Alain Vanderpoorten

Geography and host biogeography matter for understanding the phylogeography of a parasite.

The co-evolution between hosts and parasites has long been recognized as a fundamental driver of macro-evolutionary patterns of diversification. The effect of co-differentiation on parasite diversification is, however, often confounded by underlying geographic patterns of host distribution. In order to disentangle the confounding effects of allopatric versus host speciation, the mitochondrial cytochrome b (cyt b) gene was sequenced in seventy individuals of the parasitic nematode genus Heligmosomoides sampled in the six Apodemus mice species common in the western Palearctic region. The nuclear internal transcribed spacers (ITS) 1 and 2 were also sequenced in fifteen parasites to confirm the mitochondrial data. All lineages differentiated according to a geographic pattern and independently from the sampled host species. This suggests that host speciation did not involve concurrent parasite speciation. However, the geographic distribution range of some parasite lineages mirrors that of A. sylvaticus lineages in SW Europe, and that of A. flavicollis lineages in the Balkans and in the Middle East. Thus, regional co-differentiation likely occurred between the parasite and the two sister Apodemus hosts in different parts of their distribution range. We suggest that differences in regional abundances of A. sylvaticus and A. flavicollis are responsible for generating this pattern of regional co-differentiation. This study highlights the importance of integrating both geography and biogeographic information from potential hosts to better understand their parasite phylogeography.

ORIGIN AND EVOLUTION OF THE NORTHERN HEMISPHERE DISJUNCTION IN THE MOSS GENUS HOMALOTHECIUM (BRACHYTHECIACEAE)

Competing hypotheses that rely either on a stepping-stone dispersal via the North Atlantic or the Bering land bridges, or more recent transoceanic dispersal, have been proposed to explain the disjunct distribution of Mediterranean flora in southern Europe and western North America. These hypotheses were tested with molecular dating using a phylogeny of the moss genus Homalothecium based on ITS, atpB-rbcL, and rpl16 sequence data. The monophyly of two main lineages in Western Palearctic (Europe, central Asia and north Africa) and North America is consistent with the ancient vicariance hypothesis. The monophyly of Madeiran H. sericeum accessions supports the recognition of the Macaronesian endemic H. mandonii. A range of absolute rates of molecular evolution documented in land plants was used as probabilistic calibration prior by a Bayesian inference implementing a relaxed-clock model to derive ages for the nodes of interest. Our age estimates for the divergence of the American and Western Palearctic Homalothecium clade (5.7 Ma, IC 3.52-8.26) and the origin of H. mandonii (2.52 Myr IC 0.86-8.25) are not compatible with the ancient vicariance hypothesis. Age estimates suggests that species distributions result from rare instances of dispersal and subsequent sympatric diversification. The calibrated phylogeny indicates that Homalothecium has undergone a fast radiation during the last 4 Myr, which is consistent with the low levels of morphological divergence among sibling species.

Does Macaronesia exist? Conflicting signal in the bryophyte and pteridophyte floras

Macaronesia, which includes five mid-Atlantic archipelagos (Azores, Madeira, Selvagems, Canaries, and Cape Verdes), has been traditionally recognized as a distinct biogeographic unit whose circumscription has been intimately associated with the hypothesis that the flora is a relict of a formerly broadly distributed subtropical Tertiary flora. The concept of Macaronesia is revisited here using parsimony and Bayesian analyses of floristic data sets for the moss, liverwort, and pteridophyte floras. All analyses reject the monophyly of Macaronesia s.l., resolving the Cape Verdes with tropical Africa. Of the other Macaronesian archipelagos, the liverwort and pteridophyte analyses support, or could not reject, an Azorean-Madeiran-Canarian clade (hereafter Macaronesia s.s.), but the moss analysis resolves the Canary Islands as sister to North Africa, thus rejecting the concept of Macaronesia s.s. for this group. Dynamic interchange of taxa with neighboring continental areas rather than relictualism best explains the relationships of the Cape Verde cryptogamic flora and the Canary Island moss flora. In contrast, relictualism is consistent with a monophyletic Macaronesia s.s. for liverworts and pteridophytes. However, from the limited information available on relationships of endemic cryptogams, this explanation alone may be unsatisfactory. Spatially congruent patterns may, in fact, conceal a complex mixture of relictual distributions and more recent speciation and dispersal events.

Extant diversity of bryophytes emerged from successive post-Mesozoic diversification bursts

Unraveling the macroevolutionary history of bryophytes, which arose soon after the origin of land plants but exhibit substantially lower species richness than the more recently derived angiosperms, has been challenged by the scarce fossil record. Here we demonstrate that overall estimates of net species diversification are approximately half those reported in ferns and ∼30% those described for angiosperms. Nevertheless, statistical rate analyses on time-calibrated large-scale phylogenies reveal that mosses and liverworts underwent bursts of diversification since the mid-Mesozoic. The diversification rates further increase in specific lineages towards the Cenozoic to reach, in the most recently derived lineages, values that are comparable to those reported in angiosperms. This suggests that low diversification rates do not fully account for current patterns of bryophyte species richness, and we hypothesize that, as in gymnosperms, the low extant bryophyte species richness also results from massive extinctions.

RANGE DISJUNCTIONS, SPECIATION, AND MORPHOLOGICAL TRANSFORMATION RATES IN THE LIVERWORT GENUS LEPTOSCYPHUS

Bryophytes and angiosperms exhibit similar intercontinental disjunct distributions that have traditionally been explained by continental drift. Such disjunct distributions are, however, typically observed at the species level in bryophytes, whereas they occur at much higher taxonomic level in angiosperms. The corollary of this observation is that morphological evolution in bryophytes is exceedingly slow. These hypotheses can now be explicitly tested with the advent of molecular dating. In this article, we show that the trans-Atlantic disjunctions observed in the mostly tropical liverwort genus Leptoscyphus date back to 5.5 Myr, thus largely postdating the opening of the South Atlantic. The temporal calibration of the phylogeny allows us to estimate for the first time the absolute timing of morphological evolution in bryophytes. The time frame necessary for shifts to occur between character states was estimated on average at ca. 4.05 ± 1.86 Myr. As opposed to the traditional view that bryophyte evolution has been triggered by episodic shifts in habitat conditions, our analyses furthermore suggest that morphological and molecular divergence gradually accumulated in the genus, which contrasts with the rapid diversification documented in some tropical trees.

Oceanic islands are not sinks of biodiversity in spore-producing plants

Islands have traditionally been considered as migratory and evolutionary dead ends for two main reasons: island colonizers are typically assumed to lose their dispersal power, and continental back colonization has been regarded as unlikely because of niche preemption. The hypothesis that islands might actually represent dynamic refugia and migratory stepping stones for species that are effective dispersers, and in particular, for spore-producing plants, is formally tested here, using the archipelagos of the Azores, Canary Islands, and Madeira, as a model. Population genetic analyses based on nuclear microsatellite variation indicate that dispersal ability of the moss Platyhypnidium riparioides does not decrease in the island setting. The analyses further show that, unlike island populations, mainland (southwestern Europe and North Africa) populations underwent a severe bottleneck during the last glacial maximum (LGM). Our results thus refute the traditional view of islands as the end of the colonization road and point to a different perception of North Atlantic archipelagos as major sources of biodiversity for the postglacial recolonization of Europe by spore-producing plants.

When morphology and molecules tell us different stories: a case-in- point with Leptodon corsicus, a new and unique endemic moss species from Corsica

Abstract Leptodon corsicus (Neckeraceae) is described as the first endemic moss species from Corsica. It strikingly differs from the other species of the genus by the lack of a dense and pinnate to bipinnate mode of branching; about 10 times smaller shoots that do not inroll upon drying; the lack of paraphyllia; and few, occasional small pseudoparaphyllia. Due to its small size and several leaf characters, L. corsicus shares at first glance more similarities with Homalia webbiana and Neckera besseri than with Leptodon. Yet, phylogenetic analysis of chloroplast and nuclear DNA sequences unambiguously shows that L. corsicus is deeply nested within L. smithii. The numerous morphological characters that distinguish L. corsicus from L. smithii cannot be attributed to plasticity. Consequently, we interpret the phylogenetic position of L. corsicus as the result of a recent speciation process, involving mutations at one or a few coding loci or differences in gene expression, which have tremendous consequences for phenotypic appearance, and retention of ancestral polymorphism in the non-coding sequences used for phylogenetic reconstruction. Such an explanation might also apply to other species of mosses, which exhibit a striking morphology, and yet share identical non-coding sequences with the common species they derive from. The notion of species in mosses is discussed in this context.

The ghosts of Gondwana and Laurasia in modern liverwort distributions

Recent advances in phylogenetics and, in particular, molecular dating, indicate that transoceanic dispersal has played an important role in shaping plant and animal distributions, obscuring any effect of tectonic history. Taxonomic sampling in biogeographic studies is, however, systematically biased towards vertebrates and higher plants and the possibility remains that a much stronger signature of ancient vicariance might be evident among other organisms, particularly among basal land plants. Here, an explicit Bayesian model‐based approach was used to investigate global‐scale biogeographic patterns among liverwort genera and to determine whether the patterns identified are consistent with the expectations of vicariance or dispersal scenarios. The distribution of each genus was mapped onto the phylograms describing the floristic affinities among areas in order to define the synapomorphic transitions supporting the observed groupings. The probabilities of change in a branch were calculated by implementing the Markov model of BayesTraits. The consistent ambiguity in ancestral state reconstructions returned by the unconstrained, two‐rate model indicated that the overall signal in the data was weak, leading us to test the performance of competing, explicit models. The analyses resolved clades of geographic areas that are mostly consistent with the kingdoms traditionally identified for plants and animals, but with strikingly lower rates of endemism. The major split observed in the phylograms is into almost entirely Laurasian and Gondwanan clades. Other patterns recovered by the analyses, including Wallaces line and the South Atlantic Disjunction, have also traditionally been interpreted in terms of vicariance. These observations contrast with the idea that, in spore‐dispersed organisms like bryophytes and pteridophytes, dispersal obscures evidence of vicariance. However, some discrepancies between the liverwort trees and expectations from a continental drift scenario were observed, such as the sister‐group relationship of the Australian and New Zealand floras, which is supported by the co‐occurrence of many genera, often endemic to these two areas. Together with an interpretation of the results within a phylogenetic context, our analyses suggest that patterns, which are at first sight consistent with an ancient vicariance hypothesis, may, in fact, conceal a complex mixture of relictual distributions and more recent, asymmetrical dispersal events. Our results provide a framework for testing specific evolutionary hypotheses concerning the extremely low levels of endemism in bryophytes and in particular, the significance of dispersal and cryptic diversification.

Mapping Uncertainty and Phylogenetic Uncertainty in Ancestral Character State Reconstruction: An Example in the Moss Genus Brachytheciastrum

The evolution of species traits along a phylogeny can be examined through an increasing number of possible, but not necessarily complementary, approaches. In this paper, we assess whether deriving ancestral states of discrete morphological characters from a model whose parameters are (i) optimized by ML on a most likely tree; (II) optimized by ML onto each of a Bayesian sample of trees; and (III) sampled by a MCMC visiting the space of a Bayesian sample of trees affects the reconstruction of ancestral states in the moss genus Brachytheciastrum. In the first two methods, the choice of a single- or two-rate model and of a genetic distance (wherein branch lengths are used to determine the probabilities of change) or speciational (wherein changes are only driven by speciation events) model based upon a likelihood-ratio test strongly depended on the sampled trees. Despite these differences in model selection, reconstructions of ancestral character states were strongly correlated to each others across nodes, often at r > 0.9, for all the characters. The Bayesian approach of ancestral character state reconstruction offers, however, a series of advantages over the single-tree approach or the ML model optimization on a Bayesian sample of trees because it does not involve restricting model parameters prior to reconstructing ancestral states, but rather allows a range of model parameters and ancestral character states to be sampled according to their posterior probabilities. From the distribution of the latter, conclusions on trait evolution can be made in a more satisfactorily way than when a substantial part of the uncertainty of the results is obscured by the focus on a single set of model parameters and associated ancestral states. The reconstructions of ancestral character states in Brachytheciastrum reveal rampant parallel morphological evolution. Most species previously described based on phenetic grounds are thus resolved of polyphyletic origin. Species polyphylly has been increasingly reported among mosses, raising severe reservations regarding current species definition.

New national and regional bryophyte records, 26

26 L T Ellis, H Bednarek-Ochyra, R Ochyra, Silvia Calvo Aranda, Maria T Colotti, Maria M Schiavone, Michail V Dulin, P Erzberger, Tulay Ezer, Recep Kara, Rosalina Gabriel, Lars Hedenas, David T Holyoak, P Odor, B Papp, M Sabovljevic, R D Seppelt, V R Smith, Andre Sotiaux, E Szurdoki, Alain Vanderpoorten, J van Rooy, J Żarnowiec Department of Botany, The Natural History Museum, UK, Laboratory of Bryology, Institute of Botany, Polish Academy of Sciences, Krakow, Poland, 3 Departamento de Biodiversidad y Biologia Evolutiva, Museo Nacional de Ciencias Naturales, Madrid, Spain, Facultad de Ciencias Naturales e IML, San Miguel de Tucuman, Argentina, Institute of Biology Komi Science Centre UB RAS, Syktyvkar, Komi Republic, Russia, Belziger Str. 37, D-10823 Berlin, Germany, Faculty of Science and Arts, Department of Biology, Nigde University, Turkey, Departamento de Ciencias Agrarias, Universidade dos Acores, Angra do Heroismo, Portugal, Department of Cryptogamic Botany, Swedish Museum of Natural History, Stockholm, Sweden, Quinta da Cachopa, Barcoila, Cabecudo, Portugal, Department of Plant Taxonomy and Ecology, Lorand Eotvos University, Budapest, Hungary, Botanical Department, Hungarian Natural History Museum, Budapest, Hungary, Institute of Botany and Botanical Garden, Faculty of Biology, University of Belgrade, Serbia, Australian Antarctic Division, Channel Highway, Kingston, Tasmania, Australia, Department of Botany, University of Stellenbosch, Matieland, South Africa, National Botanic Garden of Belgium, Domein van Bouchout, Meise, Belgium, Hungarian Natural History Museum, Budapest, Hungary, University of Liege, Institute of Botany, Belgium, National Herbarium, South African National Biodiversity Institute, Pretoria, South Africa, Department of Ecology and Nature Conservation, Institute of Environmental Protection and Engineering, University of Bielsko-Biala, Poland