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.

Revision of the Lima clade (Miconia sect. Lima, Miconieae, Melastomataceae) of the Greater Antilles.

Miconia sect. Lima is an entirely Greater Antillean clade that consists of 19 known species of shrubs and small trees, which were previously recognized under the polyphyletic genera Leandra and Ossaea. The highest species richness in the clade is represented on Cuba (10 species), followed by Hispaniola (8 species) and then Jamaica (1 species). Here we present a taxonomic revision of the clade based on the study of species in the field, herbarium specimens, as well as a DNA-based phylogeny reconstruction. The Lima clade most likely originated on Cuba and then spread to Jamaica once and Hispaniola multiple times. Species of this clade can be recognized by the well developed bulla-based hairs of the adaxial leaf surface, as well as the clavate-dendritic hairs produced along the primary, secondary and tertiary veins of the adaxial leaf surface, mostly towards the leaf base, terminal inflorescences, acute petal apices, slightly bulla-based hairs produced subapically along the petal abaxial surface, and anthers with a dorso-basal appendage and a single, dorsally oriented pore. Descriptions, synonymies, along with distribution maps and illustrations/figures, are given for each species. Miconia pagnolensissp. nov. is newly described in this revision.

Miconia bullotricha and M. hirtistyla, two new species of Miconia sect. Lima (Miconieae, Melastomataceae) from eastern Cuba.

We describe two new species in Miconia sect. Lima, Miconia bullotricha Bécquer & Majure and Miconia hirtistyla Majure & Judd, from eastern, Cuba. We also provide illustrations and distribution maps for the two species, as well as a key to members of the Lima clade on Cuba.

ResumenDescribimos dos nuevas especies de Miconia sect. Lima, Miconia bullotricha Bécquer & Majure y Miconia hirtistyla Majure & Judd, del este de Cuba. También, proveemos ilustraciones y mapas de distribución para las dos especies, y una clave para distinguir los miembros del clado Lima en Cuba.

A revised circumscription for the Blakeeae (Melastomataceae) with associated nomenclatural adjustments.

Systematic investigations and phylogenetic analyses of the Blakeeae (Melastomataceae) have indicated that Topobea should be synonymized under Blakea, and Huilaea under Chalybea. Presented here is a detailed description of the Blakeeae, a key to its two accepted genera, and a listing of 62 new combinations, including 3 new names, necessitated by the transfer of Topobea as follows: Blakea acuminata (Wurdack) Penneys & Judd, comb. nov., Blakea adscendens (E.Cotton & Matezki) Penneys & Judd, comb. nov., Blakea albertiae (Wurdack) Penneys & Almeda, comb. nov., Blakea amplifolia (Almeda) Penneys & Almeda, comb. nov., Blakea arboricola (Almeda) Penneys & Almeda, comb. nov., Blakea asplundii (Wurdack) Penneys & Judd, comb. nov., Blakea barbata (Gleason) Penneys & Judd, comb. nov., Blakea brenesii (Standl.) Penneys & Almeda, comb. nov., Blakea brevibractea (Gleason) Penneys & Judd, comb. nov., Blakea bullata (E.Cotton & Matezki) Penneys & Judd, comb. nov., Blakea calcarata (L.Uribe) Penneys & Judd, comb. nov., Blakea calophylla (Almeda) Penneys & Almeda, comb. nov., Blakea calycularis (Naudin) Penneys & Almeda, comb. nov., Blakea castanedae (Wurdack) Penneys & Judd, comb. nov., Blakea clavata (Triana) Penneys & Judd, nom. nov., Blakea cordata (Gleason) Penneys & Almeda, comb. nov., Blakea cuprina Penneys & Judd, nom. nov., Blakea cutucuensis (Wurdack) Penneys & Judd, comb. nov., Blakea dimorphophylla (Almeda) Penneys & Almeda, comb. nov., Blakea discolor (Hochr.) Penneys & Judd, comb. nov., Blakea dodsonorum (Wurdack) Penneys & Almeda, comb. nov., Blakea eplingii (Wurdack) Penneys & Judd, comb. nov., Blakea ferruginea (Gleason) Penneys & Judd, comb. nov., Blakea fragrantissima (Almeda) Penneys & Almeda, comb. nov., Blakea gerardoana (Almeda) Penneys & Almeda, comb. nov., Blakea glaberrima (Triana) Penneys & Judd, comb. nov., Blakea henripittieri (Cogn.) Penneys & Almeda, comb. et nom. nov., Blakea hexandra (Almeda) Penneys & Almeda, comb. nov., Blakea horologica Penneys & Judd, nom. nov., Blakea induta (Markgr.) Penneys & Judd, comb. nov., Blakea inflata (Triana) Penneys & Judd, comb. nov., Blakea insignis (Triana) Penneys & Judd, comb. nov., Blakea intricata (Almeda) Penneys & Almeda, comb. nov., Blakea killipii (Wurdack) Penneys & Judd, comb. nov., Blakea lentii (Almeda) Penneys & Almeda, comb. nov., Blakea longiloba (Wurdack) Penneys & Judd, comb. nov., Blakea longisepala (Gleason) Penneys & Judd, comb. nov., Blakea macbrydei (Wurdack) Penneys & Judd, comb. nov., Blakea maguirei (Wurdack) Penneys & Judd, comb. nov., Blakea maurofernandeziana (Cogn.) Penneys & Almeda, comb. nov., Blakea mcphersonii (Almeda) Penneys & Almeda, comb. nov., Blakea modica (Wurdack) Penneys & Judd, comb. nov., Blakea mortoniana (Wurdack) Penneys & Judd, comb. nov., Blakea muricata (Lozano) Penneys & Judd, comb. nov., Blakea pascoensis (Wurdack) Penneys & Judd, comb. nov., Blakea pluvialis (Standl.) Penneys & Almeda, comb. nov., Blakea sessilifolia (Triana) Penneys & Judd, comb. nov., Blakea setosa (Triana) Penneys & Judd, comb. nov., Blakea standleyi (L.O.Williams) Penneys & Almeda, comb. nov., Blakea stephanochaeta (Naudin) Penneys & Judd, comb. nov., Blakea steyermarkii (Wurdack) Penneys & Judd, comb. nov., Blakea suaveolens (Almeda) Penneys & Almeda, comb. nov., Blakea subbarbata (Wurdack) Penneys & Judd, comb. nov., Blakea subscabrula (Triana) Penneys & Judd, comb. nov., Blakea subsessiliflora (Wurdack) Penneys & Judd, comb. nov., Blakea superba (Naudin) Penneys & Judd, comb. nov., Blakea tetramera (Almeda) Penneys & Almeda, comb. nov., Blakea tetroici (Wurdack) Penneys & Judd, comb. nov., Blakea toachiensis (Wurdack) Penneys & Judd, comb. nov., Blakea trianae (Cogn.) Penneys & Judd, comb. nov., Blakea verrucosa (Wurdack) Penneys & Judd, comb. nov., Blakea watsonii (Cogn.) Penneys & Almeda, comb. nov.

Phylogeny of Opuntia s.s. (Cactaceae): clade delineation, geographic origins, and reticulate evolution.

PREMISE OF THE STUDY: The opuntias (nopales, prickly pears) are not only culturally, ecologically, economically, and medicinally important, but are renowned for their taxonomic difficulty due to interspecific hybridization, polyploidy, and morphological variability. Evolutionary relationships in these stem succulents have been insufficiently studied; thus, delimitation of Opuntia s.s. and major subclades, as well as the biogeographic history of this enigmatic group, remain unresolved. METHODS: We sequenced the plastid intergenic spacers atpB-rbcL, ndhF-rpl32, psbJ-petA, and trnL-trnF, the plastid genes matK and ycf1, the nuclear gene ppc, and ITS to reconstruct the phylogeny of tribe Opuntieae, including Opuntia s.s. We used phylogenetic hypotheses to infer the biogeographic history, divergence times, and potential reticulate evolution of Opuntieae. KEY RESULTS: Within Opuntieae, a clade of Tacinga, Opuntia lilae, Brasiliopuntia, and O. schickendantzii is sister to a well-supported Opuntia s.s., which includes Nopalea. Opuntia s.s. originated in southwestern South America (SA) and then expanded to the Central Andean Valleys and the desert region of western North America (NA). Two major clades evolved in NA, which subsequently diversified into eight subclades. These expanded north to Canada and south to Central America and the Caribbean, eventually returning back to SA primarily via allopolyploid taxa. Dating approaches suggest that most of the major subclades in Opuntia s.s. originated during the Pliocene. CONCLUSIONS: Opuntia s.s. is a well-supported clade that includes Nopalea. The clade originated in southwestern SA, but the NA radiation was the most extensive, resulting in broad morphological diversity and frequent species formation through reticulate evolution and polyploidy.

Cytogeography of the Humifusa clade of Opuntia s.s. Mill. 1754 (Cactaceae, Opuntioideae, Opuntieae): correlations with pleistocene refugia and morphological traits in a polyploid complex.

Ploidy has been well studied and used extensively in the genus Opuntia to determine species boundaries, detect evidence of hybridization, and infer evolutionary patterns. We carried out chromosome counts for all members of the Humifusa clade to ascertain whether geographic patterns are associated with differences in ploidy. We then related chromosomal data to observed morphological variability, polyploid formation, and consequently the evolutionary history of the clade. We counted chromosomes of 277 individuals from throughout the ranges of taxa included within the Humifusa clade, with emphasis placed on the widely distributed species, Opuntia humifusa (Raf.) Raf., 1820 s.l. and Opuntia macrorhiza Engelm., 1850 s.l. We also compiled previous counts made for species in the clade along with our new counts to plot geographic distributions of the polyploid and diploid taxa. A phylogeny using nuclear ribosomal ITS sequence data was reconstructed to determine whether ploidal variation is consistent with cladogenesis. We discovered that diploids of the Humifusa clade are restricted to the southeastern United States (U.S.), eastern Texas, and southeastern New Mexico. Polyploid members of the clade, however, are much more widely distributed, occurring as far north as the upper midwestern U.S. (e.g., Michigan, Minnesota, Wisconsin). Morphological differentiation, although sometimes cryptic, is commonly observed among diploid and polyploid cytotypes, and such morphological distinctions may be useful in diagnosing possible cryptic species. Certain polyploid populations of Opuntia humifusa s.l. and Opuntia macrorhiza s.l., however, exhibit introgressive morphological characters, complicating species delineations. Phylogenetically, the Humifusa clade forms two subclades that are distributed, respectively, in the southeastern U.S. (including all southeastern U.S. diploids, polyploid Opuntia abjecta Small, 1923, and polyploid Opuntia pusilla (Haw.) Haw., 1812) and the southwestern U.S. (including all southwestern U.S. diploids and polyploids). In addition, tetraploid Opuntia humifusa s.l., which occurs primarily in the eastern U.S., is resolved in the southwestern diploid clade instead of with the southeastern diploid clade that includes diploid Opuntia humifusa s.l. Our results not only provide evidence for the polyphyletic nature of Opuntia humifusa and Opuntia macrorhiza, suggesting that each of these represents more than one species, but also demonstrate the high frequency of polyploidy in the Humifusa clade and the major role that genome duplication has played in the diversification of this lineage of Opuntia s.s. Our data also suggest that the southeastern and southwestern U.S. may represent glacial refugia for diploid members of this clade and that the clade as a whole should be considered a mature polyploid species complex. Widespread polyploids are likely derivatives of secondary contact among southeastern and southwestern diploid taxa as a result of the expansion and contraction of suitable habitat during the Pleistocene following glacial and interglacial events.

Angiosperm phylogeny: 17 genes, 640 taxa.

PREMISE OF THE STUDY: Recent analyses employing up to five genes have provided numerous insights into angiosperm phylogeny, but many relationships have remained unresolved or poorly supported. In the hope of improving our understanding of angiosperm phylogeny, we expanded sampling of taxa and genes beyond previous analyses. METHODS: We conducted two primary analyses based on 640 species representing 330 families. The first included 25260 aligned base pairs (bp) from 17 genes (representing all three plant genomes, i.e., nucleus, plastid, and mitochondrion). The second included 19846 aligned bp from 13 genes (representing only the nucleus and plastid). KEY RESULTS: Many important questions of deep-level relationships in the nonmonocot angiosperms have now been resolved with strong support. Amborellaceae, Nymphaeales, and Austrobaileyales are successive sisters to the remaining angiosperms (Mesangiospermae), which are resolved into Chloranthales + Magnoliidae as sister to Monocotyledoneae + [Ceratophyllaceae + Eudicotyledoneae]. Eudicotyledoneae contains a basal grade subtending Gunneridae. Within Gunneridae, Gunnerales are sister to the remainder (Pentapetalae), which comprises (1) Superrosidae, consisting of Rosidae (including Vitaceae) and Saxifragales; and (2) Superasteridae, comprising Berberidopsidales, Santalales, Caryophyllales, Asteridae, and, based on this study, Dilleniaceae (although other recent analyses disagree with this placement). Within the major subclades of Pentapetalae, most deep-level relationships are resolved with strong support. CONCLUSIONS: Our analyses confirm that with large amounts of sequence data, most deep-level relationships within the angiosperms can be resolved. We anticipate that this well-resolved angiosperm tree will be of broad utility for many areas of biology, including physiology, ecology, paleobiology, and genomics.

Molecular phylogeny, biogeography, and systematics of Dicerandra (Lamiaceae), a genus endemic to the southeastern United States.

Dicerandra, an endemic mint of the southeastern United States, comprises nine species, all of which are threatened or endangered and restricted to sandhill vegetation and a mosaic of scrub habitats. Molecular phylogenetic analyses of Dicerandra based on data from the nuclear and plastid genomes for all 13 taxa of the genus, identified two strongly supported clades, corresponding to the four annual and to the five perennial species of Dicerandra. However, the nuclear and plastid trees were incongruent in their placement of two perennial taxa, D. cornutissima and D. immaculata var. savannarum, perhaps due to ancient hybridization or to lineage sorting. Based on these analyses, the widespread D. linearifolia is not monophyletic, with populations of D. linearifolia var. linearifolia falling into either western or eastern clades. The western clade, comprising populations of D. linearifolia var. linearifolia and var. robustior, occurs in an area drained by rivers flowing toward the Gulf of Mexico, whereas the eastern clade, comprising populations of D. linearifolia var. linearifolia, D. densiflora, D. odoratissima, and D. radfordiana (i.e., all the annual species), occupies a region drained by rivers flowing to the Atlantic Ocean. Although this pattern of genetic differentiation between populations from these two river drainages has been documented in several animal species, it has not previously been reported for plants. A revised subgeneric classification is presented to reflect the annual and perennial clades.

Infrageneric phylogeny of Schoenocaulon (Liliales: Melanthiaceae) with clarification of cryptic species based on ITS sequence data and geographical distribution.

As currently defined, the 24 species of Schoenocaulon occur in three disjunct areas: north central Florida (one species, S. dubium), southern Peru (portion of the range of S. officinale), and the region from southeastern New Mexico-Texas south to Venezuela; the 20 species endemic to Mexico are geographically restricted. Species delimitations, often based on tepal morphology, have been problematic. Our analyses of ITS sequence data for all 27 species and infraspecific taxa support recognition of two new species and recircumscription and placement of elements of the polyphyletic S. ghiesbreghtii and S. mortonii complexes. For taxa with adequate sampling, our data also indicate 11-12 cladospecies and 3-6 metaspecies according to the apomorphic species concept. The resolved phylogeny, correlated with geography and morphology, allows insight into biogeographical diversification and the evolution of some unusual morphological characters within the genus, such as nectary differentiation and tepal margin type.

A survey of tricolpate (eudicot) phylogenetic relationships.

The phylogenetic structure of the tricolpate clade (or eudicots) is presented through a survey of their major subclades, each of which is briefly characterized. The tricolpate clade was first recognized in 1989 and has received extensive phylogenetic study. Its major subclades, recognized at ordinal and familial ranks, are now apparent. Ordinal and many other suprafamilial clades are briefly diagnosed, i.e., the putative phenotypic synapomorphies for each major clade of tricolpates are listed, and the support for the monophyly of each clade is assessed, mainly through citation of the pertinent molecular phylogenetic literature. The classification of the Angiosperm Phylogeny Group (APG II) expresses the current state of our knowledge of phylogenetic relationships among tricolpates, and many of the major tricolpate clades can be diagnosed morphologically.

An extinct genus of Salicaceae based on twigs with attached flowers, fruits, and foliage from the Eocene Green River Formation of Utah and Colorado, USA.

A newly recovered twig with attached leaves and flowers from the Eocene Green River Formation of Utah provides the basis for recognizing a new, extinct genus of Salicaceae sensu lato (s.l.). Pseudosalix handleyi gen. et sp. nov. has alternate lanceolate leaves with pinnate, semicraspedodromous venation and a serrate margin with glandular teeth. The inflorescence is terminal on the twig and is unisexual, composed of flowers organized in a paniculoid cyme, with lateral paraclades of pedicellate flowers. The attached pistillate flowers have four prominent sepals that are valvate in bud, spreading but basally fused at anthesis; the single pistil of each flower is ovoid with three or four longitudinal sutures, indicating development to a capsular fruit. Three or four recurved styles radiate from the apex of the pistil, each with a distal globose stigma. The infructescence, verified by attachment to twigs with the same kind of leaves, bore capsular fruits of three and four valves. Associated but unattached, staminate flowers also have four well-developed, basally connate sepals. They are pedicellate and bear several stamens, each with a short filament and globose anther. The available morphological characters place the fossil species within the Salicaceae s.l. as an immediate sister to the clade containing Populus and Salix. Although the likely outgroup genera (including Itoa, Poliothyrsis, Carrierea, and Idesia) to tribe Saliceae all occur in Asia today and not North America, the occurrence of both Pseudosalix and Populus in the Eocene of Utah raises the possibility of a North American origin for the Saliceae.