Bryophyte colonisation in experimental microcosms: the role of nutrients, defoliation and vascular vegetation

A three-year multi-factorial microcosm experiment simulating fertilisation, defoliation and the composition of vascular vegetation in a dry grassland succession was used to test four hypotheses concerning the establishment and survival of bryophytes in grassland vegetation. H₁: bryophyte cover may be used to predict bryophyte species richness. H₂: bryophyte richness is suppressed at high nutrient levels and promoted by defoliation of vascular plants. H₃: species richness of bryophytes is influenced by the species composition of the vascular vegetation. H₄: bryophyte species richness is negatively correlated with vascular plant biomass.
The relationship between bryophyte richness and bryophyte cover was found to follow the classical species-area richness curve. Bryophyte species richness responded positively to defoliation and negatively to fertilisation. The species composition of vascular vegetation had no significant effect on bryophyte richness. Bryophyte species richness was lower at high vascular plant biomass and vascular plant dry weight above 400 g m⁻² appeared fatal to bryophytes. At high nutrient levels, defoliation increased bryophyte richness, but defoliation did not fully compensate for the negative effect of fertilisation. The study reinforces the concern for short lived shuttle bryophytes in the agricultural landscape.     

Is vascular plant species diversity a predictor of bryophyte species diversity in Mediterranean forests?

This study aimed to (i) investigate the congruence among the species composition and diversity of bryophytes and vascular plants in forests; (ii) test if site prioritization for conservation aims by the maximization of the pooled number of vascular plant species is effective to maximize the pooled number of bryophyte species. The study was performed in six forests in Tuscany, Italy. Four-hundred and twenty vascular plant species (61 of which were woody) and 128 bryophyte species were recorded in 109 plots. Despite the good predictive value of the compositional patterns of both woody plants and total vascular with respect to the compositional pattern of bryophytes, the species richness of the latter was only marginally related to the species richness of the former two. Bryophyte rare species were not spatially related to rare plant species and neither coincided with the sites of highest plant species richness. The species accumulation curves of bryophytes behaved differently with respect to those of woody plants or total vascular plants. Reserve selection analysis based on the maximization of the pooled species richness of either woody plants or total vascular plants were not effective in maximizing the pooled species richness of bryophytes. This study indicates that species diversity of vascular plants is not likely to be a good indicator of the bryophyte species diversity in Mediterranean forests.     

Biological legacies soften pine plantation effects for bryophytes

Biological legacies are organic structures and patterns remaining after a disturbance that may contribute to the complexity of the recovering vegetation. Legacies may, in turn, reduce the impacts of human disturbances such as logging and habitat transformation on elements of biodiversity. To examine the effects of biological legacies on biotic responses after disturbance, we surveyed 32 sites for bryophytes in an area subject to large-scale conversion of native eucalypt forest to exotic Pinus radiata D. Don plantations in eastern Australia. We sampled bryophyte and substrate diversity (log, bare ground, upturned tree/log, and trees) in eight sites in each of four landscape context classes: pine plantation stands, elliptical-shaped remnants, strip-shaped remnants, and controls in a large area of contiguous, unmanaged eucalypt forest. We found a muted response by individual species of bryophyte to landscape context. We attribute this, in part, to the presence of logs in the intensively managed pine plantation sites. The boost in bryophyte diversity from species on logs meant that some pine sites supported similar species composition to the continuous eucalypt forest controls. Our findings also underline the importance of local controls and structural variation, including leaving logs and native trees in plantations, for enhancing bryophyte species richness in managed landscapes.     

Plant diversity in wooded moderate-rich fens across boreal western Canada: an ecoregional perspective

Ecoregions are increasingly being used as a framework for conservation planning. The Mid-Boreal Uplands Ecoregion stretches across Canada from Manitoba to British Columbia. From the perspective of conservation and to understand the dynamics of plant diversity and community composition in a common wetland type, we examined the plant communities and environmental variables in 80 wooded moderate-rich fens within this ecoregion. Regional diversity totalled 273 species, with 86 bryophytes and 187 vascular plants. Total diversity was greatest in Manitoba and decreased in a longitudinal trend west through Saskatchewan and Alberta. This may be related, in part, to orographic precipitation at Manitoba sites and a gradient of growing degree days. Richness of locally rare vascular plants exhibited a clear west to east gradient. Ten species of provincially rare vascular plants were observed across the ecoregion, but without pattern. Ordinations and other analyses revealed distinct plant communities for all three locations, with vascular plant assemblages more discrete than bryophyte assemblages. Bryophyte diversity increased with latitude and longitude, whereas vascular plant diversity decreased. Additionally, elevation, precipitation, surface water alkalinity, water temperature, percent overstory density, and peat organic C played a role in determining species richness and community composition. Overall, species composition and diversity in a single wetland type exhibited continuous change across multiple political jurisdictions at the ecoregion scale. Conservation plans for wetlands at the ecodistrict scale may be preferable.     

异质城市景观中苔藓植物群落多样性的研究

苔藓植物是一类敏感的环境指示植物。通过对上海市市区及市郊5种景观类型中苔藓植物群落的取样调查,从丰富性、均匀性、相似性等方面进行了多种α和β多样性指数分析。结果表明,5种生境中的苔藓群落多样性指数呈现出明显的梯度变化,对景观异质性有较好指示作用;同时苔藓植物平均株高和大型藓类比例在不同景观中也出现规律性变化.     

Impact of Land-Use Intensity and Productivity on Bryophyte Diversity in Agricultural Grasslands

While bryophytes greatly contribute to plant diversity of semi-natural grasslands, little is known about the relationships between land-use intensity, productivity, and bryophyte diversity in these habitats. We recorded vascular plant and bryophyte vegetation in 85 agricultural used grasslands in two regions in northern and central Germany and gathered information on land-use intensity. To assess grassland productivity, we harvested aboveground vascular plant biomass and analyzed nutrient concentrations of N, P, K, Ca and Mg. Further we calculated mean Ellenberg indicator values of vascular plant vegetation. We tested for effects of land-use intensity and productivity on total bryophyte species richness and on the species richness of acrocarpous (small & erect) and pleurocarpous (creeping, including liverworts) growth forms separately. Bryophyte species were found in almost all studied grasslands, but species richness differed considerably between study regions in northern Germany (2.8 species per 16 m²) and central Germany (6.4 species per 16 m²) due environmental differences as well as land-use history. Increased fertilizer application, coinciding with high mowing frequency, reduced bryophyte species richness significantly. Accordingly, productivity estimates such as plant biomass and nitrogen concentration were strongly negatively related to bryophyte species richness, although productivity decreased only pleurocarpous species. Ellenberg indicator values for nutrients proved to be useful indicators of species richness and productivity. In conclusion, bryophyte composition was strongly dependent on productivity, with smaller bryophytes that were likely negatively affected by greater competition for light. Intensive land-use, however, can also indirectly decrease bryophyte species richness by promoting grassland productivity. Thus, increasing productivity is likely to cause a loss of bryophyte species and a decrease in species diversity.     

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

高海拔幼林地表苔藓组成与结构——皆伐与造林实践的影响评估

 对壤塘县二林场4个不同时间皆伐和造林后形成的幼林地, 开展了苔藓植物和维管植物多样性调查, 采用方差分析法对苔藓植物特征进 行差异性检验, 对苔藓植物结构特征和环境因子作偏相关性分析。结果表明: 1)幼林地4个阶段的发展进程中苔藓物种组成变化显著。优势种相 似, 为喜光耐旱的侧蒴藓物种; 一些耐阴喜湿的种类种群扩大成为林下局部生境优势物种; 同时由于干扰造成地表裸露后产生耐旱土生或石生 性广泛分布的藓类。2)随着幼林地植被恢复进程, 苔藓物种丰富度(22～37)和多样性指数(0.37～1.08)呈增长趋势。结构特征在4个幼林地表现 出较强的差异性。3)乔木与灌木层盖度与苔藓植物的发育呈正相关, 灌木层盖度的发育明显促进了苔藓植物丰富度的增加, 而基于喜光先锋的 禾草为主的草本发育程度很高, 与凋落物生物量一起抑制了苔藓的发育。随幼林地发展, 乔、灌、凋落物发育性质及其相互作用发生动态变化 。因此加速幼林地乔、灌层片的发育对于促进地表苔藓植物多样性的恢复与层片发育具有重要作用。     

Bryophyte and vascular plant species richness in boreo-nemoral moist forests and mires

We compare species richness of bryophytes and vascular plants in Estonian moist forests and mires. The material was collected from two wetland nature reserves. Bryophyte and vascular plant species were recorded in 338 homogeneous stands of approximately 1 ha in nine forest and two mire types. Regional species pools for bryophytes and vascular plants were significantly correlated. The correlations between the species richnesses of bryophytes and vascular plants per stand were positive in all community types. The relative richnesses (local richness divided by the regional species pool size) were similar for bryophyte species and for vascular plant species. This shows that on larger scales, conservation of the communities rich in species of one taxonomic plant group, maintains also the species richness of the other. The minimum number of stands needed for the maintenance of the regional species pool of typical species for the every community type was calculated using the species richness accumulation curves. Less stands are needed to maintain the bryophyte species pools (300–5300 for bryophytes and 400–35 000 for vascular plants).     

Epigeic bryophytes do not improve bioindication by Ellenberg values in mountain forests

Based on a stratified random sample of 93 vegetation plots and coincident measurements of ecological conditions in mountain forests of the Bavarian Alps, the degree to which species composition and Ellenberg indicator values derived thereof were related to measured environmental variables was assessed for vascular understorey plants and epigeic bryophytes. According to Mantel tests vascular composition contained ca. 30% more ecological information than bryophyte composition. When expressed as average Ellenberg or Düll values, vascular plant-based values reflected 60% more of measured variables than bryophyte-based values. The differences remained after rarefaction of the vascular matrix to the gamma diversity of bryophytes, showing that indication is not a function of indicator richness. Analysing vascular plants and bryophytes combined yielded very similar, or even slightly less stringent relationships with the environment than using vascular plants only.Bivariate relationships of indicator values with corresponding ecological measurements confirmed the specific potential of the values to estimate ecological factors from both plant groups, but vascular plants performed better for all factors. Bryophyte indication was particularly poor for light, temperature and base saturation. Bryophyte-based indicator values did not significantly predict the residuals of measured ecological variables against vascular plant-based Ellenberg values.For the study region, it is concluded that indicator values of vascular forest understorey should be used without consideration of Düll's indicator values for epigeic bryopyhtes. There appears to be potential to improve bioindication by recalibrating indicator values of epigeic bryophytes based on ecological measurements and vascular plant indicator values.     

The natural exclusion of red deer from large boulder grazing refugia and the consequences for saxicolous bryophyte and lichen ecology

Large boulder grazing refugia permitted comparison of saxicolous bryophyte and lichen assemblages with those boulder tops accessible to red deer (Cervus elaphus) on a sporting estate in northwest Scotland. Plant succession was predicted to occur unchecked by grazing on the tops of these large boulders with cascading effects on bryophytes and lichens—assuming boulders had been in place over the same time period. Fifty pairs of boulders (one ≥2 m and the other accessible to red deer) were selected at random from various locations below north-facing crags. Percentage cover of each bryophyte and lichen species was estimated from three randomly placed quadrats on each boulder top. Due consideration was given to the influence of island biogeography theory in subsequent model simplification. Mean shrub cover and height, leaf-litter, bryophyte cover and bryophyte species richness were significantly higher within quadrats on large boulder tops that naturally excluded red deer. Lichen cover and lichen species richness were significantly higher on boulder tops accessible to red deer. Lichen cover was in a significant negative relationship with bryophyte cover, shrub cover and litter cover. Bryophyte cover showed a significant positive relationship with shrub height but there was an optimum shrub cover. Natural exclusion of red deer from the tops of large boulders has facilitated plant succession. The results suggest that grazing arrests the lithosere on boulder tops accessible to red deer at an early plagioclimax favouring saxicolous lichens. The results are relevant to situations where red deer might be excluded from boulder fields that hold lichen assemblages of conservation value.     

Species richness of vascular plants, bryophytes and lichens in dry grasslands: The effects of environment, landscape structure and competition

We studied the relative importance of local habitat conditions and landscape structure for species richness of vascular plants, bryophytes and lichens in dry grasslands on the Baltic island of Öland (Sweden). In addition, we tested whether relationships between species richness and vegetation cover indicate that competition within and between the studied taxonomic groups is important. We recorded species numbers of vascular plants, bryophytes and lichens in 4 m² plots (n=452), distributed over dry grassland patches differing in size and degree of isolation. Structural and environmental data were collected for each plot. We tested effects of local environmental conditions, landscape structure and vegetation cover on species richness using generalized linear mixed models. Different environmental variables explained species richness of vascular plants, bryophytes and lichens. Environmental effects, particularly soil pH, were more important than landscape structure. Interaction effects of soil pH with other environmental variables were significant in vascular plants. Plot heterogeneity enhanced species richness. Size and degree of isolation of dry grassland patches significantly affected bryophyte and lichen species richness, but not that of vascular plants. We observed negative relationships between bryophyte and lichen species richness and the cover of vascular plants. To conclude, effects of single environmental variables on species richness depend both on the taxonomic group and on the combination of environmental factors on a whole. Dispersal limitation in bryophytes and lichens confined to dry grasslands may be more common than is often assumed. Our study further suggests that competition between vascular plants and cryptogams is rather asymmetric.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

Endozoochory by slugs can increase bryophyte establishment and species richness

Herbivory can affect plant community composition and diversity by removing biomass and reducing light competition. Herbivory may particularly benefit low growing species such as bryophytes, which are frequently limited by light competition. Gastropods are important herbivores of seed plants and cryptogams, furthermore, they can disperse propagules such as seeds and spores via endozoochory. However, whether gastropod herbivory can reduce the dominance of vascular plants and thereby promote the germination and establishment of endozoochorously dispersed bryophyte spores has never been tested experimentally. Moreover, it is unclear whether these possible interacting effects can influence bryophyte species richness. Here, we tested for endozoochorous spore dispersal by slugs, in combination with sowing of vascular plants, in a fully factorial common garden experiment. Enclosures contained either slugs previously fed with bryophyte sporophytes, control slugs, or no slugs. After 21 days the bryophyte cover was on average 2.8 times higher (3.9% versus 1.4%) and after eight months the bryophyte species richness 2.6 times higher (5.8 versus 2.2) in enclosures containing slugs previously fed with bryophyte sporophytes than in the other treatments. Furthermore, after eight months high vascular plant cover reduced bryophyte diversity. On average enclosures without seed sowing harboured 1.6 times more bryophyte species than the ones with seed sowing (4.2 versus 2.6), indicating competitive effects of vascular plants on bryophytes. Our findings suggest that slugs are important dispersal vectors for bryophytes and that they can increase bryophyte populations and maintain bryophyte diversity by reducing the dominance of vascular plants.     

Explaining bird species composition and richness in eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the factors influencing the distribution of birds in remnants in a fragmented agricultural landscape. Location Forty‐seven eucalypt remnants and six sites in continuous forest in the subhumid Midlands region of Tasmania, Australia. Methods Sites were censused over a two‐year period, and environmental data were collected for remnants. The avifauna of the sites was classified and ordinated. The abundances of bird species, and bird species composition, richness, abundance and diversity were related to environmental variables, using simple correlation and modelling. Results There were two distinct groups of sample sites, which sharply differed in species composition, richness, diversity and bird abundance, separated on the presence/absence of noisy miner (Manorina melanocephala Latham) colonies, remnant size, vegetation structural attributes and variables that reflected disturbance history. The approximate remnant size threshold for the change from one group to another was 20–30 ha. Remnant species richness and diversity were most strongly explained by remnant area and noisy miner abundance, with contributions from structural and isolation attributes in the second case. Segment richness was explained by precipitation, logging history and noisy miner abundance. Bird abundance was positively related to precipitation and negatively related to tree dieback. The 28 individual bird species models were highly individualistic, with vegetation structural variables, noisy miner abundance, climatic variables, variables related to isolation, area, variables related to floristics, disturbance variables, the nature of the matrix and remnant shape all being components in declining order of incidence. Age of the remnant did not relate to any of the dependent variables. Main conclusions Degraded and small remnants may have become more distinct in their avifaunal characteristics than might otherwise be the case, as a result of the establishment of colonies of an aggressive native bird, the noisy miner. The area, isolation and shape of remnants directly relate to the abundance of relatively few species, compared to vegetation attributes, climate and the abundance of the noisy miner. The nature of the matrix is important in the response of some species to fragmentation.     

Effect of fragmentation on the community structure of epixylic bryophytes in Atlantic Forest remnants in the Northeast of Brazil

The effects of fragmentation and edge effects on the floristic composition, richness, diversity and abundance of epixylic bryophytes (growing on decaying wood) were investigated in ten fragments of Atlantic Forest remnants in the Northeast of Brazil. In each fragment, four perpendicular 100 m transects were demarcated. Along these transects, samples of bryophytes growing on decaying wood were collected. The forest fragments were grouped in three size classes (small: <100 ha; medium: 100–500 ha; large: >500 ha). Correlation and multivariate analysis were undertaken between bryophyte flora and fragment metrics (size, form, isolation, altitude variation, nuclear area and secondary vegetation percentage and distance from the edge). A total of 99 species of bryophytes, 52 liverworts and 47 mosses were registered. The statistical results confirming fragment size is an important factor in epixylic community structure. Therefore, composition, richness, diversity and abundance can be better explained by a junction of all studied landscape factors. Bryophyte richness, the percentage of samples with the greatest coverage of decaying wood and shade-tolerant species’ distribution, were not correlated to distance from the forest edge. This suggests that edge effects are not linear or can be detected beyond 100 m from the edge, which is very important for inclusion in future studies.     

Structural equation modelling detects unexpected differences between bryophyte and vascular plant richness along multiple environmental gradients

Aim The aim of this study was to develop and evaluate a structural equation model explaining the species richness of bryophytes and vascular plants along multiple environmental gradients. Our primary interests in developing this model were: (1) to evaluate the effect of tree canopy along an altitudinal gradient on bryophyte and vascular plant richness; (2) to determine to what extent lithology was able to explain richness in the two groups of plants; (3) to assess whether anthropogenic disturbances decrease richness; and (4) to explore, comparing competing models, the causal links connecting spring area, discharge and spring complexity and to assess how these variables are related to richness. Location Eighty‐six springs were sampled in the south‐eastern Alps of Italy, from the lowlands to high‐mountain regions, and on different lithologies. Methods A structural equation model (SEM) was used to test certain hypotheses about the direct and indirect effects of altitude, canopy, lithology, disturbance, spring complexity, discharge and spring area on bryophyte and vascular plant richness. Competing models were evaluated and bootstrap simulation was used to determine the stability of parameter estimates. Results SEM analysis made it possible to disentangle the different effects of canopy and lithology on bryophyte and vascular plant richness: for the former it was demonstrated that the increased richness with altitude was related primarily to lithology, whereas the latter increased because of the reduction of canopy cover. In addition, the model predicted that spring discharge determined the size of the spring area and even the complexity of the spring; these latter two variables influenced to different degrees both bryophyte and vascular plant richness. Anthropogenic disturbances affected the richness of bryophytes more than that of the vascular plants. Main conclusions Our study demonstrates that several similarities in the patterns of bryophyte and vascular plant richness are in fact induced by different environmental variables.     

Cryptogam communities on decaying deciduous wood – does tree species diversity matter?

Bryophyte and fungal communities were investigated on fallen trees representing seven deciduous tree species in a mixed near natural nemoral forest. Bryophytes were represented by 41 taxa, including several very frequent species. Of the 296 fungal species, most were recorded with very low frequency and the share of high frequent species was much lower than among the bryophytes. Species turnover was bigger in the fungal communities, compared to the bryophyte communities, and related to a higher extent to measured differences in environmental conditions. Tree species diversity was found to be an important factor for fungal species composition, while only small differences in bryophyte species composition were found between the different tree species. On the other hand bryophyte species richness showed distinct relations to tree species and microclimatic variables, a tendency which was not evident for fungal diversity. It is concluded that the two organism groups to some extent differ in their conservation demands. Thus, conservation of wood-inhabiting bryophytes requires prioritising of large, coherent forest stands in which a stable humid microclimate and a reasonable supply of dead wood is secured. Successful conservation of fungi requires that substantial amounts of dead wood are left for natural decay in a variety of natural forest environments representing different tree species, so that heterogeneity in dead wood types is secured.     

The impact of land abandonment on the plant diversity of olive groves

We investigated the effect of woody species’ encroachment on plant diversity changes with regard to vascular plants and bryophytes in traditional olive groves of the Maremma Regional Park (Tuscany, Italy) and assessed cross-taxon correlation between these two taxa. We classified the olive groves into four land use types, representing different successional stages. To describe the evenness of species distribution within a community, we plotted rank-abundance diagrams for each taxon and each land use type. The relationship between the number and cover of vascular plants, therophytes, bryophytes, colonists and phanerophytes in each plot was examined using linear regression. The effects of land use type on vascular plant and bryophyte richness and assemblages were assessed by permutational uni- and multivariate analysis of variance. The congruence in species composition between the two taxa was evaluated using Procrustes analysis. The number of vascular plants, bryophytes and therophytes decreased linearly with increased phanerophyte species cover. The number of species belonging to Thero-Brachypodietea progressively decreased throughout succession. Rank-abundance diagrams and multivariate analysis showed differences between the land use types, which were statistically significant for vascular plants between the traditional olive groves and the other land use types, and for bryophytes between the traditional olive groves and woodlands. PROTEST analysis and NMDS graphs showed a correlation between vascular plant and bryophyte communities. The results suggested that conservation measures are needed in the study area in order to ensure both the maintenance of traditional olive groves of conservation interest and high levels of environmental heterogeneity.     

Bryophyte diversity and distribution along an altitudinal gradient on a lava flow in La Réunion

Non‐vascular plant distribution patterns were examined in three microhabitats along an altitudinal gradient on a recent lava flow of the Piton de la Fournaise volcano (La Réunion, Mascarene archipelago). The uniform nature of the lava flow provides an excellent system to study the relationship between altitude and species diversity and distribution, and at the same time avoiding confusing multiple effects of substrate and vegetation heterogeneity. Non‐vascular plants were surveyed with quadrats within an altitudinal range from 250 m to 850 m a.s.l. Fine‐scale variations in bryophyte communities between three ecological microhabitats (the ground and on the rachises of two fern species) were investigated. Three specific questions were addressed: (1) What is the species diversity of bryophyte communities on a 19‐year‐old lava flow? (2) How does altitude influence the diversity and distribution of bryophytes on a lava flow? (3) Does microhabitat variation control bryophyte diversity? In our study, bryophyte diversity increased with altitude. Unexpectedly, species richness was very high; 70 species of bryophytes were recorded including nine new records for the island. Diversity was also controlled by ecological microhabitats. Bryophyte species were structured into six categories according to altitude and microhabitat preferences. Results suggested that the high diversity of these cryptic organisms on this lava flow is fostered in part by their host substrate and their adaptative strategies on new substrates. On a broader scale, it was concluded that lava flows as primary mineral environments are important to conserve, as they support a high diversity of pioneer organisms that constitute the early stages of the development of La Réunion's remnant lowland rainforest.