Recent Bryological Literature, 58

Abstract

Recent field work in Mexico reveals fifteen cleistocarpic moss species in the Neovolcanic Belt, several of which represent new state records for Hidalgo, Michoacán, Puebla or Tlaxcala. The distribution of these species suggests that they have been able to cross the Neovolcanic Belt, but their method of dispersal remains obscure. Local dispersal may be enhanced by forest disturbance, agriculture, grazing and/or by their ability to shed the capsules with ease.     

Phylogeography and speciation of the morphologically variable, widespread species Bufo valliceps, based on molecular evidence from mtDNA

The widespread, lowland toad Bufo valliceps has an unusual distribution in North and Middle America that straddles two major biogeographic areas; previous morphological studies of this species suggested the existence of two species. We used mitochondrial DNA sequences to examine the phylogeography of this species and discovered the existence of two distinct clades. We recognize these as two species: B. valliceps and B. nebulifer. These molecular data support morphological data from previous studies. Our results show low levels of molecular variation in a morphologically uniform temperate species (B. nebulifer) and high levels of molecular variation in a morphologically variable tropical species (B. valliceps), providing an example of molecules matching morphology. Two biogeographic hypotheses are tested to explain the current distribution of these species, based on a calibrated rate of evolution and the percent sequence divergence between the two species. A more recent Pleistocene dispersal event, followed by vicariance associated with rising sea level, is rejected in favor of an earlier Miocene-Pliocene vicariant hypothesis associated with the formation of the Trans-Mexican Neovolcanic Belt.     

Identifying habitat linkages to maintain connectivity for corridor dwellers in a fragmented landscape

Anthropogenic habitat fragmentation typically precedes conservation planning; maintaining remaining linkages among core habitat areas can thus become a key conservation objective. Identifying linkages for dispersal and ensuring those linkages have long-term protection and management are challenging tasks for wildlife managers. These tasks can be especially daunting for smaller species with low mobility, termed corridor dwellers, which must maintain sustainable populations within corridors. Between May 2007 and June 2009, we collected occurrence locations for a corridor dweller, the Palm Springs pocket mouse (Perognathus longimembris bangsii), from museums, previous research, and our own field sampling. We used those data to model their suitable niche space and then identify suitable linkages between proposed conservation areas. We used a partitioned Mahalanobis D² statistic to create a spatially explicit niche model describing the distribution of a suitable niche space, and we validated the model statistically, with live trapping and with burrowing owl (Athene cunnicularia) diets. Our model identified soil characteristics, topographic ruggedness, and vegetation as variables delimiting Palm Springs pocket mouse habitat; sand content of the soils was an especially important characteristic. Our historic distribution model identified 120,000–90,000 ha as historically potential Palm Springs pocket mouse habitat; roughly 39% of that has been lost to more recent development. Most of the remaining suitable habitat occurred in the northwestern portion of the valley. We modeled habitat within core reserves as well as within proposed linkages between those reserves as having high similarity to known occupied habitats. Live trapping in areas with high (≥0.95) Habitat Suitability Index (HSI) values resulted in captures at 66% of those locations and, along with burrowing owl diets, refined a qualitative model as to what constituted a suitable Palm Springs pocket mouse corridor. While most corridor analyses have focused on mobile species which may traverse corridors in hours, days, or weeks, linkages for corridor dwellers must include habitat for sustaining multi-generational populations. This requires evaluating whether continuous suitable habitat exists within proposed corridors. Our research demonstrates how niche modeling can provide a landscape-scale view of the distribution of suitable habitat to evaluate conservation objectives for connectivity. © 2011 The Wildlife Society.     

On Barbula munyensis R.S.Williams (Pottiaceae), an overlooked species from the tropical montane forest of South America

Abstract

Barbula munyensis R.S.Williams, a neglected moss species known only from the type material collected in central Peru in 1923, is newly reported for the moss floras of Argentina and Bolivia. The species is described, illustrated, mapped, and its habitat preferences are provided. Diagnostic characters and differentiation from some closely related South American taxa, with which it may be confused, are discussed. Barbula orizabensis Müll.Hal., a similar taxon, is excluded from the South American moss flora.     

Experiment meets biogeography: plants of river corridor distribution are not more stress tolerant but benefit less from more benign conditions elsewhere

Aims Factors limiting distributions of species are fundamental to ecology and evolution but have rarely been addressed experimentally for multiple species. The conspicuous linear distribution patterns of plant species confined to river corridors in the Central European lowlands constitute an especially long-standing distribution puzzle. We experimentally tested our novel hypothesis that the tolerance of species to river corridor conditions is independent of the degree of confinement to river corridor habitats, but that species not confined to river corridors are better able to take advantage of the more benign non-river corridor conditions.Methods We grew 42 herbaceous species differing in their confinement to river corridors in a common garden experiment on loamy soil typical for river corridor areas and sandy soil typical for non-river corridor areas, and with and without a flooding period. For a subset of species, we grew plants of both river corridor and non-river corridor origin to test for adaptation to river corridor conditions.Important findings Species more confined to river corridor areas benefited less from the more benign non-flooded and non-river corridor soil conditions than species of wider distributional range did. For subsets of 7 and 12 widespread species, the response to flooding and soil origin, respectively, did not differ between plants from river corridor sites and plants from other sites, suggesting that the habitat tolerance of widespread species is due to phenotypic plasticity rather than to local adaptation. Overall, we found clear support for our novel hypothesis that species not confined to river corridors are more able to take advantage of the more benign non-river corridor conditions. Our study provides a general hypothesis on differences between species confined to stressful habitats and widespread species out for test in further multispecies comparative experiments.     

厦门湾中华白海豚潜在生态廊道识别及人类活动干扰评估

生态廊道具有维持或恢复生态连通性的功能,对于连接生物栖息地、保护物种多样性具有重要意义。现有的生态廊道研究主要集中于陆地,而海洋生态系统具有水体广泛连通、缺乏直观的景观斑块等特点,导致海洋生态廊道的研究成为长期以来的科学难题。以栖息地位于厦门湾的国家一级保护动物中华白海豚(Sousa chinensis)为对象,尝试基于物种分布模型和最小成本路径分析法建立海洋生态廊道的识别方法。研究采用物种分布模型识别厦门湾内中华白海豚的适宜生境分布区和节点,并利用模型产出的生境适宜性结果生成海洋中的阻力表面,模拟计算节点与节点间在阻力表面上的最小成本路径,从而生成物种扩散网络。研究结果显示,厦门湾中华白海豚的分布主要受到航道距离、到岸线距离和叶绿素浓度三项因素的影响,主要适宜生境位于西海域至九龙江口和大嶝海域。潜在的核心生态廊道面积93.19km²,次级生态廊道面积170.41km²,九龙江口-鼓浪屿南侧-黄厝-大小嶝岛沿线可能是厦门湾中华白海豚的主要迁移路线。在此基础上,从用海空间重叠和桥梁影响两方面开展了人类活动对廊道的干扰评估。评估结果显示旅游活动和...     

Comparative ecology of a lowland and a subalpine species of Mnium in the northern Rocky Mountains

Factors that set the altitudinal limits of plants have been relatively well explored for many land plant groups, but not for bryophytes. Bryophytes typically represent a significant portion of alpine floras with many species specific to highland systems. Differences between highland and lowland bryophytes have been underinvestigated. In the present study spanning three growing seasons, a subalpine and a lowland moss were both reciprocally planted as apical fragments and transplanted as adults between sites at 1400 m and 2000 m in the Front Ranges of the Rocky Mountains, Alberta. The lowland species, Mnium spinulosum, was less tolerant of conditions at 2000 m than the subalpine species, M. arizonicum, was to conditions at 1400 m. In particular, M. spinulosum had lower establishment from both apical fragments and spores at higher elevation sites. Both species had significantly lower establishment during the abnormally cold growing season of 1999, but fragments of M. arizonicum were better able to adjust their investment in establishment. The effect of a dominant feather moss, Hylocomium splendens, on establishment and transplant health was tested for M. arizonicum. Establishment of M. arizonicum was lower in Hylocomium mats than on bare humus regardless of site elevation suggesting allelopathy; however, stem survival in adult transplants was higher in Hylocomium mats than in Mnium dominated microsites at the higher elevation suggesting facilitation. Competition, rather than a lack of physiological plasticity, probably determines the lower elevation limit of the subalpine moss, while poor establishment ability at low temperatures accounts for the upper elevation limit of the montane moss. This revised version was published online in June 2006 with corrections to the Cover Date.     

Species richness of moss landscapes unaffected by short-term fragmentation

Theory predicts that habitat fragmentation and varying corridor length and width will affect animal populations in adjoining habitat patches due to varying migration rates. Previous work on the moss/microarthropod microcosm showed that connecting moss patches with moss corridors maintained species richness and individual species abundance. By contrast, in this study there was little evidence for differences in species richness between landscapes of varying connectivity and corridor length and width. The γ diversity, the cumulative species richness of entire connected systems, followed the same pattern. Similar non-significant results were obtained for species abundance. Contrary to a previous study, I found no evidence that populations of predators were more affected by fragmentation than non-predators. Since this experiment ran during temperate environmental conditions and the previous experiments ran during more extreme conditions, I hypothesise that corridors may be more useful in reducing species extinction during extreme environmental conditions.     

Phylogeny of Chaetanthera (Asteraceae: Mutisieae) reveals both ancient and recent origins of the high elevation lineages

Penalized likelihood analysis of previously published chloroplast DNA (cpDNA) ndhF sequences suggests that the central-southern Andean genus Chaetanthera diverged ca. 16.5 million years (my) ago, well before the uplift of the Andes to their present heights. Penalized likelihood analysis based on new nuclear ribosomal DNA (rDNA) internal transcribed spacer (ITS) sequences indicates that the most relictual lineages occupy high elevation Andean habitats that did not exist until some 10my later. This result is contrary to the expectation that younger habitats should be occupied by phylogenetically younger lineages. The results are interpreted with respect to the development of aridity in lowland habitats during the Miocene and Pliocene, which presumably extinguished the lowland relatives of the high elevation taxa or, in effect, forced them upwards in search of adequate moisture. As the more northerly lineages were being displaced upward, others diversified in the mediterranean-type climate area of central Chile, giving rise to additional high elevation taxa again, at an early date, as well as lowland taxa. Some species of Chaetanthera from lowland central Chile appear as the phylogenetically youngest taxa, suggesting secondary adaptation to lowland aridity. At the same time, at least two high elevation species, Chaetanthera peruviana and Chaetanthera perpusilla, appear to have been derived recently from a lower elevation ancestor, while some middle to low elevation taxa seem to have evolved recently out of a high elevation complex. The results suggest that the younger high elevation habitats have served as both "cradle" and "museum" for Chaetanthera lineages.     

Moss endemism in the Mexican flora

Although the moss flora of Mexico consists of nearly 1000 species, only 77 are endemic. The country has many poorly collected or unexplored areas, but the number of endemic mosses is not expected to undergo a substantial increase; percent endemism has in fact decreased with taxonomic revisions and monographs and better exploration in other countries. Literature and herbarium records (n?=?584) were used to obtain an updated list of endemic mosses and their state distribution in Mexico. Cluster analysis and mapping indicate that there are three main areas of endemism: Lowland areas in various states, the mountain area along 19–20°N lat., and the highlands in Oaxaca and Chiapas. Similarity by province shows that Trans-Mexican Volcanic Belt, Sierra Madre Oriental, Chihuahuan Desert, and Sierra Madre del Sur have the highest numbers of endemic species. Five monotypic genera and 76 species (including two infraspecific taxa), many of which have comparatively narrow geographical ranges, suggest that speciation is recent, that species have had little time to disperse, and were formed by Pleistocene environmental climates in the highlands; older speciation may be represented by widespread disjunct species that still are found in the highlands of Mexico.     

Wide corridors with much environmental heterogeneity best conserve high dung beetle and ant diversity

Landscape ecological networks (ENs) consist of landscape-scale conservation corridors that connect areas of high natural value within a production mosaic with protected areas (PAs). In South Africa, ENs have been implemented on a large spatial scale to offset the negative impacts of plantation forestry on indigenous grasslands. We focus on corridor width as a factor for conserving dung beetle and ant diversity within an EN. We also investigate the importance of natural environmental heterogeneity (elevation, vegetation type) and habitat quality (soil hardness, invasive alien plant density). We sampled dung beetles and ants in 30 corridors of different sizes, and at ten sites in a nearby PA. In addition, we also analysed dung beetles according to their feeding guild. Tunnelling dung beetle species richness increased with corridor width. Rolling dung beetle species richness was higher in the PA than in the corridors of the EN. The dung beetle assemblage within the EN differed from that within the PA. Corridors of various widths differed in ant composition but not in species richness. Furthermore, the PA and the EN differed in environmental variables, which contributed to differences in dung beetle species richness and assemblage composition. Within the EN, environmental heterogeneity across the landscape was more important than corridor width for driving species diversity of both dung beetles and ants. When planning future ENs, wide corridors (>280 m) that encompass as much natural heterogeneity across the landscape as possible will best conserve the range of local insect species.     

Elevational gradients in species richness and endemism of selected plant groups in the central Bolivian Andes

I analyzed the distribution of Acanthaceae, Araceae, Bromeliaceae, Cactaceae, Melastomataceae, and Pteridophyta in 62 vegetation plots of 400 m² along an elevational transect between 500 m and 2450 m, and at a nearby lowland site in western Santa Cruz department, Bolivia. These groups were selected because they are physiognomically distinctive, have high species numbers, are comparatively easy to identify, adequately reflect overall floristic relationships, include a wide range of life forms, and are small. The transect was located in the Tucumano-Boliviano biogeographic zone and included drought-deciduous (<850–1000 m), mixed evergreen (850–1000 m to 1800 m), and evergreen Podocarpus-dominated (>1800 m) forests. Elevational patterns of species richness were group-specific and probably related to the ecophysiological properties of each group. Species richness in Pteridophyta and Melastomataceae was correlated with moss cover (i.e., humidity), with elevation (i.e., temperatures) in Acanthaceae and epiphytic Bromeliaceae, with potential evapotranspiration (i.e., ecosystem productivity) in Araceae, and with light availability at ground level in terrestrial Bromeliaceae and Cactaceae. Community endemism generally increased with elevation, but showed a maximum at 1700 m for terrestrial Pteridophyta, and a nonsignificant decline for epiphytic Bromeliaceae and Cactaceae. Endemism was higher for terrestrial than for epiphytic taxa, and was lower among Pteridophyta compared to all other groups, reflecting different dispersal ability among taxonomic and ecological groups. Elevational zonation, tested against a null-model of random distribution of elevational limits, revealed a significant accumulation of upper and lower elevational range boundaries at 900–1050 m and at 1500–1850 m, corresponding to the elevational limits of the main physiognomic vegetation types.     

The ranging patterns of elephants in Marsabit protected area,Kenya: the use of satellite‐linked GPS collars

We investigated the ranging patterns of elephants in the Marsabit protected area, north eastern Kenya, to ascertain the range of bachelor and female family herds in different seasons, and to identify corridor and noncorridor areas. Data were acquired for five bachelor and four female family herds equipped with satellite‐linked geographical positioning system collars, and monitored from December 2005 to December 2007. Distinct dry (about 260 km²) and wet seasons (about 910 km²) ranges were observed, with connecting corridors (north‐eastern corridor: about 90 km long, about 2‐7 km wide; southern corridors: about 10‐20 km long, about 2‐3 km wide). The dry season range corresponded with Marsabit evergreen forest, while the wet season range matched with dry deciduous lowland shrubs. The ranging elephants moved at speed of about 0.2‐20 kmh^?1. Bachelor herds moved faster than female family herds. Elephants moved fast during the intermediate and wet seasons than during the dry season. The speed of ranging elephants was over 1 kmh^?1 in the corridor areas and about 0.2 to less than 1 kmh^?1 in the non‐corridor areas. Expansion of settlements towards corridor areas needs to be controlled to avoid future blocking of connectivity between wet and dry season elephant ranges.     

Modelling potential Pleistocene habitat corridors between Afromontane forest regions

The unusually high floral and faunal similarity between the different regions of the Afromontane archipelago has been noted by biogeographers since the late 1800s. A possible explanation for this similarity is the spread of montane habitat into the intervening lowlands during the glacial periods of the Pleistocene, allowing biotic exchange between mountain ranges. In this study, we sought to infer the existence and most likely positions of these potential habitat corridors. We focused on sixteen Afromontane endemic tree, shrub, and bird species in the Cameroon Volcanic Line, East African Rift and Great Escarpment. Species were chosen based on distribution above 1200–1500 m in at least two of the major Afromontane regions. Ecological niche models were developed for each species in the present and projected to the mid-Holocene and the last glacial maximum (LGM). Models were thresholded to create binary maps of presence/absence and then summed across taxa to estimate potential LGM and mid-Holocene distributions. We found widespread climatic suitability for our montane taxa throughout the lowlands of Central Africa during the LGM, connecting all regions of the Afromontane archipelago except the Ethiopian Highlands and the Dahomey Gap. During the mid-Holocene, we noted more limited climatic suitability for fewer species in lowland areas. Although we set out to test predictions derived from alternatively hypothesized corridors, we instead found widespread climatic suitability connecting Afromontane regions across the entire Congo Basin for all species.

     

Evaluating Landscape Options for Corridor Restoration between Giant Panda Reserves

The establishment of corridors can offset the negative effects of habitat fragmentation by connecting isolated habitat patches. However, the practical value of corridor planning is minimal if corridor identification is not based on reliable quantitative information about species-environment relationships. An example of this need for quantitative information is planning for giant panda conservation. Although the species has been the focus of intense conservation efforts for decades, most corridor projects remain hypothetical due to the lack of reliable quantitative researches at an appropriate spatial scale. In this paper, we evaluated a framework for giant panda forest corridor planning. We linked our field survey data with satellite imagery, and conducted species occupancy modelling to examine the habitat use of giant panda within the potential corridor area. We then conducted least-cost and circuit models to identify potential paths of dispersal across the landscape, and compared the predicted cost under current conditions and alternative conservation management options considered during corridor planning. We found that due to giant panda''s association with areas of low elevation and flat terrain, human infrastructures in the same area have resulted in corridor fragmentation. We then identified areas with high potential to function as movement corridors, and our analysis of alternative conservation scenarios showed that both forest/bamboo restoration and automobile tunnel construction would significantly improve the effectiveness of corridor, while residence relocation would not significantly improve corridor effectiveness in comparison with the current condition. The framework has general value in any conservation activities that anticipate improving habitat connectivity in human modified landscapes. Specifically, our study suggested that, in this landscape, automobile tunnels are the best means to remove current barriers to giant panda movements caused by anthropogenic interferences.     

Forecasting the future suitable growth areas and constructing ecological corridors for the vulnerable species Ephedra sinica in China

Ephedra sinica is a rare and vulnerable species in China, and the habitat of Ephedra sinica is seriously threatened (by climate change and human activities). Predicting the suitable growth areas and constructing ecological corridors for Ephedra sinica in China will help to protect it scientifically. Based on 306 valid distribution records and 13 selected environmental factors, the maximum entropy (MaxEnt) model was used to simulate the potential current habitat zones and future (2050 and 2070) habitat zones of Ephedra sinica under four climate change scenarios. The minimum cumulative resistance (MCR) model was applied to extract important ecological corridors of Ephedra sinica. The results indicate that: (1) Under the current environment, the total area of the suitable habitat for Ephedra sinica in China is 42.24 × 10⁵ km², mainly distributed in Northwest China and North China. (2) Suitable area increases as the RCP rises. The center of mass of the habitat zone moved northward from Shaanxi Province to Ordos City in Inner Mongolia Autonomous Region. (3) Of the 13 environmental factors selected, the primary factor was elevation (20.8 %), followed by wettest month precipitation (18.2 %) and temperature seasonality (15.2 %). (4) Built 19 ecological corridors, with a total corridor length of 430.2 km, including seven long-distance passages and 12 short-distance corridors. All corridors are far from the artificial surface, mostly near high-altitude areas. The 19 ecological corridors constructed using the MCR model will also provide considerable importance for the survival of Ephedra sinica on a longer time scale in the future.     

Chorology of New Caledonian palms and possible evidence of Pleistocene rain forest refugia.

The distribution of the 36 palm species of New Caledonia is studied in relation to several parameters: elevation, rainfall, geological substrate, phytogeographical sectors and vegetation types. The climate (thermal gradient and rainfall) appears to be the principal factor influencing the distribution of palms, the substrate effect being subordinate to the climatic pattern. Nearly all palm populations are included within the 1,500 mm isoline. Maximum levels of species richness and endemism are located in four areas receiving more than 3,000 mm of annual rainfall. We interpret these areas as former Pleistocene refugia of lowland rain forest based on three lines of evidence: 1) all locally endemic lowland palm species and genera are restricted to these areas; 2) local endemics occur on east-facing slopes receiving the highest rainfall and most likely to have sustained rain forests during the driest periods; and 3) several pairs of sister species are disjunct between the southeastern and northeastern high rainfall areas.     

A numerical analysis of the mesoscale distribution patterns of vascular plants in the subarctic Kevo Nature Reserve, northern Finland

Different numerical techniques were used to detect and describe the major ecological-biogeographical patterns of vascular plant distributions at the meso-scale level in a subarctic region in Finland. The distribution patterns of 231 native taxa in 362 1 km² grid squares of the Kevo Nature Reserve were analysed by two-way indicator species analysis and detrended correspondence analysis, and were subsequently related to twenty-eight geographical, topographical, geological, and vegetational variables using simple discriminant functions and canonical correspondence analysis with associated Monte Carlo permutation tests.
The floristic variation detected reflects variations in environmental factors operative at the regional and local scales. No major broad-scale coherent geographical patterns were detected; instead, the spatial distribution of the grids with a similar floristic composition shows a scattered distribution. All the numerical techniques reveal a major gradient from alpine areas to lowland sites with rivers and rocky outcrops, and the most important explanatory variables for predicting the main floristic variation are all associated with altitude. The floristic patterns represented by the second ordination gradient mainly correlate with the abundance of mires. Partial ordinations indicate that both the geographical and geological variables explain relatively little of the species distributional patterns.
Although the meso-scale approach reveals much about the plant-environment relationships in the study area, the floristic variation appears to be determined mainly by fine-scale factors. In the most heterogeneous grids, the grid size used fails to detect accurately the ecological patterns of the species present.     

Mountain roads shift native and non‐native plant species' ranges

Roads are known to act as corridors for dispersal of plant species. With their variable microclimate, role as corridors for species movement and reoccurring disturbance events, they show several characteristics that might influence range dynamics of both native and non‐native species. Previous research on plant species ranges in mountains however seldom included the effects of roads. To study how ranges of native and non‐native species differ between roads and adjacent vegetation, we used a global dataset of plant species composition along mountain roads. We compared average elevation and range width of species, and used generalized linear mixed models (GLMMs) to compile their range optimum and amplitude. We then explored differences between roadside and adjacent plots based on a species’ origin (native vs non‐native) and nitrogen and temperature affinity. Most non‐native species had on average higher elevational ranges and broader amplitudes in roadsides. Higher optima for non‐native species were associated with high nitrogen and temperature affinity. While lowland native species showed patterns comparable to those in non‐native species, highland native species had significantly lower elevational ranges in roadsides compared to the adjacent vegetation. We conclude that roadsides indeed change the elevational ranges of a variety of species. These changes are not limited to the expansion of non‐native species along mountain roads, but also include both upward and downward changes in ranges of native species. Roadsides may thus facilitate upward range shifts, for instance related to climate change, and they could serve as corridors to facilitate migration of alpine species between adjacent high‐elevation areas. We recommend including the effects of mountain roads in species distribution models to fine‐tune the predictions of range changes in a warming climate.     

A comparison of distribution patterns in British and Irish mosses and liverworts

Abstract

We classified 747 species of British and Irish mosses into 10 clusters, based on their recorded distribution in 10×10 km grid squares (hectads). We generated the clusters in a two-stage process using the CLUSTASPEC program, the method that we had earlier used for British and Irish liverworts and hornworts. The clusters are named after the species with distributions which are most similar to those of the clusters as a whole. Clusters of widespread species (Bryum capillare), southern, lowland species (Rhynchostegium confertum), widespread calcifuges (Pleurozium schreberi), upland species (Blindia acuta), and montane calcifuges (Kiaeria falcata) closely match clusters recognised in the liverworts. The remaining clusters (Tortella flavovirens, Weissia longifolia, Mnium stellare, Encalypta alpina, Mnium lycopodioides) are less similar. The classification of mosses into 15 and 20 clusters generates additional clusters of hyperoceanic and montane mosses which also resemble liverwort clusters. The influence of calcareous bedrock has a more marked effect in determining moss distributions and, unlike the liverworts, the 10 moss clusters include one which is predominantly coastal. Mosses tend to be a less upland group than liverworts; a smaller proportion of their species have northern and western distributions and the lowland clusters are characterised by more extreme environmental conditions. As with the liverworts, geographically restricted clusters of species with predominantly Mediterranean-Atlantic, Arctic-montane and Boreo-arctic Montane world ranges include marked concentrations of threatened species, and species which are not recorded as fruiting in the British Isles.