Finn Borchsenius

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.

Twenty-five years of progress in understanding pollination mechanisms in palms (Arecaceae)

BACKGROUND With more than 90 published studies of pollination mechanisms, the palm family is one of the better studied tropical families of angiosperms. Understanding palm-pollinator interactions has implications for tropical silviculture, agroforestry and horticulture, as well as for our understanding of palm evolution and diversification. We review the rich literature on pollination mechanisms in palms that has appeared since the last review of palm pollination studies was published 25 years ago. SCOPE AND CONCLUSIONS Visitors to palm inflorescences are attracted by rewards such as food, shelter and oviposition sites. The interaction between the palm and its visiting fauna represents a trade-off between the services provided by the potential pollinators and the antagonistic activities of other insect visitors. Evidence suggests that beetles constitute the most important group of pollinators in palms, followed by bees and flies. Occasional pollinators include mammals (e.g. bats and marsupials) and even crabs. Comparative studies of palm-pollinator interactions in closely related palm species document transitions in floral morphology, phenology and anatomy correlated with shifts in pollination vectors. Synecological studies show that asynchronous flowering and partitioning of pollinator guilds may be important regulators of gene flow between closely related sympatric taxa and potential drivers of speciation processes. Studies of larger plant-pollinator networks point out the importance of competition for pollinators between palms and other flowering plants and document how the insect communities in tropical forest canopies probably influence the reproductive success of palms. However, published studies have a strong geographical bias towards the South American region and a taxonomic bias towards the tribe Cocoseae. Future studies should try to correct this imbalance to provide a more representative picture of pollination mechanisms and their evolutionary implications across the entire family.

Patterns of plant species endemism in Ecuador

Distribution patterns of plant species endemic to Ecuador and adjacent parts of southern Colombia and northern Peru are analysed on the basis of information in the Flora of Ecuador. A total of 827 restricted-range species were found, many of which are known from extremely small areas, often only one or a few localities. A total of 27% of the species treated in the Flora of Ecuador are endemic to that country. The overall proportion of endemic and restricted-range species is greater in the Andes than in the lowland areas on either side of these mountains; particularly the southern Andes appears to be very rich in endemic species. Spatial analysis of distribution data results in the recognition of 15 floristic elements and 18 geographical endemism regions in Ecuador, the characteristics of which are discussed. Comparison with distribution patterns of restricted-range bird species show a general correspondence, with the main difference that birds tend to be more widely distributed than plants along the Andes. Comparison of the results with the location of national parks and other protected areas shows that the endemic floras in the northern and eastern parts of the country are much better protected than those of the southern and western parts.

Landscape genetics, historical isolation and cross-Andean gene flow in the wax palm, Ceroxylon echinulatum (Arecaceae)

Knowledge of the role of landscapes in shaping genetic connectivity and divergence is essential for understanding patterns of biogeography and diversity. This is particularly relevant for the Andes region, a major biodiversity hotspot of relatively recent origin. We examined the phylogeography and landscape genetics of the Andean wax palm Ceroxylon echinulatum (Arecaceae) that occurs in two narrow bands of montane forests on each side of the Andes in Ecuador and northeastern Peru. First, we tested the hypothesis of C. echinulatum being a geographic cline species crossing the Andes in the Amotape–Huancabamba zone (AHZ) of southern Ecuador/northern Peru, as indicated by observations on fruit morphology. Second, we assessed the timeframe of cross‐Andean divergence, and third, we investigated the impact of contemporary and historical landscape features on observed spatio‐genetic patterns. Individual‐based Bayesian clustering (BC) identified a northeastern, southeastern, southwestern, and northwestern cluster, with areas of genetic discontinuity coinciding with the Andes and the Giron–Paute deflection. F‐statistics derived from BC suggested an east‐to‐west dispersal history. Population‐based analyses revealed strong genetic structuring at both small and large geographic scales. Interpopulation relationships and Mantel tests strongly supported the cline model with cross‐Andean dispersal in the AHZ. Along the cline, gene flow measured as FST was mainly limited by distance, with less but significant impact of climatic friction. Coalescent analysis revealed that cross‐Andean divergence took place during the Quaternary. Significant historical isolation (RST > FST) was found in the southwestern population. The current study illustrates a joint effect of founder dynamics, divergence by distance and historical isolation on patterns of Andean diversity and distribution.

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.

Vegetation structure and diversity of an ancient temperate deciduous forest in SW Denmark

Bolderslev Skov (113 ha) is the largest contiguous ancient forest remnant in Denmark. The forest has been preserved since 1998 as a strict non-intervention forest reserve. We studied vegetation structure, floristic gradients, and diversity of the forest in 50 plots of 100 m2 placed according to a restricted random sampling design. Dominant tree species were Fagus sylvatica, Fraxinus excelsior, Tilia cordata and Quercus robur, which in most parts of the forest form a mixed canopy. Most stands appeared to be of moderate age, 55-80 years old, and large old trees were rare. pH in association with light and thickness of the litter layer were the most important factors in explaining floristic gradients in the forest. Soil moisture (dry to mesic) was not strongly correlated with DCA axes for neither tree nor field layer, but had a significant effect on the distribution of a number of herb species. Forest structure was not important in explaining species distributions. Field layer species richness showed a positive relationship with the pH-gradient. At the scale of 1m2 plots we also found a highly significant negative relationship between field layer species richness and the plot-wise Ellenberg indicator value for nitrogen availability. Structure of the tree layer had little effect on field layer species richness. The mixed composition, long continuity, and presence of a high proportion of the regional species pool assigns a high conservation value to Bolderslev Skov and makes it an important site for future studies of the dynamic properties, niche preferences, and inter-specific competition of temperate deciduous forest species.

Flowering biology ofGeonoma irena andG. cuneata var.sodiroi (Arecaceae)

The phenology and flowering of two sympatric understory palms from western Ecuador, until recently regarded as conspecific, is described.Geonoma irenaBorchs. flowers throughout the year. Anthesis lasts 11–14 weeks per inflorescence, with overlapping male and female phases. Flowers open in the midmorning, and are visited mainly by meliponid and halictid bees.Geonoma cuneata var.sodiroi (Burret)Skov flowers from December to April, with a distinct peak in Feburary. Anthesis lasts 6–8 days per inflorescence, with non-overlapping male and female phases. Flowers open at dawn, and are visited mainly by drosophilid and sphaerocerid flies. The flowering pattern of the two species match different specific behavioural features of their insect visitors. The study provides an example of how differences in reproductive biology may act as an important barrier to gene flow between related, co-occurring taxa.

Molecular Phylogeny and Redefined Generic Limits of Calathea (Marantaceae)

Abstract Calathea, with an estimated 285 species, is the largest genus of Marantaceae and an important component of Neotropical herbaceous diversity. The genus is also of high importance for horticulture as species are cultivated for their showy, patterned leaves. Previous molecular phylogenetic studies indicated that the genus is polyphyletic, but have not provided a basis for redefining generic limits due to incomplete taxon sampling. To address this problem we analyzed DNA sequence data from three plastid markers (matK with flanking 3′ trnK intron, trnL intron and trnL-trnF intergenic spacer) and one nuclear marker (ITS) under a maximum parsimony criterion for a large and representative taxon sample covering all previously proposed infrageneric entities, and representing the full range of morphological variation known in the genus. Our results confirm that Calathea is polyphyletic. One clade, including subgenus Calathea, the C. lanicaulis group, and the genus Sanblasia, is sister to a clade formed by Ischnosiphon and Pleiostachya. The genus Monotagma is placed as sister to this clade. The remaining species form a second strongly supported clade as sister to a clade containing these other genera. Based on these findings Calathea is recircumscribed in a narrow sense and Sanblasia is placed in synonymy. The genus Goeppertia is resurrected and redefined to include all members of the second Calathea clade. Morphological characters defining each genus are provided. A total of 246 new combinations are made.

Contrasting palm species and use diversity in the Yucatan Peninsula and the Ecuadorian Amazon

We analyze the relationship between palm species diversity and diversity of palm use in two areas (Amazonian Ecuador; Yucatan Peninsula) of equivalent size but with contrasting characteristics in palm species diversity and morphology, and in the phylogenetic composition of palm flora. The areas also differ in their cultural and socioeconomic contexts. Palm use diversity is significantly higher in the Ecuadorian Amazon than in the Yucatan Peninsula and the lower species diversity of palms in the Yucatan Peninsula is not matched by a more intensive use of the fewer species found there. The taxonomic composition of the palm flora is a poor predictor of extent of use and morphological characteristics of palm species affect usefulness only in the Ecuadorian Amazon. The Yucatecans’ more limited reliance on forest products might explain the observed patterns. Ornamental palm use is an indicator of a general change in plant use patterns associated with tourism and macro-economic development in the Yucatan Peninsula. We find a positive relationship between ecosystem plant diversity and plant use diversity, but socioeconomic factors such as market integration strongly influence the use of local biodiversity. Palms represent a diverse and important natural resource that deserves further investigation to secure its sustainable management and conservation in the two studied sites, regardless of their degree of market integration.

The effect of nitrogen deposition on the species richness of acid grasslands in Denmark: a comparison with a study performed on a European scale.

The effect of atmospheric nitrogen deposition on the species richness of acid grasslands was investigated by combining data from a large Danish monitoring program with a large European data set, where a significant non-linear negative effect of nitrogen deposition had been demonstrated (Stevens et al., 2010). The nitrogen deposition range in Denmark is relatively small and when only considering the Danish data a non-significant decrease in the species richness with nitrogen deposition was observed. However, when both data sets were combined, then the conclusion of the European survey was further corroborated by the results of the Danish monitoring. Furthermore, by combining the two data sets a more comprehensive picture of the threats to the biodiversity of acid grasslands emerge; i.e., species richness in remnant patches of acid grassland in intensively cultivated agricultural landscapes is under influence not only from nitrogen deposition, but also from current and historical land use.