Leaf adaptations and species boundaries in North American Cercis: implications for the evolution of dry floras.

PREMISE OF THE STUDY: The North American Cercis clade spans dry to mesic climates and exhibits complex morphological variation. We tested various proposed species classifications of this group and whether aspects of leaf morphology, particularly the "drip-tip" in some regional populations, are adaptive and/or linked with phylogeny. METHODS: We made measurements on over 1100 herbarium specimens from throughout North America and analyzed the data with univariate and multivariate approaches. We analyzed phylogenetically DNA sequence data from nuclear ITS and three plastid regions from 40 samples, and estimated divergence times with a relaxed-clock Bayesian analysis. We used climate and geographic position data to predict the variation observed in leaf size and shape by using stepwise multiple linear regressions. KEY RESULTS: Morphometric analyses yielded a pattern of continuous and often clinal character variation across North America, without correlated gaps in character states. Conversely, phylogenetic and divergence time analyses yielded distinct clades from California, the interior west, and eastern North America separated by between ~12 and 16 million years. Multiple regressions yielded highly significant correlations between leaf apex shape and precipitation of the warmest quarter. CONCLUSIONS: Despite a pattern of continuous morphological character variation, the long period of geographic and presumably genetic isolation warrants the delimitation of three species. Predictive modeling supports the adaptive value of acuminate apices or "drip-tips" in mesic habitats. This suggests that Cercis leaves change more rapidly than inferred from parsimony reconstruction, which has implications for the evolution of the dry floras of North America and Eurasia.

Phylogeny of Cercis based on DNA sequences of nuclear ITS and four plastid regions: implications for transatlantic historical biogeography.

The disjunct genus Cercis has been used to test models of Northern Hemisphere historical biogeography. Previous phylogenetic estimates employing DNA sequences of the ITS region and (in one study) those of ndhF recovered a well supported clade of North American and western Eurasian species that was nested within a paraphyletic group of Chinese species. Resolution and clade support within the tree were otherwise low and the monophyly of Cercis canadensis was uncertain. Here we conduct a phylogenetic analysis of Cercis with a higher number of regions (ITS, ndhF, rpoB-trnC, trnT-trnD, and trnS-trnG) and samples than in previous studies. Results corroborate the initial divergence between the Chinese species Cercis chingii and the rest of the genus. Support is newly found both for a clade of the two North American species as sister to the western Eurasian species, and for the monophyly of C. canadensis. As in a previous study, divergence between North American and western Eurasian Cercis was estimated as mid-Miocene (ca. 13 million years ago), and the ancestor in which this divergence occurred was inferred to be xerophytic. Contrary to previous studies, however, our data infer strictly east-to-west vicariance. The timing of the transatlantic divergence in Cercis is too recent to be explained by a postulated continuous belt of semi-arid vegetation between North America and Europe in the Paleogene, suggesting instead the presence of a Miocene North Atlantic corridor for semi-arid plants. In the absence of strong evidence from other sources, the possibility that Cercis has been able to quickly adapt from mesophytic antecedents to semi-arid conditions whenever the latter have arisen in the Northern Hemisphere can be considered a plausible alternative, although parsimony optimization renders this scenario two steps longer.

Reticulate evolution, cryptic species, and character convergence in the core East Asian clade of Gaultheria (Ericaceae).

Phylogenetic relationships of 84 samples representing 30 species in the core East Asian clade of the wintergreen group of Gaultheria (Angiospermae: Ericaceae: Gaultherieae) were estimated from separate and combined DNA sequence data from five genic regions (ITS, matK, rpl16, trnL-trnF, and trnS-trnG) with parsimony, likelihood and Bayesian analyses. Two major clades were recovered, one comprising several sections and series with leaves generally more than 1 cm long [the ser. Leucothoides sensu lato (s.l.) clade] and another comprising the species of ser. Trichophyllae, with leaves generally less than 1 cm long. The ITS region yielded little phylogenetic resolution, whereas in the combined chloroplast analysis the samples from individual morphospecies in both clades were often nonmonophyletic. This was postulated to result from reticulate evolution in the ser. Leucothoides s.l. clade, particularly in two specific cases of hybridization and a crown clade with likely chloroplast capture following localized introgression. In the ser. Trichophyllae clade, such nonmonophyly was largely attributed to cryptic species and character convergence resulting at least partly from extreme morphological reduction. The relatively low-elevation habitats in which the species of the ser. Leucothoides s.l. clade generally grow are thought to have promoted opportunities for sympatry and reticulation, whereas the high-alpine habitats of ser. Trichophyllae are more likely to have spawned isolated populations and narrow endemism. As in other Sino-Himalayan plant groups, overall low sequence divergence and reticulate evolution suggest rapid radiation in the core East Asian clade of Gaultheria.

Phylogeny of Caragana (Fabaceae) based on DNA sequence data from rbcL, trnS-trnG, and ITS.

Phylogenetic relationships of 48 species of Caragana (Fabaceae: tribe Hedysareae) and one representative each of Astragalus, Calophaca, Halimodendron, and Hedysarum are estimated from DNA sequences of the rbcL gene, trnS-trnG intron and spacer, and ITS region. At least one representative of all five sections and 12 series within Caragana are included. Analyses yielded strongly supported clades corresponding to sections Caragana, Bracteolatae, and Frutescentes. The species of section Jubatae are distributed among three strongly supported clades, i.e., one with the species of section Bracteolatae, another with two species of section Spinosae, and a third as sister to section Frutescentes. All but the last of these six clades are corroborated by at least one unambiguously traced morphological character. The placement of the other four species of section Spinosae are not well supported and lack unambiguous morphological synapomorphies, and the samples of Calophaca and Halimodendron nest within Caragana with weak support.

Phylogeny of Sinojackia (Styracaceae) based on DNA sequence and microsatellite data: implications for taxonomy and conservation.

BACKGROUND AND AIMS: The genus Sinojackia consists of eight species, all endemic to China. All species of Sinojackia are endangered or threatened owing to poor recruitment within populations. Information on molecular phylogenetics is critical for developing successful conservation strategies for this genus. METHODS: Combined DNA sequence data from the nuclear ribosomal internal transcribed spacer regions and plastid psbA-trnH intergenic spacer and microsatellite data were used to infer a phylogeny of the genus. KEY RESULTS: Parsimony analysis of the combined sequence data and multivariate analysis based on fruit characters indicated that Sinojackia dolichocarpa is monophyletic and genetically well separated from the other Sinojackia species, thus supporting its rank at the generic level as Changiostyrax. Phylogenetic relationships within Sinojackia sensu stricto are unresolved from the combined sequence data. A UPGMA dendrogram based on seven microsatellite loci of 96 individual plants yielded a first-diverging cluster of all individuals of S. microcarpa. The remaining species form another cluster without any definitive patterns corresponding to current species circumscriptions, suggesting either extensive hybridization or incipient speciation. CONCLUSIONS: The results suggest that there are too many species recognized within Sinojackia sensu stricto, but this must be further assessed with comprehensive morphological and taxonomic revisionary work. The implications of the phylogenetic data for conservation are discussed.

Phylogeny and circumscription of the near-endemic Brazilian tribe Microlicieae (Melastomataceae).

The members of tribe Microlicieae in the flowering plant family Melastomataceae are nearly all endemic to the cerrado biome of Brazil. Traditional classifications of the Melastomataceae have attributed between 15 and 17 genera to the Microlicieae, but subsequent revisions have circumscribed the tribe more narrowly. The monophyly and intergeneric relationships of the Microlicieae were evaluated through phylogenetic analyses with molecular and morphological data sets. Incorporation of DNA sequences from the intron of the chloroplast gene rpl16 into a previously generated family-wide data set yielded a clade comprising Chaetostoma, Lavoisiera, Microlicia, Rhynchanthera, Stenodon, and Trembleya ("core Microlicieae"), with Rhynchanthera as the first-diverging lineage. The other four genera of Microlicieae sampled are placed in other clades: Eriocnema with Miconieae; Siphanthera with Aciotis, Nepsera, and Acisanthera of Melastomeae; Castratella as sister to Monochaetum of Melastomeae; and Cambessedesia as part of an unresolved polytomy in a large clade that includes most Melastomataceae. Analyses of the chloroplast genes rbcL and ndhF that included three core genera produced similar results, as did the combined analysis of all three data sets. Combined parsimony analyses of DNA sequences from rpl16 and the nuclear ribosomal intercistronic transcribed spacer (ITS) region of 22 species of core Microlicieae yielded generally low internal support values. Lavoisiera, recently redefined on the basis of several morphological characters, was strongly supported as monophyletic. A morphological phylogenetic analysis of the Microlicieae based on 10 parsimony-informative characters recovered a monophyletic core Microlicieae but provided no further resolution among genera. Penalized likelihood analysis with two calibration time windows produced an age estimate of 3.7 million years for the time of initial divergence of strictly Brazilian core Microlicieae. This date is in general agreement with the estimated age of the most active stage of development of cerrado vegetation and implies an adaptive shift from hydric to seasonally dry habitats during the early evolution of this group.

Phylogeny and infrageneric classification of Symplocos (Symplocaceae) inferred from DNA sequence data.

Symplocos comprises ∼300 species of woody flowering plants with a disjunct distribution between the warm-temperate to tropical regions of eastern Asia and the Americas. Phylogenetic analyses of 111 species of Symplocos based on the nuclear ribosomal internal transcribed spacer (ITS) region and the chloroplast genes rpl16, matK, and trnL-trnF yielded topologies in which only one of the four traditionally recognized subgenera (Epigenia; Neotropics) is monophyletic. Section Cordyloblaste (subgenus Symplocos; eastern Asia) is monophyletic and sister to a group comprising all other samples of Symplocos. Section Palura (subgenus Hopea; eastern Asia) is sister to a group comprising all other samples of Symplocos except those of section Cordyloblaste. Symplocos wikstroemiifolia (eastern Asia) and S. tinctoria (southeastern United States), both of subgenus Hopea, form a clade that groups with S. longipes (tropical North America) and the species of subgenus Epigenia. The remaining samples of subgenus Hopea (eastern Asia) form a clade. Section Neosymplocos (subgenus Microsymplocos; Neotropics) is well nested within a clade otherwise comprising the samples of section Symplocastrum (subgenus Symplocos; Neotropics). Section Urbaniocharis (subgenus Microsymplocos; Antilles) groups as sister to the clade comprising Symplocastrum and Neosymplocos. The data support the independent evolution of deciduousness among section Palura and S. tinctoria. The early initial divergence of sections Cordyloblaste and Palura from the main group warrants their recognition at taxonomic levels higher than those at which they are currently placed. An inferred eastern Asian origin for Symplocos with subsequent dispersal to the Americas is consistent with patterns from other phylogenetic studies of eastern Asian-American disjunct plant groups but contrary to a North American origin inferred from the earliest fossil occurrences of the genus.