The long journey of Orthotrichum shevockii (Orthotrichaceae, Bryopsida): From California to Macaronesia.

Biogeography, systematics and taxonomy are complementary scientific disciplines. To understand a species' origin, migration routes, distribution and evolutionary history, it is first necessary to establish its taxonomic boundaries. Here, we use an integrative approach that takes advantage of complementary disciplines to resolve an intriguing scientific question. Populations of an unknown moss found in the Canary Islands (Tenerife Island) resembled two different Californian endemic species: Orthotrichum shevockii and O. kellmanii. To determine whether this moss belongs to either of these species and, if so, to explain its presence on this distant oceanic island, we combined the evaluation of morphological qualitative characters, statistical morphometric analyses of quantitative traits, and molecular phylogenetic inferences. Our results suggest that the two Californian mosses are conspecific, and that the Canarian populations belong to this putative species, with only one taxon thus involved. Orthotrichum shevockii (the priority name) is therefore recognized as a morphologically variable species that exhibits a transcontinental disjunction between western North America and the Canary Islands. Within its distribution range, the area of occupancy is limited, a notable feature among bryophytes at the intraspecific level. To explain this disjunction, divergence time and ancestral area estimation analyses are carried out and further support the hypothesis of a long-distance dispersal event from California to Tenerife Island.

Climate threat on the Macaronesian endemic bryophyte flora.

Oceanic islands are of fundamental importance for the conservation of biodiversity because they exhibit high endemism rates coupled with fast extinction rates. Nowhere in Europe is this pattern more conspicuous than in the Macaronesian biogeographic region. A large network of protected areas within the region has been developed, but the question of whether these areas will still be climatically suitable for the globally threatened endemic element in the coming decades remains open. Here, we make predictions on the fate of the Macaronesian endemic bryophyte flora in the context of ongoing climate change. The potential distribution of 35 Macaronesian endemic bryophyte species was assessed under present and future climate conditions using an ensemble modelling approach. Projections of the models under different climate change scenarios predicted an average decrease of suitable areas of 62-87% per species and a significant elevational increase by 2070, so that even the commonest species were predicted to fit either the Vulnerable or Endangered IUCN categories. Complete extinctions were foreseen for six of the studied Macaronesian endemic species. Given the uncertainty regarding the capacity of endemic species to track areas of suitable climate within and outside the islands, active management associated to an effective monitoring program is suggested.

Approximate Bayesian Computation Reveals the Crucial Role of Oceanic Islands for the Assembly of Continental Biodiversity.

The perceived low levels of genetic diversity, poor interspecific competitive and defensive ability, and loss of dispersal capacities of insular lineages have driven the view that oceanic islands are evolutionary dead ends. Focusing on the Atlantic bryophyte flora distributed across the archipelagos of the Azores, Madeira, the Canary Islands, Western Europe, and northwestern Africa, we used an integrative approach with species distribution modeling and population genetic analyses based on approximate Bayesian computation to determine whether this view applies to organisms with inherent high dispersal capacities. Genetic diversity was found to be higher in island than in continental populations, contributing to mounting evidence that, contrary to theoretical expectations, island populations are not necessarily genetically depauperate. Patterns of genetic variation among island and continental populations consistently fitted those simulated under a scenario of de novo foundation of continental populations from insular ancestors better than those expected if islands would represent a sink or a refugium of continental biodiversity. We, suggest that the northeastern Atlantic archipelagos have played a key role as a stepping stone for transoceanic migrants. Our results challenge the traditional notion that oceanic islands are the end of the colonization road and illustrate the significant role of oceanic islands as reservoirs of novel biodiversity for the assembly of continental floras.

Effect of forest clear-cutting on subtropical bryophyte communities in waterfalls, on dripping walls, and along streams.

Forested freshwater ecosystems worldwide are threatened by a number of anthropogenic disturbances, such as water pollution and canalization. Transient or permanent deforestation can also be a serious threat to organisms in forested watersheds, but its effects on different types of freshwater systems has been little studied. We investigated lotic bryophyte communities on rock and soil in subtropical cloud laurel forests on La Gomera Island in the Canary Islands, Spain, and asked whether the response to forest clear-cutting varied among the communities associated with dripping walls, streams, and waterfalls. We compared three successional forest stages: ancient forests (> 250 years), young forests (20-50 years after clear-cutting), and open stands (5-15 years after clear-cutting). In each of 56 study sites we sampled general vegetation and substrate data in a 0.01-ha plot and took composition data of bryophyte species in 3 + 3 subplots of 1 x 1 m. The general pattern of decline in species richness and change in species composition after forest clear-cutting was stronger for streamside assemblages compared to assemblages on dripping walls and in waterfalls. The change in species numbers on rocks was larger than that on soils, because a guild of species growing on soil (but not on rocks) were favored by disturbance and thus increased in the disturbed sites. Most of the sensitive species could be classified as typical laurel forest species. Mosses were generally more tolerant to forest clear-cutting than were liverworts. We suggest that streamsides are more sensitive to disturbance than waterfalls and dripping walls because of a larger variation in microclimate before than after clear-cutting and because they are more easily invaded by early-successional species (both bryophytes and highly competitive vascular plants). We propose that special care should be taken along small streams within disturbed watersheds if bryophyte assemblages and threatened species should be protected. The susceptibility to anthropogenic pressures is probably rather high in ecosystems that do not regularly experience large-scale stand-replacing disturbances, especially on oceanic islands because of isolation and a small total habitat area for focal organisms.

And if Engler was not completely wrong? Evidence for multiple evolutionary origins in the moss flora of Macaronesia.

The Macaronesian endemic flora has traditionally been interpreted as a relict of a subtropical element that spanned across Europe in the Tertiary. This hypothesis is revisited in the moss subfamily Helicodontioideae based on molecular divergence estimates derived from two independent calibration techniques either employing fossil evidence or using an Monte Carlo Markov Chain (MCMC) to sample absolute rates of nucleotide substitution from a prior distribution encompassing a wide range of rates documented across land plants. Both analyses suggest that the monotypic Madeiran endemic genus Hedenasiastrum diverged of other Helicodontioideae about 40 million years, that is, well before Macaronesian archipelagos actually emerged, in agreement with the relict hypothesis. Hedenasiastrum is characterized by a plesiomorphic morphology, which is suggestive of a complete morphological stasis over 40 million years. Macaronesian endemic Rhynchostegiella species, whose polyphyletic origin involves multiple colonization events, evolved much more recently, and yet accumulated many more morphological novelties than H. percurrens. The Macaronesian moss flora thus appears as a complex mix of ancient relicts and more recently dispersed, fast-evolving taxa.