Increased bumblebee abundance along the margins of a mass flowering crop: evidence for pollinator spill‐over

Although the practice of cultivating mass flowering crops (MFCs) is seen as a means of counteracting the widespread decline of insect pollinators, no study to‐date has evaluated whether the increased pollinator abundance associated with MFCs influences pollinator visits to assemblages of adjacent native, non‐crop plant species (pollinator ‘spill‐over’). In this study we quantified bumblebee abundance along hedgerow transects in MFC (field bean) vs non‐MFC (wheat) margins. Surveys were conducted on east and west‐facing margins twice daily (morning and afternoon) for three or four days during the main MFC flowering period (June) over four years involving paired bean and wheat fields in Devon and Cornwall, southwest England. Although no single bumblebee species showed any consistent change in relative frequency, when taken across all years and bumblebee species combined, we observed twice as many bumblebees visiting flowers adjacent to the MFC. However, when we compared bumblebee activity along hedgerow transects two weeks after bean flowering, there was no difference between crops. We thus conclude that although there is evidence for pollinator spill‐over from the bean MFC to nearby semi‐natural habitat, the effect is comparatively short lived and not specific to any single bumblebee species. We also suggest that while pollinator services to native plant species might be briefly enhanced by MFC cultivation, the possible repercussions of bumblebee spill‐over from MFCs to semi‐natural habitats and the other pollinator groups they support should be evaluated before MFC cultivation is considered as a means of conserving declining bumblebee populations.     

Cytophotometric differentiation of high elevation spruces: physiological and ecological implications

Red and black spruce and their hybrids can be determined by morphological indices; however, the criteria are somewhat subjective and increasingly difficult to use at higher elevations. Although the chromosome number is identical (2n = 24), red spruce has twice as much nuclear DNA (48 pg) than black spruce (24 pg) and thus the species and their hybrids can also be separated by cytophotometry. This is relevant to spruce decline studies because black spruce is much more resistant to high elevation environmental stresses, both natural and anthropogenic. It also has implications for the effect of climatic changes on the composition of high elevation spruce-fir forests because red spruce can outcompete black spruce under more mesic conditions. Four elevation transects sampling spruce on the east and west sides of Mount Washington (New Hampshire) and Camels Hump (Vermont) and a single transect on the southwest side of Whiteface Mountain (New York) were made to investigate the degree of hybridization and introgression between these two species. A positive correlation was found between increased elevation and increased black spruce genes on Mount Washington and Camels Hump. Pure black spruce was found on Mount Washington from 1356 m to 1582 m. No pure black or red spruce was found on Camels Hump although the proportion of red spruce alleles was significantly greater on Camels Hump. All trees sampled at all elevations on Whiteface Mountain were pure red spruce. Thus the proportion of black spruce alleles in high elevation spruce populations decreases from east to west. This closely parallels the increase in spruce decline which increases from east to west.     

Cytophotometric Differentiation Of HIGH Elevation Spruces: Physiological and Ecological Implications

Red and black spruce and their hybrids can be determined by morphological indices; however, the criteria are somewhat subjective and increasingly difficult to use at higher elevations. Although the chromosome number is identical (2n = 24), red spruce has twice as much nuclear DNA (48 pg) than black spruce (24 pg) and thus the species and their hybrids can also be separated by cytophotometry. This is relevant to spruce decline studies because black spruce is much more resistant to high elevation environmental stresses, both natural and anthropogenic. It also has implications for the effect of climatic changes on the composition of high elevation spruce-fir forests because red spruce can outcompete black spruce under more mesic conditions. Four elevation transects sampling spruce on the east and west sides of Mount Washington (New Hampshire) and Camels Hump (Vermont) and a single transect on the southwest side of Whiteface Mountain (New York) were made to investigate the degree of hybridization and introgression between these two species. A positive correlation was found between increased elevation and increased black spruce genes on Mount Washington and Camels Hump. Pure black spruce was found on Mount Washington from 1356 m to 1582 m. No pure black or red spruce was found on Camels Hump although the proportion of red spruce alleles was significantly greater on Camels Hump. All trees sampled at all elevations on Whiteface Mountain were pure red spruce. Thus the proportion of black spruce alleles in high elevation spruce populations decreases from east to west. This closely parallels the increase in spruce decline which increases from east to west.     

Primate population dynamics over 32.9 years at Ngogo, Kibale National Park, Uganda

We present census data for eight primate species spanning 32.9 years along the same transect at Ngogo, Kibale National Park, Uganda, demonstrating major changes in the composition of the primate community. Correlated with an estimated decline of ～89% in the red colobus population was an increase in encounter rates with chimpanzee parties. Our data, along with the unusually high rates of predation by chimpanzees on red colobus at Ngogo and the fact that the chimpanzee community at Ngogo is the largest ever recorded, support the conclusion that the red colobus decline was caused primarily by chimpanzee predation. This seems to be the first documented case of predation by one nonhuman primate causing the population decline in another. We evaluated disease and interspecific competition as other possible causes of the red colobus decline, but judged them to be relatively insignificant compared with predation by chimpanzees. Notable changes in encounter rates with other primate species may have resulted from forest expansion. Those for mangabeys, redtails, and black and white colobus increased significantly. Encounter rates increased for l'Hoest's monkeys too, but the increased sightings may have been an artifact of increased habituation. Sightings of blue monkey and baboon groups declined. There was no significant change in encounter rates for all species combined. The Ngogo primate community seemed to be in a nonequilibrium state, changing from one dominated by two species, a folivore (red colobus) and a frugivorous omnivore (redtails), to one dominated by three species of frugivorous omnivores (redtails, mangabeys, and chimpanzees). This study demonstrates the importance of long-term monitoring in understanding population dynamics and the role of intrinsic variables in shaping the species composition of a community.     

Is decline in high altitude eucalypt forests related to rainforest understorey development and altered soil bacteria following the long absence of fire?

Abstract The objective of this study was to identify attributes of the understorey vegetation, soil root biomass, soil chemistry and microbial community that may be associated with tree decline in high altitude eucalypt forests in Tasmania. The sites studied were in healthy eucalypt forest, forest in decline and forest containing dead eucalypts dominated by rainforest, in north‐east (Eucalyptus delegatensis forest) and in north‐west (Eucalyptus coccifera forest) Tasmania. In both regions bare ground, rock and shrubby species were associated with healthy sites whereas decline sites were associated with moss and a tall understorey with a high percentage cover of rainforest species. Healthy sites had low root biomass in the top 10 cm of the soil profile relative to decline and rainforest sites. Seedlings of high altitude species were grown in rainforest soil (0.314% N and 0.060% P) and healthy eucalypt soil (0.253% N and 0.018% P). The four eucalypt species studied had similar root to shoot ratio in the two soils, but the rainforest species, Nothofagus cunninghamii and Leptospermum lanigerum, had higher root to shoot ratio in the healthy eucalypt than in the rainforest soil. We produced three soil filtrates: (i) fungi and bacteria present; (ii) bacteria only present and; and (iii) sterile, from healthy, decline and rainforest sites in north‐east and in north‐west Tasmania and used linseed as a germination bioassay. Filtrates from the north‐east decline and rainforest sites induced a significantly greater dysplastic germination response than healthy sites in (i) and (ii) filtrates, but this was not found in filtrates from sites in the north‐west. We conclude that while the development of a rainforest understorey and elevated soil root biomass in the long absence of fire is generally associated with high altitude eucalypt decline, altered bacterial and/or chemical attributes of soil are not always associated with high altitude eucalypt decline.     

Skuas on the Eurasian tundra; relative occurrence of species, ages and colour phases

The breeding ranges of the three closely related skua species in the genus Stercorarius are highly sympatric on the Arctic tundra. During the Swedish-Russian Tundra Ecology Expedition, 1994, the ages and colour phases of Pomarine Skuas Stercorarius pomarinus , Arctic Skuas Stercorarius parasiticus and Long-tailed Skuas Stercorarius longicaudus were recorded at 15 sites along a transect across 140̀ of longitude from the Kola Peninsula in the west to Wrangel Island in the east. An index of the lemming numbers was also measured. Pomarine Skua comprised 52% of the 1587 skuas seen, with 38% Long-tailed Skua and 10% Arctic Skua. The Arctic Skua occurred at low densities all along the transect. The Pomarine Skua was most common at the northernmost sites in this lemming year. It was absent from all sites with low rodent densities but also from two more southerly sites with high lemming numbers. The Long-tailed Skua showed a distribution sympatric with that of the Pomarine Skua but occurred at lower densities, and it also bred at the more southerly sites. No skuas in second-year plumage were observed. Older subadult skuas were observed in increasing numbers with age. The proportion of subadults was markedly higher in the Pomarine Skua (10%) compared with the two other species. The proportion of dark Pomarine Skuas was about 8% all along the transect. In the Arctic Skua, there was an extremely marked shift from 64% dark birds on the Kola Peninsula to an almost complete dominance by the light phase in the rest of Arctic Russia. I suggest that dark skuas are more efficient kleptoparasites over the sea while the light phase is at an advantage hunting over the tundra.     

A migration-selection model for the morph frequency variation in the peppered moth over England and Wales

A migration-selection model for the spatial and temporal variation of morph frequencies over England and Wales for the peppered moth has been constructed. The morph frequencies have been obtained by computer simulation using the model for various assumptions regarding selection and migration within the boundaries of experimental observation. The selection, in this case, has been assumed to be wholly caused by bird predation and non-visual selection is taken to be same for all phenotypes. The results are compared with available experimental data along three transects. Reasonable agreement with the data is obtained for the Manchester-North Wales transect and the South Wales-London transect. The simulation fails, on the other hand, to reproduce observations along the Central Wales-Birmingham-East Anglia transect. Thus it is concluded that the hypothesis ol balance of selection and migration is untenable as it stands. Either the visual selection pressures are incorrectly or incompletely specified, or else non-visual selection is also operating.     

Studies of the behaviour of Artioposhia triangulata (Platyhelminthes; Tricladida), a predator of earthworms

A significant negative correlation between the numbers of earthworms and the flatworm was demonstrated in fromalin-sampled lawns. Their distributions, recorded by a transect from a shrub border where the flatworm is suspected to have been introduced, indicated that predation may lead to earthworm extinction. Two different vermarium designs demonstrated a gradation of earthworm vulmerability to the flatworm due to depth within the soil and burrow width rather than perference for specific earthworm species. A strong correlation between the level of earth worm predation and flatworm movement implied active hunting. A predation rate was found to be 0.67 of an earthworm per flatworm per week with a metabolic conversion rate of 10%.     

Genetic diversity in butterflies: Interactive effects of habitat fragmentation and climate-driven range expansion

Some species are expanding their ranges polewards during current climate warming. However, anthropogenic fragmentation of suitable habitat is affecting expansion rates and here we investigate interactions between range expansion, habitat fragmentation and genetic diversity. We examined three closely related Satyrinae butterflies, which differ in their habitat associations, from six sites along a transect in England from distribution core to expanding range margin. There was a significant decline in allozyme variation towards an expanding range margin in Pararge aegeria, which has the most restricted habitat availability, but not in Pyronia tithonus whose habitat is more widely available, or in a non-expanding 'control species' (Maniola jurtina). Moreover, data from another transect in Scotland indicated that declines in genetic diversity in P. aegeria were evident only on the transect in England, which had greater habitat fragmentation. Our results indicate that fragmentation of breeding habitats leads to more severe founder events during colonization, resulting in reduced diversity in marginal populations in more specialist species. The continued widespread loss of suitable habitats in the future may increase the likelihood of loss of genetic diversity in expanding species, which may affect whether or not species can adapt to future environmental change.     

The Niobrara River Valley,a postglacial migration corridor and refugium of forest plants and animals in the grasslands of central North America

The Niobrara River Valley of northern Nebraska contains numerous bryophyte, vascular plant, and animal species more typical of forests far to the east, north, and west than to other forests in the grasslands that surround the Valley. Some species are probably relicts of cooler glacial and early postglacial times, when much of what is now grassland was covered by boreal and cool-temperate forests. Others entered the Valley from the east in postglacial times, and some entered from the west as the climate became semi-arid. There is a steep decline in total number of vascular plant species from the mouth of the Missouri River up through the Niobrara Valley, suggesting an environmental gradient and differential migration and extirpation of species at various times since the Pleistocene.     

Tabanidae and other Diptera on Camel's Hump Vermont: Ecological Observations

A canopy trap and aerial nets led to finding 8 species of Tabanidae. There was an abundance of calyptrate muscoid flies. Camel's Hump is in the Green Mountains of western New England, USA. Discovering Diptera on Camel's Hump involved sixteen visits over 40 years. Upwards of 23 other Diptera species are listed. Habitats on the east side and above 762 m (2500 ft) elevation on Camel's Hump differ from the west slope but the boreal forest on both sides is influenced by cloud and fog precipitation on trees. The cliffs just above the 900 m level along the east side are often overlooked, are not seen from the summit and provide access to morning sun for insects. Recent visits explored the role of polarized skylight in relation to the canopy trap, the boreal forest environment and flies found there.     

The decline of Common Kestrels Falco tinnunculus in a forested area of northern England: the role of predation by Northern Goshawks Accipiter gentilis

We have previously documented the decline of the Common Kestrel Falco tinnunculus over a 23‐year period in a large coniferous forest in northern England. Kestrels fed predominantly on Field Voles Microtus agrestis, which were most abundant in young plantations (1–11 years old). Over the 23 years, voles remained abundant in the study area, but their numbers fluctuated cyclically. Here we consider whether the decline of Kestrels was linked to predation by Northern Goshawk Accipiter gentilis. Goshawks first bred in the study area in 1973 and increased until 1989, after which numbers stabilized. We use a number of approaches to explore the role of Goshawk predation, all of which are correlative, but independent. First, there was a significant negative relationship between Kestrel and Goshawk numbers after controlling for a decline in vole habitat. Short‐eared Owls Asio flammeus, which also hunt by day, declined over the same period as Kestrels. Second, numbers of Tawny Owl Strix aluco and Long‐eared Owl Asio otus did not decline as Goshawk numbers increased. These two species are also vole‐dependent, but active by night, and less vulnerable to Goshawk attack. Third, six species of raptor comprised 4.5% of 5445 Goshawk prey items during the breeding season, but more Kestrels were killed than the combined total of all other raptors. Goshawks not only killed many adult Kestrels in early spring, prior to breeding, when it would have most impact on population levels, but there was also a temporal trend for predation on Kestrels to be inversely density‐dependent. Finally, we estimated that Goshawks removed more Kestrels than were recorded each spring in the study area. We interpreted this as indicating that immigrant Kestrels were being removed continually, mostly before they could breed. We conclude that the decline of Kestrels (and possibly Short‐eared Owls) was mainly due to predation by Goshawks. This study provides some of the strongest evidence yet of the role of predation in the hierarchical structuring of raptor communities.     

Changes in traits of shrub canopies across an aridity gradient in northern Patagonia, Argentina

Low and highly variable precipitation pulses exert a strong selective pressure on plant traits and this might provide axes of ecological differentiation among plant species in arid ecosystems. We asked whether aridity contributes to maintain high diversity of species and morphotypes in shrub canopies. We selected eleven study sites evenly distributed across a 400-km transect in northern Patagonia, Argentina. Precipitation is low and highly variable within and between years but almost homogeneous across the transect (125–150 mm). Mean annual temperature varied, however, ranging from 8 °C (west) to 13.5 °C (east) creating a west–east gradient of aridity (aridity index from 3.7 to 7.3, respectively). Sheep grazing commenced in the early 1900s at a similar intensity across the transect. We recorded the richness and cover of shrubs by species and by morphotypes (drought deciduous tall shrubs, evergreen tall shrubs, medium shrubs, and dwarf shrubs), and further calculated the species and morphotype Shannon diversity index at each site. We assessed the presence of spiny leaves, leaf pubescence, thorny stems, and photosynthetic stems in shrub species of all morphotypes and collected green leaves of the dominant shrub species (more than 80% of the total shrub cover) to assess the leaf area, leaf mass per unit area, N-, lignin- and soluble phenolic-concentrations per species at each site. Richness and diversity of shrub species and morphotypes were positively associated with aridity. The richness and diversity of shrub species with pubescent leaves and thorny stems, and nitrogen concentration in green leaves of dominant shrubs increased with increasing aridity. We conclude that our findings on increased diversification in life history traits, species and morhotypes in shrub canopies with increasing aridity support the hypothesis that variability in aridity provides axes of ecological differentiation among shrub species facilitating their coexistence.     

Status and distribution of golden langurs (Trachypithecus geei) in Assam, India

The distribution of golden langurs (Trachypithecus geei) is limited to a small area of western Assam in northeast India and Bhutan between the rivers Manas in the east, Sankosh in the west, and Brahmaputra in the south. It is one of the most seriously endangered primate species of India. A comparative analysis based on satellite images taken in 1988 and 1998 showed a 50% loss of original golden langur habitat. Data on population dynamics collected using line transect and total count methods are presented here. An average group size of 8.2 (range 4.0-22.0) individuals was recorded. A total of 1,064 individuals were counted living in 130 groups. The sex ratio was 1.9-2.5 adult females for each adult male. A low percentage of juveniles and infants suggests that the population is unstable. Most of the groups had only one adult male. Small group sizes, isolated distribution, proportionately few infants and juveniles, and degrading habitat are all causes of concern. Demographic trends indicate a decline in the golden langur population.     

Declining plant species richness in the tussock grasslands of Canterbury and Otago, South Island, New Zealand

We studied vegetation change on 142 permanently marked transects spread throughout tussock grasslands of Otago and Canterbury, in areas subject to both pastoral and conservation management. The transects were established between 1982 and 1986 and remeasured between 1993 and 1999, providing a record of vegetation change at each site over an interval varying from 10 to 15 years. Each transect consisted of 50 quadrats, each 0.25m(2), in which the presence of all vascular plant species had been recorded. For each transect, we calculated the change between measurements in the mean number of species recorded per quadrat, and the change in the total number of species recorded per transect. Averaged across all transects, there was a significant decline in species richness between measurements at both the quadrat and transect scales. Small herbs (those less than or equal to2 cm tall, excluding Hieracium species) showed the greatest decline. On average, more than one quarter of the small herb species present in a quadrat at the first measurement had disappeared within 10 years. Larger herbs, ferns, rushes, sedges and grasses (excluding Chionochloa species) also declined significantly in species richness, reflecting declines in the abundance of species in these groups. Woody species richness remained constant, while species in the genera Chionochloa and Hieracium increased significantly in mean quadrat species richness, reflecting increases in the abundance of these species along transects. The rate of decline in mean quadrat species richness was unrelated to changes in the abundance of either Chionochloa or Hieracium species, or to an overall increase in total vegetation cover on transects. The rate of decline in species richness was also unrelated to the level of grazing or burning between measurements. However, the rate of decline in species richness was greater at lower elevation, on schist rock and on yellow-brown and yellow-grey soils. Our results suggest that a major compositional change is occurring in these grasslands at a rate that is independent of local variation in management and independent of the widespread invasion of these grasslands by Hieracium species.     

Copepod and barnacle nauplius distributions in the Irish Sea: relation to springtime hydrographic variability

The distributions and abundances of copepod and barnacle naupliiare described along a five-station transect in the Irish Seafrom Liverpool Bay, England, to Dundalk Bay, Ireland, in May1989. Correspondence analysis revealed that various hydrographicregions in the Irish Sea are characterized by distinct naupliusassemblages. Acartia, Centropages and Temora nauplii were associatedwith the Welsh coastal water east of the Liverpool Bay front.The western stratified water to the west of the seasonal, westernIrish Sea front was distinguished from the mixed water to theeast by abundances of Calanus, Pseudocalanus, Oithona, Microcalanusand harpacticoid nauplii. The Liverpool Bay and western IrishSea fronts are, apparently, boundaries between coastal and offshoreassemblages of crustacean nauplii.     

Studies of colour polymorphism in some marginal populations of the aposematic jersey tiger moth Callimorpha quadripunctaria

Callimorpha quadripunctaria (Lepidoptera, Arctiidae) is both genetically polymorphic for hindwing colour and well protected from predators. Polymorphic populations are, however, only found in the extreme northwest of its extensive distribution in Europe and Asia Minor. Use of a colour analyser confirmed the existence of three reasonably discrete colour classes: bright red, orange and yellow. These could each be subdivided into two further groupings of phenotypes, one of moths of pure colour and the other with a secondary colour. The species is polymorphic in SW England, where it became well established in villages and towns around the Exe estuary in the late 19th century. It now extends about 30 km east and west along the south coast of Devon and up to a similar distance inland along the river valleys. The rate of spread is consistent with the substantial dispersal of up to 500 m detected in a capture-recapture experiment performed at Exeter over the whole flight period in August 1984. This experiment also found no differences in timing of emergence, survivorship or within-habitat movement between the sexes or the colour classes. Males may show a higher rate of emigration than females. The species is polymorphic throughout S Devon. Its stronghold to the west of the Exe basin is characterized by a comparatively uniform morph frequency with 5045% red, 20–40% orange and 10–20% yellow. Yellow moths have their highest frequency in this area, red moths in the east and west and orange to the north. Adult activity is concentrated in the 2–3 h after dusk. Air temperatures of higher than 15°C are probably critical to pairing which involves sex pheromones released by females. This may be an important factor limiting the species' range. Our results suggest that climatic selection pressures involving non-visual differences in fitness account for the geographical variation. The lack of daytime activity of moths in comparison to that found by other workers on Rhodes and which is likely in other warmer climates suggests the hypothesis that expression of the polymorphism is favoured in the marginal populations of NW Europe by a relaxation of the stabilizing influence on the warning coloration exerted by visual predators.     

Prevalence of nest predators in a sub-Arctic ecosystem

High nest loss is an important driver of gallinaceous bird population dynamics. Identifying factors determining the spatial distribution of potential nest predators and thereby indirectly risk of nest losses is therefore essential. The aim of this 1-year study was to estimate relative predation rates on artificial ground nests in willow ptarmigan (Lagopus lagopus) habitats, along replicate altitudinal gradients (transects, n?=?60) spanning from sub-Arctic birch forest to the low-alpine tundra in three locations in northern Norway. In each transect, one artificial nest (track board) was placed in three different habitats: (1) birch forest, (2) edge between birch forest and low-alpine tundra and (3) low-alpine tundra. Total predation rates over all habitats within locations ranged from 47.4% to 77.5% and did not vary systematically in space and time. The average predation rate by avian predators was consistently high (58%), and mammalian predation rate was consistently low (5.6%). The consistently high level of predation inflicted by birds was mainly due to omnipresent corvids, especially the hooded crow (Corvus cornix). Analysis of species-specific predation rates showed that habitat and location effects were insignificant for all species, except for raven (Corvus corax) that showed clearly higher predation in one of the locations. The results indicate that from the perspective of the spatial distribution of potential nest predators in sub-Arctic birch forest, ground nesting birds like willow ptarmigan should not be expected to be selective with respect to nesting habitat in the ecotone between birch forest and the low-alpine tundra.     

贺兰山岩羊的数量与分布

2003年11～12月,采用样线调查法对贺兰山东坡(宁夏贺兰山国家级自然保护区)和西坡(内蒙古贺兰山国家级自然保护区)境内岩羊(Pseudois nayaur)种群的数量与分布进行了调查。共记录岩羊实体141群746只。在宁夏贺兰山国家级自然保护区和内蒙古贺兰山国家级自然保护区范围内均有岩羊分布,但东坡岩羊的平均群大小为6·03只,西坡为3·24只;东坡的遇见率为0·32只/km,西坡的遇见率为0·19只/km。卡方检验表明,贺兰山东、西坡的平均群大小和遇见率均存在极显著差异,东坡是岩羊的主要分布区。利用DISTANCE5·0软件估计贺兰山东坡岩羊的密度为5·144只/km2,种群数量为10611只;西坡的密度为2·532只/km2,种群数量为1714只;整个贺兰山地区岩羊的估计密度是4·445只/km2,种群数量为12178只。     

Occurrence of epiphytic bryophytes in a ‘tetrad’ transect across southern Britain 1. Geographical trends in abundance and evidence of recent change

Abstract

A survey was undertaken by members of the British Bryological Society of all epiphytic bryophytes encountered in 107 tetrads (2 × 2 km land units) located within a systematic selection (1-in-9) of 10 × 10 km grid squares in a belt transect aligned west-east across southem Britain. Special field cards enabled data to be collected on frequency (number of trees occupied), presence of sporophytes, types of tree colonized and position on the host trees. Totals of 112 species and 4 varieties of mosses and 23 species of liverworts were recorded on 52 species or species aggregates of woody plants. Fraxinus excelsior and Sambucus nigra were the most frequently recorded phorophytes and responsible for 34% of all epiphyte records. Only one (Orthotrichum affine) out of the eleven epiphyte taxa which were frequent throughout the transect could be regarded as an obligate epiphyte, the remainder occurring commonly in other habitats. These common epiphytes are all characterized by abundant spore production. Many other epiphytes showed restriction to a portion of the transect. About ten species that were limited to the western part of the transect probably require more frequent rainfall and are possibly excluded by higher atmospheric pollution in the east. Among several facultative epiphytes occurring mainly in the east were Ceratodon purpureus, Grimmia pulvinata and Tortula muralis which are not so restricted in other habitats. The latter two, normally epilithic, may be benefiting from newly available habitats with low competition as a result of the recent decline in atmospheric SO₂ levels and slow recolonization by many epiphytic species. Increased N deposition may also be involved. In comparison with their distributions mapped over the preceding 30 years, about 13 obligate epiphytes now appear to have increased markedly, probably in response to declining SO₂ Several other obligate epiphytes characteristic of the circumneutral bark of mature, well-lit trees, e.g. Leucodon sciuroides, have not increased or have declined sharply. Bark acidification by atmospheric pollutants and loss of mature Ulmus spp. may be important causes. Lejeunea ulicina, whose western distribution pattern appears to be static, may be extremely poor at colonizing formerly polluted territory owing to the lack of both spores and gemmae.