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The classification of the family Compositae (Asteraceae) has been much improved in the last decades by the application of molecular methods culminating in the recompilation published in 2009, Systematics, evolution, and biogeography of Compositae. Additional evidence of relationships has come from the use of high‐throughput sequencing methods. Our late colleague Vicki Ann Funk (1947–2019) was a pioneer in this line of research. Together with her team, she contributed to the achievement of a mature classification of the family, which she left outlined. In this paper, we contribute this classification including all of the recent advances at the subtribal level and review in depth all contributions to Compositae classification made since the 2009 compilation. [ABSTRACT FROM AUTHOR]

This paper accounts for all names published in Argyreia Lour., Lettsomia Roxb., and Moorcroftia Choisy, reconciles the taxonomy and nomenclature for those names based on study of type specimens for nearly all of them. In total, 135 species and 5 varieties are accepted; a significant increase in the species-level diversity relative to previously published estimates in the literature. Concise summaries of the infrageneric relationships, distribution and biogeography, taxonomy, and nomenclatural complexities are provided; relevant literature is cited throughout. Excluded species and uncertain species are treated in separate sections. One new combination is made at varietal rank and one invalid name in current use is here validated. A comprehensive index is provided to all published names, cross-referenced to the accepted name. [ABSTRACT FROM AUTHOR]

The Helichrysum-Anaphalis-Pseudognaphalium (HAP) clade is a major component of the tribe Gnaphalieae (Compositae) and includes the genera Helichrysum, Anaphalis, Achyrocline and Pseudognaphalium. Allopolyploid origins for at least two clades within the HAP clade have previously been suggested, one involving the genus Anaphalis and the Mediterranean-Asian Helichrysum species, and a second one involving part of Pseudognaphalium. In the present paper, with the use of two nuclear ribosomal and two plastid DNA markers and an extensive sampling of the HAP clade, further evidence relevant to the origin, composition and closest relatives of these clades is provided, and additional cases of incongruence are discussed. The superposition of distribution areas on the phylogeny suggests that the HAP clade originated in the Cape region of southern Africa and subsequently dispersed to and diversified in the Afromontane regions of east southern Africa, mainly the Drakensberg, before spreading northward and giving rise to several lineages in Afromontane and Afroalpine areas of central tropical Africa and in Madagascar. Allopolyploidy may have preceded the dispersal and diversification of the HAP lineage out of Africa to the Mediterranean area, and to the Americas and Asia. Finally, discussion on the distribution of several morphological characters in the phylogeny and their taxonomic relevance is also provided, with views on the need for a new generic delimitation. [ABSTRACT FROM AUTHOR]

The Neotropical region (tropical America) is the most species rich region on Earth. Several causes have been proposed to explain this extraordinary biodiversity, which may be very roughly classified into two major categories: 'biotic' (e.g. soil adaptations; biotic interactions with pollinators, dispersers and herbivores; niche conservatism; dispersal ability) and 'abiotic' (e.g. time; rainfall, temperature and area; mountain uplift; hydrological changes). In this paper we review the evidence for each of these postulated causes of diversification and provide general directions towards further testing. We highlight the need of more well-sampled and dated phylogenies and urge increased inter-disciplinary collaboration. [ABSTRACT FROM AUTHOR]

In this paper we investigate phylogenetic relationships for Caralluma R. Br. (Apocynaceae-Asclepiadoideae-Cerop- egieae), the largest genus among the stapeliads, using analyses of data from four molecular markers: two plastid regions psbA- trnH and irnT-F and two nuclear regions ITS and ncpGS. Our analyses show that the stapeliads consist of two lineages, with Caralluma dominating one of these lineages. Within this lineage, Caralluma is not monophyletic, with its species distributed among two major clades. One of these clades contains species of Carallurna and representatives of Anomallurna Plowes, Echid- nopsis Hook. f. and Rhytidocaulon PRO. Bally, while in the other dade the small genera Edithcolea N.E. Br., Frerea DaIz. and Pseudolithos PRO. Bally are embedded among species of Caralluma. We discuss morphological features that characterize some of the clades. Caralluma is mainly paleotropical in its distribution and is the most widely distributed group among the stapeliads, from the Canary Islands in Macaronesia to the Indian subcontinent. We show that the Indian subcontinent has been invaded four times by species of Caralluma, while its greatest diversity is found in North-East Africa and southern Arabia. [ABSTRACT FROM AUTHOR]

A striking feature of the Mediterranean flora concerns the high rates of narrow endemism within many regions. The prevailing paradigm for the evolution of such endemism relies on an important role of geographic isolation and allopatric speciation. Until recently there has been little work on the precise ecological differences among endemic plants and their congeners in the Mediterranean region and the potential role of ecological specialisation in isolation and speciation. This paper has two objectives. First, we evaluate general ecological differences among endemic and widespread congeners in the flora of the western Mediterranean. The results and review of the literature attest to consistent patterns of ecological differentiation among endemic species and their more widespread congeners. Second, we present results of a detailed study of morphological, molecular and ecological differentiation in a group of spring-flowering Mediterranean Cyclamen species. This study illustrates how differentiation in geographically peripheral and ecologically marginal populations may contribute to the diversification of endemic species whose contemporary distributions are disjunct from con-generic species. [ABSTRACT FROM AUTHOR]

The Lactucinae or Lactuca alliance include approximately 200 species distributed across the Northern Hemisphere and Africa. They were not recognised as a separate lineage until the late 20th century and their circumscription is still not fully settled. The generic classification of no other group of the Cichorieae has faced as many controversies as the Lactuca alliance and competing taxonomies coexist. This paper provides the first molecular phylogeny of the subtribe on a global scale as a major step towards its revised systematics. The sampling includes almost 60% of the species-level diversity and spans all species groups. Two datasets were created, one including the nrDNA ITS region, the other five concatenated non-coding plastid DNA loci. Maximum parsimony, maximum likelihood and Bayesian inference were used to produce a robust phylogenetic backbone. The diversification and expansion of the Lactucinae in a geohistorical context was reconstructed by estimating the age of their lineages using relaxed molecular clock dating and by inferring the ancestral areas using Bayesian binary analysis. The redelimited monophyletic Lactucinae are composed of seven lineages that also include Prenanthes, which is confirmed to have a single species, P. purpurea. The positions of two further lineages shift between Lactucinae and Crepidinae in the nuclear and plastid DNA phylogenies. Incongruence between the phylogenies suggests events of ancient reticulation or incomplete lineage sorting in the formation of these latter two lineages and in two of the seven other Lactucinae lineages. The phylogenetic results show a dilemma for Lactucinae systematics: most generic concepts proposed to date are highly artificial but the resolved phylogenetic lineages do not constitute practicable taxonomic entities with our current knowledge. Diversification of the subtribe is inferred to have taken place since the Middle Miocene. Biogeographic analysis proposes that the clade originated in the mountains of the landmass mediating between the European and Asian continents and delimited in the south by the Tethys Sea and in the north by the Paratethys Sea. Several independent migrations have occurred into various parts of Asia, Europe, tropical Africa and North America. [ABSTRACT FROM AUTHOR]

Grusonia is an opuntioid genus in the Cylindropuntieae, Cactaceae, endemic to the North American deserts in Mexico and the United States. The monophyletic status of the genus and the degrees of endemicity of its species make Grusonia a suitable study group in order to test biogeographic hypotheses such as vicariant or dispersal events, biogeographic barriers and time‐calibrated phylogenies in order to propose speciation events in the group. The statistical dispersal‐vicariance analysis (S‐DIVA) and a statistical dispersal‐local extinction‐cladogenesis (S‐DEC) analyses show general congruence postulating ancestral ranges for the species of Grusonia. They disagree, however, in a clade with four species from the Sonoran Desert, the Baja California Peninsula and one species endemic to the central portion of the Chihuahuan Desert Region. The time‐calibrated phylogeny here proposed shows an average age of 3.7 Ma for the genus. Grusonia has two internal clades comprising species most likely generated by allopatric speciation under a vicariant scenario. The results of this study support a young age for the North American deserts and identify the end of the Pliocene and mainly the Plio‐Pleistocene boundary as the starting point of the cladogenetic events that gave rise to the two main clades of Grusonia. [ABSTRACT FROM AUTHOR]

A densely sampled molecular phylogeny of the pantropical, predominantly forest‐dwelling tribe Marsdenieae (Apocynaceae) is presented, including almost a quarter of the ca. 740 species. Sampling focused on resolving the circumscription of Marsdenia, which ranges from Marsdenia sensu Bullock with less than 10 Asian species closely related to the type, M. tinctoria, to Marsdenia sensu Forster, with ca. 300 species distributed throughout the range of the tribe, encompassing the genera Cionura, Dregea, Gongronema, Gymnema, Leichhardtia, Stephanotis, and Wattakaka as synonyms. All analyses resolve two well‐supported clades: an Asia‐Pacific Clade that includes the large epiphytic Dischidia‐Hoya alliance and a number of Marsdenia species, among them the type M. tinctoria, and a Cosmopolitan Clade that includes most of the rest of Marsdenieae with several moderately to well‐supported subclades that span the entire geographic range of Marsdenieae. The position of two small genera, Rhyssolobium (southern Africa) and Sicyocarpus (Madagascar), is ambiguous. Given this topology, a monophyletic Marsdenia s.l. would subsume all species of Marsdenieae. Instead, we propose a classification that maintains Dischidia and Hoya as distinct genera. We revise generic circumscriptions, reducing Marsdenia s.str. to a small clade of Asian species. Two new genera, Gongreos gen. nov. and Gongronemopsis gen. nov., are described, with two and nine species, respectively. A total of twenty‐six new combinations, one new name, and six new synonyms are proposed. Three neotypes and eleven lectotypes are selected, and one 2nd step lectotypification is carried out. Diagnostic morphological characters for identified clades are discussed. Biogeographic analysis retrieves tropical Asia as most likely ancestral area for Marsdenieae, with widespread tropical Asia‐Africa almost equally likely. The first‐branching lineages in both major clades are tropical Asian; the Asia‐Pacific Clade comprises a sizeable radiation in the Pacific area, with two dispersals from Australia to New Caledonia. The Cosmopolitan Clade comprises several genera (Gongronemopsis, Gymnema, Stephanotis, Telosma) widespread in Africa and tropical Asia, and some also in Madagascar, and the New World genus Ruehssia. [ABSTRACT FROM AUTHOR]

Carex subg. Uncinia (Cyperaceae) constitutes one of six currently recognized Carex subgenera. This subgenus is mainly distributed on the American continent and in the Pacific region, and it is the only subgenus almost entirely absent from the Old World and primarily diversified in the Southern Hemisphere. It includes some of the few Carex species with clear epizoochoric traits: the representatives of C. sect. Uncinia possess utricles with an exserted and hooked rachilla that allows the diaspores to attach to feather or hair. We performed phylogenetic (ITS, ETS‐1f, matK), biogeographic, and ancestral state reconstruction analyses to elucidate the systematic structure, origin and dispersal routes, and major morphological evolutionary patterns of the different lineages within the subgenus. Our phylogenetic reconstructions revealed that the subgenus comprises seven different clades that mostly match previously recognized sections. One of the clades, however, represents a new section described herein as C. sect. Wheelerianae. Unispicate lineages evolved repeatedly from ancestors bearing multispicate inflorescences, while the presence of a rachilla, often pictured as a plesiomorphy in Carex, seems to have developed four independent times in the evolution of C. subg. Uncinia. The origin of the subgenus dates back to the beginning of the Miocene, probably in North America from where it colonized the Southern Hemisphere. It first dispersed to South America during the Early Miocene. Later, in the Middle Miocene, representatives of C. sect. Uncinia would reach the Pacific Southwest region (New Zealand, Australasia) from South America in at least two independent dispersal events. The vast majority of the biogeographic events seem to be explained by long‐distance dispersal. The remarkable dispersal ability of C. sect. Uncinia enabled by the hooked rachilla has allowed it to reach remote archipelagos in the Pacific and Subantarctic regions, probably bird‐mediated. [ABSTRACT FROM AUTHOR]

The family Saxifragaceae, the current composition of which is one of the great surprises of molecular systematics, has been subject to massive improvements in the knowledge of phylogenetic relationships. Nevertheless, developments from phylogenomic efforts have yet to be mobilized to inform biogeography and taxonomy. Here, we use a recent order‐level phylogeny for Saxifragaceae and related families covering 72% of species with a set of new analyses to assess habitat evolution and biogeography. Our results suggest a North American origin of the family in cold alpine habitats, followed by rapid recent evolution of diverse habitat tolerances. We also combine these recent phylogenomic results and a synthesis of the literature to improve generic limits and tribal classification of Saxifragaceae. We recognize 40 genera in 10 tribes, with 14 new combinations, and elevate one subtribe as well as describing four new taxa at the tribal level. Finally, we synthesize information about biogeography and morphology for the family. [ABSTRACT FROM AUTHOR]

Seasonally Dry Tropical Forests (SDTFs) are found scattered throughout the Neotropics, from northwestern Mexico to northern Argentina, and southwestern and eastern Brazil, with eastern Brazil having the largest and most isolated fragment of SDTF, known as Caatinga. Growing knowledge from dated phylogenies and ancestral reconstruction studies suggests that SDTF lineages are nested in geographically structured phylogenies with clades highly restricted to each disjunct patch. To address these hypotheses, we investigated the biogeographic history of Euphorbia sect. Brasilienses, a group of succulent spurges from eastern Brazil's SDTF. To this aim, we assembled a concatenated matrix from 126 accessions with four markers (302 sequences from previous studies and 19 newly generated): one nuclear (ITS1) and three plastid (matK, ndhF, trnL‐trnF) loci. Our results showed that clade Brasilienses and its sister group Stachydium diverged from a common ancestor during the Miocene around 16.5 Ma. Clade Stachydium was recovered with a mean crown age of 7.7 Ma, while clade Brasilienses was recovered as a relatively young group that started diversifying about 3.1 Ma. Biogeographic results showed that the ancestral range of the clade Stachydium + Brasilienses comprised the Andes and eastern Brazil's SDTF, therefore suggesting past connections between western and eastern South America. The ancestral range of clade Brasilienses was restricted to eastern Brazil SDTF, which contrasts with clade Stachydium, which was recovered with a broader ancestral range around Late Miocene. Our results suggest limited dispersal abilities, niche conservatisms and an origin for E. sect. Brasilienses pre‐dating the Pleistocene. By contributing to the understanding of the origin and diversification of this group of endemic spurges, our study provides insights into the history of this SDTF in eastern South America. [ABSTRACT FROM AUTHOR]

The genus Dracaena in Macaronesia comprises two threatened species of arborescent monocots that are often associated with one of the most intriguing biogeographic disjunctions: the Rand Flora pattern. Molecular information is, however, largely missing for the Macaronesian Dracaena taxa ("MDT", hereafter), and the biogeographic or population genetic patterns of this lineage have not yet been thoroughly assessed. To fill this gap, we generated plastid DNA sequence data of 14 Dracaena populations representing the entire natural distribution of MDT (including mainland Morocco and all recognized subspecies), 9 additional populations of subspontaneous origin, and a set of related species of the genus. We performed phylogenetic, biogeographic, and population genetic analyses at different spatial scales and conducted a comparative review on plant haplotype diversity in Macaronesian plants. The results of our phylogenetic analyses indicated the monophyly of the MDT and an origin separate from a clade of geographically distant species that so far were postulated as their closest living relatives (D. cinnabari, D. ombet, D. schizantha, D. serrulata). The results of our phylogeographic analyses indicated that diversification within D. draco occurred throughout the Pleistocene and that wild peripheral populations (Madeira, mainland Morocco) may have a recent origin from Canarian source populations. Recent dispersals, coupled with remarkably low levels of haplotype diversity, probably account for the weak phylogeographic signal observed across wild populations. However, our results suggested that human‐assisted expansion of Dracaena inflates the extant phylogeographic signal by non‐random translocation of a specific subset of haplotypes. Our study demonstrates that many of the previous biogeographic scenarios on MDT are not supported by molecular data. Instead, our results highlight (i) the impact that human activity may have on the phylogeographic pattern of island plants, and (ii) the need of a deeper taxonomic sampling in future investigations on MDT and close relatives. [ABSTRACT FROM AUTHOR]

Schefflera is the largest and most complex genus of Araliaceae, with ~600 described species (and many additional species awaiting formal description), but recent studies indicate that it is polyphyletic, forming five geographically centered clades spread across the major lineages of the family. Significant progress has been made in revising the three smallest clades, but the two largest groups, centered in Asia and the Neotropics, remain poorly understood. To advance our knowledge of these groups, a taxon set of 211 Schefflera samples and 38 samples representing other genera of Araliaceae (including all genera of the "Asian Palmate clade") was assembled, from which sequences were obtained for six DNA spacer regions, including two nuclear (ITS, ETS) and four plastid markers (rpl32‐trnL, trnF‐rpl32, trnK‐rpl16, psbA‐trnH). Results were compared to an informal system of classification based on morphology. They confirm the monophyly of both the Asian and Neotropical groups, and their inclusion within the Asian Palmate clade, but not as sister groups. In the Neotropical clade, five major subclades can be recognized, four corresponding closely to morphological groupings (Cephalopanax, Crepinella, Didymopanax, Sciodaphyllum) and one that is geographically coherent (Gleasonia). Within Asian Schefflera, there are four major subclades, corresponding closely to currently recognized informal groups (Agalma, Heptaphyllum, Heptapleurum, Hypoleucoi). Biogeographic analyses suggest that both the Asian and Neotropical groups can trace their origins to SE Asia, where Asian Schefflera is still well represented and where Chengiopanax, the possible sister group of Neotropical Schefflera, is native. Trait‐evolution analyses of a remarkable series of convergences in habit, foliar, and reproductive characters suggest multiple independent origins (and some reversals), both within and between the Neotropical and Asian clades of Schefflera. [ABSTRACT FROM AUTHOR]

Korthalsella (Santalaceae, tribe Visceae) mistletoes are hemiparasitic plants that are widespread on islands and continental regions around the Indian and Pacific Oceans. In this study, we add key taxa to a previously generated dataset to produce a more inclusive phylogenetic analysis of the genus. The resulting nuclear ITS and plastid trnL‐F phylogenies reveal that the historical sectional classifications based on morphology are not supported. Instead, it appears that widespread morphological parallelism has occurred throughout Korthalsella. Geographical distribution seems to be a better indicator of phylogenetic relatedness as species found in the same geographic region, island or island group generally are more closely related to one another than species sharing similar morphological characters in other areas. We find greater support for recognition of species as local endemics rather than wide‐ranging taxa. Given these results, taxonomic changes that recognise previously described taxa are proposed, but other changes will require further study of broadly distributed taxa. [ABSTRACT FROM AUTHOR]

Traditional Tinospora is a heterogeneous genus, distributed in the African and Asian tropics as well as Australia and adjacent islands. Molecular phylogenetics has contributed greatly to improve our understanding of the genus over the past decade, however its circumscription is yet unresolved. In this study, we present a phylogenetic analysis of Burasaieae using six molecular markers (rbcL, atpB, matK, ndhF, trnL‐F, ITS), with a focus on Tinospora. Our results indicate that sampled species of Tinospora are distributed in three different clades. Tinospora tinosporoides, endemic to Australasia, is strongly supported as sister to the South American Borismene. By integrating lines of evidence from molecular phylogeny and morphology, we resurrected the monotypic Fawcettia to include F. tinosporoides. Our estimated age for the split of Fawcettia‐Borismene was ca. 29 Ma, suggesting that trans‐Pacific long‐distance dispersal may be the most feasible explanation for the Australasian–South American disjunction. Tinospora oblongifolia is a member of Hyalosepalum, characterized by three stamens with connate filaments, whereas T. tenera, once placed in Hyalosepalum, is recovered within Tinospora, characterized by six stamens. Evolutionary inferences of morphological characters indicate that the five selected characters, traditionally used to distinguish genera in Burasaieae, are highly homoplastic. Our molecular dating and ancestral range reconstruction suggest that Tinospora originated from Africa in the middle Oligocene (ca. 28 Ma) and expanded into Asia in the Early Miocene (ca. 21 Ma). Within Tinospora, a vicariance event occurred in the Middle Miocene (ca. 17 Ma) resulting in the split of African T. tenera and its Asian allies. One dispersal from Asia to Africa was inferred in the Late Miocene (ca. 10 Ma). Australian T. smilacina and T. esiangkara originated from Asia in the early Late Miocene (ca. 11 Ma). Thus, geodispersal and subsequent vicariance, as well as transoceanic long‐distance dispersal, have been responsible for the present distribution of Tinospora. [ABSTRACT FROM AUTHOR]

We estimated phylogenetic relationships of Polygala inferred from nrITS, matK‐trnK, trnL intron, trnL‐trnF intergenic spacer, and rbcL sequence data, with emphasis on the New World clade. Our results did not support the New World and Old World clades as sister groups in Polygala. Relationships among Polygala and allied genera are being discussed, with the recovered clades compared to those of the current taxonomic delimitations. A biogeographic analysis using an ultrametric phylogenetic tree and calibrated with the fossil Paleosecuridaca as well as a reconstruction of the ancestral area are also provided. [ABSTRACT FROM AUTHOR]

Whereas the eastern North American–eastern Asian floristic connection represents one of the most widely studied biogeographical relationships in flowering plant evolution, connections between western North America and Asia have been comparatively rarely investigated, especially through genetic approaches. Stellaria irrigua is one of several plants that has been treated as an exceptionally dramatic example of a disjunction between floristically similar, high alpine biotas of the southern Rocky Mountains and south‐central Siberia. We here employ numerous new field collections and ddRADseq data to test the hypothesis that S. irrigua—a species that has been known for over 180 years—represents a long‐distance disjunction between the southern Rocky Mountains and central Asia. Extensive fieldwork, review and perusal of herbarium materials, and phylogenomic analyses indicate that S. irrigua is broadly distributed across an amphi‐Beringian arc extending from southern and central Asia, east through Beringia, and south throughout mountainous regions of western North America. Sampled Asian populations formed two clades, and North American individuals all formed a clade embedded within this broader Asian lineage. Stellaria irrigua is, however, rendered non‐monophyletic by a lineage that is embedded within the North American populations and is ecologically and morphologically distinctive from S. irrigua. The identity of this newly recognized lineage, which was in prior works attributed to S. irrigua, has been confused since plants of the former were first collected in the San Juan Mountains of Colorado in the late 1800s under Arenaria and Alsine. We provide a new name for this taxon, Stellaria sanjuanensis, a charismatic starwort of dry alpine scree slopes of the southern Rocky Mountains. Additionally, two lectotypes are designated, one holotype and one isotype are identified, and two new synonymies are proposed, to help stabilize the taxonomy and nomenclature of this long‐confused species complex. A key to the starworts of the southern Rocky Mountains is also provided, and Stellaria alsine is reported as new to the region. [ABSTRACT FROM AUTHOR]

Plantago sect. Coronopus contains our two focal species (P. coronopus L., P. crassifolia Forssk.), both with an overall conspicuous bi‐hemispheric distribution range (Mediterranean and Middle Eastern regions in the Northern Hemisphere and South Africa in the Southern Hemisphere). We have evaluated up to 27 morphological characters from 96 herbarium specimens representing five out of seven species of that section that are currently recognised using principal coordinate analysis, linear discriminant analyses, agglomerative clustering, and classification tree analyses, in order to test the current taxonomic concepts of our two focal species. Furthermore, we used 54 individuals representing six out of those seven species of P. sect. Coronopus to construct molecular phylogenetic hypotheses by sequencing nuclear ribosomal DNA from the internal transcribed spacer region (ITS), the plastid trnL‐F region, the plastid intergenic spacer region trnH‐psbA and adopting maximum likelihood and Bayesian inference analyses. In the Northern Hemisphere, an Irano‐Arabian clade initially identified as P. coronopus and shown as distinct in both morphological and phylogenetical terms fits a wider circumscription of P. crypsoides Boiss. due to lack of the prominent short and thick inflorescence scape following Boissier's description. The morphological differences between P. crassifolia from the Mediterranean region and P. crassifolia from South Africa (often named P. carnosa Lam.) were marginal, yet the molecular phylogenetic analyses of both nuclear and plastid markers clearly separated these evolutionary entities. Therefore, we re‐instated the name P. carnosa as the correct name for South African P. crassifolia. Plantago carnosa differs from P. crassifolia by the combination of having stronger woody rootstocks, which are more often branched, by broader leaves (≥1.6 mm wide) and the fact that specimens more often turn brown when dried. Our dataset provides the best sampled phylogenetic hypothesis for P. sect. (and subg.) Coronopus to date, and reveals discordance between nuclear and plastid genealogies within P. sect. Maritima, which requires further investigation. [ABSTRACT FROM AUTHOR]

Eugenia, comprising ca. 1100 species, is the largest genus of Neotropical Myrtaceae. Eugenia sect. Umbellatae (formerly referred to as "clade 9") is the most speciose lineage of Eugenia. This study aims to better delimit E. sect. Umbellatae, to identify and understand relationships between manageable subgroups of this large clade for future discrete systematic studies and to explain biogeographical patterns in the genus. In total, 103 samples were used in this study. These include representatives of the nine clades of the "Eugenia group" with a particular focus on Eugenia clade 9, representing the morphological and geographical diversity found in the genus. Phylogenetic reconstructions were performed using maximum likelihood (ML) and Bayesian inference (BI) for the combined dataset, using the markers ITS, rpl16, psbA-trnH, rpl32-trnL, trnQ-rps16. The resultant tree was fossil calibrated and used for historical biogeographical analysis using DEC implemented in RASP. The mid Oligocene is the most likely period in which the crown node of Eugenia s.l. diversified. The earliest Eugenia appear to be associated with dry biomes and to have arisen from non-tropical southern South America, as did ancestors of the earliest American Myrteae. Eugenia subg. Pseudeugenia also most likely diversified in dry biomes, while E. subg. Hexachlamys and E. subg. Eugenia are likely to have diverged in the Atlantic Forests biome. Eugenia sect. Umbellatae is morphologically very variable,some clades can be circumscribed based on morphology while some remain morphologically undiagnosable. The study presented here provides discussion of the earliest origins of Eugenia and its response to climate-driven changes in the Neotropics as humid, forest biomes became more widespread in the Miocene. In addition, important practical conclusions are drawn regarding relationships within Eugenia. Three clades are newly classified as subgenera: E. subg. Pseudeugenia (including species of E. sect. Pseudeugenia),E. subg. Hexachlamys (including E. sect. Hexachlamys) and E. subg. Eugenia (including E. sect. Umbellatae, E. sect. Jossinia, E. sect. Phyllocalyx, E. sect. Pilothecium, E. sect. Racemosae, E. sect. Schizocalomyrtus, E. sect. Speciosae and Eugenia sect. Excelsae). Two previously unidentified clades are published as E. sect. Excelsae and recognized as E. sect. Jossinia, the latter consisting entirely of Old World species. [ABSTRACT FROM AUTHOR]

The taxonomic history of the Melanthera alliance is long and convoluted with many generic name changes and requires a robust phylogeny to clarify taxonomic concepts within the group and to begin asking questions of its evolutionary history. For a time, prevailing classifications placed all species in the genus Melanthera except for a handful of tetraploids from the Hawaiian Islands being recognized as a distinct genus: Lipochaeta. Recent morphological revision has reopened debate by proposing six genera: Apowollastonia, Echinocephalum, Lipotriche, and Melanthera, and two Pacific Island genera representing diploids (Wollastonia) and tetraploids (Lipochaeta), plus four closely related genera expected to fall outside the alliance (Acunniana, Indocypraea, Lipoblepharis, Quadribractea). Here, we present the most comprehensive molecular phylogeny to date of the taxa variously associated with Melanthera in order to test these competing generic limits and explore the biogeographic history of this pan-tropical lineage. The data are consistent with six segregate genera, including the sinking of Hawaiian Islands members of Wollastonia (Melanthera) back into a broader concept of Lipochaeta, although there is currently no recognized morphological synapomorphy to distinguish Lipochaeta s.l. from Wollastonia. Our results suggest that the Melanthera alliance originated some time during the Pliocene or Pleistocene and a strong contemporary presence of the alliance and closely related Ecliptinae outgroups in the Americas suggests that this region may have been the center of origin with subsequent dispersal. We illustrate the difficulty of reconstructing the dispersal history of the remaining genera and present the most parsimonious colonization hypotheses. [ABSTRACT FROM AUTHOR]

Diplusodon is the second-largest genus of the Lythraceae with 103 species of shrubs or subshrubs endemic to the Cerrado Domain. Many species are narrow endemics, restricted to campos rupestres on quartzite rocks, while others are found in cerrado and, more rarely, in gallery, or dry forest margins. Phylogenetic relationships inferred from nuclear ITS and ETS sequences strongly support the monophyly of Diplusodon. We found four major clades, three of which are strongly correlated with endemism and are centered in the Espinhaço Range (CEP), the Plateau Goiano (PG) and the Chapada Diamantina (CD). The fourth clade was designated the Cerrado clade (C), containing species from all three centers of diversity as well as widely distributed species and several species of lower-elevation cerrado. Species with the most plesiomorphic features occur in the clades east of the São Francisco River (C, CEP, CD), while the greatest morphological diversity is found in the PG. The crown node of the genus is estimated to date to the late Pliocene, likely diversifying in the campos rupestres, where most of the later Pleistocene speciation bursts also appear to have occurred. These patterns of diversification in Diplusodon appear to agree with old stable landscapes (OSL) theory, where high levels of in situ speciation in islands of campo rupestre, possibly isolated by more intense fire regimes of the surrounding cerrado vegetation, resulted in accumulation of high numbers of microendemic species. However, the adaptability of the genus is evidenced by multiple dispersal events to lower elevation cerrado, several long-distance dispersals, notably in the C clade, and the presence, in several clades, of species found both in campo rupestre and cerrado habitats. [ABSTRACT FROM AUTHOR]

Artemisia subg. Seriphidium is one of the largest groups within Artemisia, encompassing more than one hundred species, some of them having considerable ecological and economical importance. However, the evolution of subg. Seriphidium has received less attention in comparison to other subgenera of Artemisia, probably, apart from the difficulty of sampling throughout its very large distribution area, because of the low molecular and morphological variability observed in previous studies. Here, we use thorough taxonomic sampling within both Artemisia and subg. Seriphidium to reconstruct the evolutionary history of the subgenus, employing nuclear and plastid DNA sequences as well as various phylogenetic, biogeographic and diversification dynamics tools to analyse the data. Our results show that subg. Seriphidium is not monophyletic, but segregated into two main clades: one large monophyletic group corresponding to the formerly recognised sect. Seriphidium and a second, small clade, phylogenetically distant from the first. Biogeographic and diversification analyses indicate that a rapid radiation of species within sect. Seriphidium occurred in Central Asia during the Miocene-Pliocene transition. The results of our biogeographic analysis suggest that this diversification process started around the Tian-Shan, Pamir and Hindu Kush mountain ranges, subsequently expanding into the Eurasian continent. Finally, we uncovered numerous incongruences between taxonomic and genetic information in several sect. Seriphidium species, which could be explained by morphological uniformity, hybridisation and/or incomplete lineage sorting processes. [ABSTRACT FROM AUTHOR]

Salicornioideae (Amaranthaceae/Chenopodiaceae) comprise 11 genera and ca. 100 species of succulent halophytes distributed worldwide in coastal and inland saline habitats. Most species have peculiar articulated, seemingly leafless stems and spike-like sessile thyrses with highly reduced flowers hidden by fleshy bracts. We analysed sequence data of four markers (ETS, ITS, atpB-rbcL, matK-trnK) from 57 species, 9 subspecies, 1 variety and 2 hybrids representing all curently accepted genera using maximum likelihood, Bayesian analysis, fossil-calibrated molecular dating and ancestral area analysis. All molecular markers gave similar phylogenet ic signals. Salicornioideae probably or iginated in Eurasia during the late Eocene--early Oligocene. The divergence of the early main clades took place in the middle Oligocene with the separation of Allenrolfea/Heterostachys, Halocnemum/Halopeplis/Halostachys/Kalidium and Arthrocnemum/Microcnemum/Tecticornia/Salicornia/ Sarcocornia lineages. The latter lineage diversified most and comprises 3/4 of the species of the subfamily. The Arthrocnemum macrostachyum/Microcnemum lineage diverged at the turn of the Oligocene/ Miocene -- while the Arthrocnemum subterminale, Tecticornia and Salicornia/Sarcocornia lineages were all present by the middle Miocene. Long-distance dispersal has taken place several t imes to the Amer icas (Allenrolfea/Heterostachys, Arthrocnemum subterminale, Salicornia/Sarcocornia), South Africa (Halopeplis, Salicornia/Sarcocornia) and Australia (Tecticornia, Salicornia/Sarcocornia). Most currently accepted genera were resolved as monophyletic. However, Sarcocornia is paraphyletic in relation to Salicornia, and Arthrocnemum proved to be polyphyletic. In addition, the name Arthrocnemum is a nomenclatural synonym of Salicornia. As a consequence, we propose the merging of Sarcocornia under Salicornia and provide 19 new nomenclatural combinations and one replacement name. For Salicornia (incl. Sarcocornia) we propose a new infrageneric classification to accommodate the four phylogenetically and geographically well-supported sublineages of the genus. We also propose abandoning the name Arthrocnemum in its current use, and describe two new genera, Arthrocaulon (Eurasia, Africa) and Arthroceras (North America), and provide three new nomenclatural combinations. [ABSTRACT FROM AUTHOR]

Molecular phylogenetic analyses of nuclear and chloroplast DNA sequences indicate that the rediscovered Apostates, known only from Rapa in the Austral Islands of southeastern Polynesia, represents an example of extreme long-distance dispersal (>6500 km) from southwestern North America and one of at least four disjunctions of comparable magnitude in the primarily New World Bahia alliance (tribe Bahieae). Each of the disjunctions appears to have resulted from north-to-south dispersal since the mid-Miocene; three are associated with such marked morphological and ecological change that some of the southern taxa (including Apostates) have been treated in distinct genera of uncertain relationship. Phyllotaxy within the Bahia alliance, however, evidently has been even more conservative evolutionarily than reflected by previous taxonomies, with alternate-leaved and opposite-leaved clades in Bahia sensu Ellison each encompassing representatives of other genera that share the same leal arrangements. A revised taxonomic treatment of the Bahia alliance is proposed to recognize morphologically distinctive, monophyletic genera, including the critically endangered Apostates. [ABSTRACT FROM AUTHOR]

Lotus comprises ca. 130 species of herbs, semishrubs and shrubs native to the Old World, including important pasture crops and a model legume, L. japonicus. Earlier nrlTS-based phytogenies were incongruent with all taxonomic classifications of the genus. In particular, members of the former genus Dorycnium were unexpectedly placed near species of L. sect. Lotus. The primary goal of the present study is to explore whether the unexpected placement of members of sect. Lotus and the former genus Dorycnium in earlier phylogenetic studies resulted from (1) insufficient taxon sampling and/or (2) the use of only one DNA marker. The rooting of the Lotus phytogeny, its major clades and basic biogeographic patterns are also discussed. This is the first global phylogenetic study of Lotus that uses both plastid and nuclear markers. The nrlTS region was analyzed in 155 ingroup specimens representing 98 species of Lotus. Sequences of nrlTS, nrETS, psbA-trnH spacer and rpsl6 intron were analyzed for 70 ingroup specimens representing 54 species. The placement of the segregate genera Dorycnium and Tetragonolobus in the synonymy of Lotus was confirmed. Analyses of plastid data strongly supported a basal split of Lotus into two clades, one comprising species of sect. Lotus plus those traditionally placed in Dorycnium and the other including the rest of the species. The former clade has a centre of species diversity in Europe and N Asia, and the latter in Macaronesia, Africa and S Asia. Only the "Southern" clade is resolved in analyses of nrlTS and nrETS data. Trees inferred from plastid, nrlTS and nrETS data shared the occurrence of several smaller clades corresponding to traditionally recognized infrageneric taxa or species groups as well as the occurrence of some well-supported clades that differ from traditional taxonomic concepts. Several instances of incongruence were documented between nuclear and plastid markers and between the two nuclear markers, possibly resulting from reticulate evolution. The extant geographic patterns of Lotus are likely biased by at least one round of area fragmentation followed by expansion coupled with extensive speciation associated with the complex history of the Mediterranean biome. [ABSTRACT FROM AUTHOR]

A bibliography on the subject of biogeography is presented which includes several scholarly articles including "Phytogeography: The history and formation of species," by J.C. Avise, "Dispersal and vicariance in Hawaii: Submarine slumping does not create deep inter-island channels," by R.H. Cowie and B.S. Holland and "Biogeographic patterns in the Hawaiian islands," by V.A. Funk and W.L. Wagner.

Cryptolepis is the second-largest genus in the Periplocoideae with 30 species. It is the most widely distributed genus in the subfamily and occupies a wide range o f habitats, from moist forests to stony semi-deserts. Analyses of DNA-data revealed that Cryptolepis is paraphyletic and, for monophyly to prevail, Parquetina should be included in Cryptolepis. Most of the species fall into two major clades: one confined to NE Africa and neighbouring areas; the other absent from NE Africa but widely distributed from West to South Africa. Most species o f Cryptolepis from the more arid areas are shrubby, often with reduced leaves and often with much reduced, few-flowered inflorescences. We establish that Cryptolepis arose about 21 million years ago and began its major radiation around 16 million years ago. Despite this age and its wide distribution, it has not diversified significantly, perhaps limited in the more arid regions by the absence of succulence. It is also possible that the relatively uniform floral structure also limited its response to the diversity of pollinators. Although most species of Cryptolepis are African, there are three species in SE Asia. We show that these three SE Asian species are nested among African taxa and are likely to have reached SE Asia by long-distance dispersal. [ABSTRACT FROM AUTHOR]

Lobelioideae comprise about 1200 species in 30 genera worldwide and include famous cases of parallel evolution in the "giant rosette" lobelias occurring in East Africa, Hawaii, and Bhutan. To date, no phylogeny has included the Himalayan giant lobelia, and its age and phylogenetic distance from the Hawaiian and African giant species (the latter in the same section as the Bhutan species) are therefore unknown. Asian Lobelioideae in general have received little attention, partly because of the difficult access to material. Here we use nuclear and plastid markers of Lobelioideae (with worldwide sampling) to test the monophyly of the section that includes most Asian lobelias, and we then use a reduced concatenated matrix to resolve the position of Lobelia nubigena from Bhutan and of the likewise cold‐adapted lowland species L. sessilifolia. Most sequences come from herbarium material. Lobelia nubigena diverged from its Asian lowland relatives 13.8 (12.6–15) million years ago (about the time of the Bhutan Himalaya orogeny), and in an unexpected twist, L. sessilifolia is the sister species to a huge clade (ca. 265 species) that comprises African, South American, Asian, and the entire Hawaiian Lobelioideae radiation. The African and Hawaiian high‐elevation lobelias in our sample are younger than L. nubigena, although the time when gigantism evolved in this species cannot be precisely inferred. Our results support the previously hypothesized Asian origin of (1) the Hawaiian Lobelioideae, (2) Southeast Asian islands lobelias, and (3) the African Lobelia‐Rynchopetalum group, which also reached South America. [ABSTRACT FROM AUTHOR]

Lobelioideae comprise about 1200 species in 30 genera worldwide and include famous cases of parallel evolution in the "giant rosette" lobelias occurring in East Africa, Hawaii, and Bhutan. To date, no phylogeny has included the Himalayan giant lobelia, and its age and phylogenetic distance from the Hawaiian and African giant species (the latter in the same section as the Bhutan species) are therefore unknown. Asian Lobelioideae in general have received little attention, partly because of the difficult access to material. Here we use nuclear and plastid markers of Lobelioideae (with worldwide sampling) to test the monophyly of the section that includes most Asian lobelias, and we then use a reduced concatenated matrix to resolve the position of Lobelia nubigena from Bhutan and of the likewise cold-adapted lowland species L. sessilifolia. Most sequences come from herbarium material. Lobelia nubigena diverged from its Asian lowland relatives 13.8 (12.6-15) million years ago (about the time of the Bhutan Himalaya orogeny), and in an unexpected twist, L. sessilifolia is the sister species to a huge clade (ca. 265 species) that comprises African, South American, Asian, and the entire Hawaiian Lobelioideae radiation. The African and Hawaiian high-elevation lobelias in our sample are younger than L. nubigena, although the time when gigantism evolved in this species cannot be precisely inferred. Our results support the previously hypothesized Asian origin of (1) the Hawaiian Lobelioideae, (2) Southeast Asian islands lobelias, and (3) the African Lobelia-Rynchopetalum group, which also reached South America. [ABSTRACT FROM AUTHOR]

The genus Thlaspi has been variously subdivided since its description by Linnaeus in 1753, but due to similarities in fruit shape several segregates have still not gained broad recognition, despite the fact that they are not directly related to Thlaspi. This applies especially to segregates now considered to belong to the tribe Coluteocarpeae, which includes several well‐studied taxa, e.g., Noccaea caerulescens (syn. Thlaspi caerulescens), and the widespread Microthlaspi perfoliatum (syn. Thlaspi perfoliatum). The taxonomy of this tribe is still debated, as a series of detailed monographs on Coluteocarpeae was not published in English and a lack of phylogenetic resolution within this tribe was found in previous studies. The current study presents detailed phylogenetic investigations and a critical review of morphological features, with focus on taxa previously placed in Microthlaspi. Based on one nuclear (ITS) and two chloroplast (matK, trnL‐F) loci, four strongly supported major groups were recovered among the Coluteocarpeae genera included, corresponding to Ihsanalshehbazia gen. nov., Friedrichkarlmeyeria gen. nov., Microthlaspi s.str., and Noccaea s.l. In addition, two new species of Microthlaspi, M. sylvarum‐cedri sp. nov. and M. mediterraneo‐orientale sp. nov., were discovered, which are well supported by both morphological and molecular data. Furthermore, M. erraticum comb. nov. (diploid) and M. perfoliatum s.str. (polyploid) were shown to be distinct species, phylogenetically widely separate, but with some overlap in several morphological characters. Detailed descriptions, notes on taxonomy, geographical distribution, and line drawings for the new species and each species previously included in Microthlaspi are provided. In addition, the current taxonomic state of the tribe Coluteocarpeae is briefly discussed and it is concluded that while several annual taxa are clearly distinct from Noccaea, many perennial taxa, after thorough phylogenetic and morphological investigations, may have to be merged with this genus. [ABSTRACT FROM AUTHOR]

Cool climate biomes in the Northern Hemisphere have evolved relatively recently. Temperature decline between the late Eocene and the Pliocene contributed to the evolution of many plant groups in modern cool climate biomes, such as mosses belonging to the Brachythecium novae-angliae-complex. The Brachythecium novae-angliae-complex comprises three species that have a disjunct distribution in temperate and boreal regions in the Northern Hemisphere. The core species in the complex have only recently been transferred from Bryhnia to Brachythecium and the delimitation of the complex and of species within it still remains uncertain. We reconstructed the evolutionary history of the group using molecular data and estimated the timing of the diversifications in the tree. In addition, we examined the morphology and compared morphological diversification with genetic diversity. The Brachythecium novae-angliae-complex diversified in East Asia in the middle Miocene, 9.1 Ma (HPD 95% 3.9-14.8 Myr). Bryhnia trichomitria and B. graminicolor were revealed not to be close relatives of the complex and are therefore transferred to Brachythecium and Koponeniella gen. nov., respectively. Both morphology and molecular data supported the position of Asian Brachythecium noesicum as sister to the Brachythecium novae-angliae crown elade that includes two species, B. hultenii and B. novae-angliae. The crown clade diverged at the end of the Miocene, 6.4 Ma (HPD 95% 2.4-11.6 Myr) and its taxa have a disjunct distribution in cool temperate and boreal areas in the Beringia region (B. novaeangliae subsp. hultenii), and eastern North America and northwestern Europe (B. novae-angliae subsp. novae-angliae). In the crown clade the majority of populations belong to one common haplotype present in all three disjunct parts of the distribution area. Despite low variation in DNA sequence data, morphology allows the distinction between B. novae-angliae subsp. hultenii and B. novae-angliae subsp. novae-angliae. Because of weak morphological and genetic differentiation, European Bryhnia scabrida is treated here as a synonym of Brachythecium novae-angliae subsp. novae-angliae. Glacial survival during the Quaternary and climatic oscillations during the late Tertiary and Quaternary in East Asia may have resulted in diversification of the complex and enabled adaptation to cool temperate and boreal climates. [ABSTRACT FROM AUTHOR]

Hornworts comprise ca. 220 species and are among the oldest landplant lineages, even though their precise phylogenetic position remains unclear. Deep within–hornwort divergences, highly uneven species numbers per genus, and the assumed high stem age together suggest a history of changing diversification (i.e., speciation minus extinction) rates. To study the geographic distribution of modern hornworts and their patterns of species accumulation, we generated a mitochondrial and plastid DNA matrix for 103 species representing all major groups and then applied molecular–clock dating, using a different calibration approach than in earlier work. We used the BAMM software to fit rate–variable and constant–rate birth–death diversification models to the dataset, and we also inferred ancestral areas to a time depth of 55 Ma (Early Eocene). We analyzed diversification rates for all hornworts and separately for species–rich subclades. Under BAMM's variable–rates model (which fits the data better than a constant–rate birth–death model, but still assumes that each species has the same speciation and extinction probability regardless of its age), hornworts have gradually increasing rates of speciation and a constant background extinction rate. No shifts in diversification rate could be detected. The implausible finding of a constant background extinction rate illustrates the limitations of diversification modeling especially as regards extinction rates. [ABSTRACT FROM AUTHOR]

Berberis is the largest genus in the Berberidaceae, comprising more than 500 species. It is now recognised to include all of the compound‐leaved species formally ascribed to Mahonia, as well as simple‐leaved species comprising Berberis s.str. Berberis s.l. has a mainly Northern Hemisphere distribution, with a centre of diversity in the Sino‐Himalaya region, while Berberis s.str. extends into South America where it has a secondary centre of diversity. We analyzed nuclear ITS and chloroplast ndhF sequence data from 68 accessions of Berberis s.l. The results support the monophyly of Berberis s.l., but compound‐leaved Berberis are shown to be paraphyletic. The analysis supports Berberis higginsae, a member of North American B. sect. Horridae, as sister to all other Berberis species. Our results, interpreted in the light of fossil evidence, suggest a North American origin of Berberis s.l., but the area of origin of the simple‐leaved group remains uncertain. [ABSTRACT FROM AUTHOR]

The Alyssum montanum--A. repens polyploid complex is a group of related perennial taxa with a diversity centre in the European (Sub)Mediterranean, controversial taxonomic treatments, and a poorly known evolutionary history. In the present study, morphological, ploidy level and genetic (AFLPs and chloroplast DNA sequences) data were collected to address the taxonomy and evolution of a sublineage of this complex distributed in southwestern Europe and Morocco. As a result, a new taxonomic treatment, differing substantially from recent concepts, is presented, including an identification key, synonymy and typifications. The recognition of several previously described but recently not accepted endemics is favoured (A. flexicaide, A. orophilum, A. rhodanense), whereas the existence of southern Iberian endemics is not supported. Most of the Iberian Peninsula is occupied by a single species for which the name A. fastigiatum is applicable. Populations from the summit areas of the Pyrenees represent a separate species, which is described here as A. cacuminum. Populations from coastal sand dunes in the Basque country (Bay of Biscay) and Galicia, recently recognised as two subspecies of A. loiseleurii, are elevated to species rank according to their genetic and morphological divergence. Alyssum atlanticum is resolved as a species confined to northern Africa and not reaching southern Spain. The distribution of A. montanum in a strict sense is much more restricted than previously reported, being delimited by the Pyrenees in the south, and the Alps in the northeast. The species complex studied here is composed of several polyploid stenoendemics confined to different mountain ranges or specific lowland habitats, and a few relatively widely distributed species. We infer that hybridisation and polyploidisation events, along with ecological and geographic isolation, have stimulated speciation in this complex. Hypotheses about the origin and evolutionary history of the species are discussed. [ABSTRACT FROM AUTHOR]

The internal and external transcribed spacers of nuclear ribosomal DNA were sequenced in 167 samples of Memecylon s.str. and 22 outgroup samples of Lijndenia, Mouriri, Spathandra, Votomita, and Warneckea. Maximum-likelihood analyses of ETS, ITS1, 5.8S and ITS2 alignments yielded tree topologies that are not significantly incongruent, with one exception involving the Tanzanian sample Luke 9741. Monophyly of Memecylon s.str. is strongly supported in the separate ETS and the combined ETS+1TS1+5.8S+1TS2 analyses. Also supported in all analyses except 5.8S is a sister-group relationship between a small species group from western and central Africa (Memecylon subg. Mouririoidea, ovary 4-loculed) and the remaining taxa (M. subg. Memecylon, ovary unilocular). In the combined analysis, internal branches at the base of M. subg. Memecylon are short and weakly supported, yet within this large subgenus one finds a series of monophyletic groups representing different parts of the widespread paleotropical distribution (one group in western and central Africa; two separate groups in East Africa, one of these extending to southern Africa and disjunctly to western and northern Madagascar; one species-rich group occurring exclusively on Madagascar and the neighboring Comoro and Mascarene islands; and three distinct groups in Indo-Malesia, one of these also including the Seychelles endemic M. elaeagni). Within the western and central African clade, M. sect. Polyanthema sensu Jacques-Félix is paraphyletic with respect to M. sect. Afzeliana, and at the morphological level it seems to be a group defined by symplesiomorphies (ovary unilocular, fruit globose). Three East African species ( M. fragrans, M. greenwayi, M. semseii) are returned to Memecylon s.str. after being erroneously transferred to Lijndenia by Borhidi. The Madagascan endemic M. mocquerysii is no longer considered a taxonomic synonym of the distantly related, Tanzanian M. cogniauxii. In addition to elevating M. sect. Mouririoidea to subgeneric rank, the following changes are proposed in the infrageneric classification of African Memecylon: (1) the circumscription of sect. Polyanthema is narrowed to comprise only the members of the " M polyanthemos complex" sensu Jacques-Félix; (2) Engler's sections Tenuipedunculata, Cauliflora, and Obtusifolia are re-instated with emended descriptions (in the case of sect. Cauliflora with an expanded circumscription); (3) seven new sections, Buxifolia, Diluviana, Felixiocylon, Germainiocylon, Magnifoliata, Montana, and Sitacylon, are described; (4) the purported occurrence of M sect. Pseudonaxiandra in East Africa is rejected. A key is provided to the two subgenera and twelve sections currently recognized in African Memecylon. Further study is needed toward a sectional classification of Indo-Malesian Memecylon, and for revision of the seven Madagascan sections recognized by Jacques-Félix. [ABSTRACT FROM AUTHOR]

Although the Balkan Peninsula belongs to the main European biodiversity centres, phylogeographical structure of its high-mountain flora and relationships with adjoining mountain systems remain almost unknown. Here, we analyse the evolutionary patterns in Campanula alpina s.l, which comprises C. alpina Jacq. (eastern Alps, Carpathians) and the mostly neglected, allopatric Balkan taxon C. orbelica Pancic. Using a range-wide sampling, two molecular marker systems (mostly nuclear AFLPs and chloroplast DNA sequences), flow cytometry and morphometric assessment, we (1) test earlier hypotheses assuming the impact of the Balkan lineages on the history of populations in the Alps and the Carpathians; (2) elucidate the long-lasting controversy concerning the taxonomic status of the populations in the Balkan Peninsula. The primary evolutionary break, evidenced by a substantial genetic differentiation, separated the Alpine/Carpathian and Balkan populations. Thus, the previously hypothesized influence of the Balkan gene pool on the present more northerly lineages (especially on a distinct phylogeographical group occurring in the southwestern Carpathians) was rejected. Bayesian phylogenetic analysis of cpDNA may support the Carpathians as the ancestral area within C. alpina from which the species reached the Alps, although strongly divergent AFLPs point at a relatively ancient time of this colonization. Molecular data and clear morphological segregation supported C. orbelica as a Balkan endemic species evolutionarily divergent from C. alpina. Both taxa are lectotypified, illus-trated and morphologically characterized; their diagnostic characters are outlined. [ABSTRACT FROM AUTHOR]

It has been suggested that many plants now found in the arctic originated from ancestors that occurred at high altitudes in the southern mountains of the Northern Hemisphere during the Tertiary. However, this hypothesis has rarely been tested using a molecular phylogenetic approach. Here, we present a fossil-calibrated molecular phylogeny of Lagotis, an arctic-alpine genus with the greatest diversity in the Qinghai-Tibetan Plateau (QTP) and Central Asian mountains, based on five chloroplast (matK, psbA-trnH, rpslô, trnG-S, trnL-F) and nuclear ribosomal ITS DNA markers. Within this framework, we infer the ancestral area and biogeographic history of the genus. Four major clades (A-D) within Lagotis were recovered with strong support, which largely correspond to the previous classification of the genus. Within clade A, Lagotis species from QTP were distributed among several subclades, and L. integrifolia from Central Asia was sister to L. glauca and L. minor from the arctic and subarctic region. The Bayesian molecular dating and the ancestral area reconstruction analyses suggested that Lagotis could have originated in the QTP in the Miocene (Tertiary), and that the genus radiated from the Miocene to Pleistocene. The diversification of Lagotis probably took place predominantly in the QTP and it subsequently spread to the Central Asian high-lands, followed by northward migration into the arctic. Our results support the hypothesis that the "Central Asiatic Highland Corridor" as an important route for the migration of the flora between the arctic and QTP. [ABSTRACT FROM AUTHOR]

Euphorbia subg. Athymalus consists of approximately 150 species and is one of the four main lineages that comprise the species-rich genus Euphorbia. Most species in the subgenus are stem succulents with greatly reduced leaves, but there are also leafy herbs, shrubs, trees and geophytes. The subgenus is restricted to arid regions of the Old World. Most species are found in sub-Saharan Africa, with one in Macaronesia and adjacent parts of western Africa, a few in the Arabian Peninsula (one of which extends into Iran) and one native to Madagascar. Twenty-three species are endemic to the northeastern Horn of Africa (SE Ethiopia, Socotra, Somalia), while 72 species are restricted to southern Africa (including Botswana, Lesotho, Namibia, South Africa and Swaziland). Sixty of those are endemic to South Africa alone, where they are particularly associ-ated with the semi-arid west and south of the country in the Greater Cape Floristic Region and the Nama Karoo Region. We sampled 88 species and analyzed data from the nuclear ribosomal ITS and plastid ndhF regions. Analyses of the separate and combined datasets produced phylogenies that confirm the monophyly ofE. subg. Athymalus and the placement of E. antso from Madagascar as sister to the rest of the subgenus. Our analyses also show that the subgenus consists of a grade of early-diverging lineages that are relatively poor in species and that the major radiation of succulent species in southern Africa forms a highly supported clade (E. sect. Anthacanthae). Species-level relationships within this southern African clade, however, remain largely unresolved. Our phylogenetic hypotheses allow us to propose a new classification for E. subg. Athymalus where seven sections are recognized, two of which are newly described. The large southern African E. sect. Anthacanthae is further divided into five subsections, and four series are recognized in E. subsect. Florispinae. [ABSTRACT FROM AUTHOR]

Seeds of the genus Weigela (Caprifoliaceae) are described from the Early Miocene Hannuoba Formation of Weichang County, Hebei Province, China. They are flat, elliptic, with translucent membranous wings on three sides. Comparison with extant and fossil species of Weigela indicates that these seeds are different which therefore were assigned to a new species, Weigela weichangensis. On the basis of the fossil record, the distribution of Weigela through geologic ages is shown on a series of palaeogeographic maps of the Northern Hemisphere. The oldest fossils of Weigela are seeds from the Oligocene of West Siberia, Asia. The genus later expanded into East Asia and Europe as well as into Arctic/Subarctic areas of North America via the North Atlantic Land Bridge and/or Bering Land Bridge in the Miocene. Weigela disappeared from Europe and North America probably during the Pleistocene glaciations, but survived in Asia until now. [ABSTRACT FROM AUTHOR]

Biogeography is a multidisciplinary science concerned with how and why organisms are distributed as they are on Earth. It links fields such as systematics, ecology, paleontology, and climatology, and occupies a central position in evolutionary biology, being fundamental to the study of processes such as speciation and adaptive radiation. Here we provide a brief overview of some particularly dynamic areas of inquiry and offer some perspectives on future directions for the field. We hope that some historical debates, such as those over the importance of dispersal, or the validity of molecular dating, are finally being put to rest. Over the last decade, biogeography has become increasingly integrative, and has benefited from advances in statistical methods for inferring geographic range dynamics in a phylogenetic context, molecular estimation of lineage divergence times, and modeling lineage birth and death. These are enabling greater insights into patterns of organismal diversification in time and space. In the next decade, analytical challenges are emerging on several fronts. For example, phylogenies are increasing in size and taxonomic breadth and new sequencing technologies enabling phylogenetic and phylogeographic datasets are increasingly genomic in depth. In addition, geographic occurrence data are accumulating in online repositories, yet tools for data mining and synthetic analysis are lacking for comparative multi-lineage studies. Biogeography is thus entering an era characterized by phylogenomic datasets, increasingly comprehensive sampling of clades, and interdisciplinary synthesis. We anticipate continued progress in our understanding of biodiversity patterns at regional and global scales, but this will likely require greater collabo-ration with specialists in bioinformatics and computational science. Finally, it is clear that biogeography has an increasingly important role to play in the discovery and conservation of biodiversity. Lessons learned from biogeographic studies of islands are being applied to better understand extinction dynamics as continental ecosystems become more fragmented, and phylogeog-raphy and ecological niche modeling offer innovative paths toward the discovery of previously unknown species distributions and priority areas for conservation. The future of biogeography is bright and filled with exciting challenges and opportunities. [ABSTRACT FROM AUTHOR]

The Apennines show high floristic diversity promoted by their particular role as an ice-age refugium. However, studies on the evolution of Apennine plants are rare. Potentilla rigoana from the mainly European P. heptaphylla group is an example of a largely ignored taxon originally described from this mountain range. Its evolutionary origin and taxonomic status was studied in an interdisciplinary approach combining morphometric, karyological (flow cytometry and chromosome counting), DNA-molecular (AFLP), ecogeographic (WorldClim climatic variables), and phytosociological data. Morphometric analyses revealed 16 metric and 2 nominal characters separating P. rigoana from its close relatives P. crantzii, P. heptaphylla s.l., and P. neumanniana. Additionally, P. rigoana was intermediate with respect to temperature requirements as well as altitudinal distribution compared to P. crantzii and P. heptaphylla s.l. The species was documented from 54 populations or sampling sites located in the central and southern Apennines where it occurs in montane to subalpine dry habitats. Morphometric analysis suggested its occurrence in the Balkans. Molecular and morphometric analyses further suggested reticulate relationships of the Balkan samples of P. heptaphylla s.l. with P. rigoana. Potentilla rigoana comprises five ploidy levels with the dominant diploids and tetraploids geographically centered in the southern and central Apennines, respectively. The existence of diploids and a uniformly heptaploid population indicate the occurrence of both sexuality and apomixis. The combined datasets sug-gested species status for P. rigoana. The ecogeographic distribution indicated an allopatric differentiation of the species within the P. heptaphylla group. The latitudinal separation of diploid and polyploid cytotypes coincided with a glacial refugium in the southern Apennines and more dynamic past climatic conditions in the central Apennines. [ABSTRACT FROM AUTHOR]

Extensive efforts have been made to resolve the phylogeny of the large coffee family (Rubiaceae) based on molecular data. As a result, several small tribes have been described, but the phylogenies and generic delimitations for many of these groups remain unclear. This study focuses on the small tribe Danaideae that belongs to subfamily Rubioideae and whose generic limits have not previously been addressed with molecular data. It is the sole rubiaceous tribe distributed almost entirely in the Western Indian Ocean region, with the exception of the East African Danais xanthorrhoea. The tribe consists of three genera: Danais, Payera (including the monotypic genus Coursiana), and Schismatoclada. We present the first molecular phylogenetic study of Danaideae including representatives from all three genera and using Bayesian and maximum parsimony methods and sequence data from nuclear DNA (nrlTS) and chloroplast DNA (petD, psbA-trnH, rp132-trnLUAG, rps16). Our main objectives were to rigorously test the monophyly of Danaideae as currently circumscribed and assess phylogenetic relationships within the tribe. The findings of this study shed light on the colonization history of the tribe. Our analyses reaffirm the monophyly of Danaideae and Danais but reveal the paraphyly of Payera and Schismatoclada. The close relationship between the three Danaideae genera and Coursiana is supported. However, we found very little support for the inclusion of the latter genus in Payera as proposed earlier. The tribe is resolved in two morphologically distinct major lineages, the highly supported Danais clade with lianes-cent habit (= Danais sensu Buchner & Puff) and the Payera-Schismatoclada clade with arborescent habit. The Malagasy and Mauritian specimens of Danais fragrans are not closely related, and we restrict D. fragrans to the Mauritian taxa and resurrect Danais lyallii Baker to accommodate the Malagasy D. fragrans. According to our analysis, Madagascar is the origin of all spe-cies of Danaideae occurring in the Comoro archipelago, East Africa, and Mauritius. The Mauritian and East African Danais each is the result of a single colonization event, while there were at least two independent colonization events to the Comoros. [ABSTRACT FROM AUTHOR]

We took a two-tiered approach to test monophyly of Nelsonioideae and place the group within Lamiales, and to deter-mine relationships among taxa within the group. Phylogenetic analysis of a molecular dataset (ndhF+trnL-F) for a broad sample of Lamiales supports monophyly of Nelsonioideae and places the clade with strong support as sister to a lineage composed of all other plants treated as Acanthaceae (Avicennia, Thunbergioideae, Acanthoideae). We propose to treat this entire group as Acanthaceae s.l. and hypothesize that indurate, explosively dehiscent capsules are a synapomorphy for the family, albeit with autapomorphic fruit types in Avicennia and Mendoncia. These results further support monophyly of family-level groups that have emerged from recent studies of Lamiales but are largely unsuccessful in resolving relationships among these groups, as also encountered by other workers. Our results contradict some aspects of relationships that have seemed resolved by earlier studies, notably among Byblidaceae, Scrophulariaceae, Thomandersia, and other Lamiales. Among Nelsonioideae, analysis of sequence data from rapidly evolving genie regions (trnS-G, ndhF-rp132+rp132-trnL(UAG), nrlTS) and a larger sample of nelso-nioids (i.e., all genera and multiple taxa to represent the diversity of species-rich genera) indicates that Nelsonia and Elytraria are monophyletic with strong support, but with only moderate support for Nelsonia as the first branching clade and Elytraria sister to the remaining nelsonioids. An African clade comprising monospecific Saintpauliopsis sister to Anisosepalum (two of three species sampled) is sister to a clade that includes all sampled members of pantropical Staurogyne plus New World Gynocraterium and Asian Ophiorrhiziphyllon. Gynocraterium is sister to all sampled members of New World Staurogyne; this last clade is sister to a clade comprising the other sampled Staurogyne plus Ophiorrhiziphyllon, which is nested among Asian Staurogyne. The taxonomic implications of these patterns of relationship are discussed. Our results suggest that Nelsonioideae have a complex history of inter-continental dispersals compared to other lineages of Acanthaceae of similar to much larger size in terms of number of species, making it an interesting group for biogeographic study. [ABSTRACT FROM AUTHOR]

The tribe Liabeae (Compositae) contains ca. 175 species distributed in 18 genera and its members occupy a variety of habitats in the Andes of South America as well as in Mexico, Central America, and the West Indies. The tribe is recognizable by a combination of morphological characters. DNA sequence data from the nuclear ribosomal ITS region and three chloroplast regions (trnL-F, 3' end of ndhF, matK; a total of more than 5 kb of sequence data) were used to infer a phylogeny. The data were analyzed using maximum parsimony, maximum likelihood, and Bayesian inference. The results support the monophyly of the tribe and show a consistent placement for all genera except Cacosmia. Four well-supported clades are recovered in the remainder of the tribe, all recognized as subtribes. Liabineae (Ferreyranthus, Dillandia, Oligactis, Sampera, Liabum) are the sister group of the rest of the tribe. Sinclairineae (Sinclairiopsis and Sinclairia with segregates Liabellum and Megaliabum) are the sister group of Munnoziinae (Chrysactinium nested inside Munnozia s.l.) plus Paranepheliinae (Stephenbeckia, Microliabum, Pseudonoseris, Paranephelius, Chionopappus, Philoglossa, Erato). Cacosmia is placed as the sister group of either all the rest of the tribe or of subtribe Liabineae; morphologically its characters are either autapomorphic or plesiomorphic. Bishopanthus could not be confidently placed in any of the subtribes; it is only known from scraps of the type and a molecular study is not possible. The phylogeny slightly alters previous assumptions about the biogeography and it seems that Liabeae originated in the central and northern Andes and spread north and south with several independent introductions into Mexico and Central America and one into the Caribbean. With the exception of the Liabeae (Andes) and Moquineae (Brazil), all of the tribes in the subfamily Cichorioideae are either restricted to or have their basal grade in Africa. [ABSTRACT FROM AUTHOR]

Tribe Cochlearieae is among few isolated evolutionary lineages in the mustard family and comprises only about 30 species. Earlier taxonomic concepts recognised five genera that might belong with Cochlearia in this tribe (Bivonaea, Ionopsidium, Minaea, Pastorea, Thiaspi), though Pastorea and Minaea have already been reduced to synonymy of lonopsidium. Focusing on DNA sequence information from the nuclear-encoded internal transcribed spacer regions ITSI and ITS2 and the plastid trnL-F region, the phylogenetic position of all lonopsidium species is evaluated within a detailed phylogenetic frame- work of tribe Cochlearieae. It is shown that the tribe can be divided into two major groups: (i) a central European Cochlearia dade extending its distribution far into northern and arctic regions, and (ii) a western Mediterranean dade comprising all six lonopsidium species and three Cochlearia species from the Iberian Peninsula nested within lonopsidium. Consequently, neither Cochlearia nor lonopsidium represent monophyletic genera. The deep split between the two clades dates back to the middle Miocene, approximately 13.8 million years ago. The divergence times between the different major groups within the lonopsidium clades predate or are associated with the Messinian salinity crisis in the late Miocene which also agrees with the relictual and phylogenetically isolated status of the tribe. Only taxa of Cochlearia sect. Cochlearia (Cochlearia dade) underwent a rapid and very recent radiation in the mid or late Pleistocene. Four new taxonomic combinations (Jonopsidium aragonensis (H.J. Coste & Soulié) M. Koch, I. aragonensis subsp. navarranum (P. Monts.) M. Koch, I. glastifolium (L.) M. Koch, and!. megalospermum (Maire) M. Koch) and the new tribe Bivonaeeae are proposed to reflect the deep evolutionary split within the group, which is further supported by DNA sequence variation of the nuclear-encoded, single-copy gene chalcone synthase (chs). [ABSTRACT FROM AUTHOR]

We conducted a molecular phylogenetic study of the tribe Stipeae using nine plastid DNA sequences (irnK-rnalK, ,naiK, lrnI1-p.shA, trnL-F, rps3, ndhF, rp132-trnL, rpsl6-irnK, rpsl6 intron), the nuclear ITS DNA regions, and micromorphological characters from the lemma surface. Our large original dataset includes 156 accessions representing 139 species of Stipcae representing all genera currently placed in the tribe. The maximum likelihood and Bayesian analyses of DNA sequences provide strong support for the monophyly ofStipeae; including, in phylogenetic order, Macrochloa as remote sister lineage to all other Stipeae, then a primary stepwise divergence of three deep lineages with a saw-like (SL) lemma epidermal pattern (a plesiomorphic state). The next split is between a lineage (SLI) which bifurcates into separate Eurasian and American clades, and a lineage ofthree parts; a small Pails (SL2) dade, as sister to Piptaiheruin s.str. (SL3), and the achnatheroid dade (AC). The AC exhibits a maize-like lemma epidermal pattern throughout. AC consists ofa core dade ofAustral-Eurasian distribution and a "major American dade" of North and South American distribution. The base chromosome number for Stipeae is somewhat ambiguous but based on our survey it seems most likely to be x = Il or 12. Our phylogenetic hypothesis supports the recognition ofthe following genera and groups (listed by region): Eurasia-Achnatheru,n, "Miliacea group", "Neotrinia" (monotypic), Qrthoraphiuin (monotypic), Pails (also I from North America), Piplaiheruin s.str., Psaminoch/oa (monotypic), Piilagrosiis, Siipa, "Ti,nouria group", and Trikerala; Mediterranean-A ~npeIodesnios (monotypic), Ce/tica (monotypic), Macrochloa (monotypic), and "Stipella-lnaequiglumes group"; Australasia -Anetnanthele (monotypic), and Austrosiipa; North America (NA)-"Eriocoma group", Hesperostipa, Oiyzopsis (monotypic), Piptaiheropsi.c, "Pseudoeriocoma group", and "Stillmania" (monotypic); South America-Aciachne, Amelichloa (also NA), Anatherosilpa (s.str.), Jarava (polyphyletic), Lorenzochloa, Na.cse/Ia (also NA), Oriachne, Papposilpa (also NA), and Pipiochaeiium (also NA). Monophyly of Phaenospermateae including Duthieinae is demonstrated, and its inclusion within or treatment as sister to Stipeae is rejected. [ABSTRACT FROM AUTHOR]

By reviewing 25 cases of western Eurasian-western North American disjunct flowering plant taxa, we sought to improve understanding of the origin of this type of biogeographic pattern. In nine of the groups studied, phylogenetic and other evidence (often circumstantial) was found for parallel evolutionary shifts of widespread Northern Hemisphere lineages into dry environments in western Eurasia and western North America (Asteraceae-Cichorieae, Chenopodiaceae-Betoidcac, Descurainia, Ericaceae-Arbutoideae, Papaver, Platanus, Antirrh ineae-Maurandya group, Anemone, Stprax). Lotus s.l. also represents independent evolution of dryland taxa but is polyphyletic. Six taxa (Daucus, Erodium, Oligomeris, P/wi logo, Polycarpon, Senecio) and probably four others (Chenopodiaceae-Camphorosmeae, California, Antirrhineae-Gambel ia group, Antirrhineae-Antirrhinum group) are likely examples of long-distance dispersal between western Eurasia and western North America. In Asteraceae-Gnaphalieae either parallel evolution or long-distance dispersal might explain the disjunction. Migration across a Beringian or North Atlantic Land Bridge appears a possible explanation for only three of the 25 disjunctions (Datisca, Cicendia, ZeltneralExaculum/Schenkia) based on taxon ecology and divergence times. The disjunction in Cerci,s remains unexplained. In short, independent ecological shifts to dry environments and long-distance dispersal probably account for the majority of taxon-disjunctions studied; migration of lineages (pre)adapted to dry conditions across either the Beringian or the North Atlantic Land Bridges, much discussed in the literature, only rarely needs to be invoked. Based on the similar times of origin ofdisjunctions between western Eurasian and western North American taxa and those between East Asian and eastern North American plants, we conclude that these two patterns are best regarded as parts ofan ecological, geographical, and temporal continuum. [ABSTRACT FROM AUTHOR]