Haplodontium zangii X.R.Wang & J.C.Zhao,sp. nov. (Bryaceae,Bryopsida) from Xizang,China

Haplodontium zangii X.R.Wang &; J.C.Zhao, a new moss species from Xizang, China, is described and illustrated. Previously, specimens of H. zangii have been identified as Mielichhoferia himalayana Mitt. However, H. zangii is distinctly different from M. himalayana in having excurrent costae with short awns (vs long denticulate awns), short-pyriform capsules, 0.8–1?mm (vs long-pyriform capsules, 2.5–3?mm), and densely papillose exostome teeth (vs smooth or vertically striped exostome teeth). Haplodontium zangii is similar to H. macrocarpum (Hook.) J.R.Spence, which was traditionally placed in Mielichhoferia Nees &; Hornsch. as M. macrocarpa (Hook.) Bruch &; Schimp. The main differences between H. zangii and H. macrocarpum are in the morphology of the leaves, capsules, guide cells, and stomata. Mielichhoferia himalayana and another Chinese species of Mielichhoferia, M. sinensis Dix., are also transferred to Haplodontium Hampe, a new genus in the bryoflora of China, as H. himalayanum (Mitt.) X.R.Wang &; J.C.Zhao and H. sinensis (Dix.) X.R.Wang &; J.C.Zhao. A morphological comparison and a key to the three species of Haplodontium in China as well as to H. macrocarpum, a species that is likely to be found in China, are provided.     

A proposed reclassification of Bryum,Anomobryum and Brachymenium (Musci,Bryaceae)

Abstract

The traditional generic classification in the Bryoideae (Bryaceae) is based primarily on details of the sporophyte, particularly peristome reduction and capsule inclination. Recent research, however, has suggested that closely related species in Anomobryum can have very distinct capsules, varying from complete peristomes and inclined capsules to reduced peristomes and erect capsules. The suggestion is made here that sporophyte features, especially those related to peristome features, may obscure phylogenetic relationships. A revised classification is proposed, based on analyses of 100 species of Bryum, Anomobryum, and Brachymenium and sixteen sporophytic and gametophytic characters. The revised classification groups species with similar gametophores but is supported by at least some sporophytic characters. Analysis of variance and principal components analysis indicate that both the traditional and revised classification provide effective summaries of the variation within the 100 species examined. The new classification consists of three informal groups of species which can be recognized at generic rank. One comprises Bryum species, including those with reduced peristomes and inclined capsules. A second comprises all species of Anomobryum, as well as species of Bryum and Brachymenium with similar gametophores. The third group consists of the rosulate species of Bryum and Brachymenium. Phylogenetically, the second group may be more closely related to Pohlia than to the species in the first and third groups. The third group appears to be most closely related to Rhodobryum and Plagiomnium. Suggested names are provided for groups one and three, while the name Bryum is retained for group two. No nomenclatural changes are made, however, until more research is done.     

Phylogenetic relationship of Clauseneae (Rutaceae) inferred from plastid and nuclear DNA data and taxonomic implication for some major taxa

Using sequences of the nuclear ribosomal ITS region as well as the chloroplast DNA trnL‐trnF and atpB‐rbcL regions, this study aims to provide further insight into the phylogenetic relationships within Clauseneae and its relationship to Citreae (Rutaceae). Using maximum likelihood (ML) and Bayesian inference (BI), we reconstructed the phylogeny of Clauseneae based on trnL‐F, atpB‐rbcL and ITS sequences. Our data matrix contained 91 accessions, representing 72 species and varieties from 31 genera and two outgroups, including new and extensive sampling of species and varieties representing four genera in the tribe Clauseneae. In the subfamily Aurantioideae, six major clades were resolved with strong support: 1) Micromelum clade: Micromelum; 2) Glycosmis clade: Glycosmis; 3) Bergera clade: Murraya sect. Bergera; 4) Clausena clade: Clausena; 5) Murraya clade: Murraya sect. Murraya + Merrillia; 6) Citreae clade: Citreae. Micromelum, Glycosmis, Clausena and Merrillia were confirmed as monophyletic. In contrast, Murraya s.l. was reconstructed as polyphyletic. Murraya sect. Bergera clustered with Clausena while Murraya sect. Murraya and Merrillia together formed a clade that is sister to the tribe Citreae. All members from Citreae were clustered into a natural group. The genus Micromelum was found to be primitive in this subfamily and more close to Glycosmis. Based on the phylogeny and morphological characters, we discuss the taxonomy of some members of Clauseneae and conclude that the current tribal and generic classification need further revision.     

Phylogeny of Carpinus and subfamily Coryloideae (Betulaceae) based on chloroplast and nuclear ribosomal sequence data

Phylogenetic studies were conducted for Carpinus and the subfamily Coryloideae (Betulaceae) using sequences of the chloroplast matK gene, the trnL-trnF region (trnL intron, and trnL [UAA] 3' exon-trnF [GAA] intergenic spacer) and the psbA-trnH intergenic spacer, and the nuclear ribosomal ITS regions. The combined analyses of the three chloroplast regions suggest that Coryloideae is monophyletic; Ostryopsis is sister to the Carpinus - Ostrya clade; Corylus is monophyletic and sister to the Ostrya - Carpinus - Ostryopsis clade; Ostrya is paraphyletic; and within Carpinus, species of sect. Carpinus from eastern Asia form a monophyletic group, whereas the positions of C. betulus from Europe and C. caroliniana from eastern North America are unresolved within the Carpinus clade. The cpDNA tree generated in this study is largely congruent with the previously published ITS results, but the ITS tree places Carpinus sect. Distegocarpus as sister to the Ostrya - Carpinus sect. Carpinus clade. Future work is needed to examine the relationships within the Ostrya - Carpinus clade, evaluate the generic status of Ostrya, and test the phylogenetic position of Ostryopsis.     

Phylogenetic relationships between Bryum and supposedly closely related genera

Abstract

The phylogeny of the genus Bryum was studied using cladistic analyses under the maximum parsimony criterion of evolution of anatomical and morphological characters. Three analyses were made with 32 Bryum species plus 20 species from genera supposedly closely related to Bryum, and with Amblyodon dealbatus (Sw. ex Hedw.) Bruch & Schimp., Meesia uliginosa Hedw., and Leptostomum macrocarpum (Hedw.) Bach. Pyl., as outgroups. It is here suggested that under earlier systematic concepts the genus Bryum is paraphyletic. A clade with Bryum billarderi Schwägr., B. capillare Hedw., B. donianum Grev., B. russulum Broth. & Geh., Rhodobryum giganteum (Schwägr.) Paris, and R. keniae (Müll. Hal.) Broth. are circumscribed by spathulate stem leaves that are crowded in the stem apex, suggesting that the rosulate species of Bryum are more closely related to Rhodobryum than to the rest of Bryum. Stem leaf costae without stereids and spores that mature in the winter are synapomorphies for a clade with Anomobryum julaceum (P. Gaertn. et al.) Schimp. and Bryum argenteum (Hedw.). The tropical species B. cellulare Hook. and B. flaccum Wilson ex Mitt. appear in a clade with Plagiobryum zieri (Dicks. ex Hedw.) Lindb. and Synthetodontium pringlei Cardot. In one analysis, B. limbatum Müll. Hal., Epipterygium tozeri (Grev.) Lindb., Leptobryum pyriforme (Hedw.) Wilson, and Roellia roellii (Broth. ex Röll) H.A. Crum came out in a clade with Mniobryum atropurpureum (Wahlenb.) I. Hagen, Mnium hornum Hedw., Pohlia cruda (Hedw.) Lindb., P. longicollis (Hedw.) Lindb., and Pseudopohlia didymodontia (Mitt.) A.L.Andrews. It is here suggested that gametophytic features, such as the orientation and anatomy of the stem leaves and the appearance of vegetative propagules, are important for the internal relationships within the studied ingroup, whereas characters related to the sporophyte, especially those of the peristome, may obscure phylogenetic relationships. Most of the subgenera and the sections of Bryum, as defined by earlier authors, appear to be paraphyletic. However, due to the low stability of most clades it is suggested that analyses including anatomical, morphological, and molecular data are needed.     

A tribute to Dr Royce E. Longton

Abstract

The relationships within the Bryaceae, with emphasis on the genus Bryum, were studied based on morphological and anatomical characters and using cladistic methods. The analysis was performed with thirty-six species representing the different parts of the family, and with Funaria hygrometrica Hedw., Mnium hornum (Dicks.) Lindb., and Tayloria lingulata Hedw. as outgroups. The Bryaceae, and the subfamilies Bryoideae, Mielichhoferioideae, and Pohlioideae, as defined by several earlier authors appear to be paraphyletic. The genus Bryum seems to be paraphyletic, because Leptobryum pyriforme (Hedw.) Wils., Osculatia columbica De Not., and Rhodobryum giganteum (Schwaegr.) Paris, appear as ingroups within this genus when the tree is rooted with Funaria. Mnium hornum came out as the sister taxon of a clade including Pohlia cruda (Hedw.) Lindb. and P. longicollis (Hedw.) Lindb., whereas P. drummondii (Müll. Hall.) A.L. Andrews, appears not to be closely related to the other two Pohlia species studied here, making this genus paraphyletic. Mielichhoferia mielichhoferiana (Funck.) Loeske, appears as the sister taxon of Schizymenium bryoides Harv., suggesting that both these genera are paraphyletic. Overall, the stabilities of the clades are low and it is suggested that combined analyses of morphological, anatomical, and molecular data are needed to get better resolved and more stable trees.     

Phylogenetic relationships within Pappophoreae s.l. (Poaceae: Chloridoideae): Additional evidence based on ITS and trnL-F sequence data

Historically, Pappophoreae included the genera Cottea, Enneapogon, Kaokochloa, Pappophorum and Schmidtia. Some authors consider this tribe as a well-supported monophyletic group; while other evidences reveals Pappophoreae as polyphyletic, with Pappophorum separated from the rest of the tribe. When the latter happens, it can form a clade with Tridens flavus. Molecular phylogenetic analyses of the subfamily Chloridoideae have included few species of Pappophoreae; therefore, further research involving more representatives of this tribe is needed. With the aim of providing new evidence to help clarify the phylogenetic position of Pappophorum and its relationships with other genera of the tribe and the subfamily Chloridoideae, eight new sequences of ITS and trnL-F regions of Pappophoreae species were generated. These sequences were analyzed together with other available sequence data obtained from GenBank, using maximum parsimony and Bayesian inference, for individual (trnL-F or ITS) or combined trnL-F/ITS data sets. All analyses reveal that Pappophoreae is polyphyletic, with Pappophorum separated from the rest of the tribe forming a well-supported clade sister to Tridens flavus.     

Molecular phylogenetics and generic assessment in the tribe Morindeae (Rubiaceae–Rubioideae): How to circumscribe Morinda L. to be monophyletic?

Most of the species of the family Rubiaceae with flowers arranged in head inflorescences are currently classified in three distantly related tribes, Naucleeae (subfamily Cinchonoideae) and Morindeae and Schradereae (subfamily Rubioideae). Within Morindeae the type genus Morinda is traditionally and currently circumscribed based on its head inflorescences and syncarpous fruits (syncarps). These characters are also present in some members of its allied genera, raising doubts about the monophyly of Morinda. We perform Bayesian phylogenetic analyses using combined nrETS/nrITS/trnT-F data for 67 Morindeae taxa and five outgroups from the closely related tribes Mitchelleae and Gaertnereae to rigorously test the monophyly of Morinda as currently delimited and assess the phylogenetic value of head inflorescences and syncarps in Morinda and Morindeae and to evaluate generic relationships and limits in Morindeae. Our analyses demonstrate that head inflorescences and syncarps in Morinda and Morindeae are evolutionarily labile. Morinda is highly paraphyletic, unless the genera Coelospermum, Gynochthodes, Pogonolobus, and Sarcopygme are also included. Morindeae comprises four well-supported and morphologically distinct major lineages: Appunia clade, Morinda clade (including Sarcopygme and the lectotype M. royoc), Coelospermum clade (containing Pogonolobus and Morinda reticulata), and Gynochthodes–Morinda clade. Four possible alternatives for revising generic boundaries are presented to establish monophyletic units. We favor the recognition of the four major lineages of Morindeae as separate genera, because this classification reflects the occurrence of a considerable morphological diversity in the tribe and the phylogenetic and taxonomic distinctness of its newly delimited genera.     

A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data

 Phylogenetic relationships in Primulaceae were investigated by analysis of nuclear rDNA ITS sequences. Thirty-four species of Primulaceae, two of Myrsinaceae and four outgroup taxa were analyzed. In accordance to the results of recently published papers on the phylogeny of Primulaceae we found the family to be paraphyletic and resolved the positions of some genera. Our results show (a) the rather basal position of Centunculus within Lysimachieae, the genus thus being rather distantly related to Anagallis, (b) the close relationship between Lysimachia sect. Lerouxia, Anagallis, Asterolinon, and Pelletiera, (c) the well-supported monophyly of a group consisting of the four genera Hottonia, Omphalogramma, Bryocarpum, and Soldanella, and (d) the affinity of Stimpsonia to the Myrsinaceae-Lysimachieae-Ardisiandra clade. The ITS sequence data do not provide sufficient information to resolve basal relationships within the Primulaceae s.l. There is evidence against the monophyly of the large genera Primula, Androsace, and Lysimachia. In contrast to the phylogenetic reconstructions based on plastid gene sequences, Cyclamen does not appear as a member of the Myrsinaceae-Lysimachieae clade, but its position remains unclear. Revised July 10, 2002; accepted November 21, 2002 Published online: March 20, 2003     

Molecular phylogeny of the endemic fern genera Cyrtomidictyum and Cyrtogonellum (Dryopteridaceae) from East Asia

Cyrtomidictyum Ching and Cyrtogonellum Ching are two eastern Asian endemic genera whose taxonomic affinities and phylogenetic relationships have long been controversial. The main uncertainty surrounds the separation of the two genera from the species-rich genus Polystichum. Here we present a phylogenetic study focusing on the phylogenetic relationships of these polystichoid ferns. We reconstructed the relationships based on DNA sequence variation in four chloroplast genome regions, rbcL, atpB, and the intergenic spacers (IGS) rps4-trnS and trnL-trnF. Maximum likelihood and Bayesian inference analyses confirm earlier results that were based on less comprehensive taxon sampling and either only a single gene (rbcL) or two IGS (rps4-trnS and trnL-trnF). Cyrtomidictyum is the sister of the clade of polystichoid ferns that includes Cyrtogonellum, Cyrtomium subser. Balansana and three sections of Polystichum. Cyrtogonellum groups with several species of Polystichum, and constitutes the sister taxon to Polystichum sect. Sphaenopolystichum. We support the recognition of Cyrtomidictyum as circumscribed initially, rather than expansion of the genus to include either several Polystichum species or Cyrtogonellum, some Polystichum and Cyrtomium species. The monophyly of Cyrtomidictyum is supported by morphological characters such as once-pinnate leaves, free venation, prolongated leaf apices, and exindusiate sori. Two synapomorphic indels in the chloroplast genome, one 15-bp deletion in rps4-trnS, and one 3-bp insertion in trnL-trnF further differentiate Cyrtomidictyum from other polystichoid ferns. The close affinity of Cyrtogonellum to section Sphaenopolystichum of Polystichum s.s. is highly supported by molecular data. However, no shared morphological characters or molecular indels have been detected, although the distinctness of Cyrtogonellum is shown by a 13-bp insertion in the rps4-trnS alignment.     

Synopsis of <Emphasis Type="Italic">Mickelia</Emphasis>, a newly recognized genus of bolbitidoid ferns (Dryopteridaceae)

Our recent molecular phylogenetic study revealed a previously unrecognized clade of six species that is sister to Elaphoglossum. Within this clade, four species are currently classified in Bolbitis, one in Lomagramma, and one in Acrostichum. For this clade, we propose the name Mickelia, with M. nicotianifolia as the type species. We also make new combinations for the species in our phylogenetic study shown to belong to Mickelia (M. bernoullii, M. guianensis, M. hemiotis, M. nicotianifolia, M. oligarchica, and M. scandens) and two other species believed to belong to the clade based on morphology (M. lindigii, M. pergamentacea). A new hybrid and two new species are also described (M. ×atrans, M. furcata, and M. pradoi). In total, Mickelia consists of ten species and one hybrid. It is entirely neotropical. We provide a key to the genera of bolbitidoid ferns and a synopsis of Mickelia that gives for each species a complete synonymy, geographical distribution, comparative discussion, and illustration.     

A molecular phylogeny and generic rearrangement of the stapelioid Ceropegieae (Apocynaceae-Asclepiadoideae)

 Representatives of nearly all genera of the taxon-rich stem-succulent stapeliads and most of the few related, leafy genera were analyzed. Sequence data from two non-coding molecular markers (ITS region of nrDNA and trnT-L and trnL-F spacers as well as the trnL intron of cpDNA) support the traditional tribal affiliation of the genera, which form a monophyletic group. This monophylum breaks into a basal Neoschumannia/Anisotoma/Riocreuxia/Sisyranthus nk;clade, from which the core Ceropegieae are derived. The four Ceropegia species included are not monophyletic, and their relationship to Brachystelma changes depending on the marker studied. The stem succulent taxa fall in a number of well supported, but unresolved clades, the most prominent being the predominantly southern African clade comprising Orbea, Stapelia and some other genera. The most derived taxa of NE Africa, Duvaliandra and White-sloanea, are basal to this southern African clade. The other clades comprise the more basal genera of stem-succulent stapeliads, including the members of the Caralluma complex. Of the 17 genera accepted by Plowes for the Caralluma complex, seven are recognized: Caralluma, Apteranthes, Australluma, Boucerosia, Caudanthera, Desmidorchis and Monolluma. New combinations are proposed in 15 cases; Caralluma adscendens var. geniculata is raised to specific rank. Anomalluma is reinstated, and Pseudolithos mccoyi is transfered to it. A broadened concept for Orbea (incl. Angolluma and Orbeopsis) is recognized, but Orbeanthus is kept separate. The monotypic Ballyanthus, recently separated from Orbea, is nested within Duvalia. Piaranthus (incl. Huerniopsis) is monophyletic. The bitypic Notechidnopsis is reduced to the type species, N. tessellata, while N. columnaris is transferred to a new genus, Richtersveldia. Received February 25, 2002; accepted June 17, 2002 Published online: November 7, 2002 Address of the authors: Dr. Ulrich Meve (e-mail: ulrich.meve@uni-bayreuth.de) and Prof. Dr. Sigrid Liede (e-mail: sigrid.liede@uni-bayreuth.de), Universit?t Bayreuth, Lehrstuhl für Pflanzensystematik, Universit?tsstrasse 30, D-95440 Bayreuth, Germany.     

Monophyly and relationships of the tribe Exaceae (Gentianaceae) inferred from nuclear ribosomal and chloroplast DNA sequences

Both chloroplast trnL (UAA) intron and nuclear ribosomal ITS sequences highly confirmed the monophyly of the tribes of the Gentianaceae defined by the recent classification, and revealed the tribe Exaceae as a basal clade just next to the basal-most lineage, the tribe Saccifolieae. Within the tribe Exaceae, Sebaea (except Sebaea madagascariensis) appeared as the most basal clade as the sister group to the rest of the tribe. The Madagascan endemic genera Gentianothamnus and Tachiadenus were very closely related to each other, together standing as sister to a clade comprising Sebaea madagascariensis, Ornichia, and Exacum. The saprophytic genus Cotylanthera nested deeply inside Exacum. Sebaea madagascariensis was shown closer to the Madagascan endemic genus Ornichia than to any other sampled Sebaea species. Exacum appeared as the most derived taxon within this tribe. The topology of the phylogenetic trees conform with the Gondwana vicariance hypothesis regarding the biogeography of Exaceae. However, no evidence for matching the older relationships within the family to the tectonic history could be corroborated with various divergence time analyses. Divergence dating estimated a post-Gondwana diverging of the Gentianaceae about 50 million years ago (MYA), and the tribe Exaceae as about 40 MYA. The Mozambique Channel land-bridge could have played an important role in the biogeographic history of the tribe Exaceae.     

Two new species of <Emphasis Type="Italic">Miconia</Emphasis> sect. <Emphasis Type="Italic">Sagraea</Emphasis> (Melastomataceae) from the Macaya Biosphere Reserve,Haiti, and twelve relevant new species combinations

The Sagraea clade (Melastomataceae, tribe Miconieae) is briefly characterized, typified, and formally treated as a section within Miconia. In addition, two new species of Miconia sect. Sagraea, endemic to the floristically diverse Massif de la Hotte of southwestern Haiti and discovered during the course of a systematic revision of the Caribbean species of this section, are here described and illustrated. Miconia hottensis and M. navifolia, morphologically similar and possible sister species, are compared to each other and to the widespread Caribbean species M. capillaris and the southwestern Dominican Republic endemic M. tetraptera; these four species share rectangular stems with four low ridges or wings and minute, short-stalked, peltate or pseudopeltate hairs and likely form a clade.     

Phylogenetic relationships in Blumea (Asteraceae: Inuleae) as evidenced by molecular and morphological data

The present study, based on sequences of cpDNA (trnL-F & psbA-trnH) and nrDNA (ITS) and morphology, examined the evolutionary relationships in Blumea and its position among related genera. The results confirmed that the closest relatives of Blumea are Caesulia, Duhaldea and Pentanema p.p., and showed that the monotypic genera Blumeopsis and Merrittia are nested within Blumea. In Blumea s.l., two major, well-supported clades were recognised and a single species, the widespread Blumea balsamifera, that could not be placed with certainty relative to the two main clades. The two main clades differ in habit, ecology and distribution. The Blumea densiflora clade contains shrubs and subshrubs of evergreen forests, distributed from continental Asia to New Guinea and Polynesia, whereas the Blumea lacera clade is a widespread paleotropical group that comprises mostly annual, weedy herbs of open forests and fields.     

Phylogeny ofRubiaceae-Rubieae inferred from the sequence of a cpDNA intergene region

A phylogenetic analysis of 25 species, representing eight genera of theRubieae tribe (Rubiaceae), has been made using the DNA sequence of the chloroplastatp B-rbc L intergene region. Six tropical genera from other tribes ofRubiaceae have been used as outgroups. Whatever the method of analysis (distance, parsimony or maximum likelihood), five groups are clearly separated and described as informal clades. Their relative relationships are not clearly resolved by the parsimony analysis, resulting in eight equally parsimonious trees, 327 steps long, with a consistency index (CI) of 0.749 (excluding uninformative sites). TheRubieae tribe appears monophyletic from the data available. Some new and partly unexpected phylogenetic relationships are suggested. The genusRubia forms a separate clade and appears to be the relatively advanced sister group of the remaining taxa. TheSherardia clade also includes the generaCrucianella andPhuopsis. Galium sect.Aparinoides appears closely attached to theAsperula sect.Glabella clade. The remaining taxa ofGalium are paraphyletic:Galium sect.Platygalium (in theCruciata clade) is linked to the advanced generaCruciata andValantia; the more apomorphic groups ofGalium form theGalium sect.Galium clade, including the perennial sectionsGalium, Leiogalium, andLeptogalium as well as the annual (and possibly polyphyletic) sect.Kolgyda.     

A phylogeny of Cariniana (Lecythidaceae) based on morphological and anatomical data

Cariniana as previously circumscribed is a genus of 16 species restricted to neotropical forest habitats on well-drained sites. A phylogenetic analysis of the genus based on 33 morphological and anatomical characters was undertaken. The results show that Cariniana consists of two clades: the Allantoma/Cariniana decandra clade includes Allantoma lineata and seven species of actinomorphic-flowered Cariniana and is characterized by 5-merous flowers, carnose petals, incurved petal apex, scarcely lobed calyces, eucamptodromous secondary veins, dichotomizing venation, and poorly developed areolation; the C. legalis clade is made up of nine species and is characterized by an obliquely zygomorphic androecium, reticulate tertiary venation, and anomocytic stomata. The actinomorphic-flowered Cariniana are more closely related to the monotypic Allantoma lineata than they are to the species of the C. legalis clade. In order to reflect these relationships, Cariniana is divided into two genera: species in the C. legalis clade, which includes the generic type C. legalis, remain as Cariniana while species of Cariniana in the Allantoma/Cariniana decandra clade are transferred to Allantoma. The following new combinations are proposed: Allantoma decandra, A. integrifolia, A. kuhlmannii, A. pluriflora (a nomen novum for Cariniana multiflora because Allantoma multiflora is a synonym of Couratari multiflora), A. pachyantha, A. pauciramosa, and A. uaupensis.