M. Montserrat Martínez-Ortega

A new classification of the tribe Veroniceae: problems and a possible solution

Based on the most recent evidence from phylogenetic analyses of Veronica and its related genera, we propose a new infratribal and infrageneric rearrangement for Veroniceae and Veronica. These recent analyses have settled the long dispute about the relationship of the Northern Hemisphere Veronica and the Southern Hemisphere Hebe complex and have shown the derivation of the latter from within the former. Other currently recognized genera such as Synthyris and Pseudolysimachion are also derived from within Veronica. A classification based on monophyletic genera therefore needs to either lump some well recognized genera into a large genus Veronica or split Veronica into several genera that seem impossible to separate using morphological or structural characters. We discuss the advantages and disadvantages and conclude that it is best to recognize a large genus Veronica (including Hebe, Parahebe, Chionohebe, Heliohebe, Detzneria, Derwentia, Pseudolysimachion, Synthyris and Besseya) with 13 subgenera, along with 8 other genera, Scrofella, Veronicastrum (inclusive of Calorhabdos and Botryopleuron), Lagotis, Wulfenia, Kashmiria, Picrorhiza, Wulfeniopsis and Paederota, in Veroniceae.

Species Boundaries and Phylogeographic Patterns in Cryptic Taxa Inferred from AFLP Markers: Veronica subgen. Pentasepalae (Scrophulariaceae) in the Western Mediterranean

Abstract We show the power of detailed AFLP study to address questions of species delimitation and genetic diversity in morphologically-cryptic plants. We have combined this technique with karyological studies to explore species boundaries in the Western Mediterranean representatives of Veronica subgen. Pentasepalae. Several members of this subgenus are represented in the Iberian Peninsula and North Africa by morphologically similar populations, which lead to different taxonomic hypotheses. We have analyzed genetic variation with two different objectives: first, to provide a basis on which one taxonomic hypothesis is favored against its alternatives, and second, to obtain a satisfactory interpretation of phylogeographic patterns. The usefulness of several parameters and analyses is considered in order to help rank taxa and to reveal details of their speciation modes. Some of these parameters are presented with a geographical background to facilitate phylogeographic interpretations. The data support one of the previously proposed taxonomic hypotheses, which results in the recognition of eight taxa. An east-to-west migration route within the Iberian Peninsula is hypothesized, coinciding with the general pattern proposed for Veronica subgen. Pentasepalae. Further data on genetic variability and chromosome counts for 41 populations are provided.

Veronica: Parallel morphological evolution and phylogeography in the Mediterranean

Abstract.The genus Veronica s. lat. comprises about 450 species (including about 180 species from the southern hemisphere Hebe-complex), many of which grow in the Mediterranean area. Their extreme variability in morphology, life form and habitats has led to many suggestions regarding evolution and biogeography. Difficulties arise from parallel syndromes, widespread among alpine species and lowland perennials, and particularly among annual species of the genus. We have used sequences of the plastid trnL-F region and nuclear ribosomal ITS sequences to differentiate between different clades of Veronica and reveal cases of parallel evolution. Based on this data, cases of parallel evolution have been found in biogeographical patterns among the alpine species of Veroniceae, in which species from European mountains have affinities to those in the Central Asian/Himalayan region whereas alpine species from Turkey are probably more recently derived from lowland southwestern Asian taxa. Different subspecies of Veronica bombycina gained their characteristic morphology independently and parallel in adaptation to their alpine environment. Pinnatifid leaves have been gained parallel in perennial grassland species of Veronica. Finally, parallel evolutionary trends in many characters, not only morphological but also molecular characters, are common among annual species of Veronica.

Assessing extinction-risk of endangered plants using species distribution models: a case study of habitat depletion caused by the spread of greenhouses

The species distribution models (SDMs) are useful tools for investigating rare and endangered species as well as the environmental variables affecting them. In this paper, we propose the application of SDMs to assess the extinction-risk of plant species in relation to the spread of greenhouses in a Mediterranean landscape, where habitat depletion is one of the main causes of biodiversity loss. For this purpose, presence records of the model species (Linaria nigricans, a endemic and threatened species) and the greenhouses, a dataset of environmental variables, and different only presence-based modelling algorithms (Bioclim, Domain, GARP, MaxEnt and ENFA) were used to build SDMs for L. nigricans as well as for greenhouses. To evaluate the models a modified approach of the area-under-curve ROC was applied. Combining the most accurate models, we generated an extinction-risk model of L. nigricans populations, which enabled us to assess the sustainability of the most threatened populations. Our results show that is possible to model greenhouses spreading as a “biological invasion”. The procedure explained and used in this work is quite novel, and offers an objective spatial criterion intended for the management of natural resources and for the conservation of the biodiversity in areas threatened by habitat depletion processes as particular as greenhouses expansion.

The challenge of species delimitation in the diploid-polyploid complex Veronica subsection Pentasepalae

A reliable taxonomic framework and the identification of evolutionary lineages are essential for effective decisions in conservation biodiversity programs. However, phylogenetic reconstruction becomes extremely difficult when polyploidy and hybridization are involved. Veronica subsection Pentasepalae is a diploid-polyploid complex of ca. 20 species with ploidy levels ranging from 2x to 10x. Here, DNA-ploidy level estimations and AFLP fingerprinting were used to determine the evolutionary history, and species boundaries were reviewed in an integrated approach including also previous data (mainly morphology and sequence-based phylogenetic reconstructions). Molecular analyses were performed for 243 individuals from 95 populations, including for the first time all taxa currently recognized within the subsection. Phylogenetic reconstruction identified four main groups corresponding almost completely to the four clusters identified by genetic structure analyses. Multiple autopolyploidization events have occurred in the tetraploid V. satureiifolia giving rise to octoploid entities in central Europe and north of Spain, whereas hybridization is demonstrated to have occurred in several populations from the Balkan Peninsula. Furthermore, our study has established the taxonomic status of taxa, for the most part recovered as monophyletic. Cryptic taxa within the group have been identified, and a new species, Veronica dalmatica, is fully described. This study highlights the implications of polyploidy in species delimitation, and illustrates the importance to conserve polyploid populations as potential sources of diversification due to evolutionary significance of genome duplications in plant evolution.

Systematic significance and evolutionary dynamics of the achene twin hairs in Filago (Asteraceae, Gnaphalieae) and related genera: further evidence of morphological homoplasy

The variation and evolutionary dynamics of the twin hairs covering the achenes of the external female florets in Filago and related genera are studied using scanning electron microscope (SEM) and a Bayesian reconstruction of ancestral character states. A molecular phylogenetic hypothesis based on sequences of ITS and ETS (nuclear ribosomal DNA) and plastid rpl32-trnL intergenic spacer regions was used as a framework to test the phylogenetic, evolutionary and taxonomic significance of this character in the study group. The phylogenetic position of three island-endemic species (F. petro-ianii, F. tyrrhenica and F. wagenitziana) is explored. Three types of twin hairs are described, which alone or combined define five different types of indumentum. The results suggest that glabrous achenes are the ancestral condition for “Filago group s.str.”, while the presence of long-clavate twin hairs on the achene represents the ancestral character state for the genus Filago. Several cases of parallelisms are detected. With regard to taxonomy, twin hairs provide additional support for the recognition of Bombycilaena and Logfia at the generic level and allow the taxonomic re-evaluation of F. lutescens subsp. atlantica and F. crocidion. Finally, the utility of a Bayesian Binary MCMC approach (using the software RASP) for evolutionary reconstruction of morphological characters is demonstrated.

Characterization of 12 Polymorphic SSR Markers in Veronica Subsect. Pentasepalae (Plantaginaceae) and Cross-Amplification in 10 Other Subgenera

Premise of the study: Microsatellite primers were developed in the perennial herbs of the diploid-polyploid complex Veronica subsect. Pentasepalae (Plantaginaceae) to investigate the role that hybridization has played in the evolution of the group, which includes several endangered species. Methods and Results: Twelve pairs of primers leading to polymorphic and readable markers were identified and optimized from V. jacquinii and V. orbiculata using a microsatellite-enriched library method and 454 GS-FLX technique. The set of primers amplified dinucleotide to pentanucleotide repeats, and the number of alleles per locus ranged from one to six, one to 11, and one to nine for V. orsiniana, V. javalambrensis, and V. rosea, respectively. Transferability analyses were performed in 20 species corresponding to 10 different subgenera. Conclusions: These results indicate the utility of the newly developed microsatellites across Veronica subsect. Pentasepalae, which will help in the study of gene flow patterns and genetic structure.

Designing conservation strategies to preserve the genetic diversity of Astragalus edulis Bunge, an endangered species from western Mediterranean region

Astragalus edulis (Fabaceae) is an endangered annual species from the western Mediterranean region that colonized the SE Iberian Peninsula, NE and SW Morocco, and the easternmost Macaronesian islands (Lanzarote and Fuerteventura). Although in Spain some conservation measures have been adopted, it is still necessary to develop an appropriate management plan to preserve genetic diversity across the entire distribution area of the species. Our main objective was to use population genetics as well as ecological and phylogeographic data to select Relevant Genetic Units for Conservation (RGUCs) as the first step in designing conservation plans for A. edulis. We identified six RGUCs for in situ conservation, based on estimations of population genetic structure and probabilities of loss of rare alleles. Additionally, further population parameters, i.e. occupation area, population size, vulnerability, legal status of the population areas, and the historical haplotype distribution, were considered in order to establish which populations deserve conservation priority. Three populations from the Iberian Peninsula, two from Morocco, and one from the Canary Islands represent the total genetic diversity of the species and the rarest allelic variation. Ex situ conservation is recommended to complement the preservation of A. edulis, given that effective in situ population protection is not feasible in all cases. The consideration of complementary phylogeographic and ecological data is useful for management efforts to preserve the evolutionary potential of the species.

Unravelling the phylogeny of the root-hemiparasitic genus Odontites (tribe Rhinantheae, Orobanchaceae): Evidence for five main lineages

Despite the recent publication of several phylogenies focused on Rhinantheae, which has been expanded to include three Asian endemic genera, few studies so far have dealt with particular genera within the tribe. Here, we focus on Odontites and related genera because of the high morphological variability of the group and its unclear generic boundaries. Phylogenetic analyses were performed for nrDNA (ITS) and cpDNA (trnK region and rps16 intron) datasets, using Bayesian and Parsimony analyses. Our results cast doubt on the inclusion of Pterygiella and related genera within the Rhinantheae and support the paraphyly of Phtheirospermum, making it necessary to propose three new combinations to avoid it. Odontites is recircumscribed to include Bartsiella, Bornmuellerantha, and Macrosyringion, but not Odontitella. Within Odontites, five distinct lineages are identified. These are distinguishable either by morphological synapomorphies or by a combination of several character states. Most of the Odontites species are regarded as monophyletic. In the O. vernus and O. luteus complexes, some taxonomic changes are made to avoid paraphyly, which results in three new combinations.

A nomenclatural treatment for Veronica subsect. Pentasepalae (Plantaginaceae sensu APG III) and typification of several names

The large genus Veronica has a complex evolutionary history. Within it, Veronica subsect. Pentasepalae represents an ideal system to study important evolutionary processes, such as hybridization and polyploidization. Delimitation of species boundaries within the subsection is a difficult task and the highly complicated taxonomy of the group has led to an accumulation of approximately 230 names for 22 accepted taxa. Many names have been used in different taxonomic senses even in recent Floras, scientific works, catalogues, Red Lists and internet resources. The lack of an updated taxonomic and nomenclatural framework represents a significant problem to develop further studies in several fields such as ecology, conservation, plant and evolutionary biology. A complete nomenclatural treatment for the whole subsection is provided here, that relies on results derived from the first phylogenetic analysis of V. subsect. Pentasepalae based on DNA sequence data, morphology and ploidylevel information. Nomenclatural types for 22 validly published names are designated. In total 21 lectotypes, 2 epitypes and 1 neotype are chosen. A new combination, Veronica linearis, is proposed.