Juan José Aldasoro

Reconstructing the history of Campanulaceae with a Bayesian approach to molecular dating and dispersal–vicariance analyses

We reconstruct here the spatial and temporal evolution of the Campanula alliance in order to better understand its evolutionary history. To increase phylogenetic resolution among major groups (Wahlenbergieae-Campanuleae), new sequences from the rbcL region were added to the trnL-F dataset obtained in a previous study. These phylogenies were used to infer ancestral areas and divergence times in Campanula and related genera using a Bayesian approach to molecular dating and dispersal-vicariance analyses that takes into account phylogenetic uncertainty. The new phylogenetic analysis confirms Platycodoneae as the sister group of Wahlenbergieae-Campanuleae, the two last ones inter-graded into a well-supported clade. Biogeographic and dating analyses suggest that Western Asia and the Eastern Mediterranean have played a major role as centers of migration and diversification within the Campanula alliance, probably in relation to the intense orogenic activity that took place in this region during the Late Neogene, and that could have promoted isolation and allopatric speciation within lineages. Diversification rates within several Campanula lineages would have increased at the end of the Miocene, coinciding with the Messinian Stage. Strong selective pressures from climate changes and the expansion of mountainous regions during this period are suggested to explain the adaptation to drought, cold or disturbed environments observed in many Campanula species. Several independent long-distance dispersal events to North America are inferred within the Rapunculus clade, which seem to be related to high ploidy levels.

Bayesian island biogeography in a continental setting: the Rand Flora case

We here explore the use of a Bayesian approach to island biogeography for disentangling the evolutionary origins of a continental-scale floristic pattern, the enigmatic ‘Rand Flora’. The existence of disjunct distributions across many plant lineages between Macaronesia–northwest Africa, Horn of Africa–southern Arabia and east–south Africa has long intrigued botanists, but only now can we start analysing it within a statistical framework. Phylogenetic and distributional data from 13 plant lineages exhibiting this disjunct distribution were analysed to estimate area carrying capacities and historical rates of biotic exchange between areas. The results indicate that there has been little exchange between southern Africa and the northern African region, and that this exchange occurred via east Africa. Northwest Africa–Macaronesia shows the smallest carrying capacity but highest dispersal rate with other regions, suggesting that its flora was built up by immigration of lineages, probably from the Mediterranean region and western Asia. In contrast, southern Africa shows the highest carrying capacity and lowest dispersal rate, suggesting a flora formed by in situ diversification. We discuss further improvements of the method for addressing more complex hypotheses, such as asymmetric dispersal between regions or repeated cyclical events.

Phylogeny and Historical Biogeography of Geraniaceae in Relation to Climate Changes and Pollination Ecology

Abstract Chloroplast (trnL−F and rbcL) sequences were used to reconstruct the phylogeny of Geraniaceae and Hypseocharitaceae. According to these data Hypseocharitaceae and Geraniaceae are monophyletic. Pelargonium and Monsonia are sisters to the largest clade of Geraniaceae, formed by Geranium, Erodium and California. According to molecular dating and dispersal-vicariance analysis, the split of the stem branches of Geraniaceae probably occurred during the Oligocene, in southern Africa or in southern Africa plus the Mediterranean area. However, their diversification occurred during the Miocene, coinciding with the beginning of major aridification events in their distribution areas. An ancestor of the largest clade of Geraniaceae (Geranium, Erodium, and California) colonised a number of habitats in the northern hemisphere and in South American mountain ranges. In summary, the evolution of the Geraniaceae is marked by the dispersal of ancestors from Southern Africa to cold, temperate and often disturbed habitats in the rest of world, where only generalist pollination and facultative autogamy could ensure sufficient seed production and survival.

Diversity and distribution of ferns in sub-Saharan Africa, Madagascar and some islands of the South Atlantic

Abstract Aim  This paper reports the diversity and endemism patterns of African ferns, and explores the potential role of diversity refuges and environmental and historical factors in the shaping of these patterns. Material and locations  The extant fern taxa occupying Africa south of the Sahara, Madagascar and some islands of the South Atlantic. Methods  The number of taxa in each area or operational geographical unit (OGU) was scored, and the correlation between this number and physical and climatic variables analysed by standard pairwise and stepwise multiple regression analysis (SPR and SMR). The effects of biological factors such as dispersal capacity, reproductive biology, genetic features and certain physiological adaptations were evaluated by comparing the number of species in each OGU. Floral affinities among OGUs were analysed using non‐metric multi‐dimensional scaling (NMS) and parsimonic analysis of dispersion (PAD), and compared with β‐turnover and inter‐OGU distances. Results  OGU area, elevation and the distance between refuges determined the composition of local floras, but only greater OGU area and the existence of higher maximum elevations increased species richness. The distance between refuges also affected the number of endemic species, especially on islands. The biological features studied only slightly influenced fern distribution. The main climatic predictor of species number was humidity. SPR and SMR revealed three main groups of ferns with different ecological trends. NMS and PAD analyses separated the four areas of highest diversity in Africa, three of which are inhabited by ferns with distinct ecological requirements. The fourth area was Madagascar, which shows an accumulation of endemic and relict diversity that is not easy to explain. Main conclusions  The distribution of ferns in Africa has been influenced by refuges. These probably allowed many species to recolonize the neighbouring areas after the extinctions of the Pleistocene. Three major components were detected in the African flora: Guinea‐Congolian thermophilous, cold‐tolerant Afro‐montane, and Southern drought‐tolerant elements. These are related to the three main refuge areas, i.e. the Gulf of Guinea area, the eastern tropical region, and the Cape region. Endemicity in ferns was found to be lower than that of seed plants due to the higher dispersability of fern spores. The distance between OGUs seems to be the main predictor of the number of endemic fern species these areas contain.

The Genus Sorbus (Maloideae, Rosaceae) in Europe and in North Africa: Morphological Analysis and Systematics

A multivariate morphometric study of the genus Sorbus, from 127 herbarium specimens in Europe and North Africa and nine populations in Spain, shows that twelve species may be easily recognized in the area: S. torminalis, S. latifolia, S. semiincisa, S. aria, S. intermedia, S. minima, S. sudetica, S. chamaemespilus, S. aucuparia, S. hybrida, S. meinichii and S. domestica. Twenty-seven described taxa were very difficult to dis- criminate using the standard morphological and statistical methods. Some useful characters are: petiole length, leaf-nerve angle, leaf-tooth length, bud and seed shape, fruit structure, presence of tanniferous-cell groups, large sclereids and starch grains in the fruit. The fruit of S. torminalis has a multi-layered cuticle, not previously reported in Sorbus. We detected tetraploidy in S. torminalis and two other sexuals: S. aria and S. chamaemespilus. The geography and the population structure of taxa in S and C Europe does not permit their reproductive isolation; hence a large number of intermediate forms occur. Considering the difficulty in assess- ing the reproductive isolation of microspecies and in differentiating them morphologically, the use of a taxo- nomic species concept is proposed in Sorbus. Finally, a key to the species is provided.

The uneven phylogeny and biogeography of Erodium (Geraniaceae): radiations in the Mediterranean and recent recurrent intercontinental colonization.

BACKGROUND AND AIMS The genus Erodium is a common feature of Mediterranean-type climates throughout the world, but the Mediterranean Basin has significantly higher diversity than other areas. The aim here is to reveal the biogeographical history of the genus and the causes behind the evolution of the uneven distribution. METHODS Seventy-eight new nrITS sequences were incorporated with existing plastid data to explore the phylogenetic relationships and biogeography of Erodium using several reconstruction methods. Divergence times for major clades were calculated and contrasted with other previously published information. Furthermore, topological and temporal diversification rate shift analyses were employed using these data. KEY RESULTS Phylogenetic relationships among species are widely congruent with previous plastid reconstructions, which refute the classical taxonomical classification. Biogeographical reconstructions point to Asia as the ancestral area of Erodium, arising approx. 18 MYA. Four incidences of intercontinental dispersal from the Mediterranean Basin to similar climates are demonstrated. Increases in diversification were present in two independent Erodium lineages concurrently. Two bursts of diversification (3 MYA and 0·69 MYA) were detected only in the Mediterranean flora. CONCLUSIONS Two lineages diverged early in the evolution of the genus Erodium: (1) subgenus Erodium plus subgenus Barbata subsection Absinthioidea and (2) the remainder of subgenus Barbata. Dispersal across major water bodies, although uncommon, has had a major influence on the distribution of this genus and is likely to have played as significant role as in other, more easily dispersed, genera. Establishment of Mediterranean climates has facilitated the spread of the genus and been crucial in its diversification. Two, independent, rapid radiations in response to the onset of drought and glacial climate change indicate putative adaptive radiations in the genus.

Diatom and desmid relationships with the environment in mountain lakes and mires of NW Spain

The mountain ranges in NW Spain have a large number of little known wetlands. We report the results of a study carried out on a group of 77 small lakes and mires in the Sierra Segundera and Cordillera Cantábrica. The main physical and chemical variables, and phytoplankton communities from littoral samples were studied. Cantabrian wetlands showed greater variability in all environmental variables measured as well as higher values in those related to mineralisation than the Segundera ones. Many of these ecosystems were oligotrophic and showed a high species richness. Desmids and diatoms were the two most abundant groups, both in the species number and in biovolume. Desmids were the most numerous group in taxa in Sierra Segundera, whereas diatoms were in Cordillera Cantábrica. Differences in species composition of algae communities between both mountain ecosystems were studied. Canonical Correspondence Analysis (CCA) was carried out on diatom and desmid flora composition. This analysis showed that alkalinity was the most important parameter in diatom distribution and pH the most important one in that of desmids.

Ancient vicariance and climate-driven extinction explain continental-wide disjunctions in Africa: the case of the Rand Flora genus Canarina (Campanulaceae)

Transoceanic distributions have attracted the interest of scientists for centuries. Less attention has been paid to the evolutionary origins of ‘continent‐wide’ disjunctions, in which related taxa are distributed across isolated regions within the same continent. A prime example is the ‘Rand Flora’ pattern, which shows sister taxa disjunctly distributed in the continental margins of Africa. Here, we explore the evolutionary origins of this pattern using the genus Canarina, with three species: C. canariensis, associated with the Canarian laurisilva, and C. eminii and C. abyssinica, endemic to the Afromontane region in East Africa, as case study. We infer phylogenetic relationships, divergence times and the history of migration events within Canarina using Bayesian inference on a large sample of chloroplast and nuclear sequences. Ecological niche modelling was employed to infer the climatic niche of Canarina through time. Dating was performed with a novel nested approach to solve the problem of using deep time calibration points within a molecular dataset comprising both above‐species and population‐level sampling. Results show C. abyssinica as sister to a clade formed by disjunct C. eminii and C. canariensis. Miocene divergences were inferred among species, whereas infraspecific divergences fell within the Pleistocene–Holocene periods. Although C. eminii and C. canariensis showed a strong genetic geographic structure, among‐population divergences were older in the former than in the latter. Our results suggest that Canarina originated in East Africa and later migrated across North Africa, with vicariance and aridification‐driven extinction explaining the 7000 km/7 million year divergence between the Canarian and East African endemics.

Palaeo-islands as refugia and sources of genetic diversity within volcanic archipelagos: the case of the widespread endemic Canarina canariensis (Campanulaceae)

Geographical isolation by oceanic barriers and climatic stability has been postulated as some of the main factors driving diversification within volcanic archipelagos. However, few studies have focused on the effect that catastrophic volcanic events have had on patterns of within‐island differentiation in geological time. This study employed data from the chloroplast (cpDNA haplotypes) and the nuclear (AFLPs) genomes to examine the patterns of genetic variation in Canarina canariensis, an iconic plant species associated with the endemic laurel forest of the Canary Islands. We found a strong geographical population structure, with a first divergence around 0.8 Ma that has Tenerife as its central axis and divides Canarian populations into eastern and western clades. Genetic diversity was greatest in the geologically stable ‘palaeo‐islands’ of Anaga, Teno and Roque del Conde; these areas were also inferred as the ancestral location of migrant alleles towards other disturbed areas within Tenerife or the nearby islands using a Bayesian approach to phylogeographical clustering. Oceanic barriers, in contrast, appear to have played a lesser role in structuring genetic variation, with intra‐island levels of genetic diversity larger than those between‐islands. We argue that volcanic eruptions and landslides after the merging of the palaeo‐islands 3.5 Ma played key roles in generating genetic boundaries within Tenerife, with the palaeo‐islands acting as refugia against extinction, and as cradles and sources of genetic diversity to other areas within the archipelago.

Phylogenetic Relationships and Evolution in Erodium (Geraniaceae) based on trnL-trnF Sequences

Abstract Phylogenetic reconstructions in the Mediterranean genus Erodium are for the first time performed using two matrices: one with 96 trnL-trnF sequences from Erodium (90 accessions plus four outgroups) and the other with 72 trnL-trnF sequences plus 23 morphological characters (66 species of a total of 74), using Maximum Parsimony (MP) and Bayesian Inference (BI). An association between reproductive properties (high selfing rates, flower asymmetry, insect-attraction structures), life form, and breeding system distributed in different lineages suggests multiple shifts from allogamy to autogamy in the course of evolution, whereas dioecy has occurred only once. The phylogenetic analyses revealed a remarkable capability for dispersal in Erodium because closely related species occur in different continents. Major lineages containing sublineages of species also from several continents lead us to interpret ancient dispersal activity. Establishment of Mediterranean-like climates in most continents may have been crucial in the evolution of Erodium, as manifested by occurrence of species of the Mediterranean floristic region in the four major lineages. The body of knowledge accumulated from molecular phylogenetics and morphology lead us to conclude that the Mediterranean region harbors the major center of diversity of Erodium, where active radiation in dry, disturbed environments, is still operating.