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Calluna vulgaris (Ericaceae) is the most widespread and prevalent woody species of the European dry heathland, from the African side of the Strait of Gibraltar to northern Europe. However, previous biogeographic analyses of this species have ignored the western Mediterranean region, despite its preponderant role as a major glacial refugium for European biota. Here, we explore the existence of further Pleistocene glacial refugia for C. vulgaris in the Iberian Peninsula and the Strait of Gibraltar area and provide new insights into the post-glacial colonization of Europe by this emblematic heathland species. We carried out phylogeographical analyses of C. vulgaris samples from 44 locations using four plastid markers (matK, trnH-psbA, trnL-F and rpl16). We built palaeodistribution models using MAXENT to identify possible refugia for C. vulgaris during the mid-Pliocene, Last Interglacial (LIG), Last Glacial Maximum (LGM) and mid-Holocene. Our results identified a genetic lineage exclusive to the Strait of Gibraltar area s.l. (south-western Iberian Peninsula and north-western tip of Africa) as the most probable close to the clade of origin of the species. This lineage also had more private haplotypes than more recent, Eurosiberian clades. The obtained data highlighted two main refugia for C. vulgaris during the LGM, one around the Strait of Gibraltar and another in the north-western Iberian Peninsula. The Strait of Gibraltar area is a major centre of origin or glacial refugium for the heather C. vulgaris. However, the north-western Iberian Peninsula seems to have acted as a secondary glacial refugium for the more recent, Eurosiberian lineage of this species. This study stresses the importance of the Strait of Gibraltar area in the biogeographic history and conservation of the heather and, by extension, the European heathland habitat.

Understanding in what way cold-adapted mountain species have responded to historical climatic fluctuations in southern European refugia and investigating the genetic variation of endemic species is fundamental to predict their survival under contemporary global climate change. Veronica aragonensis (Plantaginaceae) is a tetraploid species endemic to the Iberian Peninsula that survived the climatic oscillations of the Quaternary. This species is adapted to cold environments and is included in several regional Red Lists due to its small and disjunct distribution range in high-mountain habitats (Pyrenees, Pre-Pyrenees and Baetic System). With the aim of deciphering the phylogeographical processes that underlie the presently disjunct distribution pattern of V. aragonensis and to evaluate its conservation status, we used 9 microsatellite loci to genotype 324 individuals from 12 populations representing the three disjunct areas where the species is distributed. Our results suggest that range fragmentation of an ancient continuous distribution of V. aragonensis—particularly in the low elevation mountain ranges located between the Pyrenees and the Baetic System—during the Last Glacial Maximum is the most plausible explanation for the disjunct distribution pattern of the species. Lastly, the intraspecific genetic patterns are discussed to predict how this endemic species can be affected by global climate warming. Based on the genetic data obtained here, an appropriate evaluation of the conservation status of V. aragonensis and some management strategies are provided.

a The Natural History Museum, Cromwell Road, London, SW7 5BD, UK; b Department of Plant and Animal Production, Munzur University, Vocational School of Tunceli, Tunceli, Turkey; c International School of Advanced Studies, University of Camerino, Camerino (MC), Italy; d Grupo de Investigación Biología para la Conservación, Universidad Pedagógica y Tecnológica de Colombia, Colombia; e Botanical Survey of India, Central National Herbarium, Howrah, India; f Botanical Garden-Institute, Russian Academy of Science, Vladivostok, Russia; g Viale Rovereto, Trento, Italy; h Department of Botany, Kherson State University, Kherson, Ukraine; i Kherson Branch of Admiral Makarov University, Kherson, Ukraine; j Institute of Industrial Ecology Problems of the North of the Kola Science Center of RAS, Apatity, Murmansk Province, Russia; k University of Insubria, Varese, Italy; l Departamento de Biología Vegetal (Botánica), Facultad de Biología, Universidad de Murcia, E-30100, Murcia, Spain; mTeam of Ecological Survey Research, National Institute of Ecology, Seocheon, Korea; n Departamento de Biología (Botánica), Facultad de Ciencias, Universidad Autónoma de Madrid, Madrid, Spain; o Centro de Investigación en Biodiversidad y Cambio Global, Madrid, Spain; p Institute of Biology Komi Science Centre UB RAS, Kommunisticheskaja St., 28, Syktyvkar, 167982, Komi Republic, Russia; q Botanical Museum, P.O. Box 7, FI-00014 University of Helsinki, Helsinki, Finland; r Department of Landscape Architecture, Faculty of Architecture, Niğde Ömer Halisdemir University, Niğde, Turkey; s Geobotany Department, Lomonosov Moscow State University, Moscow, Russia; t Botanical Garden-Institute, Far East Branch of Russian Academy of Sciences, Vladivostok, Russia; u Departamento de Biodiversidad, Ecología y Evolución, Facultad de Biología, Universidad Complutense de Madrid, 28040 Madrid, Spain; v Departamento de Botánica y Geología, Facultad de Farmacia, Universidad de Valencia, Avda, Vicente Andrés Estellés s/n, E-46100 Burjassot, Spain; wDepartment of Systematics, Biodiversity and Evolution of Plants, Albrecht von Haller Institute, University of Göttingen, 37073 Göttingen, Germany; x Donceel, Belgium; y Conservatoire Botanique National du Massif Central, Chavaniac, Laffayette, France; z Plant Resources Division, National Institute of Biological Resources, Incheon, Korea; aaAdnan Menderes Üniversitesi, Fen Edebiyat Fakültesi, Biyoloji Bölümü, TR-09010 Kepez-Aydin, Turkey; abPolar-Alpine Botanical Garden, Kola Science Centre, Russian Acadamy of Science, Kirovsk-6, Murmansk Province 184256, Russia; acInstitute of Botany, University of Natural Resources and Life Sciences Vienna, Vienna, Austria; adDepartment of Botany, University of South Bohemia, České Budějovice, Czech Republic; aeFreie Universität Berlin, Institut für Biologie, Systematische Botanik und Pflanzengeographie, Altensteinstr, 6, D-14195 Berlin, Germany; afKomarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia; agDepartment of Botany, University of Calcutta, Kolkata, India; ahTetbury, Gloucestershire, UK; aiDepartment of Botany, University of British Columbia, Vancouver, British Columbia, Canada; ajValkenswaard, The Netherlands; akLicenciatura en Biología, Universidad Distrital Francisco José de Caldas, Torre de Laboratorios, Bogotá, Colombia; alMTA Centre for Ecological Research, Institute of Ecology and Botany, Vácrátót, Hungary; am51 Wood Street, Wallingford, Oxfordshire, OX10 0AX, UK; anDepartamento de Botânica, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais, Brazil; aoDepartment of Environmental and Life Science, University of Cagliari, Sardinia, Italy; apCera dos Pomares, Aljezur, Portugal; aqKomarov Botanical Institute of the Russian Academy of Sciences, St. Petersburg, Russia; arDepartment of Biological, Geological and Environmental Biology, University of Catania, Catania, Italy; asCSIR-National Botanical Research Institute, Rana Pratap Marg, Lucknow, India; atThe Ministry of Forests, Lands, Natural Resource Operations and Rural Development, B. C. Government, Victoria, British Columbia, Canada; auDepartment of Botany, Bijoy Krishna Girls’ College, Howrah, West Bengal, India; avCenter for Biodiversity Research and Extension in Mindanao (CEBREM), Central Mindanao University, Musuan, Bukidnon Province, 8710 Mindanao Island, Philippines; awDepartamento de Botánica y Geología, Facultad de Ciencias Biológicas, Universitat de València, Avda. Dr. Moliner 50, E-46100, Burjassot, Valencia, Spain; axVia Selvotta, Muzzana del Turgnano, Udine, Italy; ayInstitute of Environmental Sciences, Kazan Federal University, Kazan, Tatarstan Republic, Russia; azCalifornia Academy of Sciences (CAS), 55 Music Concourse Drive, Golden Gate Park, San Francisco, CA 94118, USA; baInstitute of Biology Bucharest of Romanian Academy, 296 Splaiul Independentei, 060031 Bucharest, P.O. Box 56-53, Romania; bbDepartment of Biology, Faculty of Science and Letters, Aksaray University, Aksaray, Turkey; bcDepartment of Forest Engineering, Faculty of Forestry, Çankırı Karatekin University, Çankırı, Turkey; bdInstitute of Botany and Biodiversity Research, University of Vienna, Vienna, Austria; beMissouri Botanical Garden, 4344 Shaw Boulavard, St. Louis, Missouri, 63110, USA

The South African Cape fynbos heath Erica coccinea subsp. coccinea is polymorphic for post-fire regeneration mode (seeder/resprouter), flower colour (red/yellow), flowering time (summer-autumn/winter-spring), as well as anther size. These polymorphisms tend to occur between, rather than within, populations. Here, we aimed to understand flower colour divergence in this fynbos heath species and its association with post-fire regeneration. We tested for an association between these polymorphisms. We also examined whether yellow-flowered plants lacked anthocyanins and whether the lack of anthocyanins affected the concentration of other flavonoids, both in flowers and shoots. We found a strong association between flower colour, regeneration mode and flowering phenology. We also detected larger anthers and larger pollen in yellow- than in red-flowered plants, consistently for both resprouter and seeder regeneration strategies. Finally, we ascertained that being yellow-flowered is associated with loss in the capacity of synthesizing anthocyanins in both the flower and in the vegetative tissue. Considering that pollinator availability or preferences do probably not constitute a selection pressure, we propose that both enhanced reproductive success of anthocyaninless, yellow-flowered plants and the avoidance of likely negative pleiotropic effects (early senescence) in seeders would be key to understanding the flower colour divergence in E. coccinea subsp. coccinea.

International audience; Palaeoecology is increasingly contributing to conservation biology and restoration ecology by extending the recent annual to decennial records of ecological observations to centennial and millennial time scales. One of the almost neglected potential applications of palaeoecological investigations in conservation biology is the discovering of hidden species in subrecent fossil records. Such discoveries can be common in biodiversity hotspots in which the fauna and flora are not still fully known. In this study, we present the case of the liverwort Riella in continental southwest Asia (Iranian plateau), a very vulnerable taxon growing and reproducing under very special ecological circumstances. Our study adds two new species of Riella (R. echinata and R. helicophylla) in addition to the formerly reported R. cossoniana, to the subfossil bryophyte flora of Iran. We demonstrate the important contribution that the genus has played in the history of wetland flora of the semi-arid to arid areas of Iran during the late Quaternary. We further demonstrate the possibility that it is still thriving in many wetland systems and may need to be discovered and then protected in its potential habitats. We suggest that in the biodiversity hotspots of lower latitudes, more attention has to be paid to taxonomic identification of species to reveal the hidden part of the biodiversity for defining better conservation strategies for vulnerable aquatic ecosystems such as the seasonal wetland systems.

Background: Heathlands are relatively abundant in the landscape of the western Mediterranean region, especially in the Strait of Gibraltar region, where it is locally known as herriza. They are ass...

Erica australis and Erica arborea are morphologically and ecologically similar heather species. Erica australis is restricted to the western Mediterranean Basin where it overlaps with the westernmo ...

Andreaea rothii has been recorded for the first time in Croatia. It is a boreo-temperate suboceanic species (Hill et al., 2007) relatively rare in SE Europe, since it is known only from Romania (Ellis et al., 2014d), Slovenia and Serbia (Sabovljevic´ et al., 2008 ; Hodgetts, 2015). The species was found in the Papuk Mountains, situated in the mainly lowland area of NE Croatia. In this region Papuk is the largest and highest mountain range, with peaks between 800 and 900 m a.s.l. They are characterized by high geological diversity dominated by metamorphic rocks, such as different types of schists, as well as granites. The climate is temperate, moderately warm without an explicit dry period. About 60% of the almost totally forested area is covered by different communities of beech forests. The well-developed black patches of A. rothii cover an area ca 2 m× 0.5 m on a steep north-facing cliff on the edge of an acidothermophilic sessile oak (Quercus petraea agg.) forest. The specimens of A. rothii grew on the bare rock with the following bryophyte species: Cynodontium polycarpon (Hedw.) Schimp., Dicranella heteromalla (Hedw.) Schimp., Dicranum scoparium Hedw., Polytrichum piliferum Schreb. ex Hedw. and Rhabdoweisia (cf.) fugax (Hedw.) Bruch & Schimp. The population is very small with an extremely high risk of extinction, therefore we propose CR as Red List status for the taxon in Croatia. According to the last checklist of the moss flora of Croatia (Sabovljevic´, 2006), only Andreaea rupestris Hedw., collected from just one locality in 1927 (Horvat, 1932 and ZA), was reported for the genus. The locality is very interesting from the point of view of the vegetation of Croatia, because it is within 100 m of the second stand of Fagus sylvatica L.-Sphagnum quinquefarium (Braithw.) Warnst. forest (Alegro et al., 2015). The second occurrence of Dicranum spurium Hedw. (Ellis et al., 2014d) and Rhabdoweisia fugax (Papp et al., 2013) in Croatia are also found here. Another interesting moss is S. capillifolium (Ehrh.) Hedw., that forms small red patches within the thick carpets of S. quinquefarium under the open oulder scree forest in the neighbourhood.

A higher frequency of natural selection is expected in populations of organisms with shorter generation times. In fire-prone ecosystems, populations of seeder plants behave as functionally semelparous populations, with short generation times compared to populations of resprouter plants, which are truly iteroparous. Therefore, a stronger signature of natural selection should be detected in seeder populations, favoured by their shorter generation times and higher rates of population turnover. Here we test this idea in Erica coccinea from the Cape Floristic Region, which is dimorphic for post-fire regeneration mode. We measured three floral traits supposedly subject to natural selection in seeder and resprouter populations. We then compared phenotypic trait variation with neutral genetic variation in each group of populations using PST–FST comparisons to detect signatures of natural selection in seeders and resprouters. We found a strong signature of selection in seeder populations, but not in resprouters. Furthermore, anthers of seeders were more exserted (and larger) than those of resprouters. These differences were maintained at sites where seeders and resprouters co-occurred, suggesting that phenotypic plasticity or adaptation to different growth environments are unlikely explanations for trait variation. These results provide empirical support for the hypothesis that the genetic signature of natural selection is certainly more intense in seeder than in resprouter populations, favoured by their comparatively faster generation turnovers. Increased frequency of natural selection would increase differentiation among populations, thus promoting speciation in pyrophytic seeder lineages of the Cape flora., J.G.S.-M. was supported by a ‘Ramón y Cajal’ postdoctoral contract from MICINN

Population genetic studies of widespread Mediterranean shrubs are scarce compared with those of trees and narrow endemics or studies from phylogeographical perspectives, despite the key role these species may play in Mediterranean ecosystems. Knowledge on the effect of ecological factors in shaping their genetic patterns is also limited. In this study we investigate genetic diversity and population structure across 18 populations of Rosmarinus officinalis, a Mediterranean shrubland plant. Populations were sampled along two elevational gradients, one each on calcareous and siliceous soils in a mountain system in the eastern Iberian Peninsula, to decipher the effect of ecological factors on the genetic diversity and structure based on 11 microsatellite loci. We found overall high levels of genetic diversity and weak population structure. Genetic diversity increased with elevation, whereas population differentiation was stronger among populations growing on siliceous soils. The nested analysis of elevational gradients within soil types revealed that these general patterns were mostly driven by siliceous populations, whereas calcareous populations were more homogeneous along elevational belts. Bayesian analysis of population structure revealed genetic membership of lowland and high-elevation populations to different genetic clusters and a higher admixture of intermediate-elevation populations to both clusters. High-elevation populations were less differentiated from a hypothetical ancestral cluster, suggesting the persistence of their gene pool during the Pleistocene glaciations. In contrast, lowland populations resulted from more recent divergence. We propose that life-history and reproductive traits mostly contribute to explain the high levels of genetic diversity and weak population structure, whereas ecological and historical factors mostly contribute to the stronger differentiation of siliceous populations and a rapid expansion of R. officinalis on calcareous soils possibly mediated by human landscape transformations.

The effects of the universal signal molecule, abscisic acid (ABA), with limited knowledge functions in non-tracheophyte have been studied in three selected bryophyte species: two mosses, Physcomitrella patens and Atrichum undulatum, and one liverwort, Marchantia polymorpha. While vegetative development as well as total chlorophyll and carotenoid contents tended to decrease in the three bryophyte species with increasing exogenous ABA concentration, the effect on total biomass showed less clear patterns in the bryophytes tested. These differences in response to ABA likely reflect different adaptations of these three species to conditions in situ., JGS - M was supported by a Spanish Ministry of Science and Innovation “Ramón y Cajal” postdoctoral contract.

Self‐pollination by geitonogamy is likely in self‐compatible plants that simultaneously expose a large number of flowers to pollinators. However, progeny of these plants is often highly allogamous. Although mechanisms to increase cross‐pollination have been identified and studied, their relative importance has rarely been addressed simultaneously in plant populations. We used Rosmarinus officinalis to explore factors that influence the probability of self‐fertilisation due to geitonogamy or that purge its consequences, focusing on their effects on seed germination and allogamy rate. We experimentally tested the effect of geitonogamy on the proportion of filled seeds and how it influences germination rate. During two field seasons, we studied how life history and flowering traits of individuals influence seed germination and allogamy rates of their progeny in wild populations at the extremes of the altitudinal range. The traits considered were plant size, population density, duration of the flowering season, number of open flowers, flowering synchrony among individuals within populations and proportion of male‐sterile flowers. We found that most seeds obtained experimentally from self‐pollination were apparently healthy but empty, and that the proportion of filled seeds drove the differences in germination rate between self‐ and cross‐pollination experiments. Plants from wild populations consistently had low germination rate and high rate of allogamy, as determined with microsatellites. Germination rate related positively to the length of the flowering season, flowering synchrony and the ratio of male‐sterile flowers, whereas the rate of allogamous seedlings was positively related only to the ratio of male‐sterile flowers. Rosemary plants purge most of the inbreeding caused by its pollination system by aborting the seeds. This study showed that the rates of seed germination and allogamy of the seedlings depend on a complex combination of factors that vary in space and time. Male sterility of flowers, length of the flowering season and flowering synchrony of individuals within populations all favour high rates of cross‐pollination, therefore increasing germination and allogamy rates. Flowering traits appear to be highly plastic and respond to local and seasonal conditions., Financial support was provided by CGL2009‐07262/BOS project from the Spanish Ministry of Science and Innovation (MICINN) and PROMETEO/2016/021 project from the Valencia Autonomous Government. J.G.S.‐M. was supported by a ‘Ramón y Cajal’ post‐doctoral contract from MICINN. M.C.C. was supported by a JAE‐Doc CSIC post‐doctoral scholarship.

WOS: 000348594500007, …, Institute of Environmental Technologies - 'Research and Development for Innovations' Operational Programme [CZ.1.05/2.1.00/03.0100]; Structural Funds of the European Union; state budget of the Czech Republic; National Feasibility Programme I of the Czech Republic [LO1208]; SYNTHESYS project [FR-TAF-3686]; University of Ostrava [SGS27/PRF/2014]; Polish National Centre of Science [N N 303 469 338]; Institute of Botany of the Polish Academy of Sciences; Institute of Biology Bucharest of the Romanian Academy [RO1567-IBB03/2014]; Natural Environment Research Council; Scientific and Technological Research Council of Turkey (TUBITAK) [111T359]; Spanish Ministerio de Economia y Competitividad (MINECO); Flora Briofitica Iberica from MINECO [CGL2009-09530]; MINECO; Spanish 'Ministerio de Ciencia e Innovacion' [CGL2010-15959, CGL2009-07323]; FEDER; 'Ramon y Cajal' Subprogram (MICINN-European Social Fund); 'Ministerio de Economia y Competitividad' [CGL2012-30721]; MOVECLIM project (ANR under the Net-Biome call); RSF [14-14-00903, 14-24-00037]; RFBR [13-04-01427, 12-04-01476]; 'The Survey of new and unrecorded taxa in vascular plants (NIBR)' - Ministry of Environment of the Korean Government [2013-02-001]; Natural Environment Research Council [bas0100025], The contribution by V. Plasek is part of a research project of the Institute of Environmental Technologies, reg. no. CZ.1.05/2.1.00/03.0100, supported by the 'Research and Development for Innovations' Operational Programme, and financed by the Structural Funds of the European Union and by the state budget of the Czech Republic, Project LO1208 of the National Feasibility Programme I of the Czech Republic, and SYNTHESYS project FR-TAF-3686. The contribution by L. Cihal is a part of grant project SGS27/PRF/2014 financed by University of Ostrava. The contributions by R. Ochyra have been financially supported by the Polish National Centre of Science through grant no. N N 303 469 338 and, in part, by the statutory fund of the Institute of Botany of the Polish Academy of Sciences. He also thanks Helen J. Peat, Cambridge, for kindly allowing him to study the herbarium collections from AAS. S. Stefanut, acknowledges the support by project no. RO1567-IBB03/2014 through the Institute of Biology Bucharest of the Romanian Academy. J. Vana and K. K. Newsham acknowledge the logistics support that was provided by the British Antarctic Survey's Operations Group, officers and crew of the RRS Ernest Shackleton and staff at King Edward Point research station, and thank Helen J. Peat for the loan of AAS herbarium specimens. Funding was provided by the Natural Environment Research Council. All are gratefully acknowledged.; Peter Szucs and Andras Bidlo would like to thank Peter Erzberger for examining the specimen of Campylopus introflexus, and Gyorgy Pulger for his assistance in fieldwork. Tulay Ezer and Recep Kara are indebted to the Scientific and Technological Research Council of Turkey (TUBITAK) for financial support (Project Code: 111T359).; M. J. Gil-Lopez, F. Puche, and J. G. Segarra-Moragues thank H. Kharbouch (SahaTours), F. Pacheco, J. P. Peralta, and F. Rivera for their help in field work. M. J. Gil-Lopez benefited from a FPU PhD grant from the Spanish Ministerio de Economia y Competitividad (MINECO). F. Puche acknowledges funds from project CGL2009-09530, Flora Briofitica Iberica from MINECO. J. G. Segarra-Moragues benefited from a 'Ramon y Cajal' post-doctoral contract from MINECO. Juan Larrain and Amalia B. Biasuso thank Reinaldo Vargas and Gotz Palfner for their company in the field. Marta Alonso and David Long acknowledge the financial support of the Spanish 'Ministerio de Ciencia e Innovacion' (projects CGL2010-15959 co-financed by FEDER) and 'Ramon y Cajal' Subprogram (MICINN-European Social Fund). We also thank Maria J. Cano and Juan A. Jimenez for their contribution to this work.; D. Rios, M. T. Gallego, and J. Guerra are grateful to Len Ellis for organising the loan of specimens and assistance during the visit of the first author to the BM herbarium. This research was carried out with financial support from the Spanish 'Ministerio de Ciencia e Innovacion' and 'Ministerio de Economia y Competitividad' [Projects CGL2009-07323 and CGL2012-30721, respectively].; Terry Hedderson is grateful to Jean-Yves Meyer, Ravahere Tuputuari, Claudine Ah-Peng, and Oliver Flores for field assistance and companionship and to the MOVECLIM project (ANR under the Net-Biome call) for funding the fieldwork on Tahiti. The study by E.V. Sofronova was supported by RSF (project no. 14-14-00903), that by Y.S. Mamontov was partly supported by RFBR (project no. 13-04-01427 and 12-04-01476), and that by A. D. Potemkin was supported by RSF (project no. 14-24-00037). The research by Yoon Young-Jun, Benito C. Tan and Byung-Yun Sun was supported by grants from 'The Survey of new and unrecorded taxa in vascular plants (NIBR No. 2013-02-001)' funded by the Ministry of Environment of the Korean Government.

Cultures of soil sediments collected in June 2012 from saltpans and vleis from five South African localities have revealed two new, geographically distant, populations of the endemic Riella alatospora, confirmed the presence of Riella purpureospora from Blouvlei in the Cape Town area and uncovered a new endemic species, Riella trigonospora. The new species obtained from cultures from five sampled localities is described and illustrated. It is dioicous and characterised by acuminate to rostrate, papillose involucres and by a unique spore morphology. The spores are dark purple, as in R. purpureospora, and have an irregular discontinuous wing at the equatorial plane, which suggests a morphological affinity to R. alatospora. The spores possess a combination of characters that are unique among extant species of Riella, including the triangular spore body with concave sides, a discontinuous equatorial wing, and short blunt spines on the distal face and the expanded parts of the wing at the spore angles. Such peculiar spore morphology is to some extent similar to Heerisporites variabilis, a sporomorph from Upper Miocene deposits of Switzerland and suggests links to Riellaceae from the Mediterranean basin. A lectotype is designated for R. purpureospora and a new description and illustration, incorporating additional characters derived from the cultured plants, are also presented, together with an identification key to the South African species of Riella. © 2014 South African Association of Botanists., This study was partially funded by the Spanish Ministry of Science and Innovation (project CGL2009-09530, Flora Briofítica Ibérica). JGS-M was supported by a Spanish Ministry of Science and Innovation “Ramón y Cajal” (RYC-2009-05164) postdoctoral contract.

Riella affinis Howe & Underwood is a rare species that is thus far known from eight populations worldwide. Only half of these were recorded in the last 50 years and none were recollected since first reported. Study of a Spanish population of R. cossoniana Trab., recorded in 2002 from Las Lomas, Malaga province, revealed that it belongs to R. affinis. The correct identity of the populations, which had not been available for study, was already suggested by reproductive and ecological characteristics of the plants mentioned in the original publication. The three main lagoons from the Las Lomas endorheic system, where the plants had been recorded, were sampled and living plants of R. affinis were found in Viso lagoon whereas culture of sediments yielded plants of this species from Viso and Caja lagoons. Riella affinis is new to continental Europe and the new Spanish populations are described and illustrated in detail. Riella cossoniana is excluded from Malaga province. Riella affinis is classified as critically endangered (CR) globally and for Spain; the new Iberian populations should be targeted for conservation. An identification key to the currently recognised European species of Riella Mont. is included.

We thank Victor Sorribas, Shi Ying and Clive Stace for their advice and help with the RFLP and CAPS analyses, the authorities of the Spanish National Parks of Ordesa-Monte Perdido, Aiguestortes i Estany de Sant Maurici and Picos de Europa for the permits to sample the Festuca materials in these protected areas, the curators of the G, FCO, JACA, LEB, MA, W, Z and Jochen Muller and Oscar Sanchez-Pedraja (private) herbaria for facilitating the loan of voucher specimens, Jochen Muller for fruitful nomenclatural discussion and Emily Lemonds for linguistic assistance. This work was supported by a Spanish Ministry of the Environment – National Parks Organism research project (059/2009) and by an Instituto de Estudios Altoaragoneses and Aragon Government-European FEDER Bioflora grant. P.T. and A.J.D.-P. were supported by Venezuelan CDCH postdoctoral and PhD grants, respectively. J.G.S.-M. was supported by two consecutive Aragon Government ‘Araid’ and Spanish Ministry of Science and Innovation ‘Ramon y Cajal’ postdoctoral contracts. P.C. was supported by an Aragon CAI-Programa Europa XXI fellowship.

Apart from the overwhelming cases of allopolyploidization, the impact of speciation through homoploid hybridization is becoming more relevant than previously thought. Much less is known, however, about the impact of climate changes as a driven factor of speciation. To investigate these issues, we selected Festuca picoeuropeana, an hypothetical natural hybrid between the diploid species F. eskia and F. gautieri that occurs in two different mountain ranges (Cantabrian Mountains and Pyrenees) separated by more than 400 km. To unravel the outcomes of this mode of speciation and the impact of climate during speciation we used a multidisciplinary approach combining genome size and chromosome counts, data from an extensive nuclear genotypic analysis, plastid sequences and ecological niche models (ENM). Our results show that the same homoploid hybrid was originated independently in the two mountain ranges, being currently isolated from both parents and producing viable seeds. Parental species had the opportunity to contact as early as 21000 years ago although niche divergence occurs nowadays as result of a climate-driven shift. A high degree of niche divergence was observed between the hybrid and its parents and no recent introgression or backcrossed hybrids were detected, supporting the current presence of reproductive isolation barriers between these species., Financial support was provided by the Spanish projects PI097/08 from the Aragón Government, GA-LC-012/2008 from the Aragón Government-La Caixa, 059/2009 from the Ministry of Environment (National Parks division) and a European Union ESF exchange grant (FroSpects 3142). IM received funding from the People Programme (Marie Curie Actions) of the European Union's Seventh Framework Programme (FP7/2007-2013) under REA grant agreement PIOF-GA-2011-301257. DD was supported by the Portuguese Foundation for Science and Technology (SFRH/BPD/100384/2014). IM, MLLH and PC benefitted from the Bioflora grant project cofunded by the Aragón government and the European Social Fund. JGS-M was supported by two consecutive Aragón Government "Araid" and the Spanish Ministry of Science and Innovation “Ramón y Cajal” postdoctoral contracts.

The genus Erica represents the epitome of plant biodiversity in the South African Cape fynbos with over 700 species. This genus is composed of seeder and resprouter species, but both species diversity and endemism are strongly linked to the seeder habit and concentrated in the southwestern Cape Floristic Region (CFR). Erica coccinea is a relatively abundant and widespread fynbos species whose most remarkable morphological feature is the existence of distinct seeder and resprouter forms, frequently—but not always—in disjunct populations. Both higher within-population genetic diversity and among-population differentiation have been found in seeders, most likely as a consequence of the shorter generation times and faster population turnovers. Resprouters, despite being less diverse, are suspected to be ancestral. However, no solid evidence has yet been provided for the ancestrality of the resprouter form, or for the demographic processes that have determined the current distribution of genetic diversity in both regeneration forms. Here, we used microsatellites and sequences of the nuclear ribosomal internal transcribed spacers to describe the phylogeographic structure of seeder and resprouter E. coccinea populations and provide good evidence for the ancestral status of the resprouter form and the comparatively high rates of molecular evolution in derived seeder populations. We also reveal that mixed populations, where both seeder and resprouter individuals co-occur, were originated by secondary contacts. This study highlights the role of fire in driving accelerated diversification in seeder lineages of highly speciose CFR fynbos taxa., FO strongly thanks the Spanish Ministerio de Educación (travel Grant PR2010-0365) and the UCA Plan Propio de Investigación for providing travelling bursaries to do fieldwork. KBB acknowledges a PhD scholarship (FPI, BES-2009-015443) of the Spanish Ministry of Science and Innovation (MICINN). MH acknowledges a Ramón y Cajal fellowship (RYC2009-04537) from the Spanish Ministry of Science and Education (MICINN), and funding from the Research Council of Norway (203822/E40) and the Spanish Ministry of Economy and Competitiveness (CGL2012-40129-C02-02). SCGM and MH acknowledge the support of Marie Curie Intra European Fellowships within the 7th European Community Framework Programme (FP7-PEOPLE-2012-IEF, project nos. 328146 and 329088, respectively). JGS-M acknowledges a Ramón y Cajal fellowship (RYC2009-05164) from MICINN.

Dispersal can be an essential factor affecting the biological control of pests. Tetranychus urticae Koch (Acari: Tetranychidae) is a cosmopolitan and polyphagous species that may reach the pest status in many cropping systems including clementine orchards, where it may be found both in the trees and the associated flora. In a previous study, we demonstrated that the use of a ground cover of Festuca arundinacea Schreber (Poaceae) offered a better regulation of T. urticae populations than traditional alternatives (bare soil, multifloral wild cover). Therefore, we decided to study the ambulatory dispersal of mites crawling up and down tree trunks in a clementine mandarin orchard grown in association with a F. arundinacea cover for one season. The highest ambulatory migration rate was upward from the cover to the canopy. Multivariate regressions showed that the dynamics of T. urticae populations in the trees was strongly related to that of Phytoseiidae mites, their main natural predators. Surprisingly, canopy populations were not related to those on the ground cover or to those dispersing from it. When T. urticae individuals collected from the ground cover, the tree trunk, and the canopy were subjected to molecular analyses, the optimal number of genetic clusters (demes) was two. One cluster grouped individuals dispersed from the ground cover (e.g. collected on tree trunks) and 27.5 % of individuals collected in the ground cover. The second cluster grouped all the individuals collected from trees and 72.5 % of those collected in the cover. Interestingly, none of the individuals collected from the tree canopies was grouped with the first deme. This result may be taken as indicative that grass-adapted T. urticae individuals are unable to satisfactorily colonize and establish on the trees and provides evidence that host adaptation can hamper dispersal and establishment of the ground cover deme on trees, contributing to a better natural regulation of this pest species in citrus.

The rare aquatic liverwortRiella helicophylla(Bory et Mont.) Mont., inhabitant of temporary shallow ponds around the Mediterranean basin, is considered threatened throughout its distribution range. In addition, little is known of its biology and ecology or of its role in such an important ecosystem where environmental conditions vary yearly in unpredictable ways. In these variable habitats, due to the seasonal fluctuation of water levels, there is no guarantee of yearly spore input into the spore bank. Spore germination rate and the effects of different culture media in an axenic culture establishment, as well as propagation procedures ofR. helicophylla,were tested. New insights into the ecology and biology ofR. helicophyllaare given. Spore dormancy is documented, and the protocols for the in vitro culture establishment, propagation and acclimatization of this liverwort are developed. Dry storage at 20 ± 2 °C for about three months broke the dormancy of spores, which subsequently germinated in a high percentage (over 90%). A two phase (solid and liquid) culture media system was developed for the purpose of achieving fully developed gametophytes. The liquid phase contained electrolytes simulating brackish water.

Aim: Dioscorea (Dioscoreaceae) is a predominantly pantropical genus (, This study was supported by Fundación BBVA (FBBVA) BIOCON 05-093/ 06 project (PC, JGSM) and by Instituto de Estudios Altoaragoneses project (J.V.). J.V. benefited from a PhD fellowship funded by FBBVA and two research stays at Kew Gardens funded by European SYNTHESYS GB-TAF and DGA-CAI grants. J.G.S-M. was supported by Araid and Ramón y Cajal postdoctoral contracts. P.C. was partially funded by Bioflora Research Team grant cofunded by the Aragón Government and the European Social Fund. I.S. was supported by grant CGL2012-40129-C02-01 from the Spanish Ministry of Science.

Polymorphic microsatellite markers were developed for the Ibero-North African, strict gypsophyte Lepidium subulatum to unravel the effects of habitat fragmentation in levels of genetic diversity, genetic structure and gene flow among its populations. Using 454 pyrosequencing 12 microsatellite loci including di- and tri-nucleotide repeats were characterized in L. subulatum. They amplified a total of 80 alleles (2–12 alleles per locus) in a sample of 35 individuals of L. subulatum, showing relatively high levels of genetic diversity, HO = 0.645, HE = 0.627. Cross-species transferability of all 12 loci was successful for the Iberian endemics Lepidium cardamines, Lepidium stylatum, and the widespread, Lepidium graminifolium and one species each of two related genera, Cardaria draba and Coronopus didymus. These microsatellite primers will be useful to investigate genetic diversity, population structure and to address conservation genetics in species of Lepidium.

12 páginas, 1 figura, 3 tablas., Ecological network theory predicts that in mutualistic systems specialists tend to interact with a subset of species with which generalists interact (i.e. nestedness). Approaching plant-arbuscular mycorrhizal fungi (AMF) association using network analyses will allow the generality of this pattern to be expanded to the ubiquitous plant-AMF mutualism. • Based on certain plant-AMF specificity recently suggested, networks are expected to be nested as a result of their mutualistic nature, and modular, with certain species interacting more tightly than others. Network analyses were used to test for nestedness and modularity and to compare the different contribution of plant and AMF to the overall nestedness. • Plant-AMF networks share general network properties with other mutualisms. Plant species with few AMFs in their roots tend to associate with those AMFs recorded in most plant species. AMFs present in a few plant species occur in plant species sheltering most AMF (i.e. nestedness). This plant-AMF network presents weakly interlinked subsets of species, strongly connected internally (i.e. modularity). Both plants and AMF show a nested structure, although AMFs have lower nestedness than plants. • The plant-AMF interaction pattern is interpreted in the context of how plant-AMF associations can be underlying mechanisms shaping plant community assemblages., We thank J. P. Castillo, M. Morales, C. Silva and L. Sortibrán for help with field samplings and S. Donat and M. Morales for laboratory assistance. This work was funded by AECID (projects A017475⁄08, A023461⁄09), DGAPA-UNAM (project IN-224808-3; IN-202811-3), and CYTED (acción 409AC0369). J.G.S-M. was supported by a ‘Ramón y Cajal’ postdoctoral contract from MICINN.

13 páginas, 4 figuras, 5 tablas., Aim The Chilean endemic Dioscorea biloba (Dioscoreaceae) is a dioecious geophyte that shows a remarkable 600 km northsouth disjunction in the peripheral arid area of the Atacama Desert. Its restricted present-day distribution and probable Neogene origin indicate that its populations have a history linked to that of the Atacama Desert, making this an ideal model species with which to investigate the biogeography of the region. Location Chile, Atacama Desert and peripheral arid area. Methods Two hundred and seventy-five individuals from nine populations were genotyped for seven nuclear microsatellite loci, and plastid trnLF and trnTL sequences were obtained for a representative subset of these. Analyses included the estimation of genetic diversity and population structure through clustering, Bayesian and analysis of molecular variance analyses, and statistical parsimony networks of chloroplast haplotypes. Isolation by distance was tested against alternative dispersal hypotheses. Results Microsatellite markers revealed moderate to high levels of genetic diversity within populations, with those from the southern Limari Valley showing the highest values and northern populations showing less exclusive alleles. Bayesian analysis of microsatellite data identified three genetic groups that corresponded to geographical ranges. Chloroplast phylogeography revealed no haplotypes shared between northern and southern ranges, and little haplotype sharing between the two neighbouring southern valleys. Dispersal models suggested the presence of extinct hypothetical populations between the southern and northern ranges. Main conclusions Our results are consistent with prolonged isolation of the northern and southern groups, mediated by the life-history traits of the species. Significant isolation was revealed at both large and moderate distances as gene flow was not evident even between neighbouring valleys. Bayesian analyses of microsatellite and chloroplast haplotype diversity identified the southern area of Limari as the probable area of origin of the species. Our data do not support recent dispersal of D. biloba from the southern range into Antofagasta, but indicate the fragmentation of an earlier wider range, concomitant with the Pliocene-Pleistocene climatic oscillations, with subsequent extinctions of the Atacama Desert populations and the divergence of the peripheral ones as a consequence of genetic drift., We thank E. Pérez-Collazos and L. Villar for their help during fieldwork. A. Díaz-Pérez for his valuable comments and discussion on dispersal model analysis, P. Gibbs and T. Langdon for their linguistic assistance, and the editor P. Ladiges and two anonymous referees for their valuable comments that have improved the manuscript. This study has been supported by a Fundación BBVA BIOCON 05-093/06 project grant. J.V. was supported by a Fundación BBVA PhD grant, an Instituto de Estudios Altoaragoneses project, and two research stays at Royal Botanic Gardens, Kew, funded by SYNTHESYS GB-TAF and the Diputación General de Aragón-Caja de Ahorros de la Inmaculada (DGA-CAI). J.G.S.-M. was supported by two consecutive Spanish Aragón Government ‘Araid’ and Ministry of Science and Innovation ‘Ramón y Cajal’ post-doctoral contracts.

14 páginas, 4 figuras, 3 tablas., Limonium narbonense Miller is a fertile tetraploid species with a sporophytic self-incompatibility system. This sea lavender is found in coastal salt marshes which have been under intense human pressure during the past decades resulting in significant habitat fragmentation. Eleven microsatellite loci specifically designed for this species were amplified in 135 individuals from five populations. These markers were used to investigate the polyploid nature, the levels of genetic diversity and population structure in this species. L. narbonense showed high levels of genetic diversity (A = 7.82, P = 100% H (T) = 0.446), consistent with its likely autotetraploid origin revealed in this study and obligate outcrossing breeding system. Inbreeding (F (IS)) values were low in the three southern populations (mean F (IS) = 0.062), and higher in the northern populations (mean F (IS) = 0.184). Bayesian analysis of population structure revealed that populations could be grouped into two genetic clusters, one including three southern populations and the other the two northernmost ones. Individuals from the two northernmost populations showed higher admixture of the two genetic clusters than individuals from the three southern ones. A thorough analysis of microsatellite electrophoretic patterns suggests an autotetraploid origin for L. narbonense. The genetic structure revealed in this study is attributed to a recent migration from the southern area. This result suggests a net gene flow from the south to the north, likely facilitated by migratory movements of birds visiting the temporary flooded ponds occupied by L. narbonense., We are indebted to C. Palacios, S. Rodríguez, J.A. Rosselló and the staff of the Consellería de Medio Ambiente for field assistance, to S.E. Mitchell for laboratory assistance, to C. Palacios for fruitful comments, S. Kresovich for his hospitality and support during the development of this study and F. Palero for statistical advice. This study has been supported by project PB98-1436 from DGES-MEC. MPE was supported by a PhD grant from the Generalitat Valenciana (GV). JGSM was supported by a Ramón y Cajal Postdoctoral contract from the Spanish Ministerio de Ciencia e Innovación.

A new population of the Argentinian endemic Riella choconensis Hässel (Riellaceae, Marchantiophyta) was discovered from a culture of sediments from the Laguna de los Juncos (Río Negro province, Argentina). This species was known previously from two nearby collections, made in 1984, in the type locality at the Ramos Mexía reservoir (Neuquén province) and was not recorded since. Fresh material from the new population enabled detailed morphological analyses, which are extensively described here, illustrated and compared to type material of the species. Based on the smooth female involucre, the species belongs to subgenus Riella Mont. However, cross-sections of this structure showed that R. choconensis has bistratose involucre walls, contrasting with the unistratose female involucres of all other species in the genus. The updated morphological data are integrated into a new key to the identification of the American species of Riella. Fil: Segarra Moragues, José Gabriel. Universidad de Valencia; España Fil: Puche, Felisa. Universidad de Valencia; España Fil: Gleiser, Gabriela Laura. Consejo Nacional de Investigaciones Científicas y Técnicas. Centro Científico Tecnológico Conicet - Patagonia Norte. Instituto de Investigaciones en Biodiversidad y Medioambiente. Universidad Nacional del Comahue. Centro Regional Universidad Bariloche. Instituto de Investigaciones en Biodiversidad y Medioambiente; Argentina. Universidad Nacional del Comahue. Centro Regional Universitario Bariloche. Laboratorio de Ecotono; Argentina

Understanding the processes of biological diversification is a central topic in evolutionary biology. The South African Cape fynbos, one of the major plant biodiversity hotspots out of the tropics, has prompted several hypotheses about the causes of generation and maintenance of biodiversity. Fire has been traditionally invoked as a key element to explain high levels of biodiversity in highly speciose fynbos taxa, such as the genus Erica. In this study, we have implemented a microevolutionary approach to elucidate how plant-response to fire may contribute to explain high levels of diversification in Erica. By using microsatellite markers, we investigated the genetic background of seeder (fire-sensitive) and resprouter (fire-resistant) populations of the fynbos species Erica coccinea. We found higher within-population genetic diversity and higher among-population differentiation in seeder populations and interpreted these higher levels of genetic diversification as a consequence of the comparatively shorter generation times and faster population turnover in the seeder form of this species. Considering that genetic divergence among populations may be seen as the initial step to speciation, the parallelism between these results and the pattern of biodiversity at the genus level offers stimulating insights into understanding causes of speciation of the genus Erica in the Cape fynbos.

12 páginas, 6 figuras, 1 tabla., Pollen analyses of 12 Iranian species of Typha L. were conducted through light microscopy and scanning electron microscopy. Among the twelve studied species, we found that five species, T. caspica, T. latifolia, T. lugdunensis, T. minima and T. shuttleworthii, present pollen in tetrads, whereas the remaining seven species, T. angustifolia, T. azerbaijanensis, T. domingensis, T. grossheimii, T. kalatensis, T. laxmanii and T. tourcomanica, have pollen in monads. The exine of Typha species is mostly reticulate although a few species present perforate-microreticulate and rugulate exine. Our analyses revealed that using a combination of light microscopy and scanning eletron microscopy characters the identification of species is feasible. A diagnostic key based on palynological traits is presented for all studied taxa., We thank Dr. A. A. Maas soumi for providing herbarium vouchers from TARI and for useful comments in an earlier version of this manuscript. M. Moussavi and F. Aghbeigi for providing facilities to study IRAN specimens. M. Rahmani for assistance with SEM analysis. The study on the family Typhaceae in IRAN was partially supported by Project 105 of the Islamic Azad University, Garmsar branch, Research Council to SMM Hamdi. J. G. Segarra-Moragues was supported by an ARAID (Agencia Aragonesa para la Investigación y el Desarrollo) postdoctoral contract.

13 páginas., Chromosomal analyses were conducted for the first time in all three species of the Chilean endemic Epipetrum Phil. (Dioscoreaceae; the yam family). Mitotic chromosome counts showed that all studied individuals of the rare Epipetrum bilobum and E. polyanthes, and most studied individuals of the more widespread E. humile, had 2n = 14 chromosomes. However, two populations of E. humile had 2n = 28 chromosomes. The ideograms shared a similar karyotype constitution for the three species but with a duplicated chromosome number in some populations of E. humile. Our results suggest that the Epipetrum taxa are mainly diploids, with a base number of x = 7, and the tetraploid cytotypes have arisen only in few cases. After that of Borderea (x = 6), this chromosome base number is the second lowest one found in the Dioscoreaceae. Epipetrum and Borderea are two small, relict, satellite genera of the Dioscoreaceae, endemic to Chile (South America) and the Pyrenees (Southwestern Europe), respectively. The two taxa share several morphological attributes and are adapted to rocky and mountain habitats. However, whereas Borderea is an allotetra-ploid genus of relatively recent hybrid origin, Epipetrum is diploid in common with the species found in the early branching clades of Dioscorea.

14 paginas, 3 figuras, 5 tablas., In sexually polymorphic species, the morphs are maintained by frequency-dependent selection through disassortative mating. In heterodichogamous populations in which disassortative mating occurs between the protandrous and protogynous morphs, a decrease in female fitness in one morph is hypothesized to drive sexual specialization in the other morph, resulting in dimorphic populations. We test these ideas in a population of the heterodichogamous species, Acer opalus. We assessed both prospective gender of individuals in terms of their allocations and actual parentage using microsatellites; we found that most matings in A. opalus occur disassortatively. We demonstrate that the protogynous morph is maintained by frequency-dependent selection, but that maintenance of males versus protandrous individuals depends on their relative siring success, which changes yearly. Seeds produced later in the reproductive season were smaller than those produced earlier; this should compromise reproduction through ovules in protandrous individuals, rendering them male biased in gender. Time-dependent gender and paternity analyses indicate that the sexual morphs are specialized in their earlier sexual functions, mediated by the seasonal decrease in seed size. Our results confirm that mating patterns are context-dependent and change seasonally, suggesting that sexual specialization can be driven by seasonal effects on fitness gained through one of the two sexual functions., We thank the staff of the Parque Natural Carrascal de la Font Roja for giving us permission to perform this study in the natural park. This study was funded by aMEC/CICYT (REPROFOR AGL2005-07440-C02-02/FOR) project grant to MV, a I3P/CSIC postdoctoral grant to JGS-M and a FPU/MEC Ph.D. grant to GG.

Several methods based on population biology, biogeography, ecology, and genetics have been traditionally used for the identification of units for conservation below the species level. We use a combination of two methods based on population genetic structure estimators and on probabilities of loss of rare alleles to identify the Relevant Genetic Units for Conservation (RGUCs). The aims were to assess the genetic diversity and population structure of the endemic steppe plant Boleum asperum (Brassicaceae), and to determine how many and which populations significantly represent the total genetic diversity and the rarest allelic variation. Despite the high amplified fragment length polymorphism genetic diversity values detected in B. asperum (hT = 0.744), caused probably by its hexaploidy and allogamy, moderate spatial genetic differentiation was detected among populations ( 13%), suggesting the existence of an ancestral continuous distribution range that was fragmented into separate ‘islands’ in more recent historical times. Five RGUCs, accounting for the 85.10% of the total genetic variation and representative of the entire geographical distribution of the species, were selected for in situ conservation. Ex situ conservation is proposed to complement the preservation of B. asperum. This method of objective selection of populations may be applied to other candidate taxa for conservation with prior adjustment of the threshold values of diversity required for effective protection of each particular species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 94, 341–354.

10 paginas, 3 figuras., Background and Aims Heterodichogamy (a dimorphic breeding system comprising protandrous and protogynous individuals) is a potential starting point in the evolution of dioecy from hermaphroditism. In the genus Acer, previous work suggests that dioecy evolved from heterodichogamy through an initial spread of unisexual males. Here, the question is asked as to whether the different morphs in Acer opalus, a species in which males co-exist with heterodichogamous hermaphrodites, differ in various components of male in fitness. Methods Several components of male fertility were analysed. Pollination rates in the male phase were recorded across one flowering period. Pollen viability was compared among morphs through hand pollinations both with pollen from a single sexual morph and also simulating a situation of pollen competition; in the latter experiment, paternity was assessed with microsatellite markers. It was also determined whether effects of genetic relatedness between pollen donors and recipients could influence the siring success. Finally, paternal effects occurring beyond the fertilization process were tested for by measuring the height reached by seedlings with different sires over three consecutive growing seasons. Key Results The males and protandrous morphs had higher pollination rates than the protogynous morph, and the seedlings they sired grew taller. No differences in male fertility were found between males and protandrous individuals. Departures from random mating due to effects of genetic relatedness among sires and pollen recipients were also ruled out. Conclusions Males and protandrous individuals are probably better sires than protogynous individuals, as shown by the higher pollination rates and the differential growth of the seedlings sired by these morphs. In contrast, the fertility of males was not higher than the male fertility of the protandrous morph. While the appearance of males in sexually specialized heterodichogamous populations is possible, even in the absence of a fitness advantage, it is not clear that males can be maintained at an evolutionary equilibrium with two classes of heterodichogamous hermaphrodites., We thank Jaume Tormo, Yolanda Orduna and Tono Bellido for field assistance, and the staff of the Parque Natural Carrascal de la Font Roja for giving us permission to conduct this study in the natural park. This study was funded by a MEC/CICYT (REPROFOR AGL2005–07440-C02-02/FOR) project grant to M. Verdú, a I3P/ CSIC postdoctoral grant to J.G. Segarra-Moragues and a FPU/MEC PhD grant to G. Gleiser.

14 páginas, 4 figuras, 2 tablas., Aim. Borderea pyrenaica (Dioscoreaceae) is a Tertiary relict plant endemic to the Central Pyrenees. Because of its narrow distribution in a small geographical area and the fact that it is restricted to high alpine habitats, it constitutes an ideal model species for inferring the historical dynamics of population survival and migration during and after Quaternary glaciations in the Pyrenees., Location. Central Pyrenees and pre-Pyrenees, Spain–France., Methods. Eleven primer pairs were used to amplify 18 microsatellite loci in this allotetraploid species in a sample of 804 individuals from 15 populations, revealing a total of 77 alleles. Genotypic data of individuals and populations were analysed using clustering and Bayesian methods of analysis of population structure., Results. A higher number of private alleles and a significantly higher allelic richness (A*) were found in the southern area (21, A* ¼ 2.295) than in the northern area (5, A* ¼ 1.791). Furthermore, the allelic composition of the northern area represented a subset of that from the southern area., Main conclusions. The hypothesis of in situ survival in northern Pyrenean nunataks was rejected, while peripheral refugia were considered to be restricted to the southern Pyrenees and pre-Pyrenees, where historical geographical fragmentation probably caused the divergence among southern Pyrenean populations. Molecular evidence indicates that these refugial populations probably colonized the northern area after sheet-ice retreat. Borderea pyrenaica lineages followed two migratory pathways in their northward colonization, suggesting several founder events for the populations that eventually reached the territory of the Gavarnie cirque., We thank Ivan Scotti and Joachim Kadereit for their critical review of the manuscript; Clive Stace and Samuel Pyke for corrections to the English; Alain Valadon, Delphine Fallour-Rubio, José Vicente Andrés, Raúl Andrés, Gustavo Gutiérrez, Pedro Torrecilla and the forest guards at the Parc National des Pyrénées (PNP) for sampling facilities; Ma Victoria Flores-Stols for her help in constructing Figs 1 and 4; and two anonymous referees for their thorough revision of an earlier version of the manuscript. This work was supported by a Spanish Aragón Government (DGA) project grant (P105/99-AV) to P.C., a PNP project grant (2001–64S) to P.C. and J.G.S.M., and by a DGA PhD fellowship to J.G.S.M. M.P. and F.G.C. were supported by a MEC/CICYT grant (P98-1436).

10 páginas, 1 figura, 3 tablas., Mediterranean-type ecosystems are among the most remarkable plant biodiversity "hot spots" on the earth, and fire has traditionally been invoked as one of the evolutionary forces explaining this exceptional diversity. In these ecosystems, adult plants of some species are able to survive after fire (resprouters), whereas in other species fire kills the adults and populations are only maintained by an effective post-fire recruitment (seeders). Seeders tend to have shorter generation times than resprouters, particularly under short fire return intervals, thus potentially increasing their molecular evolutionary rates and, ultimately, their diversification. We explored whether seeder lineages actually have higher rates of molecular evolution and diversification than resprouters. Molecular evolutionary rates in different DNA regions were compared in 45 phylogenetically paired congeneric taxa from fire-prone Mediterranean-type ecosystems with contrasting seeder and resprouter life histories. Differential diversification was analyzed with both topological and chronological approaches in five genera (Banksia, Daviesia, Lachnaea, Leucadendron, and Thamnochortus) from two fire-prone regions (Australia and South Africa). We found that seeders had neither higher molecular rates nor higher diversification than resprouters. Such lack of differences in molecular rates between seeders and resprouters-which did not agree with theoretical predictions-may occur if (1) the timing of the switch from seeding to resprouting (or vice versa) occurs near the branch tip, so that most of the branch length evolves under the opposite life-history form; (2) resprouters suffer more somatic mutations and therefore counterbalancing the replication-induced mutations of seeders; and (3) the rate of mutations is not related to shorter generation times because plants do not undergo determinate germ-line replication. The absence of differential diversification is to be expected if seeders and resprouters do not differ from each other in their molecular evolutionary rate, which is the fuel for speciation. Although other factors such as the formation of isolated populations may trigger diversification, we can conclude that fire acting as a throttle for diversification is by no means the rule in fire-prone ecosystems., We thank R. Ree and B. R. Moore for their advice on the KIT and Sym- meTREE softwares and H. Dopazo for helping with the Phylemon server. B. Lamont provided fire-trait information on some Banksia species. We also thank T. Barraclough, P. Catalin, S. González-Martínez, B. Lamont, J. Pannell, X. Pic6, and R. Ree for helpful comments on the manuscript. This work has been partially financed by the PERSIST project (CGL2006- 07126/BOS) to JGP and by a research grant attached to a "Ramón y Cajal" contract to FO. JGS-M was granted a contract from the I3P program.

14 páginas, 5 figuras, 2 tablas., Microsatellite markers were used to evaluate the genetic diversity and population genetic structure in the critically endangered Limonium dufourii(Plumbaginaceae), a highly endemic triploid species from the coasts of eastern Spain. Sixty-five alleles from 13 microsatellite regions were amplified in a sample of 122 individuals collected from the six extant populations. Microsatellite patterns were consistent with the triploid nature of L. dufourii. Alleles were unambiguously assigned to two different parental subgenomes in this hybrid species and the greater contribution of the diploid parental subgenome was confirmed. Eleven, 25 and 26 multilocus genotypes were recorded from the haploid, diploid and from the combined information of both subgenomes, respectively. Genetic diversity was mostly distributed among populations (72.06% of the total genetic variation). Genotypes from Marjal del Moro populations grouped into two highly structured clusters (88.41% of the total variance). The observed patterns of distribution of genetic diversity are interpreted to result from multiple hybridization events and isolation between populations. Threats to this species are mainly anthropogenic (urbanization and tourism pressure), although stochastic risks cannot be ignored. Therefore, in order to preserve extant genetic variation of L. dufourii, in situ strategies such as the preservation of its habitat are a high priority. Several recommendations in order to assist ex situ measures to guarantee the success of conservation strategies and maintain the relationships between individuals and populations are proposed., We are indebted to S. Rodríguez, J.A. Rosselló and the staff of the Consellería de Medio Ambiente for field assistance; C. Palacios for fruitful comments during the development of this study, S. Kresovich and S.M. Mitchell for technical assistance, and Andrew McCabe for help in revising our use of the English language. We thank the Generalitat Valenciana (GV) for providing permission for collecting samples. This study has been supported by project PB98-1436 from DGES-MEC. MPE was supported by a PhD grant from the GV and JGSM by an Itinerario Integrado de Inserción Profesional (I3P) postdoctoral fellowship from the Spanish Consejo Superior de Investigaciones Científicas (CSIC).

Plant–plant facilitation is a crucial ecological process, as many plant species (facilitated) require the presence of an established individual (nurse) to recruit. Some plant facilitative interactions disappear during the ontogenetic development of the facilitated plant but others persist, even when the two plants are adults. We test whether the persistence of plant facilitative interactions is explained by the phylogenetic diversity of mutualistic and non-mutualistic fungi that the nurse and the facilitated species add to the shared rhizosphere. We classify plant facilitative interactions as persistent and non-persistent interactions and quantify the phylogenetic diversity of mutualistic and non-mutualistic fungi added by the plant species to the shared rhizosphere. Our results show that the facilitated species add less phylogenetic diversity of non-mutualistic fungi when plant facilitative interactions persist than when they do not persist. However, persistent and non-persistent facilitative interactions did not differ in the phylogenetic diversity of mutualistic fungi added by the facilitated species to the shared rhizosphere. Finally, the fungal phylogenetic diversity added by the nurse to the shared rhizosphere did not differ between persistent and non-persistent interactions. This study suggests that considering the fungal associates of the plant species involved in facilitative interactions can shed light on the mechanisms of persistence for plant–plant interactions., This work was funded by the Agencia Española de Cooperación Internacional para el Desarrollo (projects A017475/08, A023461/09), the Dirección General de Asuntos del Personal Académico-Universidad Nacional Autónoma de México (projects IN-202811-3; 213414-3), the Programa Iberoamericano de Ciencia y Tecnología para el Desarrollo (Acción 409AC0369) and the Ministerio de Ciencia e Innovación (CGL2014-58333-P). A. M. N. was supported by a postdoctoral contract from the Ministerio de Economía y Competitividad (FPDI-2013-16266) and an Early Career Project Grant from the British Ecological Society (3975–4849) and J. G. S.-M. by a Ramón y Cajal postdoctoral contract from the Ministerio de Ciencia e Innovación.

Plant survival in alpine habitats is controlled, in several cases, by pollination and seed dispersal success. We have investigated the genetic structure and mating patterns of the endangered Borderea pyrenaica (Dioscoreaceae), one of the oldest herbaceous Pyrenean mountain plants. Simple sequence repeat-based genotyping was carried out on all the reproductive female and male individuals and in all the female-descendent progenies of a population of this plant. Although the offspring sampling (246) was twice the size of the adult sampling (122), the latter group showed higher levels of heterozygosity and approximately 20% more alleles than the offspring. Probabilistic spatial neighbourhood modelling of parentage analysis, based on the exponential-power type model, showed immigration rates of pollen at 63.3%. The present study also detected a strong spatial clustering; most of the sired seeds of B. pyrenaica (68.83%) occurred at distances of up to 20 m, whereas kinship coefficients of adult plants reached zero at spatial distances (d) < 5 m, and 5 < d < 10 m for females and males, respectively. These results support the hypothesis of a terrestrial ant-mediated, rather than a flying insect-mediated pollination in B. pyrenaica. © 2015 The Linnean Society of London., We thank Juan José Robledo-Arnuncio and two anonymous reviewers for their valuable comments and for critically reviewing an earlier version of the manuscript, as well as Emily Lemonds and Douglas Laing for linguistic assistance. The present study has been funded by a Fundación BBVA project grant BIOCON05/093. E. Pérez-Collazos was supported by a Postdoctoral grant from the Fundación BBVA. J. G. Segarra-Moragues was funded by two consecutive Spanish Aragón Government ‘Araid’ and Spanish Ministry of Science and Innovation ‘Ramón y Cajal’ postdoctoral contracts, respectively. The work was partially supported by a Bioflora research group grant, co-funded by the Aragón Government and the European Social Fund.

Some plant secondary compounds, such as terpenes, are very flammable; however, their role in enhancing plant flammability is poorly understood and often neglected in reviews on plant chemical ecology. This is relevant as there is growing evidence that flammability-enhancing traits are adaptive in fire-prone ecosystems. We analyzed the content of monoterpenes and sesquiterpenes, performed flammability tests and genotyped microsatellite markers, all in the same individuals of Rosmarinus officinalis, to evaluate the link between the content of terpenes, flammability and the genetic similarity among individuals. The results suggest that terpenes enhance flammability in R. officinalis, and that variability in flammability among individuals is likely to have a genetic basis. Overall our results suggest that the capacity to produce and store terpenes can be considered a flammability-enhancing trait and could have an adaptive value in fire-prone ecosystems., This work was funded by the VIRRA and TREVOL projects (CGL2009-12048/BOS, CGL2012-39938-C02-01) from the Spanish government. B. M. was supported by a grant from the Portuguese government (Fundação para a Ciência e a Tecnologia; SFRH/BPD/90277/2012), G. A. A. by a Juan de la Cierva (JDC-2009-5067) post-doc from the Spanish Ministerio de Ciencia e Innovación (MICINN), and J. G. S.-M. by a Ramón y Cajal postdoc also from MICINN.

[Background and Aims] Specificity in biotic interactions is mediated' by functional traits inducing shifts in the community species composition. Functional traits are often evolutionarily conserved, resulting in closely related species tending to interact with similar species. This tendency may initially shape the phylogenetic composition of coexisting guilds, but other intraguild ecological processes may either blur or promote the mirroring of the phylogenetic compositions between guilds. The roles of intra- and interguild interactions in shaping the phylogenetic community composition are largely unknown, beyond the mere selectivity in the interguild interactions. Plant facilitation is a phylogenetically structured species-specific process involving interactions not only between the same guild of plants, but also between plants and other guilds such as arbuscular mycorrhizal fungi (AMF). In this study it is hypothesized that reciprocal plant–AMF interactions will leave an interdependent phylogenetic signal in the community composition of both plants and AMF., [Methods] A correlation was used to test for a relationship between the phylogenetic composition of plant and AMF assemblages in a patchy xeric shrubland environment shaped by plant facilitation. In addition, a null model was used to test whether this correlation can be solely explained by selectivity in plant–AMF interactions., [Key Results] A significant correlation was observed between the phylogenetic composition of plant and AMF assemblages. Plant phylogenetic composition in a patch was related to the predominance of plant species with high nursery quality that can influence the community assembly. AMF phylogenetic composition was related to the AMF phylogenetic diversity in each patch., [Conclusions] This study shows that shifts in the phylogenetic composition of plants and AMF assemblages do not occur independently. It is suggested that besides selectivity in plant–AMF interactions, inter-related succession dynamics of plants and AMF within patches could be an ecological mechanism driving community assembly. Future lines of research might explore whether interlinked above- and below-ground dynamics could be occurring across multiple guilds simultaneously., This work was funded by AECID (Projects A017475/08, A023461/09), DGAPA-UNAM (Project IN-202811-3; IN-213414-3), CYTED (Acción 409AC0369) and MICINN (CGL2011-29585-C02-01). A.M.N. was supported by a DGAPA-UNAM post-doctoral fellowship and an Early Career Project Grant from the BES (3975-4849), and J.G.S.-M. was supported by a ‘Ramón y Cajal’ post-doctoral contract from MICINN.

Polyploidy is a common phenomenon occurring in a vast number of land plants. Investigations of patterns of inheritance and the origins of plants (i.e., autopolyploidy vs. allopolyploidy) usually involve cytogenetic and molecular studies of chromosome pairing, chromosome mapping, and marker segregation analysis through experimental crosses and progeny tests. Such studies are missing for most wild species, for which artificial crosses are difficult, not feasible, or unaffordable. We report here a Bayesian method to discriminate between alternative inheritance patterns in the two extant, tetraploid species of the monocot genus Borderea (Dioscoreaceae), which does not involve progeny array tests. Our approach is based on the screening of a large number of SSR genotypes, which were obtained from successful amplifications of 17 microsatellite regions in individuals of both B. chouardii and B. pyrenaica. We tested for tetrasomic vs. disomic modes of inheritance, using the Bayes factor test. Assignment of genotypes under both alternatives could be unequivocally done for 14 and 11 of the 17 studied microsatellite regions in B. chouardii and B. pyrenaica, respectively, totaling 9502 analyzed genotypes. The comparison of posterior probabilities for the two competing hypotheses across the surveyed loci clearly favored a disomic inheritance pattern. Linkage tests indicated that none of the studied SSR loci were in linkage disequilibrium, thus representing independent samples of the Borderea genome. These results, along with previous allozyme data, support the allotetraploid origin of this paleoendemic genus and reveal the lowest reported chromosome base number for the family of the yams.

Background and Aims Molecular markers have changed previous expectations about germplasm collections of endangered plants, as new perspectives aim at holding a significant representation of all the genetic diversity in the studied species to accomplish further conservation initiatives successfully. Borderea chouardii is a critically endangered allotetraploid dioecious member of Dioscoreaceae, known from a single population in the Iberian pre- Pyrenees. This population was reported to be highly structured into two genetically distinct groups of individuals corresponding to their spatial separation along the vertical cliff where it grows. In 1999, the Spanish Government of Aragon launched the first conservation programme for the ex situ preservation of this species, and since then a seed collection has been conserved at the Germplasm Bank of the Universidad Politecnica de Madrid. However, as some seed samples had not been labelled clearly at the time of collection, their origin was uncertain. Methods Genetic variation in germplasm accessions of B. chouardii was investigated using microsatellite (simple sequence repeat; SSR) markers. Key Results The 17 primer pairs used detected 62 SSR alleles in the 46 samples analysed from five different germplasm stocks. Eight alleles scored from the wild population were not detected in the germplasm samples analysed. The relatedness of the germplasm samples to the wild subpopulations through neighbour-joining clustering, principal coordinates analysis (PCO) and assignment tests revealed a biased higher representation of the genetic diversity of the lower cliff (43 samples) subpopulation than that of the upper cliff (three samples). Conclusions The collection of additional samples from the upper cliff is recommended to achieve a better representation of the genetic diversity of this subpopulation. It is also recommended that these stocks should be managed separately according to their distinct microspatial origin in order to preserve the genetic substructuring of the wild population.

Borderea chouardii is a relictual and dioecious, strictly sexually reproducing, long-living geophyte of the Dioscoreaceae family. Previous biological and demographic studies have indicated the existence of a uniformly distributed panmictic population of this taxon at the southernmost Spanish pre-Pyrenean mountain ranges where it occurs in rather inaccessible crevices of a single limestone cliff. However, individuals of B. chouardii are spatially subdivided into two subpopulations located, respectively, on the upper and lower parts of the cliff, and vertically separated 150 m. Because of its extreme rarity, B. chouardii was the first Iberian taxon to have a specific conservation plan and has been included in several red lists under the category of critically endangered (CR). However, no previous attempts have been conducted to analyse the fine scale evolutionary mechanisms involved in its present microspatial distribution. Genetic diversity and population structure have been investigated through the analysis of neutral hypervariable markers such as simple sequence repeats (SSRs) and randomly amplified polymorphic DNAs (RAPDs) to unravel the impact of life history traits in the differentiation of the two subpopulations. Both types of molecular markers were unequivocal in distinguishing two genetically distinct groups of individuals corresponding to their spatial separation. However, SSRs detected a higher level of subpopulation differentiation (F(ST) = 0.35, R(ST) = 0.32) than RAPDs (F(ST) = 0.21). SSR data indicated significant deviation from random dispersal of genes and genotypes between the two groups, suggesting that mating occurs mainly among individuals within subpopulations, thus, favouring the divergence between the two groups. This microevolutionary differentiation scenario might have been caused by a coupled effect of past genetic drift and reproductive isolation, as a result of strong glacial age bottlenecks and inefficient dispersal system of pollen and seeds, respectively. The identification of such genetic structure in this narrow endemic prompts a modification of the management strategies of its single extant population.