Geometric Morphometrics sheds light on the systematics affinities of two enigmatic dwarf Neotropical sedges (Carex, Cyperaceae).

Geometric morphometrics (GM) is a powerful analytical tool that enables complete quantification of shapes. Its use in Botany has a great potential for complementing plant evolutionary and ecological studies. Taxonomic delimitation in Carex has been complicated due to reduction of characters and frequent homoplasy. This problem is more marked in cases where the species exhibit dwarfism. South America is the continent with the least understood Carex flora. The systematic relationships of some bizarre-looking groups were not unraveled until molecular phylogenetic studies resolved their relationships. In particular, there are two species only known from their type material whose affinities remain uncertain: Carexherteri and C.hypsipedos. These two taxa are acaulescent plants that respectively grow in the Uruguayan pampa and Peruvian high-altitude meadows. Recently, both species were ascribed to the Carexphalaroides group (subgen. Psyllophorae, sect. Junciformes) due to superficial morphological similarities, such as the androgynous peduncled spikes. However, their character combination is also coincident for its circumscription to sect. Abditispicae species. Nevertheless, in the absence of confirmation from molecular analyses, their placement must be considered preliminary until additional data can be provided. In this work we employ for the first time geometric morphometrics (GM) tools to assess the systematic affinities of two taxonomically problematic sedge species based on fruit shape. We compared utricle morphology of C.herteri and C.hypsipedos with that of C.phalaroides group and species in sect. Abditispicae. To this end we used GM and traditional morphometric approaches. Utricle shape variation along with other morphological features support the exclusion of these two species from the C.phalaroides gr. and, at the same time, show clear affinities of C.herteri to sect. Abditispicae. Carexhypsipedos remains as an incertae sedis species. Our work shows the potential utility of GM for the exploration of systematic affinities in sedges and in other graminoids.

Carexquixotiana (Cyperaceae), a new Iberian endemic from Don Quixote's land (La Mancha, S Spain).

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 Carexsect.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 Carexquixotiana 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.

New relevant chorological and conservation data on Carex (Cyperaceae) and Hypericum (Hypericaceae) from Ecuador.

Background: Knowledge of Carex L. (true sedges) and Hypericum L. (St. John's wort) in the Neotropics is fragmentary. New information: 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.

Dramatic impact of future climate change on the genetic diversity and distribution of ecologically relevant Western Mediterranean Carex (Cyperaceae).

Anticipating the evolutionary responses of species to ongoing climate change is essential to propose effective management and conservation measures. The Western Mediterranean Basin constitutes one of the hotspots of biodiversity where the effects of climate change are expected to be more dramatic. Plant species with ecological relevance constitute ideal models to evaluate and predict the impact of climate change on ecosystems. Here we investigate these impacts through the spatio-temporal comparison of genetic diversity/structure (AFLPs), potential distribution under different future scenarios of climate change, and ecological space in two Western Mediterranean sister species of genus Carex. Both species are ecologically key in their riparian habitats, but display contrasting distribution patterns, with one widespread in the Iberian Peninsula and North Africa (C. reuteriana), while the other (C. panormitana) is a restricted, probably endangered, Central Mediterranean endemic. At present, we found a strong genetic structure driven by geography in both species, and lower values of genetic diversity and a narrower ecological space in C. panormitana than in C. reuteriana, while the allelic rarity was higher in the former than in C. reuteriana subspecies. Future projections predict an overall dramatic reduction of suitable areas for both species under all climate change scenarios, which could be almost total for C. panormitana. In addition, gene diversity was inferred to decrease in all taxa, with genetic structure reinforcing in C. reuteriana by the loss of admixture among populations. Our findings stress the need for a reassessment of C. panormitana conservation status under IUCN Red List criteria and the implementation of conservation measures.

A snapshot of progenitor-derivative speciation in Iberodes (Boraginaceae).

Traditional classification of speciation modes has focused on physical barriers to gene flow. Allopatric speciation with complete reproductive isolation is viewed as the most common mechanism of speciation. Parapatry and sympatry, by contrast, entail speciation in the face of ongoing gene flow, making them more difficult to detect. The genus Iberodes (Boraginaceae, NW Europe) comprises five species with contrasting morphological traits, habitats and species distributions. Based on the predominance of narrow and geographically distant endemic species, we hypothesized that geographical barriers were responsible for most speciation events in Iberodes. We undertook an integrative study including: (i) phylogenomics through restriction-site-associated DNA sequencing (RAD-seq), (ii) genetic structure analyses, (iii) demographic modelling, (iv) morphometrics, and (v) climatic niche modelling and niche overlap analysis. The results revealed a history of recurrent progenitor-derivative speciation manifested by a paraphyletic pattern of nested species differentiation. Budding speciation mediated by ecological differentiation is suggested for the coastal lineage, deriving from the inland widespread Iberodes linifolia during the Late Pliocene. Meanwhile, geographical isolation followed by niche shifts are suggested for the more recent differentiation of the coastland taxa. Our work provides a model for distinguishing speciation via ecological differentiation of peripheral, narrowly endemic I. kuzinskyanae and I. littoralis from a widespread extant ancestor, I. linifolia. Ultimately, our results illustrate a case of Pliocene speciation in the probable absence of geographical barriers and get away from the traditional cladistic perspective of speciation as producing two species from an extinct ancestor, thus reminding us that phylogenetic trees tell only part of the story.

Are Cenozoic relict species also climatic relicts? Insights from the macroecological evolution of the giant sedges of Carex sect. Rhynchocystis (Cyperaceae).

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.

An Evolutionary Study of Carex Subg. Psyllophorae (Cyperaceae) Sheds Light on a Strikingly Disjunct Distribution in the Southern Hemisphere, With Emphasis on Its Patagonian Diversification.

Carex subgenus Psyllophorae is an engaging study group due to its early diversification compared to most Carex lineages, and its remarkable disjunct distribution in four continents corresponding to three independent sections: sect. Psyllophorae in Western Palearctic, sect. Schoenoxiphium in Afrotropical region, and sect. Junciformes in South America (SA) and SW Pacific. The latter section is mainly distributed in Patagonia and the Andes, where it is one of the few Carex groups with a significant in situ diversification. We assess the role of historical geo-climatic events in the evolutionary history of the group, particularly intercontinental colonization events and diversification processes, with an emphasis on SA. We performed an integrative study using phylogenetic (four DNA regions), divergence times, diversification rates, biogeographic reconstruction, and bioclimatic niche evolution analyses. The crown age of subg. Psyllophorae (early Miocene) supports this lineage as one of the oldest within Carex. The diversification rate probably decreased over time in the whole subgenus. Geography seems to have played a primary role in the diversification of subg. Psyllophorae. Inferred divergence times imply a diversification scenario away from primary Gondwanan vicariance hypotheses and suggest long-distance dispersal-mediated allopatric diversification. Section Junciformes remained in Northern Patagonia since its divergence until Plio-Pleistocene glaciations. Andean orogeny appears to have acted as a northward corridor, which contrasts with the general pattern of North-to-South migration for temperate-adapted organisms. A striking niche conservatism characterizes the evolution of this section. Colonization of the SW Pacific took place on a single long-distance dispersal event from SA. The little ecological changes involved in the trans-Pacific disjunction imply the preadaptation of the group prior to the colonization of the SW Pacific. The high species number of the section results from simple accumulation of morphological changes (disparification), rather than shifts in ecological niche related to increased diversification rates (radiation).

An integrative monograph of Carex section Schoenoxiphium (Cyperaceae).

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.

Timing and ecological priority shaped the diversification of sedges in the Himalayas.

BACKGROUND: Diversification patterns in the Himalayas have been important to our understanding of global biodiversity. Despite recent broad-scale studies, the most diverse angiosperm genus of the temperate zone-Carex L. (Cyperaceae), with ca. 2100 species worldwide-has not yet been studied in the Himalayas, which contains 189 Carex species. Here the timing and phylogenetic pattern of lineage and ecological diversification were inferred in this ecologically significant genus. We particularly investigated whether priority, adaptation to ecological conditions, or both explain the highly successful radiation of the Kobresia clade (ca. 60 species, of which around 40 are present in the Himalayas) of Himalayan Carex. METHODS: Phylogenetic relationships were inferred using maximum likelihood analysis of two nuclear ribosomal DNA (nrDNA) regions (ITS and ETS) and one plastid gene (matK); the resulting tree was time-calibrated using penalized likelihood and a fossil calibration at the root of the tree. Biogeographical reconstruction for estimation of historical events and ancestral ranges was performed using the dispersal-extinction-cladogenesis (DEC) model, and reciprocal effects between biogeography and diversification were inferred using the geographic state speciation and extinction (GeoSSE) model. Climatic envelopes for all species for which mapped specimen data available were estimated using climatic data from WORLDCLIM, and climatic niche evolution was inferred using a combination of Ornstein-Uhlenbeck models of shifting adaptive optima and maximum likelihood inference of ancestral character states under a Brownian motion model. RESULTS: The Himalayan Carex flora represents three of the five major Carex clades, each represented by multiple origins within the Himalayas. The oldest Carex radiation in the region, dating to ca. 20 Ma, near the time of Himalayan orogeny, gave rise to the now abundant Kobresia clade via long-distance dispersal from the Nearctic. The Himalayan Carex flora comprises a heterogeneous sample of diversifications drawn from throughout the cosmopolitan, but mostly temperate, Carex radiation. Most radiations are relatively recent, but the widespread and diverse Himalayan Kobresia radiation arose at the early Miocene. The timing and predominance of Kobresia in high-elevation Himalayan meadows suggests that Kobresia may have excluded other Carex lineages: the success of Kobresia in the Himalayas, in other words, appears to be a consequence largely of priority, competitive exclusion and historical contingency.

Being in the right place at the right time? Parallel diversification bursts favored by the persistence of ancient epizoochorous traits and hidden factors in Cynoglossoideae.

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.

Cut from the same cloth: The convergent evolution of dwarf morphotypes of the Carex flava group (Cyperaceae) in Circum-Mediterranean mountains.

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.

Bipolar distributions in vascular plants: A review.

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 first propose a new framework for the definition 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-to-south direction of dispersal, with an estimated time of diversification 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.

Karyotypic changes through dysploidy persist longer over evolutionary time than polyploid changes.

Chromosome evolution has been demonstrated to have profound effects on diversification rates and speciation in angiosperms. While polyploidy has predated some major radiations in plants, it has also been related to decreased diversification rates. There has been comparatively little attention to the evolutionary role of gains and losses of single chromosomes, which may or not entail changes in the DNA content (then called aneuploidy or dysploidy, respectively). In this study we investigate the role of chromosome number transitions and of possible associated genome size changes in angiosperm evolution. We model the tempo and mode of chromosome number evolution and its possible correlation with patterns of cladogenesis in 15 angiosperm clades. Inferred polyploid transitions are distributed more frequently towards recent times than single chromosome gains and losses. This is likely because the latter events do not entail changes in DNA content and are probably due to fission or fusion events (dysploidy), as revealed by an analysis of the relationship between genome size and chromosome number. Our results support the general pattern that recently originated polyploids fail to persist, and suggest that dysploidy may have comparatively longer-term persistence than polyploidy. Changes in chromosome number associated with dysploidy were typically observed across the phylogenies based on a chi-square analysis, consistent with these changes being neutral with respect to diversification.

Toward an accurate taxonomic interpretation of Carex fossil fruits (Cyperaceae): a case study in section Phacocystis in the Western Palearctic.

PREMISE OF THE STUDY: Despite growing interest in the systematics and evolution of the hyperdiverse genus Carex, few studies have focused on its evolution using an absolute time framework. This is partly due to the limited knowledge of the fossil record. However, Carex fruits are not rare in certain sediments. We analyzed carpological features of modern materials from Carex sect. Phacocystis to characterize the fossil record taxonomically. METHODS: We studied 374 achenes from modern materials (18 extant species), as well as representatives from related groups, to establish the main traits within and among species. We also studied 99 achenes from sediments of living populations to assess their modification process after decay. Additionally, we characterized 145 fossil achenes from 10 different locations (from 4-0.02 mya), whose taxonomic assignment we discuss. KEY RESULTS: Five main characters were identified for establishing morphological groups of species (epidermis morphology, achene-utricle attachment, achene base, style robustness, and pericarp section). Eleven additional characters allowed the discrimination at species level of most of the taxa. Fossil samples were assigned to two extant species and one unknown, possibly extinct species. CONCLUSIONS: The analysis of fruit characters allows the distinction of groups, even up to species level. Carpology is revealed as an accurate tool in Carex paleotaxonomy, which could allow the characterization of Carex fossil fruits and assign them to subgeneric or sectional categories, or to certain species. Our conclusions could be crucial for including a temporal framework in the study of the evolution of Carex.

Taxonomic delimitation and drivers of speciation in the Ibero-North African Carex sect. Phacocystis river-shore group (Cyperaceae).

PREMISE OF THE STUDY: The Ibero-North African Carex sect. Phacocystis river-shore group is a set of perennial helophytic species with poorly defined taxonomic boundaries. In the present study, we delimited the different taxonomic units, addressed the phylogeographic history, and evaluated the drivers of differentiation that have promoted diversification of these plants. METHODS: We analyzed molecular data using statistical parsimony for plastid sequences (26 samples from 26 populations) and principal coordinate analysis, neighbor joining, and Bayesian analysis of population structure for AFLPs (186 samples from 26 populations). Chromosome numbers from 14 samples (9 populations) are newly reported. KEY RESULTS: Three species can be distinguished (C. acuta, C. elata, and C. reuteriana). Unexpectedly for rhizome-growing helophytes, the vegetative reproduction detected was incidental. The widespread C. elata was found to be a genetically poorly differentiated taxon, whereas the local C. reuteriana displayed geographical structuring. Geographical factors seem to be the main driver of differentiation for both taxa. CONCLUSIONS: Despite apparent morphological and ecological similarities, C. elata and C. reuteriana have disparate genetic structures and evolutionary histories, which may have originated from small ecological differences. Carex elata is broadly distributed throughout Europe, and its northern populations were recently founded, probably after the last glacial maximum. In contrast, C. reuteriana is an Ibero-North African endemic, with long-standing populations affected by isolation and limited gene flow. It is likely that high-density blocking effects and different gene-flow barriers act together to delimit its distribution and promote its relatively high population differentiation.

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Elementary-field expansions of genuine cross-spectral density matrices.

A necessary and sufficient nonnegative definiteness condition for the cross-spectral density matrix (CDM) is derived. It is also shown that this realizability condition allows the expansion of genuine CDMs in terms of recently introduced elementary fields, namely, mean-square coherent beams, and fields with position-independent stochastic behavior. The special case of uniformly polarized electromagnetic Schell-model sources is also analyzed.

Equivalence between optimum Young's fringe visibility and position-independent stochastic behavior of electromagnetic fields.

Stochastic electromagnetic fields characterized by optimized fringe visibility in a Young interferometric arrangement are shown to be those whose random character is position independent. The optimization procedure involves local unitary transformations, which can be implemented by using reversible anisotropic polarization devices placed at the two pinholes. It is also shown that the local degree of polarization in the optimized interferometer is constant across the superposition region and coincides with the degree of polarization at the two pinholes.