Woodland dynamics under the influence of elephants and fire in Northern Botswana

Sustained elephant browsing and intense burning could result in the loss of woodlands under conditions where elephant densities are high, such as in northern Botswana. Three woodland types dominated by Acacia erioloba, Baikiaea plurijuga and Colophospermum mopane were monitored in plots and contemporary recruitment rates of woody plants were compared with the associated local elephant densities and fire occurrences. Woodland types differed with respect to structure, extent of elephant damage and the occurrence of fire. Canonical correlations indicated that high extent of fire damage and high elephant densities did not covary within the woodland types investigated. Low tree densities in some woodland types were associated with high elephant densities and new elephant damage to plants increased with high elephant densities during the dry season. Plots with an apparent high fire frequency had lower tree densities and higher cover abundance of shrubs and seedlings.The annual rates of tree recruitment/loss in each woodland type were estimated through a model based on observed seedling recruitment, mortality and reversal to lower height classes due to combinations of fire occurrence and elephant browsing. The model suggested that elephants induce tree loss in woodlands dominated by plant species which are principal food sources. Fire however, seems to have a widespread effect across woodlands which could result in extensive tree loss.     

Is fencing enough? The short‐term effects of stock exclusion in remnant grassy woodlands in southern NSW

Summary Fencing remnant native vegetation has become a widespread activity for arresting declines in biodiversity in agricultural landscapes. However, few data are available on the effectiveness of this approach. The present study investigated the short-term effects of fencing to exclude livestock on dominant tree and shrub recruitment, plant species cover, litter and soil characteristics in remnant grassy woodlands in southern NSW. Vegetation and soil surveys were undertaken at 47 sites fenced by Greening Australia (NSW) for 2–4 years. Fenced and unfenced areas at each site were compared using split-plot sampling. Woodlands sampled were dominated by Yellow Box/Blakely's Red Gum ( Eucalyptus melliodora/Eucalyptus blakelyi ), Grey Box ( Eucalyptus microcarpa ) or White Cypress-pine ( Callitris glaucophylla ). Significantly higher numbers of tree recruits were found in the fenced sites, with tree recruitment found in 59% of fenced sites compared with 13% of unfenced sites. Fenced sites also had significantly greater cover of native perennial grasses, less cover of exotic annual species and less soil surface compaction. However, outcomes varied among woodland ecosystems and individual sites. Where tree recruitment occurred, there was significantly more tree recruitment where there was greater perennial grass cover and less regeneration where exotic annual grass cover or overstorey crown cover was dense. Few shrubs recruited in fenced or unfenced areas, reflecting the lack of mature shrubs in most sites. Fencing is an important first step for conserving threatened grassy woodlands, but more active management may be needed to enhance woodland recovery, particularly in sites where few or no recruits were found.
Key words bush regeneration, fencing, grazing exclusion, rehabilitation, woodland restoration.     

Coarse woody debris can reduce mammalian browsing damage of woody plant saplings in box‐gum grassy woodlands

The critically endangered box‐gum grassy woodlands of south‐east Australia face numerous threats including the failure of woody plant regeneration caused by over‐browsing. In the Australian Capital Territory, over‐browsing of tree and shrub saplings is likely caused by dense populations of Eastern Grey Kangaroo (Macropus giganteus) found in many nature reserves free of livestock. One possible way to protect these saplings is using coarse woody debris (CWD) as a browsing deterrent. We tested this idea by planting palatable Red Stemmed Wattle (Acacia rubida) saplings among manually applied CWD, among naturally fallen CWD, and in the open, in five woodland reserves. We recorded the proportion of saplings browsed, the number of weeks to first browsing and the browsing severity (sapling height lost). Applied CWD protected saplings from being browsed only at relatively low‐to‐moderate kangaroo browsing pressure (as measured by faecal pellet counts). At relatively high browsing pressure, the probability of a sapling being browsed among applied CWD was 100%, similar to the probability in the open treatment (no CWD). Natural CWD, in contrast, provided some protection even at high browsing pressures. Time to browsing was most affected by browsing pressure, although CWD cover also had an influence. Browsing severity was similar between the three treatments and was only affected by browsing pressure. These results indicate that without protection, palatable woody plant saplings have a high chance of being browsed by kangaroos in woodland reserves, and therefore, some protection is needed for successful regeneration. The CWD being applied to reserves has a limited capacity to protect regenerating saplings. If more protection is wanted a CWD structure more resembling natural fallen timber should be used. This could be done by artificially placing branches around plantings. However, the most important action to facilitate regeneration is to manage kangaroo populations to reduce overall browsing pressure.     

Limitations to seedling establishment in a mesic Hawaiian forest

While invasive species may be visible indicators of plant community degradation, they may not constitute the only, or even the primary, limitation to stand regeneration. We used seed-augmentation and grass-removal experiments under different canopy conditions to assess the relative importance of dispersal limitation, resource availability, and competition on seedling establishment in the understory shrubs Sophora chrysophilla, Dodonea viscosa, and Pipturus albidus in a montane mesic forest in Hawaii. The study location was an Acacia koa-Metrosideros polymorpha forest at 1000–1500 m elevation on the leeward side of Hawaii Island; it is a closed-canopy forest historically subject to logging and grazing by cattle and sheep and currently dominated by the exotic grass, Ehrharta stipoides, in the herb layer. Seedling establishment after 1 and 2 years was strongly dispersal limited in Sophora and Dodonea, but not in Acacia, a non-augmented species in which abundant seedlings established, nor in Pipterus, in which only one seedling established in 2 years. Grass cover reduced seedling establishment in Acacia, Sophora, and Dodonea and, for the latter two species, seedling establishment was substantially greater in the warmer, more moist forest at the lowest elevation. Light, moisture, and resin-captured N and P were more strongly affected by elevation and canopy composition than by grass cover, but in most cases seedling establishment was not positively correlated with resource availability. Limitations to the establishment of woody seedlings in this forest-grassland mixture vary among species; however, both dispersal limitation and competition from a shade-tolerant grass are important deterrents to regeneration in these forests.     

Facilitation by herbivore-mediated nurse plants in a threatened tree, Taxus baccata: local effects and landscape level consistency

We investigated the regeneration of a threatened tree, the yew Taxus baccata, in relation to the presence of fleshy‐fruited woody plants acting as seed dispersal foci as well as protecting yew recruits against ungulate herbivores. We seek to determine if local facilitative effects are consistent across landscape in the Cantabrian range (NW Spain). Yew seed rain by birds mostly concentrated under yew trees and beneath hollies Ilex aquifolium. Seedling emergence distributed similarly to seed rain, but first‐year seedling survival was higher beneath hollies. In one site where woody vegetation was structured as nucleation centres (multispecific patches of fleshy‐fruited plants acting as foci for seed rain) yew recruits mostly occurred in yew‐dominated centres, suggesting dispersers‐mediated facilitation. However, holly was the main nurse plant for most of these recruits, considering the nurse as the species whose canopy covered directly the yew recruit. Living beneath nurse plants reduced herbivore damage on saplings and enhanced seedling survival. A planting experiment with yew rooted‐cuttings beneath different spiny shrubs corroborated this effect. Additional evidence on yew recruitment limitation by herbivory emerged from one population where ungulates were fence‐excluded. Our results suggest that nurse plants mitigate the negative effect of herbivores on yew regeneration, by providing defence against browsing and trampling. Shelter ability related to nurse structure, cone‐shaped shrubs with branches at their bases acting better as a barrier. Paradoxically, this structure resulted from heavy browsing on nurse plants. The study of yew regeneration and habitat structure in seven sites provided evidence for the consistency of facilitation by holly at the landscape level, since local values of yew recruitment positively related to nurse ground cover. Range‐scale yew management must consider the local functioning of the interaction among avian seed‐dispersers, nurse fleshy‐fruited plants and ungulate herbivores, in combination with regional measures, targeting the habitats where facilitation emerges.     

Woodlands on farms in southern New South Wales: A longer-term assessment of vegetation changes after fencing

Summary Fencing incentive programmes have been widely used throughout Australia to assist landholders to fence remnant woodland vegetation, to control grazing and improve native vegetation condition. This study investigated vegetation and soil condition in remnant woodlands fenced for 7–9 years in the Murray catchment area in southern New South Wales. Surveys were undertaken at 42 sites, where vegetation condition was assessed in paired fenced and unfenced sites. Semi‐structured interviews were also conducted with landholders to gather management information. Woodlands surveyed were Yellow Box/Blakely's Red Gum (Eucalyptus melliodora/E. blakelyi, 15 sites), Grey Box (E. microcarpa, 13 sites) and White Cypress Pine (Callitris glaucophylla, 14 sites). Fencing resulted in a range of responses which were highly variable between sites and vegetation types. In general, fenced sites had greater tree regeneration, cover of native perennial grasses, less cover of exotic annual grasses and weeds, and less soil compaction than unfenced sites. However, there was greater tree recruitment in remnants to the west of the study area, and tree recruitment was positively correlated with time since fencing. Within sites, tree recruitment tended to occur in more open areas with a good cover of native perennial grasses, as compared to sites with a dense tree canopy, or dominated by exotic annuals grasses or weeds. Forty‐eight per cent of fenced sites had no tree regeneration. There was a significant decline in native perennial grasses, and increase of several unpalatable weeds in many fenced areas, suggesting certain ecological barriers may be preventing further recovery. However, drought conditions and associated grazing are the most likely cause of this trend. A range of grazing strategies was implemented in fenced sites which require further research as a conservation management tool. Continued long‐term monitoring is essential to detect key threats to endangered woodland remnants.     

Elephant herbivory in Majete Wildlife Reserve,Malawi

Elephants have a major influence on vegetation structure, composition and ecosystem processes, and are primary agents of habitat change in Africa. At moderate‐to‐high population densities, elephants can damage vegetation, especially when enclosed in protected areas. This study examines the effects of elephant browsing on woody trees in Majete Wildlife Reserve (WR), south‐western Malawi. Regression analysis is used to assess the associations of six factors known to drive elephant browsing in other areas and determine which ones have the most influence on browsing at Majete WR. Twenty‐four per cent of tagged trees had been subject to elephant browsing. The model with vegetation type, stem diameter and distance from permanent water correctly predicted browsing for 80% of the observations. Elephants mostly favoured riparian woodlands, followed by Acacia‐dominated woodland and Brachystegia‐dominated woodland. Browsing occurrence was negatively related to distance from permanent water and diameter at breast height(DBH). A larger number of trees, sampled at random and covering a larger portion of the reserve would provide more reliable estimates of browsing and related factors. Knowledge of time‐ and site‐specific factors affecting elephant browsing can be used to forecast future habitat transformations and manipulate the range of the elephants within the reserve.     

Remotely‐sensed foliage cover and ground‐measured stand attributes are complimentary when estimating tree hollow abundances across relictual woodlands in agricultural landscapes

Hollows, also known as tree cavities, are critical to the survival of many animal species but are too poorly mapped across landscapes to allow for their adequate consideration in regional planning. Managing cost is important, so we tested whether freely available satellite‐derived foliage projective cover and field‐measured stand attributes could be used separately or combined to predict tree hollow abundance in relictual Australian temperate woodlands. Satellite‐derived foliage projective cover revealed variation in woody vegetation densities both within mapped woodland remnants and cleared areas of the agricultural matrix. Plot‐based field assessment of the actual number of hollows in each one‐hectare site (n = 110 sites) revealed a relationship with foliage cover. Improvement of the model was achieved if site‐based estimates of the proportion of the canopy due to Eucalyptus species and the number of mature trees per hectare were included. Remotely sensed foliage cover can improve on traditional vegetation mapping for predicting hollow‐bearing tree and hollow abundances at landscape scales when managing hollow‐dependent fauna habitat across relictual woodlands in temperate Australian agricultural landscapes. At finer scales, the addition of other predictors is necessary to raise the accuracy of the predicted hollow densities.     

Vegetation Control Treatments to Favor Naturally Regenerated Betula alleghaniensis Saplings Following Seed-Tree Cut: Sapling Monitoring Two Years after Treatment

Control of competing vegetation is recommended to ensure successful Yellow birch (Betula alleghaniensis Britton) regeneration within juvenile stands that do not sustain high enough sapling densities of this species. Four contrasting vegetation control treatments were tested to determine their effect on the growth and vigor of eight‐year‐old B. alleghaniensis saplings regenerating after final cutting of a shelterwood seed cut. Vegetation control treatments were TC (total circular removal), PC (circular removal of codominant competing vegetation), TS (total semicircular removal on 180° section), and NC (no vegetation control). Two years after treatment application, diameter growth significantly improved in response to vegetation control treatments, whereas sapling height growth did not. This pattern of biomass allocation was directly related to sapling etiolation, which increased with decreasing severity of vegetation removal. As a result, application of vegetation control, especially TC and PC treatments, was valuable in reducing signs of stress in saplings. However, increasing the severity of vegetation removal also made saplings more conspicuous to herbivores, which increased browsing, especially in the TC and PC treatments. Browsing was sufficient in some plots of the TC and PC treatments to overcome the vigor and diameter growth enhancements observed when browsing was negligible. In contrast to the TC and PC treatments, the TS treatment kept browsing very low while largely removing competition. The results suggest that B. alleghaniensis saplings established after final cutting of a shelterwood seed cut do take advantage of vegetation control treatments, but the decision to apply these treatments must include consideration of local herbivore population densities.     

Managing open habitats by wild ungulate browsing and grazing: A case‐study in North‐Eastern Germany

Question: Can wild ungulates efficiently maintain and restore open habitats? Location: Brandenburg, NE Germany. Methods: The effect of wild ungulate grazing and browsing was studied in three successional stages: (1) Corynephorus canescens‐dominated grassland; (2) ruderal tall forb vegetation dominated by Tanacetum vulgare; and (3) Pinus sylvestris‐pioneer forest. The study was conducted over 3 yr. In each successional stage, six paired 4 m²‐monitoring plots of permanently grazed versus ungrazed plots were arranged in three random blocks. Removal of grazing was introduced de novo for the study. In each plot, percentage cover of each plant and lichen species and total cover of woody plants was recorded. Results: Wild ungulates considerably affected successional pathways and species composition in open habitats but this influence became evident in alteration of abundances of only a few species. Grazing effects differed considerably between successional stages: species richness was higher in grazed versus ungrazed ruderal and pioneer forest plots, but not in the Corynephorus sites. Herbivory affected woody plant cover only in the Pioneer forest sites. Although the study period was too short to observe drastic changes in species richness and woody plant cover, notable changes in species composition were still detected in all successional stages. Conclusion: Wild ungulate browsing is a useful tool to inhibit encroachment of woody vegetation and to conserve a species‐rich, open landscape.     

Aspen and Willow Restoration Using Beaver on the Northern Yellowstone Winter Range

Aspen (Populus tremuloides) on the northern Yellowstone winter range has declined over the last half‐century. Beaver (Castor canadensis) were reintroduced in Eagle Creek in 1991 in an attempt to reverse this trend. In 2005, we assessed the efficacy of this project by quantifying the long‐term effects of beaver on aspen stands and the riparian area in this drainage. Between 1990 and 2005, the canopy cover of mature aspen decreased more than 62%, whereas immature aspen cover more than tripled, resulting in a total aspen canopy cover decrease (p < 0.05) from 43 to 25% (a loss of 7.25 ha). Willow canopy cover increased from 10 to 14% during the same period. The impacts of beaver on aspen stands were estimated by comparing vegetative changes among control sites with less than 10% beaver use (n = 5), active beaver sites (n = 6), sites inactive for 1–3 years (n = 7), sites inactive for 4–6 years (n = 4), and sites inactive for 7–11 years (n = 5). Aspen sprout and sapling densities were greater (p = 0.01) on sites which were active and inactive for 1–3 years compared to the other sites. Aspen ramets were not able to grow taller than 2 m on sites without beaver activity for 4–11 years due to ungulate herbivory. Although beaver stimulated the growth of aspen sprouts and saplings, ungulate herbivory prevented successful aspen recovery in the Eagle Creek drainage of the northern Yellowstone winter range 14 years after beaver reintroduction.     

Fine‐scale habitat preferences and habitat partitioning by three mycophagous mammals in tropical wet sclerophyll forest,north‐eastern Australia

Abstract Fine‐scale habitat preferences of three co‐occurring mycophagous mammals were examined in a tropical wet sclerophyll forest community in north‐eastern Australia. Two of the three mammal species responded to fine‐scale variation in vegetation and landform around individual trap locations. At a broad scale, the northern bettong (Bettongia tropica), an endangered marsupial endemic to the Australian wet tropics region, showed a preference for ridges over mid‐slopes and gullies, irrespective of forest type. In contrast, the northern brown bandicoot (Isoodon macrourus), a widespread marsupial, displayed a preference for Eucalyptus woodland over adjacent Allocasuarina forest, irrespective of topographic category. The giant white‐tailed rat (Uromys caudimaculatus), a rodent endemic to the wet tropics, showed no particular preference for either forest type or topographic category. A multiple regression model of mammal capture success against three principal habitat gradients constructed from 21 habitat variables using principal component analysis indicated strong species‐specific preferences for fine‐scale vegetation assemblages. Bettongs preferred areas of Eucalyptus woodland with sparse ground cover, low densities of certain grass species, high density of tree stems and few pig diggings. Bandicoots, in contrast, favoured areas in both forest types with dense ground cover, fewer tree stems and greater numbers of pig diggings; that is, characteristics least favoured by bettongs. The striking differences in fine‐scale habitat preferences of these two mammals of similar body size and broad habitat requirements suggest a high degree of fine‐scale habitat partitioning. White‐tailed rats did not show preference for any of the habitat gradients examined.     

Characteristics of woods used recently and historically by Lesser Spotted Woodpeckers Dendrocopos minor in England

Lesser Spotted Woodpecker Dendrocopos minor numbers have declined greatly in England since the early 1980s for reasons that are not yet fully understood. It has been suggested that the species’ decline may be linked to the increase in Great Spotted Woodpeckers Dendrocopos major, changes in woodland habitat quality (such as deadwood abundance) and landscape‐scale changes in tree abundance. We tested some of these hypotheses by comparing the characteristics of woods in southern England where the species is still relatively numerous with those of woods used in the 1980s before the major decline. In each time period, habitat, predator and landscape information from woods known to be occupied by Lesser Spotted Woodpeckers was compared with those found to be unoccupied during surveys. Before the main period of decline, Lesser Spotted Woodpeckers used oak‐dominated, mature, open woods with a large amount of standing deadwood. Habitat use assessed from recent data was very similar, the species being present in mature, open, oak‐dominated woodlands. There was a strong relationship between wood use probability and the extent of woodland within a 3‐km radius, suggesting selection for more heavily wooded landscapes. In recent surveys, there was no difference in deadwood abundance or potential predator densities between occupied and unoccupied woods. Habitat management should focus on creating and maintaining networks of connected woodlands in areas of mature, open woods. Finer‐scale habitat selection by Lesser Spotted Woodpecker within woodlands should be assessed to aid development of beneficial management actions.     

Deciduous woodland conservation under heavy deer browsing on Devils Tower National Monument, USA

Native deciduous woodlands comprise <5% of the vegetation in the Black Hills and northern plains of the USA and appear to be in decline. Because deciduous woodlands comprise 18% of the land cover on Devils Tower National Monument (DTNM), this natural area is potentially important for the conservation of these plant communities. Lack of tree regeneration and accumulation of dead wood suggest that the native deciduous woodlands on DTNM may die out under persistent, heavy deer (Odocoileus virginianus, O. hemionus) browsing. We monitored vegetation changes for seven years (1989–1996) inside and outside 6 deer exclosures on DTNM to determine the resilience of native woodland communities when protected from browsing, and to parameterize models that evaluate specific fencing scenarios for managing woodland communities. The herbaceous woodland understory was not directly, but indirectly affected by deer browsing because graminoid biomass declined as tall shrubs recovered. Tall shrubs, dominated by Prunus spp. showed the most rapid and consistent recovery across sites. Models of shrub growth indicated that rotational fencing for short durations (10 years) produced the highest long-term increase in tall-shrub densities and the greatest evenness among size classes. Because deciduous woodlands currently are resilient and occur in small patches, fencing may provide a practical management alternative to deer culling programs.     

Landscape of fear in Europe: wolves affect spatial patterns of ungulate browsing in Białowieża Primeval Forest,Poland

Large carnivores can either directly influence ungulate populations or indirectly affect their behaviour. Knowledge from European systems, in contrast to North American systems, on how this might lead to cascading effects on lower trophic levels is virtually absent. We studied whether wolves Canis lupus via density‐mediated and behaviorally‐mediated effects on their ungulate prey species influence patterns of browsing and tree regeneration inside the Bia?owie?a National Park, Poland. Browsing intensity of tree saplings (height class < 150 cm), irrespective of tree species or forest type, was lower inside a wolf core area (50.5%) where predator presence is highest, than in the remainder of the wolf pack’s home range (58.3%). Additionally, browsing intensity was reduced when the amount of coarse woody debris (CWD), which can act as a ?ungulate escape impediment?, increased (within 5‐m radius) inside the wolf core area. No relationship existed outside the core area. As a result, the proportion of trees growing out of herbivore control increased more strongly with increasing amount of CWD inside compared to outside the wolf core area. This suggests that next to direct effects of wolves on ungulate density caused by a higher predation pressure inside the core area, risk effects are important and are enhanced by habitat characteristics. These results indicate that behaviorally‐mediated effects of predators on prey can become more important than density‐mediated effects in affecting lower trophic levels. This is the first study we are aware of, that shows CWD can create fine‐scale risk effects on ungulates with the potential for cascading effects of large predators on patterns of tree regeneration for a European forest system. This knowledge broadens the discussion on how the impact of large predators on ecosystem functioning depends on the physical landscape, by illustrating these effects for a system which largely contrasts in this respect to the North American systems.     

Effect of selective logging on tree and understory regeneration in miombo woodland in western Tanzania

Tree regeneration and understory response to selective cutting of Pterocarpus angolensis DC, and Sterculia quinqueloba Sim, was measured outside Katavi National Park, Tanzania. Contrary to expectations, a selective harvest had no effect on tree recruitment for either species in this miombo woodland. In unlogged plots along transects taken at increasing distance from a main road, the stand densities of young trees declined away from the road. Since anthropogenic activity along the road results in a high fire frequency but low native ungulate densities, the increased rate of tree regeneration near the road may result from either reduced grazing and browsing pressure by native ungulates, or reduced fire intensity and severity along roads. These effects appear to mask any possible effect of selective harvest on tree recruitment. Currently, there is no evidence of compensatory recruitment of trees of any species into the canopy to replace logged trees and this will lead to a gradual thinning in overstory stand density.     

Influence of riparian cover on stream temperatures and the growth of the mayfly <Emphasis Type="Italic">Baetis rhodani</Emphasis> in an upland stream

This research aimed to assess whether stream temperature and the size of Baetis rhodani larvae (Insecta: Ephemeroptera) differed between sites with contrasting riparian vegetation cover. Stream temperatures and Baetis sizes were monitored at sites along a 5-km section of a small (30 km²) upland Scottish stream. Some sites had riparian tree cover dominated by mature broadleaved woodland, while others were located in areas of open Calluna moorland. The 13 month data period included two contrasting summers, one typical for the region and the other unusually warm. Daytime water temperatures were generally highest at the open moorland sites. The difference between moorland and broadleaved woodland sites was most apparent in the summer months, with the woodland shading apparently acting to reduce maximum temperatures and mediate temporal fluctuations. Size data suggest that Baetis has a bivoltine life-cycle in the stream. Significant differences in the mean size of Baetis were found between moorland and woodland sites, with differences occurring primarily during the summer months. Mean size of Baetis was significantly greater in the moorland during the time of summer adult emergence. During the months that followed (growth of the next generation), Baetis larvae in the woodland sites were significantly larger. Size distribution data for the population suggest earlier adult emergence in the moorland section and different egg-to-larvae hatch times. Overall, the results indicate that subtle differences in growth and life cycles, linked to differences in riparian cover and streamwater temperature, can occur over short distances in upland streams.     

The role of protective understorey in the regeneration system of a heavily browsed woodland

The paper tests an apparent relationship between young understorey of Ilex aquifolium and the presence of seedlings of the canopy species Fagus sylvatica and Quercus species, in heavily browsed woodland in the New Forest, southern England. A small block of woodland was searched and mapped, and the information collected subject to graphical and statistical analyses.Regeneration, though of low frequency, is present and shows non-random patterns of distribution. Older seedlings appear to be located in or near areas of understorey, or near fallen branches, logs, etc. Principal components analysis of woodland characteristics suggests six main habitat types are available to seedlings, and predictions are made about likely regeneration patterns associated with these habitats, according to a model based on the hypothesis of a protective understorey and heavy browsing pressure. Chi-square tests of actual seedling occurrences versus habitat types demonstrate marked associations of regeneration with non-trampled areas, including young Ilex understorey but also other forms of protective barrier. The association is more marked for larger seedlings.The implications for a wider understanding of regeneration processes and community dynamics in heavily browsed forest are discussed. It is concluded that regeneration theories appropriate to such forests are required, since the dominant gap-phase theories are not successful in predicting regeneration patterns when browsing is a major factor.     

Impacts of spruce budworm defoliation on the habitat of woodland caribou,moose, and their main predators

Forest logging has contributed to the decline of several woodland caribou populations by causing the fragmentation of mature coniferous stands. Such habitat alterations could be worsened by spruce budworm (SBW) outbreaks. Using 6201 vegetation plots from provincial inventories conducted after the last SBW outbreak (1968–1992) in boreal forests of Québec (Canada), we investigated the influence of SBW‐caused tree defoliation and mortality on understory vegetation layers relevant to woodland caribou and its main predators. We found a positive association between severe outbreaks and the cover of most groups of understory plant species, especially in stands that were dominated by balsam fir before the outbreak, where a high canopy openness particularly benefited relatively fast‐growing deciduous plants. Such increases in early successional vegetation could provide high‐quality forage for moose, which is likely to promote higher wolf densities and increase predation pressure on caribou. SBW outbreaks may thus negatively affect woodland caribou by increasing predation risk, the main factor limiting caribou populations in managed forests. For the near future, we recommend updating the criteria used to define critical caribou habitat to consider the potential impacts of spruce budworm defoliation.     

Age-states,population structure,and seedling regeneration of Chionochloa pallens in Canterbury alpine grasslands,New Zealand

Abstract. Age-states, population structure, and seedling regeneration of the grass Chionochloa pallens were investigated in eight alpine grassland stands representing the range of canopy cover (ca. 5–75 %) and basal area (112 - 3889 m²/ha^-1) in the headwaters of the Avoca catchment, Canterbury, New Zealand. In open-canopy stands (< 50 % canopy cover, < 2000 m²/ha^-1 basal area) C. pallens individuals were clearly distinguishable. Rela-tionshipsbetweenbasal diameter, number of tillers, height, and % crown death suggested four putative age-states: ‘seedlings‘, ‘juvenile tussocks’, ‘mature tussocks’ and ‘senescent tussocks’. Four types of population structure were identified depending onthe proportions of seedlings and juvenile and mature tussocks. C. pallens seedlings were present in seven stands at densities ranging from 0.6 to 123.4 / m². They were most frequent on microsites protected from frost-heave by short herbaceous vegetation or litter, but with little competition from taller vegetation. Differences in seedling densities between stands partly reflected the proximity and abundance of seed sources and degree of protection from frost-heave. C. pallens population structures appear to be influenced by the frequency and magnitude of geomorphic disturbances and browsing by introduced animals.