First Glimpse on Spring Starflower Domestication.

The cultivation and domestication of plants are human-driven processes that change the biology and attributes of a plant. Ipheion uniflorum is a bulbous geophyte known as Spring Starflower whose cultivation dates back to the first half of the 19th century. At least seven cultivars have been developed from natural stands. However, comparative analyses of wild and cultivated materials are largely missing. In the present study, we provide a morphological evaluation and analyses of the cytological and genetic variability of I. uniflorum that reveal significant levels of differentiation and evidence of artificial selection in the Spring Starflower. Distinctive phenotypic characters in cultivated materials that are rarely found or lacking in wild plants and natural populations, such as pink or violet flowers, together with its reduced heterozygosity and starting genetic differentiation support the view of early mechanisms of domestication acting upon Spring Starflower plants. The probable geographic origin of the cultivated forms is discussed together with perspectives for plant breeding.

Genomic, karyological and morphological changes of South American garlics (Ipheion) provide insights into mechanisms of speciation in the Pampean region.

Speciation proceeds through mechanisms that promote reproductive isolation and shape the extent of genetic variation in natural populations, and thus its study is essential to understand the evolutionary processes leading to increased biodiversity. Chromosomal rearrangements are known to facilitate reproductive isolation by hybrid sterility and favour speciation events. The genus Ipheion (Amaryllidaceae, Allioideae) is unique as its species exhibit a remarkable karyological variability but lack population-level genetic data. To unveil the diversification processes acting upon the formation of new lineages within Ipheion in the Pampas of South America, we combined morphology and karyology approaches with genotyping-by-sequencing. Our phylogenomic and population genomics results supported the taxonomic division of Ipheion into three morphological and genetically well-differentiated groups. The origin of Ipheion uniflorum was traced back to its current southern distribution area in the southern Pampean region (in Argentina), from where it had expanded to the north reaching Uruguay. Our results further suggested that chromosome rearrangements and ploidy shifts had triggered speciation events, first during the origin of I. uniflorum and later during its subsequent diversification into I. recurvifolium and I. tweedieanum, in both cases reinforced by extrinsic factors and biogeographical settings. The current study illustrates the analytical power of multidisciplinary approaches integrating phylo- and population genomics with classic analyses to reveal evolutionary processes in plants.

Reconstructing the phylogenetic history of the tribe Leucocoryneae (Allioideae): Reticulate evolution and diversification in South America.

At present, the Allioideae is included within the Amaryllidaceae, which is an economically important bulb crop subfamily that includes onion, garlic, and ornamental species worldwide. The Allioideae includes four tribes geographically disjunct namely: Allieae, widespread in the northern hemisphere, tribe Tulbaghieae distributed in South Africa, and tribes Leucocoryneae and Gilliesieae are endemic to South America. Although we agree with the current tribal circumscription of the Leucocoryneae including Beauverdia, Ipheion, Latace, Leucocoryne, Nothoscordum, and Tristagma, there are still taxonomic and phylogenetic uncertainties regarding the monophyly, phylogenetic relationships, and divergence time of several lineages in a biogeographic context. In this study, a comprehensive molecular phylogeny of the tribe Leucocoryneae was inferred based on nuclear ribosomal ITS and plastid (ndhF and matK) sequences. We used Bayesian inference and maximum parsimony analyses to predict ancestor-descendant relationships. Our results confirmed the monophyly of the four tribes of subfamily Allioideae. Similarly, within the Leucocoryneae, Ipheion, Leucocoryne, and Nothosocordum Sect. Inodorum were also monophyletic; Tristagma and Nothoscordum would be monophyletic if including Ipheion and Beauverdia, respectively. Network analyses were implemented to reveal putative scenarios of reticulate evolution. Both, current and ancestral hybridization events have presumably occurred among species of Nothoscordum Sect. Nothoscordum and Beauverdia favored by spatial overlapping of populations, flowering synchrony and a puzzling pattern of cytogenetic attributes. The estimation of divergence time indicates that the tribe Leucocoryneae originated in the Late Oligocene in southern South America with possible ancestors in Africa. Most crown lineages within the tribe diversified in conjunction with biogeographical events during the Late Miocene to Pliocene. We posit that new suitable environments available after the Andean uplift and during the Age of the Southern Plains provided the favorable geographic setting for the major lineages of Leucocoryneae in southern Pampas, extra-Andean Patagonia, Andean mountains, and in Chile. Hybridization, polyploidization, and Robertsonian translocations of chromosomes have been the driving forces and major sources of speciation in the evolution of tribe Leucocoryneae.

A novel indicator of karyotype evolution in the tribe Leucocoryneae (Allioideae, Amaryllidaceae).

The tribe Leucocoryneae is taxonomically and cytogenetically complex, mainly due to its extraordinary morphological and karyological variation. Robertsonian translocations had long been recognized as a central factor contributing to karyotype diversity within the Leucocoryneae, but so far no major tendency prevailing on the observed complexity of karyotype formula among species has been identified. The assessment of nuclear DNA contents by flow cytometry using propidium iodide in 23 species, representing all genera within the tribe, showed a monoploid genome size variation of 1Cx = 9.07-30.46 pg denoting a threefolds fluctuation. A highly significant linear association between the average DNA content per chromosome arm (2C/FN) and the monoploid genome size (1Cx) is reported for the first time and identified as a novel indicator of a trend governing karyotype diversity within Leucocoryneae. This trend shows that a reduction in DNA content per chromosome arm is influencing and has shaped karyotype evolution of different monophyletic groups within the tribe despite the complex karyotype diversity and apparently contrasting patterns of genome sizes.

Breeding system diversification and evolution in American Poa supersect. Homalopoa (Poaceae: Poeae: Poinae).

BACKGROUND AND AIMS: Poa subgenus Poa supersect. Homalopoa has diversified extensively in the Americas. Over half of the species in the supersection are diclinous; most of these are from the New World, while a few are from South-East Asia. Diclinism in Homalopoa can be divided into three main types: gynomonoecism, gynodioecism and dioecism. Here the sampling of species of New World Homalopoa is expanded to date its origin and diversification in North and South America and examine the evolution and origin of the breeding system diversity. METHODS: A total of 124 specimens were included in the matrix, of which 89 are species of Poa supersect. Homalopoa sections Acutifoliae, Anthochloa, Brizoides, Dasypoa, Dioicopoa, Dissanthelium, Homalopoa sensu lato (s.l.), Madropoa and Tovarochloa, and the informal Punapoa group. Bayesian and parsimony analyses were conducted on the data sets based on four markers: the nuclear ribosomal internal tanscribed spacer (ITS) and external transcribed spacer (ETS), and plastid trnT-L and trnL-F. Dating analyses were performed on a reduced Poa matrix and enlarged Poaceae outgroup to utilize fossils as calibration points. A relaxed Bayesian molecular clock method was used. KEY RESULTS: Hermaphroditism appears to be pleisiomorphic in the monophyletic Poa supersect. Homalopoa, which is suggested to have originated in Eurasia 8·4-4·2 million years ago (Mya). The ancestor of Poa supersect. Homalopoa radiated throughout the New World in the Late Miocene-Early Pliocene, with major lineages originating during the Pliocene to Pleistocene (5-2 Mya). Breeding systems are linked to geographic areas, showing an evolutionary pattern associated with different habitats. At least three major pathways from hermaphroditism to diclinism are inferred in New World Homalopoa: two leading to dioecism, one via gynodioecism in South America and another directly from hermaphroditism in North America, a result that needs to be checked with a broader sampling of diclinous species in North America. A third pathway leads from hermaphroditism to gynomonoecism in Andean species of South America, with strictly pistillate species evolving in the highest altitudes. CONCLUSIONS: Divergence dating provides a temporal context to the evolution of breeding systems in New World Poa supersect. Homalopoa The results are consistent with the infrageneric classification in part; monophyletic sections are confirmed, it is proposed to reclassify species of sect. Acutifoliae, Dasypoa and Homalopoa s.l. and it is acknowledged that revision of the infrageneric taxonomy of the gynomonoecious species is needed.

Phylogeny of the Paniceae (Poaceae: Panicoideae): integrating plastid DNA sequences and morphology into a new classification.

Included in the PACMAD clade of the family Poaceae (Panicoideae, Arundinoideae, Chloridoideae, Micrairoideae, Aristidoideae, Danthonioideae), the tribe Paniceae s.l. is one of the largest tribes of the subfamily Panicoideae, with more than 2000 species. This tribe comprises a huge morphological, cytological and physiological diversity represented by different inflorescence types, several basic chromosome numbers, and at least four major photosynthetic pathways. The tribe Paniceae has been the subject of molecular studies that have confirmed its paraphyly: two major clades were recognized based on their basic chromosome numbers (x = 9, x = 10). The x = 10 Paniceae clade is sister to the Andropogoneae-Arundinelleae s.s. clade (x = 10), while the combined x = 10 clade is sister to the x = 9 clade that contains the remaining genera of Paniceae. As a result of a recent realignment within the tribe in terms of the phylogenetic position of minor and major Paniceae genera, a reanalysis of the whole sampling is performed and new underrepresented taxa are discussed. A total of 155 genera, currently considered within subfamily Panicoideae, are represented here by almost all genera of Paniceae s.l., representatives of Andropogoneae and Arundinelleae s.s., and the endemic and small tribe Steyermarkochloeae; we also included specimens of subfamily Micrairoideae, tribes Isachneae and Eriachneae. The sampling includes as outgroups 18 genera of the PACMAD clade (excluding Panicoideae) and four genera from the BEP clade (Bambusoideae, Ehrhartoideae, Pooideae), rooting with Bromus inermis. A matrix with 265 taxa based on the combined evidence from ndhF plastid sequences (2074 bp) and 57 morphological characters was subjected to parsimony analyses. Jackknife resampling was used to calculate group support. Most clades are characterized by morphological, cytological, anatomical, and/or physiological characters. Major tribal changes are based on the basic chromosome number; the pantropical x = 9 clade is here recognized as Paniceae s.s., while the American x = 10 Paniceae s.l. is restricted to the reinstated tribe Paspaleae. The optimization of the photosynthetic pathway for the Paspaleae-Andropogoneae-Arundinelleae s.s. clade, including the monotypic Reynaudia, shows a plesiomorphic C4 state while the ancestral state for Paniceae s.s. is ambiguous. If Reynaudia were not included or placed elsewhere, the ancestral photosynthetic pathway for both the Paspaleae-Andropogoneae-Arundinelleae s.s. clade and the Paniceae s.s. would be unambiguously C3 . In order to explore character evolution further, the morphological characters were mapped onto one of the most parsimonious trees. A relationship between photosynthetic pathways and inflorescence morphology is suggested here for the first time. Based on the optimization of morphological characters and additional data, we propose names for almost all inner clades at the rank of subtribe with a few groups as incertae sedis. With this extensive sampling, we resolved the phylogenetic relationships and the assignation of synapomorphies, and improved the support in subtribe sorting; consequently a robust circumscription of the tribe Paniceae s.l. is proposed.

Phylogenetic studies favour the unification of Pennisetum, Cenchrus and Odontelytrum (Poaceae): a combined nuclear, plastid and morphological analysis, and nomenclatural combinations in Cenchrus.

BACKGROUNDS AND AIMS: Twenty-five genera having sterile inflorescence branches were recognized as the bristle clade within the x = 9 Paniceae (Panicoideae). Within the bristle clade, taxonomic circumscription of Cenchrus (20-25 species), Pennisetum (80-140) and the monotypic Odontelytrum is still unclear. Several criteria have been applied to characterize Cenchrus and Pennisetum, but none of these has proved satisfactory as the diagnostic characters, such as fusion of bristles in the inflorescences, show continuous variation. METHODS: A phylogenetic analysis based on morphological, plastid (trnL-F, ndhF) and nuclear (knotted) data is presented for a representative species sampling of the genera. All analyses were conducted under parsimony, using heuristic searches with TBR branch swapping. Branch support was assessed with parsimony jackknifing. KEY RESULTS: Based on plastid and morphological data, Pennisetum, Cenchrus and Odontelytrum were supported as a monophyletic group: the PCO clade. Only one section of Pennisetum (Brevivalvula) was supported as monophyletic. The position of P. lanatum differed among data partitions, although the combined plastid and morphology and nuclear analyses showed this species to be a member of the PCO clade. The basic chromosome number x = 9 was found to be plesiomorphic, and x = 5, 7, 8, 10 and 17 were derived states. The nuclear phylogenetic analysis revealed a reticulate pattern of relationships among Pennisetum and Cenchrus, suggesting that there are at least three different genomes. Because apomixis can be transferred among species through hybridization, its history most likely reflects crossing relationships, rather than multiple independent appearances. CONCLUSIONS: Due to the consistency between the present results and different phylogenetic hypotheses (including morphological, developmental and multilocus approaches), and the high support found for the PCO clade, also including the type species of the three genera, we propose unification of Pennisetum, Cenchrus and Odontelytrum. Species of Pennisetum and Odontelytrum are here transferred into Cenchrus, which has priority. Sixty-six new combinations are made here.

Molecular phylogeny of the subtribe Melinidinae (Poaceae: Panicoideae: Paniceae) and evolutionary trends in the homogenization of inflorescences.

The subtribe Melinidinae (Poaceae: Panicoideae: Paniceae) includes 14 genera that present the PCK photosynthetic subtype in addition to several other unique and also common characters. The purpose of this research was (1) to test the monophyly of the subtribe Melinidinae, including 331 ndhF sequences of Panicoids and related genera, (2) to analyze the phylogenetic relationships among genera of Melinidinae using four cpDNA regions, and (3) to study evolutionary trends in the homogenization of inflorescences. As a result, the monophyly of Melinidinae is supported if Urochloa venosa is excluded from the subtribe. Alloteropsis semialata subsp. semialata, an unusual PCK species, is here confirmed within the Forest shade clade. Within Melinidinae, Urochloa and Eriochloa appeared as paraphyletic and polyphyletic genera, respectively. Finally, the general trend in the evolution of the inflorescences in Melinidinae seems to be the reduction from non-homogenized to complete homogenized inflorescences.

Phylogeny of New World Stipeae (Poaceae): an evaluation of the monophyly of Aciachne and Amelichloa.

© The Willi Hennig Society 2010. ABSTRACT: The tribe Stipeae, with nearly 550 species, includes 28 core genera, of which 13 occur in America: Achnatherum, Aciachne, Amelichloa, Anatherostipa, Hesperostipa, Jarava, Nassella, Ortachne, Oryzopsis, Pappostipa, Piptatherum, Piptochaetium and Ptilagrostis. Based on 37 species representing 14 Stipeae genera, and using four chloroplast markers and morphological characters, we provide a phylogenetic hypothesis of the New World Stipeae, with our focus on Amelichloa and Aciachne. Parsimony analyses included two approaches: (i) a multiple-sequence alignment where gaps were treated as missing or coded, (ii) using direct sequences by direct optimization as implemented by the program POY v.4.0.2870. Analyses under direct optimization were conducted using the molecular data sets independently and combined, and with morphological data. Different cost regimes were explored and the one producing the highest congruence between partitions was chosen. Among the genera considered, only Piptochaetium, Austrostipa, and Hesperostipa were resolved as monophyletic, while Achnatherum, Amelichloa s.l., Anatherostipa, Jarava and Nassella were polyphyletic, and Aciachne was polyphyletic or paraphyletic. As a result, Amelichloa can be restricted to a monophyletic group if including A. brachychaeta, A. ambigua, A. clandestina and A. caudata, or it should be considered within Nassella. The phylogenetic position of species of Aciachne suggests inbreeding and outbreeding events with species of Anatherostipa, Ortachne and Hesperostipa.

A molecular phylogeny of Panicum (Poaceae: Paniceae): tests of monophyly and phylogenetic placement within the Panicoideae.

Panicum L. is a cosmopolitan genus with approximately 450 species. Although the genus has been considerably reduced in species number with the segregation of many taxa to independent genera in the last two centuries, Panicum remains a heterogeneous assemblage, as has been demonstrated in recent years. The genus is remarkably uniform in its floral characters but exhibits considerable variation in anatomical, physiological, and cytological features. As a result, several classifications, and criteria of what the genus should really include, have been postulated in modern literature. The purpose of this research, based on molecular data of the chloroplast ndhF gene, is to test the monophyly of Panicum, to evaluate infrageneric classifications, and to propose a robust phylogenetic hypothesis. Based on the present results, previous morphological and molecular phylogenetic studies, and inferred diagnostic morphological characters, we restrict Panicum sensu stricto (s.s.) to the former subgenus Panicum and support recognition of Dichanthelium, Phanopyrum, and Steinchisma as distinct genera. We have transfered other species of Panicum to other genera of the Paniceae. Most of the necessary combinations have been made previously, so few nomenclatural changes have been required. The remaining species of Panicum sensu lato (s.l.) are included within Panicum incertae sedis representing isolated species or species grouped within monophyletic clades. Additionally, we explore the performance of the three codon position characters in producing the supported phylogeny.