Genetic legacies of mega-landslides: Cycles of isolation and contact across flank collapses in an oceanic island.

Catastrophic flank collapses are recognized as important drivers of insular biodiversity dynamics, through the disruption of species ranges and subsequent allopatric divergence. However, little empirical data supports this conjecture, with their evolutionary consequences remaining poorly understood. Using genome-wide data within a population genomics and phylogenomics framework, we evaluate how mega-landslides have impacted evolutionary and demographic history within a species complex of weevils (Curculionidae) within the Canary Island of Tenerife. We reveal a complex genomic landscape, within which individuals of single ancestry were sampled in areas characterized by long-term geological stability, relative to the timing of flank collapses. In contrast, individuals of admixed ancestry were almost exclusively sampled within the boundaries of flank collapses. Estimated divergence times among ancestral populations aligned with the timings of mega-landslide events. Our results provide first evidence for a cyclical dynamic of range fragmentation and secondary contact across flank collapse landscapes, with support for a model where this dynamic is mediated by Quaternary climate oscillations. The context within which we reveal climate and topography to interact cyclically through time to shape the geographic structure of genetic variation, together with related recent work, highlights the importance of topoclimatic phenomena as an agent of diversification within insular invertebrates.

Semi-quantitative metabarcoding reveals how climate shapes arthropod community assembly along elevation gradients on Hawaii Island.

Spatial variation in climatic conditions along elevation gradients provides an important backdrop by which communities assemble and diversify. Lowland habitats tend to be connected through time, whereas highlands can be continuously or periodically isolated, conditions that have been hypothesized to promote high levels of species endemism. This tendency is expected to be accentuated among taxa that show niche conservatism within a given climatic envelope. While species distribution modeling approaches have allowed extensive exploration of niche conservatism among target taxa, a broad understanding of the phenomenon requires sampling of entire communities. Species-rich groups such as arthropods are ideal case studies for understanding ecological and biodiversity dynamics along elevational gradients given their important functional role in many ecosystems, but community-level studies have been limited due to their tremendous diversity. Here, we develop a novel semi-quantitative metabarcoding approach that combines specimen counts and size-sorting to characterize arthropod community-level diversity patterns along elevational transects on two different volcanoes of the island of Hawai'i. We found that arthropod communities between the two transects became increasingly distinct compositionally at higher elevations. Resistance surface approaches suggest that climatic differences between sampling localities are an important driver in shaping beta-diversity patterns, though the relative importance of climate varies across taxonomic groups. Nevertheless, the climatic niche position of OTUs between transects was highly correlated, suggesting that climatic filters shape the colonization between adjacent volcanoes. Taken together, our results highlight climatic niche conservatism as an important factor shaping ecological assembly along elevational gradients and suggest topographic complexity as an important driver of diversification.

Influence of polyamide composite casings with silver-zinc crystals on the quality of beef and chicken sausages during their storage.

The influence of polyamide 6 composite casings with silver-zinc crystals powder on some of the physicochemicalphysical-chemical, microbiological and sensory indicators of beef and chicken sausages during their storage was evaluated. Beef and chicken sausages were elaborated by using the conventional technology for sausage meat thin pasta; in each case, it was maintained a control batch to compare changes during the storage (4 and 12 °C, 75%-85% RH). To estimate the shelf life was considered sensory evaluation as a criterion for rejection. The results were processed as failure incomplete data via the Weibull distribution and it was admitted 5% of deteriorated units. It did not find a significant effect (p ≤ 0.05) due to the addition of silver-zinc crystals on the values of pH, aw, color, texture and sensory attributes of sausages, but did influence TBARS results, with lower values compared to control products. It reduced the counts of aerobic mesophilic microorganisms and lactic acid bacteria during the storage. The shelf life of chicken sausage was not affected at any storage temperature; while for the beef sausage stored at 4 °C, its shelf life increased in 9 days, although at 12 °C did not exist difference among treatments.

Climate drives community-wide divergence within species over a limited spatial scale: evidence from an oceanic island.

Geographic isolation substantially contributes to species endemism on oceanic islands when speciation involves the colonisation of a new island. However, less is understood about the drivers of speciation within islands. What is lacking is a general understanding of the geographic scale of gene flow limitation within islands, and thus the spatial scale and drivers of geographical speciation within insular contexts. Using a community of beetle species, we show that when dispersal ability and climate tolerance are restricted, microclimatic variation over distances of only a few kilometres can maintain strong geographic isolation extending back several millions of years. Further to this, we demonstrate congruent diversification with gene flow across species, mediated by Quaternary climate oscillations that have facilitated a dynamic of isolation and secondary contact. The unprecedented scale of parallel species responses to a common environmental driver for evolutionary change has profound consequences for understanding past and future species responses to climate variation.

Quantification of complex modular architecture in plants.

Morphometrics, the assignment of quantities to biological shapes, is a powerful tool to address taxonomic, evolutionary, functional and developmental questions. We propose a novel method for shape quantification of complex modular architecture in thalloid plants, whose extremely reduced morphologies, combined with the lack of a formal framework for thallus description, have long rendered taxonomic and evolutionary studies extremely challenging. Using graph theory, thalli are described as hierarchical series of nodes and edges, allowing for accurate, homologous and repeatable measurements of widths, lengths and angles. The computer program MorphoSnake was developed to extract the skeleton and contours of a thallus and automatically acquire, at each level of organization, width, length, angle and sinuosity measurements. Through the quantification of leaf architecture in Hymenophyllum ferns (Polypodiopsida) and a fully worked example of integrative taxonomy in the taxonomically challenging thalloid liverwort genus Riccardia, we show that MorphoSnake is applicable to all ramified plants. This new possibility of acquiring large numbers of quantitative traits in plants with complex modular architectures opens new perspectives of applications, from the development of rapid species identification tools to evolutionary analyses of adaptive plasticity.

Evolutionary origin of the latitudinal diversity gradient in liverworts.

A latitudinal diversity gradient towards the tropics appears as one most recurrent patterns in ecology, but the mechanisms underlying this pattern remain an area of controversy. In angiosperms, the tropical conservatism hypothesis proposes that most groups originated in the tropics and are adapted to a tropical climatic regime, and that relatively few species have evolved physiological adaptations to cold, dry or unpredictable climates. This mechanism is, however, unlikely to apply across land plants, and in particular, to liverworts, a group of about 7500 species, whose ability to withstand cold much better than their tracheophyte counterparts is at odds with the tropical conservatism hypothesis. Molecular dating, diversification rate analyses and ancestral area reconstructions were employed to explore the evolutionary mechanisms that account for the latitudinal diversity gradient in liverworts. As opposed to angiosperms, tropical liverwort genera are not older than their extra-tropical counterparts (median stem age of tropical and extra-tropical liverwort genera of 24.35 ±â€¯39.65 Ma and 39.57 ±â€¯49.07 Ma, respectively), weakening the 'time for speciation hypothesis'. Models of ancestral area reconstructions with equal migration rates between tropical and extra-tropical regions outperformed models with asymmetrical migration rates in either direction. The symmetry and intensity of migrations between tropical and extra-tropical regions suggested by the lack of resolution in ancestral area reconstructions towards the deepest nodes are at odds with the tropical niche conservatism hypothesis. In turn, tropical genera exhibited significantly higher net diversification rates than extra-tropical ones, suggesting that the observed latitudinal diversity gradient results from either higher extinction rates in extra-tropical lineages or higher speciation rates in the tropics. We discuss a series of experiments to help deciphering the underlying evolutionary mechanisms.

Range size heritability and diversification patterns in the liverwort genus Radula.

Why some species exhibit larger geographical ranges than others, and to what extent does variation in range size affect diversification rates, remains a fundamental, but largely unanswered question in ecology and evolution. Here, we implement phylogenetic comparative analyses and ancestral area estimations in Radula, a liverwort genus of Cretaceous origin, to investigate the mechanisms that explain differences in geographical range size and diversification rates among lineages. Range size was phylogenetically constrained in the two sub-genera characterized by their almost complete Australasian and Neotropical endemicity, respectively. The congruence between the divergence time of these lineages and continental split suggests that plate tectonics could have played a major role in their present distribution, suggesting that a strong imprint of vicariance can still be found in extant distribution patterns in these highly mobile organisms. Amentuloradula, Volutoradula and Metaradula species did not appear to exhibit losses of dispersal capacities in terms of dispersal life-history traits, but evidence for significant phylogenetic signal in macroecological niche traits suggests that niche conservatism accounts for their restricted geographic ranges. Despite their greatly restricted distribution to Australasia and Neotropics respectively, Amentuloradula and Volutoradula did not exhibit significantly lower diversification rates than more widespread lineages, in contrast with the hypothesis that the probability of speciation increases with range size by promoting geographic isolation and increasing the rate at which novel habitats are encountered. We suggest that stochastic long-distance dispersal events may balance allele frequencies across large spatial scales, leading to low genetic structure among geographically distant areas or even continents, ultimately decreasing the diversification rates in highly mobile, widespread lineages.

Increased diversification rates follow shifts to bisexuality in liverworts.

Shifts in sexual systems are one of the key drivers of species diversification. In contrast to angiosperms, unisexuality prevails in bryophytes. Here, we test the hypotheses that bisexuality evolved from an ancestral unisexual condition and is a key innovation in liverworts. We investigate whether shifts in sexual systems influence diversification using hidden state speciation and extinction analysis (HiSSE). This new method compares the effects of the variable of interest to the best-fitting latent variable, yielding robust and conservative tests. We find that the transitions in sexual systems are significantly biased toward unisexuality, even though bisexuality is coupled with increased diversification. Sexual systems are strongly conserved deep within the liverwort tree but become much more labile toward the present. Bisexuality appears to be a key innovation in liverworts. Its effects on diversification are presumably mediated by the interplay of high fertilization rates, massive spore production and long-distance dispersal, which may separately or together have facilitated liverwort speciation, suppressed their extinction, or both. Importantly, shifts in liverwort sexual systems have the opposite effect when compared to angiosperms, leading to contrasting diversification patterns between the two groups. The high prevalence of unisexuality among liverworts suggests, however, a strong selection for sexual dimorphism.

High migration rates shape the postglacial history of amphi-Atlantic bryophytes.

Paleontological evidence and current patterns of angiosperm species richness suggest that European biota experienced more severe bottlenecks than North American ones during the last glacial maximum. How well this pattern fits other plant species is less clear. Bryophytes offer a unique opportunity to contrast the impact of the last glacial maximum in North America and Europe because about 60% of the European bryoflora is shared with North America. Here, we use population genetic analyses based on approximate Bayesian computation on eight amphi-Atlantic species to test the hypothesis that North American populations were less impacted by the last glacial maximum, exhibiting higher levels of genetic diversity than European ones and ultimately serving as a refugium for the postglacial recolonization of Europe. In contrast with this hypothesis, the best-fit demographic model involved similar patterns of population size contractions, comparable levels of genetic diversity and balanced migration rates between European and North American populations. Our results thus suggest that bryophytes have experienced comparable demographic glacial histories on both sides of the Atlantic. Although a weak, but significant genetic structure was systematically recovered between European and North American populations, evidence for migration from and towards both continents suggests that amphi-Atlantic bryophyte population may function as a metapopulation network. Reconstructing the biogeographic history of either North American or European bryophyte populations therefore requires a large, trans-Atlantic geographic framework.

Is the sword moss (Bryoxiphium) a preglacial Tertiary relict?

The disjunction of floras between East Asia, Southeast North America, West North America, and Southwest Eurasia has been interpreted in terms of the fragmentation of a once continuous mixed mesophytic forest that occurred throughout the Northern Hemisphere due to the climatic and geological changes during the late Tertiary. The sword moss, Bryoxiphium, exhibits a distribution that strikingly resembles that of the mesophytic forest elements such as Liriodendron and is considered as the only living member of an early Tertiary flora in Iceland. These hypotheses are tested here using molecular dating analyses and ancestral area estimations. The results suggest that the extant range of Bryoxiphium results from the fragmentation of a formerly wider range encompassing North America and Southeast Asia about 10 million years ago. The split of continental ancestral populations is too recent to match with a continental drift scenario but is spatially and temporally remarkably congruent with that observed in Tertiary angiosperm relict species. The timing of the colonization of Iceland from Macaronesian ancestors, about two million years ago, is, however, incompatible with the hypothesis that Bryoxiphium is the only living member of an early Tertiary flora of the island. Alaska was recurrently colonized from East Asia. The ability of Bryoxiphium to overcome large oceanic barriers is further evidenced by its occurrence on remote oceanic archipelagos. In particular, Madeira was colonized twice independently from American and East Asian ancestors, respectively. The striking range disjunction of Bryoxiphium is interpreted in terms of its mating system, as the taxon exhibits a very singular pattern of spatial segregation of the sexes.

What are the evolutionary mechanisms explaining the similar species richness patterns in tropical mosses? Insights from the phylogeny of the pantropical genus Pelekium.

As opposed to angiosperms, moss species richness is similar among tropical regions of the world, in line with the hypothesis that tropical bryophytes are extremely good dispersers. Here, we reconstructed the phylogeny of the pantropical moss genus Pelekium to test the hypothesis that high migration rates erase any difference in species richness among tropical regions. In contrast with this hypothesis, several species considered to have a pantropical range were resolved as a complex of species with a strong geographic structure. Consequently, a significant phylogeographical signal was found in the data, evidencing that cladogenetic diversification within regions takes place at a faster rate than intercontinental migration. The shape of the Pelekium phylogeny, along with the selection of a constant-rate model of diversification among species in the genus, suggests, however, that the cladogenetic speciation patterns observed in Pelekium are not comparable to some of the spectacular examples of tropical radiations reported in angiosperms. Rather, the results presented here point to the constant accumulation of diversity through time in Pelekium. This, combined with evidence for long-distance dispersal limitations in the genus, suggests that the similar patterns of species richness among tropical areas are better explained in terms of comparable rates of diversification across tropical regions than by the homogenization of species richness by recurrent migrations.

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.

Measuring spore settling velocity for an improved assessment of dispersal rates in mosses.

BACKGROUND AND AIMS: The settling velocity of diaspores is a key parameter for the measurement of dispersal ability in wind-dispersed plants and one of the most relevant parameters in explicit dispersal models, but remains largely undocumented in bryophytes. The settling velocities of moss spores were measured and it was determined whether settling velocities can be derived from spore diameter using Stokes' Law or if specific traits of spore ornamentation cause departures from theoretical expectations. METHODS: A fall tower design combined with a high-speed camera was used to document spore settling velocities in nine moss species selected to cover the range of spore diameters within the group. Linear mixed effect models were employed to determine whether settling velocity can be predicted from spore diameter, taking specific variation in shape and surface roughness into account. KEY RESULTS: Average settling velocity of moss spores ranged from 0·49 to 8·52 cm s(-1) There was a significant positive relationship between spore settling velocity and size, but the inclusion of variables of shape and texture of spores in the best-fit models provides evidence for their role in shaping spore settling velocities. CONCLUSIONS: Settling velocities in mosses can significantly depart from expectations derived from Stokes' Law. We suggest that variation in spore shape and ornamentation affects the balance between density and drag, and results in different dispersal capacities, which may be correlated with different life-history traits or ecological requirements. Further studies on spore ultrastructure would be necessary to determine the role of complex spore ornamentation patterns in the drag-to-mass ratio and ultimately identify what is the still poorly understood function of the striking and highly variable ornamentation patterns of the perine layer on moss spores.

Are there any widespread endemic flowering plant species in Macaronesia? Phylogeography of Ranunculus cortusifolius.

PREMISE OF THE STUDY: Oceanic island endemics typically exhibit very restricted distributions. In Macaronesia, only one endemic angiosperm species, Ranunculus cortusifolius, has a distribution spanning the archipelagos of the Azores, Madeira, and Canaries. Earlier work suggested possible differences between archipelagos and the multiple origins of the species. This paper tests the hypothesis that R. cortusifolius is a single widespread Macaronesian endemic species with a single origin. METHODS: Chloroplast (matK-trnK, psbJ-petA) and ITS sequences were generated from across the distribution of R. cortusifolius. Relationships were investigated using Bayesian inference and divergence times estimated using BEAST. Infraspecific variation was investigated using statistical parsimony. The general mixed Yule-coalescent model (GMYC) was further used to identify putative species boundaries based on maternally inherited plastid data. KEY RESULTS: The hypothesis of multiple independent origins of R. cortusifolius is rejected. Divergence of the R. cortusifolius lineage from a western Mediterranean sister group in the late Miocene is inferred. Distinct genotypes were resolved within R. cortusifolius that are endemic to the Azores, Madeira, and the Canaries. Four to five putative species were delimited by different versions of the GMYC model. CONCLUSION: Ranunculus cortusifolius is the result of a single colonization of Macaronesia. The large distances between archipelagos have been effective barriers to dispersal, promoting allopatric diversification at the molecular level with diversification also evident within the Canaries. Isolation has not been accompanied by marked morphological diversification, which may be explained by the typical association of R. cortusifolius with stable and climatically buffered laurel forest communities.

The anagenetic world of spore-producing land plants.

A fundamental challenge to our understanding of biodiversity is to explain why some groups of species diversify, whereas others do not. On islands, the gradual evolution of a new species from a founder event has been called 'anagenetic speciation'. This process does not lead to rapid and extensive speciation within lineages and has received little attention. Based on a survey of the endemic bryophyte, pteridophyte and spermatophyte floras of nine oceanic archipelagos, we show that anagenesis, as measured by the proportion of genera with single endemic species within a genus, is much higher in bryophytes (73%) and pteridophytes (65%) than in spermatophytes (55%). Anagenesis contributed 49% of bryophyte and 40% of endemic pteridophyte species, but only 17% of spermatophytes. The vast majority of endemic bryophytes and pteridophytes are restricted to subtropical evergreen laurel forests and failed to diversify in more open environments, in contrast with the pattern exhibited by spermatophytes. We propose that the dominance of anagenesis in island bryophytes and pteridophytes is a result of a mixture of intrinsic factors, notably their strong preference for (sub)tropical forest environments, and extrinsic factors, including the long-term macro-ecological stability of these habitats and the associated strong phylogenetic niche conservatism of their floras.

In defence of the entity of Macaronesia as a biogeographical region.

Since its coinage ca. 1850 AD by Philip Barker Webb, the biogeographical region of Macaronesia, consisting of the North Atlantic volcanic archipelagos of the Azores, Madeira with the tiny Selvagens, the Canaries and Cabo Verde, and for some authors different continental coastal strips, has been under dispute. Herein, after a brief introduction on the terminology and purpose of regionalism, we recover the origins of the Macaronesia name, concept and geographical adscription, as well as its biogeographical implications and how different authors have positioned themselves, using distinct terrestrial or marine floristic and/or faunistic taxa distributions and relationships for accepting or rejecting the existence of this biogeographical region. Four main issues related to Macaronesia are thoroughly discussed: (i) its independence from the Mediterranean phytogeographical region; (ii) discrepancies according to different taxa analysed; (iii) its geographical limits and the role of the continental enclave(s), and, (iv) the validity of the phytogeographical region level. We conclude that Macaronesia has its own identity and a sound phytogeographical foundation, and that this is mainly based on three different floristic components that are shared by the Macaronesian core (Madeira and the Canaries) and the outermost archipelagos (Azores and Cabo Verde). These floristic components are: (i) the Palaeotropical-Tethyan Geoflora, formerly much more widely distributed in Europe and North Africa and currently restricted to the three northern archipelagos (the Azores, Madeira and the Canaries); (ii) the African Rand Flora, still extant in the coastal margins of Africa and Arabia, and present in the southern archipelagos (Madeira, the Canaries and Cabo Verde), and (iii) the Macaronesian neoendemic floristic component, represented in all the archipelagos, a result of allopatric diversification promoted by isolation of Mediterranean ancestors that manage to colonize Central Macaronesia and, from there, the outer archipelagos. Finally, a differentiating floristic component recently colonized the different archipelagos from the nearest continental coast, providing them with different biogeographic flavours.

Evolutionary genomics of oceanic island radiations.

A recurring feature of oceanic archipelagos is the presence of adaptive radiations that generate endemic, species-rich clades that can offer outstanding insight into the links between ecology and evolution. Recent developments in evolutionary genomics have contributed towards solving long-standing questions at this interface. Using a comprehensive literature search, we identify studies spanning 19 oceanic archipelagos and 110 putative adaptive radiations, but find that most of these radiations have not yet been investigated from an evolutionary genomics perspective. Our review reveals different gaps in knowledge related to the lack of implementation of genomic approaches, as well as undersampled taxonomic and geographic areas. Filling those gaps with the required data will help to deepen our understanding of adaptation, speciation, and other evolutionary processes.

Long-term cloud forest response to climate warming revealed by insect speciation history.

Montane cloud forests are areas of high endemism, and are one of the more vulnerable terrestrial ecosystems to climate change. Thus, understanding how they both contribute to the generation of biodiversity, and will respond to ongoing climate change, are important and related challenges. The widely accepted model for montane cloud forest dynamics involves upslope forcing of their range limits with global climate warming. However, limited climate data provides some support for an alternative model, where range limits are forced downslope with climate warming. Testing between these two models is challenging, due to the inherent limitations of climate and pollen records. We overcome this with an alternative source of historical information, testing between competing model predictions using genomic data and demographic analyses for a species of beetle tightly associated to an oceanic island cloud forest. Results unequivocally support the alternative model: populations that were isolated at higher elevation peaks during the Last Glacial Maximum are now in contact and hybridizing at lower elevations. Our results suggest that genomic data are a rich source of information to further understand how montane cloud forest biodiversity originates, and how it is likely to be impacted by ongoing climate change.

A unified model of species abundance, genetic diversity, and functional diversity reveals the mechanisms structuring ecological communities.

Biodiversity accumulates hierarchically by means of ecological and evolutionary processes and feedbacks. Within ecological communities drift, dispersal, speciation, and selection operate simultaneously to shape patterns of biodiversity. Reconciling the relative importance of these is hindered by current models and inference methods, which tend to focus on a subset of processes and their resulting predictions. Here we introduce massive ecoevolutionary synthesis simulations (MESS), a unified mechanistic model of community assembly, rooted in classic island biogeography theory, which makes temporally explicit joint predictions across three biodiversity data axes: (i) species richness and abundances, (ii) population genetic diversities, and (iii) trait variation in a phylogenetic context. Using simulations we demonstrate that each data axis captures information at different timescales, and that integrating these axes enables discriminating among previously unidentifiable community assembly models. MESS is unique in generating predictions of community-scale genetic diversity, and in characterizing joint patterns of genetic diversity, abundance, and trait values. MESS unlocks the full potential for investigation of biodiversity processes using multidimensional community data including a genetic component, such as might be produced by contemporary eDNA or metabarcoding studies. We combine MESS with supervised machine learning to fit the parameters of the model to real data and infer processes underlying how biodiversity accumulates, using communities of tropical trees, arthropods, and gastropods as case studies that span a range of data availability scenarios, and spatial and taxonomic scales.

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.