Marc S. M. Sosef

Molecular phylogenetics reveal multiple tertiary vicariance origins of the African rain forest trees

BackgroundTropical rain forests are the most diverse terrestrial ecosystems on the planet. How this diversity evolved remains largely unexplained. In Africa, rain forests are situated in two geographically isolated regions: the West-Central Guineo-Congolian region and the coastal and montane regions of East Africa. These regions have strong floristic affinities with each other, suggesting a former connection via an Eocene pan-African rain forest. High levels of endemism observed in both regions have been hypothesized to be the result of either 1) a single break-up followed by a long isolation or 2) multiple fragmentation and reconnection since the Oligocene. To test these hypotheses the evolutionary history of endemic taxa within a rain forest restricted African lineage of the plant family Annonaceae was studied. Molecular phylogenies and divergence dates were estimated using a Bayesian relaxed uncorrelated molecular clock assumption accounting for both calibration and phylogenetic uncertainties.ResultsOur results provide strong evidence that East African endemic lineages of Annonaceae have multiple origins dated to significantly different times spanning the Oligocene and Miocene epochs. Moreover, these successive origins (c. 33, 16 and 8 million years – Myr) coincide with known periods of aridification and geological activity in Africa that would have recurrently isolated the Guineo-Congolian rain forest from the East African one. All East African taxa were found to have diversified prior to Pleistocene times.ConclusionMolecular phylogenetic dating analyses of this large pan-African clade of Annonaceae unravels an interesting pattern of diversification for rain forest restricted trees co-occurring in West/Central and East African rain forests. Our results suggest that repeated reconnections between the West/Central and East African rain forest blocks allowed for biotic exchange while the break-ups induced speciation via vicariance, enhancing the levels of endemicity. These results provide an explanation for present day distribution patterns and origins of endemicity for African rain forest trees. Moreover, given the pre-Pleistocene origins of all the studied endemic East African genera and species, these results also offer important insights for setting conservation priorities in these highly diversified but threatened ecosystems.

Exploring the floristic diversity of tropical Africa

BackgroundUnderstanding the patterns of biodiversity distribution and what influences them is a fundamental pre-requisite for effective conservation and sustainable utilisation of biodiversity. Such knowledge is increasingly urgent as biodiversity responds to the ongoing effects of global climate change. Nowhere is this more acute than in species-rich tropical Africa, where so little is known about plant diversity and its distribution. In this paper, we use RAINBIO – one of the largest mega-databases of tropical African vascular plant species distributions ever compiled – to address questions about plant and growth form diversity across tropical Africa.ResultsThe filtered RAINBIO dataset contains 609,776 georeferenced records representing 22,577 species. Growth form data are recorded for 97% of all species. Records are well distributed, but heterogeneous across the continent. Overall, tropical Africa remains poorly sampled. When using sampling units (SU) of 0.5°, just 21 reach appropriate collection density and sampling completeness, and the average number of records per species per SU is only 1.84. Species richness (observed and estimated) and endemism figures per country are provided. Benin, Cameroon, Gabon, Ivory Coast and Liberia appear as the botanically best-explored countries, but none are optimally explored. Forests in the region contain 15,387 vascular plant species, of which 3013 are trees, representing 5–7% of the estimated world’s tropical tree flora. The central African forests have the highest endemism rate across Africa, with approximately 30% of species being endemic.ConclusionsThe botanical exploration of tropical Africa is far from complete, underlining the need for intensified inventories and digitization. We propose priority target areas for future sampling efforts, mainly focused on Tanzania, Atlantic Central Africa and West Africa. The observed number of tree species for African forests is smaller than those estimated from global tree data, suggesting that a significant number of species are yet to be discovered. Our data provide a solid basis for a more sustainable management and improved conservation of tropical Africa’s unique flora, and is important for achieving Objective 1 of the Global Strategy for Plant Conservation 2011–2020.

RAINBIO: a mega-database of tropical African vascular plants distributions

Abstract The tropical vegetation of Africa is characterized by high levels of species diversity but is undergoing important shifts in response to ongoing climate change and increasing anthropogenic pressures. Although our knowledge of plant species distribution patterns in the African tropics has been improving over the years, it remains limited. Here we present RAINBIO, a unique comprehensive mega-database of georeferenced records for vascular plants in continental tropical Africa. The geographic focus of the database is the region south of the Sahel and north of Southern Africa, and the majority of data originate from tropical forest regions. RAINBIO is a compilation of 13 datasets either publicly available or personal ones. Numerous in depth data quality checks, automatic and manual via several African flora experts, were undertaken for georeferencing, standardization of taxonomic names and identification and merging of duplicated records. The resulting RAINBIO data allows exploration and extraction of distribution data for 25,356 native tropical African vascular plant species, which represents ca. 89% of all known plant species in the area of interest. Habit information is also provided for 91% of these species.

Phylogenetics, ancestral state reconstruction, and a new infrafamilial classification of the pantropical Ochnaceae (Medusagynaceae, Ochnaceae s.str., Quiinaceae) based on five DNA regions

Ochnaceae s.str. (Malpighiales) are a pantropical family of about 500 species and 27 genera of almost exclusively woody plants. Infrafamilial classification and relationships have been controversial partially due to the lack of a robust phylogenetic framework. Including all genera except Indosinia and Perissocarpa and DNA sequence data for five DNA regions (ITS, matK, ndhF, rbcL, trnL-F), we provide for the first time a nearly complete molecular phylogenetic analysis of Ochnaceae s.l. resolving most of the phylogenetic backbone of the family. Based on this, we present a new classification of Ochnaceae s.l., with Medusagynoideae and Quiinoideae included as subfamilies and the former subfamilies Ochnoideae and Sauvagesioideae recognized at the rank of tribe. Our data support a monophyletic Ochneae, but Sauvagesieae in the traditional circumscription is paraphyletic because Testulea emerges as sister to the rest of Ochnoideae, and the next clade shows Luxemburgia+Philacra as sister group to the remaining Ochnoideae. To avoid paraphyly, we classify Luxemburgieae and Testuleeae as new tribes. The African genus Lophira, which has switched between subfamilies (here tribes) in past classifications, emerges as sister to all other Ochneae. Thus, endosperm-free seeds and ovules with partly to completely united integuments (resulting in an apparently single integument) are characters that unite all members of that tribe. The relationships within its largest clade, Ochnineae (former Ochneae), are poorly resolved, but former Ochninae (Brackenridgea, Ochna) are polyphyletic. Within Sauvagesieae, the genus Sauvagesia in its broad circumscription is polyphyletic as Sauvagesia serrata is sister to a clade of Adenarake, Sauvagesia spp., and three other genera. Within Quiinoideae, in contrast to former phylogenetic hypotheses, Lacunaria and Touroulia form a clade that is sister to Quiina. Bayesian ancestral state reconstructions showed that zygomorphic flowers with adaptations to buzz-pollination (poricidal anthers), a syncarpous gynoecium (a near-apocarpous gynoecium evolved independently in Quiinoideae and Ochninae), numerous ovules, septicidal capsules, and winged seeds with endosperm are the ancestral condition in Ochnoideae. Although in some lineages poricidal anthers were lost secondarily, the evolution of poricidal superstructures secured the maintenance of buzz-pollination in some of these genera, indicating a strong selective pressure on keeping that specialized pollination system.

Revision of the African Genus Hexalobus (Annonaceae)

Abstract The genus Hexalobus consists of five species characterized by six basally fused petal lobes that are transversally folded in bud, which is unique within Annonaceae. The genus is widespread across tropical Africa occurring in rain forest, savanna, and woodland. The species H. callicarpus, occurring in Madagascar, is excluded from Hexalobus, so the genus is now restricted to continental Africa. The present revision provides a synthesis of previously published information and discussions on morphology, taxonomy, phylogeny, and palynology. Conservation status assessments are provided for each species, as well as a diagnostic key and detailed species descriptions. Furthermore, four species are illustrated and all species are mapped.

Morphological characterization of African bush mango trees (Irvingia species) in West Africa

The variation of the morphological characters of bitter and sweet African bush mango trees (Irvingia species) was investigated in the Dahomey Gap which is the West African savannah woodland area separating the Upper and the Lower Guinean rain forest blocks. African bush mangoes have been rated as the highest priority multi-purpose food trees species that need improvement research in West and Central Africa. A total of 128 trees from seven populations were characterized for their bark, fruits, mesocarp and seeds to assess the morphological differences between bitter and sweet trees and among populations. Multivariate analysis revealed that none of the variables: type of bark, mature fruit exocarp colour, fruit roughness and fresh mesocarp colour, could consistently distinguish bitter from sweet trees in the field. The analysis of the measurements of fruits, mesocarps and seeds demonstrated that bitter fruits have the heaviest seeds and this consistently distinguishes them from sweet fruits. However, the measurements of the fruit, mesocarp and seed did not have a joint effect in grouping types and populations of ABMTs. This indicates high diversity with a potential for selection existing across all phytogeographical regions investigated. The sweet trees of Couffo and those of Dassa in Benin are clearly different from all other populations. This can be attributed to traditional domestication (bringing into cultivation) and climate, respectively. The large fruits and the heavy seeds of the cultivated populations are evidence of successful on-going traditional selection of sweet trees in the Dahomey Gap.

Insights into the Influence of Priors in Posterior Mapping of Discrete Morphological Characters: A Case Study in Annonaceae

Background Posterior mapping is an increasingly popular hierarchical Bayesian based method used to infer character histories and reconstruct ancestral states at nodes of molecular phylogenies, notably of morphological characters. As for all Bayesian analyses specification of prior values is an integrative and important part of the analysis. He we provide an example of how alternative prior choices can seriously influence results and mislead interpretations. Methods/Principal Findings For two contrasting discrete morphological characters, namely a slow and a fast evolving character found in the plant family Annonaceae, we specified a total of eight different prior distributions per character. We investigated how these prior settings affected important summary statistics. Our analyses showed that the different prior distributions had marked effects on the results in terms of average number of character state changes. These differences arise because priors play a crucial role in determining which areas of parameter space the values of the simulation will be drawn from, independent of the data at hand. However, priors seemed to fit the data better if they would result in a more even sampling of parameter space (normal posterior distribution), in which case alternative standard deviation values had little effect on the results. The most probable character history for each character was affected differently by the prior. For the slower evolving character, the same character history always had the highest posterior probability independent of the priors used. In contrast, the faster evolving character showed different most probable character histories depending on the prior. These differences could be related to the level of homoplasy exhibited by each character. Conclusions Although our analyses were restricted to two morphological characters within a single family, our results underline the importance of carefully choosing prior values for posterior mapping. Prior specification will be of crucial importance when interpreting the results in a meaningful way. It is hard to suggest a statistically sound method for prior specification without more detailed studies. Meanwhile, we propose that the data could be used to estimate the prior value of the gamma distribution placed on the transformation rate in posterior mapping.

Taxonomic novelties in the genus Campylospermum (Ochnaceae)

Four new species, one with two subspecies, of the genus Campylospermum are described, all endemic or sub-endemic to Gabon. These are C. auriculatum, C. gabonensis, C. gabonensis subsp. australis, C. glaucifolium and C. occidentalis. Distribution maps and scans of the holotypes are provided as well as preliminary IUCN Red List assessments. New combinations for nine species formerly assigned to the genus Ouratea and/or Gomphia are proposed: C. andongensis, C. glomeratum, C. longestipulatum, C. lunzuensis, C. lutambensis, C. nutans, C. plicatum and C. warneckei. Finally, one taxon is raised from the variety to species level, leading to the new combination C. costatum.

Contribution to the taxonomy of Garcinia (Clusiaceae) in Africa, including two new species from Gabon and a key to the Lower Guinean species

Abstract Garcinia has some 260 species and is often regarded as a genus with a difficult taxonomy. No recent treatment is available for the botanically rich Lower Guinea phytogeographical region. This study aims at partly filling this gap. First, several taxonomic problems are solved. Garcinia chromocarpa is reduced to a variety of Garcinia quadrifaria. Garcinia gnetoides and Garcinia granulata are both synonyms of Garcinia quadrifaria. Garcinia zenkeri is a synonym of Garcinia densivenia and lectotypes are being designated for both names. Garcinia brevipedicellata is a synonym of Garcinia afzelii, as is Garcinia antidysenterica for which a lectotype is designated. Second, two new species endemic to Gabon are described: Garcinia gabonensis Sosef & Dauby and Garcinia obliqua Sosef & Dauby. Finally, an identification key to all species present in the Lower Guinea region is provided. A few remaining West African species names could not be placed with certainty, because the type material was lost or not traced yet. One is a Rutaceae while the remaining three are provisionally to be regarded as synonyms of Garcinia smeathmannii.

Phylogenetics of African Rinorea (Violaceae): Elucidating Infrageneric Relationships using Plastid and Nuclear DNA Sequences

Abstract Rinorea is a pantropical genus of shrubs and small trees within the family Violaceae. The genus is particularly diverse in Africa where species are ecologically important as they are often abundant or even dominant in particular forest types and act as larval host plants for highly specialized Cymothoe butterflies. Despite their importance, species identification of African Rinorea is difficult and a taxonomic revision is needed. Previous phylogenetic studies have suggested that neotropical taxa are sister to a palaeotropical clade, with multiple independent dispersals to Madagascar, but these were based on plastid data only. We therefore present an updated phylogeny of Rinorea with increased sampling of African taxa, using plastid as well as nuclear DNA sequences. Phylogenetic relationships inferred from nuclear DNA data were generally congruent with those based on evidence from plastid haplotypes from earlier studies. Our increased taxonomic sampling also revealed previously undiscovered African Rinorea clades, some of which warrant further taxonomic study. Ancestral state reconstructions refute previous hypotheses about the evolution of morphological characters traditionally used for Rinorea infrageneric classification. In addition, some widespread species may comprise species complexes. It is clear that African Rinorea require comprehensive taxonomic revision; our contribution to understanding Rinorea infrageneric relationships will facilitate this task.