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.     

A Bryophyte Flora of Berkshire and Oxfordshire

Abstract

In temperate North American forests, collections were made of myxomycetes (plasmodial slime molds) for which bryophytes served as the primary substrate for fruiting. Most associations among the 52 myxomycete species and 55 bryophyte species, represented in these collections, appeared to be coincidental but two species of myxomycetes (Barbeyella minutissima and Lepidoderma tigrinum) were evidently bryophilous, preferring leafy hepatics on rotten coniferous logs. Bryophytes obviously provide exposed surfaces convenient for myxomycete sporulation. Whether bryophytes also sustain the feeding phases of the myxomycete life-cycle (swarm cells, myxamoebae, and plasmodia) is not yet known.     

Ordination of epiphytic bryophyte communities in a wet-temperate coniferous forest,South-Coastal British Columbia

Three coniferous tree species (Pseudotsuga menziesii, Tsuga heterophylla, and Thuja plicata) at Capilano Canyon, British Columbia, were studied for their epiphytic communities. Quantitative data were obtained for fourteen bryophyte species by sampling at heights of 0.5 m, 1 m and 2 m above ground level. Scapania bolanderi is an abundant and widely distributed species; Dicranum fuscescens, Bazzania denudata, Lepidozia reptans and Hypnum circinale are also common. An environmentally uniform study area was chosen to minimize the effects of factors other than bark and elevation on the distribution of epiphytes. In general, tree species are similar in terms of their epiphytic assemblages, but ordination methods using quantitative data exposed compositional variation that may be explained by differences in microclimate and bark-type. In this respect, the results point the way to further studies to examine epiphyte associations in closer detail, and to relate these associations to specific factors in the microenvironment.Nomenclature for vascular plants follows Hitchcock & Cronquist (1973); bryophyte nomenclature follows Schofield (1976) for mosses and Stotler & Crandall-Stotler (1977) for hepatics.We thank Dr. W. B. Schofield for aiding in bryophyte identification.     

Beech versus spruce deadwood as forest microhabitat: does it make any difference to bryophytes?

The presence data of 153 bryophyte species, recorded on beech (BDW) and spruce (SDW) deadwood in the managed forests within a montane watershed, were analysed by controlling for different plot size, substrate pH and elevation. Only three pleurocarpous species, which were probably remnant epiphytes from live wood, showed relative specificity for BDW, but none to SDW. The few species displaying preference for BDW or SDW were generally true epixylic, mosses and hepatics, respectively. The chances of occurrence of epiphytic (early successional) and epixylic (mid-late successional) taxa on either deadwood species increased and respectively, decreased with the substrate pH. There was no consistent relationship between bryophyte preference to BDW/SDW and elevation. Neither total nor threatened species richness was related to deadwood species identity. However, at 100 cm² resolution and within certain extension range, the cumulative number of bryophyte species was significantly higher on BDW than on SDW, despite no significant difference in species pool size. Threatened species richness increased monotonically towards lower elevations, suggesting that valley bottoms might act as refugia with high air humidity. The number of threatened species and in particular, the presence of Lophozia ascendens, were good predictors of total bryophyte richness regardless of deadwood species identity.     

Bryophyte Species Diversity in Secondary Forests Dominated by the Introduced Species Spathodea campanulata Beauv. in Puerto Rico

The introduced tree species Spathodea campanulata (Bignoniaceae) forms novel forests in Puerto Rico, these having emerged after the abandonment of fields in the mid‐20th century and resulting in forests with a new species composition. We assessed bryophyte species richness in these novel forests and sought correlations with geological substrate, past land use, forest edge and patch area, forest structure, elevation, microhabitat diversity, tree species richness, and microclimatic conditions. Transects were established (edge and forest interior) in nine moist forest patches dominated by Spathodea in north‐central Puerto Rico. These Spathodea forest patches ranged from 0.6 to 9 ha. ANOVA, Chi‐square, correlation, and cluster analyses were used in data analyses. We found 57 bryophyte species. There was a significant difference in bryophyte richness among patches. Those on karst exhibited highest bryophyte richness due to microhabitat diversity, past land use, and shorter hydroperiods. Alluvial sites scored lowest in bryophyte species richness, and forest structure was important for bryophyte communities on these sites. Significant differences in temperature, relative humidity, and light intensity were observed between edge and forest interior. These appeared important for establishing bryophyte species cover but not richness and composition. Microhabitat diversity, patch area, and forest age were more related to bryophyte species richness than elevation, exposed edge, and tree species richness, regardless of geologic substrate. Collectively, Spathodea patches were similar to mature forests on the Island with respect to bryophyte species richness and composition. Novel Spathodea forests have conservation value due to their habitat suitability for bryophyte communities.     

A contribution to the bryophyte flora of southern Turkey

An initial investigation has been undertaken into the bryophyte flora in the region of southern Turkey that includes the Bolkar and Amanus Mountains and Mersin villages (square C13 on the grid-map of Henderson & Prentice, 1969). Some 29 moss taxa and 3 species of hepatics are recorded for this bryologically under-explored area.     

Fissidens monguilloni Thér. in England

Abstract

The 1996 Chagos Expedition provided an opportunity to study the archipelago's bryophyte flora: 25 of the 55 islands were ecologically investigated, some in more detail than others, and lists and representative collections of mosses, hepatics and other cryptogamic plants were assembled for many of them. Although the islands have a low biodiversity for most of these groups, as would be expected in terms of their relatively young age, remoteness and small terrestrial surface areas, those organisms that are present are often found in abundance and play significant ecological roles. Where the status of taxa on islands has been determined, there appears to be a good correlation between bryophyte biodiversity and island size. Epiphytic bryophytes were not host specific and no epiphyllous species was observed. Biogeographic details are provided, where possible, for the 19 taxa recorded, of which four are new to the Indian Ocean and seven are new to the Chagos Archipelago. A new taxon, Cololejeunea planissima var. chagosensis Pócs var. nov. is described, and notes and illustrations of a probably new Lejeunea species are provided.     

Species richness in boreal swamp forests of SE Norway: The role of surface microtopography

Question: What is the relative importance of environmental gradients and surface microtopography (variation in vertical level within sampling units) for fine‐scale plant species richness in Picea abies swamp forests? Location: 11 swamp forests in SE Norway. Methods: We recorded species richness (number of species of vascular plants, mosses, Sphagnum and hepatics), depth to water table, soil base status and vertical range (microtopographic relief) in 2400 microplots, (each 1/16 m²), in 150 1‐m² plots in the 11 swamp forests. Relationships between species richness and environmental predictors were modelled by GLMM. Results: Moss and hepatic species richness increased with increasing microtopographic relief, most strongly for wet acid sites, in which similar trends were also found for Sphagnum. Relief was a poor predictor of vascular plant species richness. Conclusions: Microtopographic relief is a good predictor of local species richness in Picea abies swamp forests, partly because larger vertical variability means higher within‐plot habitat diversity with respect to the wet‐dry gradient, and partly because qualitatively new microhabitats associated with steep slopes are added in drier sites. The relationship between species richness and microtopographic relief is context dependent, differing in complex ways among species groups and among sites with different environmental conditions.     

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.     

Changes in forest understorey vegetation in Norway related to long‐term soil acidification and climatic change

Question: Does the understorey vegetation of Norwegian boreal forests change in relation to broad‐scale, long‐term changes? Location: Norway. Methods: Permanently marked 1‐m² vegetation plots from 17 monitoring reference areas in forests dominated by Picea abies (11 areas, 620 plots) and Betula spp. (six areas, 300 plots) were analysed twice, at the start in 1988–1997 and 5 yr later (1993–2002). Species subplot frequency data were analysed separately for each area by univariate and multivariate statistical methods; 5‐yr changes in single species abundances, species number per plot and species composition were tested. Results: Two distinct patterns of change were found: 1. Abundance of several vascular plant species decreased in SE Norwegian Picea forests, most noticeably of species with a preference for richer soils, such as Oxalis acetosella. 2. Abundance of many bryophyte species as well as bryophyte species number per plot increased in forests of both types over most of Norway. Conclusions: The pattern of vascular plant changes is probably a time‐delayed response of long‐lived, mainly clonal, populations to acidified soils resulting from deposition of long‐distance airborne pollutants. The pattern bryophyte changes, with reference to the close link between climatic conditions for growth and abundance changes for Hylocomium splendens established in previous demographic studies, is related to climatic conditions favourable for bryophyte growth. We conclude that many forest understorey plants are sensitive indicators of environmental change, and that the concept used for intensive monitoring of Norwegian forests enables early detection of changes in vegetation brought about by broad‐scale, regional, impact factors.     

Population biology of the clonal moss Hylocomium splendens in Norwegian boreal spruce forests. 5. Vertical dynamics of individual shoot segments

Demographic information was obtained for 12 528 mature segments (for which dry weight was estimated and vertical position in the bryophyte carpet recorded) and 3109 regenerated growing points for the perennial clonal moss Hylocomium splendens, recorded in Norwegian boreal spruce forests during a 6‐year period. Branching frequency varied with vertical position in the bryophyte carpet. Termination risk (probability of producing no offspring) was highest (44%) for buried segments, lowest (12%) for segments at intermediate vertical positions, and also high (26%) for emergent segments (due to increasing exposure to external mortality agents). Segment size increased from low levels in the bryophyte carpet to a maximum ca 2–10 mm below the top of the bryophyte carpet. This intermediate level was interpreted as the optimal compromise between incoming radiation (attenuating downwards) and microclimatic moisture conditions (improving downwards). Size‐corrected fitness, the number of offspring emerging from a mature segment within one year after maturation after allowance for differences in size, was lower for buried and emergent segments than for segments at intermediate positions. Small emergent segments were apparently liable to suffer from vitality reductions due to desiccation. The vertical position of a daughter segment depended on that of its parent segment, but also showed considerable stochastic variation. Burial acted as a strong sink for small segments regardless of vertical position. No evidence was found for species‐specific differences in the way pleurocarpous bryophytes interact, but reduced vertical mobility of H. splendens when growing among acrocarps indicated that growth‐form is an important determinant of bryophyte interactions. Evidence was found for vertical layering of the bryophyte carpet according to dominant type of interactions among individuals: none (environmental stress) above and at top, facilitation [a (+, +) interaction] at intermediate levels because of favourable water relationships in closed stands, and amensalism [a (0, ?) interaction] from higher‐situated segments that deprive lower‐situated segments access to light at lower relative levels. The intensity of amensalism increased downwards in the bryophyte carpet as indicated by a reinforced size hierarchy. The tendency for small H. splendens segments to become buried and lost from the population by amensalism is likely to represent a general mechanism for interactions between bryophyte species and succession in bryophyte‐dominated stands. Population effects of climatic and local environmental factors (favourability vs stress), disturbance and apparently random events are discussed with reference to their impact on the relative sizes of subpopulations acting as sources (due to facilitation) and sinks (due to amensalism).     

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.     

Ecophysiology in Relation to Exposure of Pendant Epiphytic Bryophytes in the Canopy of a Tropical Montane Oak Forest1

We investigated ecophysiological and morphological traits affecting light and water use of four commercially important pendant epiphytic bryophyte species in a montane oak‐bamboo forest in Costa Rica. Predictions based mostly on ecophysiological studies of temperate bryophytes and vascular plants were experimentally tested on tropical montane bryophyte species ranked on the basis of their distributions in the canopy from the most protected (1) to the most exposed sites (4): (1) Phyllogonium viscosum; (2) Pilotrichella flexilis; (3) Dendropogonella rufescens; and (4) Frullania convoluta. As predicted, highly exposed species tended to have higher light saturation and compensation points, higher dark respiration rates, more chlorophyll, higher chlorophyll a:b ratios, and higher nitrogen concentrations. Contrary to predicted trends, exposed‐site species had lower water contents at full saturation; there were no detectable differences among species in the rate of water loss. Rates of carbon gain in all species reached asymptotes with increasing water content, but the moisture compensation point for carbon uptake of the moss D. rufescens was substantially higher than in the other species. Observed patterns result from interactions among processes related to water storage and transport operating at different scales and are determined by various morphological traits including the density, size, and disposition of phylloids, as well as by whole‐clump architecture.     

Bryophyte diversity and community structure on thatched roofs of the Holnicote Estate,Somerset, U.K.

Abstract

We investigated patterns of bryophyte species richness and community structure, and their relation to roof variables, on thatched roofs of the Holnicote Estate, South Somerset. Thirty-two bryophyte species were recorded from 28 sampled roofs, including the globally rare and endangered thatch moss, Leptodontium gemmascens. Multiple regression analyses revealed that thatch age has a highly significant positive effect on the number of species present, accounting for nearly half the observed variation in species richness after removal of outliers. Aspect has a slight and marginally significant effect on species diversity (accounting for an additional 6% of variation), with north-facing samples having slightly more species. Age also has a significant impact on total bryophyte cover after removal of outlying observations. TWINSPAN analysis of bryophyte cover data suggests the existence of at least five discrete communities. Simple Discriminant Analyses indicate that these communities occupy different ecological subspaces as defined by the measured roof variables, with pitch, aspect and thatch age emerging as especially significant attributes. Contingency Analysis indicates that some communities are disfavoured by water reed as compared to wheat straw. The findings are significant for understanding the structure of bryophyte communities, for evaluating the effect of bryophyte cover on thatch performance, and for conservation of thatch communities, especially those harbouring rare species.     

Species introduction through seeds from an old,species‐rich hay meadow: Effects of management

Abstract. A local seed mixture from plants growing in a species‐rich, traditionally managed hay meadow site at Varaldsoy, Hardanger, western Norway, where many endangered hay meadow species of the region are growing, was sown in a newly harrowed experimental field 1 km from the source meadow in order to increase the habitat area for the endangered species. Of 25 endangered species recorded in the source meadow, only one (Holcus lanatus) was present in the target meadow. After sowing, 16 of the endangered species in addition to Holcus lanatus were recorded in the new site. Six species were only present in sown plots and seven others were more frequent there, while three species might have arrived by chance or originated from the seed bank. Replacing the traditional management regime, including one late cut and grazing in spring and in autumn, with three cutting times and the creation of gaps in the sward, resulted in a higher number of endangered species in plots which were only cut, possibly because the grazing was too intensive in the small enclosures.     

皖西大别山北坡苔藓植物区系

该研究于2019-2021年4次对安徽大别山北坡进行苔藓植物野外考察和标本采集,对采集到的1 517份标本鉴定以及文献整理,统计分析该区苔藓植物物种组成、区系地理组成;并与华东、华中、华北等地的苔藓植物区系地理组成进行比较,以明确大别山北坡苔藓植物区系的归属问题。结果表明：(1)安徽大别山北坡共有苔藓植物80科180属482种(包括种以下单位);该次调查增加了12科40属206种,其中安徽省新记录种9个。(2)发现受威胁苔藓植物2种——滇西高领藓(Glyphomitrium minutissimum)属于极危(CR),瘤根真藓(Bryum bornholmense)属于易危(VU)。(3)大别山北坡苔藓植物区系拥有3个世界性单种科,14个世界性单种属,且有34个世界性寡种属(2～6种),该区单种属和寡种属(48属)占该区总属数的26.67%,表明该区苔藓植物起源古老。(4)大别山北坡苔藓植物区系地理成分复杂多样,以北温带成分(26.43%)和东亚成分(24.89%)为主,热带亚洲成分(14.54%)次之。(5)安徽大别山北坡苔藓植物区系与华东(西天目山、清凉峰、阳际峰)区系的关系最为密...     

Sporophytes of Rhodobryum laxe-limbatum (Ochi) Iwats. et Kop.

Abstract

A check-list of the bryophyte species of the Grand-Duchy of Luxembourg is given and briefly discussed. An appendix deals with infraspecific taxa. The total number of species is 517 (122 hepatics and 395 mosses); taking into account some acceptable infraspecific taxa, this small country has at least 538 different bryophytes.     

Comparisons of photosynthesis‐related traits of 27 abundant or subordinate bryophyte species in a subalpine old‐growth fir forest

Bryophyte communities can exhibit similar structural and taxonomic diversity as vascular plant communities, just at a smaller scale. Whether the physiological diversity can be similarly diverse, and whether it can explain local abundance patterns is unknown, due to a lack of community‐wide studies of physiological traits. This study re‐analyzed data on photosynthesis‐related traits (including the nitrogen, phosphorus and chlorophyll concentrations, photosynthetic capacities, and photosynthetic nutrient use efficiencies) of 27 bryophyte species in a subalpine old‐growth fir forest on the eastern Tibetan Plateau. We explored differences between taxonomic groups and hypothesized that the most abundant bryophyte species had physiological advantages relative to other subdominant species. Principal component analysis (PCA) was used to summarize the differences among species and trait values of the most abundant and other co‐occurring subdominant species. Species from the Polytrichaceae were separated out on both PCA axes, indicating their high chlorophyll concentrations and photosynthetic capacities (axis 1) and relatively high‐light requirements (axis 2). Mniaceae species also had relatively high photosynthetic capacities, but their light saturation points were low. In contrast, Racomitrium joseph‐hookeri and Lepidozia reptans, two species with a high shoot mass per area, had high‐light requirements and low nutrient and chlorophyll concentrations and photosynthetic capacities. The nutrient concentrations, photosynthetic capacities, and photosynthetic nutrient use efficiencies of the most abundant bryophyte species did not differ from co‐occurring subdominant species. Our research confirms the links between the photosynthesis‐related traits and adaptation strategies of bryophytes. However, species relative abundance was not related to these traits.     

Evidence of targeted consumption of mosses by birds in sub‐Antarctic South America

Bryophyte consumption is uncommon among bird species globally and is often presumed incidental. We sought to determine whether herbivorous bird species of the high Andes, including the white‐bellied seedsnipe (Attagis malouinus) and Chloephaga geese (C. picta and C. poliocephala), consume bryophytes, and if so, how frequently. We collected 26 seedsnipe and 22 goose droppings from alpine and sub‐alpine habitats of Navarino Island, Chile and examined their contents for bryophyte diaspores. We detected bryophyte fragments in 84.6% and 90.9% of seedsnipe and Chloephaga goose faecal samples, respectively. We also extracted DNA from three bryophyte fragments isolated from goose droppings and sequenced three chloroplast loci for each sample. We inferred through a barcoding analysis that at least one species of Chloephaga goose consumes Polytrichum strictum and Notoligotrichum trichodon. The composition of 11 collected goose droppings was >50% Polytrichaceae bryophyte fragments, suggesting that at least one Chloephaga goose species foraged deliberately on moss species of this family. These new observations suggest that bryophytes are part of the diet of some high Andean birds and that birds might disperse bryophytes internally – via endozoochory – in the sub‐Antarctic.     

The bryophyte diaspore bank and its role after small-scale disturbance in a boreal forest

Abstract. The composition of the bryophyte diaspore bank in an old-growth Picea abies forest was studied before and after experimental disturbance of forest-floor patches. 40 species, both hepatics and mosses, emerged from spores, and probably also from gemmae and moss fragments, in greenhouse cultivations of soil samples. The three most abundant taxa were Pohlia nutans, Sphagnum spp. and Polytrichum commune/longisetum. Initially, the number of species from the diaspore bank in mineral soil, 9.9 species/sample, was higher than that in humus, 6.6. Four years after the disturbance took place, the reverse was found. Several species that are typical colonisers of disturbed soil were very common in the diaspore bank. By contrast, some of the most abundant forest floor species appeared to be absent. The diaspore bank of bryophytes appears to play a role similar to that of the seed bank in vascular plants: (1) it allows species to survive unfavourable periods (temporal dispersal); (2) it facilitates rapid colonisation after disturbance; (3) it influences the post-disturbance species composition and diversity. The results emphasise that the understanding of vegetation succession patterns is, to a large extent, dependent on quantitative data on the relative importance of spatial and temporal dispersal.