Marcial Escudero

Strait of Gibraltar: an effective gene-flow barrier for wind-pollinated Carex helodes (Cyperaceae) as revealed by DNA sequences, AFLP, and cytogenetic variation.

The Strait of Gibraltar is the most important barrier disconnecting the landmasses of Europe and Africa on the western Mediterranean extreme. Carex helodes is a wind-pollinated species endemic to the western Mediterranean. Because molecular and cytogenetic data allow the inference of its evolutionary history, we analyzed variations in chromosome number, including meiotic chromosome behavior, amplified fragment length polymorphism (AFLP) fingerprints, and nucleotide substitutions in plastid and nuclear DNA sequences. Cytogeographic results showed that the African populations have stabilized at a single chromosome number of 2n = 74, whereas the most frequent cytotype in Iberia is 2n = 72. Phylogenetic reconstructions of 17 sequences from nine closely related species revealed that C. helodes is monophyletic and that the Moroccan populations are embedded in the Iberian lineages. The haplotype network is also consistent with a European origin of the northern African haplotype. AFLP analysis also revealed hierarchical levels of genetic variation compatible with a founder effect process responsible for the African populations. All sources of evidence support the hypothesis that the Strait of Gibraltar has been an effective gene-flow barrier, generating two isolated evolutionary lineages after their dispersal. Recent connections between the two lineages appear unlikely, whereas active gene flow occurs among populations within the two lineages.

The east-west-north colonization history of the Mediterranean and Europe by the coastal plant Carex extensa (Cyperaceae)

Coastal plants are ideal models for studying the colonization routes of species because of the simple linear distributions of these species. Carex extensa occurs mainly in salt marshes along the Mediterranean and European coasts. Variation in cpDNA sequences, amplified fragment length polymorphisms (AFLPs) and simple sequence repeats (SSRs) of 24 populations were analysed to reconstruct its colonization history. Phylogenetic relationships indicate that C. extensa together with the South American Carex vixdentata and the southern African Carex ecklonii form a monophyletic group of halophilic species. Analyses of divergence times suggest that early lineage diversification may have occurred between the late Miocene and the late Pliocene (Messinian crisis). Phylogenetic and network analyses of cpDNA variation revealed the monophyly of the species and an ancestral haplotype contained in populations of the eastern Mediterranean. The AFLP and SSR analyses support a pattern of variation compatible with these two lineages. These analyses also show higher levels of genetic diversity and differentiation in the eastern population group, which underwent an east‐to‐west Mediterranean colonization. Quaternary climatic oscillations appear to have been responsible for the split between these two lineages. Secondary contacts may have taken place in areas near the Ligurian Sea in agreement with the gene flow detected in Corsican populations. The AFLP and SSR data accord with the ‘tabula rasa’ hypothesis in which a recent and rapid colonization of northern Europe took place from the western Mediterranean after the Last Glacial Maximum. The unbalanced west‐east vs. west‐north colonization may be as a result of ‘high density blocking’ effect.

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.

Systematics and evolution of Carex sects. Spirostachyae and Elatae (Cyperaceae)

Carex sect. Spirostachyae comprises 25 species displaying the centre of diversity in Eurasia, while sect. Elatae comprises 22 species displaying the centre of diversity in tropical African mountainous regions. Phylogenetic analysis of the 136 ITS and 108 5′trnK intron sequences of 23 species of sect. Spirostachyae and 20 species of sect. Elatae revealed that neither section is monophyletic. With the exclusion of C. montis-eeka (sect. Spirostachyae) and C. insularis, C. iynx and C. longibrachiata (sect. Elatae), the sects. Spirostachyae and Elatae constitute a monophyletic group of 38 species, 22 of which were previously included in sect. Spirostachyae and 16 of which were in sect. Elatae (considering C. thomasii as C. mannii ssp. thomasii). Two main groups with different edaphic preferences were identified in the core Spirostachyae, in congruence with some morphological features. One group comprises primarily acidophilus species including 11 species of sect. Spirostachyae and 16 species of sect. Elatae. The other group includes 11 basophilic species of sect. Spirostachyae. Incongruence between nuclear and plastid genomes was detected, suggesting hybridization or lineage sorting processes in the evolution of the core Spirostachyae.

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.

Karyotype stability and predictors of chromosome number variation in sedges: a study in Carex section Spirostachyae (Cyperaceae).

Previous work on holocentric chromosomes in the angiosperm genus Carex demonstrates that many of the traditional sections are marked by different ranges of chromosome number, suggesting phylogenetic autocorrelation. It has been hypothesized that shifting constraints on chromosome rearrangements may limit the potential for hybridization among lineages, promoting speciation. In this study, we evaluated alternative evolutionary models to test for such transitions in Carex section Spirostachyae as well as the relative effects of several plausible drivers of intraspecific chromosome diversity. Chromosome number variation in section Spirostachyae shows significant phylogenetic signal, but no evidence of clade-specific shifts in chromosome number distribution. This gradual model of chromosome evolution contrasts with the shifting equilibrium model previously identified in a younger section of the same genus, suggesting that section Spirostachyae may have a more slowly evolving karyotype. Chromosome number variance, on the other hand, exhibits low phylogenetic signal. Average time of coalescence rather than geographic range or chromosome number itself predicts chromosome number variance, demonstrating a previously unreported relationship between population history and cytogenetic variation.

MATICCE: mapping transitions in continuous character evolution

SUMMARY MATICCE is a new software package in the R language for mapping phylogenetic transitions in organismal traits that have continuous distributions. MATICCE integrates over phylogenetic and model uncertainty and provides simulation functions for visualizing evolutionary scenarios based on estimated parameter values. AVAILABILITY AND IMPLEMENTATION MATICCE is written in the open source R language and freely available through the Comprehensive R Archive Network (http://cran.r-project.org/web/packages/maticce).

The Phylogenetic Origins and Evolutionary History of Holocentric Chromosomes

Abstract In eukaryotes, we can recognize two kinds of chromosomes, based on the location of the kinetochores. The majority of eukaryotes have monocentric chromosomes, in which kinetochoric activity is concentrated in a single locus. In several unrelated eukaryotic lineages, chromosomes are holocentric, having diffuse centromeric / kinetochoric activity along the length of the chromosome. Whether holocentric chromosomes are derived or ancestral is still under debate. This study uses the phylogenetic tree from Time Tree of Life project, comprising more than 50,000 sampled species, to reconstruct the evolution of holocentry. Asymmetrical two-state Markov (Mk2) models were compared with BiSSE models to assess sensitivity of our conclusions to possible effects of holocentry on lineage diversification rates. Our analyses based on Mk2 and BiSSE models inferred that the rate of transition from holocentric to monocentric chromosomes is two orders of magnitude higher than the reverse direction. The ancestral state of all eukaryotes is ambiguous depending on the model, inferred to be either monocentric (Mk2) or holocentric (BiSSE). Whatever the direction, the multiple transitions and high diversity of centromere organization across the tree of life are what we would expect if there are selective advantages to both chromosome types. Understanding those selective advantages is key to understanding how genetic information is organized and transmitted from one generation to the next, and why these major evolutionary transitions in centromere organization have occurred in the first place.

Biogeography of Flowering Plants: A Case Study in Mignonettes (Resedaceae) and Sedges (Carex, Cyperaceae)

Biogeography is a multidisciplinary science that studies the past and present geographic distribution of organisms and the causes behind it. The combination of historical events and evolutionary processes has usually an outstanding role when explaining the shape of a species range. As already noted by Darwin more than 150 years ago, patterns of species distribution may often be seen as clear footsteps of their evolution and diversification (Darwin, 1859). It is now also well known that geological events (i.e. continental drift, orogeny or island formation) and climatic oscillations occurred during the recent geological history of the Earth, like the cooling and aridification that took place during the Pliocene (5.3 – 2.5 million years ago, m.a) and the Pleistocene glaciations (1.8 – 0.01 m.a), prompted great range shifts. These geological and/or climatic changes caused, in some cases, the extinction of species; in many others, they provided conditions of reproductive isolation and/or genetic divergence between populations and, eventually, produced speciation, the engine of biodiversity.

Niche shifts after long-distance dispersal events in bipolar sedges (Carex, Cyperaceae)

PREMISE OF THE STUDY Bipolar species represent the greatest biogeographical disjunction on Earth, raising many questions about the colonization and adaptive processes behind such striking distribution. We investigated climatic niche differences of five Carex bipolar species in North and South America to assess niche shifts between these two regions. Moreover, we assessed potential distribution changes with future climate change. METHODS We used 1202 presence data points from herbarium specimens and 19 bioclimatic variables to assess climatic niche differences and potential distributions among the five species using ordination methods and Maxent. KEY RESULTS The niche overlap analyses showed low levels of niche filling and high climatic niche expansion between North and South America. Carex macloviana and C. maritima showed the greatest niche expansion (60% and 96%, respectively), followed by C. magellanica (45%) and C. microglochin (39%). Only C. canescens did not colonize new environments (niche expansion = 0.2%). In contrast, all species but C. magellanica had niche filling that was <40%; hence, they are absent in the south from many environments they inhabit in North America. Climate change will push all species toward higher latitudes and elevation, reducing the availability of suitable environments. CONCLUSIONS The colonization of South America seems to have involved frequent climatic niche shifts. Most species have colonized new environments from those occupied in the North. Observed niche shifts appear congruent with time since colonization and with current genetic structure within species. In these cold-dwelling species, climate change will most likely decrease their suitable environments in the future.