Angiosperms are the cornerstone of most terrestrial ecosystems and human livelihoods^1,2. A robust understanding of angiosperm evolution is required to explain their rise to ecological dominance. So far, the angiosperm tree of life has been determined primarily by means of analyses of the plastid genome^3,4. Many studies have drawn on this foundational work, such as classification and first insights into angiosperm diversification since their Mesozoic origins^5–7. However, the limited and biased sampling of both taxa and genomes undermines confidence in the tree and its implications. Here, we build the tree of life for almost 8,000 (about 60%) angiosperm genera using a standardized set of 353 nuclear genes⁸. This 15-fold increase in genus-level sampling relative to comparable nuclear studies⁹ provides a critical test of earlier results and brings notable change to key groups, especially in rosids, while substantiating many previously predicted relationships. Scaling this tree to time using 200 fossils, we discovered that early angiosperm evolution was characterized by high gene tree conflict and explosive diversification, giving rise to more than 80% of extant angiosperm orders. Steady diversification ensued through the remaining Mesozoic Era until rates resurged in the Cenozoic Era, concurrent with decreasing global temperatures and tightly linked with gene tree conflict. Taken together, our extensive sampling combined with advanced phylogenomic methods shows the deep history and full complexity in the evolution of a megadiverse clade.

Sapindales is an angiosperm order of high economic and ecological value comprising nine families, c. 479 genera, and c. 6570 species. However, family and subfamily relationships in Sapindales remain unclear, making reconstruction of the order’s spatio-temporal and morphological evolution difficult. In this study, we used Angiosperms353 target capture data to generate the most densely sampled phylogenetic trees of Sapindales to date, with 448 samples and c. 85% of genera represented. The percentage of paralogous loci and allele divergence was characterized across the phylogeny, which was time-calibrated using 29 rigorously assessed fossil calibrations. All families were supported as monophyletic. Two core family clades subdivide the order, the first comprising Kirkiaceae, Burseraceae, and Anacardiaceae, the second comprising Simaroubaceae, Meliaceae, and Rutaceae. Kirkiaceae is sister to Burseraceae and Anacardiaceae, and, contrary to current understanding, Simaroubaceae is sister to Meliaceae and Rutaceae. Sapindaceae is placed with Nitrariaceae and Biebersteiniaceae as sister to the core Sapindales families, but the relationships between these families remain unclear, likely due to their rapid and ancient diversification. Sapindales families emerged in rapid succession, coincident with the climatic change of the Mid-Cretaceous Hothouse event. Subfamily and tribal relationships within the major families need revision, particularly in Sapindaceae, Rutaceae and Meliaceae. Much of the difficulty in reconstructing relationships at this level may be caused by the prevalence of paralogous loci, particularly in Meliaceae and Rutaceae, that are likely indicative of ancient gene duplication events such as hybridization and polyploidization playing a role in the evolutionary history of these families. This study provides key insights into factors that may affect phylogenetic reconstructions in Sapindales across multiple scales, and provides a state-of-the-art phylogenetic framework for further research.

A checklist of all species of bryophytes, lycophytes, ferns, gymnosperms and angiosperms that are found in the wild (native, naturalised and casual) in Singapore is presented. We have attempted to account for all names of species and infraspecific taxa that have ever been recorded for Singapore, along with the pertinent publications that reported each of these names. For each currently accepted name, the synonyms of relevance for Singapore are included. The native or non-native status for all taxa is given, along with the most recent national conservation assessment applied to each native taxon. If we were aware that the most recent assessment required an update, the taxon is newly assessed here. The checklist includes 2654 native taxa, 479 naturalised/casual taxa and 101 cryptogenic taxa.

Premise

The economically important, cosmopolitan soapberry family (Sapindaceae) comprises ca. 1900 species in 144 genera. Since the seminal work of Radlkofer, several authors have attempted to overcome challenges presented by the family’s complex infra‐familial classification. With the advent of molecular systematics, revisions of the various proposed groupings have provided significant momentum, but we still lack a formal classification system rooted in an evolutionary framework.

Although vanilla is one of the most popular flavours in the world, there is still uncertainty concerning the native distribution of the species that produces it, Vanilla planifolia. To circumscribe the native geographical extent of this economically important species more precisely, we propose a new landscape-based approach to incorporate information from open-source databases and validate occurrences. In this approach, we include metrics to account for habitat suitability and population sustainability in terms of the biotic (co-occurrence of pollinators and dispersers) and abiotic (habitat quality) factors limiting plant distributions. To further validate occurrences within the resulting distribution, we compare the presence of morphologically similar wild relatives, assess the heterogeneity of ecological niches and verify the correct identification of herbarium specimens. Results from this approach suggest that V. planifolia has a larger geographical distribution than previously recognized; we hypothesize that populations naturally dispersed from Mesoamerica and became established in South America (with a south-eastern limit in Brazil). The recognition of an improved estimate of the distribution of this species will increase the accuracy of predictive models, promote further species circumscription, improve the efficacy of conservation strategies, and help to ensure the sustainability of a valuable, sought-after spice.

Driven by limited resources and a sense of urgency, the prioritization of species for conservation has been a persistent concern in conservation science. Gymnosperms (comprising ginkgo, conifers, cycads, and gnetophytes) are one of the most threatened groups of living organisms, with 40% of the species at high risk of extinction, about twice as many as the most recent estimates for all plants (i.e. 21.4%). This high proportion of species facing extinction highlights the urgent action required to secure their future through an objective prioritization approach. The Evolutionary Distinct and Globally Endangered (EDGE) method rapidly ranks species based on their evolutionary distinctiveness and the extinction risks they face. EDGE is applied to gymnosperms using a phylogenetic tree comprising DNA sequence data for 85% of gymnosperm species (923 out of 1090 species), to which the 167 missing species were added, and IUCN Red List assessments available for 92% of species. The effect of different extinction probability transformations and the handling of IUCN data deficient species on the resulting rankings is investigated. Although top entries in our ranking comprise species that were expected to score well (e.g. Wollemia nobilis, Ginkgo biloba), many were unexpected (e.g. Araucaria araucana). These results highlight the necessity of using approaches that integrate evolutionary information in conservation science.

Understanding how speciation relates to ecological divergence has long fascinated biologists. It is assumed that ecological divergence is essential to sympatric speciation, as a mechanism to avoid competition and eventually lead to reproductive isolation, while divergence in allopatry is not necessarily associated with niche differentiation. The impact of the spatial context of divergence on the evolutionary rates of abiotic dimensions of the ecological niche has rarely been explored for an entire clade. Here, we compare the magnitude of climatic niche shifts between sympatric versus allopatric divergence of lineages in butterflies. By combining next-generation sequencing, parametric biogeography and ecological niche analyses applied to a genus-wide phylogeny of PalaearcticPyrgusbutterflies, we compare evolutionary rates along eight climatic dimensions across sister lineages that diverged in large-scale sympatry versus allopatry. In order to examine the possible effects of the spatial scale at which sympatry is defined, we considered three sets of biogeographic assignments, ranging from narrow to broad definition. Our findings suggest higher rates of niche evolution along all climatic dimensions for sister lineages that diverge in sympatry, when using a narrow delineation of biogeographic areas. This result contrasts with significantly lower rates of climatic niche evolution found in cases of allopatric speciation, despite the biogeographic regions defined here being characterized by significantly different climates. Higher rates in allopatry are retrieved when biogeographic areas are too widely defined—in such a case allopatric events may be recorded as sympatric. Our results reveal the macro-evolutionary significance of abiotic niche differentiation involved in speciation processes within biogeographic regions, and illustrate the importance of the spatial scale chosen to define areas when applying parametric biogeographic analyses.

Abstract

This study investigates the biogeography, evolution and systematics of Benstonea Callm. & Buerki (Pandanaceae) based on six plastid DNA regions and 54 specimens representing 36 species (60% of species generic diversity). Our maximum likelihood and Bayesian phylogenetic inferences support the monophyly of Benstonea and its close relationship with the speciose Pandanus Parkinson. Benstonea is subdivided into three clades exhibiting contrasting species diversities. Clades I and II have seven species each, whereas most of the species diversity occurs in clade III with 21 species. None of the sections defined by Stone in Pandanus subgenus Acrostigma (Kurz) B.C. Stone (now Benstonea) are retrieved monophyletic by our analyses. Biogeographical inference supports the origin of Benstonea on the Sunda shelf during the Miocene and shows several subsequent exchanges between Peninsular Malaysia and Borneo. Species in Indochina and the Indian continent originated in Peninsular Malaysia and all belong to clade I. Wallacea was colonized at least twice from Borneo sometimes during the Miocene and no back-dispersals were inferred. The Sunda shelf was colonized once, most likely from Halmahera. Finally, our analyses suggest that the Fijian endemic Benstonea thurstonii (C.H. Wright) Callm. & Buerki dispersed from either Australia or New Guinea during the Pleistocene.

There is increased evidence that incorporating evolutionary history directly in conservation actions is beneficial, particularly given the likelihood that extinction is not random and that phylogenetic diversity (PD) is lost at higher rates than species diversity. This evidence is even more compelling in biodiversity hotspots, such as Madagascar, where less than 10% of the original vegetation remains. Here, we use the Leguminosae, an ecologically and economically important plant family, and a combination of phylogenetics and species distribution modelling, to assess biodiversity patterns and identify regions, coevolutionary processes and ecological factors that are important in shaping this diversity, especially during the Quaternary. We show evidence that species distribution and community PD are predicted by watershed boundaries, which enable the identification of a network of refugia and dispersal corridors that were perhaps important for maintaining community integrity during past climate change. Phylogenetically clustered communities are found in the southwest of the island at low elevation and share a suite of morphological characters (especially fruit morphology) indicative of coevolution with their main dispersers, the extinct and extant lemurs. Phylogenetically over-dispersed communities are found along the eastern coast at sea level and may have resulted from many independent dispersal events from the drier and more seasonal regions of Madagascar.

Abstract

New Caledonia is a remote archipelago of the South-West Pacific, whose flora is rich, distinctive, and disharmonic. The interest of botanists has long been attracted by the spatio-temporal origin of this flora, but little attention has been paid to the modes of colonization and the diversification processes that have led to the archipelago’s modern flora. To date, no explosive plant radiation has yet been highlighted for New Caledonia. A dated phylogenetic framework on the second richest New Caledonian genus – Psychotria s.l. and its allied genera (tribes Psychotrieae and Palicoureeae, Rubiaceae; ca. 85 species) – is provided in this study to explore its patterns of colonization and diversification in the archipelago. This study is based on a comprehensive species sampling, two nuclear and four plastid loci. Results show that New Caledonia was colonized four times by Psychotria and its allied genera during the Neogene long after its mid-Eocene re-emergence from the sea. The Pacific clade of Psychotrieae, one of the largest plant diversifications in the Pacific islands and the Indo-Pacific region, is absent from New Caledonia, possibly due to niche competition. Although the four lineages colonized New Caledonia relatively simultaneously during the Neogene, they express different evolutionary histories, as revealed by unevenness in species richness and net diversification rates. The genus Geophila has not diversified on New Caledonia, as a non-endemic single species has been documented in the archipelago. The genus Margaritopsis had a moderate level of diversification (four species) similar to that on other Pacific islands. The Psychotria clade NC1 appears to be a relictual lineage, which probably underwent a drastic extinction, with a narrow ecological habitat and dispersal limitations. The Psychotria clade NC2 is the largest and youngest New Caledonian plant radiation, and has undergone the fastest recorded diversification of any endemic lineage in the archipelago, and could be the result of a ‘non-adaptive radiation’, originating from Australian rainforests.

The paleotropical monocot Pandanaceae family comprises c. 700 species distributed into five genera: Benstonea (c. 60 spp.), Freycinetia (c. 250 spp.), Martellidendron (6 spp.), Pandanus (c. 450 spp.) and Sararanga (2 spp.). Benstonea was circumscribed to include species previously placed in Pandanus section Acrostigma (one of the four sections of Pandanus subgenus Acrostigma). New phylogenetic data show that the six species of the remaining three sections of subgenus Acrostigma (sections Epiphytica, Fusiforma and Platystigma) and a seventh species doubtfully placed in section Acrostigma (Pandanus microglottis) also belonged to Benstonea. This genus is therefore characterized by a suite of morphological characters, viz. stigmatic groove on the adaxial side of the stigma and a staminate flower reduced to 1 to 3 free stamens (sometimes joined at base). We therefore make here the necessary seven new combinations accompanied by one lectotypification in Benstonea, a genus that now reflects the view of the visionary Benjamin Stone who had already grouped these species in Pandanus subgenus Acrostigma based solely on morphology.

Background and Aims

Tecophilaeaceae (27 species distributed in eight genera) have a disjunct distribution in California, Chile and southern and tropical mainland Africa. Moreover, although the family mainly occurs in arid ecosystems, it has colonized three Mediterranean-type ecosystems. In this study, the spatio-temporal history of the family is examined using DNA sequence data from six plastid regions.

Methods

Modern methods in divergence time estimation (BEAST), diversification (LTT and GeoSSE) and biogeography (LAGRANGE) are applied to infer the evolutionary history of Tecophilaeaceae. To take into account dating and phylogenetic uncertainty, the biogeographical inferences were run over a set of dated Bayesian trees and the analyses were constrained according to palaeogeographical evidence.

Key Results

The analyses showed that the current distribution and diversification of the family were influenced primarily by the break up of Gondwana, separating the family into two main clades, and the establishment of a Mediterranean climate in Chile, coinciding with the radiation of Conanthera. Finally, unlike many other groups, no shifts in diversification rates were observed associated with the dispersals in the Cape region of South Africa.

Conclusions

Although modest in size, Tecophilaeaceae have a complex spatio-temporal history. The family is now most diverse in arid ecosystems in southern Africa, but is expected to have originated in sub-tropical Africa. It has subsequently colonized Mediterranean-type ecosystems in both the Northern and Southern Hemispheres, but well before the onset of the Mediterranean climate in these regions. Only one lineage, genus Conanthera, has apparently diversified to any extent under the impetus of a Mediterranean climate.

Abstract

The Paleotropical monocot family Pandanaceae includes ca. 700 species assigned to four genera: Pandanus (ca. 500 spp.), Freycinetia (ca. 200 spp.), Martellidendron (6 spp.) and Sararanga (2 spp.). The most speciose genus, Pandanus, was classically subdivided into eight subgenera. Previous cladistic analyses revealed that several key morphological characters might have evolved independently several times, thus highlighting the need for a robust molecular phylogenetic framework to elucidate phylogenetic relationships and infrafamilial and infrageneric classification within this group. In this study, three plastid DNA regions (matK, trnQ­rps16, trnL­trnF) and 200 individuals (representing 134 species and 609 newly produced sequences)—spanning the taxonomic and biogeographic diversity of the family—are analyzed to test the monophyly at the familial and generic levels, and to infer phylogenetic relationships within the family. Particular emphasis is devoted to Pandanus with the aim of recognizing key morphological characters that reflect the evolutionary history of the genus. Phylogenetic inferences support the monophyly of Pandanaceae and establish Sararanga as sister to the rest of the family, with Freycinetia as sister to the Pandanus­Martellidendron pair. Although relationships are not well­resolved within the latter clade, three supported lineages are retrieved: (1) the Acrostigma clade comprising taxa of P. subg. Acrostigma, (2) the Martellidendron clade including taxa assigned to the genus Martellidendron and (3) the core Pandanus clade including taxa of all other subgenera of Pandanus. Morphological and biogeographic evidence supporting clade definitions are discussed in detail. This study provides the first phylogenetic backbone for Pandanaceae, which is sufficiently robust to serve as a springboard for future research into the evolutionary history of this neglected family.

Sapindaceae (Sapindales) are a conspicuous and diversified element of the New Caledonian flora, with ca. 67 species (ca. 90 % endemic) in 13 genera (four endemic: Gongrodiscus, Loxodiscus, Podonephelium, Storthocalyx). The phylogeny of New Caledonian Sapindaceae is inferred by adding 97 new samples, encompassing the full distributional and morphological range of the archipelago's genera, to a broad plastid and nuclear DNA sequence dataset that is representative of the family worldwide. Results from phylogenetic analyses indicate that members of the family on New Caledonia belong to two major clades, the Dodonaea group (placed within subfamily Dodonaeoideae) and the Cupania group (subfamily Sapindoideae), which exhibit strikingly different species diversities (ca. 89% of the species on New Caledonia belong to the Cupania group). Results support the monophyly of all four endemic genera and most of those that also occur elsewhere, with the exception of the morphologically similar Austro‐Pacific genera Arytera and Cupaniopsis, both of which have representatives in each of two well‐supported subclades within the Cupania group, suggesting at least two dispersals to New Caledonia (most likely from Australia). The results provide a robust phylogenetic framework for ongoing taxonomic revisions of Sapindaceae genera on New Caledonia and for investigating the spatio‐temporal history of the family in this biogeographically intriguing archipelago, although expanded sampling (including from other areas) and further analyses will be required to resolve generic limits among the taxa currently placed in Arytera and Cupaniopsis.

Abstract

Pandanaceae, a palaeotropical monocot family of c. 700 species, comprises four currently recognized genera: Freycinetia Gaudich., Martellidendron (Pic. Serm.) Callm. & Chassot, Pandanus Parkinson and Sararanga Helms. Within Pandanus (c. 500 spp.), species of sect. Acrostigma Kurz [one of four sections comprising subg. Acrostigma (Kurz) B. C. Stone] possess highly distinctive morphological features (viz. sharp spiniform, linear styles with the stigmatic groove on the abaxial side of the style and a staminate flower reduced to 1 to 3 stamens) shared with two other species (likewise belonging to subg. Acrostigma but originally placed in sect. Fusiforma B. C. Stone) that separate them from all other congeners. Based on morphology, biogeography, and recent inferences from plastid DNA sequence data, we place these distinctive species in a new genus, Benstonea Callm. & Buerki, making the necessary new combinations for the 50 recognized species, accompanied by six lectotypifications, one epitytification and two neotypifications, and placing seventeen names in synonymy. A generic key is provided to facilitate distinguishing Benstonea from the four other genera of Pandanaceae. Comments are provided on the distribution, ecology and typification of each accepted species.

In the past, the circumscription of the large genus Psychotria (Rubiaceae) was difficult, until molecular phylogenetic studies revealed its considerable paraphyly, enabling the delimitation of its major lineages and the grouping of related genera, and most notably the separation of Psychotria and its relatives (former Psychotrieae) into two tribes: Psychotrieae and Palicoureeae. The genus Margaritopsis, which is included in Palicoureeae, encompasses 27 Neotropical species, and in previous studies these have been shown to be close relatives of a group of eight Psychotria species that occur over a large region extending from South‐East Asia to tropical South Pacific through Malesia (= the AMP region, defined as including South‐East Asia, Malesia, tropical Australia, Melanesia, Micronesia, New Caledonia and Polynesia). A molecular phylogenetic study, using one nuclear DNA region (ITS) and four plastid DNA regions (ndhF, rps16, trnH‐psbA, trnT‐F), is undertaken in order to test the placement of 17 AMP Psychotria species within Palicoureeae. The phylogenetic results show that they form a monophyletic clade (= clade G), which also includes the monotypic Fijian genus Readea and Hodgkinsonia frutescens from Australia. Clade G is embedded in a well‐supported grade with five Neotropical representatives of Margaritopsis. A morphological survey based on twenty characters, with the potential to circumscribe generic entities, shows that the monophyly of clade G is supported by a character combination that is similar to species of Neotropical Margaritopsis, confirming their inclusion in this genus. Taxonomic and nomenclatural work on these species is required to formalize nomenclatural implications. Based on the same set of morphological characters, but in absence of molecular data, 28 other Psychotria species from the AMP region were detected as likely candidates for inclusion in Margaritopsis, allowing the estimation of species richness of clade G to be a minimum of 47 species. Within clade G, five well‐supported subclades and a Readea lineage are delimited and each of these is generally supported by unique morphological features. Phylogenetic topologies reveal biogeographical patterns, including a main route of dispersal from western to eastern parts of the AMP region, with subsequent dispersals between archipelagos in the region.