Significance of ecological vicariance and long-distance dispersal in the diversification of Carex sect. Spirostachyae (Cyperaceae)

Plant disjunctions have provided one of the most intriguing distribution patterns historically addressed by biogeographers. Carex sect. Spirostachyae (Cyperaceae) displays an interesting pattern of disjunction to evaluate these scenarios, with species occurring in the main continental landmasses and in oceanic islands of the two hemispheres. Internal transcribed spacer and 5'-trnK intron plastid gene sequences were analyzed to determine (1) the times of diversification using penalized likelihood, and (2) reconstructions of the regions using maximum likelihood and Bayesian approaches of origin of sect. Spirostachyae and internal main lineages. The times for the diversification of sect. Spirostachyae are dated to between the end of the Eocene and the Oligocene, whereas the two main lineages are dated to between the end of the Oligocene and the beginning of Miocene. The phylogenetic analyses reveal a Mediterranean-Eurasian center of differentiation for sect. Spirostachyae and subsection Spirostachyae, whereas no clear, single ancestral area could be inferred for subsection Elatae. Both long-distance dispersal and ecological vicariance appear to have been involved in the evolutionary history of the disjunct distribution of the main lineages of sect. Spirostachyae. These organisms appear to have a special ability to colonize remote areas (through transoceanic and interhemispherical colonizations), but special long-distance dispersal mechanisms are not evident.     

Genetic diversity and endemism in North American Carex section Ceratocystis (Cyperaceae)

Factors leading to endemism, and the evolutionary implications of endemism, can be explored by studying closely related taxa with variously restricted distributions. Such a model is provided by Carex section Ceratocystis (Cyperaceae); Carex cryptolepis, Carex sp. nov., and C. lutea are North American endemics with broad, restricted, and highly restricted distributions, respectively. The prediction that levels of genetic diversity are a consequence of distribution size was tested within a phylogenetic context using population level genetic variation at 18 allozyme loci. In contrast to expectations, mean proportion of loci polymorphic, number of alleles per polymorphic locus, and expected heterozygosity were significantly greater in C. lutea than either C. cryptolepis or Carex sp. nov. Although the possibility of a shift in breeding system, past introgression, or progenitor-derivative relationships could explain the relatively high levels of variation observed in C. lutea, these were dismissed on the basis of allozyme and nuclear ribosomal sequence data. We conclude that C. lutea maintains levels of genetic diversity typical of caespitose carices despite its narrow endemism and that the low levels of genetic variation in C. cryptolepis and Carex sp. nov. are likely the result of population fluctuations during Pleistocene glacial-interglacial cycles.     

一种濒危大疣蛛的潜在分布区域以及气候变暖的影响

本研究的目的是i)确定伊比利亚半岛一种大疣蛛(Macrothele calpeiana)分布的气候相关性以预测其潜在分布,ii)详细阐述该物种在伊比利亚半岛的分布假说,iii)通过推断模型预测确认该物种在北非和整个地中海地区的适合区域,iv)预测气候变暖对蜘蛛潜在分布区的影响。基于物种的存在、远离目前环境条件下的可能缺失以及其它的气候参数,使用广义线性模型发展了潜在分布的可预测模型。蜘蛛在伊比利亚半岛上的潜在分布远大于目前已知的分布区,延伸到尚未发现蜘蛛分布的葡萄牙广大地区。本文提出了该种大疣蛛在适合分布区域内没有分布的历史因素。北非具有适合该物种的条件但却没有该属物种的分布支持了大疣蛛(Macrothele)祖先的东方起源假说。对地中海地区蜘蛛分布的推断突出显示了阿根廷地区有合适的分布区,该地区也发现有另一种欧洲大疣蛛。气候变暖将对现存于伊比利亚的M.calpeiana种群产生负面影响,因为气候变暖将减少并破碎化蜘蛛在北非的潜在栖息地。目前,急需确认在葡萄牙广大地区是否存在蜘蛛物种,发展对该属的系统发育研究以确定大疣蛛属起源和扩散史的理论。     

Karyosystematic study of Potentilla L. subgen. Potentilla (Rosaceae) in the Iberian Peninsula

Chromosome numbers in 80 populations belonging to 18 species of Potentilla L. subgen. Potentilla from the Iberian Peninsula and two of P. maura, a North African endemic taxon, have been counted. The basic number of chromosomes is always x = 7 and these chromosomes are small (between 1 and 2 μm). For three species, the number of chromosomes is reported for the first time and, for another six, this number has been established in Iberian representatives. Moreover, new ploidy levels have been obtained for P. hispanica and P. crantzii with regard to their entire distribution area, and in P. cinerea and P. neumanniana for the Iberian Peninsula. Some taxonomic, phylogenetic and phytogeographic comments are made for several species or groups of species from the West Mediterranean region. In 13 species only one ploidy level has been found, but six species have several ploidy levels. Seven ploidy levels occur in the investigated taxa. The frequency of each ploidy level represented within Iberian Potentilla is analysed and the data are compared with those available for taxa from the rest of the distribution area of the genus.     

Phylogeographical and speciation patterns in subterranean worm lizards of the genus Blanus (Amphisbaenia: Blanidae)

The peculiar lifestyle of subterranean reptiles must determine their modes of speciation and diversification. To further understand the evolutionary biology of subterranean reptiles, we studied the phylogeny of worm lizards of the genus Blanus and the phylogeography of its Iberian representatives. We used mitochondrial (ND4 and 16S rRNA) and nuclear (anonymous) partial gene sequences to resolve phylogenetic relationships within Blanus. The Eastern Mediterranean Blanus strauchi was recovered as sister group of Western Mediterranean species. Iberian and North African Blanus were recovered as reciprocally monophyletic groups. The same genes were used to determine phylogeography of 47 populations of Blanus cinereus. Mitochondrial and nuclear sequence data recovered two highly supported Iberian clades. Parapatry and high sequence divergences between them suggest that these clades may represent independent taxonomic units. A molecular clock was calibrated considering that the split between Iberian and North African Blanus was due to the re-opening of the Betic Strait in the Upper Tortonian (8-9 million years ago). Differentiation between the two Iberian clades was estimated to date back to 5.2 million years ago. The Central Iberian clade included five mitochondrial haplotype lineages (A-E). Geographical ranges of two of them broadly overlap in the central Iberian plateau. After testing alternative hypotheses, the most likely explanation for this striking phylogeographical pattern involves recent dispersal of one of the lineages (C) over the geographical range of the other (B). The inferred recent dispersal of this fossorial reptile is explained in terms of demographic advantages associated to underground lifestyle.     

Haplotype diversity and phylogenetic relationships among the Iberian barbels (Barbus,Cyprinidae) reveal two evolutionary lineages

The phylogenetic relationships and haplotype diversity of all Iberian barbels were examined by analyzing the complete mitochondrial cytochrome b gene sequence (1141 bp) of 72 specimens from 59 Iberian localities. Phylogenetic findings demonstrated a clear distinction between two mitochondrial lineages and confirmed the existence of two previously considered subgenera: Barbus and Luciobarbus: The first subgenus, Barbus, is represented on the Iberian Peninsula by Barbus haasi and Barbus meridionalis. The second subgenus, Luciobarbus, includes the remaining endemic Iberian species: Barbus comizo, Barbus bocagei, Barbus microcephalus, Barbus sclateri, Barbus guiraonis, and Barbus graellsii. Mean haplotype divergence between these subgenera was 10.40%, providing evidence of a clear subdivision within the Iberian barbels. Our results conflict with those reported in a recent study, based on 307 cytochrome b base pairs, that failed to identify any division within the genus Barbus in the Iberian Peninsula. The inclusion of nine further species belonging to this genus (used as outgroups) allowed us to establish a closer relationship of the Iberian species of the subgenus Barbus with other European taxa than with the Iberian Luciobarbus, which was found to cluster with North African, Caucasian, and Greek species. At the population level, no biogeographic structure was shown by specimens of each species (only 5.98% of the variation was attributable to differences among populations of each species). Given the discrete amount of divergence found among the Luciobarbus species, the formation of current hydrographic basins during the Plio-Pleistocene seems to have played a major role in their isolation and evolution.     

Genetic relationships within and among Iberian fescues (Festuca L.) based on PCR-amplified markers.

The genus Festuca comprises approximately 450 species and is widely distributed around the world. The Iberian Penninsula, with more than 100 taxa colonizing very diverse habitats, is one of its main centers of diversification. This study was conducted to assess molecular genetic variation and genetic relatedness among 91 populations of 31 taxa of Iberian fescues, based on several molecular markers (random amplified polymorphic DNA, amplified fragment length polymorphisms, and trnL sequences). The analyses showed the paraphyletic origin of the broad-leaved (subgenus Festuca, sections Scariosae and Subbulbosae, and subgenus Schedonorus) and the fine-leaved fescues (subgenus Festuca, sections Aulaxyper, Eskia, and Festuca). Schedonorus showed a weak relationship with Lolium rigidum and appeared to be the most recent of the broad-leaved clade. Section Eskia was the most ancient and Festuca the most recent of the fine-leaved clade. Festuca and Aulaxyper were the most related sections, in concordance with their taxonomic affinities. All taxa grouped into their sections, except F. ampla and F. capillifolia (section Festuca), which appeared to be more closely related to Aulaxyper and to a new independent section, respectively. Most populations clustered at the species level, but some subspecies and varieties mixed their populations. This study demonstrated the value in combining different molecular markers to uncover hidden genetic relationships between populations of Festuca.     

嵩草属植物硅酸体系统的研究

对国产狭义嵩草属 (K obresia,Cyperaceae) 3个组 2 7种植物进行了硅酸体系统分析研究 ,同时结合近缘的苔草属 (Carex) 2个组的植物硅酸体特征 ,深入探讨了植物硅酸体系统分析研究在莎草科苔草族 (Cariceae)中的系统分类学意义。认为 :(1 )嵩草属及其临近类群在属间、种间及种下水平的系统分类学比较研究都可以借助植物硅酸体系统研究的证据。 (2 )穗状嵩草组 (sect. Elyna)的硅酸体系统特征具有较高的一致性 ,除矮嵩草 (K.humilis)及其近缘种同其他穗状嵩草组的种类有所不同 ,它很可能有一个不同于穗状嵩草组其他类群的嵩草组(sect. K obresia)的亲缘。 (3 )从植物硅酸体系统特征的研究来看 ,拟苔草组 (sect. H emicarex)的一些种接近于穗状嵩草组 ,如高山嵩草 (K . pygmeae) ;而另一些种接近于嵩草组 ,如禾叶嵩草 (K . graminifolia) ;也有一些种类的硅酸体系统特征更接近苔草属 ,说明这个组很可能是多源的 ,即本组的单一花穗和果囊形态特征由不同类群的复合花序种类趋同演化而出。(4 )嵩草组的硅酸体系统特征具有较高的一致性 ,说明嵩草属原始类群在硅酸体系统特征方面与外部形态特征演化的某些相关性。     

Geographic genetic structure of Iberian columbines (gen. <Emphasis Type="Italic">Aquilegia</Emphasis>)

Southern European columbines (genus Aquilegia) are involved in active processes of diversification, and the Iberian Peninsula offers a privileged observatory to witness the process. Studies on Iberian columbines have provided significant advances on species diversification, but we still lack a complete perspective of the genetic diversification in the Iberian scenario. This work explores how genetic diversity of the genus Aquilegia is geographically structured across the Iberian Peninsula. We used Bayesian clustering methods, principal coordinates analyses, and NJ phenograms to assess the genetic relationships among 285 individuals from 62 locations and detect the main lineages. Genetic diversity of Iberian columbines consists of five geographically structured lineages, corresponding to different Iberian taxa. Differentiation among lineages shows particularly complex admixture patterns at Northeast and highly homogeneous toward Northwest and Southeast. This geographic genetic structure suggests the existence of incomplete lineage sorting and interspecific hybridization as could be expected in recent processes of diversification under the influence of quaternary postglacial migrations. This scenario is consistent with what is proposed by the most recent studies on European and Iberian columbines, which point to geographic isolation and divergent selection by habitat specialization as the main diversification drivers of the Iberian Aquilegia complex.     

Phylogenetic relationships of Iberian cyprinids: systematic and biogeographical implications.

The phylogenetic relationships among all Iberian endemic cyprinids were inferred using the complete nucleotide sequence of the cytochrome b gene. The inferred molecular phylogeny included representatives from Central European, Asian and North African species, and is highly congruent with previous phylogenies based on osteological characters. Iberian cyprinids were grouped into only five, very speciose lineages (with the exception of the monotypic Anaecypris): Barbus, Luciobarbus, Chondrostoma, Leuciscus and Anaecypris. The existence of such a relatively small number of Iberian cyprinid lineages can be explained by the historical isolation of the Iberian Peninsula. North African and Asian barbels are the sister group of Iberian Luciobarbus, supporting a south-eastern route of colonization of the Iberian Peninsula for this subgenus. With leuciscins, Anaecypris hispanica was considered a relict species as it could not be related to any other Iberian cyprinid. The phylogenetic relationships among the main lineages of Iberian cyprinids based on cytochrome b sequence data supported the traditional division of the Cyprinidae into two subfamilies: Cyprininae and Leuciscinae.     

Systematics and biogeography of Klasea (Asteraceae–Cardueae) and a synopsis of the genus

Klasea , traditionally treated as a section in Serratula , is now widely accepted at the generic level. A classification of the genus is presented here, accommodating the 46 species in ten sections based on nuclear ribosomal DNA external and internal transcribed spacer sequence data and morphology. New combinations for five species and ten subspecies are published, and a new hybrid species is described. The genus ranges from the Iberian Peninsula and north Africa through southern and eastern Europe, west and central Asia to the Himalayas, and the Far East of Russia and China. The ancestral area is in west Asia, most probably eastern Anatolia and northern and western Iran. In this region, representatives of all sections are present. The largest section Klasea diversified most likely in the mountains of central Asia. A key to all Klasea species is provided.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 152 , 435–464.     

Monograph of Carex section Schiedeanae (Cyperaceae): Unexpected taxonomic and ecological diversity in a Mexican sedge clade

Carex section Schiedeanae (subg. Euthyceras) has long been thought to comprise at most five species of Mexican sedges. Our morphological studies in the field and herbarium, however, supported by more recent DNA sequencing work, demonstrate that the section is, in fact, three times as species-rich, making it one of the largest sections in Mexico. Its 15 species range from Chihuahua to Tamaulipas south to Veracruz and Oaxaca; a single species, Carex muriculata, extends into the United States. The section has unusual inflorescences with simple (except in Carex complexa), uniformly androgynous, sessile spikes, utriculiform cladoprophylls, and perigynia with a unique combination of pubescence types composed of papillae; unicellular, simple hairs; and (in many species) multicellular tubercles. We recognize five previously described species (C. muriculata,Carex perstricta,Carex schiedeana, Carex stellata, and Carex vizarronensis) and describe 10 additional species (Carex angustilepis, Carex cabralii, Carex complexa, Carex dentata, Carex gypsophila, Carex mesophila, Carex paneroi, Carex planilamina, Carex revoluta, and Carex tehuacana). We have expanded the morphological features used in the section to include particularly anther, staminate scale, and style features, and more detailed micromorphology of the perigynia, especially indumentum. In total, 13 of the 15 species recognized herein, with a total of 32 individuals, were examined for ETS and ITS for phylogenetic reconstruction. Phylogenetically, the xeric species (plus C. paneroi) form a grade leading to a clade of the more mesophytic species adapted to forest understories, plus the unique cliff dwelling C. complexa.     

Evidence of a cenozoic Betic-Kabilian connection based on freshwater fish phylogeography (Luciobarbus, Cyprinidae)

The phylogenetic relationships among 26 species of the subgenus Luciobarbus were examined through comparison of the complete sequence of the mitochondrial genes ATPase 6 and 8 and cytochrome b. The monophyletic condition of the Luciobarbus subgenus was confirmed by several data treatment methods. Findings indicate a closer relationship among species inhabiting Caucasian, Greek, and North African areas than between the latter and those of the Iberian region. The isolation of these main clades is postulated to have occurred after the Messinian salinity crisis (5.5 MY ago) when the Iberian Peninsula broke away from the African continent. Based on a calibrated molecular clock proposed herein, the subsequent splitting of this subgenus corresponds to the isolation of the other three areas (Caucasian, Greek, and North African) about 4.5 MY ago, which would have interrupted any possible gene flow between lineages. However, the present data indicate that the North African Kabilies Mountain area maintained its contact with the south Iberian Peninsula for a longer period.     

Systematics and evolution of sections of the subgenus <Emphasis Type="Italic">Rohrbachia</Emphasis> (Kronf. ex Riedl) A. Krasnova of the hydrophilic genus <Emphasis Type="Italic">Typha</Emphasis> L. (Typhaceae) in Eurasia

Systematics of the three modern sections of the subgenus Rohrbachia of the genus Typha (Typhaceae) and reconstruction of evolution in different geological periods of the Cenozoic Era in Eurasia have been examined. The section Minimae was formed in the Paleogene (Eocene) on littorals of the Angarida–Beringian area; the sections Turanicae and Mongolica were formed on the border of the Paleogene–Neogene in Central Asia (Middle Asia, North China, and Manchuria).     

Epidermal micromorphology of the genus Festuca L. (Poaceae) in the Iberian Peninsula

Micromorphological features of the leaf-blade, lemma, and palea were examined under scanning electron microscopy (SEM) for 64 species of the genus Festuca L. occurring in the Iberian Peninsula. The objective of the study was to survey these characters for the first time in the whole of the genus Festuca in this region, to assess their taxonomic value, and to compare the variation with recent DNA sequence-based phylogenies. The principal features include the frequency, shape, and distribution of silica bodies, short cells, crown cells, and prickles, and the morphology of long cells. The abaxial lemma surface shows the most important taxonomical and phylogenetic characters. The variation observed was consistent with the subgenera and sections identified in molecular studies.     

Allozyme variation of African and Iberian populations of the genus Cobitis

African and Iberian populations of the genus Cobitis exhibit high morphological polymorphism and their taxonomy is unclear. In seven such populations the percentage of polymorphic loci ranged between _ P =0.08–0.48 and mean heterozygosity per locus H =0.005–0.073. The lowest Nei genetic distance between African and Iberian populations of Cobitis was D =0.250. The populations separated with the formation of the Gibraltar strait 5 million years ago. C. calderoni is a well-differentiated branch from the rest of the Iberian populations of the genus. Our analysis supports the concept that C. paludica populations and C. maroccana are referable to the subgenus Iberocobitis . These African and Iberian populations represent two taxa, C. maroccana Pellegrin, 1929 and C. paludica De Buen, 1930. C. maroccana and C. paludica populations are monophyletic and the African population is the most differentiated genetically. The Arrago population of C. paludica is genetically the most differentiated of the Iberian populations. Our allozyme data do not support the previous consideration of the Albufera lagoon population as a different species from C. paludica .     

Pollen morphology of Xanthium L. (s.l.) (Asteraceae,Asteroideae, Heliantheae,Ambrosiinae) in the Iberian Peninsula - a palynotaxonomic approach to a poisonous,allergenic and invasive genus

Abstract

The pollen morphology of the Iberian taxa of the genus Xanthium s.l. and of the non-Iberian species X. ambrosioides was studied to help to identify these poisonous, allergenic and invasive taxa and clarify their taxonomic relationships. In addition, the question of the existence and structure of a mesoaperture in the apertural system of Xanthium was also considered. The pollen grains of sections Acanthoxanthium and Xanthium clearly differ by their characters, which supports the separation of both taxa and allows for their distinction based on pollen morphology. It is suggested that the morphological and palynological differences between Xanthium and Acanthoxanthium, together with the lack of hybridisation between the two sections and the monophyly of Acanthoxanthium, could lead to the elevation of the latter to the generic status. The two species of section Xanthium present in the Iberian Peninsula (X. orientale and X. strumarium) and of section Acanthoxanthium are well separated by their palynological features. Likewise, the Iberian subspecies of Xanthium, X. orientale subsp. orientale and X. orientale subsp. italicum are palynologically distinguishable. However, no significant difference could be found between the pollen of X. strumarium subsp. strumarium and of X. strumarium subsp. brasilicum. The pollen data do not support the hypotheses of a hybrid origin for X. strumarium subsp. brasilicum. A mesoaperture intersecting the foot layer and the upper part of the endexine is present in both sections, which is congruent with the results already found for other Asteroideae.