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.
Phylogenetic studies of Carex L. (Cyperaceae) have consistently demonstrated that most subgenera and sections are para-or polyphyletic. Yet, taxonomists continue to use subgenera and sections in Carex classification. Why? The Global Carex Group (GCG) here takes the position that the historical and continued use of subgenera and sections serves to (i) organize our understanding of lineages in Carex, (ii) create an identification mechanism to break the~2000 species of Carex into manageable groups and stimulate its study, and (iii) provide a
Estimating species ability to adapt to environmental changes is crucial to understand their past and future response to climate change. The Mediterranean Basin has experienced remarkable climatic changes since the Miocene, which have greatly influenced the evolution of the Mediterranean flora. Here, we examine the evolutionary history and biogeographic patterns of two sedge sister species (Carex, Cyperaceae) restricted to the western Mediterranean Basin, but with Pliocene fossil record in central Europe. In particular, we estimated the evolution of climatic niches through time and its influence in lineage differentiation. We carried out a dated phylogenetic-phylogeographic study based on seven DNA regions (nDNA and ptDNA) and fingerprinting data (AFLPs), and modelled ecological niches and species distributions for the Pliocene, Pleistocene and present. Phylogenetic and divergence time analyses revealed that both species form a monophyletic lineage originated in the late Pliocene-early Pleistocene. We detected clear genetic differentiation between both species with distinct genetic clusters in disjunct areas, indicating the predominant role of geographic barriers limiting gene flow. We found a remarkable shift in the climatic requirements between Pliocene and extant populations, although the niche seems to have been relatively conserved since the Pleistocene split of both species. This study highlights how an integrative approach combining different data sources and analyses, including fossils, allows solid and robust inferences about the evolutionary history of a plant group since the Pliocene.
Carex section Phacocystis (Cyperaceae) is one of the most diverse and taxonomically complex groups of sedges (between 116 and 147 species), with a worldwide distribution in a wide array of biomes. It has a very complicated taxonomic history, with numerous disagreements among different treatments. We studied the biogeography and niche evolution in a phylogenetic framework to unveil the relative contribution of geographical and ecological drivers to diversification of the group. We used a large species sampling of the section (82% of extant species) to build a phylogeny based on four DNA regions, constrained with a phylogenomic HybSeq tree and dated with six fossil calibrations. Our phylogenetic results recovered section Phacocystis s.s. (core Phacocystis) as sister to section Praelongae. Ancestral area reconstruction points toward the N Pacific as the cradle for the crown diversification of section Phacocystis during the Middle Miocene. Wide distributions were recurrently inferred across deep nodes. Large Northern Hemisphere lineages with geographical congruence were retrieved, pointing toward the importance of allopatric divergence at deep phylogenetic levels, whereas withinarea speciation emerges as the predominant pattern at shallow phylogenetic level. The Southern Hemisphere (Neotropics, SW Pacific) was colonized several times from the Northern Hemisphere. The global expansion of Carex section Phacocystis did not entail major ecological changes along the inner branches of the phylogeny. Nevertheless, ecological differentiation seems to gain importance toward recent times.
Plants growing in high-mountain environments may share common morphological features through convergent evolution resulting from an adaptative response to similar ecological conditions. The Carex flava species complex (sect. Ceratocystis, Cyperaceae) includes four dwarf morphotypes from Circum-Mediterranean mountains whose taxonomic status has remained obscure due to their apparent morphological resemblance. In this study we investigate whether these dwarf mountain morphotypes result from convergent evolution or common ancestry, and whether there are ecological differences promoting differentiation between the dwarf morphotypes and their taxonomically related large, well-developed counterparts. We used phylogenetic analyses of nrDNA (ITS) and ptDNA (rps16 and 5’trnK) sequences, ancestral state reconstruction, multivariate analyses of macro- and micromorphological data, and species distribution modeling. Dwarf morphotype populations were found to belong to three different genetic lineages, and several morphotype shifts from well-developed to dwarf were suggested by ancestral state reconstructions. Distribution modeling supported differences in climatic niche at regional scale between the large forms, mainly from lowland, and the dwarf mountain morphotypes. Our results suggest that dwarf mountain morphotypes within this sedge group are small forms of different lineages that have recurrently adapted to mountain habitats through convergent evolution.
Premise Most of the Paleotropical flora widely distributed in the Western Palearctic became extinct during the Mio‐Pliocene as a result of global geoclimatic changes. A few elements from this Cenozoic flora are believed to remain as relicts in Macaronesia, forming part of the laurel forests. Although the origins of the present species assembly are known to be heterogeneous, it is unclear whether some species should be considered climatic relicts with conserved niches. An ideal group for studying such relict characteristics is a Miocene lineage of Carex sect. Rhynchocystis (Cyperaceae), which comprises four species distributed in mainland Palearctic and Macaronesia. Methods We reconstructed the current and past environmental spaces for extant mainland and Macaronesian species, as well as for Pliocene fossils. We also studied the bioclimatic niche evolution. Species distribution modeling and ensemble small modeling were performed to assess the potential distribution over time. Results Climatic niche analyses and distribution modeling revealed that the ecological requirements of Macaronesian species did not overlap with those of either mainland species or with the Pliocene fossils. Conversely, the niches of mainland species displayed significant similarity and equivalence. Conclusions Macaronesian species are not climatic relicts from the Paleotropical flora, but instead seem to have changed the ecological niche of their ancestors. By contrast, despite their ancient divergence (Late Miocene), mainland C. pendula and C. agastachys show conserved niches, with competitive exclusion likely shaping their mostly allopatric ranges.
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.
Carex section Schoenoxiphium (Cariceae, Cyperaceae) is endemic to the Afrotropical biogeographic region and is mainly distributed in southern and eastern Africa, with its center of diversity in eastern South Africa. The taxon was formerly recognized as a distinct genus and has a long history of taxonomic controversy. It has also an important morphological and molecular background in particular dealing with the complexity of its inflorescence and the phylogenetic relationships of its species. We here present a fully updated and integrative monograph of Carex section Schoenoxiphium based on morphological, molecular and cytogenetic data. A total of 1,017 herbarium specimens were examined and the majority of the species were studied in the field. Previous molecular phylogenies based on Sanger-sequencing of four nuclear and plastid DNA regions and RAD-seq were expanded. For the first time, chromosome numbers were obtained, with cytogenetic counts on 44 populations from 15 species and one hybrid. Our taxonomic treatment recognizes 21 species, one of them herein newly described (C. gordon-grayae). Our results agree with previous molecular works that have found five main lineages in Schoenoxiphium. We provide detailed morphological descriptions, distribution maps and analytical drawings of all accepted species in section Schoenoxiphium, an identification key, and a thorough nomenclatural survey including 19 new typifications and one nomen novum.
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