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.

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.

Phylogenetic analyses of the family Sapindaceae inferred from nuclear and plastid sequence data have revealed a high level of para- and polyphyly at the subfamilial, tribal, and generic levels. A phylogenetic study focusing on taxa in the southern Pacific belonging to tribe Cupanieae has shown that the two most species-rich genera, Arytera Blume and Cupaniopsis Radlk., are polyphyletic. This study aims to clarify generic limits among the taxa currently placed in these two genera by identifying morphological features that support monophyletic groups suitable for recognition at the generic level. Specimens deposited in major herbaria holding material of these taxa were examined to complement extensive field observations. Careful consideration of morphological features in light of previous taxonomic treatments and the results of phylogenetic analyses enabled us to propose a re-aligned generic framework for Cupanieae in which two new genera are described to accommodate species previously placed in Arytera and Cupaniopsis: viz., Lepidocupania Buerki, Callm., Munzinger & Lowry (21 species) and Neoarytera Callm., Buerki, Munzinger & Lowry (4 species). A total of 25 new combinations are made, lectotypes are designated for nine names (two first step and seven second-step), and one new synonym is established. A key to the newly circumscribed genera Arytera and Cupaniopsis, along with allied genera, is provided, accompanied by information on the distribution and ecology of each species.

Abstract

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.

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.

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.

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.