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Although previous cladistic analyses revealed Braunia secunda and B. andrieuxii as two lineages and valid species, they have been considered synonyms due to morphological similarities and sympatric distribution. This study was undertaken to evaluate which morphological characters best distinguish these two species. A total of 180 specimens of B. andrieuxii and 112 of B. secunda from the U.S. and Mexico were examined for morphometric analyses. Digital images of capsules, leaves, and groups of leaf cells were used to extract 31 linear measurements. Specimens were subdivided into seven geographic groups to compare levels of variation within and between species with univariate and multivariate analyses. Most characters are similar among groups within species, except the length of the revolute leaf margin and apical and upper leaf cells, which are significantly different between species (ANOVA and MRT). Partitioned Canonical Variates Analyses on eight variables of leaf cells and seven variables of vegetative leaves identified significant Mahalanobis distances between the two species. These methods also revealed that the revolute leaf margin and upper leaf cells contribute most to the distinction between species.
Previous hypotheses of relationships within Dioon Lindl. indicated the presence of two large clades within the genus. However, relationships among species still remained unresolved. In this study, molecular phylogenetic analyses were performed with individual and combined data sets from the ITS region of the ribosomal DNA and the trnL–F region of the chloroplast DNA. To explore whether indels were a source of phylogenetically informative characters, indels were analyzed by excluding them from the analyses (Coding A); including them as ambiguous data (Coding B); as multistate (Coding C) and as binary characters (Coding D). We found that the rate of mutation in the ITS region is appropriate to solve most relationships at the species level. This is in contrast with the trnL–F rgion, which showed little variation. Our results show that most clades obtained during analyses correspond with previously recognized species within Dioon. This phylogeny groups the genus into two main clades that show clear biogeographic relationships between the species occurring along the Pacific Sierra Madre Occidental and the Atlantic Sierra Madre Oriental.
Supported by detailed phenetic macro- and micromorphological characters, ecology, chorological studies, numerical and phylogenetic analysis, and all known data about Eurasian species of Carex muricata group (sect. Phaestoglochin), together with their biogeography, we have redefined this group and its subordinate aggregates. Four new taxa are described, illustrated and compared with putative related species: Carex spicata subsp. andresii, endemic to the inner part of the Iberian Peninsula, has the perigynum beak smooth and slightly corky at the base. Carex omeyica, from the southern mountains of Spain and North Africa, has erect-spreading oval perigynia and oval brown glumes without scarious margins. Two new subspecies of C. muricata are described: C. muricata subsp. cesanensis, from European mountains, has a big trullate perigynium 4.5–5.25 mm long with a short beak, and C. muricata subsp. ashokae, from the Eastern European Mountains to the Himalaya, has an oval perigynium 5–6 mm long with a longer beak. On the other hand, C. pairae is not a taxon subordinate to C. muricata. A key to the new and related taxa belonging to the aggregate is included.
Alkali grasses (Puccinellia) are temperate and arctic grasses of coastal and alkaline habitats, with ploidy levels that range from diploid to octoploid. This paper investigates the species limits of diploid alkali grasses in the North American Arctic. We used flow cytometry to confirm that four to seven of the 13–19 initially recognized taxa in the North American Arctic are diploid. Multivariate analyses of both morphological and amplified fragment length polymorphism (AFLP) data were congruent in resolving five or six diploid species: 1) P. arctica of northern and western distribution, including P. agrostidea and P. poacea; 2) a new diploid species from Banks Island, N.W.T., Canada; 3) P. tenella subsp. langeana; 4) P. alaskana, which had been previously treated as a subspecies of P. tenella; and 5) circumpolar (except coastal Beringia) P. vahliana, from which 6) the coastal Beringian endemic P. wrightii is distinguishable only on size and for which subspecies status may be more appropriate. In common garden experiments, eleven of 21 quantitative morphological characters varied significantly between field and common garden, showing phenotypic plasticity that explains much of the difficulty in identification. We present a map showing known geographic ranges of diploid Puccinellia species in the North American Arctic.
A new species of Pariana, found only in remnants of the Atlantic moist forest in the state of Espírito Santo in eastern Brazil, is described. The inflorescence structure in the new Pariana multiflora resembles those of P. lanceolata and P. carvalhoi, two species endemic to the Atlantic moist forests in southern Bahia, and all three are considered CR (critically endangered) by IUCN Red List criteria. The new species differs mainly by a greater number of inflorescences (up to five) per fertile culm and by the purple adaxial surface of the leaf blade, as well as by conical papillae and long unicellular trichomes that are present on both adaxial and abaxial leaf blade surfaces. Description of its morphology and leaf anatomy, compared with the other two mentioned species, based on optical microscopy and scanning electron microscopy, as well as illustrations and ecological data, are provided.
Four new Brazilian species from the genus Paspalum are described and illustrated:P. phaeotrichum, P. vexillarium, P. veredense, and P. clipeumPaspalum phaeotrichum is an annual with no obvious affinity to any known species of Paspalum, although it shares several characters with species of both P. subg. Ceresia and the ‘Bertoniana’ group. Paspalum vexillarium is presumably related to P. ceresia, with which it has been confused. Paspalum veredense shows affinities with both P. ellipticum and P. erianthoides. Paspalum clipeum is probably related to annual species of the ‘Plicatula’ group, although it lacks the dark brown upper florets typical of that group. Moreover, P. spissum, a species currently considered as a synonym under P. carinatum, is recognized on the basis of vegetative morphology, phenology and habitat. Keys to the treated species are provided when relevant, as well as a distribution map.
Resumo—Quatro novas espécies de Paspalum do Brasil são descritas e ilustradas:P. phaeotrichum, P. vexillarium, P. veredense, e P. clipeum. Paspalum phaeotrichum é uma espécie anual sem afinidades obvias com outras espécies do gênero, embora compartilhe vários caracteres com P. subg. Ceresia e com o grupo ‘Bertoniana'. Paspalum vexillarium é supostamente relacionada a P. ceresia, com a qual tem sido confundida. Paspalum veredense tem afinidades com P. ellipticum e P. erianthoides. Paspalum clipeum é provavelmente relacionada às espécies anuais do grupo ‘Plicatula', embora não apresente os antécios superiores de cor marrom escura, típicos desse grupo. Além disso, P. spissum, uma espécie atualmente considerada como sinônimo de P. carinatum, é reabilitada com base na morfologia vegetativa, fenologia e habitat. São fornecidas chaves para as espécies tratadas, quando relevante, assim como um mapa de distribuição.
Representative taxa of the five genera of Guaduinae, a subtribe of Neotropical woody bamboos, were sampled to investigate the phylogenetic relationships of the species of the genus Otatea using morphological and molecular (cpDNA intergenic spacer trnH-psbA and the rpl16 intron) evidence. Phylogenetic analysis of a combined data set retrieved 53 most parsimonious trees in which subtribe Guaduinae is monophyletic if two species of Anlonemia (A. clarkiae and A. fulgor) are included. They were previously classified within subtribe Arthrostylidiinae. Guaduinae is supported by the lack of papillae from the abaxial surface, by an almost solid style, a short rachis extension, and oral setae present in culm and foliage leaves. Monophyly of the genera in Guaduinae (Eremocanlon, Guadna, Apoclada, Otatea, and Olmeca) was corroborated. Otatea species formed a monophyletic clade, supported by culms with three subequal ascending branches and pubescent lemmas. Eight species in Guaduinae (the four species in Otatea, Olmeca recta, O. reflexa, Aulonemia clarkiae, and A. fulgor) are distributed in southeastern Mexico in areas determined as Pleistocene refugia. Some of them possess baccoid caryopses and long culm necks, and grow in threatened vegetation types such as cloud, tropical, and tropical deciduous forests, so they are important bamboos to preserve.
The taxonomic status of Panicum section Lorea has remained as “incertae sedis” within Panicum. To resolve its position within the Paniceae and to test the monophyly of this section, phylogenetic analyses based on chloroplast sequence data (ndhF) and morphology were conducted for the Paniceae with particular emphasis on Panicum section Lorea. The results did not support the monophyly of this section. The species of this group were resolved in two clades which are not sister groups and neither of them is closely related to Panicum s.s. As a result, two new genera are proposed and described: Apochloa and Renvoizea, which are restricted to the Guiana highlands and eastern Brazil. New combinations are: Apochloa animara, A. bahiense, A. chnoodes, A. cipoense, A. eligulata, A. euprepes, A. jauana, A. lorea, A. lutzii, A. molinioides, A. poliophylla, A. sipapoense, A. steyermarkii, A. subtiramulosa, A. tijucae, Renvoizea acicularifolia, R. durifolia, R. glaziovii, R. lagostachya, R. marauense, R. restingae, R. sacciolepoides, R. teretifolia, R. trinii, and R. vaginiviscosa.
Seventeen morphological characters are described and plotted on a phylogeny of the eudicots. The distribution of perianth characters demonstrates that the currently held view that petals originated from stamens in the core eudicots is not consistent with the predominance of bract-derived petals (bracteopetals). Petals in the core eudicots have the same morphological affinities and structure as sepals or bracts, with a few notable exceptions. Depending on the most likely topology, Berberidopsidales may represent the plesiomorphic perianth condition in the core eudicots, giving rise to pentamerous flowers with sepals and petals of a similar tepaline origin. Within the core eudicots several divergent trends have led to a wide variety of perianth morphologies involving to different degrees the presence of a calyx and corolla. Different factors contribute to the development of a biseriate perianth, including the upward shift of bracts and their absorption in the flower and a progressive differentiation of tepals through shifts in petaloidy. A number of correlations in perianth evolution are highlighted, such as petaloid sepals linked with loss of petals and presence of a hypanthium, a reversal to tepal-like petals with secondary polymery in the androecium and gynoecium, or the loss of sepals linked with the development of stamen-petal tubes. The relationship between petal morphology and the molecular basis for petaloidy is discussed.
Chloroplast (trnL−F and rbcL) sequences were used to reconstruct the phylogeny of Geraniaceae and Hypseocharitaceae. According to these data Hypseocharitaceae and Geraniaceae are monophyletic. Pelargonium and Monsonia are sisters to the largest clade of Geraniaceae, formed by Geranium, Erodium and California. According to molecular dating and dispersal-vicariance analysis, the split of the stem branches of Geraniaceae probably occurred during the Oligocene, in southern Africa or in southern Africa plus the Mediterranean area. However, their diversification occurred during the Miocene, coinciding with the beginning of major aridification events in their distribution areas. An ancestor of the largest clade of Geraniaceae (Geranium, Erodium, and California) colonised a number of habitats in the northern hemisphere and in South American mountain ranges. In summary, the evolution of the Geraniaceae is marked by the dispersal of ancestors from Southern Africa to cold, temperate and often disturbed habitats in the rest of world, where only generalist pollination and facultative autogamy could ensure sufficient seed production and survival.
Two new species of Eugenia (Myrtaceae) are described from the Masoala Peninsula in Madagascar: Eugenia williamsiana and Eugenia ambanizanensis. Each has relatively large leaves among Malagasy taxa of the genus. Both species are presently known from only two collections, but since they occur in a well-protected area they are considered Least Concern under IUCN guidelines. Features of the inflorescence and floral morphology of E. ambanizanensis are atypical for Eugenia and are discussed in light of the putatively related genera Stereocaryum, Monimiastrum, and Plinia. A tentative key is provided for species of Eugenia from Madagascar in which some or all of the mature leaves are in excess of 15 cm in length. Based on published and unpublished DNA sequencing data, five species of Monimiastrum are transferred into Eugenia.
The Old World genus Trichosanthes has flowers with strikingly fringed petals, and Linnaeus therefore placed a species from Hispaniola that he only knew from an illustration (showing such fringed petals) in that genus. The species remained hidden from the attention of subsequent workers until acquiring new relevance in the context of molecular-biogeographic work on Cucurbitaceae. Based on molecular data, it is the sister to all Sicyeae, a New World clade of about 125 species in 16 genera. We here place this species in a new genus, Linnaeosicyos, describe and illustrate it, and discuss its phylogenetic context using molecular and morphological data. Judging from Dominican amber, elements of the flora of Hispaniola date back 15–20 my, and the occurrence on the island of at least five endemic species of Cucurbitaceae (Linnaeosicyos amara, Melothria domingensis, Sicana fragrans, and the sister species Anacaona sphaerica and Penelopeia suburceolata) points to its long occupation by Cucurbitaceae.
The approximately 27 New World species of Vicia (tribe Fabeae, family Leguminosae) were classified into four stylar type groups as follows: (1) a laterally compressed and evenly hairy stylar type (Le-type) group, (2) a dorsiventrally compressed and abaxially tufted hairy stylar type (Dabt-type) group, (3) a dorsiventrally compressed and evenly hairy stylar type (De-type) group, and (4) V. leuncophaea that is characterized by an arcuate style with a dense ring of stylar hairs at a considerable distance from the stigma. In the present study, the stylar distinctions among the groups were confirmed by morphological and anatomical studies on the stylar characters of representative species of the groups. The phylogenetic positions of these groups were then assessed using molecular phylogenetic analysis based on sequences from the nuclear ribosomal DNA (nrDNA) internal transcribed spacer (ITS) region of the representative species. The New World Le-type stylar group was revealed to be a sister group to the Old World Le-type stylar group. The New World Dabt-type stylar group and V. leucophaea are nested within the Old World Dabt-type stylar group. These New World groups were nested well within the Old World Vicia, suggesting that the New World diversification occurred relatively recently during the evolution of Vicia.
Madagascar boasts a substantial radiation within Hibisceae (Malvaceae). Molecular data were used to determine the number of migrations to Madagascar, to clarify the timing of these radiations, and to assess the relationships among the endemic taxa. Sequences of matK and ndhF were obtained for representatives of all the Malagasy endemic genera in Hibisceae, as well as representative Malagasy species of Hibiscus and Kosteletzkya. Phylogenetic analysis of these data along with a sampling of outgroups and other major lineages of Hibisceae strongly supported the existence of an exclusively Malagasy clade, the /Megistohibiscus clade, which is sister to the rest of tribe Hibisceae. Humbertiella, Megistostegium, Perrierophytum, all the Malagasy Kosteletzkya, and a number of endemic species usually placed in Hibiscus are members of /Megistohibiscus and appear to be derived from a single introduction to Madagascar in the mid Miocene. Other sampled Malagasy Hibisceae, including the endemic genera Helicteropsis, Jumelleanthus, and Macrostelia, were resolved as members of the /Euhibiscus clade, implying at least one additional dispersal to the island.
The African-Malagasy genus Protorhus and the African-Malagasy species of Rhus s.l. have been suggested to be polyphyletic, but definitive evidence has been lacking. We investigated a clade of Anacardiaceae subfamily Anacardioideae in order to clarify evolutionary relationships of the African-Malagasy members of these two genera. Phylogenetic analysis of sequence data from cpDNA trnL–trnF, and nrDNA ETS and ITS provide strong statistical support for the 1) polyphyly of Protorhus; 2) monophyly of Malagasy Protorhus species; 3) polyphyly of Rhus section Baronia (Malagasy Rhus). These results support the need for new circumscriptions of these groups. Two new combinations are proposed, Searsia erosa and Searsia pendulina.
Allopolyploidy is recognized as an important mode of plant speciation. The allopolyploid origin of a taxon is often inferred from morphological intermediacy or a mixture of morphological characters from the putative parental taxa, especially if accompanied by geographical intermediacy of the polyploid. Here we show that in the case of Androsace cantabrica (Primulaceae), a narrow endemic from the central Cordillera Cantábrica (Spain), both lines of evidence are misleading. Instead of being an allopolyploid derivative of A. halleri (western Cordillera Cantábrica, eastern Pyrenees, Massif Central, Vosges) and A. laggeri (central Pyrenees), A. cantabrica is most likely an autopolyploid closely related to A. adfinis s. 1., endemic to the southwestern European Alps. This counterintuitive biogeographic connection not only adds to the group of species showing a Cantabrian-Alpine disjunction, but, together with the unexpected link between A. chaixii, also from the southwestern Alps, and A. halleri in form of the northern Spanish local endemic A. rioxana, underlines the important role of large-scale range shifts in shaping the current distribution of European mountain taxa.
Heliotrapium sect. Cochranea (Heliotropiaceae) consists of 19 species endemic to the coastal Atacama Desert of Chile and Peru. This section has one of the most restricted geographic distributions and is the largest among the South American sections of Heliotropium. We performed a phylogenetic analysis of a total of 92 species using nuclear ribosomal ITS and chloroplast ndhF, rpsl6, and trnL-trnF sequences, and estimated the divergence times of major lineages of the group. Our results suggest that Heliotropium sect. Cochranea is monophyletic. There are two main well-supported lineages within the section: one is H. pycnophyllum, which is sister to rest of the species in the section. Within this second lineage, H. filifolium, H. glutinosum, H. krauseanum, and a large polytomous group composed of 15 species form a tetratomy. The age estimates using the penalized likelihood method suggests a minimum age of 14.0 ± 2.0 My for section Cochranea, and 4.6 ± 0.9 My for the large polytomous group within it. Heliotropium sect. Cochranea may have originated in the Miocene during the major uplift of the Andes, and then radiated in the Pliocene in the Atacama Desert.
Gonolobus (Apocynaceae, Asclepiadoideae) is a New World genus comprising an estimated 100–150 species. Variation in estimated species numbers is largely the result of still poorly known tropical taxa and differences regarding generic limits. Characters historically used to delimit genera such as Gonolobus within Gonolobinae—such as laminar dorsal anther appendages and various follicle morphologies—have been controversial and their evolution remains unknown, not having been explored in a phylogenetic framework. The primary objectives of the current study were to (1) test the monophyly of Gonolobus sensu Woodson in the context of a phylogeny of New World Asclepiadeae and (2) explore the evolution of laminar dorsal anther appendages and winged follicles with respect to their potential utility in generic circumscription. Chloroplast (trnL–F, rpsl6) data are newly presented for sixty-three taxa of Gonolobinae, representing an increased sampling of the subtribe from a maximum of seven taxa in prior studies. These data were combined with a previously published dataset to form a 183 taxa matrix of New World Asclepiadeae. Nuclear (LEAFY) data were newly generated for forty-nine taxa of Gonolobinae. Evidence from parsimony and Bayesian analyses of chloroplast and nuclear data presented here supports the monophyly of both the subtribe and the genus Gonolobus in a narrow or broad sense. Laminar dorsal anther appendages are restricted to Gonolobus s.l. or s.s., although parsimony remains equivocal on whether they evolved once or twice. A transversion in the trnL–F spacer is shown synapomorphic for Gonolobus s.s. Two indels in LEAFY, as well as winged follicles, are shown synapomorphic for Gonolobus s.l.
We used DNA sequence data from five genic regions (nrlTS; chloroplast trnL-F, trnT–L, rps16, trnS–G) to study phylogenetic relationships of the Tetramerium lineage (Acanthaceae: Justicieae). From a sample of 70 species (representing 25 genera) previously affiliated with the Tetramerium lineage, 68 are included therein. Our analyses excluded Papuasian Calycacanthus and Neotropical Streblacanthus monospermus from the Tetramerium lineage; however, two species described in Justicia (J. gonzalezii and J. medranoi) and a Malagasy species of uncertain generic affinities are nested within the lineage. A monophyletic Tetramerium lineage consists of 23 currently recognized genera with at least 168 species, more than 70% of which occur in the New World. Old World Chlamydocardia and Clinacanthus are serially sister to all other members of the lineage. Other Old World taxa consist of: Ecbolium clade (all sampled species of Ecbolium plus Malagasy Papulina richardii), Megalochlamys clade (Megalochlamys, Trichaulax and the unidentified Malagasy species), and two isolated taxa (Angkalanthus and Chorisochora). All analyses strongly support monophyly of the New World Tetramerium lineage. The basal dades of New World plants, all with nototribic flowers, are: 1) the taxonomically heterogeneous but palynologically consistent Mirandea clade, and 2) the Pachystachys clade the South American Anisacanthus clade. The second is sister to all other NW plants, referred to here as the core Tetramerium lineage. We recognize five clades within the core Tetramerium lineage related as follows: (Henrya clade (Carlowrightia parviflora clade (North American Anisacanthus clade (core Carlowrightia clade Tetramerium)))). Macromorphological synapomorphies are unknown for the Tetramerium lineage and for many of its constituent clades. However, we propose sternotribic flowers as synapomorphic for the core Tetramerium lineage, and flowers with the lower-central lobe of the corolla modified into a keel as a synapomorphy for a lineage consisting of Tetramerium and the core Carlowrightia clade. Palynological characters provide putative synapomorphies for some clades (e.g. Ecbolium clade, Mirandea clade) and autapomorphies for several species (e.g. Mexacanthus mcvaughii, Trichaulax mwasumbii). An Old World origin is postulated for the Tetramerium lineage, and we posit a single dispersal event to America and subsequent extensive radiation there, especially in arid zones of Mexico and adjacent regions. Taxonomic implications of our results are extensive. Notably, many traditionally recognized genera
Trichostema (Lamiaceae) is a North American genus comprising five sections and 18 species. Phylogenetic relationships were inferred using DNA sequences (ITS, ndhF) and morphology (63 characters). The monophyly of Trichostema is well supported in four analyses (ITS, ndhF, ITS ndhF, ITS ndhF morphology), but not in the analysis based on morphological data alone. In all analyses, the monophyly of sections Chromocephalum and Trichostema is well supported. The monophyly of section Paniculatum is moderately supported, and there is conflicting evidence regarding the monophyly of section Orthopodium. Although DNA data alone suggest that T. brachiatum is not a member of section Orthopodium, its retention in section Orthopodium is equivocal in the combined DNA-morphology analyses. The submerging of the monotypic section Rhodanthum into Paniculatum by Lewis and Rzedowski is not adopted because the monophyly of this grouping is poorly supported. Biogeographic history and character evolution (life history and chromosome number) are explored in the context of the phylogeny based on the combined ITS and ndhF sequence data.
Oxypolis and Ptilimnium are two endemic genera of North America within tribe Oenantheae (Apiaceae subfamily Apioideae). Both genera are small, with 12 species currently recognized. Some members of each genus share an unusual leaf morphology. Rather than having the pinnately compound leaves that generally characterize apioid umbellifers, they have highly reduced, linear, terete, hollow, septate appendages known as rachis leaves. It has long been questioned whether the species with rachis leaves should be placed in separate genera. In this study, we use data from the internal transcribed spacer region of nuclear ribosomal DNA to explore relationships within and between these genera. A total of 147 internal transcribed spacer sequences were obtained from multiple accessions of all species of Oxypolis and Ptilimnium and for several other genera from tribe Oenantheae. These included Lilaeopsis, which also has rachis leaves, and Cynosciadium and Limnosciadium, which have rachis-like leaves. These sequence data were analyzed using maximum parsimony, maximum likelihood, and Bayesian methods. The results from each of these analyses were congruent and suggest that neither Oxypolis nor Ptilimnium as presently defined is monophyletic. Oxypolis and Ptilimnium are each separated into two dades according to leaf morphology. The rachis-leaved Oxypolis species are provisionally recognized as the genus Tiedemannia and the rachis-leaved Ptilimnium species are provisionally recognized as the genus Harperella, pending further investigation. The relationships among these four clades and the genera Cynosciadium, Daucosma, and Limnosciadium are not clear. Nevertheless, it appears that the rachis-leaf habit has evolved multiple times in the tribe. Geographic structure is apparent in the phylogenetic trees and, pending further study, may suggest new taxa; in addition, the presence of O. occidentalis on the Queen Charlotte Islands, well-separated from other populations of this species, suggests that it may have survived the last glaciation in a refugium in this area.
A new species, Perityle harkerae (Asteraceae), endemic to southern Zacatecas, Mexico is described and illustrated. Salient morphological features of the new species are compared to those of P. jaliscana, P. feddemae, and P. glaucescens, which appear to be closely related. The new species is distinguished by its canescent and ovate to deltate-ovate leaves, and by its discoid capitula with yellow corollas. A key to the species of Perityle in western Mexico is provided.
Una nueva especie, Perityle harkerae (Asteraceae), endémica al sur de Zacatecas, México es descrita e ilustrada. Algunas características morfológicas sobresalientes son comparadas con las de P. jaliscana, P. feddemae y P. glaucescens, que parecen ser las especies más cercanamente relacionadas. El nuevo taxón se distingue por sus hojas canescentes, ovadas a deltadoovadas y por sus capítulos discoides con corolas amarillas. Se presenta una clave para la identificatión de las especies de Perityle en el occidente de México.
A 3.7 kilobase region of chloroplast DNA that includes atpB, rbcL, and their intergenic spacer was sequenced in 61 samples from 45 species of South American Lobeliaceae plus two outgroup samples from Australia. A clade of four hexaploid Lobelia species from Chile is sister to a clade comprising Lysipomia, Siphocampylus, Centropogon, and Burmeistera. Lysipomia is a monophyletic group of small cushion-forming plants endemic to the high Andes, and is sister to the clade comprising the remaining three shrubby genera, which are most diverse in the Andes, but also extend to Central America, Mexico, and the West Indies. Siphocampylus has capsular fruit and is inferred to be paraphyletic relative to fleshy-fruited Centropogon and Burmeistera, but fleshy fruits have evidently evolved repeatedly, making Centropogon polyphyletiC. Burmeistera is primarily bat-pollinated and monophyletic, having evolved from one group of species in Centropogon. The phylogenetic relationships within Burmeistera indicate that this genus underwent repeated episodes of rapid diversification when organismal diversification outpaced the accumulation of mutations in this region of chloroplast DNA.
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