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Aim: Contemporary species’ distributions are shaped by both geography and historical events, such as extinction, diversification in specific areas and long-distance dispersals. In the most diverse family of lichen-forming fungi, Parmeliaceae, the Hypogymnioid clade, is an example of an evolutionary lineage comprised of species occurring in temperate to subpolar regions in both hemispheres. Here, we elucidate the timing of diversification events and the impact of historical events on the species distribution in this lineage. Location: Worldwide. Taxon: Genera Arctoparmelia, Brodoa, Hypogymnia and Pseudevernia (Parmeliaceae). Methods: Our sampling focused on the most diverse genus of the clade, Hypogymnia, including c. 70% of the described species. We reconstructed phylogenetic relationships using a multi-locus data set, estimated divergence times, and inferred ancestral distributions. Results: Our analyses suggest that the ancestor of the Hypogymnioid clade occurred in the Holarctic. In each of the four genera, all recovered as monophyletic here, diversification have occurred largely during the Miocene and Pliocene. A number of currently accepted species did not form monophyletic groups, especially in cases where specimens were collected from distinct geographic areas, with multiple, distinct clades corresponding to the geographic region of origin. Our results suggest that only a very few species in the Hypogymnioid clade have cosmopolitan distributions, all of which reproduces using vegetative propagules including both symbiotic partners. Main conclusions: While the diversification occurred predominantly in the Northern Hemisphere during the Miocene, a long-distance dispersal event from the Northern to the Southern Hemisphere, resulted in diversification of a clade of species largely restricted to the Southern Hemisphere. Similar to other groups in this diverse family, our study highlights the need for re-evaluation of species boundaries among members of the Hypogymnioid clade.

Hypotrachyna is a speciose genus of primarily tropical and oceanic lichen-forming fungi. It includes species with distinct distribution patterns, such as pantropical, restricted and disjunct species. We used a dataset of mitochondrial SSU, nuclear ITS and LSU ribosomal DNA from 89 specimens to study the historical biogeography of the genus. We employed Bayesian and maximum likelihood approaches for phylogenetic analyses, a likelihood-based approach to ancestral area estimation, and a Bayesian approach to estimate divergence times of major lineages within the genus based on molecular evolutionary rates for ITS and a secondary calibration point at the Hypotrachyna clade – Parmeliopsis split. Our analyses suggest that the genus might have originated in the Neotropics during the Eocene and that the split of major lineages happened primarily during the Eocene and Oligocene. The major diversification within those clades is estimated to have occurred during the Miocene. Pantropical species distributions are explained by long-distance dispersal. A number of currently accepted species were found to be non-monophyletic, illustrating that the delimitation of species in the genus needs attention.

From a mineralogical point of view the La Union ore field (SE Spain) can be regarded as an oddity as some of its hydrothermal, stratabound type deposits display an extremely unusual paragenesis comprising magnetite ± greenalite ± minnesotaite ± siderite and galena ± sphalerite (IOSC – LZS). Recent mineralogical studies have shown that this paragenesis is also present at the neighboring Mazarron ore field. These ore fields share a similar geologic setting, involving metamorphic and sedimentary rocks (Paleozoic s.l. to Permian) hosting late Miocene high‐K calc‐alkaline volcanic and subvolcanic rocks. The latter have andesitic to dacitic composition, and triggered hydrothermal activity and ore deposition. This study discusses the detailed mineral chemistry of magnetite samples from Mazarron and La Union and provides some hints for the origin of the IOSC – LZS paragenesis. We performed electron microprobe (EPMA) analyses in magnetite samples from La Union and Mazarron to determine the contents of minor and trace elements (Zn, Ni, Mn, Cr, V, K, Ca, Ti, Al, Si, Mg). Given that some results fell below the detection limit for the EPMA instrumental conditions we used robust regression on order statistics (robust ROS), with the NADA package in R to deal with these data sets. The Ca + Al + Mn contents in magnetites from San Cristobal and Emilia are equivalent to the mean contents of those of IOCG, Kiruna, BIF, Cu porphyry, skarn, VMS, hydrothermal and clastic Pb–Zn deposits, but they are low in Ti + V and Ni + Cr. The principal components analysis indicates that Zn, Ni, Cr, V, K, Ca, Ti, and Mg are roughly collinear, therefore correlated, being also independent from Fe. Besides, the function discriminant analysis of data shows that the magnetites from Emilia and San Cristobal cluster in differentiated groups, thus probably reflecting some differences related to the distance to the magmatic source that triggered the hydrothermal system: proximal in the case of San Cristobal and distal in Emilia.

The classification of the legume family proposed here addresses the long-known non-monophyly of the traditionally recognised subfamily Caesalpinioideae, by recognising six robustly supported monophyletic subfamilies. This new classification uses as its framework the most comprehensive phylogenetic analyses of legumes to date, based on plastid matK gene sequences, and including near-complete sampling of genera (698 of the currently recognised 765 genera) and ca. 20% (3696) of known species. The matK gene region has been the most widely sequenced across the legumes, and in most legume lineages, this gene region is sufficiently variable to yield well-supported clades. This analysis resolves the same major clades as in other phylogenies of whole plastid and nuclear gene sets (with much sparser taxon sampling). Our analysis improves upon previous studies that have used large phylogenies of the Leguminosae for addressing evolutionary questions, because it maximises generic sampling and provides a phylogenetic tree that is based on a fully curated set of sequences that are vouchered and taxonomically validated. The phylogenetic trees obtained and the underlying data are available to browse and download, facilitating subsequent analyses that require evolutionary trees. Here we propose a new community-endorsed classification of the family that reflects the phylogenetic structure that is consistently resolved and recognises six subfamilies in Leguminosae: a recircumscribed Caesalpinioideae DC., Cercidoideae Legume Phylogeny Working Group (stat. nov.), Detarioideae Burmeist., Dialioideae Legume Phylogeny Working Group (stat. nov.), Duparquetioideae Legume Phylogeny Working Group (stat. nov.), and Papilionoideae DC. The traditionally recognised subfamily Mimosoideae is a distinct clade nested within the recircum-scribed Caesalpinioideae and is referred to informally as the mimosoid clade pending a forthcoming formal tribal and/or clade-based classification of the new Caesalpinioideae. We provide a key for subfamily identification, descriptions with diagnostic charactertistics for the subfamilies, figures illustrating their floral and fruit diversity, and lists of genera by subfamily. This new classification of Leguminosae represents a consensus view of the international legume systematics community; it invokes both compromise and practicality of use.

WOS: 000365393000021, PubMed ID: 26299211, We studied the evolutionary history of the Parmeliaceae (Lecanoromycetes, Ascomycota), one of the largest families of lichen-forming fungi with complex and variable morphologies, also including several lichenicolous fungi. We assembled a six-locus data set including nuclear, mitochondrial and low-copy protein-coding genes from 293 operational taxonomic units (OTUs). The lichenicolous lifestyle originated independently three times in lichenized ancestors within Parmeliaceae, and a new generic name is introduced for one of these fungi. In all cases, the independent origins occurred c. 24 million yr ago. Further, we show that the Paleocene, Eocene and Oligocene were key periods when diversification of major lineages within Parmeliaceae occurred, with subsequent radiations occurring primarily during the Oligocene and Miocene. Our phylogenetic hypothesis supports the independent origin of lichenicolous fungi associated with climatic shifts at the Oligocene-Miocene boundary. Moreover, diversification bursts at different times may be crucial factors driving the diversification of Parmeliaceae. Additionally, our study provides novel insight into evolutionary relationships in this large and diverse family of lichen-forming ascomycetes., Spanish Ministerio de Ciencia e Innovacion [CGL2010-21646/BOS, CGL2011-25003, CGL2013-42498-P]; Universidad Complutense-Banco Santander [GR3/14]; Comunidad Autonoma de Madrid [REMEDINAL S-2009/AMB-1783]; National Science Foundation [DEB-0949147]; Swedish Research Council [VR621-2009-5372, VR 621-2012-3990]; Academy of Finland [1133858]; Estonian Science Foundation [9109]; European Regional Development Fund (Center of Excellence FIBIR); German Academic Exchange Service (DAAD); Tromso University Museum, We are indebted to Adriano A. Spielmann, Joel Mercado, Robert Egan, Udeni Jayalal and Daniel Sanchez Mata for providing a few samples, and to Rosario G. Gavilan for assisting in collecting expeditions. We also thank editors and anonymous reviewers for valuable comments and suggestions. This work was supported by the Spanish Ministerio de Ciencia e Innovacion (projects CGL2010-21646/BOS, CGL2011-25003 and CGL2013-42498-P), the Universidad Complutense-Banco Santander (GR3/14), Comunidad Autonoma de Madrid (REMEDINAL S-2009/AMB-1783), the National Science Foundation ('Hidden diversity in parmelioid lichens'; DEB-0949147), the Swedish Research Council (VR621-2009-5372 and VR 621-2012-3990), the Academy of Finland (grant 1133858), the Estonian Science Foundation (grant 9109), and the European Regional Development Fund (Center of Excellence FIBIR). G.S. was supported by a fellowship from the German Academic Exchange Service (DAAD). I.K.K.T. was supported by a grant from Tromso University Museum.

The Mediterranean region, comprising the Mediterranean Basin and the Macaronesian Islands, represents a center of diversification for many organisms. The genetic structure and connectivity of mainland and island microbial populations has been poorly explored, in particular in the case of symbiotic fungi. Here we investigated genetic diversity and spatial structure of the obligate outcrossing lichen-forming fungus Parmelina carporrhizans in the Mediterranean region. Using eight microsatellite and mating-type markers we showed that fungal populations are highly diverse but lack spatial structure. This is likely due to high connectivity and long distance dispersal of fungal spores. Consistent with low levels of linkage disequilibrium and lack of clonality, we detected both mating-type idiomorphs in all populations. Furthermore we showed that the Macaronesian Islands are the result of colonization from the Mediterranean Basin. The unidirectional gene flow, though, seemed not to be sufficient to counterbalance the effects of drift, resulting in comparatively allelic poor peripheral populations. Our study is the first to shed light on the high connectivity and lack of population structure in natural populations of a strictly sexual lichen fungus. Our data further support the view of the Macaronesian Islands as the end of the colonization road for this symbiotic ascomycete.

The importance of the anatomy and fine morphology of reproductive structures for the systematics of the familyParmeliaceaeis highlighted by the new genusNotoparmelia, described here for Australasian species ofParmelia. These species were known to form a monophyletic lineage but correlated characters for its delimitation were lacking. A major characteristic used here for the circumscription of this genus is the overlooked apothecial anatomy. The proper exciple is reduced to one layer formed exclusively by large hyphae without any clear branching pattern and embedded in an abundant polysaccharide matrix. This feature differs from the rest ofParmeliaceaewhich have a stratified three-layered proper exciple composed of a thin hyaline layer, intermediate layer and basal cortex-like structure. The anatomy of proper exciple, together with the thickness of spore walls and other morphological characters such as lobe morphology, pseudocyphellae and rhizine type, allow a precise diagnosis of the new genus. The area of distribution is also useful for characterizing this genus. Sixteen new combinations are proposed.

Se evaluaron seis regiones de ADN como posibles códigos de barras de ADN para Hongos, el segundo reino más grande de vida eucariota, por un consorcio multinacional y de laboratorios múltiples. La región de la mitocondria se utilizó la subunidad 1 de la citocromo c oxidasa como código de barras animal excluido como un marcador potencial, porque es difícil de amplificar en hongos, a menudo incluye intrones grandes y puede ser insuficientemente variable. Tres subunidades del cistrón de ARN ribosómico nuclear fueron en comparación con las regiones de tres genes de codificación proteica representativos (la subunidad más grande de la ARN polimerasa II, la segunda más grande subunidad de la ARN polimerasa II y mantenimiento de minicromosomas proteína). Aunque las regiones de genes que codifican proteínas a menudo tenían un porcentaje más alto de identificación correcta en comparación con ribosomal marcadores, baja amplificación por PCR y éxito de secuenciación eliminados ellos como candidatos para un código de barras universal para hongos. Entre la regiones del cistrón ribosómico, el espaciador transcrito interno (ITS) tiene la mayor probabilidad de identificación exitosa para la gama más amplia de hongos, con la brecha de código de barras más claramente definida entre la variación inter e intraespecífica. El nuclear subunidad ribosómica grande, un marcador filogenético popular en ciertos grupos, tuvieron una resolución de especies superior en algunos grupos taxonómicos, como los primeros linajes divergentes y las levaduras ascomicetas, pero por lo demás era ligeramente inferior al ITS. El ribosomal nuclear La subunidad pequeña tiene una resolución pobre a nivel de especie en los hongos. SU será propuesto formalmente para su adopción como el código de barras fúngico principal marcador al Consorcio para el Código de Barras de la Vida, con la posibilidad de que se desarrollen códigos de barras suplementarios para grupos taxonómicos estrechamente circunscritos., Six DNA regions were evaluated as potential DNA barcodes for Fungi, the second largest kingdom of eukaryotic life, by a multinational, multilaboratory consortium. The region of the mitochondrial cytochrome c oxidase subunit 1 used as the animal barcode was excluded as a potential marker, because it is difficult to amplify in fungi, often includes large introns, and can be insufficiently variable. Three subunits from the nuclear ribosomal RNA cistron were compared together with regions of three representative proteincoding genes (largest subunit of RNA polymerase II, second largest subunit of RNA polymerase II, and minichromosome maintenance protein). Although the protein-coding gene regions often had a higher percent of correct identification compared with ribosomal markers, low PCR amplification and sequencing success eliminated them as candidates for a universal fungal barcode. Among the regions of the ribosomal cistron, the internal transcribed spacer (ITS) region has the highest probability of successful identification for the broadest range of fungi, with the most clearly defined barcode gap between inter- and intraspecific variation. The nuclear ribosomal large subunit, a popular phylogenetic marker in certain groups, had superior species resolution in some taxonomic groups, such as the early diverging lineages and the ascomycete yeasts, but was otherwise slightly inferior to the ITS. The nuclear ribosomal small subunit has poor species-level resolution in fungi. ITS will be formally proposed for adoption as the primary fungal barcode marker to the Consortium for the Barcode of Life, with the possibility that supplementary barcodes may be developed for particular narrowly circumscribed taxonomic groups.

Parmelina tiliacea is a common, widely distributed species in south-western Europe, easily identifiable by morphology and much used as an air pollution bioindicator in many regions. A molecular phylogenetic survey of samples from many geographical areas, using Maximum Parsimony and Bayesian inference of nuITS and mtLSU rDNA regions, revealed a group of samples geographically restricted to a small region of the Iberian Peninsula and genetically separated from the other P. tiliacea specimens studied. These samples are morphologically indistinguishable from P. tiliacea, apart from subtle anatomical characters in the ascomata (hyphae of the exciple and ascospore width), which are frequently absent. Although geographically different, the two taxa occupy similar habitats and are even sympatric in some areas, indicating that they do not exchange genetic material. This previously overlooked, and apparently endemic lineage, is described as P. cryptotiliacea sp. nov., and the name Lichen tiliaceus is epitypified by a sequenced specimen to fix the application of Parmelina tiliacea to the widespread genotype. A second unexpected result was the discovery that the morphologically distinct P. pastillifera was nested within P. tiliacea. These two cases stress the need to use molecular tools to elucidate species concepts even within widespread morphologically well-characterized macrolichens. Such investigations are necessary to improve our understanding and estimation of biodiversity, and to facilitate the development of sound biodiversity conservation strategies for lichens.

The species delimitation in fungi is currently in flux. A growing body of evidence shows that the morphology-based species circumscription underestimates the number of existing species. The large and ever growing number of DNA sequence data of fungi makes it possible to use these to identify potential cases of hidden species, which then need to be studied with extensive taxon samplings. We used Parmeliaceae, one of the largest families of lichenized fungi as a model. Intra- and interspecific distances derived from maximum-likelihood phylogenetic trees inferred from 491 nuclear ITS rDNA sequences were examined for five major clades of parmelioid lichens. The intra- and interspecific distances were well separated in most cases allowing the calculation of a threshold, with exceptions of highly deviating distances in a few cases. These situations are shown to be taxa in which the current delimitation needs revision. Thus the analysis of the distance distributions is shown to be a powerful tool for identifying species complexes.

Phylogenetic relationships of Adenocarpus species were assessed by sequence analyses of the ITS and 3' end of ETS (nrDNA) regions and the trnL-trnF intergenic spacer (cpDNA). The nrDNA analysis recovered four main clades within Adenocarpus. Clades 1―3 comprise morphological and molecularly well-differentiated species that occur in the western Mediterranean realm. Clade 4 comprises three distinctive endemic species from the Canary Islands, an endangered Algerian endemic, and a set of species close to A. complicatus with distinct geographical areas but with little molecular variation between taxa. The latter species are centred in the Iberian Peninsula but also distributed along the Mediterranean basin (from the Iberian Peninsula to Turkey). The phylogenetic position of A. mannii, a morphologically distinctive species with a unique afro-tropical distribution, is not unequivocally resolved, being sister to clade 3 in the ETS tree but to clade 4 in the ITS tree. Rates of evolution and ages of divergence of major groups were estimated using nonparametric rate smoothing with a fixed calibration point of 41.2 Ma and constraining the node of the separation of Anarthrophyllum and Mediterranean Genisteae at a minimum age of 13.6 and a maximum of 25.8 Ma. The estimated ages indicate that the diversification of Adenocarpus and first separation of two lineages may have occurred during the onset and the wake of the profound Middle Miocene cooling episode. One of the lineages separated rapidly into two groups (clades 1 and 2) while the second one led to clade 3, A. mannii and clade 4. Clades 1, 2 and 3 differentiated into the present species soon after these major events whereas clade 4 radiated much later, during the Pliocene.

The monotypic genera Omphalodiella and Placoparmelia were described from Patagonia (Argentina). The latter is characterized by placodioid species, morphologically similar to some subcrustose, brown Xanthoparmelia species, whereas Omphalodiella is a peltate lichen. Analyses of ribosomal DNA sequences supported their placement in the parmelioid clade. Both genera are nested within Xanthoparmelia. Affinites to the latter genus are congruent with the presence of Xanthoparmelia-type lichenan in the hyphal cell walls and an arachiform vacuolar body in the ascospores, two key characters typical of Xanthoparmelia. Consequently we propose to place the generic names in synonymy with Xanthoparmelia and transfer the two species to the latter genus as X. patagonica and X. peltata comb. nov.

Morphological and phylogenetic relationships of the worldwide Mediterranean lichen forming fungus, Parmelina quercina, have been studied. Specimens from western Europe, western North America and southern Australia were analysed using molecular data (nuITS rDNA, nuLSU rDNA and mtSSU rDNA) and selected morphological features (upper cortex maculae, scanning electron microscopy examination of the epicortex, ascospores and conidia shape and size, and amphithecial retrorse rhizines). The results conclusively reveal that: (1) there is not one single species but four separate species in the Mediterranean or sub Mediterranean areas of the world. Parmelina quercina and Parmelina carporrhizans (Euroasiatic species), Parmelina coleae sp. nov. (North America) and Parmelina elixia sp. nov. (Australia); (2) largely debated P. carporrhizans is not a synonym of P. quercina but supported as a valid species circumscribed to Macaronesic relict sites; (3) the geographical isolation of the Australian population is correlated with a large genetic distance; (4) morphological characters (ascospores and conidial variability and thallus epicortex) correlate with the phylogenetic hypothesis; (5) the new or revalidated species within Parmelina quercina are not cryptic species but morphologically recognizable taxa. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91, 455–467.

Sections of apothecia were used to study the internal morphology of ascospores in the largest monophyletic clade within Parmeliaceae composed of Xanthoparmelia and related genera. The results were compared with fertile representative species of most other parmelioid clades. All the Xanthoparmelia species had spores with a single smooth vacuole, which was peanut-shaped, with different degrees of constriction in the equatorial plane. This differs from the ellipsoid vacuole of other parmelioids. In the Xanthoparmelia clade, sexual reproduction seems much more common than in other parmelioids. Thus, we suggest that the presence of this unique spore morphology might contribute to the evolutionary success of this monophyletic group. Further, the discovery of this useful ascospore character demonstrates that detailed ascospore morphological studies significantly enhance molecular phylogenetic analyses. Ascospore features may be more taxonomically significant in Parmeliaceae than hitherto considered.

The San Cristóbal–Perules mining site in Mazarrón in southeast Spain was subjected to about a hundred years of intense mining activity for lead, silver, and zinc. Metallurgical operations (smelting, calcination, gravity concentration) carried out during the late nineteenth century–early twentieth century induced significant land transformation, and the most conspicuous wastes of this period consist of a chaotic piling of ‘old’ tailing deposits. Later on, during the mid-twentieth century, ‘modern’ tailings resulting from froth flotation were accumulated filling small valleys; these latter valley-fill tailings rose sequentially according to the upstream construction method, progressively raising the level of the dam during the process. Once abandoned, both types of tailing deposits underwent severe erosion, resulting in a mosaic of erosional and sedimentary landforms developed upon (e.g., gully formation) and within them (e.g., piping). We made an inventory and classification of these landforms. Our study shows the geomorphic work to reestablish a new steady state between the tailings deposits and the local erosive conditions. This scenario implies several hazards related to the extremely high heavy metal contents of these tailings and the geomorphic instability of the deposits. We also quantified the tailings tonnage and erosion that occurred at one of the tailings dams (El Roble). As shown by an oblique aerial photograph taken in 1968, this dam had a terraced topography, whereas in 2013 this morphology had evolved into a badland-type relief with deep parallel gullies. By recognizing and surveying specific, remnant points along the benches and outslopes of the older terraced topography, we were able to build up a first digital elevation model (DEM1) reflecting the initial topography. A second DEM, this time showing the present topography, allowed quantification of erosion via Material Loss = DEM1 − DEM2. This yields an erosion rate (1968–2009) of 151.8 Mg (MT) ha− 1 y− 1, which matches well typical values for erosion of mined areas, commonly above 100 Mg (MT) ha− 1 y− 1. Abandoned mine tailing deposits are extremely common in the semiarid scenarios of the SW USA, Australia, Chile, and Peru. Given the similarities of these scenarios with SE Spain, the example from Mazarrón may provide useful new insights regarding the erosion and geomorphic evolution of such tailing deposits. These matters should be addressed in key environmental actions such as mine closure plans and land reclamation projects. A solution may come via restoration of these deposits through landform design involving the building up of stable mature landscapes, which in turn can withstand erosion much more easily.

Genetic distances among populations and phylogenetic hypothesis concerning origin and diversification of Cytisus purgans s.1. are inferred by analysis of the ITS and ETS regions (nrDNA) and the trnL-trnF intergenic spacer (cpDNA). Cytisus purgans s.1., a shrub of western Mediterranean mountains, consists of two evolutionary units: C. balansae (Boiss.) Ball (Morocco, Algeria, southern Spain) and C. oromediterraneus Rivas Mart. & al. (France, Portugal, Andorra, central and northern Spain). The two units are related to different Cytisus groups, in Bayesian and Maximum Parsimony analyses, and may be genetically unrelated, their morphological similarities being the result of convergence. However, the incongruence in the position of both species in the ITS tree and the ETS and trnL-trnF trees suggests an alternative explanation. Genetic differentiation could be due to lineage sorting of allelic polymorphism in the ITS region of the common ancestor, and the morphological similarities the result of morphological parallelism. Cytisus balansae conserves ITS sequences related to old Cytisus taxa; rapid radiation may have led to a western Mediterranean lineage in which C. oromediterraneus originated.

Genetic variation in 27 populations of Ulex species from southern Spain and northern Morocco (Betic-Rif arc) was assessed using 11 chloroplast microsatellite (cpSSR) markers, which revealed 47 different haplotypes. These nonrecombinant, haploid markers allow measurement of genetic variation in closely related species of Ulex where molecular phylogenetic analyses have not provided a clear view of interspecific relationships. Discriminant analysis indicates that the haplotypes are useful to differentiate among species, and analysis of molecular variance (AMOVA) shows high levels of differentiation among populations. The minimum spanning tree (MST), that represents the connections between the haplotypes, suggests that the eastern Rifean U. africanus haplotypes are more genetically related than those from southern Spain. The latter may have lost genetic diversity while colonizing new habitats, eventually differentiating into U. baeticus and U. scaber. Hybridization between these populations, followed by polyploidization, may have originated the tetraploids (U. congestus and U. borgiae) that colonized new habitats associated with acidic rocks. Separate groupings of U. scaber populations may indicate multiple origins from different stocks. Diversification in this group of Ulex species could be related to the opening of the Alboran Sea by Middle Miocene, when the populations from Morocco and Spain became isolated from each other.

Phylogenetic relationships of Genista and related genera (Teline, Chamaespartium, Pterospartum, Echinospartum, Ulex, Stauracanthus and Retama) were assessed by the analysis of sequences of the nrDNA internal transcribed spacer (ITS region), and the cpDNA trnL-trnF intergenic spacer. The tree obtained by combining both sets of data indicates the existence of three lines of diversification within Genista, that correspond to three subgenera: Genista, Phyllobotrys and Spartocarpus, however, each of these lineages encompass also species of the related genera Echinospartum, Teline, Retama, Chamaespartium, Pterospartum, Ulex, Stauracanthus. The molecular data do not support division of these subgenera into taxonomical units at the sectional level; only sections Genista and Spartocarpus are monophyletic groups. The sequences of both regions are also informative at the specific level, grouping morphologically related species (e.g. the G. cinerea aggregate). The molecular data have also helped to clarify the position of taxa whose relationships were not well established (e.g. G. valdes-bermejoi). The relationships of related genera that belong to the Genista lines of diversification have also been investigated. Echinospartum splits into two separate clades matching the separation of two ecological and caryological differentiated groups. Teline also forms two groups, both placed near to Genista subgenus Genista, but that separated from the main core of the group. Retama, morphologically well differentiated from Genista, is close to Genista subgenus Spartocarpus. Chamaespartium and Pterospartum do not form a monophyletic group. Chamaespartium is closer to Genista subgenus Genista, whereas Pterospartum stands close to: 1) Genista subgenus Spartocarpus (particularly, sect. Cephalospartum); and 2) the Ulex-Stauracanthus clade (a terminal derivative of Genista subgenus Spartocarpus). Cases of incongruence (e.g. Echinospartum, Chamaespartium, Teline) between the trees obtained from the two molecular markers, may be indicating hybridisation and/or introgression between different lines of Genisteae.

Phylogenetic relationships of Cytisus and allied genera (Argyrocytisus, Calicotome, Chamaecytisus, Cytisophyllum, and Spartocytisus) were assessed by analysis of sequences of the nrDNA internal transcribed spacer (ITS) and the cpDNA trnL-trnF intergenic spacer. Genera of the Genista-group (Chamaespartium, Echinospartum, Genista, Pterospartum, Spartium, Teline and Ulex) were included to check the position of Cytisus species transferred to Teline. The tree obtained by combining both sets of data indicates that the Genista and Cytisus groups form two separate clades. Cytisus heterochrous and C. tribracteolatus are more closely related to the Cytisus-group, thus their transfer to Teline is not supported by molecular data. Cytisus fontanesii (syn. Chronanthos biflorus) groups with Cytisophyllum sessilifolium and Cytisus heterochrous within the Cytisus-group. Similarly, Argyrocytisus battandieri falls within the Cytisus-group as a well differentiated taxon. All these taxa seem to have early diverged from the Cytisus-group. Their taxonomic rank should be reconsidered to better reflect their phylogenetic separation from Cytisus. On the contrary, Chamaecytisus proliferus and Spartocytisus supranubius enter in the main core of Cytisus, and they should better be included in sections of Cytisus (sect. Tubocytisus and Oreosparton, respectively). Sect. Spartopsis is not monophyletic and the position of several species, currently included in this section, deserves reevaluation: C. arboreus aggregate is closely related to C. villosus (sect. Cytisus) and to Calicotome; C. striatus is closely related to Cytisus sect. Alburnoides; and the position of C. commutatus (incl. C. ingramii) remains unclear. The relationships and positioning of several minor taxa (C. transiens, C. megalanthus, and C. maurus) are also discussed.

Karyological information on Cytisus species indicates at least two chromosome numbers for most of the taxa. This instability is a striking karyological feature of Cytisus . Chromosome numbers of taxa in Sect. Spartopsis and Sect. Alburnoides, both well represented in Morocco and the Iberian Peninsula, are presented here. We provide the first data on chromosome numbers for the Moroccan taxa C. grandiflorus subsp. barbarus , and subsp. haplophyllus ( n =23, 2 n =46), C. maurus (2 n =48), C. megalanthus ( n =23), C. arboreus subsp. arboreus , subsp. baeticus , and subsp. catalaunicus (2 n =50), C. valdesii ( n =23). New populations from the Iberian Peninsula have been counted C. grandiflorus subsp. grandiflorus (2 n =46), C. scoparius subsp. scoparius ( n =23), C. striatus subsp. eriocarpus ( n =23, 2 n =46), C. multiflorus ( n =23), C. oromediterraneus ( n =23, 24). Our data confirm the instability of the chromosome number in Cytisus . The presence of B chromosomes in C. valdesii and C. megalanthus , as well in other species, is discussed in relation to this instability and previous data. We suggest that instability of the chromosome number within a taxon, and even in the same population, may be related to the breakage of A chromosomes and the appearance of B chromosomes.

Although it is generally accepted that the tribe Genisteae has rather homogeneous pollen, we have found considerable variation in pollen size and morphology in Cytisus and related genera (Argyrocytisus and Chamaecytisus), which match taxonomic groups defined on morphological grounds. The results of Principal Component Analysis of pollen characteristics show the following: Cytisus Sect. Alburnoides is well-delimited, with the smallest pollen grains and the simplest pattern in the tectum, fossulate-perforate to perforate; No separation of C. villosus (Sect. Cytisus) from Sect. Alburnoides is observed; Cytisus Sect. Spartopsis, with the largest pollen grains and reticulate to reticulate-fossulate ornamentation, forms an isolated group; C. fontanesii (Sect. Heterocytisus) stands apart from other Cytisus species by its homogeneous ornamentation and the perforate instead of psilate-punctate margo. Thus, the pollen data supports its transfer to the separated genus Chronanthos; Pollen ornamentation and morphology...

Karyological information about Iberian Genista species is limited because of their wide geographical distribution. It is a major factor in the understanding of a genus whose genetic evolution is mostly related to aneuploidy and euploidy. The chromosome numbers of the following taxa have been counted G. carpetana subsp. carpetana ( n =20), G. cinerascens ( n =12), G. micrantha ( n =18), G. mugronensis subsp. rigidissima ( n =18), G. ramosissima (2 n =48) and G. tinctoria ( n =24). The previously-assumed uniformity of chromosome number ( n =24) for all the subspecies of G. cinerea is supported here by data from subspp. murcica and speciosa . In G. florida , a new chromosome number, n =23, has been found for both subspecies ( florida and polygaliphylla ). A discussion of the discrepancies between these data and previous ones is included. A new level of ploidy has been found in G. tournefortii subsp. tournefortii ( n =32). The data stress the necessity of obtaining chromosomal information from a number of plants of a taxon before its characteristic number is clear, and indicate that more chromosomal data will help the understanding of the cytological diversity within taxa in Genista .

The two subspecies of Ulex europaeus L. (subsp. europaeus and subsp. latebracteatus ) are easily differentiated by the shape and size of their bracteoles, and by their chromosome number. However, in the north-west of the Iberian Peninsula, the areas of both subspecies are not clearly delimited, the shape of the bracteoles is intermediate and the width is close to the diagnostic limit. Thus, allocation of some specimens to either subspecies is diffcult. Recently, a controversy has arisen concerning the intrapopulational uniformity of the chromosome number in U. europaeus from west France and north Spain, and the usefulness of ploidy level and chromosome number as clear-cut indicators of the two subspecies. Thus, the merits of recognizing the subspecies as independent are re-analysed here. Our results for north-west Spain show that the chromosome number is uniform and stable within populations and is clearly related to the morphological characters. Correlation and discriminant analyses show that bracteoles and stomata sizes are highly correlated and are good predictors of ploidy level. Furthermore, the results support the maintenance of two subspecies in U. europaeus , which can be delimited by a combination of morphological characters, chromosome number and geographical distribution.

In traditional morphology-based concepts many species of lichenized fungi have world-wide distributions. Molecular data have revolutionized the species delimitation in lichens and have demonstrated that we underestimated the diversity of these organisms. The aim of this study is to explore the phylogeography and the evolutionary patterns of the Xanthoparmelia pulla group, a widespread group of one of largest genera of macrolichens. We used a dated phylogeny based on nuITS and nuLSU rDNA sequences and performed an ancestral range reconstruction to understand the processes and explain their current distribution, dating the divergence of the major lineages in the group. An inferred age of radiation of parmelioid lichens and the age of a Parmelia fossil were used as the calibration points for the phylogeny. The results show that many species of the X. pulla group as currently delimited are polyphyletic and five major lineages correlate with their geographical distribution and the biosynthetic pathways of secondary metabolites. South Africa is the area where the X. pulla group radiated during the Miocene times, and currently is the region with the highest genetic, morphological and chemical diversity. From this center of radiation the different lineages migrated by long-distance dispersal to others areas, where secondary radiations developed. The ancestral range reconstruction also detected that a secondary lineage migrated from Australia to South America via long-distance dispersal and subsequent continental radiation.

Summary The pollen morphology in the Genisteae is very homogeneous despite the great range of variability displayed by this tribe in macromorphology, chromosome numbers and ploidy levels. However, the wall structure of pollen grains of Ulex, Stauracanthus and Genista shows important differences which allow recognition of four types of exine structure. These types closely correlate with the ploidy levels. Type I, found in most of the diploids, is characterized by a compact structure of the pollen wall, and an infratectum ill- developed and mainly composed of granules. Type II, characterized by an infratectum composed of both granules and columellae. This structure is commonly found in pollen grains of tetraploids. Type III, showing an infratectum composed of well developed columellae which are often formed of pillared granules. Pollen grains of diploids, tetraploids and hexaploids show this type of exine structure. In type IV, all the pollen wall layers are formed of densely packed granules which often for...

There is a long-standing debate on the extent of vicariance and long-distance dispersal events to explain the current distribution of organisms, especially in those with small diaspores potentially prone to long-distance dispersal. Age estimates of clades play a crucial role in evaluating the impact of these processes. The aim of this study is to understand the evolutionary history of the largest clade of macrolichens, the parmelioid lichens (Parmeliaceae, Lecanoromycetes, Ascomycota) by dating the origin of the group and its major lineages. They have a worldwide distribution with centers of distribution in the Neo- and Paleotropics, and semi-arid subtropical regions of the Southern Hemisphere. Phylogenetic analyses were performed using DNA sequences of nuLSU and mtSSU rDNA, and the protein-coding RPB1 gene. The three DNA regions had different evolutionary rates: RPB1 gave a rate two to four times higher than nuLSU and mtSSU. Divergence times of the major clades were estimated with partitioned BEAST analyses allowing different rates for each DNA region and using a relaxed clock model. Three calibrations points were used to date the tree: an inferred age at the stem of Lecanoromycetes, and two dated fossils: Parmelia in the parmelioid group, and Alectoria. Palaeoclimatic conditions and the palaeogeological area cladogram were compared to the dated phylogeny of parmelioid. The parmelioid group diversified around the K/T boundary, and the major clades diverged during the Eocene and Oligocene. The radiation of the genera occurred through globally changing climatic condition of the early Oligocene, Miocene and early Pliocene. The estimated divergence times are consistent with long-distance dispersal events being the major factor to explain the biogeographical distribution patterns of Southern Hemisphere parmelioids, especially for Africa-Australia disjunctions, because the sequential break-up of Gondwana started much earlier than the origin of these clades. However, our data cannot reject vicariance to explain South America-Australia disjunctions.

The tribe Genisteae includes genera of great ecological importance in Mediterranean countries because they are dominant elements of many plant communities. Genetic variation and diversification patterns in Stauracanthus (Genisteae) provide information relevant for the study of the processes of diversification in relation to the environmental history of the western Mediterranean. Nineteen populations of S. boivinii and S. genistoides were assessed by 11 chloroplast microsatellite markers, revealing 44 haplotypes. Both species had different haplotypes and contrasting patterns of karyological, morphological, and genetic variation. In the minimum spanning tree of the haplotypes, AMOVA analysis, and nested clade analysis, S. boivinii had high levels of differentiation and restricted gene flow among populations. Allopatric differentiation occurred between the Moroccan and Iberian populations of S. genistoides, although S. genistoides subsp. spectabilis and subsp. vicentinus had high levels of differentiation among populations (F(ST)), whereas S. genistoides subsp. genistoides had a low F(ST). Genetic patterns are discussed in relation to the Messinian salinity crisis (MSC): hard conditions drove plants to refuge habitats along the Atlantic coast and higher altitude areas in the Moroccan mountains (S. genistoides subsp. spectabilis and S. boivinii). After the MSC, S. boivinii underwent polyploidization and expansion, whereas S. genistoides expanded and continued diversifying into S. genistoides subspp. genistoides and vicentinus.

The Mazarrón epithermal ore deposits (SE Spain) formed in Miocene time in relation to the emplacement of dacitic–rhyodacitic domes. The Pb–(Ag)–Zn ore deposits are of the high sulphidation vein and stockwork type. There are two main mining sites: San Cristóbal–Perules (adjacent to the town of Mazarrón) and Pedreras Viejas. The area is located in the southern realm of a Miocene marine basin surrounded by mountains in its western, eastern and southern flanks. A main seasonal river (Rambla de las Moreras) crosses the basin from North to South and is strongly affected by contaminated sediments from the San Cristóbal– Perules mining site. The mine tailings are extremely rich in Pb (mean=12,400 μg g−1) and Zn (mean=6100 μg g−1), whereas As concentrations are also very high (mean=650 μg g−1). Our geochemical survey also covered anthropic soils, stream sediments and the so-called almagres deposits (red alum calcines). All these environmental matrices have also very high concentrations of Pb, Zn, As, and Cd compared to our local or world baseline concentrations. The calcines are rich in As (mean=450 μg g−1), and the main environmental concern relates to the wrong use of these materials as agricultural soils at discrete sites in the district. Besides, the anthropic soils of the district are very rich in Pb (mean=2550 μg g−1) and Zn (mean=1870 μg g−1). On the other hand, the seasonal river Rambla de las Moreras receives contaminated sediments from the San Cristóbal–Perules old mining site and mine wastes, which results in high concentrations of As (mean=120 μg g−1), Pb (mean=2700 μg g−1) and Zn (mean=1090 μg g−1) in the sediments. These high concentrations of dangerous contaminants in an area close to a town of 35,000 inhabitants and adjacent to significant agricultural activity may pose a risk to human health.

Pollen grains of Ulex are isopolar, 3-zonocolporate with an ill-defined endoaperture, and tectateperforate to reticulate. The pollen grains have distinct internal folds due to the thickening of the endexine along the colpi. The general morphology does not vary substantially from one taxon to another, but the pollen wall structure is very variable. Three types have been recognized based on the stratification: Type I: a compact structure with an ill-developed infratectum mainly composed of granules; Type II: a transitional infratectal layer composed by granules and columellae; and finally, Type III: with a well differentiated columellar infratectum. These structural types correlate to a high degree with the different ploidy levels found in Ulex. Most of the diploid taxa have Type I, most of the tetraploids have Type II, and all of the hexaploids have Type III.

The Rodalquilar mineral deposits (SE Spain) were formed in Miocene time in relation to caldera volcanic episodes and dome emplacement phenomena. Two types of ore deposits are recognized: (1) the El Cinto epithermal, Au–As high sulphidation vein and breccia type; and (2) peripheral low sulphidation epithermal Pb–Zn–Cu–(Au) veins. The first metallurgical plants for gold extraction were set up in the 1920s and used amalgamation. Cyanide leaching began in the 1930s and the operations lasted until the mid 1960s. The latter left a huge pile of ~900,000–1,250,000 m3 of abandoned As-rich tailings adjacent to the town of Rodalquilar. A frustrated initiative to reactivate the El Cinto mines took place in the late 1980s and left a heap leaching pile of ~120,000 m3. Adverse mineralogical and structural conditions favoured metal and metalloid dispersion from the ore bodies into soils and sediments, whereas mining and metallurgical operations considerably aggravated contamination. We present geochemical data for soils, tailings and wild plant species. Compared to world and local baselines, both the tailings and soils of Rodalquilar are highly enriched in As (mean concentrations of 950 and 180 μg g−1, respectively). Regarding plants, only the concentrations of As, Bi and Sb in Asparagus horridus, Launaea arborescens, Salsola genistoides, and Stipa tenacissima are above the local baselines. Bioaccumulation factors in these species are generally lower in the tailings, which may be related to an exclusion strategy for metal tolerance. The statistical analysis of geochemical data from soils and plants allows recognition of two well-differentiated clusters of elements (As–Bi–Sb–Se–Sn–Te and Cd–Cu–Hg–Pb–Zn), which ultimately reflect the strong chemical influence of both El Cinto and peripheral deposits mineral assemblages.

Thallus morphology has traditionally played a major role in the classification of lichenised fungi. We have used a combined dataset of nuITS, nuLSU and mtSSU rDNA sequences to evaluate the phylogenetic relationships between the subcrustose genus Karoowia and the mostly foliose genus Xanthoparmelia. Our phylogenetic analyses using maximum parsimony, maximum likelihood and a Bayesian approach show that Karoowia species do not form a monophyletic group but cluster in different clades nested within Xanthoparmelia. The monophyly of Karoowia either as a separate clade from Xanthoparmelia, or nested within Xanthoparmelia is significantly rejected using alternative hypothesis testing. These results suggest that the usefulness of the phenotypic features used to define Karoowia has been overestimated because the subcrustose growth form has evolved independently in several clades within Xanthoparmelia. Other characters used to circumscribe Karoowia, such as the presence of cylindrical conidia, also occur in Xanthoparmelia, and the differences in rhizine morphology are minimal. Consequently, we propose to reduce Karoowia to synonymy with Xanthoparmelia. The enlarged genus is characterised by the presence of Xanthoparmelia-type lichenan in the hyphal cell walls and the presence of an arachiform vacuolar body in the ascospores. Fifteen new combinations in Xanthoparmelia and the new name Xanthoparmelia mucinae for Karoowia squamatica are made.

Polystichunz Xbicknellii (Christ) Hahne, the hybrid between P. aculeatum (L.) Roth and P. setiferum (Forsk.) Woynar, has been found in many European countries (Sleep, 1975). In Spain this hybrid has been mostly found in the Pyrenees, Iberian Ranges, and Cantabrian Ranges (Salvo et al., 1986). Both fertile plants and their hybrid are morphologically close, and can be identified by differences in frond morphology (Fig. 1) and perispore pattern (Sleep, 1971, 1975; Salvo et al., 1986). Plants of P. setiferum, P. aculeatum and P. Xbicknellii were found growing together in the Spanish Central Pyrenees, thus giving a good opportunity to study the influence of both parents on the hybrid's spore morphology (structure and external morphology of the perispore). The plants are from: Huesca, Cai6n de Aiisclo, 750 m, growing in a beechwood on base-rich soil, 24.6.1987, leg. Fernandez Gonzalez et al. Voucher specimens are kept in the Herbarium of the Faculty of Pharmacy, Universidad Complutense, Madrid (P. setiferum, MAF 135851; P. aculeatum, MAF 135852; P. Xbicknellii, MAF 135853). Spores of the three specimens were suspended in water and cut by means of a freezing microtome following the method of Muller (1973). Sections were transferred to slides and mounted in gum chloral for light microscopy observations, and air dried, mounted on an aluminum stub and gold coated for SEM studies. The differences in the perispore morphology of P. setiferum and P. aculeatum reflect the different structures of their spore wall. The spores of P. setiferum (Fig. 2, A-D) show

WOS: 000285566000008, Parmelioid lichens are a diverse and ubiquitous group of foliose lichens. Generic delimitation in parmelioid lichens has been in a state of flux since the late 1960s with the segregation of the large, heterogeneous genus Parmelia into numerous smaller genera. Recent molecular phylogenetic studies have demonstrated that some of these new genera were monophyletic, some were not, and others, previously believed to be unrelated, fell within single monophyletic groups, indicating the need for a revision of the generic delimitations. This study aims to give an overview of current knowledge of the major clades of all parmelioid lichens. For this, we assembled a dataset of 762 specimens, including 31 of 33 currently accepted parmelioid genera (and 63 of 84 accepted genera of Parmeliaceae). We performed maximum likelihood and Bayesian analyses of combined datasets including two, three and four loci. Based on these phylogenies and the correlation of morphological and chemical characters that characterize monophyletic groups, we accept 27 genera within nine main clades. We re-circumscribe several genera and reduce Parmelaria to synonymy with Parmotrema. Emodomelanelia Divakar & A. Crespo is described as a new genus (type: E. masonii). Nipponoparmelia (Kurok.) K.H. Moon, Y. Ohmura & Kashiw. ex A. Crespo & al. is elevated to generic rank and 15 new combinations are proposed (in the genera Flavoparmelia, Parmotrema, Myelochroa, Melanelixia and Nipponoparmelia). A short discussion of the accepted genera is provided and remaining challenges and areas requiring additional taxon sampling are identified., Spanish Ministerio de Ciencia e Innovacion [CGL2008-01125-E/BOS, CGL2007-64652/BOS]; Ramon y Cajal [RYC02007-01576]; Field Museum; Swedish Research Council [VR 629-2001-5756, VR 621-2003-303, VR 621-2006-3760], We wish to thank Allison Knight, Gary Perlmutter, Ken Sweat, James Lendemer and Gennadii Urbanavichius for providing us with fresh material for our studies. We thank John Pormann and Sean Dilda from the Duke Shared Cluster Resource and Michael Nuhn from the Nano+Bio Center for their support with installing and running the necessary software. This study has been supported financially by the Spanish Ministerio de Ciencia e Innovacion (CGL2008-01125-E/BOS, CGL2007-64652/BOS), and Ramon y Cajal grant (RYC02007-01576) to PKD, a start up fund of The Field Museum to Thorsten Lumbsch, the Swedish Research Council grants VR 629-2001-5756, VR 621-2003-303 and VR 621-2006-3760 to Mats Wedin. We thank the Galapagos National Park for permission to analyze their material. This is publication no. 2001 of the Charles Darwin Research Station.