Association between bryophytes and seedlings in Nothofagus sub-Antarctic forests

Bryophytes, which include mosses, liverworts and hornworts, indeed play a significant role in the natural dynamics of native forests. They contribute to fundamental ecological processes such as tree regeneration. In this context, the objective was to analyse the association of bryophytes with Nothofagus seedlings, as well as whether the substrates where bryophytes grew influenced bryophyte-tree seedling association, in sub-Antarctic forests at two contrasting landscapes (coastal and mountain) of Tierra del Fuego (Argentina), to better understand their impacts on the natural regeneration processes. We hypothesized that bryophytes act as a substrate for seed germination and initial regeneration growth in pure and mixed forests of N. pumilio and N. betuloides, resulting in widespread tree seedlings–bryophyte associations in all landscapes. In each forest and landscape location, 60 transects were established (10 m in length) to evaluate bryophyte cover (by point intercept method) and substrate type (bare soil, decaying wood and litter cover) where they were growing. An adaptation of the relative interaction index (RII) based on tree seedling cover associated with bryophyte species (%) and tree seedling cover in the absence of bryophytes (%) was calculated and analysed to assess competition or facilitation between bryophytes and tree seedlings. Nothofagus pumilio seedlings were less abundant in bryophytes compared to other substrates, suggesting an inhibitory effect on the germination and/or survival of N. pumilio. In contrast, the seedlings of N. betuloides, in both pure and mixed forests, exhibited higher abundance on bryophyte substrates, particularly in mosses at mountain landscape (RII = 0.83 ± 0.08 in litter and 0.62 ± 0.11 in decaying wood). These findings suggest that bryophytes play a facilitating role for N. betuloides seedlings during germination and initial phases of growth, mainly in the mountain. Therefore, their conservation promotes N. betuloides forest continuity, both in pure and mixed structures.     

生态恢复方式对喀斯特生境苔藓植物物种组成的影响

为探究生态恢复方式对苔藓植物的影响，研究了自然弃耕和退耕还林等生态恢复方式对重庆市中梁山喀斯特地区不同生境类型下苔藓植物物种组成特征的影响。结果表明，研究区有苔藓植物15科26属41种，其中藓类39种，苔类2种，美灰藓(Eurohypnum leptothollum)为主要优势物种；研究区石生藓类植物占比较大，生活型以平铺型和丛集型为主，各生境物种多样性及相似性指数整体偏低，但相比弃耕杂草地、灌木林和经济型林地，竹林和马尾松林中苔藓植物具有更高的物种多样性；冗余分析表明，郁闭度和光照强度是影响苔藓植物分布的关键环境因子，乔木林为苔藓植物的生长提供了更适宜的生境。因此，相比于自然恢复和经济性林地等恢复方式，退耕还林是喀斯特退化生境更为理想的生态恢复方式。     

Assemblages of native bryophytes in secondary forests with introduced Quercus rubra

We have studied the diversity of bryophytes in planted Polish post‐agricultural forests dominated by the native Scots pine Pinus sylvestris and the alien (North American) red oak Quercus rubra. The planted sites would be suitable for a mesic coniferous forest (abbreviation: CFS) or mesic broadleaved forest (abbreviation: BFS). We analysed the structure and composition of the bryophyte assemblages in relation to forest site and substrate availability. Special attention was paid to the introduced Q. rubra as a host species for native bryophytes. A total of 54 bryophyte species (9 liverworts and 45 mosses) were found in the 90 plots ( = phytosociological relevés, 10 × 10 m in area; 45 at each forest site) studied. DCA analysis showed that the bryophyte assemblages of the P. sylvestris–Q. rubra secondary forest community differed significantly between CFS and BFS sites; the similarity of the composition of bryophyte species was 36.8%. The substrate preferences (epigeic, epixylic, epiphytic), as well as the growth form and life form of the recorded bryophytes, also differed between CFS and BFS, while the proportion of bryophytes that had a particular life strategy was very similar. The introduced Q. rubra was inhabited by 28 bryophyte species, including two liverworts. This tree hosted 64% of the CFS and 47% of BFS bryophyte flora and as a host for epiphytes the species successfully fulfilled the functional role of the native oaks (Q. robur and Q. petraea). Thus, the introduction of Q. rubra may contribute to the restoration of post‐agricultural forests and to the conservation of epiphytic bryophyte species. On the other hand, the negative impact of Q. rubra observed on the ground flora (including bryophytes) puts the benefits of Q. rubra for the conservation of native biodiversity in general in question.     

Species richness and distribution of understorey bryophytes in different forest types in Colombian Amazonia

Abstract

The first bryophyte survey results from Colombian Amazonia are presented. Bryophyte species, differentiated into mosses and liverworts, and further into four life-form classes, were sampled in 0.1-ha plots. These plots were distributed over four landscape units in the middle Caquetá area: floodplains, swamps, terra firme forests and white-sand areas. The total numbers of bryophyte species in the units were 50, 45, 45 and 32, respectively. The plots in floodplains and swamps were richer in moss species than the terra firme and white-sand plots, suggesting that coexistence of many moss species is favoured by high humidity. Moss species with fan life-forms preferred floodplains. On the other hand, liverwort species richness was highest in white-sand plots, which suggests that light incidence controls liverwort species-richness, perhaps more than humidity. All plots from the floodplain of the Caquetá River differed remarkably in species composition (of both mosses and liverworts) from the other landscape units. This may be due to the unique properties of this varzea system where, during yearly flooding events, soil, dead logs and stems are covered with a fresh layer of nutrient-rich fine silt, enhancing the surface for colonization and improving the conditions for productive bryophyte growth compared with elsewhere in the middle Caquetá area.     

Bryophyte Mats Inhibit Germination of Non-native Species in Burnt Temperate Native Grassland Remnants

Species-rich native grasslands in western Victoria, Australia, are often small, have a high perimeter to area ratio and are surrounded by non-native species. Few non-native species, however, have invaded them. A feature of species-rich grasslands is the presence of a bryophyte mat (composed of mosses and liverworts) that carpets the intertussock spaces. I assessed the role of these mats in plant invasions by sowing three non-native species (Briza maxima, Hypochoeris radicata, Plantago lanceolata) in replicated disturbed (mats removed) and undisturbed (mats intact) microsites at three grassland remnants (two recently burnt, one unburnt for 3 years) and followed seedling emergence, survival and growth for 5 months. Three native species were also sown for comparison. The rate of germination and total percent germination of non-native species were significantly enhanced at both burnt sites when the mat was disturbed. The large-seeded Briza maxima failed to germinate at both burnt sites in the absence of soil disturbance. The native species generally did not show a strong germination or growth response to soil disturbance in burnt areas. At the unburnt site, where monthly percent soil moisture was highest, final percent germination of the non-native and native species was greatest of any site in both microsites, and germination was not significantly affected by soil disturbance. Differences in the seed morphology of native and non-native species may play an important role in their ability to establish on bryophyte mats in moisture-limiting environments. Any activity that disrupts the mats in the frequently burnt, species-rich grassland remnants is likely to significantly enhance the germination and subsequent growth by non-natives. However, where burning is infrequent, germination of some non-native species may be expected, regardless of disturbance, although growth will likely be favoured in disturbed areas.     

苔藓植物化石的研究进展及应用

苔藓植物化石为探索植物界物种进化发展和重建古气候环境提供了重要的科学基础。该文通过大量的化石资料,结合当今前沿科学技术和热点问题,从苔藓植物起源分化及物种多样性分布、形态表现及分子进化及古气候环境的重建等方面对该领域的研究成果进行了综述和总结。(1)苔藓植物起源于早古生代的赤道低纬度区,至少在奥陶纪时期就有了苔类与藓类的分化;其物种多样性总体上呈增加趋势,并具有低纬度区物种多样性高的特点,但个别地质年代苔类与藓类的物种多样性具有明显差异,凸显了二者对气候环境的敏感程度不同。(2)苔藓植物具有缓慢的形态和分子进化,其中苔类的进化速率相对较快。(3)对于显生宙古气候环境的变迁及重建,苔藓植物化石自身的形态特征、孢粉学和古分子化石的分析、以及碳和氧同位素的测量均提供了重要的科学依据。针对苔藓植物化石研究中存在的问题,作者提出了自己的观点。     

Diversity of saproxylic bryophytes in old‐growth and managed beech forests in the central Balkans

Abstract

The diversity of saproxylic bryophyte species in beech forest stands from the wide region of the central Balkans (i.e. Serbia and Montenegro) was studied, and this study is the first of such a type in SE Europe. Comparison of preserved old‐growth and managed forests were made. Bryophyte species diversity is higher in primeval forest stands where the spectra of dead wood in various decaying stages of its dynamics are present. The ecological group of epixylic specialists is predominant among the bryophytes recorded. Threatened bryophyte species occur in old‐growth beech stands. The dead wood as habitat together with some other factors are extremely important for the surviving of epixylic bryophyte; so these species can be used as bioindicator bryophyte species of old‐growth or managed and structured forest ecosystems.     

New national and regional bryophyte records, 31

Abstract

Seasonal patterns in growth and dynamics of 13 epixylic bryophyte species (six liverworts and seven mosses) were studied within permanent plots in the ?ofínský prales old-growth forest in south-eastern Bohemia, Czech Republic. Transparencies were used to record positions and cover of bryophyte colonies within each plot twice a year (in spring and autumn) for periods of 1–4 years. Percentage growth of species populations and growth dynamics of their colonies were analysed in order to identify seasonality in their growth. Establishment of bryophyte colonies and their expansion, as well as pair-wise overgrowth of adjacent colonies of different species were compared in order to identify competitive interactions between bryophyte species. Bryophytes grew faster during the winter season (October–April) than during summer, and the various species did not differ in their percentage growth. The growth dynamics of bryophyte colonies was highly seasonal and the species differed in their dynamics pattern. The expansion and retreat (local extinction) of bryophyte species were greatest in winter, and the stability of colonies was, therefore, higher in summer. Interspecific competition among the species studied appears to be low.     

Disentangling the roles of chance,abiotic factors and biotic interactions among epiphytic bryophyte communities in a tropical rainforest (Yunnan,China)

• Epiphytes offer an appealing framework to disentangle the contributions of chance, biotic and abiotic drivers of species distributions. In the context of the stress-gradient theory, we test the hypotheses that (i) deterministic (i.e., non-random) factors play an increasing role in communities from young to old trees, (ii) negative biotic interactions increase on older trees and towards the tree base, and (iii) positive interactions show the reverse pattern.
• Bryophyte species distributions and abiotic conditions were recorded on a 1.1 ha tropical rainforest canopy crane site. We analysed co-occurrence patterns in a niche modelling framework to disentangle the roles of chance, abiotic factors and putative biotic interactions among species pairs.
• 76% of species pairs resulted from chance. Abiotic factors explained 78% of non-randomly associated species pairs, and co-occurrences prevailed over non-coincidences in the remaining species pairs. Positive and negative interactions mostly involved species pairs from the same versus different communities (mosses versus liverworts) and life forms, respectively. There was an increase in randomly associated pairs from large to small trees. No increase in negative interactions from young to old trees or from the canopy to the base was observed.
• Our results suggest that epiphytic bryophyte community composition is primarily driven by environmental filtering, whose importance increases with niche complexity and diversity. Biotic interactions play a secondary role, with a very marginal contribution of competitive exclusion. Biotic interactions vary among communities (mosses versus liverworts) and life forms, facilitation prevailing among species from the same community and life form, and competition among species from different communities and life forms.

     

Effects of plant leachates from four boreal understorey species on soil N mineralization, and white spruce (Picea glauca) germination and seedling growth

BACKGROUND AND AIMS: Natural regeneration of white spruce (Picea glauca) after disturbance has been reported to be very poor. Here a study was made to determine whether C compounds released from understorey species growing together with white spruce could be involved in this regeneration failure, either by (1) changing soil nutrient dynamics, (2) inhibiting germination, and/or (3) delaying seedling growth. METHODS: Foliage leachates were obtained from two shrubs (Ledum palustre and Empetrum hermaphroditum) and one bryophyte (Sphagnum sp.) with high phenolic compound concentrations that have been reported to depress growth of conifers in boreal forests, and, as a comparison, one bryophyte (Hylocomium splendens) with negligible phenolic compounds. Mineral soil from a white spruce forest was amended with plant leachates to examine the effect of each species on net N mineralization. Additionally, white spruce seeds and seedlings were watered with plant leachates to determine their effects on germination and growth. KEY RESULTS: Leachates from the shrubs L. palustre and E. hermaphroditum contained high phenolic compound concentrations and dissolved organic carbon (DOC), while no detectable levels of C compounds were released from the bryophytes Sphagnum sp. or H. splendens. A decrease in net N mineralization was determined in soils amended with L. palustre or E. hermaphroditum leachates, and this effect was inversely proportional to the phenolic concentrations, DOC and leachate C/N ratio. The total percentage of white spruce germination and the growth of white spruce seedlings were similar among treatments. CONCLUSIONS: These results suggest that the shrubs L. palustre and E. hermaphroditum could negatively affect the performance of white spruce due to a decrease in soil N availability, but not by direct effects on plant physiology.     

Enhanced sexual reproduction in bryophytes at high latitudes in the maritime Antarctic

Abstract

Contrary to the generally accepted hypothesis that bryophyte fertility decreases with increasing latitude and therefore climatic severity, a detailed study of bryophyte reproductive strategies at sites in the southern maritime Antarctic (68–72°S) has revealed that an unexpectedly high proportion of species is capable of producing sporophytes. Of the regional bryoflora, 43% (19 species; 17 mosses, 2 liverworts) in Marguerite Bay and 47% (17 species; 16 mosses, 1 liverwort) in Alexander Island are known to produce sporophytes, although the number fruiting at comparable latitudes on the colder and more arid Antarctic continent is less (33%). These numbers and proportions are much greater than were previously known at such relatively high southern polar latitudes. Sporophytes of several species are recorded for the first time within the Antarctic biome, while those of two liverworts (Lophozia excisa and Cephaloziella varians) are reported for the first time south of 62°S. High incidence of sporophyte production is attributed to locally favourable microclimatic conditions producing small-scale 'oases'. The large majority of fertile mosses are monoecious short acrocarps growing on rather calcareous soils. Spore production and size data for a number of species are compared with previously published studies of the same or closely related species from the northern maritime Antarctic and sub-Antarctic, but no consistent trends are revealed between species over the latitudinal gradient.     

Allelopathic effects of <Emphasis Type="Italic">Ceratiola ericoides</Emphasis> (Empetraceae) on germination and survival of six Florida scrub species

Allelopathic inhibition of germination by Florida scrub plants has been demonstrated in the greenhouse and lab, but not in the field. We studied the allelopathic effects of Florida rosemary (Ceratiola ericoides) roots, leaves, and litter leachates on field germination and three-month survival of six Florida scrub species, three habitat generalists (Lechea deckertii, Palafoxia feayi, and Polygonella robusta), and three rosemary scrub specialists (Hypericum cumulicola, Lechea cernua, and Polygonella basiramia). We used AIC and model averaging to evaluate support for a series of non-exclusive hypotheses. Species varied in germination (2.7–24.6%) and survival (39.2–71%) percentages, and in their sensitivity to leachates. Germination of scrub species was most negatively affected by leaf > root > litter leachates, although not all species followed the overall trend. Additional germination suppression by leachate combinations (relative to single leachates) was minimal. Sites did not vary in germination, but seedling survival did differ among sites. This study further documents the negative impact of Florida rosemary leachates on the germination of co-occurring plant species. Allelopathy may be partly responsible for bare sand gaps in Florida rosemary scrub, and therefore be one of the forces structuring Florida rosemary scrub ecosystems.     

Bryophyte Red List of Serbia and Montenegro

The bryophyte Red List of Serbia and Montenegro comprises 254 species (212 mosses and 42 liverworts). Serbia and Montenegro have 39.50% of threatened bryoflora. One moss species is considered as extinct (Encalypta serbica Katic). In the threatened categories there are 20 critically endangered (CR), 35 endangered (EN) and 100 vulnerable (VU) bryophyte species. Seventy-two species are considered to be of low risk, and 36 are too data deficient to place them in any category, but potentially with further investigation will enter one of the threat categories in the Red List.     

Mosses and liverworts show contrasting elevational distribution patterns in an oceanic island (Terceira,Azores): the influence of climate and space

Due to ongoing global changes, it is essential to establish a baseline record from which to determine future shifts in species distributions and community assembly patterns. In this context, we used digitised historical bryophyte distribution data along a 1021?m elevational gradient in Terceira Island (Azores) to determine how bryophyte species distribution varies with elevation and which spatial and climatic drivers contribute to this variation. We used ordinary least squares analysis to test for climatic and spatial data as explanatory variables for bryophyte richness and Mid-Domain Null simulations to assess the influence of spatial constraints on species distributions. Bryophyte richness follows a hump-shaped pattern, with mosses predominating in the first half of the gradient and liverworts in the second half. While moss richness did not correlate to any climatic variables and responded weakly to the presence of forest cover, liverwort presence was related to temperature, rainfall, humidity and the occurrence of native forest areas, suggesting that these plants are more sensitive to changes in their environment and can thus be used as better indicator species for climate change. Despite their inherent biases, our study shows that historically compiled data can be a valuable tool for preliminary assessment of macroecological patterns.     

PRELIMINARY INFERENCES OF THE PHYLOGENY OF BRYOPHYTES FROM NUCLEAR-ENCODED RIBOSOMAL RNA SEQUENCES

Ribosomal RNA sequences and cladistic analysis were used to infer a phylogeny for eight bryophyte taxa. Portions of the cytoplasmic large (26S-like) and small (18S-like) subunit ribosomal RNA genes were sequenced for three marchantioid liverworts (Asterella, Conocephalum, and Riccia), three mosses (Atrichum, Fissidens, and Plagiomnium), and two hornworts (Phaeoceros and Notothylas). Cladistic analysis of these data suggests that the hornworts are the sister group to the mosses, the mosses and hornworts form a clade that is sister to the tracheophytes, and the liverworts form a clade sister to the other land plants. These results differ from previous cladistic analyses based on morphology, ultrastructure, and biochemistry, wherein the mosses alone are sister group to the tracheophytes. We conclude that cladistic analysis of molecular data can provide an independent data set for the study of bryophyte phylogeny, but the differences between the molecular and morphological results are a topic for further investigation.     

Phytogeography of the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica

Aim Central America is a biogeographically interesting area because of its location between the rich and very different biota of North and South America. We aim to assess phytogeographical patterns in the bryophyte floras of oak forests and páramo of the Cordillera de Talamanca, Costa Rica. Location Tropical America, in particular the montane area of Cordillera de Talamanca, Costa Rica. Methods The analysis is based on a new critical inventory of the montane bryophyte flora of Cordillera de Talamanca. All species were assigned to phytogeographical elements on the basis of their currently known distribution. Absolute and percentage similarities were employed to evaluate floristic affinities. Results A total of 401 species [191 hepatics (liverworts), one hornwort, 209 mosses] are recorded; of these, 251 species (128 hepatics, one hornwort, 122 mosses) occur in oak forests. Ninety‐three per cent of all oak forest species are tropical in distribution, the remaining 7% are temperate (4%) and cosmopolitan (3%) species. The neotropical element includes almost 74% of the species, the wide tropical element (pantropical, amphi‐atlantic, amphi‐pacific) only 19%. A significant part of the neotropical species from oak forests are species with tropical Andean‐centred ranges (27%). As compared with bryophyte species, vascular plant genera in the study region are represented by fewer neotropical, more temperate and more amphi‐pacific taxa. Bryophyte floras of different microhabitats within the oak forest and epiphytic bryophyte floras on Quercus copeyensis in primary, early secondary and late secondary oak forest show a similar phytogeographical make‐up to the total oak forest bryophyte flora. Comparison of oak forest and páramo reveals a greater affinity of the páramo bryophyte flora to temperate regions and the great importance of the páramo element in páramo. Surprisingly, oak forests have more Central American endemics than páramo. Main conclusions (1) Providing first insights into the phytogeographical patterns of the bryophyte flora of oak forests and páramo, we are able to confirm general phytogeographical trends recorded from vascular plant genera of the study area although the latter were more rich in temperate taxa. (2) Andean‐centred species are a conspicuous element in the bryophyte flora of Cordillera de Talamanca, reflecting the close historical connection between the montane bryophyte floras of Costa Rica and South America. (3) High percentages of Central American endemics in the bryophyte flora of the oak forests suggest the importance of climatic changes associated with Pleistocene glaciations for allopatric speciation.     

广东黑石顶森林苔藓群落特征初探

通过对5个面积为2500m2的样地中树附生和地面苔藓植物的定量调查,初步研究了广东黑石顶森林中苔藓植物多样性特征。本次调查共发现苔藓植物50种,包括30种苔类和20种藓类。样地中树附生苔藓植物40种,地面苔藓植物24种,前者远高于后者。各样地中地面苔藓植物的种类为8-13种,盖度0.59%-1.12%,坡度及小生境的多样性对地面苔藓植物物种多样性及盖度均有很大影响;树附生苔藓植物12-20种,盖度0.63%-1.63%。结果显示30a的次生林中苔藓植物的物种丰富度及盖度均可恢复到与成熟阔叶林接近的水平。相似性分析表明,成熟阔叶林内苔藓植物种类组成与针阔叶混交林及30a的次生阔叶林差异不明显,但前者的苔藓植物群落结构与后者的差异较大。     

Estimation of stand-level leaf area for boreal bryophytes

Bryophytes dominate the carbon and nitrogen cycling of many poorly drained terrestrial ecosystems and are important in the vegetation-atmosphere exchange of carbon and water, yet few studies have estimated their leaf area at the stand scale. This study quantified the bryophyte-specific leaf area (SLA) and leaf area index (LAI) in a group of different-aged boreal forest stands in well and poorly drained soils. Species-specific SLA (for three feather mosses, four Sphagnum spp. and Aulacomnium palustre mixed with Tomentypnum nitens) was assessed by determining the projected area using a flatbed scanner and cross-sectional geometry using a dissecting microscope. The hemisurface leaf area was computed as the product of SLA and live biomass and was scaled by coverage data collected at all stands. Pleurozium schreberi dominated the spatial coverage, biomass and leaf area in the well-drained stands, particularly the oldest, while S. fuscum and A. palustre were important in the poorly drained stands. Live moss biomass ranged from 47 to 230 g m⁻² in the well-drained stands dominated by feather mosses and from 102 to 228 g m⁻² in the poorly drained stands. Bryophyte SLA varied between 135 and 473 cm² g⁻¹, for A. palustre and S. capillifolium, respectively. SLA was strongly and significantly affected by bryophyte species, but did not vary between stands; in general, there was no significant difference between the SLA of non-Sphagnum mosses. Bryophyte LAI increased with stand age, peaking at 3.1 and 3.7 in the well and poorly drained stands, respectively; this represented approximately 40% of the overstory LAI in the well-drained stands and 100–1,000% in the poorly drained stands, underscoring the important role bryophytes play in the water and carbon budgets of these boreal forests.     

Does the width of isolated ravine forests influence moss and liverwort diversity and composition?—A study of temperate forests in South Africa

The evidence of microclimatic edge effects through forest/open area ecotones is firm. How this affects the species composition near edges is less well understood and documented. In south-western South Africa shrub-dominated regularly burnt vegetation (i.e. fynbos) is the most common vegetation with indigenous temperate forests naturally occurring mostly in ravines on mountain slopes. The size and width of these forest patches varies considerably. In order to understand how the width of forest patches affects species composition of mosses and liverworts we investigated 20 forest patches of different width. In each forest patch we compiled a total species list, and collected data on selected environmental variables, for a plot of 10×20 m. No significant relationship was found between patch width and the number of either moss or liverwort species. However, the species composition of mosses (but not liverworts) changed along the gradient of patch width. The variation in number of species was large (8–29 mosses and 11–33 liverworts) among the plots and we propose that factors other than distance to the forest edge may be more important in structuring the bryophyte community of these ravine forests. One such factor that is likely to be important is the climatic difference among the plots. There is variation in features, such as precipitation, summer temperature, cloudiness, and the amount of fog, for which we do not have data but that could overrule the microclimatic differences due to edge effects.     

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.