Pulled Diversification Rates, Lineages-Through-Time Plots, and Modern Macroevolutionary Modeling.

Estimating time-dependent rates of speciation and extinction from dated phylogenetic trees of extant species (timetrees), and determining how and why they vary, is key to understanding how ecological and evolutionary processes shape biodiversity. Due to an increasing availability of phylogenetic trees, a growing number of process-based methods relying on the birth-death model have been developed in the last decade to address a variety of questions in macroevolution. However, this methodological progress has regularly been criticized such that one may wonder how reliable the estimations of speciation and extinction rates are. In particular, using lineages-through-time (LTT) plots, a recent study has shown that there are an infinite number of equally likely diversification scenarios that can generate any timetree. This has led to questioning whether or not diversification rates should be estimated at all. Here, we summarize, clarify, and highlight technical considerations on recent findings regarding the capacity of models to disentangle diversification histories. Using simulations, we illustrate the characteristics of newly proposed "pulled rates" and their utility. We recognize that the recent findings are a step forward in understanding the behavior of macroevolutionary modeling, but they in no way suggest we should abandon diversification modeling altogether. On the contrary, the study of macroevolution using phylogenetic trees has never been more exciting and promising than today. We still face important limitations in regard to data availability and methods, but by acknowledging them we can better target our joint efforts as a scientific community. [Birth-death models; extinction; phylogenetics; speciation.].

A sequential diversification with Miocene extinction and Pliocene speciation linked to mountain uplift explains the diversity of the African rain forest clade Monodoreae (Annonaceae).

BACKGROUND AND AIMS: Throughout the Cenozoic, Africa underwent several climatic and geological changes impacting the evolution of tropical rain forests (TRF). African TRF are thought to have extended from East to West in a 'pan-African' TRF, followed by several events of fragmentation during drier climate periods. During the Miocene, climate cooling and mountain uplift led to the aridification of tropical Africa and open habitats expanded at the expense of TRF, which likely experienced local extinctions. However, in plants, these drivers were previously inferred using limited taxonomic and molecular data. Here, we tested the impact of climate and geological changes on diversification within the diverse clade Monodoreae (Annonaceae) composed of 90 tree species restricted to African TRF. METHODS: We reconstructed a near complete phylogenetic tree, based on 32 nuclear genes, and dated using relaxed clocks and fossil calibrations in a Bayesian framework. We inferred the biogeographic history and the diversification dynamics of the clade using multiple birth-death models. KEY RESULTS: Monodoreae originated in East African TRF ca. 25 million years ago (Ma) and expanded toward Central Africa during the Miocene. We inferred range contractions during the middle Miocene and document important connections between East and West African TRF after 15-13 Ma. Our results indicated a sudden extinction event during the late Miocene, followed by an increase in speciation rates. Birth-death models suggested that African elevation change (orogeny) is positively linked to speciation in this clade. CONCLUSION: East Africa is inferred as an important source of Monodoreae species, and possibly for African plant diversity in general. Our results support a "sequential scenario of diversification" where increased aridification triggered extinction of TRF species in Monodoreae. This was quickly followed by rain forests fragmentation, subsequently enhancing lagged speciation resulting from vicariance and improved climate conditions. In contrast to previous ideas, the uplift of East Africa is shown to have played a positive role in Monodoreae diversification.

Cradles and museums of generic plant diversity across tropical Africa.

Determining where species diversify (cradles) and persist (museums) over evolutionary time is fundamental to understanding the distribution of biodiversity and for conservation prioritization. Here, we identify cradles and museums of angiosperm generic diversity across tropical Africa, one of the most biodiverse regions on Earth. Regions containing nonrandom concentrations of young (neo-) and old (paleo-) endemic taxa were identified using distribution data of 1719 genera combined with a newly generated time-calibrated mega-phylogenetic tree. We then compared the identified regions with the current network of African protected areas (PAs). At the generic level, phylogenetic diversity and endemism are mainly concentrated in the biogeographically complex region of Eastern Africa. We show that mountainous areas are centres of both neo- and paleo-endemism. By contrast, the Guineo-Congolian lowland rain forest region is characterized by widespread and old lineages. We found that the overlap between centres of phylogenetic endemism and PAs is high (> 85%). We show the vital role played by mountains acting simultaneously as cradles and museums of tropical African plant biodiversity. By contrast, lowland rainforests act mainly as museums for generic diversity. Our study shows that incorporating large-scale taxonomically verified distribution datasets and mega-phylogenies lead to an improved understanding of tropical plant biodiversity evolution.

An updated and extended version of the Melastomataceae probe set for target capture.

Premise: A probe set was previously designed to target 384 nuclear loci in the Melastomataceae family; however, when trying to use it, we encountered several practical and conceptual problems, such as the presence of sequences in reverse complement, intronic regions with stop codons, and other issues. This raised concerns regarding the use of this probe set for sequence recovery in Melastomataceae. Methods: In order to correct these issues, we cleaned the Melastomataceae probe set, extended it with additional sequences, and compared its performance with the original version. Results: The final probe set targets 396 putative nuclear loci represented by 6009 template sequences. The probe set has been made available, along with details on the cleaning process, for reproducibility. We show that the new probe set performs better than the original version in terms of sequence recovery. Discussion: This updated, extended, and cleaned probe set will improve the availability of phylogenomic resources across the Melastomataceae family. It is fully compatible with sequence recovery and extraction pipelines. The cleaning process can also be applied to any plant-targeting probe set that would need to be cleaned or updated if new genomic resources for the targeted taxa become available.

Phylogenomic inference of the African tribe Monodoreae (Annonaceae) and taxonomic revision of Dennettia, Uvariodendron and Uvariopsis.

Monodoreae (Annonaceae) is a tribe composed of 11 genera and 90 species restricted to the tropical African rain forests. All the genera are taxonomically well circumscribed except the species rich genera Uvariodendron and Uvariopsis which lack a recent taxonomic revision. Here, we used a robust phylogenomic approach, including all the 90 currently accepted species, with several specimens per species, and based on more than 300 Annonaceae-specific nuclear genes, to infer the phylogenetic tree of the Monodoreae and test the limits between the genera and species. We recover all the genera as monophyletic, except the genus Uvariopsis for which the species Uvariopsistripetala falls outside this clade. We thus reinstate the monotypic genus Dennettia for its single species Dennettiatripetala. We also erect a new tribe, Ophrypetaleae trib. nov., to accommodate the genera Ophrypetalum and Sanrafaelia, as we recover them excluded from the Monodoreae tribe with good support. Below the genus level, the genera Isolona, Monodora, Uvariastrum, Uvariodendron and Uvariopsis show weakly supported nodes and phylogenetic conflicts, suggesting that population level processes of evolution might occur in these clades. Our results also support, at the molecular level, the description of several new species of Uvariodendron and Uvariopsis, as well as several new synonymies. Finally, we present a taxonomic revision of the genera Dennettia, Uvariodendron and Uvariopsis, which contain one, 18 and 17 species respectively. We provide a key to the 11 genera of the Monodoraeae and describe four new species to science: Uvariodendronkimbozaense Dagallier & Couvreur, sp. nov., Uvariodendronmossambicense Robson ex Dagallier & Couvreur, sp. nov., Uvariodendronpilosicarpum Dagallier & Couvreur, sp. nov. and Uvariopsisoligocarpa Dagallier & Couvreur, sp. nov., and provide provisional descriptions of three putatively new species. We also present lectotypifications and nomenclatural changes implying synonymies and new combinations (Uvariodendroncitriodorum (Le Thomas) Dagallier & Couvreur, comb. et stat. nov., Uvariodendronfuscumvar.magnificum (Verdc.) Dagallier & Couvreur, comb. et stat. nov., Uvariopsiscongensisvar.angustifolia Dagallier & Couvreur, var. nov., Uvariopsisguineensisvar.globiflora (Keay) Dagallier & Couvreur, comb. et stat. nov., and Uvariopsissolheidiivar.letestui (Pellegr.) Dagallier & Couvreur, comb. et stat. nov.).

RésuméLa tribu des Monodoreae (Annonaceae) est composée de 11 genres et 90 espèces des forêts tropicales humides d'Afrique. Tout les genres sont taxonomiquement bien résolus, à part les genres Uvariodendron et Uvariopsis qui manquent d'une révision taxonomique récente. Ici, nous avons utilisé une approche phylogénomique robuste pour estimer l'arbre phylogénétique des Monodoreae, et tester les limites de genres et d'espèces. Pour cela, nous avons inclut les 90 espèces acceptées, et avons séquencé plus de 300 gènes. Tous les genres sont retrouvés monophylétiques, à part le genre Uvariopsis pour lequel l'espèce Uvariopsistripetala se retrouve exclue. Nous rétablissons donc le genre monotypique Dennettia et son unique espèce Dennettiatripetala. Nous érigeons une nouvelle tribu, les Ophrypetaleae trib. nov., pour accueillir les genres Ophrypetalum et Sanrafaelia, car nous les retrouvons exclus de la tribu des Monodoreae avec un bon support. Au niveau infra-générique, les genres Isolona, Monodora, Uvariastrum, Uvariodendron et Uvariopsis montrent de faibles supports de noeuds et des conflits phylogénétiques, ce qui suggère que des processus d'évolution se déroulent au niveau des populations. Nos résultats soutiennent également, sur un plan moléculaire, la description de plusieurs nouvelles espèces d'Uvariodendron et d'Uvariopsis, de même que plusieurs synonymies. Enfin, nous présentons une révision taxonomique des genres Dennettia, Uvariodendron et Uvariopsis, qui contiennent respectivement un, 18 et 17 espèces. Nous fournissons une clé des 11 genres de Monodoreae, et décrivons quatre nouvelles espèces pour la science: Uvariodendronkimbozaense Dagallier & Couvreur, sp. nov., Uvariodendronmossambicense Robson ex Dagallier & Couvreur, sp. nov., Uvariodendronpilosicarpum Dagallier & Couvreur, sp. nov. et Uvariopsisoligocarpa Dagallier & Couvreur, sp. nov., et fournissons une description provisoire de trois autres potentielles. Nous effectuons des lectotypifications et des changements nomenclaturaux tels que des synonymies et des nouvelles combinaisons (Uvariodendroncitriodorum (Le Thomas) Dagallier & Couvreur, comb. et stat. nov., Uvariodendronfuscumvar.magnificum (Verdc.) Dagallier & Couvreur, comb. et stat. nov., Uvariopsiscongensisvar.angustifolia Dagallier & Couvreur, var. nov., Uvariopsisguineensisvar.globiflora (Keay) Dagallier & Couvreur, comb. stat. nov., et Uvariopsissolheidiivar.letestui (Pellegr.) Dagallier & Couvreur, comb. stat. nov.).

A Taxonomically-verified and Vouchered Checklist of the Vascular Plants of the Republic of Guinea.

The Checklist of the Vascular Plants of the Republic of Guinea (CVPRG) is a specimen-based, expert-validated knowledge product, which provides a concise synthesis and overview of current knowledge on 3901 vascular plant species documented from Guinea (Conakry), West Africa, including their accepted names and synonyms, as well as their distribution and status within Guinea (indigenous or introduced, endemic or not). The CVPRG is generated automatically from the Guinea Collections Database and the Guinea Names Backbone Database, both developed and maintained at the Royal Botanic Gardens, Kew, in collaboration with the staff of the National Herbarium of Guinea. A total of 3505 indigenous vascular plant species are reported of which 3328 are flowering plants (angiosperms); this represents a 26% increase in known indigenous angiosperms since the last floristic overview. Intended as a reference for scientists documenting the diversity and distribution of the Guinea flora, the CVPRG will also inform those seeking to safeguard the rich plant diversity of Guinea and the societal, ecological and economic benefits accruing from these biological resources.

Uvariopsis dicaprio (Annonaceae) a new tree species with notes on its pollination biology, and the Critically Endangered narrowly endemic plant species of the Ebo Forest, Cameroon.

BACKGROUND: The Ebo Forest area is a highly threatened centre of diversity in the Littoral Region of Cameroon, globally important for conservation with many threatened species including 68 threatened species of plant, yet not formally protected. The tropical African evergreen forest tree genus Uvariopsis Engl. & Diels (Annonaceae) is characterised by unisexual, usually cauliflorous flowers with a uniseriate corolla of four petals, and two sepals. Cameroon is the centre of diversity of the genus with 14 of the 19 known species. METHODS: The herbarium collection MacKinnon 51 from Ebo is hypothesized to represent a new species to science of Uvariopsis. This hypothesis is tested by the study of herbarium specimens from a number of herbaria known to hold important collections from Cameroon and surrounding countries. RESULTS: We test the hypothesis that MacKinnon 51 represents a new species to science, using the most recent dichotomous identification key, and comparing it morphologically with reference material of all known species of the genus. We make a detailed comparative morphological study focussing on three other Cameroonian species, Uvariopsis solheidii, U. korupensis and the sympatric U. submontana. In the context of a review of the pollination biology of Uvariopsis, we speculate that in a genus otherwise with species with dull, flesh-coloured (pink, red to brown) flowers pollinated (where known) by diptera, orthoptera and blattodea (flies, crickets and cockroaches), the glossy, pale yellow-green flowers of Uvariopsis dicaprio, with additional traits unique in the genus, may be pollinated by nocturnal moths. Based on MacKinnon 51, we formally name Uvariopsis dicaprio Cheek & Gosline (Annonaceae) as new to science, and we describe, and illustrate, and map it. Restricted so far to a single site in evergreen forest in the Ebo Forest, Littoral Region, Cameroon, Uvariopsis dicaprio is provisionally assessed as Critically Endangered using the IUCN, 2012 standard because the forest habitat of this species remains unprotected, and there exist imminent threats of logging and conversion to plantations. DISCUSSION: We show that the highest density of species of the genus (12), and of narrow endemics (5), is found in the Cross-Sanaga Interval of SE Nigeria and Western Cameroon. A revised key to the 14 Cameroonian species of Uvariopsis is presented. We review the other seven narrowly endemic and threatened species unique to the Ebo forest of Cameroon and discuss the phytogeographic affinities of the area. CONCLUSIONS: Uvariopsis dicaprio adds to the growing list of species threatened with extinction at Ebo Forest due to current anthropogenic pressures.

Flora of Cameroon - Annonaceae Vol 45.

Annonaceae is a major pantropical family with 113 genera and about 2550 species. Cameroon is one of the most biodiverse countries in Africa but its flora remains incompletely known. In this volume of the Flora of Cameroon, we describe 166 native taxa representing 163 species in 28 native genera within the family Annonaceae. A total of 22 species (about 13%) are endemic to the country. We provide keys to all native genera, species, and infraspecific taxa. For each species a detailed morphological description and a map of its distribution in Cameroon are provided. Distribution maps and diversity analyses are based on a taxonomically verified database of 2073 collections. Across Africa, Cameroon is a center of diversity for Annonaceae harboring one of the highest numbers of species and genera. For example, Cameroon harbors the highest number of African species for the only pantropical genus of Annonaceae, Xylopia. Annonaceae are found across all 10 administrative regions of Cameroon but diversity is concentrated within the tropical rain forest areas situated in the south and South-West. The areas around Bipindi and Mount Cameroon show the highest levels of diversity, but this is correlated with collection effort. Line drawings and/or photographs accompany most species. One species new to science Uvariopsisetugeana Dagallier & Couvreur sp. nov. is described. We also undertake a number of nomenclatural changes such as lectotypifications, six new synonymies and two new combinations (Uvariaanisotricha (Le Thomas) Couvreur, comb. nov.; Uvariodendronfuscumvar.giganteum (Engl.) Dagallier & Couvreur, comb. nov.).

RésuméLes Annonacées sont une grande famille pantropicale avec 113 genres et 2550 espèces. Le Cameroun est l'un des pays les plus riches en biodiversité d'Afrique mais sa flore reste incomplètement connue. Dans ce volume de la Flore du Cameroun, nous décrivons 166 taxons représentant 163 espèces dans 28 genres au sein de la famille des Annonaceae. Au total, 22 espèces (environ 13%) sont endémiques du pays. Nous fournissons une clé de tous les genres et espèces et des infra-espèces au sein des genres. Pour chaque espèce une description morphologique détaillée et une carte de sa répartition au Cameroun sont fournies. Les cartes de distribution et les analyses de diversité sont basées sur une base de données taxonomiquement vérifiée de 2073 collections. À travers l'Afrique, le Cameroun est un centre de diversité pour les Annonacées abritant l'un des plus grands nombres d'espèces et de genres. Par exemple, le Cameroun abrite le plus grand nombre d'espèces africaines pour le seul genre pantropical d'Annonaceae, Xylopia. Les Annonaceae sont présentent dans les 10 régions du Cameroun, mais la plus grande diversité est concentrée dans les régions abritant la forêt tropicale humide située au sud et au sud-ouest. Les zones autour de Bipindi et du Mont Cameroun présentent les niveaux de diversité les plus élevés, mais cela est corrélé à l'effort de collecte. Des dessins et/ou des photographies accompagnent la plupart des espèces. Une espèce nouvelle pour la science Uvariopsisetugeana Dagalier & Couvreur sp. nov. est décrite. Nous entreprenons également un certain nombre de changements nomenclaturaux tels que des lectotypifications, six nouvelles synonymies et deux nouvelles combinaisons (Uvariaanisotricha (Le Thomas) Couvreur, comb. nov.; Uvariodendronfuscumvar.giganteum (Engl.) Dagalier & Couvreur, comb. nov.).

Three new species of Uvariodendron (Annonaceae) from coastal East Africa in Kenya and Tanzania.

East Africa is a hotspot of biodiversity with many endemic plant species. We describe three new species of the genus Uvariodendron (Annonaceae) from the coastal forests of Kenya and Tanzania. Uvariodendron mbagoi Dagallier & Couvreur, sp. nov. is endemic to Tanzania and unique within the genus by its strong bergamot scent and its tomentose fruits having regular tufts of higher hair density. Uvariodendron dzomboense Dagallier, W.R.Q. Luke & Couvreur, sp. nov. is endemic to Dzombo Hill in Kenya and is rendered distinct by its small leaves and very densely pubescent carpels. Uvariodendron schmidtii W.R.Q. Luke, Dagallier & Couvreur, sp. nov. is endemic to Shimba Hills in Kenya and differs by its small flowers and fused sepals forming a ring. Following IUCN criteria we assessed U. mbagoi and U. dzomboense as endangered (EN) while U. schmidtii is assessed as Vulnerable (VU). We also propose a new combination: Polyceratocarpus oligocarpus (Verdc.) Dagallier, comb. nov. The description of these three new species underlines the richness in endemics in East Africa and that new discoveries might arise from further botanical exploration of this region.

A large-scale species level dated angiosperm phylogeny for evolutionary and ecological analyses.

Phylogenies are a central and indispensable tool for evolutionary and ecological research. Even though most angiosperm families are well investigated from a phylogenetic point of view, there are far less possibilities to carry out large-scale meta-analyses at order level or higher. Here, we reconstructed a large-scale dated phylogeny including nearly 1/8th of all angiosperm species, based on two plastid barcoding genes, matK (incl. trnK) and rbcL. Novel sequences were generated for several species, while the rest of the data were mined from GenBank. The resulting tree was dated using 56 angiosperm fossils as calibration points. The resulting megaphylogeny is one of the largest dated phylogenetic tree of angiosperms yet, consisting of 36,101 sampled species, representing 8,399 genera, 426 families and all orders. This novel framework will be useful for investigating different broad scale research questions in ecological and evolutionary biology.