐individuals hypothesis revisited: the role of community abundance in species richness regulation and the productivity–diversity relationship

Species richness increases with energy availability, yet there is little consensus as to the exact processes driving this species–energy relationship. The most straightforward explanation is the more‐individuals hypothesis (MIH). It states that higher energy availability promotes a higher total number of individuals in a community, which consequently increases species richness by allowing for a greater number of species with viable populations. Empirical support for the MIH is mixed, partially due to the lack of proper formalisation of the MIH and consequent confusion as to its exact predictions. Here, we review the evidence of the MIH and evaluate the reliability of various predictions that have been tested. There is only limited evidence that spatial variation in species richness is driven by variation in the total number of individuals. There are also problems with measures of energy availability, with scale‐dependence, and with the direction of causality, as the total number of individuals may sometimes itself be driven by the number of species. However, even in such a case the total number of individuals may be involved in diversity regulation. We propose a formal theory that encompasses these processes, clarifying how the different factors affecting diversity dynamics can be disentangled.     

The role of climate and plant functional trade‐offs in shaping global biome and biodiversity patterns

Aim Two of the oldest observations in plant geography are the increase in plant diversity from the poles towards the tropics and the global geographic distribution of vegetation physiognomy (biomes). The objective of this paper is to use a process‐based vegetation model to evaluate the relationship between modelled and observed global patterns of plant diversity and the geographic distribution of biomes. Location The global terrestrial biosphere. Methods We implemented and tested a novel vegetation model aimed at identifying strategies that enable plants to grow and reproduce within particular climatic conditions across the globe. Our model simulates plant survival according to the fundamental ecophysiological processes of water uptake, photosynthesis, reproduction and phenology. We evaluated the survival of an ensemble of 10,000 plant growth strategies across the range of global climatic conditions. For the simulated regional plant assemblages we quantified functional richness, functional diversity and functional identity. Results A strong relationship was found (correlation coefficient of 0.75) between the modelled and the observed plant diversity. Our approach demonstrates that plant functional dissimilarity increases and then saturates with increasing plant diversity. Six of the major Earth biomes were reproduced by clustering grid cells according to their functional identity (mean functional traits of a regional plant assemblage). These biome clusters were in fair agreement with two other global vegetation schemes: a satellite image classification and a biogeography model (kappa statistics around 0.4). Main conclusions Our model reproduces the observed global patterns of plant diversity and vegetation physiognomy from the number and identity of simulated plant growth strategies. These plant growth strategies emerge from the first principles of climatic constraints and plant functional trade‐offs. Our study makes important contributions to furthering the understanding of how climate affects patterns of plant diversity and vegetation physiognomy from a process‐based rather than a phenomenological perspective.     

Soil and vegetation features determine the nested pattern of ant–plant networks in a tropical rainforest

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Predicting shifts in the functional composition of tropical forests under increased drought and CO2 from trade‐offs among plant hydraulic traits

Tropical forest responses are an important feedback on global change, but changes in forest composition with projected increases in CO₂ and drought are highly uncertain. Here we determine shifts in the most competitive plant hydraulic strategy (the evolutionary stable strategy or ESS) from changes in CO₂ and drought frequency and intensity. Hydraulic strategies were defined along a spectrum from drought avoidance to tolerance by physiology traits. Drought impacted competition more than CO₂, with elevated CO₂ reducing but not reversing drought‐induced shifts in the ESS towards more tolerant strategies. Trait plasticity and/or adaptation intensified these shifts by increasing the competitive ability of the drought tolerant relative to the avoidant strategies. These findings predict losses of drought avoidant evergreens from tropical forests under global change, and point to the importance of changes in precipitation during the dry season and constraints on plasticity and adaptation in xylem traits to forest responses.     

Foliar Extrafloral Nectar of Humboldtia brunonis (Fabaceae), a Paleotropic Ant‐plant,is Richer than Phloem Sap and More Attractive than Honeydew

The ant‐plant Humboldtia brunonis secretes extrafloral nectar (EFN) despite the lack of antiherbivore protection from most ants. EFN was richer in composition than phloem sap and honeydew from untended Hemiptera on the plant, suggesting that EFN could potentially distract ants from honeydew, since ants rarely tended Hemiptera on this plant.     

Signatures of competition and strain structure within the major blood‐stage antigen of Plasmodium falciparum in a local community in Ghana

The concept of niche partitioning has received considerable theoretical attention at the interface of ecology and evolution of infectious diseases. Strain theory postulates that pathogen populations can be structured into distinct nonoverlapping strains by frequency‐dependent selection in response to intraspecific competition for host immune space. The malaria parasite Plasmodium falciparum presents an opportunity to investigate this phenomenon in nature, under conditions of high recombination rate and extensive antigenic diversity. The parasite's major blood‐stage antigen, PfEMP1, is encoded by the hyperdiverse var genes. With a dataset that includes thousands of var DBLα sequence types sampled from asymptomatic cases within an area of high endemicity in Ghana, we address how var diversity is distributed within isolates and compare this to the distribution of microsatellite allelic diversity within isolates to test whether antigenic and neutral regions of the genome are structured differently. With respect to var DBLα sequence types, we find that on average isolates exhibit significantly lower overlap than expected randomly, but that there also exists frequent pairs of isolates that are highly related. Furthermore, the linkage network of var DBLα sequence types reveals a pattern of nonrandom modularity unique to these antigenic genes, and we find that modules of highly linked DBLα types are not explainable by neutral forces related to var recombination constraints, microsatellite diversity, sampling location, host age, or multiplicity of infection. These findings of reduced overlap and modularity among the var antigenic genes are consistent with a role for immune selection as proposed by strain theory. Identifying the evolutionary and ecological dynamics that are responsible for the nonrandom structure in P. falciparum antigenic diversity is important for designing effective intervention in endemic areas.