Wolf L. Eiserhardt

Geographical ecology of the palms (Arecaceae): determinants of diversity and distributions across spatial scales

BACKGROUND The palm family occurs in all tropical and sub-tropical regions of the world. Palms are of high ecological and economical importance, and display complex spatial patterns of species distributions and diversity. SCOPE This review summarizes empirical evidence for factors that determine palm species distributions, community composition and species richness such as the abiotic environment (climate, soil chemistry, hydrology and topography), the biotic environment (vegetation structure and species interactions) and dispersal. The importance of contemporary vs. historical impacts of these factors and the scale at which they function is discussed. Finally a hierarchical scale framework is developed to guide predictor selection for future studies. CONCLUSIONS Determinants of palm distributions, composition and richness vary with spatial scale. For species distributions, climate appears to be important at landscape and broader scales, soil, topography and vegetation at landscape and local scales, hydrology at local scales, and dispersal at all scales. For community composition, soil appears important at regional and finer scales, hydrology, topography and vegetation at landscape and local scales, and dispersal again at all scales. For species richness, climate and dispersal appear to be important at continental to global scales, soil at landscape and broader scales, and topography at landscape and finer scales. Some scale-predictor combinations have not been studied or deserve further attention, e.g. climate on regional to finer scales, and hydrology and topography on landscape and broader scales. The importance of biotic interactions - apart from general vegetation structure effects - for the geographic ecology of palms is generally underexplored. Future studies should target scale-predictor combinations and geographic domains not studied yet. To avoid biased inference, one should ideally include at least all predictors previously found important at the spatial scale of investigation.

Cenozoic imprints on the phylogenetic structure of palm species assemblages worldwide

Despite long-standing interest in the origin and maintenance of species diversity, little is known about historical drivers of species assemblage structure at large spatiotemporal scales. Here, we use global species distribution data, a dated genus-level phylogeny, and paleo-reconstructions of biomes and climate to examine Cenozoic imprints on the phylogenetic structure of regional species assemblages of palms (Arecaceae), a species-rich plant family characteristic of tropical ecosystems. We find a strong imprint on phylogenetic clustering due to geographic isolation and in situ diversification, especially in the Neotropics and on islands with spectacular palm radiations (e.g., Madagascar, Hawaii, and Cuba). Phylogenetic overdispersion on mainlands and islands corresponds to biotic interchange areas. Differences in the degree of phylogenetic clustering among biogeographic realms are related to differential losses of tropical rainforests during the Cenozoic, but not to the cumulative area of tropical rainforest over geological time. A largely random phylogenetic assemblage structure in Africa coincides with severe losses of rainforest area, especially after the Miocene. More recent events also appear to be influential: phylogenetic clustering increases with increasing intensity of Quaternary glacial-interglacial climatic oscillations in South America and, to a lesser extent, Africa, indicating that specific clades perform better in climatically unstable regions. Our results suggest that continental isolation (in combination with limited long-distance dispersal) and changing climate and habitat loss throughout the Cenozoic have had strong impacts on the phylogenetic structure of regional species assemblages in the tropics.

A set of 100 chloroplast DNA primer pairs to study population genetics and phylogeny in monocotyledons.

Chloroplast DNA sequences are of great interest for population genetics and phylogenetic studies. However, only a small set of markers are commonly used. Most of them have been designed for amplification in a large range of Angiosperms and are located in the Large Single Copy (LSC). Here we developed a new set of 100 primer pairs optimized for amplification in Monocotyledons. Primer pairs amplify coding (exon) and non-coding regions (intron and intergenic spacer). They span the different chloroplast regions: 72 are located in the LSC, 13 in the Small Single Copy (SSC) and 15 in the Inverted Repeat region (IR). Amplification and sequencing were tested in 13 species of Monocotyledons: Dioscorea abyssinica, D. praehensilis, D. rotundata, D. dumetorum, D. bulbifera, Trichopus sempervirens (Dioscoreaceae), Phoenix canariensis, P. dactylifera, Astrocaryum scopatum, A. murumuru, Ceroxylon echinulatum (Arecaceae), Digitaria excilis and Pennisetum glaucum (Poaceae). The diversity found in Dioscorea, Digitaria and Pennisetum mainly corresponded to Single Nucleotide Polymorphism (SNP) while the diversity found in Arecaceae also comprises Variable Number Tandem Repeat (VNTR). We observed that the most variable loci (rps15-ycf1, rpl32-ccsA, ndhF-rpl32, ndhG-ndhI and ccsA) are located in the SSC. Through the analysis of the genetic structure of a wild-cultivated species complex in Dioscorea, we demonstrated that this new set of primers is of great interest for population genetics and we anticipate that it will also be useful for phylogeny and bar-coding studies.

Testing the Water-Energy Theory on American Palms (Arecaceae) Using Geographically Weighted Regression

Water and energy have emerged as the best contemporary environmental correlates of broad-scale species richness patterns. A corollary hypothesis of water–energy dynamics theory is that the influence of water decreases and the influence of energy increases with absolute latitude. We report the first use of geographically weighted regression for testing this hypothesis on a continuous species richness gradient that is entirely located within the tropics and subtropics. The dataset was divided into northern and southern hemispheric portions to test whether predictor shifts are more pronounced in the less oceanic northern hemisphere. American palms (Arecaceae, n = 547 spp.), whose species richness and distributions are known to respond strongly to water and energy, were used as a model group. The ability of water and energy to explain palm species richness was quantified locally at different spatial scales and regressed on latitude. Clear latitudinal trends in agreement with water–energy dynamics theory were found, but the results did not differ qualitatively between hemispheres. Strong inherent spatial autocorrelation in local modeling results and collinearity of water and energy variables were identified as important methodological challenges. We overcame these problems by using simultaneous autoregressive models and variation partitioning. Our results show that the ability of water and energy to explain species richness changes not only across large climatic gradients spanning tropical to temperate or arctic zones but also within megathermal climates, at least for strictly tropical taxa such as palms. This finding suggests that the predictor shifts are related to gradual latitudinal changes in ambient energy (related to solar flux input) rather than to abrupt transitions at specific latitudes, such as the occurrence of frost.

Climate‐driven extinctions shape the phylogenetic structure of temperate tree floras

When taxa go extinct, unique evolutionary history is lost. If extinction is selective, and the intrinsic vulnerabilities of taxa show phylogenetic signal, more evolutionary history may be lost than expected under random extinction. Under what conditions this occurs is insufficiently known. We show that late Cenozoic climate change induced phylogenetically selective regional extinction of northern temperate trees because of phylogenetic signal in cold tolerance, leading to significantly and substantially larger than random losses of phylogenetic diversity (PD). The surviving floras in regions that experienced stronger extinction are phylogenetically more clustered, indicating that non-random losses of PD are of increasing concern with increasing extinction severity. Using simulations, we show that a simple threshold model of survival given a physiological trait with phylogenetic signal reproduces our findings. Our results send a strong warning that we may expect future assemblages to be phylogenetically and possibly functionally depauperate if anthropogenic climate change affects taxa similarly.

Dispersal and niche evolution jointly shape the geographic turnover of phylogenetic clades across continents

The turnover of phylogenetic clades across space is a fundamental biodiversity pattern that may depend on long-term evolutionary processes, and that has downstream effects on other aspects of diversity including species richness and community structure. Limited niche evolution and limited dispersal are two major processes causing spatial restriction, and thus turnover, of clades. We studied the determinants of clade turnover within the Worlds richest floristic kingdom, the Neotropics, using the palm family (Arecaceae) as a model. We show that continental-scale clade turnover is driven by a combination of limited niche evolution — with respect to temperature and soil tolerances — and limited dispersal. These findings are consistent with strong dispersal barriers within the Neotropics, and the observation that some palm lineages are most diverse in certain biomes or climates. The importance of such deep-time effects suggest that palms might be slow to adapt or disperse in response to anthropogenic climate change.

An all-evidence species-level supertree for the palms (Arecaceae)

Several attempts have been made to generate complete species-level phylogenies for large clades, enabling comprehensive analyses of ecological or evolutionary hypotheses at the species level. No such phylogeny has, however, been generated for any major plant group yet, but here we generate such a phylogeny for the palm family (Arecaceae). We do this using a novel Bayesian approach, estimating the validity of intra-generic taxonomic groupings as topological constraints to assist in placing species without genetic or morphological data. From these we implement those that are supported by genetic or morphological data for a given genus or for related genera. The intergeneric relationships in our new phylogeny are surprisingly different from earlier phylogenies in the placement of genera within tribes, but largely identical to previous findings in the deeper branches in the phylogeny, pointing to the need for incorporating phylogenetic uncertainty in analyses based on this phylogeny. Initial analyses of the new phylogeny suggest non-constancy in diversification rates over time within genera, with an apparent increase in diversification rate over time, but no evidence for any geographic variation in the magnitude of this increase. We hope that our study will stimulate further evolutionary or ecological studies using palms as study organisms as well as discussions of the optimal way to place the many species without genetic or morphological data.

Assembly of forest communities across East Asia - insights from phylogenetic community structure and species pool scaling

Local communities are assembled from larger-scale species pools via dispersal, environmental filtering, biotic interactions, and local stochastic demographic processes. The relative importance, scaling and interplay of these assembly processes can be elucidated by comparing local communities to variously circumscribed species pools. Here we present the first study applying this approach to forest tree communities across East Asia, focusing on community phylogenetic structure and using data from a global network of tropical, subtropical and temperate forest plots. We found that Net Relatedness Index (NRI) and Nearest Taxon Index (NTI) values were generally lower with geographically broad species pools (global and Asian species pools) than with an East Asian species pool, except that global species pool produced higher NTI than the East Asian species pool. The lower NRI for the global relative to the East Asian species pool may indicate an important role of intercontinental migration during the Neogene and Quaternary and climatic conservatism in shaping the deeper phylogenetic structure of tree communities in East Asia. In contrast, higher NTI for the global relative to the East Asian species pool is consistent with recent localized diversification determining the shallow phylogenetic structure.

Strong effects of variation in taxonomic opinion on diversification analyses

Summary The effects of variation in taxonomic opinion between ‘splitters’ and ‘lumpers’ have been debated intensely in conservation biology and have also been discussed in macroecology. However, the impact on diversification analyses has received little attention and has largely been ignored by many end-users of macroevolutionary analyses. Using simulated data sets, we analysed the effects of variation in taxonomic opinion on lineage diversification analysis, focusing on two measures of phylogenetic asymmetry (β and IC), a measure of the relative age composition of nodes (γ), and the MEDUSA algorithm, which searches for clades with unique diversification histories. All measures were biased by variation in taxonomic opinion, but to different degrees. The γ-estimate and the MEDUSA algorithm were found to be especially sensitive, with error rates high enough to make it likely that false increases in diversification rates over time or clades with apparently, but not actually, unique diversification histories would be present in many published analyses. In contrast, for the two measures of phylogenetic asymmetry the biases were likely too small to substantially change conclusions in studies ignoring variation in taxonomic opinion. Our results highlight that variation in taxonomic opinion can be an important source of error in diversification analyses. We therefore suggest that all end-users of diversification analyses should consider whether the approaches used are sensitive to this issue and, if so, whether any of their results could be a consequence of different taxonomic treatment of different lineages rather than real biological differences.

Snow cover consistently affects growth and reproduction of Empetrum hermaphroditum across latitudinal and local climatic gradients

Arctic ecosystems face strong changes in snow conditions due to global warming. In contrast to habitat specialists, species occupying a wide range of microhabitats under different snow conditions may better cope with such changes. We studied how growth and reproduction of the dominant dwarf shrub Empetrum hermaphroditum varied among three habitat types differing in winter snow depth and summer irradiation, and whether the observed patterns were consistent along a local climatic gradient (sub-continental vs. sub-oceanic climates) and along a latitudinal gradient (northern Sweden vs. central Norway). Habitat type explained most of the variation in growth and reproduction. Shoots from shallow snow cover and high summer irradiation habitats had higher numbers of flowers and fruits, lower ramet heights, shorter shoot segments, lower numbers of lateral shoots and total biomass but higher leaf density and higher relative leaf allocation than shoots from habitats with higher snow depth and lower summer irradiation. In addition, biomass, leaf allocation and leaf life expectancy were strongly affected by latitude, whereas local climate had strong effects on seed number and seed mass. Empetrum showed high phenotypic trait variation, with a consistent match between local habitat conditions and its growth and reproduction. Although study areas varied strongly with respect to latitude and local climatic conditions, response patterns of growth and reproduction to habitats with different environmental conditions were consistent. Large elasticity of traits suggests that Empetrum may have the potential to cope with changing snow conditions expected in the course of climate change.