Evolutionary persistence in Gunnera and the contribution of southern plant groups to the tropical Andes biodiversity hotspot.

Several studies have demonstrated the contribution of northern immigrants to the flora of the tropical Andes-the world's richest and most diverse biodiversity hotspot. However, much less is known about the biogeographic history and diversification of Andean groups with southern origins, although it has been suggested that northern and southern groups have contributed roughly equally to the high Andean (i.e., páramo) flora. Here we infer the evolutionary history of the southern hemisphere plant genus Gunnera, a lineage with a rich fossil history and an important ecological role as an early colonising species characteristic of wet, montane environments. Our results show striking contrasts in species diversification, where some species may have persisted for some 90 million years, and whereas others date to less than 2 Ma since origination. The outstanding longevity of the group is likely linked to a high degree of niche conservatism across its highly disjunct range, whereby Gunnera tracks damp and boggy soils in cool habitats. Colonisation of the northern Andes is related to Quaternary climate change, with subsequent rapid diversification appearing to be driven by their ability to take advantage of environmental opportunities. This study demonstrates the composite origin of a mega-diverse biota.

Floral development of Sabia (Sabiaceae): evidence for the derivation of pentamery from a trimerous ancestry.

UNLABELLED: • PREMISE OF THE STUDY: Flowers of Sabiaceae diverge from basal eudicots in combining pentamery with superposed whorls of sepals, petals, and stamens and are therefore crucial in understanding origins of core eudicot flowers. Different hypotheses are tested using floral developmental evidence, whether the pentamerous flower is derived from a spiral, trimerous, or dimerous progenitor.• METHODS: The floral development of two species of Sabia was investigated with the scanning electron microscope to understand their unusual floral morphology and the origin of pentamery.• KEY RESULTS: The species show major developmental differences in their inflorescence morphology and organ initiation sequence. In S. limoniacea, flowers are subtended by a pherophyll preceding two prophylls, one of which encloses a younger flower; floral organs arise in a continuous spiral sequence without interruption between different organs. The ovary is oriented in an oblique-median position. In S. japonica, one prophyll replaces one of the sepals, and there is a disruption in the spiral sequence. As a result, the ovary is inserted more or less transversally.• CONCLUSIONS: The flower of Sabiaceae is structurally best interpreted as derived from a trimerous progenitor, and a derivation from a dimerous or spiral progenitor is less likely. One organ of each median adaxial whorl is interpreted as lost (from K3+3 C3+3 A3+3 G3 to K3+2 C3+2 A3+2 G2). The number of sepals is variable as pherophylls, prophylls, and sepals cannot be distinguished by shape and intergrade with each other. The floral organization of Sabia is reminiscent of trimerous Ranunculales and supports an earlier divergence of Sabiaceae relative to Proteales.

Floral development and anatomy of Salvadoraceae.

UNLABELLED: BACKGROUND AND AIMS; This study is an investigation into the floral development and anatomy of two genera of the small family Salvadoraceae, which belongs to the Brassicales in a clade with Batis and Koeberlinia. Salvadoraceae remains little known, despite its wide distribution in arid areas of the globe. Floral morphological data are scarce, and information on floral anatomy is limited to a single study, although morphological and anatomical characters are now used increasingly as a counterpart of molecular data. There remain a number of controversial morphological questions, such as the fusion of the petals, the number of carpels and the nature of the nectaries. METHODS: Floral anatomy and ontogeny were studied in two species of Salvadora and one species of Dobera. Only for S. persica could a full floral developmental sequence be done. KEY RESULTS: The floral development demonstrates that the ovary of Salvadoraceae is basically bicarpellate and pseudomonomerous with a single locule and parietal placenta. The ovary of Dobera resembles Azima tetracantha in the presence of a false apical septum. Evidence for a staminodial nature of the nectaries is not decisive. In Salvadora petals and stamens are lifted by a short hypanthium. CONCLUSIONS: Salvadoraceae share several morphological and developmental synapomorphies with Batis (Bataceae) and possibly Koeberlinia (Koeberliniaceae), supporting their close relationship as indicated by molecular phylogeny.

Perianth evolution in the sandalwood order Santalales.

Flowers of Santalales remain largely unexplored with several questions of homology unanswered despite the large size of the order. Morphological and ontogenetic floral studies have the potential to identify new informative characters. We studied floral development in species of Loranthaceae, Santalaceae, Opiliaceae, and "Olacaceae" with scanning electron microscopy to clarify the origin and evolution of the perianth in Santalales. The perianth is either dichlamydeous or when monochlamydeous interpreted as sepals, petals, or tepals. A girdling calyculus of debatable origin is found in some clades. We show that species of Diogoa and Heisteria in "Olacaceae" have a dichlamydeous perianth, unlike Olax in which the calyx is replaced by a calyculus. The calyculus arises by development of two lateral primordia, supporting the hypothesis of bracteole origin. A calyculus with similar development is present in species of Loranthaceae and possibly of Opiliaceae, suggesting a position of Olax closer to these families than to traditional genera of "Olacaceae". The monochlamydeous perianth in Santalaceae is shown to correspond to petals of other members of Santalales. Flower ontogenetic evidence suggests a repeated loss of the calyx, replacement by a calyculus, and further loss, leading to monochlamydeous perianths in Santalaceae.

Flower development of Meliosma (Sabiaceae): evidence for multiple origins of pentamery in the eudicots.

Flower developmental studies are a complement to molecular phylogenetics and a tool to understand the evolution of the angiosperm flower. Buds and mature flowers of Meliosma veitchiorum, M. cuneifolia, and M. dilleniifolia (Sabiaceae) were investigated using scanning electron microscopy to clarify flower developmental patterns and morphology, to understand the origin of the perianth merism, and to discuss the two taxonomic positions proposed for Sabiaceae, among rosids or in the basal grade of eudicots. Flowers in Meliosma appear pentamerous with two of the five sepals and petals strongly reduced, three staminodes alternating with two fertile stamens opposite the small petals, and a two-carpellate gynoecium. The flower development in Meliosma is spiral without distinction between bracteoles and sepals. Because of this development, sepals, petals, and stamens are almost opposite and not alternating as expected in cyclical pentamerous flowers. In four-sepal flowers the direction of petal initiation is reversed. The symmetry of the flower appears to be transversally zygomorphic, although this is hidden by the almost equal size of the larger petals. Evidence points to a unique pentamerous origin of flowers in Meliosma, and not to a trimerous origin, as earlier suggested, and adds support to multiple origins of pentamery in the eudicots.

Wax plants disentangled: a phylogeny of Hoya (Marsdenieae, Apocynaceae) inferred from nuclear and chloroplast DNA sequences.

Hoya (Marsdenieae, Apocynaceae) includes at least 200 species distributed from India to the Pacific Islands. We here infer major species groups in the genus based on combined sequences from the chloroplast atpB-rbcL spacer, the trnL region, and nuclear ribosomal DNA ITS region for 42 taxa of Hoya and close relatives. To assess levels of ITS polymorphism, ITS sequences for a third of the accessions were obtained by cloning. Most ITS clones grouped by species, indicating that speciation in Hoya usually predates ITS duplication. One ITS sequence of H. carnosa, however, grouped with a sequence of the morphologically similar H. pubicalyx, pointing to recent hybridization or the persistence of paralogous copies through a speciation event. The topology resulting from the combined chloroplast and nuclear data recovers some morphology-based sections, such as Acanthostemma and Eriostemma, as well as a well-supported Australian/New Guinean clade. The combined data also suggest that morphological adaptations for ant-symbiosis evolved at least three times within Hoya.