Morphological Synapomorphies Research Articles

Chapter 6. Phylogenetic Relationships of Harpyionycterine Megabats (Chiroptera: Pteropodidae)

Abstract After almost 70 years of stability following publication of Andersen's (1912) monograph on the group, the systematics of megachiropteran bats (Chiroptera: Pteropodidae) was thrown into flux with the advent of molecular phylogenetics in the 1980s—a state where it has remained ever since. One particularly problematic group has been the Austromalayan Harpyionycterinae, currently thought to include Dobsonia and Harpyionycteris, and probably also Aproteles. In this contribution we revisit the systematics of harpyionycterines. We examine historical hypotheses of relationships including the suggestion by O. Thomas (1896) that the rousettine Boneia bidens may be related to Harpyionycteris, and report the results of a series of phylogenetic analyses based on new as well as previously published sequence data from the genes RAG1, RAG2, vWF, c-mos, cytb, 12S, tVal, 16S, and ND2. Despite a striking lack of morphological synapomorphies, results of our combined analyses indicate that Boneia groups with Aprotele...

An investigation into the cladistic relationships and monophyly of therocephalian therapsids (Amniota: Synapsida)

A comprehensive phylogenetic investigation was performed to elucidate the cladistic relationships and possible monophyly of therocephalian therapsids (Amniota: Synapsida). The phylogenetic positions of 30 therapsid taxa were examined under maximum parsimony, including 23 therocephalian genera. The analysis incorporated 110 cranial and postcranial characters in order to assess the interrelationships of basal therocephalians and eutherocephalians and their relationships to Cynodontia, representing the most complete review of therocephalian phylogeny to date. The analysis supports the hypothesis that Therocephalia represents the monophyletic sister taxon to Cynodontia, with as many as 15 morphological synapomorphies, in contrast with other recent analyses of lesser taxon sampling. The results also support the hypothesis that Scylacosauridae is more closely related to Eutherocephalia than to the basal therocephalian family Lycosuchidae, supporting a ‘Scylacosauria’ clade. The taxa suggested here to be neotenic forms (e.g. Ictidosuchoides and Ictidosuchops) are positioned near the base of a monophyletic Baurioidea. Neotenic development of the therocephalian feeding apparatus and evolutionary parallelism with cynodonts are suggested to have been important trends in the early evolution of baurioid therocephalians into the Late Permian and Early Triassic.

Morphological evolution in the variable resin-producing Detarieae (Fabaceae): do morphological characters retain a phylogenetic signal?

Previous molecular phylogenetic studies disagree with the informal generic-level taxonomic groups based on morphology. In this study morphological characters in the caesalpinioid clade Detarieae are evaluated within a phylogenetic framework as a means of better understanding phylogenetic relationships and morphological evolution. Morphological characters were observed and scored for representative species of Detarieae focusing on the resin-producing genera. Phylogenetic analyses were carried out with morphological characters alone and then combined with DNA sequences. Despite a high level of homoplasy, morphological data support several clades corresponding to those recovered in molecular phylogenetic analyses. The more strongly supported clades are each defined by at least one morphological synapomorphy. Several characters (e.g. apetaly) previously used to define informal generic groups evolved several times independently, leading to the differences observed with the molecular phylogenetic analyses. Although floral evolution is complex in Detarieae some patterns are recovered. New informal taxonomic groupings are proposed based on the present findings. Floral evolution in the diverse Detarieae clade is characterized by a repeated tendency toward zygomorphy through the reduction of lateral petals and toward complete loss of petals.

Phylogeny of the bee subtribe Caenohalictina Michener (Hymenoptera, Apidae s.l., Halictinae s.l.)

Gonçalves, R. B. & Melo, G. A. R. (2009). Phylogeny of the bee subtribe Caenohalictina Michener (Hymenoptera, Apidae s.l., Halictinae s.l.). —Zoologica Scripta, 39, 187–197. Recently, Caenohalictina Michener, 1954 was formally proposed as a group within Halictini based on a phylogenetic analysis using DNA data. Morphological synapomorphies for the subtribe are not stabilized and the relationships among its attributed genera were not completely reconstructed. Also, there are different propositions for the scope and classification of the genera. This study investigates the monophyly of Caenohalictina based on morphological characters and presents a hypothesis for the relationships among its genera and subgenera. For this purpose, a matrix of 36 terminals and 96 characters was coded. The cladistic analysis using character equal weighting resulted in one most parsimonious tree with 189 steps. According to this hypothesis, the subtribe is monophyletic with five synapomorphies supporting it. The relationships among Caenohalictina taxa are: Habralictus(Caenohalictus (((Agapostemon s.s. + Notagapostemon) (Agapostemonoides (Rhinetula (Paragapostemon + Dinagapostemon)))) + ((Ruizanthedella (Ruizantheda + Oragapostemon)) + (Brasilagapostemon (Pseudagapostemon s.s. + Neagapostemon))))). The subgenus Habralictus s.s. is paraphyletic with respect to Habralictus (Zikaniella) crassipes, hence both subgenera are not recognized. Comments on alternative generic classifications and biogeographic patterns are presented.

Phylogenetic analysis of Rhabdepyris (Hymenoptera: Bethylidae) and redefinition of generic limits based on morphological characters

Rhabdepyris (Epyrinae) is a cosmopolitan genus comprised of 132 species. No morphological synapomorphies are known for the genus and the genus is characterized by a combination of characters common to most Epyrini. Herein, we performed a cladistic analysis based on morphological characters to test the monophyly of Rhabdepyris. The three known subgenera of Rhabdepyris (Chlorepyris, Rhabdepyris s. str., and Trichotepyris) and other Epyrini (Anisepyris, Bakeriella, Calyozina, Epyris, Laelius, Trachepyris) were included in the ingroup. The cladistic analysis of 48 taxa (46 ingroup species and two outgroup species) and 81 structural characters yielded 72 cladograms under equal weights, and one under successive weighting. Rhabdepyris was found to be polyphyletic; the subgenus Trichotepyris was closely related to Anisepyris whereas Rhabdepyris str. s. was closely related to Laelius. The subgenus Chlorepyris is paraphyletic. Morphological characters are discussed in the light of the new phylogeny; novel characters are proposed and illustrated, and a new classification of Rhabdepyris and Epyrini is proposed. The following nomenclatural changes are proposed: Trichotepyris is synonymized under Anisepyris (syn. n.); Chlorepyris is recognized as a separated genus (stat. rev.); all 12 American species of the subgenus Rhabdepyris are transferred to Laelius; 22 species of Trichotepyris are transferred to Anisepyris; 58 species are transferred to Chlorepyris. A remaining total of 40 species are now recognized in Rhabdepyris. The holotype of Rhabdepyris, R. myrmecophilus Kieffer, the type species of Rhabdepyris, is redescribed.

Phylogeny and evolution of morphological characters in tribe Chamaedoreeae (Arecaceae)

AbstractWithin the palm family, molecular analyses have resulted in a need to re­evaluate characters previously considered useful to establish hypotheses of relationships among groups. Recent phylogenetic analyses of tribe Chamaedoreeae have shown that characters traditionally used as strong indicators of relationship, such as presence of solitary flowers and dioecy, are homoplasious within this tribe. In the largest genus, Chamaedorea, published molecular analyses recover well­supported groups not previously proposed based on morphological characters and at the same time resolve some of the current subgenera as polyphyletic. In this study we further explore the phylogenetic relationships in Chamaedoreeae and search for morphological synapomorphies for the monophyletic groups recovered. Phylogenetic analyses of morphological, nuclear (PRK, RPB2), and plastid (matK, ndhF, trnD­trnT, rps16 intron, trnL­trnF) data are performed and morphological characters are subsequently optimized on the resulting topologies. Although most of the morphological characters included are highly homoplasious, the inclusion of morphological characters in the phylogenetic analyses improves the resolution within Chamaedoreeae, particularly among species of Chamaedorea. With the exception of Synechanthus and Wendlandiella, all the genera and subgenera in Chamaedoreeae are defined by a combination of homoplasious characters, none of which is unique within the tribe. Two of the seven subgenera of Chamaedorea included in our analyses, C. subg. Eleutheropetalum and subg. Stephanostachys, are supported as monophyletic with a number of morphological synapomorphies also used in the original descriptions of these subgenera. Some morphological characters not used in the traditional subgeneric classification of Chamaedorea, such as leaf sheath structure and pistillode, gynoecium and filament connation, though homoplasious, are resolved as synapomorphies and, therefore, potentially useful when defining subgenera within this genus.

Neogastropod phylogenetic relationships based on entire mitochondrial genomes

BackgroundThe Neogastropoda is a highly diversified group of predatory marine snails (Gastropoda: Caenogastropoda). Traditionally, its monophyly has been widely accepted based on several morphological synapomorphies mostly related with the digestive system. However, recent molecular phylogenetic studies challenged the monophyly of Neogastropoda due to the inclusion of representatives of other caenogastropod lineages (e.g. Littorinimorpha) within the group. Neogastropoda has been classified into up to six superfamilies including Buccinoidea, Muricoidea, Olivoidea, Pseudolivoidea, Conoidea, and Cancellarioidea. Phylogenetic relationships among neogastropod superfamilies remain unresolved.ResultsThe complete mitochondrial (mt) genomes of seven Neogastropoda (Bolinus brandaris, Cancellaria cancellata, Conus borgesi, Cymbium olla, Fusiturris similis, Nassarius reticulatus, and Terebra dimidiata) and of the tonnoidean Cymatium parthenopeum (Littorinimorpha), a putative sister group to Neogastropoda, were sequenced. In addition, the partial sequence of the mitochondrial genome of the calyptraeoidean Calyptraea chinensis (Littorinimorpha) was also determined. All sequenced neogastropod mt genomes shared a highly conserved gene order with only two instances of tRNA gene translocation. Phylogenetic relationships of Neogastropoda were inferred based on the 13 mt protein coding genes (both at the amino acid and nucleotide level) of all available caenogastropod mitochondrial genomes. Maximum likelihood (ML) and Bayesian inference (BI) phylogenetic analyses failed to recover the monophyly of Neogastropoda due to the inclusion of the tonnoidean Cymatium parthenopeum within the group. At the superfamily level, all phylogenetic analyses questioned the taxonomic validity of Muricoidea, whereas the monophyly of Conoidea was supported by most phylogenetic analyses, albeit weakly. All analyzed families were recovered as monophyletic except Turridae due to the inclusion of Terebridae. Further phylogenetic analyses based on either a four mt gene data set including two additional Littorinimorpha or combining mt and nuclear sequence data also rejected the monophyly of Neogastropoda but rendered rather unresolved topologies. The phylogenetic performance of each mt gene was evaluated under ML. The total number of resolved internal branches of the reference (whole-mt genome) topology was not recovered in any of the individual gene phylogenetic analysis. The cox2 gene recovered the highest number of congruent internal branches with the reference topology, whereas the combined tRNA genes, cox1, and atp8 showed the lowest phylogenetic performance.ConclusionPhylogenetic analyses based on complete mt genome data resolved a higher number of internal branches of the caenogastropod tree than individual mt genes. All performed phylogenetic analyses agreed in rejecting the monophyly of the Neogastropoda due to the inclusion of Littorinimorpha lineages within the group. This result challenges morphological evidence, and prompts for further re-evaluation of neogastropod morphological synapomorphies. The important increase in number of analyzed positions with respect to previous studies was not enough to achieve conclusive results regarding phylogenetic relationships within Neogastropoda. In this regard, sequencing of complete mtDNAs from all closely related caenogastropod lineages is needed. Nevertheless, the rapid radiation at the origin of Neogastropoda may not allow full resolution of this phylogeny based only on mt data, and in parallel more nuclear sequence data will also need to be incorporated into the phylogenetic analyses.

Floral development and anatomy of Salvadoraceae

BACKGROUND AND AIMS; This study is an investigation into the floral development and anatomy of two genera of the small family Salvadoraceae, which belongs to the Brassicales in a clade with Batis and Koeberlinia. Salvadoraceae remains little known, despite its wide distribution in arid areas of the globe. Floral morphological data are scarce, and information on floral anatomy is limited to a single study, although morphological and anatomical characters are now used increasingly as a counterpart of molecular data. There remain a number of controversial morphological questions, such as the fusion of the petals, the number of carpels and the nature of the nectaries. Floral anatomy and ontogeny were studied in two species of Salvadora and one species of Dobera. Only for S. persica could a full floral developmental sequence be done. The floral development demonstrates that the ovary of Salvadoraceae is basically bicarpellate and pseudomonomerous with a single locule and parietal placenta. The ovary of Dobera resembles Azima tetracantha in the presence of a false apical septum. Evidence for a staminodial nature of the nectaries is not decisive. In Salvadora petals and stamens are lifted by a short hypanthium. Salvadoraceae share several morphological and developmental synapomorphies with Batis (Bataceae) and possibly Koeberlinia (Koeberliniaceae), supporting their close relationship as indicated by molecular phylogeny.

Revised classification of the family Hesperiidae (Lepidoptera: Hesperioidea) based on combined molecular and morphological data

Abstract We propose a revised higher classification for the genera of Hesperiidae (skipper butterflies) of the world. We have augmented our published DNA data matrix with 49 morphological characters in order to infer relationships for taxa not sampled in the molecular study. We use the results of a combined analysis to identify morphological synapomorphies of the suprageneric clades of Hesperiidae, and to hypothesize a phylogenetic classification of the world’s genera of Hesperiidae, the first of its kind for this diverse group. Monophyly of the family Hesperiidae is strongly supported, as are some of the traditionally recognized subfamilies. The results presented here largely corroborate those of our molecular study, but differ in several details. The Australian endemic Euschemon rafflesia is given subfamily status, as is Eudaminae. We recognize seven subfamilies of Hesperiidae: Coeliadinae, Euschemoninae (confirmed status), Eudaminae (new status), Pyrginae, Heteropterinae (confirmed status), Trapezitinae and Hesperiinae. We treat Pyrrhopygini, Tagiadini, Celaenorrhinini, Carcharodini, Achlyodidini, Erynnini and Pyrgini as tribes of Pyrginae. Circumscriptions of Achlyodidini and Pyrgini require further elucidation. Tribes of Hesperiinae include Aeromachini, Baorini, Taractrocerini, Thymelicini, Calpodini (reinstated status), Anthoptini (new tribe), Moncini and Hesperiini. The tribal placement of many Old World hesperiine genera remains ambiguous.

A new species of casque-headed tree frog, genus Aparasphenodon Miranda-Ribeiro (Amphibia: Anura: Hylidae), from the Atlantic Rainforest of southern Bahia, Brazil

A new species of Aparasphenodon is described from patches of arboreal restinga within the Atlantic Forest Biome, in a region known as Baixo Sul in southern Bahia, northeastern Brazil. Aparasphenodon arapapa sp. nov. is promptly diagnosed from other Aparasphenodon mainly by having small size (male snout-vent length 57.4–58.1 mm), loreal region flattened and wide, and canthus rostralis rounded and poorly elevated. The wide and flattened snout resembles that found in Triprion and Diaglena, and possibly is a parallelism (homoplasy) related to the phragmotic behavior of casque-headed tree frogs to their microhabitat usage. The decision to allocate the new species in the genus Aparasphenodon is discussed in detail, as the single morphological synapomorphy of the genus, the presence of a prenasal bone, is insufficient to morphologically relate the new species to Aparasphenodon, Triprion, or Diaglena.

Historical biogeography and character evolution of Cistaceae (Malvales) based on analysis of plastid rbcL and trnL-trnF sequences

Cistaceae consist of eight genera and about 180 species. Some taxonomic limits and relationships within the family remain unresolved when relying exclusively on morphological data. In the present study, a phylogeny was reconstructed and divergence times were estimated for 47 species representing various groups in Cistaceae and using coding ( rbcL) and spacer ( trnL-trnF) sequences of plastid DNA. The firm set of morphological synapomorphies that indicates the monophyly of the family is supported by both Bayesian and parsimony analyses. Five major lineages can be distinguished within the Cistaceae: (1) an early-diverging lineage containing Fumana species; (2) the New World Lechea clade; (3) the Helianthemum s.l. clade, containing two sister groups, one of species from the New World ( Crocanthemum, Hudsonia) and the other with species from the Old World ( Helianthemum s. str.); (4) the Tuberaria clade; and (5) a cohesive complex consisting of Halimium and Cistus species. Evolutionary shifts in 12 key characters of Cistaceae are inferred based on the most plausible phylogenetic hypothesis. Reconstructing the evolution of ovule position supports anatropous ovules as the ancestral condition within the Cistaceae, which is currently found only in Fumana. The Cistus-Halimium assemblage is consistently obtained as a natural clade and further supported by a cytological synapomorphy (chromosome number n=9). Optimisation of ancestral distribution areas and estimates of divergence times reveal an early divergence (10.17–18.51 Ma) of the Mediterranean-European genera, which may be related to subtropical vegetation, as complemented by paleobotanical data. In addition, the occurrence of multiple, independent migration events from the Old World to America between the Middle Miocene (8.44–14.7 Ma; Lechea) and the Upper Miocene (5.15–9.20 Ma; Crocanthemum/Hudsonia), and to the Canary Islands in the Pleistocene is inferred. We argue that the Mediterranean basin has been the main centre of differentiation of Cistaceae.

Phylogenetic relationships of Piptochaetium Presl (Poaceae: Stipeae) and related genera reconstructed from nuclear and chloroplast sequence datasets

ABSTRACTPiptochaetium (Poaceae, Pooideae, Stipeae) is a New World genus of grasses distributed primarily in temperate South America, with several species extending into Central and southern North America. The genus comprises 36 species and 2 varieties. Morphological synapomorphies for the genus include involute lemma margins that fit into the palea groove, a bi‐keeled palea longer than the lemma and protruding from the lemma apex, and fundamental cell sidewalls that are regularly dentate. The closely related genera Nassella (116 species) and Jarava (59 species) are also primarily South American, with a few species found in Central and North America. Published molecular studies to date have focused upon generic‐level relationships within the tribe Stipeae, but none have included more than a few representatives of each genus. We present here results of phylogenetic analyses of these genera based upon combined DNA sequences of both nuclear (ITS) and cpDNA (rpl16, trnL‐F) markers. Piptochaetium is supported as monophyletic in all studies, whereas Nassella is recovered as monophyletic only if Jarava plumosa is included. Infrageneric relationships of Piptochaetium are also compared to a recent morphological phylogeny of the genus.

Molecular phylogenetic interrelationships of the south Asian cyprinid genera Danio, Devario and Microrasbora (Teleostei, Cyprinidae, Danioninae)

Molecular analysis of mitochondrial cytochrome b sequences from 159 species of the family Cyprinidae supports the subfamily Danioninae, of which Rasborinae is shown to be a junior synonym. Analysis of combined cytochrome b and a fragment of the nuclear rhodopsin gene from 68 species, including 43 species representing the subfamily Danioninae, supports phylogenetic distinctness of Danio and Devario. In the combined molecular analysis Microrasbora rubescens, Chela, Laubuca, Devario, and Inlecypris form a clade with M. gatesi, M. nana and M. kubotai being in sister group position to the rest. The sister group of this Devario clade is Danio. Inlecypris is synonymized with Devario. Microdevario, new genus, is proposed for M. gatesi, M. nana and M. kubotai, supported by morphological characters. In the cytochrome b analysis, M. rubescens falls outside Devario, and there is no morphological support for including M. rubescens in Devario. In the cytochrome b analysis Esomus + Danionella is the sister group of Danio and Devario clades, whereas in individual rhodopsin and combined analyses Esomus is the sister group of Danio, and of Danio and the Devario clade, respectively. Sundadanio presents at least one strong morphological synapomorphy with Danio, but is positioned in molecular trees either as a member of the Cyprininae or as sister group of the remaining Danioninae. In the morphological analysis, small‐sized species grouped together based on shared reductions that are not necessarily synapomorphies. In the molecular analysis, small‐sized species such as Danionella and Sundadanio possess long branches and their position varies, but they did not group together. This suggests morphological homoplasy, but phylogenetic positions are not well supported in the molecular analyses

Species diversity and distributions of microdrile earthworms (Annelida, Clitellata, Enchytraeidae) from South America

A comprehensive biodiversity database of Enchytraeidae produced 66 nominal species reported to date from South America (Achaeta, 6 species; Buchholzia, 1; Cognettia, 1; Enchytraeus, 3; Fridericia, 5; Grania, 1; Guaranidrilus, 10; Hemienchytraeus, 12; Henlea, 2; Lumbricillus, 7; Marionina, 12; Stephensoniella, 1; Timmodrilus, 1; Tupidrilus, 4). Almost 76% of this fauna (50 species) is endemic to the South American continent. The remaining 16 species are more or less largely distributed. Detailed South American occurrences are provided. This is the first reassessment of South American enchytraeids in 27 years. The group is conspicuoulsy absent from the northeast region of Brazil. Enchytraeidae are microdrile earthworms, representing the sister group of the Crassiclitellata, or megadrile earthworms. This relationship is supported by molecular phylogenies, ontogenetic transformations, ecological evidence, and several morphological synapomorphies (dorsal pores, contractil dorsal vessel, lateral position of dorsal chaetal bundles, and shape of chaetae).

Phylogeny of the tribe Indigofereae (Leguminosae–Papilionoideae): Geographically structured more in succulent‐rich and temperate settings than in grass‐rich environments

This analysis goes beyond many phylogenies in exploring how phylogenetic structure imposed by morphology, ecology, and geography reveals useful evolutionary data. A comprehensive range of such diversity is evaluated within tribe Indigofereae and outgroups from sister tribes. A combined data set of 321 taxa (over one-third of the tribe) by 80 morphological characters, 833 aligned nuclear ribosomal ITS/5.8S sites, and an indel data set of 33 characters was subjected to parsimony analysis. Notable results include the Madagascan dry forest Disynstemon resolved as sister to tribe Indigofereae, and all species of the large genus Indigofera comprise just four main clades, each diagnosable by morphological synapomorphies and ecological and geographical predilections. These results suggest niche conservation (ecology) and dispersal limitation (geography) are important processes rendering signature shapes to the Indigofereae phylogeny in different biomes. Clades confined to temperate and succulent-rich biomes are more dispersal limited and have more geographical phylogenetic structure than those inhabiting tropical grass-rich vegetation. The African arid corridor, particularly the Namib center of endemism, harbors many of the oldest Indigofera lineages. A rates analysis of nucleotide substitutions confirms that the ages of the oldest crown clades are mostly younger than 16 Ma, implicating dispersal in explaining the worldwide distribution of the tribe.

Phylogenetic relationships of the spider family Tetragnathidae (Araneae, Araneoidea) based on morphological and DNA sequence data.

The monophyly of Tetragnathidae including the species composition of the family (e.g., Are Nephila and their relatives part of this lineage?), the phylogenetic relationships of its various lineages, and the exact placement of Tetragnathidae within Araneoidea have been three recalcitrant problems in spider systematics. Most studies on tetragnathid phylogeny have focused on morphological and behavioral data, but little molecular work has been published to date. To address these issues we combine previous morphological and behavioral data with novel molecular data including nuclear ribosomal RNA genes 18S and 28S, mitochondrial ribosomal RNA genes 12S and 16S and protein-coding genes from the mitochondrion [cytochrome c oxidase subunit I (COI)] and from the nucleus (histone H3), totaling ca. 6.3 kb of sequence data per taxon. These data were analyzed using direct optimization and static homology using both parsimony and Bayesian methods. Our results indicate monophyly of Tetragnathidae, Tetragnathinae, Leucauginae, the "Nanometa clade" and the subfamily Metainae, which, with the exception of the later subfamily, received high nodal support. Morphological synapomorphies that support these clades are also discussed. The position of tetragnathids with respect to the rest of the araneoid spiders remains largely unresolved but tetragnathids and nephilids were never recovered as sister taxa. The combined dataset suggests that Nephilidae is sister to Araneidae; furthermore, the sister group of Nephila is the clade composed by Herennia plus Nephilengys and this pattern has clear implications for understanding the comparative biology of the group. Tetragnathidae is most likely sister to some members of the "reduced piriform clade" and nephilids constitute the most-basal lineage of araneids.

Phylogeny and classification of the species‐rich pantropical showy genus Ixora (Rubiaceae‐Ixoreae) with indications of geographical monophyletic units and hybrids

Species-rich genera often have various conflicting circumscriptions from independent regional flora treatments. Testing the monophyly of these groups of plants is an important step toward the establishment of a phylogenetic classification. The genus Ixora of the tribe Ixoreae in the subfamily Ixoroideae (coffee family or Rubiaceae) is a species-rich pantropical genus of ca. 500 species. Phylogenetic analyses of Ixoreae based on combined sequence data from one nuclear (nrETS) and two chloroplast (rps16 and trnT-F) markers reveal the paraphyly of Ixora as presently delimited and also show that the tribe can be subdivided into three major clades: the Mascarene/neotropical/Malagasy/African clade, the Pacific clade, and the Asian clade. Given the lack of morphological synapomorphies supporting the different Ixora clades and the morphological consistency of the ingroup taxa, we propose a broad circumscription of Ixora including all its satellite genera: Captaincookia, Doricera, Hitoa, Myonima, Sideroxyloides, Thouarsiora, and Versteegia. The current infrageneric classification of Ixora is not supported. The different Ixora subclades represent geographical units. Nuclear and chloroplast tree topologies were partially incongruent, indicating at least four potential natural hybridization events. Other conflicting positions for the cultivated species are most likely due to anthropogenic hybridization.

TrnL-F and rpl16 Sequence Data and Dense Taxon Sampling Reveal Monophyly of Unilocular Anthered Gomphrenoideae (Amaranthaceae) and an Improved Picture of Their Internal Relationships

The Amaranthaceae-Chenopodiaceae alliance has been the focus of several phylogenetic studies, but major questions concerning the internal relationships of Amaranthaceae still remain unanswered. This study aims to test the monophyly of the subfamily Gomphrenoideae and to examine the generic relationships within this group. Parsimony and Bayesian analyses of trnL-F and rpl16 show that the subfamily Gomphrenoideae is monophyletic. The clade of Gomphrenoideae represents a large and diverse radiation of the Amaranthaceae in the New World. Unilocular anthers are a morphological synapomorphy for Gomphrenoideae that was derived from bilocular anthers. Three major clades are resolved: the Gomphrenoids, the Alternantheroids, and the Iresinoids, which are largely supported by pollen morphology. The Iresinoid clade is sister to Alternantheroids plus Gomphrenoids, rendering metareticulate pollen as the synapomorphy for the latter two clades. Tribes and subtribes delimited by androecium and inflorescence characters are poly- or paraphyletic. Several genera are monophyletic including the large genus Alternanthera, whereas Gomphrena is polyphyletic. Irenella and Woehleria are resolved within Iresine whereas Blutaparon and Lithophila fall within the polyphyletic Gomphrena. The trnL-F and rpl16 sequence data are the most variable chloroplast regions examined to date for the family and are highly effective in resolving relationships in Amaranthaceae.

Molecular phylogenetic and morphological evidence for the congeneric status of Goniothalamus and Richella (Annonaceae)

AbstractMaximum parsimony, maximum likelihood and Bayesian phylogenetic analyses of selected Goniothalamus and Richella species (Annonaceae) are presented, based on sequences of three chloroplast markers (matK, psbA‐trnH intergenic spacer, and trnL‐trnF intergenic spacer). The two genera are shown to be congeneric, and the nomenclatural consequences of this are discussed. Morphological synapomorphies for individual clades are evaluated, although a detailed interpretation of morphological evolution will require more extensive species sampling.

Generic classification of the Verrucariaceae (Ascomycota) based on molecular and morphological evidence: recent progress and remaining challenges

AbstractRecent molecular phylogenetic analyses and morphological studies have shown that it is necessary to revise the present morphology‐based generic delineation of the lichen family Verrucariaceae in order to account for evolutionary relatedness between species. Consequently, several genera were recently described or resurrected, and others were re‐circumscribed. As an additional step toward this generic revision, three new genera (Hydropunctaria, Parabagliettoa, Wahlenbergiella) and eleven new combinations are proposed here. A summary of the current taxonomic and morphological circumscription of all genera investigated so far is also presented. Several monophyletic groups are identified for which further taxonomical changes will be required, but for which taxon and gene sampling is presently viewed as insufficient. Clear morphological synapomorphies were found to be rare for newly delimited genera. In some cases (reduced morphology or plesiomorphism), even the combinations of slightly homoplasious phenotypic characters do not allow a clear morphological generic circumscription. Molecular features are envisioned as characters for delimiting these taxa.