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.     

Long-term change in growth, mortality and regeneration of trees in Denny Wood, an old-growth wood-pasture in the New Forest (UK)

Long-term changes in stand composition and structure were recorded in Denny Wood (New Forest, UK) by means of a permanent transect covering 2 ha. Denny is an ancient, mixed deciduous wood-pasture dominated by beech (Fagus sylvatica), pedunculate oak (Quercus robur) and holly (Ilex aquifolium) whose canopy trees ranged in age from approximately 70 years to over 300 years when the study began in 1956. Individual trees, shrubs and saplings were mapped and measured at irregular intervals until 1996.

During the 40 years of observations, storms and drought disrupted the stand. Considerable volumes of dead wood accumulated, and canopy gaps extended to 30% of the transect area. Small trees and saplings were severely damaged by ponies and grey squirrels. Regeneration ceased after 1964, due principally to heavy grazing and browsing by deer and ponies. Despite the disturbances, most mortality was due to competitive exclusion within well-stocked parts of the stand.

Historical records from the 17th century onwards demonstrate a long-term change from oak dominance with groups of beech before 1800 to beech dominance in the late 20th century. The stand through which the transect now runs was enclosed in 1870, and this allowed beech to regenerate abundantly, but in the nearby unenclosed part of Denny Wood holly regenerated more abundantly than beech.

The patterns of growth, mortality and regeneration are compared with natural temperate deciduous woodland. The long-term relationship between beech and oak is likely to involve periodic oak regeneration after major disturbances, interspersed with steady increases in the proportion of beech. The implications for managing and monitoring the “Ancient and Ornamental Woods” of the New Forest are considered.     

Germination of native and introduced plants from scats of Fallow Deer (Dama dama) and Eastern Grey Kangaroo (Macropus giganteus) in a south‐eastern Australian woodland landscape

Introduced deer occur in many forests and woodlands in Australia and potentially play an important role in influencing the floristics and structure of these landscapes through eating plants and disseminating seeds. In a glasshouse trial, we tested whether field‐collected scats of Fallow Deer (Dama dama) contained viable plant seeds. Scats of deer obtained from a woodland study area in Kosciuszko National Park, New South Wales, produced seedlings from a range of native and introduced plant species. Forbs and herbs were dominant in these samples, confirming the grazing behaviour of deer at the time scats were collected. Samples of scats from Eastern Grey Kangaroos (Macropus giganteus), collected contemporaneously from the same sites as deer scats, also produced plant germinants. By volume, deer scats produced a greater diversity of plant germinants, including native and weed species, than did kangaroo scats. Although no weed species emanating from deer or kangaroo scats were of national significance, several species were of regional environmental significance, including Common Mullein (Verbascum thapsis), which was only found germinating out of deer scat, Stinkgrass (Eragrostis cilianensis) and Purpletop (Verbena bonariensis). In addition to dispersing viable seeds, Fallow Deer may also influence vegetation structure through their browsing. Further research is necessary to elucidate their respective role in dispersing native and introduced plants as well as any impacts that foraging behaviour might be having on woodland landscapes, to better inform management of the resident deer population.     

Exotic deer diminish post-fire resilience of native shrub communities on Santa Catalina Island, southern California

Browsing by exotic mule deer on Santa Catalina Island (SCI) off the coast of southern California may diminish the post-fire resilience of native shrublands. To assess this, deer exclosures were established following a wildfire to monitor post-fire recovery of three dominant, native shrub species (Heteromeles arbutifolia, Rhus integrifolia, and Rhamnus pirifolia). Post-fire resprout growth, mortality, and tissue water status as well as pre- and post-fire shrub density and cover were measured inside and outside of deer exclosures. We found that deer browsing significantly limited post-fire resprout growth and led to increased mortality of resprouting H. arbutifolia shrubs (88 % mortality outside compared to 11 % inside exclosures). Post-fire resprouts maintained favorable water status during the study despite drought conditions, indicating that water stress was not a proximate cause of resprout mortality. Deer browsing resulted in a >93 % reduction in canopy coverage of dominant shrub species. The dramatic reduction of native shrubs at this site may create opportunities for displacement by exotic species, resulting in eventual vegetation-type conversion. The observed link between intense browsing and post-fire shrub mortality provides much needed information concerning the environmental impact of exotic deer on SCI and illustrates the interaction between exotic herbivores and fire on an island system.     

Diversity measurements in shrubland communities of Galicia (NW Spain)

Diversity was studied in 10 communities, including the understory of native oak woodland, planted woodlands (pine and eucalypt), and shrublands in the strict sense (heathlands, broom shrublands, gorse shrublands).In each community, species richness, diversity, dominance and evenness were analysed. Differences were observed among communities with regard to species composition, richness in annual herbs, perennial herbs and shrubs, dominant plant families (Ericaceae, Papilionaceae) and diversification of shrub species.The possible relations between environmental stress and/or human influences on differences in diversity are discussed.     

Recruitment of Quercus agrifolia in central California: the importance of shrub-dominated patches

Abstract. Many perennial plants strongly enhance the survival of seedlings of other species. We studied patterns of long-term recruitment of Quercus agrifolia (Coastal live oak) associated with shrub-dominated communities by counting Q. agrifolia recruits on a time sequence of historical aerial photographs and comparing recruitment among mapped patches of coastal sage scrub, chaparral, and grassland in an 1120-ha landscape. Because we could not identify new recruits in existing woodlands with aerial photographs, we studied the recruitment of Q. agrifolia in this vegetation type indirectly by comparing population size structures and the spatial relationships between shrubs and recruits among woodlands that varied in understory community type. At the landscape scale, recruitment was higher in coastal sage scrub vegetation than predicted by the extent of its coverage, commensurate with the spatial coverage of chaparral, and very low in grassland. Recruitment within woodland communities also varied considerably. In woodland communities on sheltered, north-oriented topography with understories dominated by shrubs, there were large numbers of small Q. agrifolia, and recruits were not significantly spatially associated with shrubs within plots. In woodlands with herbaceous understories there were few individuals in the small size classes, and recruits were strongly spatially associated with shrubs within plots. Woodlands with shrub-dominated understories have population structures that appear to be stable, but woodlands with herbaceous understories exhibit size structures associated with declining populations. Quercus recruitment into shrub-dominated patches corresponds with previous documentation of facilitative relationships between shrubs and oak seedlings, and suggests the occurrence of an unusual form of patch dynamics in these landscapes.     

Eurasian badger habitat selection in Mediterranean environments: Does scale really matter?

It is widely believed that spatial scale affects habitat selection, and should influence management options, especially for species with wide geographic distribution or large territories. Eurasian badger habitat selection has been well studied throughout most of its European distribution range, but never at multiple spatial scales. We used compositional analysis to assess habitat selection of Eurasian badgers in southern Portugal at four spatial scales (1, 4, 25, and 100 km²). We assessed habitat use from setts, latrines and footprints presence, and road kills. Oak woodlands with understorey were selected at all scales, being the most preferred habitat at 3 scales (1, 4, and 100 km²). Pastures were most selected at the scale of the 25 km² cell, but their use was not significantly different from oak woodland with understorey. Shrubs and pastures were also secondly important at the majority of scales. Contrary to findings at northern latitudes, deciduous forests decreased in importance as cell size increased. In the highly humanized and fragmented landscape of southern Portugal, Eurasian badgers are selecting the matrix of oak woodlands interspersed with patches of pastures, shrubs and riparian vegetation. In these oak woodlands, scale does not have a marked effect. Management for badgers should provide, for at least, 30% of oak woodland cover at all scales. Our study illustrates the across-scale importance of maintaining the historically human altered, sustainable and unique landscape and land use system – the montado.     

Habitat change and woodland birds in Britain: implications for management and future research

There is increasing evidence that changes in habitat structure in the form of reduced understorey and loss of open habitats, both probably a result of increases in shading and deer browsing, may be responsible for causing recent changes in the composition of breeding bird communities in many lowland British woods and forests. In contrast, management of upland coniferous forests may prevent the attainment of mature and old-growth structures which would benefit community development in these new ecosystems. We suggest that the key challenge for woodland conservation policy is to create larger areas of both young-growth and old-growth habitat. These objectives need not necessarily conflict with each other, or with other multipurpose forestry objectives, provided that they inform strategic plans and are targeted at appropriate locations and scales. In the lowlands, the current large stock of middle-aged, often unmanaged and species-poor woodland provides an opportunity to restore or create new woodland habitats of high biodiversity value. The development of woodfuel markets may effectively increase the amount of young-growth but it is unclear exactly what habitat structures might be created. In the uplands, allowing more natural development of native woodland in mosaics with other habitats may provide opportunities for both old- and young-growth species. For the foreseeable future, deer impacts will continue to be widespread and probably increasingly severe in many areas. In view of the uncertainty about the implications of climate change for woodland ecosystems, a strong case can be made for attempting to buffer valued wildlife communities against damaging effects by creating and maintaining high structural diversity at a range of scales and increasing the area of woodland.     

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.     

Woodland biodiversity, palaeo-human ecology and some implications for conservation management

This paper appraises the role of prehistoric human societies in modifying the natural woodland biodiversity of several regions throughout Scotland. It draws attention to the likelihood that, contrary to popular belief, biodiversity was enhanced by interactions between agricultural communities and woodlands. The possible purposefulness of such interactions is discussed. Mindful of the present concern to re-establish ‘native’ woodlands in many parts of northern Britain, we then review three approaches to the identification of native tree types in Scotland, and conclude that insufficient attention in reafforestation schemes is being paid to (a) the palaeoecological record as a record of native woodland or (b) the likely former high taxonomic diversity of woodlands. Both these failings will lead to the creation of new woodlands lacking in species diversity.     

Spontaneous development of urban woody vegetation on differing soils

The general aim of the study was to follow the development of spontaneous woody vegetation on man-made urban habitats in Berlin, Germany. In spring 1968, the vegetation was removed completely in five groups of plots, which had been filled with five different transferred soil material in 1928. The spontaneous vegetation was monitored annually from 1968 to 2006. Especially, the development of shrubs and trees was followed. In addition the growth of the most important herbaceous species, Solidago canadensis, was investigated.

The soils varied from moderately rich sandy soils to rich sandy or loamy soils. On the two sandy soils S. canadensis developed poorly. Tree species dominated (>50% of total cover) on these plots after 14–18 years. On the soils rich in nutrients, S. canadensis developed strongly. Tree species did not dominate there before 21–25 years. The diversity of woody species increased over time and reached 33 species altogether. Indigenous Acer, Quercus and Betula species prevailed. Alien species, typical for urban ruderal sites, were rare.

The results support the idea of establishing urban woodlands by spontaneous succession as a cheap way to develop near-natural plant communities rich in species.     

Landscape effects on birds in urban woodlands: an analysis of 34 Swedish cities

Aim To compare bird abundances in woodlands along gradients from the city centre to the peri‐urban area. To evaluate the importance of the proportion of woodland within the city and in the peri‐urban landscape to forest bird communities breeding in urban woodlands. To test whether fragmentation effects on birds were linked to the type of peri‐urban matrix. Location A total of 34 Swedish cities with > 10,000 inhabitants in south‐central Sweden. The study area covered 105,000 km², in which 84% of the Swedish population of 9.1 million lives. Methods Repeated point count surveys were conducted in 2004 in a total of 474 woodlands. General linear models were used to test for possible differences in abundance along urban to peri‐urban gradients, and to regress bird abundances in local urban woodlands on: (1) total woodland cover in the city, (2) total woodland cover in the peri‐urban landscape, (3) the interaction between woodland cover in the city and in the peri‐urban area, (4) region, and (5) human density. Results More than 12,000 individuals of 100 forest bird species were recorded. Of the 34 most common species detected, 13 bird species had higher abundances in urban than in peri‐urban woodlands, and seven species showed the opposite trend. The bird community of urban woodlands was characterized by species associated with deciduous forests and tree nesters, whereas the bird community of peri‐urban woodlands was characterized by species associated with coniferous woodland and ground nesters. Twelve species were significantly linearly associated with the proportion of urban woodland and/or the proportion of peri‐urban woodland, and a further eight species were associated with the interaction between these two factors. Local breeding bird abundances of four species were significantly positively associated with the proportion of urban woodland only in farmland‐dominated landscapes. Main conclusions Fragmentation effects on some urban birds are linked to the type of peri‐urban matrix. In farmland landscapes, peri‐urban woodlands may have been too scarce to act as a source of bird immigrants to fragmented urban woodlands. To maintain populations of specialized forest birds within cities in landscapes dominated by agriculture, it is of paramount importance to conserve any remaining urban woodlands.     

Woodland recovery after suppression of deer: cascade effects for small mammals, wood mice (Apodemus sylvaticus) and bank voles (Myodes glareolus)

Over the past century, increases in both density and distribution of deer species in the Northern Hemisphere have resulted in major changes in ground flora and undergrowth vegetation of woodland habitats, and consequentially the animal communities that inhabit them. In this study, we tested whether recovery in the vegetative habitat of a woodland due to effective deer management (from a peak of 0.4-1.5 to <0.17 deer per ha) had translated to the small mammal community as an example of a higher order cascade effect. We compared deer-free exclosures with neighboring open woodland using capture-mark-recapture (CMR) methods to see if the significant difference in bank vole (Myodes glareolus) and wood mouse (Apodemus sylvaticus) numbers between these environments from 2001-2003 persisted in 2010. Using the multi-state Robust Design method in program MARK we found survival and abundance of both voles and mice to be equivalent between the open woodland and the experimental exclosures with no differences in various metrics of population structure (age structure, sex composition, reproductive activity) and individual fitness (weight), although the vole population showed variation both locally and temporally. This suggests that the vegetative habitat--having passed some threshold of complexity due to lowered deer density--has allowed recovery of the small mammal community, although patch dynamics associated with vegetation complexity still remain. We conclude that the response of small mammal communities to environmental disturbance such as intense browsing pressure can be rapidly reversed once the disturbing agent has been removed and the vegetative habitat is allowed to increase in density and complexity, although we encourage caution, as a source/sink dynamic may emerge between old growth patches and the recently disturbed habitat under harsh conditions.     

Large herbivores affect the stoichiometry of nutrients in a regenerating woodland ecosystem

Previous studies on the effects of herbivores on nutrient cycling have given little consideration to the relationship between soil nitrogen (N) and phosphorus (P) availability. Here we examined how browsing by red deer influences the relative availability of N and P in a regenerating woodland ecosystem. We found that removal of browsing by fencing for 14 years led to a shift from N toward P limitation of the dominant tree species Betula pubescens . This was evidenced by a significant increase in foliar N:P ratio of B. pubescens as a result of removal of browsing; mean N:P ratio of foliage from browsed areas was 13.2 suggesting that trees growing in browsed areas were N limited, whereas foliage from unbrowsed areas had a mean N:P ratio of 15.8, suggesting that these areas were more P limited. Further evidence of a shift toward P limitation in unbrowsed areas came from the finding that root uptake of labelled ³²P was significantly greater in roots collected from unbrowsed than browsed trees. Soil phosphatase activity did not significantly differ between browsed and unbrowsed areas. Our data indicate therefore that herbivores have the potential to significantly affect the stoichiometry of N and P in forest ecosystems.     

Seasonal change in browsing by sika deer on hinoki cypress trees on Mount Takahara,central Japan

Browsing by sika deer (Cervus nippon Temminck) on hinoki cypress (Chamaecyparis obtusa (Sieb. et Zucc.) Endl.) was monitored for approximately 3years on 300 trees at three sites on Mount Takahara, central Japan. We investigated the seasonal change in browsing on hinoki cypress in relation to the availability of other food plants (dominated by Sasa nipponica Makino et Shibata) and preferred food plants (deciduous broad leaves). Browsing occurred mostly from November to April, while no browsing damage occurred from June to October. Browsing was observed in November, even when the amount of other food plants did not decrease. Therefore, the cause of browsing cannot be fully explained by a decreased availability of other food plants. Comparison of fecal composition and food availability showed that deer had a preference for deciduous broad leaves. The availability of these preferred plants decreased in November and increased in May. Chemical analysis of the hinoki cypress and deciduous broad leaves (crude protein and neutral detergent fiber) showed that the quality of deciduous broad leaves decreased in November, and as a result the quality of hinoki became higher than that of the deciduous broad leaves. Browsing on hinoki cypress occurred when the availability of deciduous broad leaves decreased, and vice versa. These results suggest that the availability of preferred food plants influences browsing on hinoki cypress by sika deer.     

Relations between the floristic composition of isolated young woods and their proximity to ancient woodland

Abstract. Recent studies indicate that, in the present-day agricultural landscape, the floristic composition of young woodland communities can be fully developed if the woods are situated adjacent to ancient woodlands. Four 70-yr-old deciduous woods in the Carpathian foothills were examined in relation to three adjacent ancient oak-hornbeam and oak-pine woodlands, which are the nearest source of woodland species diaspores. On the basis of data from 208 plots, the frequencies of various species groups in the field layer of the woods were analysed. The dependence of vegetation differentiation within the recent woods on (a) distance to the border with the ancient woodlands and (b) light intensity was examined by Partial Detrended Canonical Correspondence Analysis (DCCA). A significant relation between distance to ancient woodland and species composition was found for recent woods on rich brown soils. The vegetatively propagating species, myrmecochores and small autochores attained higher cover values near ancient woodland; endozoochores and anemochores were most abundant further away. Within recent, more open woods on poor podzolic and leached brown soils, colonisation is strongly inhibited by dense growth of Carex brizoides; here, vegetation regeneration is much slower than in woods on rich soils much further away from the source of diaspores.     

Effects of burning and rainfall on former agricultural land with remnant grassy woodland flora

Grassy woodlands have been extensively cleared for agricultural land uses; land managers need to know whether restoration of biodiversity on such sites requires further interventions beyond simply stopping agricultural land use. Cumberland Plain Woodland occurs on shale‐derived soils in western Sydney; former Cumberland Plain Woodland sites can range from grasslands cleared for agricultural use to regenerated woodlands. An experiment was established in Scheyville National Park to determine what effect repeated burning would have in this system. Four blocks were established (three in grassy areas, one in woodland) and plots in each block were either burnt in 2001 and 2005 or left unburnt. Native plant species richness was initially lower in the grassy blocks than in the woodland, and this ranking remained on unburnt plots over time. The first fire increased species richness of both natives and exotics on the grassy blocks, with the largest increases observed for native and exotic forbs, and lesser increases for grasses (native only), gramminoids and shrubs. Native species richness changed very little with burning in the woodland. Fire effects on species richness were still apparent 3 years later on the grassy blocks; the difference between the grassy blocks and the woodland was not significant on burnt plots at this stage. Changes in native species richness were far less after the second fire on the grassy blocks, with grasses and gramminoids showing increases; native species richness remained higher in the burnt treatment. The first fire reduced the initial differences in native species richness between the grassy blocks and the woodland, and the second fire maintained the benefit through time. Fire also increased exotic species richness; the proportion of total species as natives was not altered by the two fires. On unburnt grassy plots, native species richness and prior cumulative rainfall were positively related; a decline in native species richness on unburnt plots corresponded to increasingly drier conditions over the study.     

Top-down and bottom-up consequences of unchecked ungulate browsing on plant and animal diversity in temperate forests: lessons from a deer introduction

Debate on the relative importance of competition for resources and trophic interactions in shaping the biological diversity of living communities remains unsettled after almost a century. Recently, dramatic increases in ungulate populations have provided a useful quasi-experiment on the effects of unrestrained ungulates on forest ecology. The islands of Haida Gwaii (Canada) offer a unique situation to investigate the potential of large herbivores to control temperate forest community structure and diversity. Black-tailed deer Odocoileus hemionus Merriam, native to adjacent mainland areas of British Columbia, were introduced in 1878 and spread to all but a few islands. Because deer were not native to the archipelago, islands that still lack deer provide a rare instance of temperate forest vegetation and fauna that developed in the absence of large herbivores. The colonisation of different islands at different times, and the absence of significant predation allow us to assess whether and how a large herbivore can exert “top-down” control on vegetation and its associated fauna. We studied plant communities in forest interior and shoreline, on seven small islands of varying browse history. Three islands were untouched by deer, deer had been resident for about 15 years on two, and on another two deer had been present for more than 50 years. Without deer, vegetation in the understorey and/or shrub layer was dense or very dense. Structure and composition varied markedly within and between shoreline and interior communities. Without deer, shoreline communities were dominated by species absent from islands with deer. Where deer had been present for less than 20 years most plant species characteristic of shorelines on islands without deer were already absent or scarce, but in the forest interior species richness was less affected and extensive shrub thickets remained. On islands where deer had been present for >50 years vegetation below the browse line was extremely simplified, converging in both forest interior and shoreline towards an open assemblage of a few deer-tolerant species, basically two coniferous trees. This top down effect on the plant community reflected up the food chain so that understorey invertebrate and shrub-dependent songbird communities became simplified. In contrast, species densities of litter arthropods (especially weevils and millipedes) were highest where deer were present for >50 years. Canopy birds were unaffected by deer presence. In the absence of predators, major climatic stress or other means to control the herbivore, deer browsing created greatly simplified plant and animal communities.     

Indirect effects of deer herbivory on local nitrogen availability in a coastal dune ecosystem

Herbivores can have indirect effects on local nutrient availability if their direct effects on plants lead to changes in the amount or chemical composition of litter reaching the soil surface. Using two exclosure experiments, we evaluated this possibility for black-tailed deer ( Odocoileus hemionus columbianus ) feeding on silver bush lupine ( Lupinus chamissonis ) in a coastal dune system in northern California. Our first experiment assessed the effects of deer herbivory on 360 lupines that were tracked from newly germinated seedlings in 1996 and 1997 until 2000, when most surviving individuals had reached reproductive age. Results from this experiment showed that browsing by deer significantly reduced growth rates and seed production of lupines and delayed the onset of reproductive maturity. Although deer had no effect on the density of litter accumulating underneath shrubs (g m⁻²), browsing significantly decreased C:N ratios of leaf tissue, due primarily to increased nitrogen content of leaves. Deer browsing also caused significantly increased net nitrogen mineralization rates in the soil under shrubs, although pools of ammonium and nitrate were uninfluenced. The second experiment examined 72 established bush lupines from 1997 to 2000, and showed that deer browsing significantly decreased seed production but had no effect on shrub growth. We also detected trends for browsed shrubs to have reduced pools of ammonium and nitrate underneath their canopies, although no such patterns emerged for nitrogen mineralization rates. Collectively, these data suggest that changes in litter quality, more so than litter quantity, contributed to the indirect effects of deer herbivory on local nitrogen dynamics. We hypothesize that deer browsing induced chemical changes in lupine plant tissue, which increased the nitrogen content of litter reaching the soil surface and subsequently increased rates of nitrogen mineralization.