Fossil berries reveal global radiation of the nightshade family by the early Cenozoic.

Fossil discoveries can transform our understanding of plant diversification over time and space. Recently described fossils in many plant families have pushed their known records farther back in time, pointing to alternative scenarios for their origin and spread. Here, we describe two new Eocene fossil berries of the nightshade family (Solanaceae) from the Esmeraldas Formation in Colombia and the Green River Formation in Colorado (USA). The placement of the fossils was assessed using clustering and parsimony analyses based on 10 discrete and five continuous characters, which were also scored in 291 extant taxa. The Colombian fossil grouped with members of the tomatillo subtribe, and the Coloradan fossil aligned with the chili pepper tribe. Along with two previously reported early Eocene fossils from the tomatillo genus, these findings indicate that Solanaceae were distributed at least from southern South America to northwestern North America by the early Eocene. Together with two other recently discovered Eocene berries, these fossils demonstrate that the diverse berry clade and, in turn, the entire nightshade family, is much older and was much more widespread in the past than previously thought.

Flowers of Apocynaceae in amber from the early Eocene of India.

PREMISE: Early Eocene ambers of the Cambay lignite in Gujarat, India, are well known for their diverse insect fauna and dispersed pollen, but the included flowers have received limited attention. The fossil record of Apocynaceae is relatively poor, and the distinctive floral characters of this family have not been recognized in the fossil record before. METHODS: Remains of tiny flowers in amber were studied by micro-CT scanning, reflected light, and epifluorescence microscopy. RESULTS: Flowers of Maryendressantha succinifera gen. et. sp. n. have actinomorphic, pentamerous, tubular corollas 2.2-2.3 mm wide, and 1.7-2.1 mm deep with sinistrorse aestivation and androecia consisting of a whorl of five stamens attached by short filaments to the lower half of the corolla tube. Anthers are ovate, rounded basally and apically tapered with their connectives convergent with one another in a conical configuration. The pollen is globose, psilate, tricolporate, and very small (10-11 µm). The combined characters indicate a position within the grade known as subfamily Rauvolfioideae. CONCLUSIONS: These fossils, as the oldest remains of Rauvolfioids, complement the fossil records of Apocynoid and Asclepioid fossil seeds from other regions, demonstrating that the Apocynaceae were well established by the early Eocene, mostly consistent with prior divergence estimates for the phylogeny of this family. Potential pollinators, also preserved in the Cambay amber, include mosquitos, gnats, small moths, and stingless bees.

Boreotropical range expansion and long-distance dispersal explain two amphi-Pacific tropical disjunctions in Sabiaceae.

Sabiaceae comprises three genera and ca. 80 species with an amphi-Pacific tropical disjunct distribution. It has been unclear whether the family is monophyletic, where the family belongs within the angiosperm phylogeny, and when and how is present-day disjunct distribution originated. To address these questions, we conducted a phylogenetic analysis of Sabiaceae with comprehensive sampling of the family and basal eudicots using six chloroplast DNA loci (atpB, rbcL, matK, ndhF, atpB-rbcL and trnL-trnF). Our results support the monophyly of Sabiaceae s. l. that includes three genera: Meliosma Blume, Ophiocaryon Endl. and Sabia Colebr. The placement of Sabiaceae as sister to Proteales receives moderate bootstrap support, and is corroborated by various alternative hypothesis tests. Within Sabiaceae, Ophiocaryon and Sabia were resolved as strongly supported clades, whereas Meliosma was paraphyletic with Ophiocaryon nested within it. The biogeographically disjunct accessions of Meliosma alba (which is alternatively known as Kingsboroughia alba (Schltdl.) Liebm.) sampled from southwestern China and Mexico form a monophyletic group. Molecular dating and ancestral area reconstruction suggest a Eurasian origin of Sabiaceae in the late Cretaceous and a boreotropical range expansion during Paleogene. Southward migrations were inferred from continental Eurasia to the Malesian region in Sabia and in the Asian Meliosma, and from Central America to South America in the Neotropical clade of Meliosma in response to climatic cooling after the late Miocene. A long distance dispersal from Central America to tropical Asia was suggested during the time at the Neogene and Quaternary boundary in Meliosma alba (now recognized as Kingsboroughia alba). Our results also support the recognition of Kingsboroughia Liebm. as a distinct genus to maintain the monophyly of each of the genera: Meliosma, Ophiocaryon and Sabia. Kingsboroughia along with Meliosma and Ophiocaryon constitutes the subfamily Meliosmoideae Mast., while Sabia is the sole genus of Sabioideae Y.W. Law & Y.F. Wu.

Fruit Morphology and Anatomy of the Spondioid Anacardiaceae.

The Spondioideae subfamily of the Anacardiaceae is widely distributed today in tropical regions. Recent molecular phylogenetic investigations indicate that the Spondioideae are not monophyletic, but rather comprise at least two separate clades that are difficult to distinguish using vegetative and floral characters. Nevertheless, the syndrome of fruit characters traditionally used in identifying the subfamily is useful in discriminating genera of these clades and for identification of both modern and fossil anacardiaceous fruits. Here we document the morphology and anatomy of endocarps for representatives of all extant genera traditionally treated as Spondioideae, plus two genera that have been placed close to them in molecular investigations, Buchanania and Campnosperma. All genera are characterized by drupe-like fruits with sclerified stones that vary from uni- to multilocular depending on the genus. Germination modes vary throughout the Spondioideae. Some have characteristic plug-like opercula; others have recessed bilabiate germination valves, and still others open by apical flaps or simple slits. Although most currently recognized genera appear to be monophyletic, fruit morphology indicates that current circumscriptions of Cyrtocarpa, Poupartia and Tapirira are in need of revision.

Icacinaceae from the eocene of Western North America.

PREMISE OF THE STUDY: The Icacinaceae are a pantropical family of trees, shrubs, and climbers with an extensive Paleogene fossil record. Our improved understanding of phylogenetic relationships within the family provides an excellent context for investigating new fossil fruit and leaf material from the Eocene of western North America. METHODS: We examined fossils from early and middle Eocene sediments of western Wyoming, northeastern Utah, northwestern Colorado, and Oregon and compared them with extant species of Iodes and other icacinaceous genera as well as previously described fossils of the family. KEY RESULTS: Three new fossil species are described, including two based on endocarps (Iodes occidentalis sp. nov. and Icacinicaryites lottii sp. nov.) and one based on leaves (Goweria bluerimensis sp. nov.). The co-occurrence of I. occidentalis and G. bluerimensis suggests these might represent detached organs of a single species. A new genus, Biceratocarpum, is also established for morphologically distinct fossil fruits of Icacinaceae previously placed in Carpolithus. Biceratocarpum brownii gen. et comb. nov. resembles the London Clay species "Iodes" corniculata in possessing a pair of subapical protrusions. CONCLUSIONS: These fossils increase our knowledge of Icacinaceae in the Paleogene of North America and highlight the importance of the Northern Hemisphere in the early diversification of the family. They also document interchange with the Eocene flora of Europe and biogeographic connections with modern floras of Africa and Asia, where Icacinaceae are diverse today. The present-day restriction of this family to tropical regions offers ecological implications for the Eocene floras in which they occur.

Assessing the Fossil Record of Asterids in the Context of Our Current Phylogenetic Framework.

The fossil record provides good evidence for the minimum ages of important events in the diversification and geographic spread of Asteridae, with earliest examples extending back to the Turonian stage of the Late Cretaceous (~89 Ma). Some of the fossil identifications accepted in previous considerations of asterid phylogeny do not stand up to careful scrutiny. Nevertheless, among major clades of asterids, there is good evidence for a range of useful anchor points. Here we provide a synopsis of fossil occurrences that we consider as reliable representatives of modern Asterid families and genera. In addition, we provide new examples documented by fossil dispersed pollen investigated by both light and scanning electron microscopy studies including representatives of Loranthaceae, Amaranthaceae, Cornaceae (incl. Nyssa L., Mastixia Blume, Diplopanax Hand.-Mazz.), Sapotaceae, Ebenaceae, Ericaceae, Icacinaceae, Oleaceae, Asteraceae, Araliaceae, Adoxaceae and Caprifoliaceae from Paleogene sites in Greenland, western North America, and central Europe, and of Lamiaceae and Asteraceae from the middle to late Miocene northeastern China. We emphasize that dispersed pollen, taken along with megafossil and mesofossil data, continue to fill gaps in our knowledge of the paleobotanical record.

First occurrence of Cedrelospermum (Ulmaceae) in Asia and its biogeographic implications.

Cedrelospermum (Ulmaceae) is an extinct genus with extensive fossil records in Europe and North America. However, no fossil of the genus has been reported from Asia. Here we describe Cedrelospermum asiaticum L.B. Jia, Y.J. Huang et Z.K. Zhou sp. nov. based on compressed fruits from the late Miocene of Yunnan, southwestern China. The fossil fruits are characterized by an ovate fruit body adjoined by double wings, with the veins on the primary wing converging toward a stigmatic area. According to the historical geographic distribution of the genus, we hypothesize that Cedrelospermum originated in North America where both single-winged and double-winged fruits were reported. The single-winged form subsequently spread into Europe via the North Atlantic land bridge and the double-winged form dispersed into Asia via the Bering land bridge. From the Eocene to Oligocene, a southward retreat of the genus distribution probably took place, which coincided with the global surface cooling initiated during the Eocene-Oligocene transition. The extinction of Cedrelospermum from Asia may be related to the intensification of the East Asian monsoon.

Alnus subgenus Alnus in the Eocene of western North America based on leaves, associated catkins, pollen, and fruits.

PREMISE OF THE STUDY: The fossil record of alder (Alnus) is well known in the Cenozoic deposits throughout the northern hemisphere, based on numerous reports of the distinctive pollen, cone-like infructescences, staminate inflorescences, and leaves. However, our understanding of the systematic position of these fossils relative to the modern phylogeny of the genus has been limited because most fossils were described from only one organ. METHODS: We examined well-preserved leaves and associated fruiting and staminate catkins from the middle Eocene, Clarno Formation, Oregon, USA by stereomicroscopy. In situ and dispersed pollen were cleaned with HF and acetolized for light and scanning electron microscopy. KEY RESULTS: We reconstructed a new extinct species based on multiple organs and discuss significant phytogeographic and phylogenetic implications for Alnus. Alnus clarnoensis sp. nov. is described based on serrate leaves with 1-4 small teeth between each primary tooth, associated cone-like fruiting catkins with fruits in situ, and associated slender pollen catkins bearing in situ 3- to 6-pored pollen with arci between the pores. Combined investigations of each organ indicate that they probably derive from the same species and can be confidently attributed to subgenus Alnus Furlow based on leaf architecture and pollen pore number frequency. CONCLUSIONS: The Clarno fossils are most similar to the extant North American species of subgenus Alnus rather than to those from Asia and Europe, indicating that this modern subgenus was already distinct by the middle Eocene and that the intercontinental migration likely occurred earlier.

Ozakia, a new genus of winged fruit shared between the Miocene of Japan and western North America.

A new genus is recognized based on winged fruits with a single species shared between the Miocene of southwestern Honshu, Japan, and the Miocene of Oregon and Idaho, USA. Calyces of Ozakia emryi gen. et sp. n. were formerly attributed to Heptacodium (Caprifoliaceae) and Amelanchier (Rosaceae); however, newly recovered specimens reveal additional characters that contradict these assignments. The pedicellate fruits are obovate, tapering basally and truncate apically, with about 10 longitudinal ribs, a prominent epigynous synsepalous calyx of five lobes, each with a midvein and a pair of weaker, ascending intramarginal primary veins. The single style has a capitate stigma. Ozakia is considered to represent an extinct eudicot genus, the familial affinities of which remain uncertain. The eastern Asian-western North American disjunction of Ozakia occurrences suggests that this plant traversed the Beringia land bridge during or prior to the Middle Miocene. Relatively few extinct angiosperm genera are known as late as the Miocene.

Cenozoic seeds of Vitaceae reveal a deep history of extinction and dispersal in the Neotropics.

The remarkably diverse plant communities of the Neotropics are the result of diversification driven by multiple biotic (for example, speciation, extinction and dispersal) and abiotic (for example, climatic and tectonic) processes. However, in the absence of a well-preserved, thoroughly sampled and critically assessed fossil record, the associated processes of dispersal and extinction are poorly understood. We report an exceptional case study documenting patterns of extinction in the grape family (Vitaceae Juss.) on the basis of fossil seeds discovered in four Neotropical palaeofloras dated between 60 and 19 Ma. These include a new species that provides the earliest evidence of Vitaceae in the Western Hemisphere. Eight additional species reveal the former presence of major clades of the family that are currently absent from the Neotropics and elucidate previously unknown dispersal events. Our results indicate that regional extinction and dispersal have substantially impacted the evolutionary history of Vitaceae in the Neotropics. They also suggest that while the Neotropics have been dynamic centres of diversification through the Cenozoic, extant Neotropical botanical diversity has also been shaped by extensive extinction over the past 66 million years.

Oldest fruits of the grape family (Vitaceae) from the Late Cretaceous Deccan cherts of India.

PREMISE OF THE STUDY: Despite the inferred Cretaceous origin of the Vitaceae, fossils of the grape family are relatively young, with the oldest previously known examples limited to the Paleocene of Europe and North America. New fossil evidence indicates that the family was already present in India in the Late Cretaceous (Maastrichtian), about 10-15 million years before the tectonic collision of India with Eurasia. • METHODS: Fruits and seeds were investigated by serial sections and peels of chert from the Deccan Intertrappean beds of central India, and compared anatomically with those of extant genera. • KEY RESULTS: Indovitis chitaleyae gen. et sp. n. is described based on immature fruits bearing four to six seeds, and isolated mature seeds. The seeds possess paired ventral infolds and a dorsal chalaza, features diagnostic of the order Vitales. Characters of chalaza shape, infold morphology, and seed coat anatomy place I. chitaleyae within Vitaceae and favor a phylogenetic position either sister to the Vitis-Ampelocissus clade or sister to the Ampelopsis-Clematicissus-Rhoicissus clade. • CONCLUSIONS: Presence of the oldest known vitaceous fossils in the latest Cretaceous of India indicates a previously undocumented Gondwanan history and a possible southern hemisphere origin for the Vitales. An "out-of-India" scenario might explain the relatively sudden appearance of diverse Vitaceae in the Late Paleocene and Early Eocene of the Northern Hemisphere.

Fruits of Koelreuteria (Sapindaceae) from the Cenozoic throughout the northern hemisphere: their ecological, evolutionary, and biogeographic implications.

PREMISE OF THE STUDY: Koelreuteria (Sapindaceae) has four extant deciduous tree species, disjunctly distributed in eastern Asia and the Fiji Islands. While K. paniculata is widely cultivated, the biogeographic origin and evolutionary history of Koelreuteria remain unclear. METHODS: Fruits, pollen, wood, and leaves of closely related extant taxa were examined in comparison with fossil remains to evaluate the fossil record and biogeographic history of Koelreuteria. KEY RESULTS: Overall, characters of capsular fruits are more diagnostic than other organs for this genus. We describe two new species of fruit remains from the Eocene, K. taoana sp. nov. (northeastern China and far eastern Russia) and K. dilcheri sp. nov. (western United States), and give emended descriptions of three species: K. allenii (Lesq.) W. N. Edwards (early-late Eocene of the United States), K. macroptera (Kováts) W. N. Edwards (late Oligocene-early Pliocene of Europe), and K. miointegrifoliola Hu et R. W. Chaney (Miocene of eastern Asia). CONCLUSIONS: Reliable fossil records of capsules and ring-porous wood indicate that Koelreuteria may have originated in North Pacific-Rim area of the northern hemisphere by the early Eocene, representing an early temperate lineage in Sapindaceae adapted for wind dispersal. The fossils herein place a minimum age (ca. 52 Ma) for the divergence of Koelreuteria from tropical genera that appear more basal in the molecular phylogeny of Sapindaceae. Regional extinctions after the Eocene in North America and the Pliocene in Europe, reduced the range of Koelreuteria to eastern Asia, where three species occur today. The present distribution of another species in the Southern Pacific may be explained by long-distance dispersal.

Dated molecular phylogenies indicate a Miocene origin for Arabidopsis thaliana.

Dated molecular phylogenies are the basis for understanding species diversity and for linking changes in rates of diversification with historical events such as restructuring in developmental pathways, genome doubling, or dispersal onto a new continent. Valid fossil calibration points are essential to the accurate estimation of divergence dates, but for many groups of flowering plants fossil evidence is unavailable or limited. Arabidopsis thaliana, the primary genetic model in plant biology and the first plant to have its entire genome sequenced, belongs to one such group, the plant family Brassicaceae. Thus, the timing of A. thaliana evolution and the history of its genome have been controversial. We bring previously overlooked fossil evidence to bear on these questions and find the split between A. thaliana and Arabidopsis lyrata occurred about 13 Mya, and that the split between Arabidopsis and the Brassica complex (broccoli, cabbage, canola) occurred about 43 Mya. These estimates, which are two- to threefold older than previous estimates, indicate that gene, genomic, and developmental evolution occurred much more slowly than previously hypothesized and that Arabidopsis evolved during a period of warming rather than of cooling. We detected a 2- to 10-fold shift in species diversification rates on the branch uniting Brassicaceae with its sister families. The timing of this shift suggests a possible impact of the Cretaceous-Paleogene mass extinction on their radiation and that Brassicales codiversified with pierid butterflies that specialize on mustard-oil-producing plants.

Evolution of the intercontinental disjunctions in six continents in the Ampelopsis clade of the grape family (Vitaceae).

BACKGROUND: The Ampelopsis clade (Ampelopsis and its close allies) of the grape family Vitaceae contains ca. 43 species disjunctly distributed in Asia, Europe, North America, South America, Africa, and Australia, and is a rare example to study both the Northern and the Southern Hemisphere intercontinental disjunctions. We reconstruct the temporal and spatial diversification of the Ampelopsis clade to explore the evolutionary processes that have resulted in their intercontinental disjunctions in six continents. RESULTS: The Bayesian molecular clock dating and the likelihood ancestral area analyses suggest that the Ampelopsis clade most likely originated in North America with its crown group dated at 41.2 Ma (95% HPD 23.4-61.0 Ma) in the middle Eocene. Two independent Laurasian migrations into Eurasia are inferred to have occurred in the early Miocene via the North Atlantic land bridges. The ancestor of the Southern Hemisphere lineage migrated from North America to South America in the early Oligocene. The Gondwanan-like pattern of intercontinental disjunction is best explained by two long-distance dispersals: once from South America to Africa estimated at 30.5 Ma (95% HPD 16.9-45.9 Ma), and the other from South America to Australia dated to 19.2 Ma (95% HPD 6.7-22.3 Ma). CONCLUSIONS: The global disjunctions in the Ampelopsis clade are best explained by a diversification model of North American origin, two Laurasian migrations, one migration into South America, and two post-Gondwanan long-distance dispersals. These findings highlight the importance of both vicariance and long distance dispersal in shaping intercontinental disjunctions of flowering plants.

Palynological composition of a Lower Cretaceous South American tropical sequence: climatic implications and diversity comparisons with other latitudes.

PREMISE OF THE STUDY: Reconstruction of floristic patterns during the early diversification of angiosperms is impeded by the scarce fossil record, especially in tropical latitudes. Here we collected quantitative palynological data from a stratigraphic sequence in tropical South America to provide floristic and climatic insights into such tropical environments during the Early Cretaceous. METHODS: We reconstructed the floristic composition of an Aptian-Albian tropical sequence from central Colombia using quantitative palynology (rarefied species richness and abundance) and used it to infer its predominant climatic conditions. Additionally, we compared our results with available quantitative data from three other sequences encompassing 70 floristic assemblages to determine latitudinal diversity patterns. KEY RESULTS: Abundance of humidity indicators was higher than that of aridity indicators (61% vs. 10%). Additionally, we found an angiosperm latitudinal diversity gradient (LDG) for the Aptian, but not for the Albian, and an inverted LDG of the overall diversity for the Albian. Angiosperm species turnover during the Albian, however, was higher in humid tropics. CONCLUSIONS: There were humid climates in northwestern South America during the Aptian-Albian interval contrary to the widespread aridity expected for the tropical belt. The Albian inverted overall LDG is produced by a faster increase in per-sample angiosperm and pteridophyte diversity in temperate latitudes. However, humid tropical sequences had higher rates of floristic turnover suggesting a higher degree of morphological variation than in temperate regions.

Rosid radiation and the rapid rise of angiosperm-dominated forests.

The rosid clade (70,000 species) contains more than one-fourth of all angiosperm species and includes most lineages of extant temperate and tropical forest trees. Despite progress in elucidating relationships within the angiosperms, rosids remain the largest poorly resolved major clade; deep relationships within the rosids are particularly enigmatic. Based on parsimony and maximum likelihood (ML) analyses of separate and combined 12-gene (10 plastid genes, 2 nuclear; >18,000 bp) and plastid inverted repeat (IR; 24 genes and intervening spacers; >25,000 bp) datasets for >100 rosid species, we provide a greatly improved understanding of rosid phylogeny. Vitaceae are sister to all other rosids, which in turn form 2 large clades, each with a ML bootstrap value of 100%: (i) eurosids I (Fabidae) include the nitrogen-fixing clade, Celastrales, Huaceae, Zygophyllales, Malpighiales, and Oxalidales; and (ii) eurosids II (Malvidae) include Tapisciaceae, Brassicales, Malvales, Sapindales, Geraniales, Myrtales, Crossosomatales, and Picramniaceae. The rosid clade diversified rapidly into these major lineages, possibly over a period of <15 million years, and perhaps in as little as 4 to 5 million years. The timing of the inferred rapid radiation of rosids [108 to 91 million years ago (Mya) and 107-83 Mya for Fabidae and Malvidae, respectively] corresponds with the rapid rise of angiosperm-dominated forests and the concomitant diversification of other clades that inhabit these forests, including amphibians, ants, placental mammals, and ferns.

Phytogeographic implications of fossil endocarps of Menispermaceae from the Paleocene of Colombia.

PREMISE OF THE STUDY: Fossil leaves of Menispermaceae were previously described from the Paleocene of Colombia. Because of strong homoplasy of leaf characters, the fossils could not be placed more specifically within recognized clades, and additional data were needed to specify intrafamilial and paleogeographic relationships during the Paleocene. METHODS: Fossil endocarps of Menispermaceae were collected from the Cerrejón Formation, the recently discovered Bogotá flora, and Wyoming (∼60 Ma). We surveyed the endocarp morphology of almost all extant genera, conducted character optimization, a molecular scaffold analysis, and critically reviewed the related fossil genera. KEY RESULTS: Parallel syndromes of fruit characters have appeared in unrelated clades of the family according to current phylogenetic reconstructions. However, mapping selected endocarp characters across those clades that contain horseshoe-shaped endocarps facilitates identification and phylogenetic assessment of the fossils. Three fossil species are recognized. One of them belongs to the extant genus Stephania, which today grows only in Africa and Australasia. Palaeoluna gen. nov. is placed within the pantropical clade composed of extant Stephania, Cissampelos, and Cyclea; this morphogenus is also recognized from the Paleocene of Wyoming. Menispina gen. nov. shows similarity with several unrelated clades. CONCLUSIONS: The new fossils from Colombia reveal a complex paleobiogeographic history of the recognized clades within Menispermaceae, suggesting a more active exchange among neotropical, paleotropical, North American, and European paleoforests than previously recognized. In addition, the new fossils indicate that neotropical forests were an important biome for the radiation and dispersal of derived lineages in Menispermaceae after the Cretaceous-Paleogene boundary.

Biogeographic Overview of Ulmaceae: Diversity, Distribution, Ecological Preferences, and Conservation Status.

The elm family (Ulmaceae) is a woody plant group with important scientific, societal, and economic value. We aim to present the first biogeographic synthesis investigating the global diversity, distribution, ecological preferences, and the conservation status of Ulmaceae. A literature review was performed to explore the available data for all extant species. Our study made it possible to map the actual global distribution of Ulmaceae with high precision, and to elucidate the centers of diversity, located mainly in China and in the southeastern USA. A detailed comparative analysis of the macroclimatic niche for each species was produced, which shows the general biogeographic pattern of the family and pinpoints the outlier species. The results corroborate recent molecular analyses and support the division of Ulmaceae into two taxonomically, biogeographically, and ecologically well-differentiated groups: the so-called temperate clade with 4 genera and 43 species and the tropical clade with 3 genera and 13 species. The elm family is often described as a typical temperate plant group, however the diversity peak of all Ulmaceae is located in the subtropical zone, and a non-negligible part of the family is exclusively distributed in the tropics. We also noticed that a high proportion of Ulmaceae is linked to humid macro- or microhabitats. Finally, we highlighted that nearly 25% of all Ulmaceae are threatened. Fieldwork, conservation efforts, and research activities are still necessary for this family, particularly for the tropical members and the most endangered species.

Phylogeny and biogeography of Alangiaceae (Cornales) inferred from DNA sequences, morphology, and fossils.

Alangiaceae, in the basal Asterid clade Cornales, consists of only one genus, Alangium. The genus has approximately 24 species distributed mainly in tropical and subtropical regions of the Old World and is well represented in the Tertiary fossil record of the northern hemisphere. We conducted phylogenetic and biogeographic analyses for Alangium by integrating data from DNA sequences, morphology, and fossils to evaluate systematic and biogeographic hypotheses. The results largely agree with the traditional classification of four sections within the genus, and suggest sects. Conostigma and Rhytidandra are successive sister taxa to a clade containing sects. Marlea and Alangium. Our results also indicate that the widespread species A. chinense consists of at least two lineages meriting recognition as distinct species. Biogeographic analysis using DIVA and divergence time dating with the Bayesian method (MULTIDIVTIME) resolved the ancestor of Alangium as being in S.E. Asia in the Late Cretaceous. Several intercontinental migrations involving the margin of the Tethys seaway (TESW), the North Atlantic land bridge (NALB) or the Bering land bridge (BLB), and long-distance dispersals are suggested. The results support TESW for plant migration of thermophilic (including evergreen) taxa in the early Tertiary.

Correction: Reinvestigating an enigmatic Late Cretaceous monocot: morphology, taxonomy, and biogeography of Viracarpon.

[This corrects the article DOI: 10.7717/peerj.4580.].