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Polyploidy presents a challenge to those wishing to delimit the species within a group and reconstruct the phylogenetic relationships among these taxa. A clear understanding of the tree-like relationships among the diploid species can provide a framework upon which to reconstruct the reticulate events that gave rise to the polyploid lineages. In this study we apply this “diploids-first” strategy to the fern genus Astrolepis (Pteridaceae). Diploids are identified using the number of spores per sporangium and spore size. Analyses of plastid and low-copy nuclear sequence data provide well-supported estimates of phylogenetic relationships, including strong evidence for two morphologically distinctive diploid lineages not recognized in recent treatments. One of these corresponds to the type of Notholaena deltoidea, a species that has not been recognized in any modern treatment of Astrolepis. This species is resurrected here as the new combination Astrolepis deltoidea. The second novel lineage is that of a diploid initially hypothesized to exist by molecular and morphological characteristics of several established Astrolepis allopolyploids. This previously missing diploid species is described here as Astrolepis obscura.
The nomenclature related to Tectaria subpedata (Harr.) Ching (Tectariaceae) is clarified. The misidentification of T. subpedata is documented by a comprehensive study of herbarium specimens, wild and cultivated plants, spore morphology, gametophyte development, and chromosome count. The type of T. subpedata is confirmed to be a young frond of T. polymorpha (Wall. ex Hook.) Copel. A synonym of T. subpedata, namely Nephrodium morsei Baker, represents a separate species and is here recognized as Tectaria morsei (Baker) P. J. Edwards ex S. Y. Dong. Based on the sporophyte morphology, gametophyte development and distribution, the affinity and position of T. morsei is briefly discussed.
Typhonium was recognized as a monophyletic genus in the tribe Areae of the subfamily Aroideae (Araceae) until a recent molecular phylogenetic study indicated that the genus was paraphyletic respective to other Areae genera. However, that phylogenetic study did not discuss the details of infrageneric relationships due to the limited numbers of Typhonium samples. To elucidate the phylogenetic relationships in Typhonium sensu lato, we conducted phylogenetic analyses based on the combined DNA sequences of six chloroplast regions (3′trnL—trnF, rp120—5′rps12, psbB—psbH, trnG intron, rpoC2—rps2, and trnK 3′intron) from 18 representative Typhonium species and additional samples from related genera. The resultant tree strongly suggests that Typhonium is not a monophyletic group and that it comprises at least two separate lineages, with other Areae genera nested within, and that Typhonium sensu lato may be subdivided into several monophyletic groups. These groups are distinguishable based on the stem-type of shoot organization as well as other morphological characters, which mostly correspond to traditionally recognized taxa. Based on molecular phylogeny and morphology, we proposed a revision of the Areae, wherein Typhonium sensu lato is divided into four genera: Typhonium sensu stricto, Sauromatum, and three new genera, Diversiarum, Hirsutiarum, and Pedatyphonium, which are described here which results in the following combinations: Diversiarum diversifolium, Diversiarum alpinum, Pedatyphonium horsfieldii, Pedatyphonium larsenii, Pedatyphonium kunmingense, Pedatyphonium calcicolum, Pedatyphonium omeiense, Hirsutiarum hirsutum, Hirsutiarum brevipilosum, and Sauromatum giganteum.
During a recent survey of Atlantic central African orchids, we collected four orchid specimens in Rio Muni (Equatorial Guinea) that share the general morphology at Angraecum gabonense, the most frequent member of Angraecum section Pectinaria in Central Africa, but differ in leaf shape and flower size. Further inspection of specimens deposited at the Wageningen herbarium and cultivated in their greenhouse led to the discovery of additional specimens from the Monts Doudou area in Gabon. Comparison with other Angraecum specimens indicate that these collections represent a new species, which we describe here as Angraecum atlanticum, the fifth species of Angraecum section Pectinaria recorded in central Africa. The new species is restricted to submontane forests covering the mountain chain situated along the coasts of Gabon and Equatorial Guinea. The distinguishing features of the species include its leaves, which are more widely spaced than A. gabonese, and are 2.2 mm wide; its petals and sepals, which are slightly longer than the lip; its spur, which is somewhat inflated in the middle; and its larger ovary. Information on the ecology, phenology and distribution of Angraecum atlanticum is presented, along with a preliminary conservation assessment using the IUCN Red List Categories and Criteria.
Eleocharis (Cyperaceae) is a morphologically and physiologically diverse lineage of 250 species with a cosmopolitan distribution. We here explore phylogenetic relationships in this lineage using maximum parsimony and Bayesian inference analyses of nrDNA ITS and cpDNA tmC-ycf6 and ycf6-psbM sequence data with the goals of comparing our phylogenetic hypotheses to previous classifications, morphological variation, and photosynthetic pathway variation. Our results suggest that in Eleocharis C4 photosynthesis has been derived at least three times, with several cases of possible reversion to C3-like or intermediate pathways and several additional origins of C3–C4 intermediate photosynthetic pathways, as inferred by carbon isotope ratio measurements. Many classification units currently recognized in Eleocharis are not monophyletic, however, E. subgenus Limnochloa and E. subgenus Scirpidium are monophyletic. Other classification units largely corresponding to clades include E. subgenus Zinserlingia, E. subseries Chaetariae, and E. series Maculosae. Problems with species circumscription and morphological variation in several groups are discussed in light of the phylogeny, particularly in the context of species membership of seven focal clades found in the analyses.
Ormosia limae (Leguminosae, Papilionoideae, Sophoreae s. 1., Genistoid clade) is described and illustrated as a new species of Ormosia sect. Unicolores. It resembles the Amazonian species O. holerythra, but is morphologically and geographically distinct. The relationship of O. limae along with O. holerythra clearly lies with species of Ormosia sect. Unicolores because of the shared distinctive four to nine pairs of arcuate secondary leaflet veins, fruits that are dehiscent and glabrous at maturity, and seeds that are uniformly red in color and with a small, ca. three mm long, elliptic hilum. An identification key to sect. Unicolores is provided in order to enumerate all of the constituent species and to suggest relationships among them. Ormosia limae is known only from few collections in the Atlantic forest of southern Bahia, Brazil.
Four new species of Mimosa are described and illustrated: M. kalunga, M. pseudosetosa, M. viperina, and M. diminuta (all from section Habbasia, series Pachycarpae). All of them are apparently narrowly restricted endemics from the Chapada dos Veadeiros in Goiás, central Brazil. In addition, more complete data and an illustration are presented for M. pycnocoma Benth., previously incompletely known only from the type collection. The new species described here, as well as the new data on M. pycnocoma, provide interim increments to our knowledge of a group of around 55 Mimosa species, combining Barneby's (1991) series Setosae and Pachycarpae, that remains poorly and incompletely understood.
A new combination, Coursetia greenmanii (Leguminosae, Papilinioideae, tribe Robinieae), is proposed because analysis of nucleotide sequence data from the nuclear ribosomal internal transcribed spacer region resolves a clade that is phenotypically distinct and ecologically and geographically centered in lowland seasonally dry tropical vegetation of the Yucatan Peninsula region. The well supported monophyly of the ribosomal sequences sampled from Coursetia greenmanii suggests that this species has had a long history independent of its close relatives. Coalescence of intraspecific samples of ribosomal sequences from this geographically confined species is strongly suggestive of an evolutionary persistent small effective population size. This is the general case for species of Coursetia and related genera of the tribe Robinieae. Coalescence of nuclear ribosomal sequences sampled from geographically restricted species in this group further suggests that evolutionary persistence could be the result of high levels of dispersal limitation in seasonally dry Neotropical vegetation that is rich in cacti and other succulent species. This is the ecological setting to which Robinieae shows phylogenetic niche conservatism.
Daniellia (Leguminosae: Caesalpinioideae) is a genus of ten species of medium to large trees from tropical and subtropical Africa. In contrast to earlier accounts, D. mortehanii is treated here as a synonym of D. pymertii, and D. oblonga, which previously has been treated by some authors as a synonym of D. thurifera, is accepted. In addition, we describe as new Daniellia glandulosa. Quantitative characters such as receptacle length, petiole width, sepal length, and largest leaflet length have been used to aid species differentiation following the results of recent morphometric studies that revealed their taxonomic utility. Micromorphological features such as pollen surface and leaf glands were studied in all species, many for the first time, and the number, position, and shape of the glands present in the leaves are shown to be taxonomically useful in distinguishing species. A dichotomous key and a table of key characters is provided, as well as detailed descriptions and distribution maps for each species. All species are illustrated, D. oblonga and D. glandulosa for the first time. Three lectotypes and a neotype are designated.
A taxonomic revision of the recently described genus Wiborgiella is presented. Nine species are recognized within the genus, two of which are newly described, W. dahlgrenii and W. vlokii. The genus is endemic to the Cape region of South Africa and most species are highly localized and rare. Wiborgiella species can be distinguished from other genera in the tribe Crotalarieae by a combination of brown young branches (early formation of bark in the stems of the perennial species), laminar trifoliolate leaves, glabrous petals, a 4 6 anther arrangement, and oblong, wingless, inflated fruit. Anatomical studies revealed that all species of the genus have dorsiventral leaves (mesophyll differentiated into palisade parenchyma adaxially and spongy parenchyma abaxially) with mucilage cells in the epidermis which distinguishes Wiborgiella from other closely related genera, such as Calobota which has isobilateral leaves without mucilage cells. The fruits of Wiborgiella species are all thin-walled, have highly sclerified mesocarps and mucilage cells are present in some species. These fruit character states are also present in other genera of the Crotalarieae. The taxonomic treatment of the genus includes a key to the species, descriptions, illustrations, nomenclature, typification, and geographical distribution of each species.
Psiguria is a genus of lianas found throughout the Neotropics from sea level to 2,100 m, in dry or wet forests, in both light gaps and shade. Psiguria and its sister genera, Gurania and Helmontia, are unique among Cucurbitaceae having brightly colored flowers and butterfly and hummingbird pollinators. Historically, taxonomic revisions have described as many as 29 species, but this number has been inflated by misleading characteristics such as variable leaf and flower morphology. A thorough morphological investigation of 758 herbarium specimens and individuals from both greenhouse and natural settings, combined with a recent molecular phylogeny, supports the delineation of six species. In this revision, a review of over 35 yr of publications addressing ecological and natural history studies focusing on Psiguria is presented. A description and a distribution map of each species is provided along with two taxonomic keys: one that utilizes male flowers, and another that incorporates leaf characteristics and geographic distribution but uniquely identifies species according to DNA barcodes. With a list of characters that distinguish all pairs of geographically overlapping species, botanists and ecologists finally have the tools to confidently identify species of Psiguria.
The phylogeny of Celastraceae subfamily Salacioideae (ca. 255 species in the Old and New World tropics) and tribe Lophopetaleae (ca. 29 species in southern Asia and the Austral-Pacific) was inferred using morphological characters together with plastid (matK, trnL-F) and nuclear (ITS and 26S rDNA) genes. Brassiantha, a monotypic genus endemic to New Guinea, is inferred to be more closely related to the clade of Dicarpellum (New Caledonia) and Hypsophila (Queensland, Australia) than it is to Hippocrateoideae or Salacioideae. This unambiguously supported resolution indicates that a nectary disk positioned outside the stamens has been convergently derived in these two lineages. The clade of Kokoona and Lophopetalum is resolved as more closely related to Brexia and Elaeodendron than it is to Hippocrateoideae or Salacioideae. Sarawakodendron, a monotypic genus endemic to Borneo, is resolved as sister to Salacioideae. Salacioideae are inferred to have an Old World origin that was followed by a single successful radiation within Central and South America. We infer that capsular fruits are primitive within the clade of Hippocrateoideae Sarawakodendron Salacioideae, with berries a synapomorphy for Salacioideae. Based on the resolution of Sarawakodendron as sister to Salacioideae, we hypothesize that the filaments of Sarawakodendron arils are homologous to the spiral filaments in the mucilagenous pulp of Salacioideae.
The genus Calycolpus (Myrtaceae), a mainly northern South American group of 15 species, is revised based on a study of over 600 plant collections, many duplicated in more than one herbarium. The relationships of Calycolpus to other genera are discussed and a key distinguishing it from its relatives is provided. All species are described, mapped, and illustrated with line drawings and/or photographs. The relationships among the species are discussed and a key distinguishing them is provided. Lectotypes are chosen for Myrtus goetheana DC., Calycolpus chnoiophyllus Riley, Calycolpus cordatus L. Riley, Calycolpus kegelianus O. Berg, Calycolpus schomburgkianus var. speciosus O. Berg, Eugenia excisa Urb., Eugenia nipensis Urb., Psidiopsis moritziana O. Berg, and Psidium reversum Urb. Neotypes are chosen for Calycolpus kegelianus var. gracilis O. Berg, Calycolpus warszewiczianus O. Berg, and Myrtus revoluta Schauer. Five of these species, one third of the total, were described by the author since 1989.
Sarcocornia comprises ca. 20–24 perennial, halophytic herb and shrub species. The genus is distinct from other genera in the Salicornioideae in having flowers that are more or less equal in size, arranged in a row, and with seeds that have a membranous hairy testa and lack perisperm. Sarcocornia is distributed worldwide, mainly in regions characterized by warm-temperate and, to a lesser extent, subtropical climates. The representatives of this genus are found in habitats such as estuarine salt marshes, tidal mud flats, coastal cliffs, inland salt pans, and salt-laden alluvia of intermittent semi-desert and desert streams. Some South African taxa also occur in inland (semi-desert) quartz patches while some South American species occur on saline soils of dry, continental high plateaus in the Andes. The genus reaches its highest species diversity in the Greater Cape Floristic Region of South Africa, where twelve species and one subspecies are known. This revision of southern African (South Africa, Namibia, and Mozambique) Sarcocornia comprises an identification key, the descriptions of the genus, species, and subspecies. One new combination, S. dunensis, is proposed. The former varieties recognized within S. natalensis are here treated as subspecies, S. natalensis subsp. natalensis and subsp. affinis. Morphological characters of high taxonomic value in the genus are habit (growth-form), segment morphology, and testa micromorphology. Our revision also features information on the distribution and ecology of the studied taxa.
Boraginaceae tribe Trigonotideae comprises a heterogenous assemblage of taxa, many of which have been shown to belong to widely divergent lineages in Boraginaceae in the recent past, with some taxa now assigned to three of the four currently recognized tribes of the Boraginaceae s. s., namely the Cynoglosseae, Echiochileae, and Lithospermeae. The systematics of Moritzia and Thaumatocaryon, the only endemic South American genera of Boraginaceae, have been controversially discussed in the past, and their most recent placement was in Trigonotideae. The present study investigates the phylogenetic relationships of “Trigonotideae” based on micromorphology and molecular data (ITS including 5.8S rRNA, and the trnL-trnF spacer). Molecular data show that “Trigonotideae” are polyphyletic, and none of its members is at all closely related to Trigonotis itself. Moritzia and Thaumatocaryon are closely allied to each other and are the sister group of the Old World Boragineae. Flowers, pollen, and fruit morphology strongly support this systematic placement. Extant (native) Boragineae are absent from North America and had not previously been reported from South America, whereas members of the Cynoglosseae, Echiochileae, and Lithospermeae have been reported from both continents. Moritzia and Thaumatocaryon are thus the only native representatives of Boragineae in the Americas and represent an unexpected western Eurasian/South American disjunction. However, several (widespread and abundant) fossil taxa from the Cenozoic of North America (especially species of †Prolithospermum) can be confidently placed into Boragineae. Extant Moritzia /Thaumatocaryon likely go back to a lineage which reached North America from Europe and then migrated into South America, with subsequent extinction in North America.
The identity of Witheringia sellowiana, a narrow endemic species from São Paulo (Brazil), is reconsidered. After analysis of the protologue and the morphological characters of the type material and recent flowering collections, a new combination is proposed (Aureliana sellowiana (Sendtn.) Barboza & Stehmann). A lectotype and epitype are designated here. A detailed description and illustration, an analysis of the karyotype of A. sellowiana, and a discussion of its closest relatives are provided. A key to differentiate the species of Aureliana is also given.
Previous attempts to resolve relationships among the primary lineages of Campanulidae (e.g. Apiales, Asterales, Dipsacales) have mostly been unconvincing, and the placement of a number of smaller groups (e.g. Bruniaceae, Columelliaceae, Escalloniaceae) remains uncertain. Here we build on a recent analysis of an incomplete data set that was assembled from the literature for a set of 50 campanulid taxa. To this data set we first added newly generated DNA sequence data for the same set of genes and taxa. Second, we sequenced three additional cpDNA coding regions (ca. 8,000 bp) for the same set of 50 campanulid taxa. Finally, we assembled the most comprehensive sample of campanulid diversity to date, including ca. 17,000 bp of cpDNA for 122 campanulid taxa and five outgroups. Simply filling in missing data in the 50-taxon data set (rendering it 94% complete) resulted in a topology that was similar to earlier studies, but with little additional resolution or confidence. In contrast, the addition of the ca. 8,000 bp of sequence data provided resolution and support for a number of relationships. With the addition of many more campanulid taxa we recovered a well-resolved phylogeny for Campanulidae where, with the exception of the placement of Escalloniaceae, support was high for all major clades. In addition, we were able to confidently place a number of enigmatic taxa, some whose relationships were previously unresolved and some never before included. In light of these results, we briefly expand the discussion of floral symmetry and provide a phylogenetic classification of Campanulidae with phylogenetic definitions for all well-supported major clades.
Although recent molecular phylogenetic studies of Dipsacales support a Caprifolieae clade containing Leycesteria, Triosteum, Symphoricarpos, and Lonicera, relationships among these taxa remain uncertain. Likewise, phylogenetic studies sampling more than just a handful of the species of Symphoricarpos has not been pursued. Here I present evidence based on both chloroplast and nuclear DNA to help resolve the phylogenetic relationships of both the Caprifolieae, as well as Symphoricarpos. For nuclear markers, I sequenced the ITS region in addition to the geranylgeranyl reductase (Chlp) gene, which appears to be present in only a single copy in Caprifolieae. Although Chlp does not provide a great deal of resolution by itself, it does provide additional phylogenetically informative characters. From these data, there appears to be strong congruence between both the genomes with respect to the relationships within Symphoricarpos, however, statistical support for any given hypothesis is weak. These additional data do not further support or clarify our understanding about the relationships among members of the Caprifolieae, showing conflicting results between the chloroplast and nuclear genomes.
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