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PREMISE Pollinators are thought to exert selective pressures on plants, mediating the evolution of convergent floral shape often recognized as pollination syndromes. However, little is known about the accuracy of using petal shape for inferring convergence in pollination mode without a priori pollination information. Here we studied the genus Erythrina L. as a test case to assess whether ornithophyllous pollination modes (hummingbirds, passerines, sunbirds, or mixed pollination) can be inferred based on the evolutionary analysis of petal shape. METHODS We characterized the two-dimensional dissected shape of standard, keel, and wing petals from 106 Erythrina species using geometric morphometrics and reconstructed a phylogenetic tree of 83 Erythrina species based on plastid trnL-F and nuclear ribosomal ITS sequences. We then used two phylogenetic comparative methods based on Ornstein-Uhlenbeck models, SURFACE and l1OU, to infer distinct morphological groups using petal shape and identify instances of convergent evolution. The effectiveness of these methods was evaluated by comparing the groups inferred to known pollinators. RESULTS We found significant petal shape differences between hummingbird- and passerine-pollinated Erythrina species. Our analyses also revealed that petal combinations generally provided better inferences of pollinator types than individual petals and that the method and optimization criterion can affect the results. CONCLUSIONS We show that model-based approaches using petal shape can detect convergent evolution of floral shape and relatively accurately infer pollination modes in Erythrina. The inference power of the keel petals argues for a deeper investigation of their role in the pollination biology of Erythrina and other bird-pollinated legumes.

The extent to which phylogenetic biome conservatism vs biome shifting determines global patterns of biodiversity remains poorly understood. To address this question, we investigated the biogeography and trajectories of biome and growth form evolution across the Caesalpinia Group (Leguminosae), a clade of 225 species of trees, shrubs and lianas distributed across the Rainforest, Succulent, Temperate and Savanna Biomes. We focused especially on the little-known Succulent Biome, an assemblage of succulent-rich, grass-poor, seasonally dry tropical vegetation distributed disjunctly across the Neotropics, Africa, Arabia and Madagascar. We reconstructed a time-calibrated phylogeny, assembled species occurrence data and assigned species to areas, biomes and growth forms. These data are used to estimate the frequency of transcontinental disjunctions, biome shifts and evolutionary transitions between growth forms and test for phylogenetic biome conservatism and correlated evolution of growth forms and biome shifts. We uncovered a pattern of strong phylogenetic Succulent Biome conservatism. We showed that transcontinental disjunctions confined within the Succulent Biome are frequent and that biome shifts to the Savanna, Rainforest and Temperate Biomes are infrequent and closely associated with shifts in plant growth forms. Our results suggest that the Succulent Biome comprises an ecologically constrained evolutionary arena spanning large geographical disjunctions across the tropics.

The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK gene sequences, and including near-complete sampling of genera (698 of the currently recognised 765 genera) and ca. 20% (3696) of known species. The matK gene region has been the most widely sequenced across the legumes, and in most legume lineages, this gene region is sufficiently variable to yield well-supported clades. This analysis resolves the same major clades as in other phylogenies of whole plastid and nuclear gene sets (with much sparser taxon sampling). Our analysis improves upon previous studies that have used large phylogenies of the Leguminosae for addressing evolutionary questions, because it maximises generic sampling and provides a phylogenetic tree that is based on a fully curated set of sequences that are vouchered and taxonomically validated. The phylogenetic trees obtained and the underlying data are available to browse and download, facilitating subsequent analyses that require evolutionary trees. Here we propose a new community-endorsed classification of the family that reflects the phylogenetic structure that is consistently resolved and recognises six subfamilies in Leguminosae: a recircumscribed Caesalpinioideae DC., Cercidoideae Legume Phylogeny Working Group (stat. nov.), Detarioideae Burmeist., Dialioideae Legume Phylogeny Working Group (stat. nov.), Duparquetioideae Legume Phylogeny Working Group (stat. nov.), and Papilionoideae DC. The traditionally recognised subfamily Mimosoideae is a distinct clade nested within the recircum-scribed Caesalpinioideae and is referred to informally as the mimosoid clade pending a forthcoming formal tribal and/or clade-based classification of the new Caesalpinioideae. We provide a key for subfamily identification, descriptions with diagnostic charactertistics for the subfamilies, figures illustrating their floral and fruit diversity, and lists of genera by subfamily. This new classification of Leguminosae represents a consensus view of the international legume systematics community; it invokes both compromise and practicality of use.

The generic affiliation of the Andean species Caesalpinia trichocarpa, C. mimosifolia, and their close relatives has remained uncertain in all recent studies of Caesalpinia s.l. (Leguminosae, subfamily Caesalpinioideae). A new densely sampled phylogeny based on four DNA sequence regions (rps16, trn D-trnT, ycf6- psbM, ITS) strongly supports the monophyly of an Andean clade. We propose that despite the lack of obvious diagnostic morphological synapomorphies, this Andean group should be considered as a distinct genus, here described as the new genus Arquita. Phylogenetic analyses also suggest a problem with species delimitation in this group. Within C. trichocarpa, accessions from disjunct geographic areas in Argentina, Bolivia and Peru each form a robustly supported, unresolved clade that includes C. mimosifolia. The morphological and genetic cohesiveness of the C. trichocarpa complex is investigated using morphometric phenetic analyses of qualitative and quantitative flower and leaf traits, and reconstruction of a densely sampled phylogeny using three plastid and one nuclear ribosomal DNA sequence loci. Our results suggest that the most geographically isolated of these clades, narrowly endemic to two inter-Andean valleys in central-north Peru and separated by ∼1350 km, and extensive high Andean cordilleras above 4000 m, from the nearest populations in Bolivia, represents a genetically highly distinct and morphologically cryptic lineage here described as a new species (Arquita grandiflora). A full taxonomic account of the new genus Arquita and its component species is provided, with a distribution map and a key to the species.

Rosa rugosa , a vigorous ornamental shrub introduced from Asia in the 19th century, is now naturalized in coastal northeastern North America, where it occasionally grows in sympatry with the native R. blanda . To document hybridization between these species, evaluate its extent across the area of sympatry, and examine the use of morphology as a fi eld monitoring tool, we sampled 179 individuals of parental species and putative hybrids in 13 pure and 11 mixed populations. We developed allele-specifi c primers to assay single nucleotide polymorphisms (SNPs) markers from one chloroplast region and four low-copy nuclear introns. Our results revealed frequent bidirectional hybridization and infrequent introgression in sympatric populations of these species. The recurrent presence of F 1 hybrids in mixed populations indicated the weakness of early-acting reproductive barriers. Morphological data were concordant with molecular data and provided additional evidence for the presence of a few backcrosses. Morphological analyses yielded diagnostic characters for identifying hybrids and monitoring the hybrid zone. Such hybridization could ultimately lead to the genetic assimilation of R . blanda in mixed populations and to the formation of invasive hybrid genotypes, a phenomenon that is of economic and ecological concern because of the increasing number of exotic species worldwide.

This study investigates the impact of hybridization and polyploidy in the evolution of eastern North American roses. We explore these processes in the Rosa carolina complex (section Cinnamomeae), which consists of five diploid and three tetraploid species. To clarify the status and origins of polyploids, a haplotype network (statistical parsimony) of the glyceraldehyde 3-phosphate dehydrogenase (GAPDH) nuclear gene was estimated for polyploids of the complex and for diploids of section Cinnamomeae in North America. A genealogical approach helped to decipher the evolutionary history of polyploids from noise created by hybridization, incomplete lineage sorting, and allelic segregation. At the diploid level, species west of the Rocky Mountains are distinct from eastern species. In the east, two groups of diploids were found: one consists of R. blanda and R. woodsii and the other of R. foliolosa, R. nitida, and R. palustris. Only eastern diploids are involved in the origins of the polyploids. Rosa arkansana is derived from the blanda-woodsii group, R. virginiana originated from the foliolosa-nitida-palustris group, and R. carolina is derived from a hybrid between the two diploid groups. The distinct origins of these polyploid taxa support the hypothesis that the three polyploids are separate species.

The chloroplast trnL-F region was used as an independent data set for phylogenetic analysis of 118 aroid taxa. We investigated the intergeneric relationships of subfamily Monsteroideae (Araceae) and used this as a basis for an interspecific phylogenetic study of Rhaphidophora Hassk., the largest genus of the Monsteroideae. Results of the molecular tree were useful for inferring subfamilial and tribal circumscription and evolution in Araceae. Our results show that family Araceae consists of five clades that correspond to the subfamilies traditionally recognized. Starting from the most basal clade, these correspond to subfamilies Gymnostachydoideae and Orontioideae (proto aroids), with Lemna sp. (Lemnaceae) embedded in the Araceae and sister to the true aroids consisting of Pothoideae sister to Monsteroideae, immediately sister to Lasioideae and Aroideae. There is less agreement with existing tribal classifications. Complex relationships exist between members of the Monsteroideae. Our results show that Rhaphidophora and Epipremnum are paraphyletic with species of Rhaphidophora sampled, forming three informal groups with other genera of the Monstereae. Phylogenetic results may be used to suggest taxonomic changes to the current systematics of the monsteroids.

It has been asserted that in order to avoid circularity in phylogenetic tests of ecological hypotheses, one must exclude from the cladistic analysis any characters that might be correlated with that hypothesis. The argument assumes that selective correlation leads to lack of independence among characters and may thus bias the analysis. This argument conflates the idea of independence between the ecological hypothesis and the phylogeny with independence among characters used to construct the tree. We argue that adaptation or selection does not necessarily result in the non-independence of characters, and that characters for a cladistic analysis should be evaluated as homology statements rather than functional ones. As with any partitioning of data, character exclusion may lead to weaker phylogenetic hypotheses, and the practice of mapping characters onto a tree, rather than including them in the analysis, should be avoided. Examples from pollination biology are used to illustrate some of the theoretical and practical problems inherent in character exclusion.

Erythrina L. (Leguminosae: Phaseoleae) is a pantropical genus of over 100 species, all of which are either hummingbird or passerine pollinated. Phylogenetic hypotheses based on morphological and chloroplast DNA restriction site characters suggest that shifts from passerine to hummingbird pollination have occurred a minimum of four times in the genus. In hummingbird-pollinated species the inflorescences are held upright, the flowers are arranged radially along the axis, and the narrow standard petal is conduplicately folded to form a pseudotube. In most of the passerine-pollinated species, the inflorescences are held horizontally, the flowers are secund, and the standard petal is open so that the nectar and androecium are easily visible and accessible. Nectar amino acid concentrations and sucrose to hexose ratios are closely associated with pollination mode. Despite the general resemblance in flower and inflorescence morphology among species with the same pollination type, homology assessment reveals that petal morphology and size, and calyx and pollen morphology differ. Morphological characters, even if comprising modifications associated with adaptive pollination systems, therefore provide useful phylogenetic information.

Apios americana Medicus (Leguminosae; Phaseoleae), a tuberous, herbaceous vine, consists of diploid and triploid clones which are morphologically nearly identical. Although A. americana is widespread in eastern North America and flowers profusely throughout its range, fruit set is low. In part, this is because populations north of Connecticut consist almost exclusively of sterile triploid individuals. Although triploid populations occur south of this region, diploids predominate. Fruit set is also low in diploids. We attribute this to partial self-incompatibility and to a low rate of floral visits by leaf-cutter bees (Megachile spp.), the only legitimate pollinator. Visitation is easily measured because flowers have a nonrepetitive explosive tripping mechanism which must be triggered by insects. In two Connecticut populations only 10% of 299 flowers were tripped, and on average only about one-halfof these set fruit. We conclude that A. americana maintains populations by vegetative propagation, but expands its range and retains variability through sexual reproduction of diploids.