A revision of the moss genus Erythrophyllopsis (Pottiaceae) recognized two species: E. andina and E. zanderi sp. nov. Barbula fuscula, Erythrophyllopsis boliviana, Glyphomitrium ferrugineum, Tortula semivaginata, and Trichostomum juniperinum are synonymized with E. andina. Erythrophyllopsis zanderi includes Zander's (1993) concept of Erythrophyllastrum andinum. Lectotypes for Barbula fuscula, Erythrophyllopsis boliviana, Trichostomum juniperinum, and Tortula semivaginata are here proposed. A new key, detailed descriptions, illustrations, and a distribution map are provided. Maximum parsimony and Bayesian analyses of cpDNA sequences (trnL-trnF and trnG regions) suggest that Erythrophyllopsis is nested within Bryoerythrophyllum. A denser sampling and the use of other phylogenetic markers are required to better understand the generic and suprageneric relationships of Erythrophyllopsis.

A new fern species, Pteris xiaoyingae (sect. Pteris, Pteridaceae), is described from a karst cave in southern Guizhou, China. The new species is a small-sized limestone rock-dwelling plant. It has trifoliate leaves with two lateral pinnae each mostly divided at the base into two almost free basiscopic lobes. Morphologically it is similar to P. gallinopes and P. deltodon that also occur on shady limestone walls often inside or near karst caves. Pteris xiaoyingae differs from P. gallinopes in its middle pinna being ovate-lanceolate with blunt apex and lateral pinnae and their lobes oblong-obovate with round or obtuse apex. Pteris xiaoyingae also has wider pinnae (more than 7 mm) than P. gallinopes (2–4 mm). Pteris xiaoyingae is distinguishable from P. deltodon by being smaller in stature (only 4–7 cm high in P. xiaoyingae vs. 10–40 cm high in P. deltodon) and by having lamina more or less pedate in outline (for the majority of leaves with lobed lateral pinnae) and pinnae and lobes with round or blunt apices (vs. with short or long acuminate apices in P. deltodon). Palynologically, P. xiaoyingae has conic-deltate spores with verrucate sculpture while both P. gallinopes and P. deltodon have deltate spores with vermiculate-rugulate sculpture. Morphological distinctiveness of this new fern and its relationships with allied species are discussed and a key to these and other morphologically similar species is given.

Recent phylogenetic analyses of grammitid ferns (Polypodiaceae) have demonstrated that some genera recognized within this clade are not monophyletic. We focus on resolving the circumscriptions of some large Neotropical genera by focusing on a clade, identified in previous analyses, that includes the monophyletic genus Melpomene, plus portions of two polyphyletic genera, Lellingeria and Terpsichore. Phylogenetic analyses using chloroplast markers atpB, rbcL, and trnL-F, along with 109 morphological characters, resolve this ingroup as monophyletic and sister to a clade that includes Ceradenia, Enterosora, and Zygophlebia. We use morphological characters included in the analysis to circumscribe clades. Melpomene is clearly monophyletic, but forms a polytomy with Lellingeria s. l. in a strict consensus of MP trees. Ingroup species of Terpsichore form three well supported monophyletic groups that together are paraphyletic with respect to Melpomene plus Lellingeria. Two clades corresponding to species groups of Terpsichore will be recognized as new genera.

Terpsichore, a Neotropical genus of about 70 species, was shown in previous molecular phylogenetic studies to be polyphyletic. The present study assesses this conclusion using morphology and also examines the phylogenetic relationships of five informal species groups originally described within Terpsichore. A morphological matrix was constructed for 109 qualitative characters and 129 terminals. Maximum parsimony was used to analyze the complete data set and three different partitions of that data set which excluded characters of either leaf shape or indumentum position or both. Terpsichore is polyphyletic in all analyses, and the characters originally used to define the genus are either plesiomorphic or homoplastic. Whereas the analysis of the complete matrix recovered a clade similar to the original circumscription, analyses of the partitioned data sets yielded results similar to those of molecular phylogenetic studies. In all analyses, two of the five infrageneric groups of Terpsichore are resolved as monophyletic, whereas the others are either paraphyletic or polyphyletic. In all analyses, species from the Terpsichore taxifolia Group are recovered as diphyletic, a result corresponding to that found in the previous molecular phylogenetic studies. By scoring morphological characters for this study, it was found that two vascular bundles are present in many grammitid petioles and are not unique to Luisma, mesophyll composed of long-armed stellate parenchyma evolves at least three times and is not unique to Enterosora, and perforated dictyosteles are common in grammitids and not restricted to Zygophlebia.

A review of Ephedra species of the Western Himalayan region led to the recognition of two new species, Ephedra kardangensis Sharma & Uniyal sp. nov. and E. khurikensis Sharma & Uniyal sp. nov. The most distinctive characters observed are shape and size of male strobilus, number of bracts, length of synangiophore with synangia, shape and size of female strobilus, number of bracts, size and color of seeds, and several features of the micropyle for morphological and statistical analysis (Parametric MANOVA and Principal Components Analysis). These species are most similar to E. intermedia.

Pollination by water (hydrophily) is a highly specialized mechanism that occurs rarely among aquatic angiosperms, which mainly retain the reproductive systems reminiscent of their terrestrial ancestors. Hydrophily is abiotic and typically associated with unisexual flowers, factors that predictably would promote xenogamy and outcrossing. Yet, there have been few reports of hybridization involving waterpollinated plants (hydrophiles), with no firm evidence of natural interspecific hybridization. The genus Najas comprises about 40 species of submersed aquatic plants, all characterized by subsurface hydrophily. Hybridization in this genus has been suspected, but verified previously only among infraspecific taxa. In this study we document the first instance of interspecific hybridization in Najas using genetic evidence from three populations that were identifiable as N. guadalupensis but yielded polymorphic DNA sequence profiles. To facilitate our analysis we first conducted a phylogenetic survey of New World Najas taxa using nuclear and chloroplast markers. Alleles cloned from a biparentally-inherited locus (ITS) in these aberrant populations associated with two distinct but phylogenetically sister species (N. guadalupensis subsp. olivacea and N. flexilis) thus confirming their hybrid origin. In all cases the chloroplast markers associated with N. guadalupensis subsp. olivacea, implicating it as the maternal parent. The hybrid Najas plants occur at the edge of the sympatric range of the parental species. They possess no readily distinctive morphological features and require genetic analysis for confident detection. One population grows aggressively, raising concerns that at least some hybrid Najas plants represent a potential conservation threat. The possible hybrid ancestry of the endemic N. guadalupensis subsp. muenscheri also was assessed, but could not be confirmed or refuted by the data evaluated.

Hechtia chichinautzensis and H. colossa, morphologically related to H. podantha and to H. tehuacana, respectively, are described and illustrated. Hechtia chichinautzensis is endemic to the Mexican state of Morelos and is distinguished from H. podantha by its flexuous and lepidote rachis, ovate-triangular primary bracts, sparsely lepidote pedicels, and brownish to blackish anthers; anatomically its leaves are characterized by their square adaxial epidermal cells with undulate walls, one or two layers of adaxial hypodermic cells, discontinuous chlorenchyma, oblong palisade cells, and the presence of raphides. Hechtia colossa, from the states of Oaxaca and Puebla, differs from H. tehuacana by its larger habit, the lepidote peduncle and primary bracts, the length of the inflorescense branches and the larger foliar sheaths, blades, and floral bracts; anatomically its leaves are characterized by their irregular adaxial epidermal cells and by the number of vascular bundles.

Classification within Eragrostis, a large genus of ca. 400 species in Poaceae subfamily Chloridoideae, has long been problematic. While spikelet disarticulation characters have been the most common basis of infrageneric classifications, they have not been demonstrated to describe monophyletic groups. An alternative classification system linked to leaf blade anatomy characters is explored using phylogenies based on DNA sequence data from three loci (nuclear GBSSI, plastid rps16, and the plastid trnL-F region) in this study. Eragrostis possesses a great deal of variation in leaf blade anatomy, ranging from NAD-ME-like in structure (subgenus Eragrostis) to PCK-like (subgenus Caesiae), with some intermediates between these extremes. This study reveals that a great deal of variation exists in those species traditionally classified as “intermediate” in leaf blade anatomy. It also indicates that the PCK-like species form a monophyletic group in phylogenetic analyses based on plastid data, GBSSI sequences, and simultaneous analysis of all loci. NAD-ME-like and intermediate species do not form monophyletic groups, but phylogenies based on plastid sequences indicate some evolutionary conservation of the various types of intermediate anatomy. This study also evaluates the leaf anatomy type in allopolyploids. A high proportion of these allopolyploids have intermediate leaf anatomy, potentially reflecting their derivation from progenitors with distinct anatomical types.

Moraceae is a large (∼1,050 species) primarily tropical family with several economically and ecologically important species. While its monophyly has been well supported in recent studies, relationships within the family at the tribal level and below remain unresolved. Delimitation of the tribe Artocarpeae has been particularly difficult. Classifications based on morphology differ from those based on phylogenetic studies, and all treatments include highly heterogeneous assemblages of genera that seem to represent a cross section of the family. We evaluated chloroplast and nuclear DNA sequence data for 60 Moraceae taxa representing all genera that have been included in past treatments of Artocarpeae and also included species from several other Moraceae tribes and closely related families as outgroups. The data were analyzed using maximum parsimony and maximum likelihood methods and indicate that none of the past treatments of Artocarpeae represent a monophyletic lineage. We present the most complete phylogenetic hypothesis for Artocarpeae and the genus Artocarpus to date. Inflorescence evolution and pollination are briefly discussed and the phylogenetic reconstructions are used to inform a revised treatment of Artocarpeae and the Artocarpus subgenera. The following new combinations are proposed: the genus Prainea is reduced to Artocarpus subgenus Prainea, and the series Cauliflori is raised to Artocarpus subgenus Cauliflori.

With nearly 2,000 species, all restricted to the Neotropics, Miconieae represent the largest radiation within Melastomataceae. Generic limits and relationships within the tribe have long been contested. Our molecular phylogenetic analyses of the tribe, based on sequences of nrITS and ndhF, resulted in consistent support for a clade comprising Bellucia, Loreya, Henriettea, and Henriettella, which is neither nested within nor sister to the Miconieae. Furthermore, and based upon a series of presumably synapomorphic anatomical and morphological characters, e.g. megastyloid crystals, absence of both medullary and cortical vascular bundles, plinerved leaves, axillary or cauliflorous inflorescences, we consider three additional genera, Kirkbridea, Llewelynia, and Myriaspora, to belong to this clade, a newly recognized tribe, Henrietteeae. These seven genera had formerly been assigned to the Miconieae based on their having exappendiculate anthers (except Kirkbridea) and berry fruits. Henriettella and Llewelynia are relegated to synonymy under Henrietten (25 new combinations are made under that genus: Henriettea boliviensis, H. bracteosa, H. caudata, H. duckeana, H. fissanthera,H. glabra,H. cogniauxiana, H. goudotiana,H. heteroneura, H. hondurensis, H. ininiensis, H. lawrancei, H. lundellii, H. maguirei, H. manarae, H. odorata, H. ovata, H. prancei, H. rimosa, H. steyermarkii, H. tachirensis, H. tobagensis, H. tovarensis, H. trachyphylla, H. williamsii), and Loreya and Myriaspora are synonymized under Bellucia (with nine new combinations under that genus: Bellucia egensis, B. gracilis, B. klugii, B. nigricans, B. ovata, B. riparia, B. strigosa, B. subandina, B. wurdackiana). A description of Henrietteeae along with a key to and brief descriptions of its constituent genera are provided, in addition to necessary new combinations.

Section Macrantha is a monophyletic group with 18–22 species, all of which are distributed in Asia except for Acer pensylvanicum in eastern North America. Nuclear sequences from nrDNA ITS, LEAFY, and GBSSI (two putative copies), along with morphological characters, recognize two clades (series Crataegifolia and Micrantha). However, chloroplast phylogenies based on ndhF, trnL-F, and psbA-trnH are significantly incongruent with the nuclear trees, showing two major clades with species from the Asian continent excluding the northeast (AS - N) in one clade and those from Asian Pacific Islands plus northeast Asia in the other (API N). Ancient hybridization and chloroplast capture may have occurred in the two series in both geographic areas. The two series diverged at 46.3 mya, which is before the formation of the Japanese Sea in the early Miocene (32–18 mya). The Japanese Sea may be the barrier for the seed exchange between the two geographic areas, and was responsible for the formation of the disjunct distribution of section Macrantha between the AS N and API N areas, and between eastern Asia and North America.

Photosynthetic pathway innovations have had a large impact on patterns of diversification of angiosperm lineages and the biogeographic distribution of ecological assemblages. C₄ photosynthesis has been one of the most studied processes in plants with respect to function, structure, occurrence, and response to climatic conditions. One of the most promising areas of research of C₄ photosynthesis is in the Cleomaceae. Here we explore the phylogenetic origins of the C₄ pathway in the Cleomaceae using maximum parsimony, maximum likelihood, and Bayesian inference analyses of nrDNA ITS sequences. As has been found previously, commonly recognized genera including Buhsia, Cleomella, Dactylaena, Gynandropsis, Isomeris, Oxystylis, Podandrogyne, Polanisia, and Wizlizenia are derived from within a paraphyletic Cleome. The phylogenetic results presented here indicate that there are likely at least five separate origins of carbon concentrating mechanisms in the Cleomaceae, including at least three separate origins of C₄ species. Analyses of historical biogeography suggest Cleomaceae originated in central Asia.

The Hawaiian Islands are home to a quarter of the named diversity of sandalwoods in the genus Santalum. There has been considerable confusion on how to best interpret the variation patterns of the red-flowered Hawaiian sandalwoods, S. freycinetianum and S. haleakalae, and particularly of the endangered S. freycinetianum var. lanaiense in an appropriate taxonomy. In this study, nrDNA (ITS, ETS) and cpDNA (3′ trnK intron) sequence, microsatellite, and morphological data are integrated to appropriately revise the taxonomy of this group by better understanding the genetic and morphological diversity within and between populations. Results reveal that populations of S. freycinetianum from O‘ahu are genetically distinct from populations on Moloka‘i, Lana‘i, and Maui. Santalum freycinatianum is now considered only to occur on O‘ahu. The East Maui endemic S. haleakalae intergrades morphologically and is not genetically distinct from populations of S. freycinetianum var. lanaiense based on the sequence and microsatellite data gathered thus far. We combine them here into a single species, S. haleakalae, with two varieties (var. haleakalae and var. lanaiense, comb nov.). Lastly, examination of populations of S. freycinetianum var. pyrularium suggest it is best treated at specific rank as S. pyrularium. Some populations that are sympatric with S. pyrularium and S. ellipticum in the coastal cliffs and valleys of northern Kaua‘i, are morphologically similar to S. pyrularium but are more closely related to the white-flowered S. ellipticum clade according to both nrDNA and cpDNA data. However, at least three synapomorphic sites in the nrDNA data indicate that its origin may have been the result of an ancient hybridization event with the red-flowered clade. The morphological characteristics of this inferred ancient hybrid lineage appear to correspond with the species S. involutum described by H. St. John.

A phylogenetic investigation of American members of the tribe Atripliceae (Chenopodiaceae) was conducted to examine evolutionary patterns and ecophysiological change in the North American lineages. Molecular phylogenetic analyses including representatives of all American genera of Atripliceae and data from 18S–26S nuclear ribosomal (ITS, ETS) and cpDNA (3′trnK intron) sequences suggest that neither Atriplex, as traditionally recognized, nor the North American members of Atriplex constitute a monophyletic group. Congruence between well-supported nuclear and cpDNA clades and results of expanded leaf anatomical and isotopic analyses indicate that American taxa belong to two distantly related lineages with different photosynthetic pathways; only one shift from C₃ to C₄ photosynthesis is required to explain photosynthetic pathway evolution in sampled lineages of Atripliceae. Conservatism in photosynthetic pathway evolution in the tribe is exemplified by the endemic, C₃ North American Atriplex taxa, which are nested among other genera (e.g. Grayia, Holmbergia, and Zuckia) within a robust C₃ clade. One North American C₃ genus, Suckleya, often included within Atripliceae, should be excluded from the tribe. A previous hypothesis of variation in photosynthetic pathway within Atriplex phyllostegia sensu Hall and Clements is rejected. A revised taxonomy employing only monophyletic groups reflects this new understanding of relationships within the tribe. Extriplex includes E. californica (comb, nov.) and E. joaquinana (comb, nov.) and has not been recognized as a natural group by previous authors. Stutzia is proposed to replace the name Endolepis Torr. (1860), a later homonym of Endolepis Schleid. (1846), and includes S. covillei (comb, nov.) and S. dioica (comb. nov). Grayia (G. plummeri, comb. nov, and G. arizonica, comb. nov.) is expanded to include Zuckia. Recognition of Proatriplex as distinct from Atriplex is supported. Holmbergia is retained in Atripliceae. The improved phylogenetic understanding of Atripliceae should allow for more meaningful comparative studies of physiological and other functional adaptations of Atriplex, especially in North America.

A cladistic analysis including for the first time all taxa of Anulocaulis, five species and four varieties, is presented. Sixteen potentially related taxa classified within Boerhavia, Commicarpits, Cyphomeris, Nyctaginia, and Okenia were used as outgroup to test the monophyly and phylogenetic position of Anulocaulis; Mirabilis glabrifolia was used to root the tree. Forty-one characters were included in the matrix, which was analyzed using the parsimony criterion. Our results are in partial agreement with previous studies using molecular data. Here, the monophyly of Anulocaulis is corroborated, but the sister group remains ambiguous depending on whether the multistate characters are treated as additive or nonadditive. In the first case, Anulocaulis is sister to Nyctaginia-Okenia, while in the second it is sister to Boerhavia anisophylla. The morphological characters which have been used in the traditional taxonomy and classification of the genus are analyzed and discussed under a phylogenetic context. Species recognition, infraspecific taxa and infrageneric classification for Anulocaulis are discussed. A taxonomic synopsis for this xerophytic genus including keys and descriptions based on our cladistic results is presented.

A phylogeny of Cantua was estimated using parsimony analyses of DNA sequences from the nuclear ribosomal ITS region and the cpDNA spacers trnT-tmL, tmL-trnF, and partial ndhF coding region. One of the two species formerly included in Huthia was included in these analyses; the other is known only from the type. Data from the phylogenetic analyses strongly supported the paraphyly of Cantua to this taxon, Cantua volcanica (= Huthia coerulea). This result supports the earlier inclusion of the two species formerly placed in Huthia within Cantua. Cantua volcanica was most closely related to C. quercifolia and these two species together were the sister group to the rest of Cantua. The remaining taxa, C. flexuosa, C. cuzcoensis, C. bicolor, C. candelilla, C. buxifolia, and C. pyrifolia were well supported as a monophyletic group. For most species, multiple individuals were sampled. In each case, except for the individuals of C. buxifolia, the multiple individuals formed a clade. Precise relationships between C. buxifolia and other species remain unresolved. Cantua mediamnis (= Huthia longiflora) and C. dendritica were not included in the phylogenetic analyses, however based on morphological characters the former is clearly allied with C. volcanica and the latter with C. flexuosa.

The Leptostemonum clade of Solanum contains approximately 350–450 species, including the cultivated eggplant, S. melongena. This clade is characterized by the presence of prickles and apically attenuate anthers. Solanum section Androceras, the focus of this study, is a group of ca. 12 species belonging to the Leptostemonum clade. This section is unusual in the genus because of its mostly north temperate distribution and distinctive zygomorphic, heterantherous, and enantiostylous flowers. We infer phylogenetic relationships among 43 Solanum taxa, including 11 species and all varieties of sect. Androceras, using DNA sequence data from two nuclear regions (ITS and the granule-bound starch synthase gene [GBSSI or waxy]) and the chloroplast region trnT-F. The combined phylogenetic tree supports sect. Androceras as a monophyletic group sister to Solanum sect. Crinitum. Only one of the three series proposed by previous taxonomists, ser. Pacificum, is supported as monophyletic. Solanum tenuipes from the northern Chihuahua Desert is sister to the remaining species in sect. Androceras. Species-level relationships were also examined and it was found that two species, S. heterodoxum and S. citrullifolium, are not monophyletic. The ancestral flower color in sect. Androceras appears to be violet, with white and yellow flowers restricted to more derived clades. Characters formerly used to diagnose ser. Androceras, such as exclusively branched hairs and lack of complex foliar flavonoids, appear to have evolved more than once in the section.

Herbarium specimens are an underutilized information resource on historic vegetation patterns. We record habitat loss and present evidence that visits to specimen collecting localities can document environmental change. The taxonomic study of Solanum in Africa continues to be plagued by confusion and ignorance. Despite the almost universal occurrence of Solanum in the flora of Kenya, there continues to be a significant gap in floristic knowledge of this important genus. As a result of studies of herbarium collections and visits to collection localities across Kenya, we have identified three new species and compiled a list of native species of spiny Solanum. We describe Solanum polhillii from the Acacia-Commiphora savanna, Solanum phoxocarpum from the Kenyan highlands, and Solanum malindiense from the coastal vegetation.

Ptilostemon is a fine example of the representatives of the eastern groups of the Cardueae that have diversified in the western Mediterranean. Relationships to Cynara, which exhibits a similar distribution, and Lamyropsis, which is morphologically closer according to previous studies, are investigated using Bayesian analysis of DNA sequences of the plastid intergenic spacer ycf3-trnS and two nuclear regions, the ETS and ITS spacers. The sectional classification and biogeography of Ptilostemon are also revised in the light of the molecular phylogeny Our results suggest that Cynara is the most plausible sister genus to Ptilostemon. Some paralogous copies of the ETS region found among species of the three genera by cloning are interpreted as incomplete lineage sorting of ancestral polymorphisms. The current sectional classification of Ptilostemon shows excessive fragmentation, which does not agree with our phylogeny, and therefore a more synthetic classification is proposed. The present distribution of Ptilostemon indicates that there were two colonization events in the western Mediterranean region, paralleling a similar pattern of successive waves already suggested for Cynara.