The physiological basis of bryophyte production

In the main features of their carbon metabolism and physiological responses, bryophytes behave as normal C_:) plants. However, their small size and frequent poikilohydric habit have important effects on the context in which these characteristics are expressed, and on their environmental physiology. Many are tolerant of drying out to low water contents (c. 5–10%, of dry weight). Photosynthesis declines rapidly with water loss, and resumes with greater or lesser delay on remoistening. The rate and completeness of recovery depend on the intensity and duration of desiccation, and on drought-hardening (perhaps largely related to protection of cell components from oxidative damage) which lakes place as the bryophyte dries. Most bryophytes, including species of well-illuminated habitats, function in effect as shade plants, with low chlorophyll a/b ratios, and become light-saturated at relatively low irradiance. Boundary-layer resistance is critically important in determining water loss from bryophytes in many situations. The time for which a poikilohydric species can photosynthesize after rain is determined by storage capacity and rate of water loss, both strongly influenced by growth-form. In sheltered habitats with extensive bryophyte cover water loss is largely determined by radiation balance, and may be very slow in deeply shaded places. Bryophyte growth-forms must represent an adaptive balance between water economy and needs for light capture and carbon and mineral nutrient acquisition.     

哀牢山湿性常绿阔叶林地生、树干及树枝 附生苔藓生活型组成及其水分特性

为探讨林内不同垂直高度生境下及不同生活型苔藓植物水分特征的差异,该文对哀牢山湿性常绿阔叶林内地生苔藓、林下树干附生和林冠层树枝附生苔藓优势或常见种的生活型组成、持水力、失水特征和水分利用效率进行了研究。结果表明:地生、树干附生和树枝附生苔藓植物类群分别以交织型、扇型、悬垂型苔藓植物占据优势;地生、树干附生、枝条附生苔藓植物的饱和持水率分别为476%DW、210.98%DW、238.95%DW;地生苔藓植物的持水率和失水速率均高于附生苔藓,树干附生苔藓植物持水率低于树枝附生苔藓,而失水速率高于树枝附生苔藓。在不同生活型苔藓的水分特性上,交织型苔藓具有较高的持水率和失水速率,保水性能较弱,其次为悬垂型苔藓,扇型苔藓的持水率最小,失水速率也较快;地生和树干附生苔藓的水分利用效率均显著高于树干附生苔藓,交织型和扇型苔藓的水分利用效率显著高于悬垂型苔藓。在三种生境下,地生苔藓持水力高,水分利用效率较高而保水能力低;枝条附生苔藓持水力低,水分利用效率低而保水能力较高;树干附生苔藓水分利用效率较高而持水力和保水能力均较差。因此,不同生境下苔藓植物生活型组成及其水分变化特性在一定程度上反映了它们对不同生境的适应策略。     

Differential proteomics of dehydration and rehydration in bryophytes: evidence towards a common desiccation tolerance mechanism

All bryophytes evolved desiccation tolerance (DT) mechanisms during the invasion of terrestrial habitats by early land plants. Are these DT mechanisms still present in bryophytes that colonize aquatic habitats? The aquatic bryophyte Fontinalis antipyretica Hedw. was subjected to two drying regimes and alterations in protein profiles and sucrose accumulation during dehydration and rehydration were investigated. Results show that during fast dehydration, there is very little variation in protein profiles, and upon rehydration proteins are leaked. On the other hand, slow dehydration induces changes in both dehydration and rehydration protein profiles, being similar to the protein profiles displayed by the terrestrial bryophytes Physcomitrella patens (Hedw.) Bruch and Schimp. and, to what is comparable with Syntrichia ruralis (Hedw.) F. Weber and D. Mohr. During dehydration there was a reduction in proteins associated with photosynthesis and the cytoskeleton, and an associated accumulation of proteins involved in sugar metabolism and plant defence mechanisms. Upon rehydration, protein accumulation patterns return to control values for both photosynthesis and cytoskeleton whereas proteins associated with sugar metabolism and defence proteins remain high. The current results suggest that bryophytes from different ecological adaptations may share common DT mechanisms.     

Production and microtopography of bog bryophytes: response to warming and water-table manipulations

Boreal peatlands, which contain a large fraction of the world's soil organic carbon pool, may be significantly affected by changes in climate and land use, with attendant feedback to climate through changes in albedo, fluxes of energy or trace gases, and soil carbon storage. The response of peatlands to changing environmental conditions will probably be dictated in part by scale-dependent topographic heterogeneity, which is known to interact with hydrology, vegetation, nutrients, and emissions of trace gases. Because the bryophyte community can contribute the majority of aboveground production in bogs, we investigated how microscale topography affects the response of bryophyte species production and cover to warming (using overhead infrared lamps) and manipulations of water-table height within experimental mesocosms. We removed 27 intact peat monoliths (2.1-m² surface area, 0.5-0.7 m depth) from a bog in northern Minnesota, USA, and subjected them to three warming and three water-table treatments in a fully crossed factorial design. Between 1994 and 1998, we determined annual production of the four dominant bryophyte taxa within three microtopographic zones (low, medium, and high relative to the water table). We also estimated species cover and calculated changes in topography and roughness of the bryophyte surface through time. Total production of all bryophytes, and production of the individual taxa Polytrichum strictum, Sphagnum magellanicum, and Sphagnum Section Acutifolia, were about 100% greater in low microtopographic zones than in high zones, and about 50% greater in low than in medium zones. Production of bryophytes increased along the gradient of increasing water-table heights, but in most years, total production of bryophytes was negatively correlated with height above the set water table only for the wettest water-table treatment. Although bryophyte production was unaffected by the warming treatments, the bryophyte surface flattened in proportion to the degree of warming. These results indicate that production of bryophytes is driven most strongly by the absolute and relative height of the bryophyte surface above the water table. Predicted changes in water-table height commensurate with changes in surface temperature may thus affect both production and superficial topography of bryophyte communities.     

The importance of colony structure versus shoot morphology for the water balance of 22 subarctic bryophyte species

Questions: What are the water economy strategies of the dominant subarctic bryophytes in terms of colony and shoot traits? Can colony water retention capacity be predicted from morphological traits of both colonies and separate shoots? Are suites of water retention traits consistently related to bryophyte habitat and phylogenetic position? Location: Abisko Research Station, North Sweden. Methods: We screened 22 abundant subarctic bryophyte species from diverse habitats for water economy traits of shoots and colonies, including desiccation rates, water content at field capacity, volume and density (mg cm^?3) of water‐saturated and oven‐dried patches, evaporation rate (g·m^?2·s^?1) and cell wall thickness. The relationships between these traits and shoot and colony desiccation rates were analysed with Spearman rank correlations. Subsequent multivariate (cluster followed by PCA) analyses were based on turf density, turf and shoot desiccation rate, cell wall thickness and amount of external and internal water. Results: Individual shoot properties, i.e. leaf cell wall properties, water retention capacity and desiccation rate, did not correspond with colony water retention capacity. Colony desiccation rate depended on density of water‐saturated colonies, and was marginally significantly negatively correlated with species individual shoot desiccation rate but not related to any other shoot or colony trait. Multivariate analyses based on traits assumed to determine colony desiccation rate revealed six distinct species groups reflecting habitat choice and phylogenetic relationships. Conclusions: General relationships between shoot and colony traits as determinants of water economy will help to predict and upscale changes in hydrological function of bryophyte‐dominated peatlands undergoing climate‐induced shifts in species abundance, and feedbacks of such species shifts on permafrost insulation and carbon sequestration functions.     

Drag coefficients of stream bryophytes: experimental determinations and ecological significance

1. Drag coefficients ( C D) of bryophyte-covered rocks were measured to see whether these differed between species. Replicate rocks, each supporting one of six bryophyte taxa, were attached to a base plate mounted on Teflon bearings in a flow tank. The drag force exerted on rocks with and without bryophyte material was measured by a strain gauge at different water velocities.
2. The difference in C D between rocks with and without bryophyte material was calculated for each plant, and expressed as a percentage change ( δC D). This varied significantly from 0 in three of the six taxa. The cushion-shaped moss Bryum blandum increased the C D by around 10%. The moss Blindia lewinskyae and liverwort Cryptochila grandiflora decreased the C D by around 40 and 30%, respectively, presumably reflecting their streamlined growth.
3. Drag characteristics of aquatic bryophytes may help explain differences in their resistance to fast flows. Furthermore, we suggest that some aquatic bryophytes can increase substrate stability by streamlining rocks, rendering them less prone to movement.     

On the influence of water conductivity,pH and climate on bryophyte assemblages in Catalan semi-natural springs

Bryophytes are some of the most sensitive biological indicators of environmental change. Springs have a significant presence of bryophytes and so are ideal habitats for studying their relationship with the environment. We tested whether bryophyte assemblages can be explained with macro-, meso- and micro-ecological variables (i.e. seasonal climate, altitude, water pH and conductivity) sampling bryophytes from 198 semi-natural springs distributed along montane regions in the north-eastern Iberian Peninsula. We tested the influence of environmental variables on bryophyte assemblages in springs using sparse Partial Least Squares. Our results show that variability in bryophyte assemblages is explained by seasonal climate (temperature and precipitation from winter, spring, summer and autumn and temperature and precipitation seasonality), altitude and water conductivity. The results obtained by the present study will be useful for predicting bryophyte diversity in springs using simple and easy to obtain variables such as climate, water pH and conductivity.     

干旱与半干旱地区苔藓植物生态学研究综述

苔藓植物一般生活在阴湿的地方,但也有不少种类具有极强的耐旱性,在干旱和半干旱地区也有分布,在干旱和半干旱地区,影响苔藓植物分布的环境因子包括降水,pH值,CaCO3含量,植被盖度,有机质含量以及土壤质地等,其中起沙,固沙和生态小环境的改善中发挥着不可替代的作用。     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

Decomposition rates of bryophytes in managed boreal forests: influence of bryophyte species and forest harvesting

The slow decomposition rate of boreal forest floor bryophytes contributes both to maintaining high soil C reserves as well as affecting conditions for tree growth by maintaining excessively high soil water content, cooling the soil and slowing nutrient cycles. In this study, mass loss of three bryophyte species (Pleurozium schreberi, Sphagnum capillifolium, S. fuscum) was measured in unharvested, partial cut and low-retention cut forest blocks. Mesh decomposition bags containing the three species and wood sticks were placed at two depths in colonies of either P. schreberi or S. capillifolium (environment) in the three harvest treatments and retrieved after two growing seasons. Mass loss was primarily related to substrate type (P. schreberi > S. capillifolium > wood sticks > S. fuscum) and secondarily to depth. Harvest treatment and environment (P. schreberi or S. capillifolium) only weakly affected sphagna mass loss. The weak effect of harvest treatment suggests that conditions created by low retention cuts do not to stimulate decomposition in this system and are not important enough to stimulate carbon loss, or to counteract paludification. On the other hand, the strong effect of bryophyte type indicates that conditions affecting bryophyte colonization and succession are of great importance in driving carbon and nutrient cycles.     

Bryophyte dispersal inferred from colonization of an introduced substratum on Whiteface Mountain, New York

A long-standing debate in bryophyte biogeography concerns the frequency of long-distance spore dispersal. The diversity of bryophytes on mortared rock walls along the Veterans Memorial Highway on Whiteface Mountain, New York, USA, was studied to document the recruitment of species over the 65 years since the highway was constructed. The highway is situated in the Adirondack Mountains, a relatively unpopulated region with a largely acidic flora. The introduction of mortar has increased the bryophyte diversity by 50% above that of native lithic substrata on the mountain. The composition of the native and mortar floras differed greatly, suggesting that the walls were not colonized by locally abundant ruderal species. Many of the species sampled on the walls are typically found only in lower elevation forested sites, distant (～5 km or more) from the highway, and not on anthropogenic calcium carbonate. These results suggest that a bryophyte community consisting of common and uncommon species assembled from distant sites at the rate of at least one species per year in the last 65 years. These data provide the ecological context for experimental and phylogeographic studies and suggest that some bryophytes may be capable of routine dispersal over distances of at least 5 km.     

Change in the bryophyte diversity and species composition of Central European temperate broad-leaved forests since the late nineteenth century

The diversity and species composition of the bryophyte flora colonizing the forest floor, live trees and deadwood in semi-natural broad-leaved forests of northern Germany was compared between a recent survey in 2013 and historical records from around 1900. The survey was based on the comparison of presence/absence data and their interpretation with help of ecological indicator values. Total species richness has declined in bryophytes growing on the forest floor but not in the other guilds, whereas a dramatic species turnover was found for all three guilds. This turnover was apparently primarily driven by the increased atmospheric load of reactive nitrogen from anthropogenic emissions. Sensitive species were replaced by more eutrophication-tolerant bryophytes; liverworts are overrepresented among the sensitive and thus declining species. Promotion of the competitive strength of vascular plants due to increased nitrogen levels is the likely cause of the reduction of bryophyte species richness on the forest floor. Former acidification by high atmospheric sulfur dioxide loads has left an imprint in the bryophyte vegetation by having favored acidophytic species and discriminating against basiphytic species. An increase in the mean indicator value for temperature suggests a beginning effect of climate warming on the bryophyte vegetation. Change in forest structure had an apparently smaller imprint on the bryophyte diversity of the studied semi-natural forests than atmospheric chemistry and climate. In this respect, bryophytes differ from the ecologically similar lichens, where published studies from the same region showed a dramatic decline of species richness and a stronger susceptibility to forest management.     

The contribution of bryophytes to the carbon exchange for a temperate rainforest

Bryophytes blanket the floor of temperate rainforests in New Zealand and may influence a number of important ecosystem processes, including carbon cycling. Their contribution to forest floor carbon exchange was determined in a mature, undisturbed podocarp‐broadleaved forest in New Zealand, dominated by 100–400‐year‐old rimu (Dacrydium cupressimum) trees. Eight species of mosses and 13 species of liverworts contributed to the 62% cover of the diverse forest floor community. The bryophyte community developed a relatively thin (depth <30 mm), but dense, canopy that experienced elevated CO₂ partial pressures (median 46.6 Pa immediately below the bryophyte canopy) relative to the surrounding air (median 37.6 Pa at 100 mm above the canopy). Light‐saturated rates of net CO₂ exchange from 14 microcosms collected from the forest floor were highly variable; the maximum rate of net uptake (bryophyte photosynthesis – whole‐plant respiration) per unit ground area at saturating irradiance was 1.9 μmol m^?2 s^?1 and in one microcosm, the net rate of CO₂ exchange was negative (respiration). CO₂ exchange for all microcosms was strongly dependent on water content. The average water content in the microcosms ranged from 1375% when fully saturated to 250% when air‐dried. Reduction in water content across this range resulted in an average decrease of 85% in net CO₂ uptake per unit ground area. The results from the microcosms were used in a model to estimate annual carbon exchange for the forest floor. This model incorporated hourly variability in average irradiance reaching the forest floor, water content of the bryophyte layer, and air and soil temperature. The annual net carbon uptake by forest floor bryophytes was 103 g m^?2, compared to annual carbon efflux from the forest floor (bryophyte and soil respiration) of ?1010 g m^?2. To put this in perspective of the magnitude of the components of CO₂ exchange for the forest floor, the bryophyte layer reclaimed an amount of CO₂ equivalent to only about 10% of forest floor respiration (bryophyte plus soil) or ～11% of soil respiration. The contribution of forest floor bryophytes to productivity in this temperate rainforest was much smaller than in boreal forests, possibly because of differences in species composition and environmental limitations to photosynthesis. Because of their close dependence on water table depth, the contribution of the bryophyte community to ecosystem CO₂ exchange may be highly responsive to rapid changes in climate.     

Could the Canopy Structure of Bryophytes Serve as an Indicator of Microbial Biodiversity? A Test for Testate Amoebae and Microcrustaceans from a Subtropical Cloud Forest in Dominican Republic

The mechanisms that ultimately regulate the diversity of microbial eukaryotic communities in bryophyte ecosystems remain a contentious topic in microbial ecology. Although there is robust consensus that abiotic factors, such as water chemistry of the bryophyte and pH, explain a significant proportion of protist and microcrustacean diversity, there is no systematic assessment of the role of bryophyte habitat complexity on such prominent microbial groups. Water-holding capacity is correlated with bryophyte morphology and canopy structure. Similarly, canopy structure explains biodiversity dynamics of the macrobiota suggesting that canopy structure may also be a potential parameter for understanding microbial diversity. Canopy roughness of the dominant bryophyte species within the Bahoruco Cloud Forest, Cachote, Dominican Republic, concomitant with their associated diversity of testate amoebae and microcrustaceans was estimated to determine whether canopy structure could be added to the list of factors explaining microbial biodiversity in bryophytes. We hypothesized that smooth (with high moisture content) canopies will have higher species richness, density, and biomass of testate amoebae and higher richness and density of microcrustaceans than rough (desiccation-prone) canopies. For testate amoebae, we found 83 morphospecies with relative low abundances. Species richness and density differed among bryophytes with different bryophyte canopy structures and based on non-metric multidimensional scaling, canopy roughness explained 25% of the variation in species composition although not as predicted. Acroporium pungens (low roughness, LR) had the lowest species richness (2 ± 0.61 SD per gram dry weight bryophyte), and density (2.1 ± 0.61 SD individual per gram of dry weight bryophyte); whereas Thuidium urceolatum (high roughness) had the highest richness (24 ± 10.82 SD) and density (94 ± 64.30 SD). The fact that the bryophyte with the highest roughness had the highest levels of diversity for testate amoebae suggests that moisture levels at the level of the bryophyte canopy may not represent a biodiversity driver in a cloud forest with high relative humidity; however, high roughness could generate a dynamic and fluctuating moisture environment with concomitant alternating microbial communities. A total of 26 microcrustacean morphospecies were found across 11 bryophytes; however, no bryophyte canopy effect was detected on their richness and density. Microcrustacean mean density was low ranging from less than one individual per 50 cm2 of bryophyte in Leucobryum (LR) to a maximum of 6 ± 3.37 SD individuals/50 cm2 in Monoclea (LR). This lack of pattern suggests that possible explanatory variables may be related to larger scale processes than those examined in this study.     

WATER-REGULATORY BEHAVIOUR IN TERRESTRIAL GASTROPODS

1. Terrestrial snails and slugs are exceedingly susceptible to dehydration due to evaporative water loss from the integument and lung surface, and the deposition of a dilute mucous trail. Active slugs can lose 30–40% of their initial body weight (IBW) within 2 h. 2. Both field and laboratory studies have revealed that habitat selection by snails and slugs is well correlated with the availability of water. In addition, numerous species display homing behaviour, returning directly to their moist secluded daytime resting sites at dawn. 3. Several aspects of locomotor activity are affected by body hydration and environmental conditions such as relative humidity. Moist conditions result in termination of aestivation in snails and a generally higher level of activity in both snails and slugs. In contrast, severe dehydration initiates aestivation in snails and an increase in the intensity and duration of circadian locomotor activity in slugs. 4. Huddling behaviour is a specialized example of the general preference of slugs for moist habitats. When groups of slugs are exposed to dry environmental conditions, they form closely packed aggregations. This response results in a decrease in the rate of dehydration of the individual slugs. 5. When slugs have been dehydrated to about 90% IBW, rhythmic closures of the pneumostome are initiated. As dehydration progresses there is a reduction in the open diameter of the pneumostome. These responses reduce the total exposure of the lung surface and thereby evaporative water loss. In slugs dehydrated to about 80 % IBW, these responses can result in a 7 % reduction in water loss. 6. When slugs have been dehydrated to about 65% IBW (67·6 ± 4·3% IBW) they move on to a moist surface, assume a characteristic flattened posture and remain quiescent while water is absorbed through the surface of the foot. Once they are rehydrated (to 93·5 ± 12·4% IBW) they move off the moist surface. Thus there exists a specific dehydration threshold for the initiation of contact-rehydration and a rehydration set-point at which the response is terminated. 7. Both initiation and termination of contact-rehydration are controlled by variations in haemolymph osmotic pressure. The behaviour can be experimentally initiated by injection of hyperosmotic mannitol solution and terminated by injection of dilute saline. 8. Contact-rehydration involves bulk flow of water through an epithelial paracellular pathway in the integument of the foot. The rate of absorption of [¹⁴C]inulin during contact-rehydration is similar to that of water. The efficacy of water movement through the pathway is modulated by body hydration, the pathway being opened only in dehydrated slugs. 9. By means of the dual-limit control of contact-rehydration slugs can behaviourally regulate their body hydration and haemolymph osmolality within the tolerable hydration range described by the upper and lower limits.     

Phytogeography of bryophyte and lichen vegetation in the Windmill Islands,Wilkes Land,Continental Antarctica

The distribution and frequency of bryophyte and lichen vegetation on ice-free regions of the Windmill Islands are presented using data derived from aerial photography and ground surveying. The qualitative and quantitative plant cover of sites are listed and related to the topography and major soil characteristics of each site. The richest associations of macrolichens and bryophytes occurred on the metamorphic northern peninsulas. Species richness and frequency was generally reduced on the charnockitic southern peninsula and the islands which have been deglaciated longer. Salinity varied significantly throughout the region with the highest levels in the northern islands reflecting the presence of penguin colonies. In such sites bryophytes and lichens were virtually absent. Wind blown sea-spray contributed far less salts than direct excretion from penguins. On the peninsulas snow cover and site exposure appeared to delimit plant distribution. Higher salt levels from sea-spray on the northern aspects of the peninsulas seemed to have negligible impact on vegetation patterns with the possible exception ofBiatorella cerebriformis which was encountered only inland. The total phosphorus and nitrogen levels of the skeletal soils were generally low except in eutrophic sites adjacent to penguin colonies. The vegetation patterns are discussed in terms of the climate, topography and species autecology.     

Interception and retention of macroscopic bryophyte fragments by branch substrates in a tropical cloud forest: an experimental and demographic approach

Although the significance of canopy plant communities to ecosystem function is well documented, the process by which such communities become established in trees remains poorly known. Colonization of tree surfaces by canopy-dwelling plants often begins with the establishment of bryophytes, so the conditions that affect the dispersal of bryophytes in the forest canopy merit study. We assessed success rates of one mechanism of bryophyte propagation, the aerial dispersal of macroscopic fragments, using an experimental approach. We quantified interception and retention of marked fragments released from a 36 cm×36 cm grid 50 cm above branches of saplings and mature trees of the species Ocotea tonduzii in a montane cloud forest in Costa Rica. Only 1% of bryophyte fragments dropped over sapling crowns in this manner were retained for the 6-month duration of the study, while branches in the forest canopy with intact epiphyte loads and branches that had been stripped of their epiphytes retained 24% and 5%, respectively. Our results suggest that larger-diameter branches and the presence of other epiphytes can both improve the retention of bryophyte fragments on canopy branches. Further work will be needed to address the relative roles of other dispersal mechanisms (spores, gemmae, microscopic bryophyte fragments) and the dynamics of growth and establishment of macroscopic bryophyte fragments following their interception. Received: 1 March 1999 / Accepted: 24 August 1999     

Protective management of trees against debarking by deer negatively impacts bryophyte diversity

When wildlife populations become too large, they impact other flora and fauna within the ecosystems that they inhabit. For example, the recent rise in population numbers of sika deer in Japan has led to the stripping of bark from tree overstories in forested areas. This has led to protective management actions, such as wrapping the trunks of trees in wire mesh. The present study investigates the impact of this management action on epiphytic diversity at Mt. Ohdaigahara, which is one of the hotspots for bryophyte diversity in Japan. The correlation between the diversity of epiphytic bryophytes and environmental variables was examined, including the presence/absence of wire mesh protection. A generalized linear model showed that species richness and bryophyte cover was significantly correlated with both tree diameter (at 1.5 m height) and tree density (P < 0.01), but negatively correlated with wire mesh protection. Inductively coupled plasma-mass spectrometry analysis showed a significant 3- to 6-fold higher concentration of zinc in bryophytes occupying tree bark under wire mesh protection than for those without wire mesh. Hence, the high sensitivity of bryophytes to zinc accumulation, as a result of toxicity caused by galvanized iron mesh, has led to the loss of species richness and bryophyte cover on tree trunks. Furthermore, other heavy metals found in wire mesh may also contribute to the negative effect on bryophytes. Therefore, to establish best practices for biodiversity conservation that include bryophytes, materials that are free of heavy metals should be preferentially used for tree protection.     

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.