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Knowledge of Carex L. (true sedges) and Hypericum L. (St. John's wort) in the Neotropics is fragmentary. As a result of a fieldwork campaign in Ecuador and revision of herbarium collections (K, QCA and QCNE), we present here relevant records of twelve Carex (Cyperaceae) and four Hypericum (Hypericaceae) species. Regarding Carex, we present the novel report for South America of C. aztecica, as well as the first Ecuadorian records for C. brehmeri, C. collumanthus, C. fecunda, C. melanocystis and C. punicola. The three later records have additional biogeographical significance, as they represent the new northern limit of these species. We also include observations for another five species included in the Ecuadorian Red List of Endemic Plants. As a result, the list of native Carex reported for Ecuador would now include 52 taxa. With regard to Hypericum, we include the new report of H. sprucei for the province of Bolívar, and the confirmation of the presence of three rare species (H. acostanum, H. matangense, H. prietoi) in their type localities, although with extremely low population sizes. We discuss their conservation status and implications.

Relevant chorological notes of genus Carex L. (Cyperaceae) for the flora of the Balkans are provided, with an emphasis on Albania and adjacent countries (North Macedonia and Montenegro). Our findings include new national species records and/or confirmations for Albania, Montenegro and North Macedonia (C. agastachys, C. atrata, C. curvula, C. demissa, C. hispida, C. parviflora), as well as other interesting records of rare and/or endangered Carex species in Albania (C. castroviejoi, C. myosuroides). Eventually, we provide relevant comments in order to clarify the taxonomy, distribution and/or ecology of Carex sections Rhynchocystis (C. agastachys, C. pendula) and Aulocystis (C. ferruginea, C. kitaibeliana, C. lazarei) in the Balkans.

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.

Despite centuries of work, the basic taxonomic knowledge of the flora of the Iberian Peninsula is still incomplete, especially for highly diverse and/or difficult genera such as Carex. In this study, we conducted an integrative systematic study based on molecular, morphological and cytogenetic data to elucidate the taxonomic status of several problematic Carex populations from La Mancha region (S Spain) belonging to Carex sect. Phacocystis. These populations have been traditionally considered of uncertain taxonomic adscription, but close to C. reuteriana due to their morphological appearance and ecological preferences. A detailed morphological and cytogenetic study was performed on 16 La Mancha’s problematic populations (Sierra Madrona and Montes de Toledo) to compare them with the other Iberian sect. Phacocystis species. In addition, a phylogenetic analysis was conducted using two nuclear (ITS, ETS) and two plastid (rpl32‐trnLUAG, ycf6‐psbM) DNA regions, including representatives from all species of sect. Phacocystis. We found a significant degree of molecular and morphological differentiation that supports the recognition of La Mancha’s problematic populations as a new Iberian endemic species, described here as Carex quixotiana Ben.Benítez, Martín-Bravo, Luceño & Jim.Mejías. Our results reveal that C. quixotiana, unexpectedly, is more closely related to C. nigra than to C. reuteriana on the basis of phylogenetic relationships and chromosome number. These contrasting patterns reflect the taxonomic complexity in sect. Phacocystis and highlight the need for integrative systematic approaches to disentangle such complicated evolutionary scenarios.

In this contribution, new data concerning the distribution of native vascular flora in Italy are presented. It includes new records, confirmations, exclusions, and status changes to the Italian administrative regions. Nomenclatural and distribution updates, published elsewhere, and corrigenda are provided as Suppl. material 1.

— Populations of Carex sect. Rhynchocystis (Cyperaceae) from the Macaronesian archipelagos (Azores and Madeira) have traditionally been treated either as a variety of the widely distributed Western Palearctic C. pendula, or directly synonymized under it. However, recent phylogenetic studies have shown that Azorean populations of C. pendula display a certain degree of differentiation from mainland plants, while the phylogenetic relationships of Madeiran populations remain unclear. Here we perform an integrated systematic study focused on the Macaronesian populations of Carex sect. Rhynchocystis to elucidate their phylogenetic relationships and taxonomic status. We reconstructed a molecular phylogeny based on five DNA regions and conducted a multivariate morphological analysis. Divergence time estimates show that the Macaronesian populations can be traced back to a Plio-Pleistocene origin. Our results suggest that these island populations of C. pendula are better treated as two distinct species within Carex sect. Rhynchocystis (i.e. C. leviosa from the Azores and C. sequeirae from Madeira). We provide morphological characters to differentiate the new species from C. pendula s. s., detailed descriptions of the three taxa, a revised key for the entire section, as well as detailed analytical drawings of the two newly described species. We also perform a critical evaluation of the taxonomic diversity of Carex in the Azores and Madeira. Finally, we informally assessed the conservation status of the new species at a global scale under IUCN categories and criteria, resulting in the proposal of the categories Least Concern for C. leviosa and Critically Endangered for C. sequeirae.

Publisher Copyright: © This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited In this contribution, new data concerning the distribution of vascular flora alien to Italy are presented. It includes new records, confirmations, exclusions, and status changes for Italy or for Italian administrative regions. Nomenclatural and distribution updates published elsewhere are provided as Suppl. material 1.

Carex microcarpa (sect. Rhynchocystis) is a Central Mediterranean endemic known from Corsica, Sardinia and the Tuscan archipelago, that has been also reported from scattered localities in mainland Italy (Abruzzo, Lazio and Apulia). We tested by molecular means the identification of three specimens classified as C. microcarpa from continental Italy, as well as a voucher from the Montecristo island (Tuscany) and one from Sardinia, whose taxonomic identity could not be determined with certainty by morphological examination. We performed a DNA barcoding study based on four DNA regions (two nuclear, two plastid). Our results reveal that none of previous mainland Italian records corresponds to C. microcarpa, but to C. pendula, which also belong to section Rhynchocystis, and in other cases possibly to C. acutiformis, from a different species group (sect. Paludosae). The specimen from Montecristo island was confirmed to be C. microcarpa. We conclude that C. microcarpa is likely a Tyrrhenian endemic exclusively known with certainty in Corsica, Sardinia and the Tuscan Archipelago. The presence of C. microcarpa in the Italian peninsula remains highly doubtful and we recommend its exclusion from checklists. Our study highlights the utility of DNA barcoding approaches in Carex when a reliable morphology-based identification is not possible. Supplemental data for this article is available online at https://doi.org/10.1080/11263504.2021.1897703 .

—We describe a new remarkable dwarf and apparently acaulescent species of Carex (Cyperaceae) from the Andes of northern Chile: Carex phylloscirpoides. Morphological and molecular data (two nuclear and three plastid DNA regions) were used to study the phylogenetic placement and systematic relationships of this species, which resulted in its assignment to Carex section Racemosae. However, despite being related to the other three species of the section present in the Southern Cone based on phylogenetic evidence, it displays a number of unusual morphological features. In addition, it is geographically disjunct and genetically differentiated from them. A preliminary conservation assessment was performed which resulted in the proposal of the Critically Endangered category at the global level under IUCN guidelines. We discuss the biogeographic and ecological peculiarities of the new species and its allies, with emphasis in the high number of acaulescent sedges present in South America. The features of C. phylloscirpoides emphasize its unique evolutionary position and its rarity highlights its conservation importance, especially in the context of the South American Andean flora.

This is the first work of a series of thorough studies into the taxonomy and systematics of Carex in South America. Here, we present the systematic placement, biogeographic insights, taxonomic accounts, formal typifications and summarized distributions for the 24 species of Carex subgenus Vignea with androgynous spikes in the continent. We performed a phylogenetic study using the barcode markers ETS, ITS and matK to check the placement of 22 of these species on a previous tree with > 1000 Carex spp. We examined > 600 specimens from 38 herbaria and conducted an exhaustive nomenclatural survey consulting all pertinent literature. South American androgynous species of subgenus Vignea are grouped in seven distinct lineages, corresponding with the same number of inferred colonizations. Most groups seem to have colonized South America during the Plio-Pleistocene, except for species of section Bracteosae, a local radiation that dates back to the Late Miocene. All colonizations by native species seem to have originated from North America. The two putatively introduced taxa are western Palaearctic species. An identification key is provided for the studied species. We make 19 new typifications and a new species (Carex pedicularis) is described.

Aim: Madagascar is renowned for its unparalleled biodiversity and endemism. With many ecosystems under threat, research is urgently needed on its unique plant diversity. This applies both to Madagascar's forests and treeless vegetation types. Sedges (Cyperaceae) are among the top 10 species-richest angiosperm families in Madagascar (310 native species, 38% endemic), of which two-thirds occur in open habitats. We aimed to infer the evolutionary history of sedges in Madagascar, by estimating the number, age and origins of endemic lineages, and how they diversified on the island. We tested contrasting hypotheses of (a) few colonizations but important in situ radiations against (b) a high number of anagenetic colonizations. Location: Madagascar and the surrounding Indian Ocean islands, integrated within a global dataset. Taxon: Sedge family Cyperaceae. Methods: We estimated time-calibrated molecular phylogenies encompassing a large proportion of Madagascar's known sedge flora (incl. 55% of native species), integrating sequence data for 1,382 accessions representing almost 25% of the c. 5,600 sedge species worldwide, combined with ancestral area reconstruction, diversification analyses and Bayesian stochastic mapping. Results: Cyperaceae lineages arrived in Madagascar from c. 40 Mya with many arriving more recently. About 20 endemic lineages of Cyperaceae occur on the island, of which only six encompass more than five species. All except one of the endemic lineages that diversified in Madagascar use the C3 photosynthetic pathway. The main biogeographical links of Madagascar's sedge flora are to Southern and Tropical Africa. Main conclusions: The biogeographical history of Cyperaceae in Madagascar is a chronicle of relatively recent multiple in and out processes of long-distance dispersal colonizations constrained by distance. Also, the Madagascar region is not only a ‘sink’ for immigrant taxa, in situ diversification and dispersal to other regions also occurred. Some of the most diverse endemic lineages show clear adaptation to local environments.

A morphological and molecular study was carried out on two species of Carex from the former genus Uncinia from mainland South America and Juan Fernández Islands: C. firmula (Kük) J.R. Starr and C. subsacculata (G.A.Wheeler & Goetgh.) J.R.Starr. Both species share the particular character of having articulated pistillate scales that leave behind on the spike rachis a saccate stipe-like appendage after falling. We detected within C. firmula morphological differences between mainland South American specimens and Juan Fernández specimens, mainly regarding flower density and utricle size. On the contrary, the molecular analysis showed no molecular differentiation between the two groups of populations. Whereas, strong molecular and morphological support were found for C. subsacculata. As such, we recommend that the populations of C. firmula found on Juan Fernández and those found in the mainland should be recognised as different subspecies: island Carex firmula subsp. firmula and mainland C. firmula subsp. tenuis (Poepp. ex Kunth) R.Ridley & Jim.-Mejías. We discuss that the morphological differences may be a result of isolation in an oceanic island environment and have effect on seed dispersal. Taxonomic descriptions, images of diagnostic characters and a key are included.

The field of systematics is experiencing a new molecular revolution driven by the increased availability of high-throughput sequencing technologies. As these techniques become more affordable, the increased genomic resources have increasingly far-reaching implications for our understanding of the Tree of Life. With c. 2000 species, Carex (Cyperaceae) is one of the five largest genera of angiosperms and one of the two largest among monocots, but the phylogenetic relationships between the main lineages are still poorly understood. We designed a Cyperaceae-specific HybSeq bait kit using transcriptomic data of Carex siderosticta and Cyperus papyrus. We identified 554 low-copy nuclear orthologous loci, targeting a total length of c. 1 Mbp. Our Cyperaceae-specific kit shared loci with a recently published angiosperm-specific Anchored Hybrid Enrichment kit, which enabled us to include and compile data from different sources. We used our Cyperaceae kit to sequence 88 Carex spp., including samples of all the five major clades in the genus. For the first time, we present a phylogenetic tree of Carex based on hundreds of loci (308 nuclear exon matrices, 543 nuclear intron matrices and 66 plastid exon matrices), demonstrating that there are six strongly supported main lineages in Carex: the Siderostictae, Schoenoxiphium, Unispicate, Uncinia, Vignea and Core Carex clades. Based on our results, we suggest a revised subgeneric treatment and provide lists of the species belonging to each of the subgenera. Our results will inform future biogeographic, taxonomic, molecular dating and evolutionary studies in Carex and provide the step towards a revised classification that seems likely to stand the test of time.

Sedges (Cyperaceae) are commonly pollinated by wind, but some transitions from wind to insect pollination have been reported in tropical sedges. Despite being a temperate sedge, the pollination strategy of Rhynchospora alba may partly depend on insects, as suggested by its white-colored inflorescences. Here we quantify the contribution of insects to the reproductive fitness of R. alba as an exploration of the success of entomophily in temperate sedges. Honeybees and hoverflies seem to be the primary visitors of R. alba in the study site, and pollen of R. alba was recovered from the mouthparts and legs of the insects. Treatments excluding insects led to a lower reproductive fitness (regarding fruit weight and fruit viability) than those not excluding them. Therefore, insects seem to contribute significantly to the pollination of R. alba despite moderate morphological adaptation. We highlight the role of entomophily as an overlooked reproductive mechanism in cold-temperate sedges.

We identify specimens belonging to the original material of the names Onosmodium molle Michaux (Boraginaceae) and Verbena riparia Raf. ex Small & Heller (Verbenaceae). We designate the lectotype for O. molle. On the other hand, we present evidence that original material of V. riparia is indeed the holotype, which supersedes a previous lectotype designation.

Carex phleoides Cav. is the largest species of the former genus Uncinia Pers. It is widely distributed in the Neotropics and is also known from the Juan Fernandez archipelago off the coast of Chile. A second taxon, U. trichocarpa C. A. Mey., is often considered a synonym of C. phleoides, but also sometimes a variety of that species or even a distinct species. We performed a morphometric study using representative material covering the entire distribution and morphological variation of C. phleoides s.l. to clarify its taxonomy. Our analyses revealed that U. trichocarpa is distinct from C. phleoides. Accordingly, we coined the name C. quinquin Jim.-Mejias & Dorr to accommodate this species under Carex L. because combinations of other available names are all blocked. Carex phleoides displayed two extremes of variation with geographical correspondence, a southern form in the Southern Cone north to Bolivia, and a northern one ranging from Bolivia to Mexico. We propose to recognize two subspecies within C. phleoides, the nominate subspecies being the southern one, and the northern one based on the epithet koyamae. Plants from the Juan Fernandez archipelago were shown to be morphologically intermediate with the endemic C. plurinervata J. R. Starr, a result that might point to introgression between these two species. We selected lectotypes for nine names, including two of the accepted ones, as well as their synonyms.

International audience; Molecular phylogenetic studies based on Sanger sequences have shown that Cyperaceae tribe Fuireneae s.l. is paraphyletic. However, taxonomic sampling in these studies has been poor, topologies have been inconsistent, and support for the backbone of trees has been weak. Moreover, uncertainty still surrounds the morphological limits of Schoenoplectiella, a genus of mainly small, amphicarpic annuals that was recently segregated from Schoenoplectus. Consequently, despite ample evidence from molecular analyses that Fuireneae s.l. might consist of two to four tribal lineages, no taxonomic changes have yet been made. Here, we use the Angiosperms353 enrichment panel for targeted sequencing to (i) clarify the relationships of Fuireneae s.l. with the related tribes Abildgaardieae, Eleocharideae, and Cypereae; (ii) define the limits of Fuireneae s.s., and (iii) test the monophyly of Fuireneae s.l. genera with emphasis on Schoenoplectus and Schoenoplectiella. Using more than a third of Fuireneae s.l. diversity, our phylogenomic analyses strongly support six genera and four major Fuireneae s.l. clades that we recognize as tribes: Bolboschoeneae stat.nov., Fuireneae s.s., Schoenoplecteae, and Pseudoschoeneae tr. nov. These results are consistent with morphological, micromorphological (nutlet epidermal cell shape), and embryo differences detected for each tribe. At the generic level, most sub-Saharan African perennials currently treated in Schoenoplectus are transferred to Schoenoplectiella. Our targeted sequencing results show that these species are nested in Schoenoplectiella, and their treatment here is consistent with micromorphological and embryo characters shared by all Schoenoplectiella species. Keys to recognized tribes and genera are provided.

Cyperaceae (sedges) are the third largest monocot family and are of considerable economic and ecological importance. Sedges represent an ideal model family to study evolutionary biology because of their species richness, global distribution, large discrepancies in lineage diversity, broad range of ecological preferences, and adaptations including multiple origins of C4 photosynthesis and holocentric chromosomes. Goetghebeur’s seminal work on Cyperaceae published in 1998 provided the most recent complete classification at tribal and generic level, based on a morphological study of Cyperaceae inflorescence, spikelet, flower and embryo characters plus anatomical and other information. Since then, several family‐level molecular phylogenetic studies using Sanger sequence data have been published. Here, more than 20 years after the last comprehensive classification of the family, we present the first family‐wide phylogenomic study of Cyperaceae based on targeted sequencing using the Angiosperms353 probe kit sampling 311 accessions. Additionally, 62 accessions available from GenBank were mined for overlapping reads and included in the phylogenomic analyses. Informed by this backbone phylogeny, a new classification for the family at the tribal, subtribal and generic levels is proposed. The majority of previously recognized suprageneric groups are supported, and for the first time we establish support for tribe Cryptangieae as a clade including the genus Koyamaea. We provide a taxonomic treatment including identification keys and diagnoses for the 2 subfamilies, 24 tribes and 10 subtribes and basic information on the 95 genera. The classification includes five new subtribes in tribe Schoeneae: Anthelepidinae, Caustiinae, Gymnoschoeninae, Lepidospermatinae and Oreobolinae. This article is protected by copyright. All rights reserved.

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 Macaronesian Laurel Forest. 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 to study such relict characteristics is the Miocene lineage of Carex sect. Rhynchocystis, which comprises four species distributed in the mainland Palearctic and Macaronesia. METHODS We reconstructed the current and past environmental spaces for extant mainland and Macaronesian species, and for the occurrence of 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. KEY RESULTS 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. The ecological differentiation within Macaronesia and similarity across the continent were evaluated in bioclimatic evolution and distribution modeling analyses. CONCLUSIONS Macaronesian species are not climatic relicts from the Paleotropical flora, but instead seems have changed the ecological niche from 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. This article is protected by copyright. All rights reserved.

Here, we present relevant records of three species that are new at national level (Carex jamesonii Boott and C. pseudomacloviana G.A.Wheeler for Bolivia, and C. ruthsatziae G.A.Wheeler for Argentina), six new regional records for Bolivia (C. acutata Boott, C. bonariensis Desf. ex Poir., C. crassiflora Kuk., C. feddeana H.Pfeiff., C. fuscula d’Urv., and C. pachamamae Jim.Mejias & Reznicek), and additional records of rare plants (C. pachamamae and C. ruthsatziae). We describe C. giovanniana Jim.Mejias as a new species previously misidentified as C. subdivulsa (Kuk.) G.A.Wheeler. In addition, we perform the typification for four names (C. acutata, C. crassiflora, C. jamesonii, and C. tessellata Spruce ex C.B.Clarke). Comments on the taxonomy of C. ruthsatziae are provided. We synonymise C. obtusisquama (Gross) G.A.Wheeler & S.Beck to C. crassiflora, and C. tessellata to C. acutata.

Geography and climate have been the main drivers of evolution in recent geological epochs. While new lineages of species have been formed in the last millions of years (speciation) and others have vanished as a result of historical climate changes (extinction), some ancient lineages appear to have persisted to the present day without net diversification. In this paper, evolution of ancient lineages is addressed by combining phylogenetic and conservation approaches to test the concept of ‘endangered living fossil’ (ELF). Using endangered, monospecific genera as starting point, we propose three criteria to identify ELFs (in order): (1) scarcity and narrow distribution of populations, i.e. the species (and thus the genus) is categorised as either ‘endangered’ or ‘critically endangered’ using IUCN criteria; (2) evolutionary distinctiveness, i.e. phylogenetic singularity of a single-species lineage as a result of a null net diversification rate; (iii) ancient divergence, i.e. split from the closest extant relatives predating the dramatic climate changes of particular geological epochs (specifically changes since the Miocene-Pliocene boundary). The vascular flora of the Iberian Peninsula offers a suitable study system to reliably test the ELF concept. Indeed, time-calibrated phylogenies revealed that five of the six critically endangered, monospecific genera endemic to the Iberian Peninsula are ELFs. These five genera appear to have diverged from their closest relatives in the Oligocene (Gyrocaryum), Miocene (Avellara, Castrilanthemum, Gadoria) and around the Miocene-Pliocene boundary (Naufraga). This result entails long-term survival (with no net diversification) through at least three dramatic climate changes: the Messinian Salinity Crisis (late Miocene), the establishment of the mediterranean climate (Pliocene), and the glacial-interglacial cycles (Pleistocene). Using results from the literature, we found examples of ELFs for the mediterranean floras of California (Dodecahema), Chile (Avellanita, Gomortega, Legrandia) and other Mediterranean areas of Europe (Petagnaea, Phitosia). ELFs are unique and threatened lineages representing an exceptional evolutionary heritage, and therefore they should be prioritised in biodiversity research and conservation programs.

Aim To evaluate how Cenozoic climate changes shaped the evolution and distribution of Carex section Rhynchocystis. Location Western Palaearctic and Afrotropical regions (Rand Flora pattern). Methods DNA regions ITS, ETS (nuclear), matK and rpl32-trnLUAG (plastid) were amplified for 86 samples of species from section Rhynchocystis. Phylogenetic and phylogeographical relationships were inferred using maximum parsimony, Bayesian inference and coalescent-based species tree approaches. Divergence times and ancestral areas were also inferred. Results Carex section Rhynchocystis is a clade that diversified during the middle Miocene in Europe. Most cladogenesis events date to the middle and late Miocene. The Afrotropical group seems to have originated from a colonization event from Europe that occurred in the late Miocene. Main conclusions Species of the section Rhynchocystis in the Western Palaearctic are Miocene relicts. Late Miocene-Pliocene aridification of the Mediterranean rather than the more commonly reported Pleistocene glaciations seems to have shaped the phylogeography of the group. Putative Miocene-Pliocene refugia were probably located in the Mediterranean peninsulas and islands, as well as in the eastern shores of the ancient Paratethys Sea. The colonization of Africa could have been facilitated by Miocene-Pliocene global cooling.

Notas taxonómicas y de distribución de Carex (Cyperaceae) en el Neotrópico. Presentamos datos relevantes acerca de la taxonomía y distribución de 20 especies del género Carex en el Neotrópico(de la region Caribe a la Patagonia). Aportamos un total de 19 registros nuevos, dos de ellos para especies hasta el momento no registradas para Sudamérica, diez especies citadas como novedad para diferentes países y siete especies aquí reportadas como novedades regionales en Argentina, Bolivia y Chile. Lectotipificamos de nueve nombres de especies (C. brehmeri, C. catharinensis, C. ecuadorica, C. firmicaulis, C. fuscula, C. mandoniana, C. phleoides, C. polysticha y C. subantarctica) y epitipificamos uno de ellos (C. fuscula). Finalmente, proponemos a Carex koyamae como nuevo sinónimo de Carex phleoides.

On the basis of previously published molecular data, and morphological observations, Carex nigra subsp. drukyulensis from the eastern Himalayas (Bhutan) is raised to specific rank as Carex drukyulensis.

Premise of the Study: Long-distance dispersal (LDD) syndromes, especially endozoochory, facilitate plant colonization of new territories that trigger diversification. However, few studies have analyzed how epizoochorous fruits influence both range distribution and diversification rates. We examined the evolutionary history of a hyperdiverse clade of Boraginaceae (subfamily Cynoglossoideae, eight tribes, ~60 genera, ~1100 species) and the evolution of fruit traits. We evaluated the evolutionary history of diaspore syndromes correlated with geographic distribution and diversification rates over time. Methods: Plastid DNA regions and morphological traits associated with dispersal syndromes were analyzed for 71 genera (226 species). We employed trait-dependent diversification analysis (HiSSE) and biogeographic reconstruction (Lagrange) using a time-calibrated phylogeny. Key Results: Our results indicate that (1) the earliest divergence events in Cynoglossoideae occurred in the central-northeastern Palearctic during the Paleogene (early to middle Eocene); (2) an epizoochorous trait (specialized hooks named glochids) is ancestral and has been maintained long term; and (3) glochids are correlated with increased diversification rates in two distantly related clades (Rochelieae and Cynoglossinae). Rapid speciation occurred for these two groups in the same area (central-eastern Palearctic) and same period (Oligocene-Miocene: Rochelieae, 30.82–13.69 mya; Cynoglossinae, 33.10–15.21 mya). Lower diversification rates were inferred for the remaining four glochid-bearing clades. Conclusions: One more example of “biogeographic congruence” in angiosperms is supported by a shared geographic (central-northeastern Palearctic) and temporal (28.60–21.59 mya, late Oligocene) opportunity window for two main clades’ diversification. Epizoochorous traits (fruit glochids) had an effect in higher diversification rates only with the joint effect of other unmeasured factors.

Biogeographic dispersal is supported by numerous phylogenetic results. In particular, transoceanic dispersal, rather than vicariance, is suggested for some plant lineages despite current long distances between America and Europe. However, few studies on the biogeographic history of plants have also studied the role of diaspore syndromes in long-distance dispersal (LDD). Species of the tribe Omphalodeae (Boraginaceae) offer a suitable study system because the species have a wide variety of diaspore traits related to LDD and different lineages conform to patched worldwide distributions on three distant continents (Europe, America and New Zealand). Our aim is to reconstruct the biogeographical history of the Omphalodeae and to investigate the role of diaspore traits favoring LDD and current geographic distributions. To this end, a time-calibrated phylogeny with 29 of 32 species described for Omphalodeae was reconstructed using biogeographical analyses (BioGeoBEARS, Lagrange) and models (DEC and DIVA) under different scenarios of land connectivity. Character-state reconstruction (SIMMAP) and diversification rate estimations of the main lineages were also performed. The main result is that epizoochorous traits have been the ancestral state of LDD syndromes in most clades. An early diversification age of the tribe is inferred in the Western Mediterranean during late Oligocene. Colonization of the New World by Omphalodeae, followed by fast lineage differentiation, took place sometime in the Oligocene-Miocene boundary, as already inferred for other angiosperm genera. In contrast, colonization of remote islands (New Zealand, Juan Fernández) occurred considerably later in the Miocene-Pliocene boundary., This study was supported by the Fundación General CSIC and Banco Santander as part ofthe project titled “Do all endangered species hold the samevalue?: origin and conservation of living fossils of floweringplants endemic to Spain

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., This work was carried out with financial support by the National Science Foundation (Award #1255901 to ALH and Award #1256033 to EHR), the Spanish Ministry of Economy and Competitiveness (project CGL2016–77401‐P to SM-B and ML), the USDA National Institute of Food and Agriculture (McIntire Stennis project 1018692 to DS) as well as postdoctoral fellowships towards SM‐B (Universidad Pablo de Olavide, PP16/12‐APP), and PJ‐M (National Science Foundation, Award #1256033, and the Smithsonian Postdoctoral Fellowship program).

Despite previous efforts to unify the terminology for Cyperaceae, two different terms, perigynium and utricle, are in common use for the prophyllar bract enclosing the female flower of Carex. Use of these terms is divided largely on geographic lines (mainly North American versus European and other authors, respectively). The recent merging of Kobresia with Carex requires a single term to refer to both the open prophyll of Kobresia and the laterally closed one of Carex. However, even when authors use utricle for Carex species, these same authors do not refer to the open prophyll of Kobresia as a utricle. We show that perigynium was apparently coined earlier than utricle, at the end of the 18th century, but the term utricle became more widespread than perigynium by the 20th century. Neither of the two terms is unambiguous, as both have also been used for other structures in other plant groups. Given this background, we propose revised definitions of both terms based on two facts: 1) the greater sem...

We present the first large-scale phylogenetic hypothesis for the genus Carex based on 996 of the 1983 accepted species (50.23%). We used a supermatrix approach using three DNA regions: ETS, ITS and matK. Every concatenated sequence was derived from a single specimen. The topology of our phylogenetic reconstruction largely agreed with previous studies. We also gained new insights into the early divergence structure of the two largest clades, core Carex and Vignea clades, challenging some previous evolutionary hypotheses about inflorescence structure. Most sections were recovered as non-monophyletic. Homoplasy of characters traditionally selected as relevant for classification, historical misunderstanding of how morphology varies across Carex, and regional rather than global views of Carex diversity seem to be the main reasons for the high levels of polyphyly and paraphyly in the current infrageneric classification.

Despite the existence of a rich record, fossils of Carex have been largely neglected by neobotanists. Here we present an exhaustive critically commented checklist of the so-far published pre-Pleistocene records ascribed to Carex. We have made every attempt to include all pertinent paleobotanical literature. The extensive collection of the Museum fur Naturkunde (Berlin, Germany) has also been studied. We found that up to 83 names belonging to different Carex groups may be applied to reliable remains, plus another 23 names of doubtful remains. These fossils come from more than 550 sites, mainly from C and E Europe and the Russian Federation. The age of the reliable fossil record ranges from the Eocene to recent times. Problems detected on the taxonomy, age and naming of these fossils are discussed. We hope that this work will stimulate neobotanical and paleobotanical communities to work together on broad-scope systematic projects as this one.

Critically endangered species representing ancient, evolutionarily isolated lineages must be given priority when allocating resources for conservation projects. Sound phylogenetic analyses and divergence time estimations are required to detect them, and studies on their population genetics, ecological requirements and breeding system are needed to understand their evolutionary history and to design efficient conservation strategies. Here we present the paradigmatic case of Avellara, a critically endangered monotypic genus of Compositae inhabiting a few swamps in the west-southwest Iberian coastal plains. Our phylogenetic and dating analyses based on nuclear (ITS) and plastid (matK) DNA sequences support a Miocene (>8.6 Ma) divergence between Avellara and closely related genera, resulting in marked morphological and ecological differentiation. We found alarmingly low levels of genetic diversity, based on AFLPs and plastid DNA sequences, and confirmed the prevalence of clonal reproduction. Species distribution modelling suggested a large macroclimatically suitable area for Avellara in the western Iberian Peninsula, but its apparently narrow microecological requirements restrict its distribution to peatlands with low-mineralised waters. Although five populations have been recorded from Spain and Portugal in the past, its current distribution may be reduced to only one population, recurrently found in the last decade but threatened by herbivory and habitat degradation. All this confirms the consideration of Avellara as a threatened species with high phylogenetic singularity, and makes it a flagship species for plant conservation in both Spain and Portugal that should be given priority in the design of in situ and ex situ conservation programmes.

Linum bicolor is a problematic North African taxon associated with L. setaceum. A consensus on the taxonomic status of a putative form (L. setaceum var. bicolor f. robusta) has not been achieved yet. We conducted a morphological study based on herbarium and field collections, combined with nuclear (ITS) and plastid (ndhF5–8 and trnL-F) phylogenies to clarify its status. The phylogenetic analysis did not reveal molecular divergence, but a comparative morphological study revealed substantial differences in traits previously used to distinguish the two taxa (stems, leaves, corolla and calyx size). In addition, we found that the indumentum of sepals, petal colour, and the arrangement of anthers and stigmas differed so clearly between the form robusta and L. setaceum s. stricto, that recognizing the former taxon as an independent species was justified. We provide an identification key to the yellow- and white-flowered Linum species in NW Africa. We also revised the unplaced name L. bicolor Schoubs. ex DC. and lectotypified it to avoid nomenclatural problems.

An updated inventory of the vascular flora alien to Italy, providing details on the occurrence at regional level, is presented. The checklist includes 1597 species, subspecies, and hybrids, distributed in 725 genera and 152 families; 2 taxa are lycophytes, 11 ferns and fern allies, 33 gymnosperms, and 1551 angiosperms. 157 taxa are archaeophytes and 1440 neophytes. The alien taxa currently established in Italy are 791 (570 naturalized and 221 invasive), while 705 taxa are casual aliens, 4 are not assessed, 7 are of unknown regional distribution, 47 have not been confirmed in recent times, 3 are considered extinct or possibly extinct in the country, and 40 are doubtfully occurring in Italy. This checklist allows to establish an up-to-date number (9792) of taxa constituting the whole (native and alien) Italian flora.

We are grateful to the John D. and Catherine T. MacArthur Foundation for funding of the Biodiversity Synthesis Group of the Encyclopedia of Life (EOL) project, which funded our BioSynC Synthesis meeting at the Field Museum in Chicago in September 2011, when the Global Carex Group was formed. We also thank the US National Science Foundation (NSF) for funding our continuing international collaborative work on the phylogeny and classification of Carex under grants DEB 1255901 to ALH and MJW, and DEB 1256033 to EHR. We also acknowledge with thanks funding for nomenclatural research and for attendance at our second meeting during the Monocots V conference in New York in July, 2013, from the Natural Sciences and Engineering Research Council, Canada (NSERC) to MJW and JRS; University of Mainz to BG; JSPS KAKENHI Grant no. 25840136 to OY; Korea National Arboretum to SK; CGL2012- 38744 project from the Spanish Ministry of Economy and Competitiveness to ML; project 30870178 from the National Natural Science Foundation of China to SRZ, and a University of Wisconsin-Madison Raper Travel Grant to DS. The figures were prepared with invaluable technical advice from H. C. Rimmer.

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.

The Spanish Ministry of Science and Technology supported this research through the project CGL2012-38744 and CGL2016- 77401-P to M. Luceño (Principal Investigator), E. Maguilla, M. Escudero, S. Martín-Bravo, and T. Villaverde and an FPU fellowship (AP2012-2189) from the Spanish Government (Ministry of Education, Culture, and Sport) and a Synthesys grant (GB-TAF-2523) by the European Community Research Infrastructure Program to E.Maguilla.

Castrilanthemum debeauxii is a critically endangered annual plant narrowly distributed in mountains of SE Iberia. It occurs in spiny scrublands on limestone soils dominated by Erinacea anthyllis at medium altitudes. Phylogenetic results indicate that the monotypic Castrilanthemum constitutes a paleoendemic lineage that diverged from its closest extant relatives in the Miocene. To achieve a better understanding of its recent evolution and conservation status, we performed an analysis of genetic variation and structure of four subpopulations using plastid haplotypes and AFLPs. We also investigated additional characteristics (ecological requirements, reproductive biology, pre-dispersal seed predation) that are relevant to evaluate the conservation status of the species. Castrilanthemum was found to be a facultative selfer with no major germination limitations. We detected moderate levels of genetic diversity based on AFLPs and plastid DNA sequences. The low genetic structure found for both nuclear and plastid markers, together with species distribution modeling results, pointed to high gene flow, which might counteract processes of genetic erosion caused by the low number of individuals. We found significant levels of pre-dispersal seed predation by the burrowing larvae of a tephritid fly, which may have contributed to the decline of the populations. Based on our results, we suggest directions for the management of Castrilanthemum. In particular, we recommend seeding and collection of seeds for banks, and discuss the possibilities of population reinforcement and reintroduction in localities where the species is currently extinct., This research was funded by Banco Santander, the Spanish National Research Council (CSIC) and Fundación General CSIC (FGCSIC) through a Proyecto Cero on endangered species 2010 (“Do all endangered species hold the same value?: origin and conservation of living fossils of flowering plants endemic to Spain”).

Bipolar disjunct distributions are a fascinating biogeographic pattern exhibited by about 30 vascular plants, whose populations reach very high latitudes in the northern and southern hemispheres. In this review, we fi rst propose a new framework for the defi nition of bipolar disjunctions and then reformulate a list of guiding principles to consider how to study bipolar species. Vicariance and convergent evolution hypotheses have been argued to explain the origin of this fragmented distribution pattern, but we show here that they can be rejected for all bipolar species, except for Carex microglochin . Instead, human introduction and dispersal (either direct or by mountain-hopping) — facilitated by standard and nonstandard vectors — are the most likely explanations for the origin of bipolar plant disjunctions. Successful establishment after dispersal is key for colonization of the disjunct areas and appear to be related to both intrinsic (e.g., self-compatibility) and extrinsic (mutualistic and antagonistic interactions) characteristics. Most studies on plant bipolar disjunctions have been conducted in Carex (Cyperaceae), the genus of vascular plants with the largest number of bipolar species. We found a predominant north-tosouth direction of dispersal, with an estimated time of diversifi cation in agreement with major cooling events during the Pliocene and Pleistocene. Bipolar Carex species do not seem to depend on specialized traits for long-distance dispersal and could have dispersed through one or multiple stochastic events, with birds as the most likely dispersal vector., Th is research was supported by the Spanish Ministry of Economy and Competitiveness (projects CGL2012-3874 and CGL2016-77401-P).

Narrow endemics constitute the cornerstone of Mediterranean plant diversity. Naufraga balearica (Apiaceae) is a critically endangered, extremely narrow endemic plant from the western Mediterranean island of Majorca. Because the species belongs to a monotypic genus, N. balearica was hypothesized to be a palaeoendemism. Here we conducted phylogenetic dating, population genetic and climatic niche analyses in order to understand the evolutionary history and conservation perspectives of this flagship species. Phylogenetic dating analysis of nuclear and plastid DNA sequences revealed a late Miocene to early Pliocene divergence between Naufraga and its sister genus Apium, supporting the palaeoendemic status of the former. Amplified fragment length polymorphism (AFLP) markers and plastid DNA sequences of the five Naufraga populations revealed moderate genetic diversity. This diversity is in line with that of other palaeoendemisms from western Mediterranean islands, as revealed by a comparison with 22 other narrow endemic species from this region. Despite the fact that all Naufraga populations are located at a maximum distance of 10. km in a straight line, a strikingly strong population differentiation was found for AFLP markers, which is explained by long-term isolation likely related to short-range pollination and dispersal strategies of the species. While the species is not genetically impoverished, species distribution modelling and microclimatic monitoring revealed that narrow ecological requirements underlie the current extreme rarity of Naufraga and may jeopardize its long-term survival. Our results indicate that a multidisciplinary approach provides powerful tools to develop conservation strategies for evolutionarily singular lineages. © 2014 Geobotanisches Institut ETH, Stiftung Ruebel., This research was funded by Banco Santander, the Spanish National Research Council (CSIC) and Fundación General CSIC (FGCSIC) through a Proyecto Cero on endangered species 2010 (“Do all endangered species hold the same value?: origin and conservation of living fossils of flowering plants endemic to Spain”)

Carex flava is a species complex with frequent hybridization. Within its Old World distribution, the group reaches the Iberian Peninsula, North Africa and Macaronesia, where the taxonomy of the group is particularly controversial. We performed a biometric study using quantitative characters and statistical multivariate and univariate techniques to test taxonomic relationships within previously defined genetic lineages, and to study the clinal morphological variation between those lineages. Our analytical approach allowed us to clearly discriminate between five well-defined morphogroups (C. demissa s. s., C. flava, C. lepidocarpa, C. viridula and a fifth group composed of C. nevadensis and populations from the Atlas region) and to establish morphological relationships for problematic morphotypes. The Pyrenean-Cantabrian populations predominantly displayed intermediate morphologies between C. demissa and C. lepidocarpa. Conversely, the Atlantic-Iberian morphotype, considered a hybrid between C. dem...