The genus Mantisalca (Asteraceae) in southwest Europe

The diversity of the genus Mantisalca in southwest Europe is the subject of much controversy. The present taxonomic revision of the genus on the Iberian Peninsula allows four species to be recognized, whose morphological variability is described and various synonyms are typified. A new species is described, M. cabezudoi Ruiz de Clavijo and Devesa, endemic to southeast Spain. The name Microlonchus spinulosus Rouy is reinstated for a species endemic to the central and east Iberian Peninsula, for which a new combination is made: Mantisalca spinulosa (Rouy) Ruiz de Clavijo and Devesa. A diagnostic key to the taxa is given.     

Ecological differentiation and cladogenesis of Baldellia (L.) Parl. (Alismataceae)

The Iberian Peninsula is home to a large number of endemic species that often show strong genetic subdivisions which indicate possible isolation of populations in the past. Numerous phylogeographic and phylogenetic studies on Iberian flora and fauna have revealed this territory to be the origin of genetic differentiation during the Pleistocene Ice Age. To better understand the influence of Plio-Pleistocene climatic oscillations on the cladogenesis and population structuring of refugial taxa restricted to freshwaters of the Iberian Peninsula, we studied the evolutionary history of the small European genus Baldellia, both from a phylogenetic/phylogeographic perspective and regarding its ecological characteristics. Molecular analyses were based on four plastid regions and nuclear ITS2 ribosomal spacer from individuals belonging to geographically isolated Baldellia populations, while the relationships between climate and Baldellia species distribution was investigated by redundancy analysis applied to the climatic data with the occurrence of Baldellia species as external variables. The absence of a clear phylogenetic signal most likely reflects a common and recent cladogenesis for Baldellia. In contrast, the more rapidly evolving plastid sequences indicate a recent phylogeographic history of refugial repartition and ecological segregation probably occurring during the last glaciation, a scenario well supported by marked differences in species?? ecological preferences and climate analysis.     

Ecology, genetic diversity and phylogeography of the Iberian endemic plant Jurinea pinnata (Lag.) DC. (Compositae) on two special edaphic substrates: dolomite and gypsum

Background and aims

Jurinea pinnata is an Iberian vascular plant which only grows on gypsum and dolomite, two types of rocks associated with their exclusive endemic floras. In addition, the plant has an island-like distribution which could affect the differentiation and the genetic variability of wild populations. Thus, the species provides a unique opportunity for comparing (bio)geographical and ecological (edaphic) differentiation by means of molecular markers.

Methods

For our investigation we took 24 soil samples paired with a similar number of foliar samples for nutritional analyses. Our molecular-marker approach (AFLPs) involved 16 populations.

Results

The edaphic parameters revealed significant dissimilarities between dolomitic and gypsum soils. These differences are also found in the mineral composition of the leaves. However, molecular data revealed that the differentiation between populations correlates better with geographical isolation than with the substrate character.

Conclusions

The populations showing the greatest genetic diversity are those of the East Baetic territory where the species grow on both substrates and its populations are closer together. The plant tolerance to gypsum and dolomite can be explained either as a result of common adaptive mechanisms or of a more general adaptation to arid environments.     

Appearances can be deceptive: different diversification patterns within a group of Mediterranean earthworms (Oligochaeta, Hormogastridae)

Many recent studies on invertebrates have shown how morphology not always captures the true diversity of taxa, with cryptic speciation often being discussed in this context. Here, we show how diversification patterns can be very different in two clades of closely related earthworms in the genus Hormogaster stressing the risk of using nonspecific substitution rate values across taxa. On the one hand, the Hormogaster elisae species complex, endemic to the central Iberian Peninsula, shows morphological stasis. On the other hand, a clade of Hormogaster from the NE Iberian Peninsula shows an enormous morphological variability, with 15 described morphospecies. The H. elisae complex, however, evolves faster genetically, and this could be explained by the harsher environmental conditions to which it is confined—as detected in this study, that is, sandier and slightly poorer soils with lower pH values than those of the other species in the family. These extreme conditions could be at the same time limiting morphological evolution and thus be responsible for the observed morphological stasis in this clade. Contrarily, Hormogaster species from the NE Iberian Peninsula, although still inhabiting harsher milieu than other earthworm groups, have had the opportunity to evolve into a greater morphological disparity. An attempt to delimit species within this group following the recently proposed general mixed Yule‐coalescent method showed a higher number of entities than expected under the morphospecies concept, most probably due to the low vagility of these animals, which considerably limits gene flow between distant conspecific populations, but also because of the decoupling between morphological and genetic evolution in the H. elisae complex.     

High levels of population subdivision in a morphologically conserved Mediterranean toad (Alytes cisternasii) result from recent,multiple refugia: evidence from mtDNA,microsatellites and nuclear genealogies

Pleistocene glaciations often resulted in differentiation of taxa in southern European peninsulas, producing the high levels of endemism characteristic of these regions (e.g. the Iberian Peninsula). Despite their small ranges, endemic species often exhibit high levels of intraspecific differentiation as a result of a complex evolutionary history dominated by successive cycles of fragmentation, expansion and subsequent admixture of populations. Most evidence so far has come from the study of species with an Atlantic distribution in northwestern Iberia, and taxa restricted to Mediterranean‐type habitats remain poorly studied. The Iberian Midwife toad (Alytes cisternasii) is a morphologically conserved species endemic to southwestern and central Iberia and a typical inhabitant of Mediterranean habitats. Applying highly variable genetic markers from both mitochondrial and nuclear genomes to samples collected across the species’ range, we found evidence of high population subdivision within A. cisternasii. Mitochondrial haplotypes and microsatellites show geographically concordant patterns of genetic diversity, suggesting population fragmentation into several refugia during Pleistocene glaciations followed by subsequent events of geographical and demographic expansions with secondary contact. In addition, the absence of variation at the nuclear β‐fibint7 and Ppp3caint4 gene fragments suggests that populations of A. cisternasii have been recurrently affected by episodes of extinction and recolonization, and that documented patterns of population subdivision are the outcome of recent and multiple refugia. We discuss the evolutionary history of the species with particular interest in the increasing relevance of Mediterranean refugia for the survival of genetically differentiated populations during the Pleistocene glaciations as revealed by studies in co‐distributed taxa.     

Isolation and characterization of two dinucleotide and four tetranucleotide polymorphic microsatellite loci in the Iberian midwife toad Alytes cisternasii

The Iberian midwife toad Alytes cisternasii is an endemic species of the southwestern and central parts of the Iberian Peninsula. Allozyme analysis of Iberian midwife toads indicated no inter‐ or intrapopulation variation and a more sensitive method is required to quantify genetic differentiation and gene flow between populations of this species. We describe here the development of primers for the amplification by the polymerase chain reaction (PCR) of six polymorphic microsatellite loci in A. cisternasii.     

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

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

A review of the Iberian Sphingonotini with description of two novel species (Orthoptera: Acrididae: Oedipodinae)

The genus Sphingonotus Fieber, 1853 is one of the most species‐rich grasshopper genera in the world. We studied the morphology of c. 1000 individuals from the Iberian Peninsula to review the taxonomy of the genus and its relatives. Moreover, we inferred a molecular phylogeny of the Iberian Sphingonotini based on two mitochondrial genes. The Iberian and north‐west African Sphingonotini comprise two recent radiations, within which the genetic relationships are not fully resolved. A multivariate morphometric analysis showed that S. azurescens (Rambur, 1838) and S. morini (Defaut, 2005) can be clearly discriminated, supporting their species status. Based upon the combined data, the genus Granada Koçak & Kemal, 2008 is synonymized with Sphingonotus Fieber, 1853 and its type species re‐assigned to Sphingonotus imitans Brunner von Wattenwyl, 1882 comb. rev . The data also supported species rank for Sphingonotus lusitanicus Ebner, 1941 comb. rev. Oedipoda callosa Fieber, 1853 is considered as nomen dubium. Two novel species are described: Sphingonotus ( Neosphingonotus ) almeriense Llucià‐Pomares sp. nov . and Sphingonotus (Neosphingonotus) nodulosus Llucia‐Pomares sp. nov. from the southern part of the Iberian Peninsula. The new species are compared with other Iberian Sphingonotini and a key to the species is provided. © 2013 The Linnean Society of London     

A study of Sarcocornia A.J. Scott (Chenopodiaceae) from Western Mediterranean Europe

The latest publications on Sarcocornia taxonomy and phylogeny recognize six taxa in this genus on the Iberian Peninsula: S. perennis, S. fruticosa, S. alpini, S. alpini subsp. carinata, S. hispanica, and S. pruinosa. The present study represents a comprehensive revision of the different taxa in the Sarcocornia genus present in Western Mediterranean Europe by means of morphological, micromorphological and phylogenetic internal transcribed spacer (ITS) analysis. Morphological and micromorphological data were studied from Sarcocornia samples from 113 populations in coastal salt marshes and inland salt pans in Portugal, France, Spain and Italy. Sixteen new ITS sequences were obtained from Mediterranean Sarcocornia species and analysed together with previous reported data. Published karyological, ecological and biogeographical data from Western Mediterranean Europe were also reviewed. The results indicate the presence of a new species, S. lagascae, found growing in coastal Mediterranean areas of the Iberian Peninsula. The species S. fruticosa was found to be absent from the Iberian territories.     

Phylogeography and species delineation of the genus Phoxinus Rafinesque, 1820 (Actinopterygii: Leuciscidae) in the Iberian Peninsula

The genus Phoxinus is comprised of at least 15 currently recognized species inhabiting Eurasia. Morphological traits have been traditionally used to delineate species in Phoxinus; however, the high level of phenotypic plasticity observed in the genus has confounded this process. Molecular genetic analyses have revealed a higher than expected genetic structure within Phoxinus. Here, we analyzed both nuclear and mitochondrial molecular genetic markers to infer the phylogeography and divergence times of Phoxinus in the Iberian Peninsula. Our results show that the Iberian lineages of Phoxinus were polyphyletic. They also support the co‐existence of three species in the Iberian Peninsula, two corresponding to two previously recognized species (Phoxinus bigerri and Phoxinus septimaniae) and a third undescribed species (Phoxinus sp.). Phoxinus bigerri is structured into western Cantabrian, eastern Cantabrian, and Artibai basins. We hypothesize that this structure is a consequence of glaciation–deglaciation cycles during the Pleistocene. While the presence of P. septimaniae in the Iberian Peninsula is possibly the result of human translocation, that of Phoxinus sp. in lower Ebro rivers may be attributed to past fluvial captures. Our study represents the first report to show a relationship among Phoxinus populations from central Pyrenean rivers of Spain and France. Furthermore, we found genetic hybridization between Phoxinus sp. and P. septimaniae in the shared localities, a likely consequence of anthropogenic activities. Overall, our findings provide insight into the genetic structure of Iberian Phoxinus populations, including the presence of an undescribed species and the putative introduction of some species that may have implications for conservation.     

Survival and long-term maintenance of tertiary trees in the Iberian Peninsula during the Pleistocene: first record of Aesculus L. (Hippocastanaceae) in Spain

The Italian and Balkan peninsulas have been places traditionally highlighted as Pleistocene glacial refuges. The Iberian Peninsula, however, has been a focus of controversy between geobotanists and palaeobotanists as a result of its exclusion from this category on different occasions. In the current paper, we synthesise geological, molecular, palaeobotanical and geobotanical data that show the importance of the Iberian Peninsula in the Western Mediterranean as a refugium area. The presence of Aesculus aff. hippocastanum L. at the Iberian site at Cal Guardiola (Tarrasa, Barcelona, NE Spain) in the Lower–Middle Pleistocene transition helps to consolidate the remarkable role of the Iberian Peninsula in the survival of tertiary species during the Pleistocene. The palaeodistribution of the genus in Europe highlights a model of area abandonment for a widely-distributed species in the Miocene and Pliocene, leading to a diminished and fragmentary presence in the Pleistocene and Holocene on the southern Mediterranean peninsulas. Aesculus fossils are not uncommon within the series of Tertiary taxa. Many appear in the Pliocene and suffer a radical impoverishment in the Lower–Middle Pleistocene transition. Nonetheless some of these tertiary taxa persisted throughout the Pleistocene and Holocene up to the present in the Iberian Peninsula. Locating these refuge areas on the Peninsula is not an easy task, although areas characterised by a sustained level of humidity must have played an predominant role.     

Paleobiogeography of an Iberian endemic species,Luciobarbus sclateri (Günther, 1868) (Actinopterygii,Cyprinidae), inferred from mitochondrial and nuclear markers

Since the Cenozoic Era, the southern Iberian Peninsula has undergone a series of complex geological and climatic changes that have shaped the hydrographic configuration of the freshwater network, influencing the present‐day distribution of primary freshwater species and favoring a high level of local endemicity. The cyprinid species Luciobarbus sclateri (Günther, 1968) is an endemic species confined to the southern Iberian Peninsula and characterized by a complex evolutionary history. Previous studies linked the structure of L. sclateri populations to the effects of climate change during glaciations and were not able to explain the genetic discordance found between nuclear and mitochondrial markers. The results of this study show that the structure of L. sclateri populations is a reflection of diversification processes linked to the geological history of the region. Thus, we found three main mitochondrial phylogroups: the first one corresponding to small basins in southern Iberian Peninsula, a second one in eastern Iberian Peninsula, corresponding to Segura population, and a third one including the rest of the basins where the species is distributed. The southern group began diverging in the Pliocene as result of tectonic dynamics characterized by the emersion of the basins around the Strait of Gibraltar. The other two groups began diverging with the formation of the current Iberian hydrographic system during Pleistocene. So, the isolation of the hydrographic basins was the main factor driving intraspecific differentiation, followed by recent secondary contacts, admixture, and re‐isolation of the populations.     

Climatic and geographic effects on the spatial genetic pattern of a landbird species (Alectoris rufa) on the Iberian Peninsula

Understanding the spatial pattern of genetic diversity may be pivotal to adaptive conservation management of a given taxon. The red-legged partridge (Alectorisrufa, Linnaeus 1758) is naturally widely distributed from the Mediterranean to humid temperate zones. According to a recent study, the genetic structure of this species comprises five clusters, three of which are in the Iberian Peninsula (glacial refugia). Partridge demographic expansion events and climatic shifts during Pleistocene glaciations have been used to test the hypotheses concerning Iberian red-legged partridge distribution. We tested the existence of climatic and geographic relationships on genetic diversity/distances. We employed markers from two different genetic systems, such as part of the mitochondrial DNA control region (n = 113) and 20 species-specific microsatellite DNA loci (n = 377), including climatic and geographic factors from the 14 Iberian localities where A. rufa populations were sampled. Our results showed a mitochondrial genetic diversity pattern associated with a thermic gradient, and a decrease of genetic diversity in peripheral populations that concurred with the ‘abundant centre’ hypothesis. Overall, current climatic variables reliably described genetic variation and differentiation in the red-legged partridge, which may be a result of local species adaptation.     

Morpho-environmental characterization of the genus Baldellia Parl. (Alismataceae) in the Iberian Peninsula,Balearic islands and North Morocco

Abstract

The genus Baldellia Parl. has always been a complex taxon. Three species and three subspecies have been proposed for the Iberian Peninsula. The morphological variation of this genus in the Iberian Peninsula, Balearic islands and northern Morocco, and its thermo-pluviometric and altitudinal correlation are the focus of this contribution. Twenty-eight morphological variables were examined and subjected to a multivariate statistical analysis. The three morphotypes observed (Form 1, which includes B. ranunculoides subsp. repens and B. alpestris; Form 2, which includes B. ranunculoides and Form 3 – for the new combination B. ranunculoides subsp. ranunculoides var. tangerina (Pau) J. Rocha, A. Crespí, M. García-Barriuso, R. Almeida, J. Honrado, comb. nova proposed here for the first time) seem to represent different reproductive strategies. Morphological variables related with the presence or absence of stolons and the architecture of the inflorescence; the size and number of fruits are the most discriminating variables. Form 1 represents the asexual morphotype; Forms 2 and 3 correspond to morphotypes for which sexual reproduction is preferential. The environmental approach revealed that the asexual form (Form 1) grows in temperate, in more humid conditions, and within a broad altitudinal range. In contrast, the two sexual forms are more common in warmer and drier conditions, and occur over a narrower interval of altitudes.     

Karyology of theScorzonera (Compositae) species from the Iberian Peninsula

A karyological study of 15 taxa ofScorzonera L. from the Iberian Peninsula has been made. The chromosome numbers found inS. hispanica var.pinnatifida, S. baetica, S. reverchonii, S. angustifolia, S. laciniata var.calcitrapifolia and var.subulata (2n = 14) are new. Diploid cytotypes with 2n = 14 and 2n = 12 prevail, andS. hispanica var.crispatula is the only taxon which exhibits autopolyploidy (2n = 14, 28). x = 7 is considered to be the base chromosome number within the genus, with x = 6 being derived from it by translocation. This and detailed karyotype analyses allow to group the Iberian Peninsula species ofScorzonera into three groups.     

MtDNA markers reveal the existence of allopatric evolutionary lineages in the threatened lampreys <Emphasis Type="Italic">Lampetra fluviatilis</Emphasis> (L.) and <Emphasis Type="Italic">Lampetra planeri</Emphasis> (Bloch) in the Iberian glacial refugium

The Iberian Peninsula has been identified as an important glacial refugium during the Pliocene and Pleistocene epochs for the genus Lampetra, providing intermittent refuge and postglacial opportunities for colonization and expansion. We used mitochondrial DNA markers to investigate the processes that have shaped present-day genetic constitution of the genus Lampetra within the Iberian Peninsula. We surveyed 1,173 bp of the cytochrome b gene and 829 bp of the genes ATPase subunits 6 and 8 in 233 individuals of Lampetra fluviatilis (L.) and Lampetra planeri (Bloch) from 21 localities along their distribution range in the Iberian Peninsula. We identified four highly divergent allopatric evolutionary lineages that evolved by fragmentation during the Pliocene and Pleistocene likely driven by environmental factors, namely regional geomorphic events. The high level of genetic divergence between the four lineages suggests that sufficient time has apparently passed by to originate a complex of incipient or cryptic resident species and allows the definition of four evolutionary significant units (ESUs) for L. planeri and one ESU for L. fluviatilis. These findings have important consequences for the interpretation of refugia biological diversity and have major implications on the conservation of these threatened species.     

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

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

Nymphal feeding habits of Perla madritensis Rambur 1842 (Plecoptera: Perlidae)

The diet of Perla madritensis, endemic species of the northern half of the Iberian Peninsula, is described for the first time by means of a study carried out in north-western Spain. As other species of the genus Perla, this taxon behaves mainly as predator, with Chironomidae, followed by Baetidae, being the most abundant prey in its gut. Shifts in diet composition were detected in relation to size in this species, with smaller prey being replaced by larger ones such as Simuliidae and Leptophlebiidae when the nymphs become larger.     

Phylogeography of Iberian Discoglossus (Anura: Discoglossidae)

The genus Discoglossus (Anura, Discoglossidae) comprises five morphologically similar species. Molecular analyses have found high values of genetic differentiation among these cryptic taxa and confirmed the existence of two Iberian endemic lineages ( Discoglossus galganoi and D. jeanneae ) and a distinct lineage of uncertain origin representing a third species in NE Spain. In order to analyse the pattern of geographical distribution of haplotype diversity within Iberian Discoglossus and test the hypothesis of an Algerian origin for populations in NE Spain sequence data has been resolved from 35 populations of D. galganoi and D. jeanneae on the Iberian Peninsula and samples from NE Spain and north Africa. I analysed 959 bp corresponding to partial sequences of cytochrome b and nad 4 and found extremely low values of sequence divergence among populations of D. jeanneae . Three clades can be recognized within D. galganoi , however, and a statistically significant association with geography can be explained as a result of past fragmentation. Sequence data strongly support an Algerian origin for populations of Discoglossus from NE Spain. Finally, the taxonomic status of Discoglossus in Morocco is discussed and its elevation to species status as Discoglossus scovazzi Camerano, 1878 is suggested.