Protecting more than the wetland: The importance of biased sex ratios and habitat segregation for conservation of the Hine’s emerald dragonfly, Somatochlora hineana Williamson

Within species habitat use may depend on age, season or sex of an individual. The distribution of males and females may vary both temporally and spatially due to differences in the costs of reproduction and the distribution of critical resources. Conservation of a species requires knowledge of the habitat use of both sexes in order to predict the population size and protect all habitats that a species requires. Adult dragonfly populations often have highly male-biased sex ratios at the breeding habitat. This bias has been attributed to females using alternative habitats to avoid male harassment, or to high female mortality. We monitored adult Hine’s emerald dragonfly (Somatochlora hineana Williamson) populations, in breeding and non-breeding habitats in Door County, Wisconsin and found significant differences in habitat use between males and females. Males primarily used wetland habitats, while females primarily used dry meadows and marginal breeding habitats, only coming into wetlands to lay-eggs or find mates. We assessed food resources in the different habitats and found that high quality insect prey (primarily adult Diptera) were more available in the wetland habitat, indicating that these areas were likely a more productive foraging area for adult dragonflies. The fact that females appear to avoid the wetland habitat is consistent with the hypothesis that male harassment alters female distribution patterns. Consideration of the patterns of habitat use by S. hineana indicates the need to develop a broader understanding of the importance of non-wetland areas in the conservation of wetland species.     

Saiga antelope calving site selection is increasingly driven by human disturbance

Many terrestrial mammalian species aggregate to give birth. Such aggregations are likely to be a response to changing resource and water availability, for predator swamping and avoidance of disturbance. The critically endangered saiga antelope (Saiga tatarica) is one such species. We analysed spatio-temporal locations of saiga calving aggregations in Kazakhstan over the last four decades obtained from aerial and ground surveys, to identify the factors determining the selection of calving sites within the species’ range as well as any changes in these locations over time. Generalized mixed models were employed in a use - availability framework to assess the factors distinguishing calving from random sites and predict suitable areas for calving. Saigas selected sites, with lower than average productivity and low year to year variability in productivity, at an intermediate distance from water sources, and away from human settlements. A significant change in calving locations was observed during the last decade, with calving areas occurring further north and further away from settlements than previously. The results demonstrate that the choice of calving areas is largely driven by environmental factors. However, disturbance also has a significant impact on calving site selection and in recent decades, its influence overrides that of environmental factors. This increase in the influence of disturbance coincides with a precipitous decline in saiga numbers due to poaching, as well as substantial reductions in the intensity of land use for livestock grazing following the breakup of the Soviet Union. Predictive models based on such studies can improve species conservation by guiding the stratification of sampling for effective monitoring and deployment of rangers to protect the females at this critical time.     

The habitat requirements of the rufous treecreeper (Climacteris rufa). 1. Preferential habitat use demonstrated at multiple spatial scales

Determining the habitat requirements of a species is fundamental to effective conservation, particularly if the species is declining in areas where its habitat is being modified. Multi-scaled investigations of habitat use are essential because different selection processes may operate at different scales. I examined the habitat use of a declining woodland passerine, the rufous treecreeper (Climacteris rufa), at three spatial scales (landscape, woodland and territory) in the wheatbelt of Western Australia. Preferential habitat use was exhibited at all scales. At the landscape scale, wandoo (Eucalyptus wandoo) woodland was used at a significantly greater rate than three other common vegetation types. Territory use within woodlands was positively related to the density of hollow-bearing logs, the density of nest sites, and tree age. Within an individual territory, nest sites (hollows) were favoured if they had a spout angle of ?50° to the horizontal and an entrance size of between 5 and 10 cm. The rufous treecreeper preferentially used habitat with traits characteristic of old-growth wandoo woodland. Degradation of wandoo through habitat modification (e.g. grazing, logging, fire and removal of deadwood) represents a significant threat to the persistence of treecreepers.     

Impacts of urbanization on Florida Key deer behavior and population dynamics

Rapid human population growth and urbanization have had a negative impact on species biodiversity. As competition for resources between man and wildlife continues, it is important to understand the effects of urbanization on species. Endangered Key deer (Odocoileus virginianus clavium) are endemic to the Florida Keys which have undergone rapid human population growth and development over the past 30 years. Our study objectives were to evaluate the impacts of urban development on Key deer habitat use, population dynamics, behavior, and body mass. We used data from two comprehensive studies on Key deer spanning 30 years to evaluate these changes. Our results suggest that Key deer have become more urbanized, using urban areas more today than they did 30 years ago. Contrary to our predictions, survival was higher for more urban deer than for less urban deer. Problems still exist with mortality factors heavily impacting some portions of the deer population including lower survival associated with less urban male deer. Analysis of Key deer body mass also was converse to our predictions as deer weights appear to have increased over time. Collectively, our results suggest that over the past 30 years Key deer have become more urbanized and that deer plasticity has allowed them to adapt and persist in an urbanizing environment. However, the future ability of Key deer to persist in an environment with continued urban development is unknown. At some threshold, urban development would become unsustainable and unlike other forms of habitat change or environmental disturbances, urban development is in most cases irreversible requiring careful planning in habitat conservation strategies.     

Reed harvest and summer drawdown enhance bittern habitat in the Camargue

The vulnerable Eurasian bittern Botaurus stellaris has recently benefited from conservation actions in several European countries. The Camargue in Mediterranean France holds 8000 ha of reed marshes and 50% of the French bittern population estimated at 300-350 booming males. Although reed cutting in Europe is a current practice, we provide the first analyses on habitat selection relative to reed exploitation. The contribution of reed harvesting and water management was assessed at a fine scale (digitized aerial photographs of 50-cm resolution) based on multi-modelling data analyses. GLMs applied to random and bittern points of 50-m radius showed that bitterns preferred areas with less than 10% open water enclosing 1-year reeds. Only in harvested reedbeds had the water management a significant impact on habitat selection, with bitterns avoiding areas dried during less than two months and flooded since less than 8 months. Water edge played a role in non-harvested reeds only, with bittern seeking for areas with the highest ratio water edge to open water. Although reed cutting is generally considered as detrimental in the short term, two 4-year experiments showed a marked increase in bitterns the first spring after cutting was interrupted. Based on these results, we propose various management scenarios that conciliate economic constraints of reed harvesters and protection of Eurasian bittern populations.     

Habitat associations of bats in Northern Ireland: implications for conservation

Agricultural intensification in Northern Ireland has brought about large-scale changes to the landscape with a detrimental effect on biodiversity. Between 1996 and 1998, we surveyed a stratified random sample of 1 km squares for bats using a spot-sample technique and time expansion bat detector to establish linear and area habitat associations. Bats strongly selected water bodies with bankside vegetation, treelines, and deciduous and mixed woodland edge, avoiding open areas such as upland/unimproved grassland and improved grassland. Of three sympatric pipistrelle species (Pipistrellus pipistrellus, P. pygmaeus and P. nathusii), two were shown to forage in different habitats; P. pipistrellus being more of a ‘generalist’, foraging in a wider variety of habitats than P. pygmaeus, a ‘specialist’ which foraged in a smaller range of habitats. Nyctalus leisleri selected parkland/amenity grassland, deciduous woodland edge and rivers/canals and avoided improved grassland, hedgerows, whilst Myotis spp. selected deciduous woodland centre, lake/reservoir margins and rivers/canals. Reduction in area and quality of inland water, deciduous woodlands and field boundaries in Northern Ireland may impact on bat populations. Habitat management should focus on improving those habitats which are selected by bats, including the maintenance and enhancement of connecting linear habitats.     

Habitat selection by the pale-headed brush-finch (Atlapetes pallidiceps) in southern Ecuador: implications for conservation

The pale-headed brush-finch (Atlapetes pallidiceps) is threatened with extinction due to habitat loss, but very little is known about its ecological requirements. We used multiple logistic regression to study habitat selection of this species at landscape, territory, and nest site scales in order to make recommendations about effective management. Habitat selection by the sympatric stripe-headed brush-finch (Buarremon torquatus) was examined with the same methods in order to analyse interspecific resource partitioning and potential competition. The pale-headed brush-finch selected semi-open habitat types with intermediate scrub heights, and avoided forests. Nest sites depended on the presence of vines or bamboo. By contrast, the stripe-headed brush-finch chose dense habitat with low ground cover under tall vegetation and avoided semi-open habitat. The two species had overlapping territories but differed significantly in microhabitat use and the use of vegetation strata. We found no convincing evidence that the stripe-headed brush-finch displaces the pale-headed brush-finch from optimal habitat. The preservation of semi-open scrubland maintained by low-intensity grazing is suggested for future conservation of the pale-headed brush-finch.     

Impacts of grazing on montane heath vegetation in Wales and implications for the restoration of montane areas

The impact of long-term heavy grazing pressure on montane vegetation and soils was investigated on the Carneddau plateau in North Wales. At this site, historical information suggests that formerly extensive Racomitrium- and Vaccinium-dominated heaths have become degraded and reduced in extent over the last 40-50 years. This coincides with a period when both sheep numbers and atmospheric deposition of nitrogen have greatly increased. Vegetation composition, plant tissue chemistry, soil chemistry and sheep presence were compared between areas of ‘good’ condition Racomitrium and Vaccinium heath and degraded areas. Comparison of species composition data with sheep presence and environmental data showed no significant link between current grazing distribution and species composition in the areas investigated, with all habitats having sheep presence above the recommended maximum for this habitat. Soil chemistry showed several differences between degraded and ‘good’ condition habitats, suggesting that degraded areas had experienced loss of soil organic matter and upper organic horizons. Comparison of soil and vegetation nitrogen content with data from other UK sites suggests a build up of nitrogen has occurred in Carneddau vegetation and soils, possibly related to high levels of nitrogen deposition. It is concluded that restoration of severely degraded areas of montane vegetation may be retarded by deleterious changes in soil properties which could slow recolonisation by montane species. Reductions in stocking density aiming to restore montane communities might have more immediate impact on less degraded areas.     

The habitat requirements of the rufous treecreeper (Climacteris rufa). 2. Validating predictive habitat models

Examining the predictive capability of statistical models with data independent from that used to derive the model is a vital step in the iterative procedure of assessing model performance. I derived logistic regression models of the habitat use of the rufous treecreeper (Climacteris rufa) at two spatial scales: woodland (territory selection model) and territory (nest-site selection model). The performance of these models was assessed in relation to the original data collected and validated with new, independent data. When applied to the original data, the territory model had a high predictive capability correctly classifying 90% of sites (n=100) that were either occupied or unoccupied by treecreepers. Correct classification rate was reduced to 70% (n=50) when the model was applied to the validation data. Model performance was generally robust when probability of occurrence values for the species were varied. In contrast, the nest-site model had lower predictive capabilities correctly classifying between 66 and 68% of sites, and performed relatively poorly when probability values were varied. The performance of the models differed slightly between the original and validation data, and substantially between the spatial scales examined. Territory use by rufous treecreepers could be predicted with some confidence indicating that the territory model may be a useful tool for habitat management. Nest-site use could not be predicted with confidence probably as a result of the high abundance of suitable, but unused, nest sites in the study area.     

Reproduction and pre-dispersal survival of Iberian lynx in a subpopulation of the Doñana National Park

Little is known about the reproduction of the Iberian lynx (Lynx pardinus) even though it is the most endangered felid in the world. We studied during 9 years several reproductive parameters of the Iberian lynx in one of the subpopulations situated in Doñana National Park (south-western Spain), by means of radio-tracking, direct observations and photo-trapping. The potential breeding subpopulation was usually composed of 3 adult females, which bred 83% of the total possible 29 female-reproductive year. The minimum total number of cubs born during the study was 64 (7.7 ± 0.69 per year). There was no correlation between the number of cubs born or number of breeding females and population size of European rabbit (Oryctolagus cuniculus, the main prey of the lynx). All known births (n = 16) occurred in March except one in April and another in June. Mean litter size was 3.0 ± 0.16 (n = 16, range = 2-4). Sex-ratio (n = 59 cubs) was 1.03:1.00 (females:males). Nevertheless, the commonest picture was a female with 2 cubs older than 3 months. Altogether, at age of 3 months, 75% of cubs survived. Number of lynx alive at 10 months old and before dispersing was 69% and 57%, respectively. Sex did not affect survival for any age. Normally, it was not possible to know the causes of the death of cubs younger than 3 months. Survival at this age was not related with mother, mother age, or rabbit abundance. All females that bred were older than 3 years. The age of last reproduction was 9 years. For 3 females that were tracked during almost their complete reproductive life, the life time reproductive output was between 11 and 19 cubs. Iberian lynx reproductive parameters did not respond to wide changes in prey abundance during the study. Conservation plans considering the extraction of cubs with a low survival probability should be considered by managers, for instance, in translocation campaigns.     

Assessing disease risk at the wildlife–livestock interface: A study of Sierra Nevada bighorn sheep

Despite evidence that domestic sheep diseases threaten the persistence of bighorn sheep populations, the economic consequences of restricting domestic sheep grazing has polarized the debate, with some arguing that disease risk posed by domestic sheep has been exaggerated and grazing restrictions should be eased. We constructed a model to assess how different management strategies (grazing allotment closures, grazing time reductions, and reduced probability of stray domestic sheep) affect the risk of respiratory disease transmission from domestic sheep to endangered Sierra Nevada bighorn sheep, and to predict population-level impacts of an outbreak. Even when management strategies reduced risk of interspecies contact to less than 2% per year, our model predicted a 50% probability of a catastrophic respiratory disease outbreak during the next 10 bighorn sheep generations. If an outbreak occurs in the near future, the model predicts that the smallest Sierra Nevada bighorn sheep population would have a 33% probability of quasi-extinction. To eliminate all risk of contact and potential disease transmission, domestic sheep cannot be grazed on allotments that overlap with areas utilized by Sierra Nevada bighorn sheep. Where wildlife and domestic animal populations share limited habitat, and there is documented evidence of a substantial disease threat and extinction risk, stakeholders must recognize that the only way to eliminate contact and risk of disease transmission is to give priority to one species or the other. If conservation is the priority, difficult decisions will need to be made to balance trade-offs between economic livelihoods and species conservation.     

A priori valuation of land use for the conservation of black rhinoceros (Diceros bicornis)

The appropriateness of using carrying capacity (CC) estimates to indicate habitat utilisation for a particular species, and thus as a tool for conservation population planning, has been questioned. We argue individual fitness is driven by resource availability, and we therefore assume individuals select habitats with a higher quality, abundance, and availability of key resources. In the past such selection has been related to the CC of a habitat. We tested whether we can use CC estimates to indicate habitat selection by individuals using a selective forager, black rhinoceros Diceros bicornis, for which CC approaches underpin species conservation plans. We tested for correlation of individuals’ habitat selection with predicted CC values at three spatial scales of selection. Individual selection was not related to the value of the habitat according to our CC estimates for any of the three scales we tested at. We discuss how density-dependence, environmental variables, scale of selection, individual variation and intra- and inter-specific dynamics may have influenced these results. Following this, we question the use of a priori calculations of potential resource quality and abundance of habitats (CC estimates), which do not take into account the various factors that influence an animal’s selection of a habitat, as an indicator of species habitat selection. We raise caution regarding the use of such CC models to determine optimal population numbers for an area.     

Habitat use and spatial dynamics of takhi introduced to Hustai National Park, Mongolia

The successful introduction of captive bred takhi or Przewalski’s horse, Equus ferus przewalskii, into Mongolia in the 1990s is a good example of the benefits of ex situ conservation and one of the few examples of the recovery of an animal after it became extinct in the wild. This is also particularly interesting because virtually nothing was known about how takhi lived before they died out, and the introductions have enabled us to study how they have settled, and their ecology and behaviour within their former natural range. In this paper, we describe the movement, home range size and shape, and habitat use of takhi at one of the release areas, the 570 km² Hustai National Park in Mongolia. Harem home ranges varied between 129 and 2399 ha, with 80% core areas of between 61 and 1196 ha. There was no relationship between range size and harem size, or length of time since release. Initially, harems stayed near their release enclosures, but over time they established home ranges further away. There was little overlap between home ranges of different harems, but neither was there evidence of exclusive range use. The more nutritious vegetation at lower elevations was preferentially selected. Thus the present situation looks good, but, as the population continues to grow, we anticipate that there will be potential problems related to intraspecific competition for water and vegetation resources, and the potential for hybridisation with domestic horses belonging to the local people. We consider the time it may take for takhi to reach carrying capacity within Hustai National Park and emphasise that continual monitoring of the population is essential because interventional management is likely to be required in the future.     

Assessing habitat use of the endangered marine mollusc Patella ferruginea (Gastropoda, Patellidae) in northern Africa: preliminary results and implications for conservation

The limpet Patella ferruginea Gmelin, 1791, endemic to the Mediterranean, is the most endangered marine species on the list of the European Council Directive 92/43/EEC on the Conservation of Natural Habitat of Wild Fauna and Flora (1992). Its Mediterranean range has progressively contracted to a few restricted areas and the species is now threatened with extinction. Seventeen stations were sampled along the littoral zone on the coast of Ceuta, North Africa, for quantifying environmental factors (water movement, siltation and suspended solids), and 70 transects were selected for P. ferruginea sampling. The study revealed the presence of a well-established population of P. ferruginea with values of density and size of 0.67 individuals m⁻¹ ±0.96 and 48.94 mm±11.61 respectively (mean±standard error of mean). The biggest densities of P. ferruginea were found inside the harbour of Ceuta. The limpets were more abundant on artificial harbour stones than on natural rocky shores, and the areas under the highest human pressure were characterised by the lowest densities and the smallest specimens probably due mainly to the predation for food and fishing. Human pressure is probably the main contributing factor to the currently endangered status of P. ferruginea. Programmes of environmental education to avoid its collection for fishing, food or for fun as decorative objects should be conducted, and further experimental studies dealing with the reproductive biology of this species are needed to properly assess the future programmes of conservation.     

Sperm storage and low incidence of multiple paternity in the European pond turtle, Emys orbicularis: A secure but costly strategy?

The freshwater pond turtle, Emys orbicularis, has recently suffered from population declines throughout its range, mainly due to habitat destruction. The mating strategies of this species were studied using genetic data from successive clutches within and between years. To test for the occurrence and frequency of multiple paternity and sperm storage, genetic paternity at six microsatellite markers was assessed in 114 embryos and hatchlings from single and subsequent clutches of 11 females (including clutches from the same or consecutive years). Multiple paternity was rare and only found in two out of 20 clutches from 11 females. All annual successive clutches and 58% of the clutches in the next year, were fertilized with sperm from the same male. The use of stored sperm is thus a frequent strategy in E. orbicularis. However, hatching rate, hatchling mass, and hatchling length decreased in clutches fertilized by stored sperm, suggesting sperm depletion or deterioration through time. The occurrence of stored sperm despite an associated reduced reproductive output indicated that mating and/or the fertilization process is costly to females. The low incidence of multiple paternity may simply be the residual consequence of the capacity to store viable sperm. These results provide important and innovative insights for the conservation of E. orbicularis. In threatened populations, management strategies may aim to enhance effective copulations in order to increase the reproductive output of females.     

Managing landscapes for the little bustard Tetrax tetrax: lessons from the study of winter habitat selection

More than half of the world's population of the little bastard Tetrax tetrax lives in the Iberian Peninsula, where it is mostly dependent for survival on extensive agricultural areas. The species has declined dramatically, chiefly due to changes in agricultural practices, and is now globally ‘Near Threatened’. Knowledge of its habitat requirements is crucial to reverse the trend of habitat deterioration. Winter habitat preferences were studied in a region dominated by extensive cereal farming in Southern Portugal, comparing the characteristics of sites used by 54 flocks with those of randomly selected sites within the study area. The birds preferred recent fallows and grassy vegetation of mid-size (about 11-20 cm) and mid-density (about 11-50% cover). They tended to concentrate on the tops of hills and to avoid disturbed areas near roads and houses. Overall, the observed preferences suggest that predator avoidance is a significant factor in habitat selection. To improve habitat suitability for little bustard, managers of key wintering areas should minimize permanent sources of human disturbance, encourage rotations with frequent fallows, and favour moderate levels of grazing to manipulate vegetation height and cover. Particular attention should be given to the upper parts of the hills.     

Linking process to pattern: Causes of stream-breeding amphibian decline in urbanized watersheds

Sufficient data have been collected to document negative effects of urbanization on many abiotic parameters, aquatic insects, and fish. Recently, stream-breeding salamanders have been observed to decrease in density in urban areas. We employed a two-step process to determine the mechanism(s) most likely responsible for salamander decline in disturbed streams. First, we conducted a 2-year survey of larval two-lined salamanders (Eurycea cirrigera) and abiotic variables among 12 streams in western Georgia, USA surrounded by land disturbed by urbanization and pasture. From these surveys we constructed hypotheses that might explain differences in observed larval salamander density among streams. A model in which increased watershed impervious surface causes an increase in spate (i.e., high water flow) frequency and magnitude, which then leads to decreased larval density had the most support. This analysis led to a second step in which we conducted an experiment using artificial streams lined with substrates commonly found in urban and forested streams (sand/sand with detritus and gravel-pebble/pebble-cobble, respectively). We placed salamander larvae in the artificial stream channels and subjected them to increasing water velocity. Larvae on sand-based substrates common to urban streams were flushed from the artificial stream at significantly lower velocities than larvae on rock-based substrates. The experimental data were consistent with the hypothesis generated from field sampling that high water velocities in urban streams result in decreased larval retention in streams. Our result emphasizes the need for upland habitat restoration in urban areas, which will reduce run-off and improve stream habitat quality.     

Disturbance from roads negatively affects Andean condor habitat use

Human disturbance can be a severe problem for some animal species. Behavioural ecology theory predicts that sensitivity to mortality risks, and thereby to disturbance, will be related to life-history characteristics. Long-lived species with low reproductive rates are expected to have a high cost of predation and therefore avoid risks strongly. In this paper we quantify the effect of roads on the Andean condor (Vultur gryphus) using behavioural indicators. We used a pair wise design, offering condors standardized food patches near and far from roads simultaneously. The patches consisted of dead adult sheep. We recorded condors’ behaviour when visiting each patch. This study establishes the versatility of behavioural indicators and their usefulness to study conservation problems. We found that, although near and far patches were discovered after similar times and there were similar numbers of condors in the vicinity of near and far patches, the use of patches differed strongly. In the patches far from roads many more condors came to feed, the average time spent per individual was longer, the proportion of time spent vigilant was lower, and the amount of food left uneaten on the carcasses was lower. These results strongly suggest that Andean condors clearly avoid roads, and behave in a way predicted from theory if they trade off safety for energy. This study shows that roads have stronger influence on the habitat use of Andean condors, than previously thought, and may be a significant problem for the species in populated areas.     

Habitat selection of endangered and endemic large flying-foxes in Subic Bay, Philippines

Large flying-foxes in insular Southeast Asia are the most threatened of the Old World fruit bats due to high levels of deforestation and hunting and effectively little local conservation commitment. The forest at Subic Bay, Philippines, supports a rare, large colony of vulnerable Philippine giant fruit bats (Pteropus vampyrus lanensis) and endangered and endemic golden-crowned flying-foxes (Acerodon jubatus). These large flying-foxes are optimal for conservation focus, because in addition to being keystone, flagship, and umbrella species, the bats are important to Subic Bay’s economy and its indigenous cultures. Habitat selection information streamlines management’s efforts to protect and conserve these popular but threatened animals. We used radio telemetry to describe the bats’ nighttime use of habitat on two ecological scales: vegetation and microhabitat. The fruit bats used the entire 14,000 ha study area, including all of Subic Bay Watershed Reserve, as well as neighboring forests just outside the protected area boundaries. Their recorded foraging locations ranged between 0.4 and 12 km from the roost. We compared the bats’ use to the availability of vegetative habitat types, riparian areas, and bat trees. The fruit bats’ locations showed a preference for undisturbed forest types and selection against disturbed and agricultural areas. Bat locations also showed selection for particular fruiting/flowering bat trees. The bats showed strong preference for riparian areas; locations were in riparian areas over four times more than expected. From these results we recommend that management focus flying-fox conservation efforts on undisturbed forest and riparian areas.     

Rehabilitation of degraded Calluna vulgaris (L.) Hull-dominated wet heath by controlled sheep grazing

Many upland areas of the British Isles have seen declines in the area and condition of heather (Calluna vulgaris)-dominated heathland vegetation. To reverse this decline, management regimes must be designed to rehabilitate areas that have seen this decline. As most of this heathland vegetation is primarily managed by grazing, such management has to determine what stocking levels can maintain the vegetation in a desired state. This paper describes how to reverse this decline through suitable grazing management.A degraded ‘wet-heath’ system, previously grazed at 2.1 sheep ha⁻¹, was subject to a range of grazing treatments over a 5-year period. Treatments varied in intensity (0-1.4 sheep ha⁻¹) and timing (summer only, winter only, or year round) of grazing. Grazing levels were maintained at 2.1 sheep ha⁻¹ outside the fenced areas. Vegetation composition remained stable outside the fenced treatments. All the fenced treatments showed an increase in the relative frequency of the evergreen Calluna vulgaris, with the greatest increase being in the ungrazed treatment, and the least in the year round 1.4 sheep ha⁻¹ treatment. This increase was in line with a reduction in heather utilisation to relatively low and sustainable levels. Other species that benefited from reduced grazing included Carex nigra, Deschampsia flexuosa and to a lesser extent Galium saxatile and Erica tetralix, whereas a range of moss species including Hypnum jutlandicum and Rhytidiadelphus loreus were more frequent at higher grazing levels. Though the recovery of heather was similar in the two seasonally grazed treatments, the vegetation showed different overall trajectories. Winter only grazed allowed a substantial increase in the cover of the deciduous Molinia caerulea, whereas this species was kept in check by summer only grazing. A stocking level of between 0.7 and 1.4 sheep ha⁻¹ appears to be appropriate to maintain and even enhance the cover of heather on degraded wet heath. Complete removal is not necessary. Grazing restricted to the winter period is inappropriate in areas where M. caerulea occurs. Setting appropriate stocking levels to maintain the condition of the vegetation must take into account site conditions, especially the presence of species that can affect the utilisation of heather.