Monitoring of pollen rain in moss polsters and mats in view of their suitability as controls for determining the air-borne component of pollen spectra in herbivore faeces

Pollen has been extracted from six bryophyte and two lichen taxa, sampled from three habitats: upland heath, lowland bog and mixed woodland, in both west and east Scotland. The highest number of pollen taxa was recorded in bothSphagnum andThuidium; the lowest numbers of pollen taxa were recorded in two of the mosses and both lichens. For pollen rain control purposes in herbivore faecal studies, woodland species of bryophytes are not suitable due to the sheltering effects from rain provided by the overhead canopy. Open habitats such as moorland heath and bog provide polsters/mats which normally trap pollen rain more effectively for control purposes. The aim of this study was twofold: to observe the effectiveness of selected polsters and lichens as traps for pollen rain and to establish eventually, whether or not such vegetative traps could be used as controls in pollen spectra analyses of herbivore faeces for the determination of nutritional regime and habitat. From a comparison with recent studies in north west Spain of three taxa in common with Scottish samples,Polytrichum demonstrated a consistently higher number of pollen taxa trapped.Thuidium andCladonia all yielded between one and a half and three times the number of pollen taxa as in the same three bryophyte/lichen taxa sampled in Scotland. Investigation relating the pollen rain trapped in polsters to that in faecal material are continuing, and will be reported in due course.This paper is based on one given by E.C. at the 5th International Aerobiology Conference, held at Bangalore, India, August 1994.     

Comparisons of moss polster and pollen trap data: a pilot study

Modern pollen-vegetation relationships have been explored in recent years through analysis of either pollen trap (= absolute results) or moss polster contents (= relative results). In the past few attempts were made to compare data from these two sources. It was the aim of the present study to achieve this comparison through more exact sampling of moss polsters. A pilot study was carried at Morton Lochs, Fife, Scotland where pollen traps have been sampled continuously for ten years. Moss samples were collected from cushions adjacent to two of the pollen traps during May 1985, 1986 and 1987. Moss polsters were sampled usingan equivalent surface cross-section area to that of the traps. In comparing the pollen results from the two sources some similarities can be identified, but in general there are more inconsistencies. These may be traced to difficulties in the precision of moss sampling, variable rates of moss growth and local deposition factors. It is considered that further study, in which these difficulties are minimised, would be justified. Ultimately it is hoped that precise sampling of mosses will produce sufficiently reliable modern pollen rain results to enable pollen-vegetation studies to move away from relative (percentage) values without requiring long-term artificial pollen trapping techniques.     

Evaluation of Swedish woodland key habitats using red-listed bryophytes and lichens

All Swedish forest land is at present being surveyed with the aim of mapping woodland key habitats which have an estimated number of ca. 70 000. An untested requirement of these habitats is that they should contain red-listed species. In order to investigate if they fulfill their aim, an inventory of nationally red-listed bryophytes and lichens was performed in ca. 120 randomly selected woodland key habitats distributed throughout the country. The species were recorded in line transects, covering the whole surface of the habitats. The mean number of red-listed species per habitat was 0.7 for bryophytes and 1.4 for lichens with 11 species at the richest site and 71% of the sites having at least one species. Nemoral forests and coniferous forests were the most species-rich habitat types. The habitats of northern Sweden were more species rich than the southern ones. There were weak correlations between presence of bryophytes and lichens. The study shows that a majority of the woodland key habitats contain red-listed bryophyte and lichen species. Further studies are needed in order to record more organism groups and to compare the woodland key habitats with the surrounding matrix forests.     

Pollen productivity estimates and relevant source area of pollen for selected plant taxa in a pasture woodland landscape of the Jura Mountains (Switzerland)

Relevant source area of pollen (RSAP) and pollen productivity for 11 key taxa characteristic of the pasture woodland landscape of the Jura Mountains, Switzerland, were estimated using pollen assemblages from moss polsters at 20 sites. To obtain robust pollen productivity estimates (PPEs), we used vegetation survey data at a fine spatial-resolution (1 × 1 m²) and randomized locations for sampling sites, techniques rarely used in palynology. Three Extended R value (ERV) submodels and three distance-weighting methods for plant abundance calculation were applied. Different combinations of the submodels and distance-weighting methods provide slightly different estimates of RSAP and PPEs. Although ERV submodel 1 using 1/d (d = distance in meters) best fits the dataset, PPE values for heavy pollen types (e.g. Abies) were sensitive to the method used for distance-weighting. Taxon-specific distance-weighting methods, such as Prentice’s model, emphasize the intertaxonomic differences in pollen dispersal and deposition, and are thus theoretically sound. For the dataset obtained in this project, Prentice’s model was more appropriate than other distance-weighting methods to estimate PPEs. Most of the taxa have PPEs equal to (Fagus, Plantago media and Potentilla-type), or higher (Abies, Picea, Rubiaceae and Trollius europaeus) than Poaceae (PPE = 1). Acer, Cyperaceae, and Plantago montana-type are low pollen producers. This set of PPEs will be useful for reconstructing heterogeneous, mountainous pasture woodland landscapes from fossil pollen records. The RSAP for moss polsters in this semi-open landscape region is ca. 300 m.     

Comparative study of different methods for capturing airborne pollen,and effects of vegetation and meteorological variables

The atmosphere of Mar del Plata city (Argentina) was monitored using three different pollen traps—Burkard, Rotorod, and Tauber—during a flowering season (mid-winter to springtime) in order to compare the diversity and amounts of pollen grains collected in each trap. Sixty three percent of taxa diversity was simultaneously recorded by all three apparatuses. The five most abundant types of pollen in the three traps were Platanus, Ulmus, Quercus, Poaceae, and Fraxinus. The Tauber trap contained the most pollen from taxa with the largest grains, and exclusively trapped ten pollen types with entomophilous characteristics. The amount of pollen in each trap and local/extra-local plant abundance were compared to elucidate the effect of the method used on the pollen–vegetation relationship. The correlation coefficients for total pollen were positive and significant between all pairs of traps. The maximum peak was observed first in the Burkard trap, in agreement with an increase in temperature, but the general pattern in the three traps was similar, with a maximum in early spring. Opposite deviations from the mean value were observed for the Burkard and Tauber data; these deviations were associated negatively with wind speed and relative humidity, respectively. This paper discusses whether in some cases the pollen records could be used in an equivalent manner, although with some limitations.     

Do moss samples,pollen traps and modern lake sediments all collect pollen in the same way? A comparison from the forest limit area of northernmost Europe

Moss polsters, pollen traps and lake surface sediment samples are commonly used as climate calibration data or as modern analogues for reconstructing vegetation from fossil profiles, but the differences in pollen content between these media have received little attention. This study aims to analyse how the three media differ in reflecting individual vegetation types and spatial differences in vegetation. 119 modern samples (64 moss polsters, 37 lake surface sediment samples and 18 pollen traps from which a collection was made annually) were taken from northern Fennoscandia and the Kola Peninsula as a broad transect crossing the northernmost forest limits of Betula pubescens ssp. czerepanovii (mountain birch), Pinus sylvestris (Scots pine) and Picea abies (Norway spruce). The pollen assemblages from these samples were compared with the surrounding vegetation visually and via PCA (principle components analysis) and cluster analysis. Both comparisons allow a correct distinction between pollen assemblages of arctic/alpine heath, mountain birch dominated areas, and boreal coniferous forests. The differences between the vegetation zones are stronger than the differences between the sampling media. Nevertheless, lake sediment samples from the mountain birch woodland zone tend to overestimate pine and underestimate birch. Pollen traps are biased towards lower tree pollen percentages and higher values of shrubs, herbs and Cyperaceae. This bias is especially strong in traps that have missing years in the data. Irrespective of the vegetation zone, pollen traps tend to have lower Pinus pollen percentages than in the adjacent moss polsters.     

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

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

Bryophyte and lichen diversity: A comparative study

Abstract We describe the regional species richness, variation in species richness and species turnover of bryophytes and lichens from 36 sites in lowland forests of southeastern Australia. The analyses subdivided the two major taxa into their constituent sub-groups: mosses, liverworts, and crustose, fruticose and foliose lichens. They also explored correlations between selected environmental variables and patterns of diversity. On a regional scale, there were 77 species of bryophytes and 69 species of lichens, giving a total of approximately one-third of the total number of vascular plant species in the region. Mean species richness was higher for lichens than bryophytes. Also, the two taxa were negatively correlated because lichens favoured dry sites and bryophytes favoured moist ones. Species turnover was greater for bryophytes than lichens, largely due to the distribution of liverwort species. Foliose lichens showed higher levels of turnover than crustose lichens. Multiple regression and canonical correspondence analysis showed that both taxa and all sub-groups responded to the same three variables: vascular plant cover, time since last fire and topographic position. Other variables, including time since logging and intensity of logging, explained little variation in bryophyte or lichen diversity. The data suggest that the strategies for the conservation of bryophyte and lichen biodiversity will be different, to reflect the different patterns of species richness and species turnover.     

Modern pollen–vegetation relationships in subarctic southern Greenland and the interpretation of fossil pollen data from the Norse landnám

Aim The objective of this paper is to explore the relationships that exist between vegetation and modern pollen rain in the open, largely treeless landscape of subarctic Greenland. The implications of these results for the interpretation of fossil pollen assemblages from the time of the Norse landnám are then examined. Location The study area is the sheep farming district of Qassiarsuk in the subarctic, subcontinental vegetational and climatic zone of southern Greenland (61° N, 45° W). Between c.ad 1000–1500 this region was contained within the Norse Eastern Settlement. Methods Detrended Correspondence Analysis (DCA) of harmonized plant–pollen data sets is used to compare plant cover in 64 vegetation quadrats with pollen assemblages obtained from moss polsters at matching locations. Presence/absence data are also used to calculate indices of association, over‐ and under‐representation for pollen types. Results Good correspondence between paired vegetation–pollen samples occurs in many cases, particularly in locations where Salix glauca–Betula glandulosa dwarf shrub heath is dominant, and across herbaceous field boundaries and meadows. Pollen samples are found to be poor at reflecting actual ground cover where ericales or Juniperus communis are the locally dominant shrubs. Dominant or ubiquitous taxa within this landscape (Betula, Salix and Poaceae) are found to be over‐represented in pollen assemblages, as are several of the ‘weeds’ generally accepted as introduced by the Norse settlers. Main conclusions Due to their over‐representation in the pollen rain, many of the Norse apophytes and introductions (e.g. Rumex acetosa and R. acetosella) traditionally used to infer human activity in Greenland should be particularly sensitive indicators for landnám, allowing early detection of Norse activity in fossil assemblages. Pteridophyte spores are found to be disassociated with the ground cover of ferns and clubmosses, but are over‐represented in pollen assemblages, indicating extra‐local or regional sources and long residence times in soil/sediment profiles for these microfossils. A pollen record for Hordeum‐type registered in close proximity to a field containing barley suggests that summer temperatures under the current climatic regime are, at least on occasion, sufficient to allow flowering.     

Changing climate and historic‐woodland structure interact to control species diversity of the ‘Lobarion’ epiphyte community in Scotland

Question: How will changing climate and habitat structure interact to control the species diversity of lichen epiphytes? Location: Scotland. Method: Species richness (=diversity) of the epiphyte lichen community known as Lobarion (named after Lobaria pulmonaria) was quantified for 94 Populus tremula stands across Scotland, and compared in a predictive model to seven climate variables and eight measures of woodland structure. An optimum model was selected and used to project Lobarion diversity over the geographic range of the study area, based on IPCC climate change scenarios and hypothetical shifts in woodland structure. Results: Species diversity of the Lobarion community was best explained by three climate variables: (1) average annual temperature; (2) autumn and winter precipitation; in combination with (3) historic‐woodland extent. Projections indicate a positive effect of predicted climate change on Lobarion diversity, consistent with the physiological traits of cyanobac‐terial lichens comprising the Lobarion. However, the general response to climate is modified significantly by the effect on diversity of historic‐woodland extent. Conclusions: Historic‐woodland extent may exert an important control over local climate, as well as impacting upon the metapopulation dynamics of species in the Lobarion. In particular, a temporal delay in the response of Lobarion species to changed woodland structure is critical to our understanding of future climate change effects. Future Lobarion diversity (e.g. in the 2050s) may depend upon the interaction of contemporary climate (e.g. 2050s climate) and historic habitat structure (e.g. 1950s woodland extent). This is supported by previous observations for an extinction debt amongst lichen epiphytes, but suggests an extension of simple climate‐response models is necessary, before their wider application to lichen epiphyte diversity.     

The natural exclusion of red deer from large boulder grazing refugia and the consequences for saxicolous bryophyte and lichen ecology

Large boulder grazing refugia permitted comparison of saxicolous bryophyte and lichen assemblages with those boulder tops accessible to red deer (Cervus elaphus) on a sporting estate in northwest Scotland. Plant succession was predicted to occur unchecked by grazing on the tops of these large boulders with cascading effects on bryophytes and lichens—assuming boulders had been in place over the same time period. Fifty pairs of boulders (one ≥2 m and the other accessible to red deer) were selected at random from various locations below north-facing crags. Percentage cover of each bryophyte and lichen species was estimated from three randomly placed quadrats on each boulder top. Due consideration was given to the influence of island biogeography theory in subsequent model simplification. Mean shrub cover and height, leaf-litter, bryophyte cover and bryophyte species richness were significantly higher within quadrats on large boulder tops that naturally excluded red deer. Lichen cover and lichen species richness were significantly higher on boulder tops accessible to red deer. Lichen cover was in a significant negative relationship with bryophyte cover, shrub cover and litter cover. Bryophyte cover showed a significant positive relationship with shrub height but there was an optimum shrub cover. Natural exclusion of red deer from the tops of large boulders has facilitated plant succession. The results suggest that grazing arrests the lithosere on boulder tops accessible to red deer at an early plagioclimax favouring saxicolous lichens. The results are relevant to situations where red deer might be excluded from boulder fields that hold lichen assemblages of conservation value.     

Lichens — the biodiversity value of western woodlands

Summary

The Atlantic broad-leaved woodlands of Britain are of international renown for their lichen floras. They are inhabited by 517 lichens, representing 28.3% of the total lichen flora and 73.2% of all British woodland lichens, and they are the main habitat for 165 species. Of these, 31 have a marked southern distribution and do not reach Scotland, whereas 26 species are found in Scotland, but not England or Wales. Their British Red-listed species are outnumbered by the 86 species for which Britain has International Responsibility.

Within the Atlantic broad-leaved woodlands, only 30 lichens show a preponderance for oak. With the exception of some ancient oakwoods in southern England, a high lichen biodiversity is rarely dependent on a dominance of oak in the woodland canopy, more usually it is the result of a long ecological continuity, often a varied tree and shrub composition, a varied canopy density, and good air quality. Consequently, the oak stands within former ‘industrial’ woodlands have a much lower lichen biodiversity compared with woodlands that have a history as ‘pasture woodland’ or, as with some ravine woodlands, have otherwise escaped intensive management.

The life-history of an oak tree is considered in relation to the niches it provides for lichen colonisation with time. Some management scenarios are provided with the enhancement of the lichen interest of former ‘industrial’ oakwoods as an objective.     

Pollen dispersal and representation in a neotropical rain forest

Results of 3 years of pollen trapping on Barro Colorado Island, Panama, are presented. Minimum pollen dispersal distances are estimated for the most abundant pollen taxa. Dispersal distances for some taxa appear to be as low as 5 m, while for other taxa at least 50% of the trapped pollen had travelled more than 40 m. No consistent pattern of spatial variation was reflected in the pollen rain, however, samples close to large canopy gaps had exceptional abundances of 'disturbance' taxa. A preliminary analysis of the representation of canopy components indicates that as much as 19% of pollen caught in the traps was derived from large tree species.     

Acidobacteria in microbial communities of the bog and tundra lichens

Bacterial communities of the lichens from a Sphagnum bog (Karelia) and tundra (Vorkuta oblast) were investigated. Members of the phylum Acidobacteria were numerous in the thallus of living and decaying lichens (3.8 × 10⁸ cells/g), constituting 6 to 32% of the total bacterial number. Pure cultures of acidobacteria were isolated from the samples of living and decaying lichen thallus. Ten of them were identified and classified as members of subgroup 1 of the Acidobacteria. The hydrolytic activity of two strains isolated from the living and decomposing zones of the thallus was investigated. They were capable of growth on xylan, starch, pectin, laminarin, and lichen extract. Acidobacteria were shown to be a stable and numerous component of microbial communities of the bog and tundra lichens.     

Comparing pollen spectra from modified Tauber traps and moss samples: examples from a selection of woodlands across Europe

This paper compares pollen spectra derived from modified Tauber traps and moss samples from a selection of woodland types from Bulgaria, the Czech Republic, Georgia, Greece, Poland, Switzerland and Wales. The study examines the representation of individual taxa in the two sampling media and aims to ascertain the duration of pollen deposition captured by a moss. The latter aim was pursued through the calculation of dissimilarity indexes to assess how many years of pollen deposited in a pollen trap yield percentage values that are most similar to those obtained from the moss. The results are broadly scattered; the majority of moss samples being most similar to several years of pollen deposition in the adjacent trap. For a selection of samples, a comparison of the pollen accumulation rate in pollen traps with the pollen concentration in the moss per unit surface indicates that the entrapment and/or preservation of individual pollen types in the moss differ from that in the pollen trap. A comparison of the proportion of different taxa in the moss with the pollen spectrum of 2 years of pollen deposition in the trap also revealed large differences. There is a tendency for bisaccate grains such as Pinus and Picea to have a higher representation in moss than in traps but there is considerable regional variation. The results indicate that pollen proportions from moss samples often represent the pollen deposition of one area over several years. However, bisaccate pollen grains tend to be over-represented in moss samples compared to both pollen traps and, potentially, lake sediments.     

Tree and stand level variables influencing diversity of lichens on temperate broad-leaved trees in boreo-nemoral floodplain forests

Tree and stand level variables affecting the species richness, cover and composition of epiphytic lichens on temperate broad-leaved trees (Fraxinus excelsior, Quercus robur, Tilia cordata, Ulmus glabra, and U. laevis) were analysed in floodplain forest stands in Estonia. The effect of tree species, substrate characteristics, and stand and regional variables were tested by partial canonical correspondence analysis (pCCA) and by general linear mixed models (GLMM). The most pronounced factors affecting the species richness, cover and composition of epiphytic lichens are acidity of tree bark, bryophyte cover and circumference of tree stems. Stand level characteristics have less effects on the species richness of epiphytic lichens, however, lichen cover and composition was influenced by stand age and light availability. The boreo-nemoral floodplain forests represent valuable habitats for epiphytic lichens. As substrate-related factors influence the species diversity of lichens on temperate broad-leaved trees differently, it is important to consider the effect of each tree species in biodiversity and conservation studies of lichens. Nomenclature Randlane et al. (2007) for lichens; Leht (2007) for vascular plants.     

Herbivory on calcicolous lichens: different food preferences and growth rates in two co-existing land snails

A total of 32 calcicolous lichen species, one alga and one bryophyte were recorded on a limestone wall in the grassland Great Alvar on the Baltic island of Öland, Sweden. Fourteen (41%) of these 34 species and free-living cyanobacteria showed herbivore damage, most probably due to grazing by the land snails Chondrina clienta and Balea perversa which inhabited the limestone wall. Three laboratory experiments were conducted to examine the food preferences of individuals of C. clienta and B. perversa collected at this site and to evaluate any association between their preference and the net food quality of the lichens to the snails. Chondrina clienta and B. perversa exhibited food preferences, which differed significantly between species. Within species, variation in food choice was similar among individuals. This indicates that snail populations may be composed of polyphagous individuals with similar food preferences. Different lichen species were of different net food quality to the snails as indicated by growth rate differences. In both snail species the most preferred lichen species of the choice experiment caused the largest weight increase in juveniles, viz. Caloplaca flavovirescens for C. clienta and Aspicilia calcarea for B. perversa. This suggest that the snail species studied differ in their abilities to deal with secondary compounds and physical characteristics of certain lichens or that they can utilize the energy and nutrients of these lichens to a different extent. It is suggested that differential food preferences might reduce the intensity of interspecific competition for resources (lichens) between C. clienta and B. perversa.     

Growth of reindeer lichens and effects of reindeer grazing on ground cover vegetation in a Scots pine forest and a subarctic heathland in Finnish Lapland

Reindeer lichens are an important component of northern ecosystems. The aim of this study was to measure the growth rate of terricolous lichens as it is a key parameter involved in productivity of these ecosystems and an important part of lichen tolerance to reindeer grazing. Furthermore, the natural succession and the long-term effects of reindeer grazing on lichen community characteristics in two contrasting habitats were investigated as well as the interactions between lichen cover and mosses and vascular plants. Biomass and coverage measurements were conducted in a lichen woodland and in a subarctic heath with grazed and ungrazed areas in northern Finland. Measurements spanning over 13 yr of undisturbed development show that the growth rate of Cladina stellaris can be as high as >0.17 g g⁻¹ produced annually, although in average growth rates were much lower. During the succession of ground vegetation, C. stellaris , C. rangiferina , C. mitis and Cetraria nivalis increased in biomass in fenced areas and were reduced most in biomass by reindeer in unprotected areas. Reindeer grazing and trampling seem to change the vegetation towards a type that is dominated by small dwarf shrubs, bare soil and minute-cup lichens ( Cladonia spp.). Removing the lichen layer by reindeer may reduce natural regeneration of pine trees as implied by increasing numbers of pine seedlings with increasing lichen cover.     

A short pollen diagram from rainforest in highland eastern Victoria

Pollen data from the Rooty Breaks swamp – a Sphagnum bog surrounded by Atherosperma-Elaeocarpus rainforest – in the eastern Victorian highlands appears to reflect changes in the mode of pollen and spore deposition at the study site, from the time sediments began accumulating at about 5500 BP until the present day. In the lower half of the deposit, pollen of rainforest taxa is more common than it is in the top half, while Eucalyptus pollen reflects the reverse situation. Although the pollen changes may reflect climatic change it is suggested that most of the pollen and spores in the lower sediment were stream-borne. Later the stream moved away from the site and the records became more representative of atmospheric pollenrain. Throughout the period covered by the pollen diagram the Rooty Breaks swamp itself was dominated by Leptospermum grandifolium. The pollen record shows that the association of important taxa in rainforests of the highlands of eastern Victoria was established by at least 5500 BP. The presence of sparse old eucalypts throughout the area dominated by rainforest suggests that sclerophyll forest was replaced by rainforest in the recent past. However, this change is not evidenced in the pollen diagram.     

Scots pine growing on forested mires in Abernethy Forest,Strathspey, Scotland

Summary

A substantial proportion of the Abernethy Forest Reserve has Scots pine (Pinus sylvestris) growing on the surfaces of a variety of mires. The hydrology of the mires has been affected by drainage and peat cutting but this area is unusual in having had a long period of protection from grazing by domestic stock. There are three main types of pine populations found on these mires. Woodland bog comprises predominantly bog vegetation with abundant pine seedlings due to the heavy seed rain from the surrounding woodland. Only a few very small trees survive, which are stunted, heavily diseased and have very low seed production. Wooded bog also comprises predominately bog vegetation but there are scattered mature trees of a moderate height with an open canopy. The trees are fertile and can form uneven aged stands with regeneration. Bog woodland is a predominantly woodland vegetation with tall, dense tree cover on deep peat. The trees are well grown with a dense canopy. A few remnants of bog vegetation remain in the ground flora although most have been replaced by woodland bryophytes and shrubs. Each of these three types is described and their development is discussed.