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.
Asteraceae are the largest family of dicotyledonous plants and have long been known for their taxonomie complexity. The ubiquitous parallelisms in morphology within the family have made phylogenetic reconstruction and tribal circumscription an area of long debate. In this study we explored the utility of using two relatively short non-coding chloroplast DNA sequences, the trnL intron and IrnUtrnV intergenic spacer, to resolve phylogenetic relationships among the tribes. The results of the phylogenetic analysis produced trees that are topologically congruent with prior phylogenetic hypotheses based on both morphological and molecular data sets. The Asteroideae are a monophyletic group, but the Cichorioideae are paraphyletic. The primary clades of the Cichorioideae are the Mutisieae-Cardueae, Liabeae-Vernonieae, and of the Asteroideae, the Inuleae-Plucheeae, Astereae-Anthemideae, Senecioneae-Gnaphalieae, and the helianthoid clade (Helenieae, Heliantheae s.str.. and Eupatorieae). The Inuleae-Plucheeae clade is sister to the remainder of the Asteroideae. and the paraphyly of the Inuleae 8.1. (Gnaphalieae. Inuleae s. str., and Plucheeae) is firmly supported by our analysis. Our study illustrates the utility of the trnL intron and trnUV intergenic spacer for resolving relationships among tribes of the Asteraceae. Using approximately 874 bp, we were able to produce a phylogeny of comparable resolution to phylogenies based on well-known coding regions such as rbc\, and ndhY. For phylogenetic inference at the family level the trnL intron and trnUV spacer provide similar levels of resolution to longer coding sequences (e.g.. rbcL, ndhF), while having the advantage of being much easier to amplify and sequence due to their short lengths and universal primers. The numerous insertions and deletions commonly found in this region are easily aligned and are phylogenetically informative, thus adding considerably to the information content per base pair sequenced.
We investigate the species discriminatory power of a subset of the proposed plant barcoding loci (matK, rbcL, rpoC1, rpoB, trnH-psbA) in Carex, a cosmopolitan genus that represents one of the three largest plant genera on earth (c. 2000 species). To assess the ability of barcoding loci to resolve Carex species, we focused our sampling on three of the taxonomically bestknown groups in the genus, sections Deweyanae (6/8 species sampled), Griseae (18/21 species sampled), and Phyllostachyae (10/10 species sampled). Each group represents one of three major phylogenetic lineages previously identified in Carex and its tribe Cariceae, thus permitting us to evaluate the potential of DNA barcodes to broadly identify species across the tribe and to differentiate closely related sister species. Unlike some previous studies that have suggested that plant barcoding could achieve species identification rates around 90%, our results suggest that no single locus or multilocus barcode examined will resolve much greater than 60% of Carex species. In fact, no multilocus combination can significantly increase the resolution and statistical support (i.e., ³ 70% bootstrap) for species than matK alone, even combinations involving the second most variable region, trnH-psbA. Results suggest that a matK barcode could help with species discovery as 47% of Carex taxa recently named or resolved within cryptic complexes in the past 25 years also formed unique species clusters in UPGMA trees. Comparisons between the nrDNA internal transcribed spacer region (ITS) and matK in sect. Phyllostachyae suggest that matK not only discriminates more species (50-60% vs. 25%), but it provides more resolved phylogenies than ITS. Given the low levels of species resolution in rpoC1 and rpoB (0-13%), and difficulties with polymerase chain reaction amplification and DNA sequencing in rbcL and trnH-psbA (alignment included), we strongly advocate that matK should be part of a universal plant barcoding system. Although identification rates in this study are low, they can be significantly improved by a regional approach to barcoding.
Phylogenetic reconstruction for Carex and relatives in tribe Cariceae is complicated by species richness and nearly cosmopolitan distribution. In this investigation, our main objective was to estimate evolutionary relationships in tribe Cariceae using DNA sequence data from two spacer regions in nuclear ribosomal genes (ITS and ETS-1f) combined with noncoding chloroplast DNA (trnL intron, trnL-trnF intergenic spacer, and trnE-trnD intergenic spacers). Parsimony analyses of separate and combined data and Bayesian analysis of the combined data matrix revealed strong support for monophyly of tribe Cariceae and for monophyly of two major lineages, one comprising principally Carex subgen. Carex and Vigneastra, and the other representing subgen. Vignea. A third clade with representatives from Kobresia and Uncinia, along with Cymophyllus fraserianus, Carex curvula, and several unispicate Carex received weak-to-moderate support. A small clade comprising Schoenoxiphium and two unispicate carices was placed as sister to the clades comprising multispicate Carex species in the parsimony analysis, but sister to the clade of Kobresia, Uncinia, and unispicate Carex in the Bayesian analysis. Two large widespread groups within subgen. Carex, sect. Hymenochlaenae and sect. Physocarpae s.l. (''bladder sedges''), were highly polyphyletic, while ten clades that grouped species from two or more sections were each strongly supported as monophyletic. Within subgen. Vignea, three sections were strongly supported as monophyletic while sects. Phaestoglochin and Vulpinae were polyphyletic. Adding the variable ETS-1f region improved resolution and bootstrap support values over previous studies, but many of the characters supporting major branches came from the trnL region.
Aim Across angiosperm families, the area occupied by a family is strongly correlated with its richness. We explore the causes of this area‐richness correlation using the cosmopolitan family, Cyperaceae Juss., as a model. We test the hypothesis that, despite a proposed tropical origin, temperate lineages in the family diversified at elevated rates. We test the hypothesis that the area‐richness correlation is maintained within intrafamilial clades, and that this relationship could be described as a function of niche space. We also test the hypothesis that the partitioning of geographical and ecological space, not the extent of this space, is the factor most closely associated with clade richness. Location Cosmopolitan. Methods We use molecular data from four genes sequenced in 384 taxa to develop a chronogram of Cyperaceae. We then develop a model of ancestral ranges and measure rates of diversification throughout the history of the family. Integrating data from over 4,800,000 digitized herbarium records, we characterize the range and niche of more than 4500 species and test for correlations of the species richness maintained within clades with range size, range partitioning, range overlap, niche, clade age and rate of diversification. Results Cyperaceae originated in South America in the late Cretaceous and subsequently dispersed throughout the globe. Of three increases in diversification rate, two occurred in the temperate Northern Hemisphere. The variable most closely associated with clade richness is the partitioning of geographical space by species within each clade. Main conclusions We show that species‐rich clades in Cyperaceae are not only more widespread, occupy more niche space, and diversify more quickly, but also exhibit patterns that are consistent with the partitioning of geographical and ecological space as a major correlate to diversification.
Since the Monocots II meeting in 1998, significant new data have been published that enhance our systematic knowledge of Cyperaceae. Phylogenetic studies in the family have also progressed steadily. For this study, a parsimony analysis was carried out using all rbcL sequences currently available for Cyperaceae, including data for two new genera. One of the four subfamilies (Caricoideae) and seven of the 14 tribes (Bisboeckelereae, Cariceae, Cryptangieae, Dulichieae, Eleocharideae, Sclerieae, Trilepideae) are monophyletic. Subfamily Mapanioideae and tribe Chrysitricheae are monophyletic if, as the evidence suggests, Hellmuthia is considered a member of Cypereae. Some other features of our analysis include: well-supported Trilepideae and Sclerieae-Bisboeckelereae clades; a possible close relationship between Cryptangieae and Schoeneae; polyphyletic tribes Schoeneae and Scirpeae; the occurrence of Cariceae within the Dulichieae-Scirpeae clade, and a strongly supported clade, representing Cyperus and allied genera in Cypereae, sister to a poorly supported Ficinia-HellmuthiaIsolepis-Scirpoides clade. Such patterns are consistent with other studies based on DNA sequence data. One outcome may be that only two subfamilies, Mapanioideae and Cyperoideae, are recognized. Much further work is needed, with efforts carefully coordinated among researchers. The work should focus on obtaining morphological and molecular data for all genera in the family.
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