Biodiversity and vegetation dynamics in the coastal heathlands of western Norway

The heaths of Lygra, western Norway, were investigated, and 19 communities at the association level were distinguished. Distinct ecotypes of Ranunculus acris and Poa humilis and microspecies of Euphrasia were restricted to the grass heaths. No distinct ecotypes were found in the Calluna -dominated vegetation. It is suggested that the regular burning, which has been part of the management of the area, has activated the seed bank of the ericaceous heath, and thus maintained genetic diversity and slowed down ecotype differentiation. The grass heaths, with no combustible material, have been unaffected in this way. Permanent plots in the two dominant heath types, dry heath (Vaccinio-Callunetum) and wet heath (Ericetum tetralicis) were followed over six years and the data analysed using a modification of Jaccard's index, allowing for species abundances.     

The effect of heather beetle Lochmaea suturalis on vegetation in a wet heath in NE Scotland

An outbreak of heather beetle Lochmaea suturalis on a wet heath in north-east Scotland was monitored. In many respects, the outbreak was similar to those described in lowland heaths; it started at a particular locus and when the Calluna vulgaris was depleted, spread throughout the heath. Population size then decreased annually due to some undetermined exogenous factor. A contributory factor may be the high incidence (35%) of parasitoid infestation which is density independent and, thus, not controlling.
There is no evidence that the hairiness of the var. hirsuta of Calluna has an effect on herbivory by Lochmaea. The distribution of larvae through the canopy is neither influenced by the morphology of Calluna , nor by the associated species, and appears to be at random. This probably represents the distribution of eggs since little lateral migration of the larvae takes place.
The response of the vegetation to herbivory is not simple. Associated with decreased cover of Calluna is an increase in Sphagnum plumulosum and Hypnum jutlandicum and reduction in S. compactum and Pleurozium schreberi. Changes seem to depend on initial vegetation and other parameters not measured (e.g. proximity of propagules, microclimate). Regeneration of Calluna does occur, largely by layering; although seedling occurrence increases in some areas, especially in Sphagnum , it seems unlikely that this will make a major contribution to the recovery of Calluna.     

Early stages of Calluna vulgaris regeneration after burning of coastal heath in central Norway

Questions: How does Calluna vulgaris regenerate after burning wet and dry heaths of different age? Location: Central Norway. Methods: Patches of Calluna dominated old and young wet heath and old dry heath were burned. The mean temperatures reached in the categories of heath were measured. Cover, frequency, height and number of seedlings of Calluna were studied in plots. Results: The mean burning temperatures varied from 680 to 740°C, and were highest in wet heath that had not been burned for at least 50 years. Regeneration of Calluna was evident in the year of burning, but only from seeds. Even heath with a short burning interval lacked vegetative regeneration. Calluna cover increased yearly and exceeded 50 % three years after burning. Regeneration was evident in the year of burning, and was highest in heath with a short burning interval. The height increased regularly at all sites, but was most rapid in dry heath, probably because of better microclimatic conditions. There were more seedlings in the heath with a short burning interval. Conclusions: Calluna only regenerates from seeds afterburning in central Norway. The density of Calluna increased rapidly, which is positive for the conservation and management of coastal heaths in central Norway.     

Geographical separation of two Ulex species at three spatial scales: does competition limit species' ranges?

It is a common assumption that species' ranges are limited by their physiological tolerances to climatic factors, Biotic factors, such as competition, are rarely considered. We investigated the distributions of Ulex minor and U. gallii at three spatial scales from geographic ranges to individual heaths - to examine whether the species are negatively associated, as predicted by the hypothesis that the ranges of the species are limited by competition with each other. Distribution maps for the British Isles and France (100 400 km² survey units) show the two species have largely separated, but slightly overlapping ranges. A region of range overlap on the heaths of Dorset, southern England was mapped using 4 ha survey squares. There was strong negative association between the species, and the heaths could be divided into zones where one species was dominant. There was some indication of edaphic differences between the U. minor -dominated zones and the U. gallii zones. The few heaths where the species co-occurred were surveyed using 4 m² quadrats placed along transects. Usually one species was widespread over the heath, while the other occurred in patches. The species were strongly negatively associated in all transects. Therefore, the two species showed strong negative associations at three mapping scales. Apparent co-occurrences detected at one spatial scale largely disappeared when species were mapped at finer scales, emphasising the fractal nature of distributions. This provides evidence that the distributions of the two species are not independent and that they cannot coexist, and therefore that their ranges are limited by competition. Over their ranges, competitive superiority is probably determined by the climate. At the range boundaries in the region of overlap, climate is not important, but other physical factors such as edaphic conditions may determine the outcome of competition.     

Genetic and morphological divergence among Gravel Bank Grasshoppers, <Emphasis Type="BoldItalic">Chorthippus pullus</Emphasis> (Acrididae), from contrasting environments

Gravel Bank Grasshopper (Chorthippus pullus) populations inhabit two contrasting environments, pebbly gravel banks with scarce vegetation cover in mountainous areas along the Alps and lowland grasslands dominated by Common Heather (Calluna vulgaris). Heath populations of C. pullus have been rediscovered only recently, and show a distribution scattered across Central Europe. The wings are reduced in this species; thus, it has low potential for long-distance dispersal. We used sequence data on a newly developed non-coding nuclear marker from three gravel-bank and four heath populations to test whether grasshoppers from the two environments represent distinct lineages. Gravel-bank populations were studied in southern Germany (Bavaria), heath populations in eastern Germany (Brandenburg and Saxony) and Ukraine. We compared those genetic data with an analysis of variation in a suite of morphometric traits. Finally, we combined genetic and morphometric data to reconstruct a plausible scenario for the ecological shift observed in C. pullus. Our newly developed marker did not sort populations from contrasting environments in two monophyletic lineages. Nevertheless, we found a general lack of gene flow between the gravel-bank and heath populations. There was pronounced variation among populations in morphometric traits. That variation was partially partitioned by habitat type, and populations from the same habitat tended to be more similar than those from different habitats. Our data suggest that heath populations originated through northward expansion from multiple southern European refugia, and that the gravel-bank populations represent one of these sources. Patterns of genetic and morphometric divergence suggest that gravel-bank and heath populations may be in the process of incipient speciation.     

The Lepidoptera fauna associated with Calluna vulgaris: effects of plant architecture on abundance and diversity

Abstract .1. The effects of plant architecture on the Lepidoptera assemblage associated with heather Calluna vulgaris were studied at four locations in northern England and southern Scotland in 1992 and 1993. The study areas were Calluna vulgaris – Vaccinium myrtillus heaths, where management by rotational burning had created a mosaic of stands of different-aged Calluna . Lepidoptera were sampled in the larval stage.
2. Larval abundance was considered in relation to Calluna age, height, cover, green shoot density, and flower density. For most study areas and species, multiple regression selected Calluna height as the variable that explained most variation in larval density, although the explanatory variables were intercorrelated.
3. After compensating for the effects of different study locations, the slope of the regression between the logarithm of larval density and the logarithm of Calluna height was common to the macrolepidoptera, microlepidoptera, geometrid, and noctuid larval groups.
4. There was a significant progressive increase in larval diversity with increase in Calluna height, due to the presence of uncommon moth species in the samples from taller Calluna, and a change in the contribution of common species to the community in different height zones.
5. The significance of these results is discussed in relation to other heathland invertebrate studies and management practices.     

Plant coexistence in coastal heaths: Between- and within-habitat effects of competition,disturbance and predation in the post-fire environment

Abstract Previous experiments on post-fire establishment of two obligate seeders and two resprouting species suggested that initial establishment processes differ between wet- and dry-heath habitats. Disturbance, interspecific competition between seedlings and between seedlings and adults, and access for potential predators were manipulated in the field after a fire to identify mechanisms to explain between- and witm'n-habitat species coexistence. When soil surfaces are disturbed or seeds are buried, dry-heath species can establish in the wet heath. Under natural conditions, however, wet heath is rarely disturbed by animal foragers and lack of safe sites may preclude establishment of dry-heath species. In contrast, dry heath is often disturbed by ground foragers such as bandicoots and safe sites are plentiful for establishment of all heath species. Nevertheless, while wet-heath species can establish in dry heath, their seedlings are apparently unable to survive there due to the drier conditions. No evidence for competition among species was detected in either type of habitat during the first 3 years of the experiment, even at elevated seedling densities. Lack of early competition and unsaturated seed-banks may promote coexistence of species. Seedling predation from vertebrates was low and did not appear to influence patterns of coexistence within or between habitats. In terms of community theory we suggest that several interacting mechanisms are responsible for community structure in heaths after a fire. Chance and microsite disturbance are important during dispersal, while resources are critical during establishment and early survival of seedling populations. In these heath systems where fires burn extensive areas, seed-banks may be more important in temporal storage of species than patch or spatial storage as a mechanism for maintaining coexistence and plant diversity.     

The phytogeography of European and Mediterranean heath species (Ericoideae, Ericaceae): a quantitative analysis

The geographic ranges of heath species in Europe and the Mediterranean and their relationships with environmental (climatic and ecogeographic) variables and biological features are analysed by means of multivariate methods. In particular, twinspan classifications into floristic elements and floristic regions, DCA floristic ordinations, CCA environment-constrained ordinations and CCA biology-constrained ordinations are carried out. Results of the analyses show a correspondence with conventional regionalization analyses based on broader criteria, and less correspondence with numerical analyses of other taxonomic groups at a similar scale. This lack of fit depends on the particular history and ecology of the taxonomic groups under study. A number of climatic (temperature and water stress) and geographic (coast length) variables are associated with different types of heaths according to their geographical ranges (continental, Mediterranean, Atlantic). Biological features of heaths account for a small part of the variation in range, but the association of temperate heaths with a preference for acid soils, of Mediterranean heaths with pubescence, and of Atlantic heaths with plant height is of interest. Heath species richness throughout Europe and the Mediterranean is analysed by multiple regression analyses and, apart from a strong influence of area size, a significant effect of water conditions, temperature and proximity to sea is detected. The area with the highest heath species richness is Western Mediterranean. Additional classification, ordination and multiple regression analyses of heaths in the Iberian Peninsula (Spain and Portugal, the areas with the highest heath diversity) revealed similar patterns to those found in Europe and the Mediterranean. The effect of heterogeneity of the studied units at this latter scale is removed in the Iberian analysis because of the relative homogeneity of the units considered at this scale.     

Chloroplast DNA diversity in Calluna vulgaris (heather) populations in Europe

Variation in the chloroplast genome of Calluna vulgaris (heather), the dominant species of northwest European heath communities, was analysed using polymerase chain reaction-restriction fragment length polymorphisms (PCR-RFLPs) and microsatellites. No length polymorphisms were detected in the 100-200 base pair (bp) fragments amplified by the conserved microsatellite primers, and sequencing revealed that the repeat regions were interrupted relative to the corresponding sequence in Nicotiana tabacum. In contrast, PCR-RFLP analysis revealed high levels of haplotype diversity within populations (hS = 0.443, hT = 0.842), as well as substantial differentiation between populations (GST = 0.473). Diversity and differentiation were higher in southern Europe than in northern Europe. Interpreted in the light of data from allozyme studies and pollen core records, the results suggest that the main glacial refugia for C. vulgaris were located in southwest Europe, including northern Spain, the Pyrenees and the Massif Central region of France. There is also evidence for diffuse survival of the species at more northerly latitudes.     

Plant coexistence in coastal heaths: Habitat segregation in the post-fire environment

Abstract Previous studies of heaths on Pleistocene coastal sands showed consistent variation in patterns of floristic composition between areas on ridges and slopes but not within them. A large wild-fire that swept this system provided an opportunity to observe temporal processes in habitat segregation of species. Ridges and slopes were found to differ in species richness of seedlings in 0.0625 m2 quadrats and there was no evidence that the species richness in the two habitats was converging over 3 years. This suggests that initially these processes differ in space between the two habitats with more species in the wet heath being packed into a smaller area than in the dry heath. Seed-banks of species did not saturate available space for recruitment in either habitat, but seedling densities differed asymmetrically between habitats across two pairs of species studied. Experimental manipulation of seeds among habitats also showed distinct differences in establishment and survival among representative species from different habitats. The presence of seed-banks of wet-heath species in dry heath suggested that recruitment of their seedlings may occur there, but experiments showed that seedlings that arise from them do not survive there. Conversely, although no seed-bank of either dry-heath species studied was found in wet heath, our manipulations showed that their seedlings could grow there. Thus, superficially, physiological tolerance appeared to limit regeneration of wet-heath species in dry heath. However, there appeared to be no physiological limit for dry-heath plants to survive in wet heath. This paper shows that segregation of species may operate at either of two stages of the life cycle: at dispersal when safe sites for establishment are required, and at recruitment after seedlings have established. These findings highlight the importance of the regeneration niche in structuring community composition in coastal heaths, and contrast with traditional explanations of species segregation invoking physiological tolerance and competition at later life-history stages.     

Regeneration capacity of oceanic-montane liverworts: implications for community distribution and conservation

Abstract

Scotland’s mountains are home to a rare and unique liverwort community, ‘the oceanic-montane liverwort-rich heath’, but its component species are absent from regions where they could potentially thrive. Many biological characteristics of these species are unknown, making it impossible to explain the reasons for their rarity; however, they have not been observed to produce sporophytes within Britain. We use ex situ cultivation of whole liverworts and fragments, and in situ cultivation of fragments, to assess the growth rate and the potential for vegetative reproduction of several species. Most of the species grew from both fragments and as whole plants, indicating that the rarity of the liverwort heath is not due to poor powers of regeneration. We propose that growth rate and the potential to regenerate from fragments are important factors structuring the liverwort heath community, at least locally. Furthermore, this study demonstrates that there is potential for ex situ conservation of rare liverwort species, in situ enhancement of existing populations, and creation of new ones.     

Does recent habitat fragmentation affect the population genetics of a heathland specialist, <Emphasis Type="Italic">Andrena fuscipes</Emphasis> (Hymenoptera: Andrenidae)?

Habitat fragmentation is believed to be a key threat to biodiversity as it decreases the probability of survival of populations, reduces gene flow among populations and increases the possibility of inbreeding and loss of genetic diversity within populations. Heathlands represent excellent systems to study fragmentation effects as the spatial and temporal course of fragmentation is well documented for these habitats. At the beginning of the nineteenth century, heathlands were widespread in northern Germany, but they became increasingly fragmented at the end of the nineteenth century until only few fragments had been left. As many insect species are strongly specialized on heathland habitats, they represent ideal study systems to test the genetic effects of such recent fragmentation processes. The solitary bee Andrena fuscipes is strongly specialized on heather (Calluna vulgaris) and, therefore, occurs exclusively in heathland habitats. The species is red-listed in Germany and other parts of Europe. Here, we present an analysis of the genetic structure of 12 populations of A. fuscipes using eight microsatellite loci. The populations showed little geographical structure and the degree of genetic differentiation was low. Compared to related bee species, inbreeding coefficients were relatively low and seem to be mainly affected by the bees’ solitary nesting behaviour.     

The reproductive potential and importance of key management aspects for successful Calluna vulgaris rejuvenation on abandoned Continental heaths

The abandonment of traditional pastoralism as well as the use of heath areas for military purposes has had a major impact on dry heaths in the Continental biogeographical region of Europe, causing severe degradation of its key species Calluna vulgaris (L.) HULL. The reproductive potential of this species in a Continental climate is assumed to be low, although there is yet no observational or experimental evidence for this. More knowledge is also needed about cost‐effective and sustainable measures to restore abandoned dry heaths in this biogeographical region, because traditional management options are often too expensive (e.g., sod‐cutting) or restricted due to environmental laws and the danger of unexploded ammunition (e.g., burning). Using as an example an 800 ha Continental heathland in Germany that has been abandoned for about two decades, we studied the reproductive potential (seed production, soil seed bank, and germination ability) of degenerate C. vulgaris stands. In addition, we conducted a comprehensive field experiment to test the effects of low‐intensity, year‐round grazing by Heck cattle and Konik horses as well as one‐time mowing and patchy exposure of bare soil on the generative rejuvenation (i.e., recruitment and survival) of degenerate C. vulgaris stands over 3 years. We used generalized linear mixed models for statistical analyses. Seed production of degenerate C. vulgaris stands was high as well as the germination ability of their seeds, being similar to Atlantic heathlands. However, soil seed‐bank densities were lower than those found in managed or abandoned Atlantic heaths. Overall seedling recruitment in the field was considerably lower in comparison with Atlantic heaths. Low‐intensity grazing or one‐time mowing did not induce a substantial increase in C. vulgaris recruitment, whereas an additional one‐time creation of bare soil patches or the one‐time creation of bare soil without subsequent management significantly facilitated seedling recruitment and survival in the first year. However, from the second year on, the positive effect of the creation of bare soil without subsequent management was no longer present. In the third year, survival of juveniles was significantly supported by low‐intensity grazing in combination with shallow soil disturbances as well as in combination with one‐time mowing and shallow soil disturbances, whereas mowing alone resulted in marginally significant lower survival. The extremely low seedling recruitment requires a careful choice of suitable management measures to promote the survival of sufficient numbers of Calluna individuals. Therefore, we recommend low‐intensity grazing with free‐ranging robust breeds and the combination of this with one‐time mowing as an effective means of supporting generative rejuvenation of C. vulgaris in degraded heaths. However, at the beginning of the restoration process, the creation of bare soil patches for seedling recruitment is crucial. For implementation into practice, we present different strategies to enhance the proportion of bare soil after long‐term abandonment of heaths when traditional management options are no longer feasible.     

The age of Calluna stands moderates post‐fire regeneration rate and trends in northern Calluna heathlands

Questions: Does stand age influence the direction and rate of post‐fire successional dynamics in coastal Calluna heaths and can old degraded heath vegetation be restored through reintroduction of fire? Location: Coastal heaths in the Tarva archipelago, central Norway. Methods: We investigated revegetation dynamics after experimental fires set in young (8 years since last fire) and old (>50 years since last fire) grazed heath stands. A repeated measures design was used, with floristic data recorded in permanent plots in the post‐fire successions (n=12) over a 7‐year period. The data were analysed using multivariate ordination techniques (PCA, RDA and PRC) and mixed effects models. Results: The age of Calluna stands strongly influenced post‐fire succession, different trends due to age explained 10.4% of variation in floristic data. Young heath showed faster succession towards pre‐fire community composition than old heath, and this could partially be explained by succession‐related factors: young heath had lower cover of mosses and lichens in the pre‐burned vegetation, and lower cover of litter early in succession. Young heath had a less pronounced overall community response to fire than old heath. Vegetative regeneration of C. vulgaris was absent in both old and young heath, but Calluna still re‐established as the dominant species within 5–7 years in both young and old stands. Regeneration dynamics were also affected by habitat conditions, different trends due to habitat explained 6% of variation. Conclusions: Our study demonstrates that old stands do develop characteristic heathland vegetation and structure after fire, and while potential invasives into the system such as trees and rhizomatous species are present, they do not impair Calluna regeneration or vegetation development towards the target heathland community composition and structure. Further, as our young stands are only in their second fire rotation after restoration, we suggest that characteristic dynamics of managed heathlands can re‐establish relatively rapidly, even in severely degenerated sites (>50 years since last fire). Site‐specific factors also need to be considered. We conclude that there is restoration potential in old heaths, despite slow dynamics in the first rotation.     

Is Calluna vulgaris a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems?

In the face of ongoing atmospheric nutrient loads the employment of management measures to remove nutrients from heathland ecosystems has increased in importance. The present study is the first to analyse whether Calluna vulgaris is a suitable bio-monitor of management-mediated nutrient pools in heathland ecosystems. If Calluna vulgaris proves to be an appropriate indicator, its bio-indicative usage may prove to be a helpful tool for an assessment of management success in heathland ecosystems. In the Lüneburger Heide nature reserve (NW Germany) we analysed the impacts of grazing, mowing, prescribed burning, choppering and sod-cutting on the nutritional status of Calluna vulgaris by measuring nutrient contents (N, P, Ca, Mg, K) of current year's shoots 1 and 5 years after application of management measures. Results were related to management-induced nutrient flows and nutrient pools at the focal heath sites. Our results indicate that the less the physical environment of a heath site was affected by management measures the better the nutrient contents of current year's shoots of Calluna vulgaris mirrored changes in nutrient pools. For low-intensity measures (i.e. grazing, mowing, prescribed burning), shoot nutrient contents were a suitable indicator for changes in nutrient pools, particularly for nutrients with conservative cycles such as P. At grazed and mown sites high output rates of P caused by these measures were well reflected by decreased shoot P content. At burned sites, Calluna vulgaris proved to be a good indicator of changes in nutrient pools of the organic layer, mainly attributable to the deposition of nutrients with ash. In contrast, at sites subjected to high-intensity measures, shoot nutrient contents did not reflect management-mediated shifts in nutrient pools, despite the high nutrient losses caused by choppering and sod-cutting. At these sites, shoot nutrient contents mirrored only the effects of altered mineralisation rates attributable to changes in the physical environment following high-intensity measures. As plant growth and competition in heathlands is considered to be controlled by N or P, shoot N:P ratios are recommended as a tool to indicate whether plant growth tends to be limited by N, by P or by N and P. This, in turn, allows for an assessment of long-term effects of both atmospheric nutrient loads and management-mediated shifts in N and P pools at a focal heath site.     

Ecological distribution of four co-occurring Mediterranean heath species

Erica australis, E. scoparia, E. arborea and Calluna vulgaris are the most abundant heath species on acid, sandstone-derived soils of the Strait of Gibraltar region (southern Spain and northern Morocco). Despite their apparently similar ecological requirements, these four species are somewhat ecologically segregated. Erica australis is abundant only on poor, shallow soils, with a high content in soluble aluminium, generally on mountain ridges and summits. Erica scoparia becomes dominant on deeper sandstone soils with lower aluminium. Calluna vulgaris coexists with these Erica species in communities under low or no tree cover. In the Spanish side of the Strait (Algeciras), Erica arborea tends to be relegated to communities under moderate to dense tree cover, whereas this species is more abundant and widespread in the Moroccan side (Tangier). Tolerance to extreme physical conditions - high aluminium and dense tree cover - and interspecific competition seem to explain the ecological distribution of these four heath species in the Strait of Gibraltar region. The more fragmented pattern of sandstone patches and higher disturbance levels in Tangier might account for the differences in the patterns of ecological distribution of these four heath species between both sides of the Strait of Gibraltar.     

Genetic differentiation among populations of Sesleria albicans Kit. ex Schultes (Poaceae) from ecologically different habitats in central Europe

As observed for many other plant species, the populations of Sesleria albicans in Central Europe are located in habitats, which differ to a high degree from each other with regard to ecological factors such as nutrients, light and water as well as in type of land use. The species colonizes limestone cliffs, pavements, screes, grazed and mown grasslands, heaths, fens and open woodlands. In this study, we used random amplified polymorphic DNA (RAPD) analysis to investigate the genetic differentiation among 25 populations of S. albicans from six different types of habitat (beech forests, alpine and lowland rocky ridges, lowland screes, fens, calcareous grasslands). With RAPD analysis, 344 fragments could be amplified, of which 95.9% were polymorphic. The level of polymorphism ranged from 29.7 to 56.7% polymorphic bands per population and was correlated with population size. In an analysis of molecular variance (AMOVA), used to detect variation among individuals within populations, among populations from the same habitat and among different habitats, most of the genetic variation was found within populations (62.06%) and among populations from the same habitat (33.36%). In contrast, only a very low level of differentiation could be observed among different habitats (4.58%). The results of our study give only little evidence for an ecotypic differentiation of Sesleria albicans. This differentiation is principally conceivable, but obviously not related to the investigated RAPD loci.     

Characterization of microsatellite loci in Coenonympha hero (Lepidoptera: Nymphalidae)

Microsatellite markers are suitable tools for studying dispersal pattern among local populations. I report on the characterization of seven polymorphic microsatellite loci in the scarce heath butterfly (Coenonympha hero), from one unenriched and one enriched genomic library. Number of alleles ranged from two to 20 when 108 individuals from seven populations were screened. H_O ranged from 0.140 to 0.889. Primers were also tested on three other butterfly species. Amplification was successful for all loci in Erebia triaria, while only one gave products in Maculinea alcon and Maculinea rebeli. To my knowledge, these are the first microsatellite markers published in the Nymphalidae subfamily Satyrinae.     

Vegetative Production and Performance of

Aspects of the production and vegetative performance of Calluna vulgaris (heather) were examined in four areas of New Zealand between 1981 and 1983; Tongariro National Park in the North Island, together with Mount Cook National Park, the Wilderness Scientific Reserve, and Ben Callum peat bog in the South Island. The height and height/width quotient of Calluna bushes, the diameter increment of woody stems and the amount of flowers/stem all decreased with altitude. Other age-adjusted components of stem biomass were positively related to soil depth, soil moisture, soil organic content and soil pH. Biomass by direct harvest gave values between 300 g per m and 2200 g per m for plant communities with Calluna. These are similar Values to recorded from British and European heaths, although the contribution of Calluna to the total biomass was often less in New Zealand. Calluna from New Zealand has a higher proportion of wood relative to total biomass than is normally found in Britain and Europe and this is probably related to the management of British and European heaths by burning. Multiple regressions indicated that increased woodiness is associated with lower altitudes and deeper, moister soils. Woodiness tended to decrease on more organic and more acidic soils, with a corresponding increase in the proportion of green shoots. Flowering declined with increased altitude, while the proportion of green shoots relative to total biomass increased. It is suggested that the upward extension of Calluna within Tongariro National Park is limited mainly by constraints on flowering and sexual reproduction.