The megadiverse genus Carex (c. 2000 species, Cyperaceae) has a nearly cosmopolitan distribution, displaying an inverted latitudinal richness gradient with higher species diversity in cold‐temperate areas of the Northern Hemisphere. Despite great expansion in our knowledge of the phylogenetic history of the genus and many molecular studies focusing on the biogeography of particular groups during the last few decades, a global analysis of Carex biogeography and diversification is still lacking. For this purpose, we built the hitherto most comprehensive Carex‐dated phylogeny based on three markers (ETS–ITS–matK), using a previous phylogenomic Hyb‐Seq framework, and a sampling of two‐thirds of its species and all recognized sections. Ancestral area reconstruction, biogeographic stochastic mapping, and diversification rate analyses were conducted to elucidate macroevolutionary biogeographic and diversification patterns. Our results reveal that Carex originated in the late Eocene in E Asia, where it probably remained until the synchronous diversification of its main subgeneric lineages during the late Oligocene. E Asia is supported as the cradle of Carex diversification, as well as a “museum” of extant species diversity. Subsequent “out‐of‐Asia” colonization patterns feature multiple asymmetric dispersals clustered toward present times among the Northern Hemisphere regions, with major regions acting both as source and sink (especially Asia and North America), as well as several independent colonization events of the Southern Hemisphere. We detected 13 notable diversification rate shifts during the last 10 My, including remarkable radiations in North America and New Zealand, which occurred concurrently with the late Neogene global cooling, which suggests that diversification involved the colonization of new areas and expansion into novel areas of niche space.
The goal of this study was to review the impact of DNA sequence analyses on our understanding of Cariceae phylogeny, classification and evolution. To explore character evolution, 105 taxa from four different studies were included in an nrDNA ITS + ETS 1f analysis of all recognized genera (Carex, Cymophyllus, Kobresia, Schoenoxiphium, Uncinia) and Carex subgenera (Carex, Psyllophora, Vignea, Vigneastra). As in previous analyses, four major Cariceae clades were recovered: (1) a "Core Carex Clade" (subg. Carex, Vigneastra, Psyllophora p.p); (2) A "Vignea Clade" (subg. Vignea, Psyllophora p.p.); (3) a "Schoenoxiphium Clade" (Schoenoxiphium, subg. Psyllophora p.p.), and (4) a "Core Unispicate Clade" (Uncinia, Kobresia, subg. Psyllophora p.p.). All studies provide strong support (86-100% BS) for the Core Carex and Vignea Clades, but only weak to moderate support (<50%-78% BS) for the Core Unispicate and Schoenoxiphium Clades. The relationships of these groups are unresolved. Studies suggest that Carex is either paraphyletic with respect to all Cariceae genera or to all genera except Schoenoxiphium. Kobresia is a grade, but Uncinia and possibly Schoenoxiphium are monophyletic. The monotypic Cymophyllus is indistinct from Carex subg. Psyllophora species. Character analyses indicate that inflorescence proliferation and reduction have occurred in all major clades, and that the Cariceae's unisexual flowers have evolved from perfect flowers. The ancestor to Cariceae possessed a multispicate inflorescence with cladoprophylls and female spikelets with tristigmatic gynoecia and closed utricles. This morphology is most similar to extant Carex subg. Carex species, which contradicts the nearly unanimous assumption that the highly compound inflorescences of Schoenoxiphium are primitive. Since taxonomic sampling and statistical support for phylogenies have generally been poor, we advocate the temporary maintenance of the four traditional Carex subgenera with androgynous unispicate species placed within subg. Psyllophora and dioecious and gynaecandrous unispicate species distributed amongst subgenera Carex and Vignea. A collective effort focused on developing new nuclear markers, on increasing taxonomic and geographic sampling, and on studying development within the context of phylogeny, is needed to develop a phylogenetic classification of Cariceae.
Wide speculation surrounds the origin and phylogenetic relationships of the most highly reduced sections in the genus Carex. In order to gain a better understanding of phylogeny in Carex, the relationship of the reduced sect. Phyllostachys to 12 putatively related sections, representing all four subgenera (Primocarex, Indocarex, Carex, Vignea), was inferred from sequences of the ITS (internal transcribed spacer) region of nrDNA. Phylogenetic reconstructions identified two main clades: (1) a "compound" clade composed of sections from subg. Indocarex, Primocarex, and a portion of subg. Carex, and (2) a "reduced" clade consisting of sections from subg. Carex (Phyllostachys) and Primocarex (Filifoliae and Firmiculmes). Subgenus Indocarex was paraphyletic within the "compound" clade supporting classifications that have merged it within a wider subg. Indocarex/Carex/Primocarex line. Subgenus Primocarex was polyphyletic. This result was consistent with theories that extreme reduction has occurred along several different evolutionary lines in Carex. Phylogenetic theories inferred from the presence or abnormal growth of the rachilla were not supported by tree topologies. Difficult sectional circumscriptions, such as the separation of sections Laxiflorae and Careyanae, were strongly upheld by sequence data. The ITS region is an effective tool for defining sectional limits and for estimating relationships among sections in Carex, but does not provide enough phylogenetic information to fully resolve relationships below the sectional level.
A population genetic approach was used to explore the evolutionary biology of the parasitic angiosperm Arceuthobium americanum Nutt. ex Engelm. (Viscaceae). Arceuthobium americanum infects three principal hosts and has the most extensive geographical range of any North American dwarf mistletoe. Based on the lack of apparent morphological and phenological differences between populations of A. americanum, past researchers have found no evidence for recognizing infraspecific taxa. In this study, molecular analysis using amplified fragment length polymorphism (AFLP) analysis indicated that A. americanum is divided into three distinct genetic races, each associated with a different host taxon in regions of allopatry: (i) Pinus banksiana in western Canada; (ii) Pinus contorta var. murrayana in the Sierra Nevada and Cascade Mountain ranges in the western US; and (iii) Pinus contorta var. latifolia in the western US and Canada. These observations suggest that host identity, geographical isolation and environmental factors have contributed to race formation in A. americanum. The lack of fine-scale patterning within each of the A. americanum races is attributed to random dispersal of seeds over long distances by animal vectors. Historical factors such as glaciations and founder events have also influenced structuring and genetic diversity in A. americanum populations. Given sufficient time, it is possible that these races will become reproductively isolated and undergo speciation.
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