Herbivores Influence the Growth,Reproduction, and Morphology of a Widespread Arctic Willow

Shrubs have expanded in Arctic ecosystems over the past century, resulting in significant changes to albedo, ecosystem function, and plant community composition. Willow and rock ptarmigan (Lagopus lagopus, L. muta) and moose (Alces alces) extensively browse Arctic shrubs, and may influence their architecture, growth, and reproduction. Furthermore, these herbivores may alter forage plants in such a way as to increase the quantity and accessibility of their own food source. We estimated the effect of winter browsing by ptarmigan and moose on an abundant, early-successional willow (Salix alaxensis) in northern Alaska by comparing browsed to unbrowsed branches. Ptarmigan browsed 82–89% of willows and removed 30–39% of buds, depending on study area and year. Moose browsed 17–44% of willows and browsed 39–55% of shoots. Browsing inhibited apical dominance and activated axillary and adventitious buds to produce new vegetative shoots. Ptarmigan- and moose-browsed willow branches produced twice the volume of shoot growth but significantly fewer catkins the following summer compared with unbrowsed willow branches. Shoots on browsed willows were larger and produced 40–60% more buds compared to unbrowsed shoots. This process of shoot production at basal parts of the branch is the mechanism by which willows develop a highly complex “broomed” architecture after several years of browsing. Broomed willows were shorter and more likely to be re-browsed by ptarmigan, but not moose. Ptarmigan likely benefit from the greater quantity and accessibility of buds on previously browsed willows and may increase the carrying capacity of their own habitat. Despite the observed tolerance of willows to browsing, their vertical growth and reproduction were strongly inhibited by moose and ptarmigan. Browsing by these herbivores therefore needs to be considered in future models of shrub expansion in the Arctic.     

Environmental variation and moose Alces alces density as determinants of spatio-temporal heterogeneity in browsing

Understanding temporal variation in habitat selection and browsing intensity by large herbivores is fundamental because of their large impact on the ecosystems. I studied the annual variation in winter browsing pressure on young trees and habitat selection by moose Alces alces over a ten year period. Specifically, the relationships between browsing pressure on Scots pine Pinus sylvestris and two birch species ( Betula ssp.) and three explanatory variables – 1) availability of forage, 2) moose density (estimated by pellet group counts) and 3) snow cover was studied. At a larger spatial scale (forest stand level) the relationship between moose habitat selection between three different habitat types (forest <30 yr, forest>30 yr and mire) and two explanatory variables, 1) snow condition and 2) moose density, were studied. Browsing pressure on Scots pine, the dominating food plant, was related to forage availability, moose density and snow condition. No significant relationships between any of the three explanatory variables and browsing pressure on the two birch species were found. Moose selection for certain habitats varied between years and was affected by number of days with >0.10 m of snow.
Habitat selection was not significantly related to moose density and the relationship between overall moose density and habitat specific moose densities was proportional within the studied density range. These findings have implications for understanding varying browsing patterns – and will affect both the ability to predict herbivores' effect on the forest ecosystem. A snow dependent browsing pattern also indicates that one can expect a long term decrease in browsing pressure on the tree and shrub layer as a consequence of the ongoing large-scale climate change.     

Reindeer herbivory reduces willow growth and grouse forage in a forest-tundra ecotone

Browsing mammals strongly modify the structure of vegetation of forest-tundra ecotones. We investigated the impact of reindeer browsing on growth and morphology of an arctic willow, Salix glauca, by studying plants inside and outside fenced areas in a tundra habitat at the tree line. We also studied if reindeer feeding has an effect on the forage availability for willow grouse, a herbivore sharing the same food plant. Analyses of 6-year data show that reindeer strongly reduces the growth of tundra willow and changes plant morphology to a stunted growth form. Intense reindeer browsing on willow limited the forage availability for willow grouse and grouse fed less on reindeer-browsed willow than willows protected from reindeer browsing. The results of this study imply that herbivores can counteract the increase of shrubs in forest-tundra ecotones, as has been predicted in some studies discussing the effects of climatic warming on vegetation. Furthermore, trophic interactions should be incorporated in modelling vegetation changes as a response to increased temperatures.     

Browsing‐mediated shrub canopy changes drive composition and species richness in forest‐tundra ecosystems

Changes in climate and in browsing pressure are expected to alter the abundance of tundra shrubs thereby influencing the composition and species richness of plant communities. We investigated the associations between browsing, tundra shrub canopies and their understory vegetation by utilizing a long‐term (10–13 seasons) experiment controlling reindeer and ptarmigan herbivory in the subarctic forest tundra ecotone in northwestern Fennoscandia. In this area, there has also been a consistent increase in the yearly thermal sum and precipitation during the study period. The cover of shrubs increased 2.8–7.8 fold in exclosures and these contrasted with browsed control areas creating a sharp gradient of canopy cover of tundra shrubs across a variety of vegetation types. Browsing exclusions caused significant shifts in more productive vegetation types, whereas little or no shift occurred in low‐productive tundra communities. The increased deciduous shrub cover was associated with significant losses of understory plant species and shifts in functional composition, the latter being clearest in the most productive plant community types. The total cover of understory vegetation decreased along with increasing shrub cover, while the cover of litter showed the opposite response. The cover of cryptogams decreased along with increasing shrub cover, while the cover of forbs was favoured by a shrub cover. Increasing shrub cover decreased species richness of understory vegetation, which was mainly due to the decrease in the cryptogam species. The effects were consistent across different types of forest tundra vegetation indicating that shrub increase may have broad impacts on arctic vegetation diversity. Deciduous shrub cover is strongly regulated by reindeer browsing pressure and altered browsing pressure may result in a profound shrub expansion over the next one or two decades. Results suggest that the impact of an increase in shrubs on tundra plant richness is strong and browsing pressure effectively counteracts the effects of climate warming‐driven shrub expansion and hence maintains species richness.     

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.     

Slashing Phragmites (Phragmites australis) prior to planting does not promote native vegetation establishment

Phragmites or Common Reed (Phragmites australis) is a tall rhizomatous cosmopolitan grass. While native to Australia, it can be invasive in wetlands, forming dense monocultures and reducing their ecological integrity. We assessed the potential for the cutting of Phragmites reeds prior to planting to promote the establishment of indigenous shrubs that might ultimately outcompete Phragmites. We established ten 5 m × 5 m quadrats in an area dominated by Phragmites, brush‐cut the reeds to ground level in five of them and left five uncut as controls. Within each quadrat, we planted 20 plants (~40 cm tall) of each of five indigenous shrub species, unguarded (4 plants/m²). We surveyed the plants one month after planting and annually for the following four years for survival, height and browsing damage. Browsing damage to plants was common (>50%) and unaffected by cutting. After four years, overall plant survival rates were ~25% and mean plant heights for the five shrub species ranged between 120 and 174 cm. Cutting of Phragmites had no positive effect on plant survival or height. In fact, two Melaleuca species grew taller in the uncut quadrats. Cutting of Phragmites reed beds prior to planting is unlikely to promote the establishment of woody plantings. However, planting within established Phragmites with or without prior brush‐cutting is worthy of further trialling as a potential tool for reinstating native diversity at Phragmites‐dominated sites.     

Demography of the invasive shrub Scotch broom (Cytisus scoparius) at Barrington Tops,New South Wales: insights for management.

The exotic shrub Scotch Broom (Cytisus scoparius) has invaded large areas of eucalypt woodland at Barrington Tops, New South Wales, where it forms dense stands that have significant impacts on vegetation structure, flora and fauna. Data are presented from four 25 m² plots, which have been studied since 1985. Two plots were located in uniform broom thickets of different ages, and two were located across the margins of broom stands, which have since expanded to cover the entire plots. All broom plants in the plots (other than young seedlings, which were counted) were mapped, tagged and monitored annually. New seedlings appeared annually, but there was no relationship between their numbers (varying between years) and subsequent recruitment of older plants. The probability of seedlings reaching first flowering was less than 2%, and of surviving to mature size (>10 cm² basal area) was negligible. Seedlings mainly died through suppression (shade). Individuals less than 50 cm high were also browsed. Recruitment occurred only where light levels were high, either before closure of the broom canopy or after senescence had led to canopy opening. From approximately 12–30 years after initial invasion, broom stands underwent self‐thinning of mature plants, accelerated by collapse of plants on to each other. Recruitment of new, maturing plants, after this period produced a stand that was less dense than that found after initial invasion. Broom is creating more disturbance‐prone environments due to its impacts on other biota, likely alterations to the fire regime, and by harbouring feral pigs. Further disturbance favours broom, and elsewhere it has resulted in massive seedling regeneration. While fire or other disturbance can be used to stimulate germination, and thereby reduce a large part of the soil seed bank, denser broom infestations are likely to result unless follow‐up treatments can be applied over long time periods. A wiser management option, at least in the short term, may be avoidance of all disturbance, especially for stands of mature broom.     

Test of Localized Management for Reducing Deer Browsing in Forest Regeneration Areas

ABSTRACT White-tailed deer (Odocoileus virginianus) browsing in forest regeneration sites can affect current and future stand structure and species composition. Removal of deer social units (localized management) has been proposed as a strategy to alleviate deer overbrowsing in forest systems. We conducted an experimental localized removal in a high-density deer population in the central Appalachians of West Virginia, USA, during winter 2002. We removed 51 deer within a 1.1-km² area that encompassed 2 forest regeneration sites (14 ha). During the summer following removal, we detected decreases in distance from the removal area in 8 of 30 (26.7%) adult females having pretreatment mean telemetry locations <2.5 km from the center of the removal area. We measured browsing rates during the summers of 2001–2004 from forest regeneration sites to examine efficacy of localized management. Browsing rates declined annually in both removal and control areas, due in part to increased timber harvesting on the larger study site, suggesting that increasing forage availability may be more effective at reducing impacts on forest regeneration than localized reductions in deer populations. Three years after the initial removal, we removed an additional 31 deer from the original 1.1-km² removal area. Home range shifts of adjacent deer coupled with the large number of animals collected in the second removal suggests that localized management only produces temporary voids within high-density deer herds. Localized management may not effectively reduce negative impacts of deer in areas of high deer density.     

Growth and architecture of small honey mesquites under jackrabbit browsing: overcoming the disadvantage of being eaten

Browsing is an important mortality factor in seedlings and small plants. However, the induced changes in the architecture of plant survivors may influence subsequent browsing, opening the possibility of compensating for the damage done. How jackrabbit (Lepus californicus) browsing affects the growth and architecture of small individuals of honey mesquite, Prosopis glandulosa var. torreyana, a tree/shrub that produces spines at every node, was explored. Naturally established mesquites of unknown age were selected in one site, and 2-year-old mesquites were transplanted in another site. In both cases, half of them were exposed to jackrabbits and the other half were excluded as controls. After 4 years, shoot production (height, length and number of derived shoots) and plant growth (height and cover) increased 1.4-2.5-fold in naturally established controls relative to exposed plants, depending on the measured variable. In the transplant experiment, the increases were 2.8-7.1-fold in controls relative to exposed plants 2 years after initiation of the experiment. The net loss of biomass in treatment vs. control plants in these experiments suggests a negative response to browsing which has been defined as under-compensation. Alternative architectures in honey mesquites were evident at the end of the exclusion experiments: controls had long branches and an extended crown cover, while exposed plants had short branches and a compact crown cover. Results indicated that mesquites were able to grow under browser pressure by packing many stems in a compact matrix armed with spines and producing one or more shoots tall and wide enough to escape from jackrabbits.     

Tolerance of an Expanding Subarctic Shrub,Betula glandulosa,to Simulated Caribou Browsing

Densification of the shrub layer has been reported in many subarctic regions, raising questions about the implication for large herbivores and their resources. Shrubs can tolerate browsing and their level of tolerance could be affected by browsing and soils productivity, eventually modifying resource availability for the caribou. Our objective was to assess the compensatory growth potential of a subarctic shrub, Betula glandulosa Michx., in relation with caribou browsing and nutriment availability for the plants. We used a simulated browsing (0, 25 and 75% of available shoots) and nitrogen-fertilisation (0 and 10 g m⁻²) experiment to test two main hypotheses linking tolerance to resource availability, the Compensatory Continuum Hypothesis and the Growth Rate Hypothesis as well as the predictions from the Limiting Resource Model. We seek to explicitly integrate the relative browsing pressure in our predictions since the amount of tissues removed could affect the capacity of long-lived plants to compensate. Birches fully compensated for moderate browsing with an overall leaf biomass similar to unbrowsed birches but undercompensated under heavy browsing pressure. The main mechanism explaining compensation appears to be the conversion of short shoots into long shoots. The leaf area increased under heavy browsing pressure but only led to undercompensation. Fertilisation for two consecutive years did not influence the response of birch, thus we conclude that our results support the LRM hypothesis of equal tolerance under both high and low nitrogen availability. Our results highlight that the potential for compensatory growth in dwarf birch is surpassed under heavy browsing pressure independently of the fertilisation regime. In the context of the worldwide decline in caribou herds, the reduction in browsing pressure could act synergistically with global climate change to promote the current shrub expansion reported in subarctic regions.     

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.     

Long‐term response of temperate canopy trees to removal of browsing from an invasive arboreal herbivore in New Zealand

Defoliation of forest tree canopies by herbivores and other agents, leading to tree mortality and reduced productivity, threatens the ecological stability of forests globally. This study shows that long‐term control of a mammalian arboreal folivore (brushtail possums; Trichosurus vulpecula Phalangeridae) reduces crown dieback and increases foliage cover in browsing‐damaged canopy trees. We monitored indices of possum density, possum browsing, tree foliage cover and crown dieback for 20 years following initiation of possum control in 1994 that repeatedly reduced possum densities to near zero every 5–6 years and kept the population below 35% of pre‐control levels over the entire period. Observable possum browsing was recorded on 20–49% of individuals of three palatable tree species at the time of first control. Those percentages fell to zero after control and never exceeded 2–10% for individual species over the next 19 years. We recorded significant increases in foliage cover attributable to recovery from defoliation by possums for all three species during the first 10 years. Large increases in foliage cover occurred on individuals that were heavily browsed in 1994 (mean increases: 36–89%), but mean population increases were modest (3–19%) because only 10–19% of trees were initially heavily browsed. Twenty‐year mortality rates were similar for plants with, or without, initial possum browsing, indicating no residual impact of pre‐control browsing on tree mortality. Times for full recovery of crown foliage cover varied from 10 years for the youngest trees and faster growing species to more than 20 years for mature individuals of the slowest growing species.     

Herbivory of clonal populations: cattle browsing affects reproduction and population structure of Yucca elata

The hypothesis that cattle browsing on inflorescences of the soaptree Yucca elata reduces reproductive success was investigated by comparing recruitment and population structure in six populations protected from grazing, six grazed during the flowering season and five grazed outside the flowering season. Cattle consumed 98% of inflorescences, which were found to be highly nutritious. Reduced recruitment in flower-grazed populations could not be attributed to reduced flower survival, as recruitment in non-flower grazed populations was also reduced. Changes in population structure were due to cattle browsing small caudices, including both genets and ramets. An alternate hypothesis of limited germination in soils compacted by cattle was not supported. Cattle browsing of inflorescences reduced reproductive effort, which may be due to the inability of the plants to resorb nutrients after flowering. Browsing also increased branching, probably through lack of apical dominance, whereas physical trampling increased procumbency. Cattle browsing implies a lack of genetic recruitment, possible local extinction of the yucca moth Tegeticula yuccasella, the exclusive pollinator of Y. elata, and local reduction in insect and bird biodiversity.     

The ontogeny of handling hard‐to‐process food in wild brown capuchins (Cebus apella apella): evidence from foraging on the fruit of Maximiliana maripa

We examined age‐related differences in wild brown capuchins' foraging efficiency and the food‐processing behaviors directed toward maripa palm fruit (Maximiliana maripa). A detailed comparison of the different foraging techniques showed that plucking the fruit from the infructescence constituted the main difficulty of this task. Foraging efficiency tended to increase with age, with a threshold at which sufficient strength allowed immatures by the age of three to reach adult‐level efficiency. Youngsters spent more time than older individuals browsing the infructescence and pulling the fruit in an attempt to harvest it. Infants tried to compensate for their inability to pluck fruit by adopting alternative strategies but with low payback, such as gnawing unplucked fruit and opportunistically scrounging others' partially processed food. Although around 2 years of age, young capuchins exhibited all of the behaviors used by adults, they did not reach adult‐level proficiency at feeding on maripa until about 3 years (older juveniles). We compared this developmental pattern with that of extractive foraging on beetle larvae (Myelobia sp.) hidden in bamboo stalks, a more difficult food for these monkeys [Gunst N, Boinski S, Fragaszy DM. Behaviour 145:195–229, 2008]. For maripa, the challenge was mainly physical (plucking the fruit) once a tree was encountered, whereas for larvae, the challenge was primarily perceptual (locating the hidden larvae). For both foods, capuchins practice for years before achieving adult‐level foraging competence, and the timeline is extended for larvae foraging (until 6 years) compared with maripa (3 years). The differing combinations of opportunities and challenges for learning to forage on these different foods illustrate how young generalist foragers (i.e. exploiting a large number of animal and plant species) may compensate for their low efficiency in extractive foraging tasks by showing earlier competence in processing less difficult but nutritious foods, such as maripa fruit. Am. J. Primatol. 72:960–973, 2010. © 2010 Wiley‐Liss, Inc.     

Near Infrared Spectroscopy Facilitates Rapid Identification of Both Young and Mature Amazonian Tree Species

Precise identification of plant species requires a high level of knowledge by taxonomists and presence of reproductive material. This represents a major limitation for those working with seedlings and juveniles, which differ morphologically from adults and do not bear reproductive structures. Near-infrared spectroscopy (FT-NIR) has previously been shown to be effective in species discrimination of adult plants, so if young and adults have a similar spectral signature, discriminant functions based on FT-NIR spectra of adults can be used to identify leaves from young plants. We tested this with a sample of 419 plants in 13 Amazonian species from the genera Protium and Crepidospermum (Burseraceae). We obtained 12 spectral readings per plant, from adaxial and abaxial surfaces of dried leaves, and compared the rate of correct predictions of species with discriminant functions for different combinations of readings. We showed that the best models for predicting species in early developmental stages are those containing spectral data from both young and adult plants (98% correct predictions of external samples), but even using only adult spectra it is still possible to attain good levels of identification of young. We obtained an average of 75% correct identifications of young plants by discriminant equations based only on adults, when the most informative wavelengths were selected. Most species were accurately predicted (75–100% correct identifications), and only three had poor predictions (27–60%). These results were obtained despite the fact that spectra of young individuals were distinct from those of adults when species were analyzed individually. We concluded that FT-NIR has a high potential in the identification of species even at different ontogenetic stages, and that young plants can be identified based on spectra of adults with reasonable confidence.     

Ungulate Browsing Maintains Shrub Diversity in the Absence of Episodic Disturbance in Seasonally-Arid Conifer Forest

Ungulates exert a strong influence on the composition and diversity of vegetation communities. However, little is known about how ungulate browsing pressure interacts with episodic disturbances such as fire and stand thinning. We assessed shrub responses to variable browsing pressure by cattle and elk in fuels treated (mechanical removal of fuels followed by prescribed burning) and non-fuels treated forest sites in northeastern Oregon, US. Seven treatment paddocks were established at each site; three with cattle exclusion and low, moderate and high elk browsing pressure, three with elk exclusion and low, moderate and high cattle browsing pressure, and one with both cattle and elk exclusion. The height, cover and number of stems of each shrub species were recorded at multiple plots within each paddock at the time of establishment and six years later. Changes in shrub species composition over the six year period were explored using multivariate analyses. Generalized Linear Mixed Models were used to determine the effect of browsing pressure on the change in shrub diversity and evenness. Vegetation composition in un-browsed paddocks changed more strongly and in different trajectories than in browsed paddocks at sites that were not fuels treated. In fuels treated sites, changes in composition were minimal for un-browsed paddocks. Shrub diversity and evenness decreased strongly in un-browsed paddocks relative to paddocks with low, moderate and high browsing pressure at non-fuels treated sites, but not at fuels treated sites. These results suggest that in the combined absence of fire, mechanical thinning and ungulate browsing, shrub diversity is reduced due to increased dominance by certain shrub species which are otherwise suppressed by ungulates and/or fuels removal. Accordingly, ungulate browsing, even at low intensities, can be used to suppress dominant shrub species and maintain diversity in the absence of episodic disturbance events.     

Patterns and trends in coral reef macroalgae browsing: a review of browsing herbivorous fishes of the Indo-Pacific

Browsing fishes have been identified as an important component of coral reef resilience, because in contrast to other herbivorous fishes they are able to feed on established macroalgae. Climate change and local anthropogenic impacts have contributed to phase shifts in many coral reefs from coral to macroalgae dominance, and recent research suggests the potential ability of browsers to reverse such phase shifts. However, there is high variation among studies and some contradicting findings exist. Here, we review the relevant literature to assemble a list of species currently known to contribute to browsing in the Indo-Pacific. Furthermore, we identify spatial and temporal patterns, outline factors influencing browsing, and discuss the probability of phase shift reversal. We formulate research recommendations addressing the identified gaps in knowledge about the interactions of browsing fishes and their environment. To date, 37 species of fishes have been observed consuming macroalgae in the Indo-Pacific. The most important groups are the family Siganidae, the subfamily Scarinae (Labridae), and the subfamily Nasinae (Acanthuridae). Browsing species vary between studies depending on location, season and macroalgae species examined. Several influencing factors, such as structural complexity, palatability of macroalgae and ecosystem connectivity have been suggested to cause these discrepancies. The most promising avenues for future research are the effect of structural complexity, the importance of mobile link species and influences of food availability on the selectivity of browsing species. Increasing our knowledge in these fields will provide a better basis for successful management strategies directed at increasing the resilience of coral reefs.     

Ordination of vegetation change in Guadalupe Mountains,New Mexico,USA

Vegetation change over a nine-year period was studied in Guadalupe Mountains, New Mexico. Permanent transects in desert shrub vegetation were sampled in 1972 and 1980. Emphasis was given to shrubs because of their importance to big game diets. Univariate paired t-tests and reciprocal averaging ordination were used to detect and display coordinated changes in species composition over time. Despite apparently less browsing pressure in desert shrub vegetation in 1980 there were few significant changes in species composition. In addition, preferred forage species showed reduced reproduction while species of intermediate and poor forage value dis-played increased reproduction during this time. These data do not support traditional rangeland succession theory which states that enhanced reproduction of preferred species should follow grazing or browsing pressure reduction.     

Relationships between Scotch broom (Cytisus scoparius), soil nutrients, and plant diversity in the Garry oak savannah ecosystem

Scotch broom (Cytisus scoparius), is a leguminous shrub, native to the Mediterranean, which has invaded most of the remaining Garry oak savannah ecosystems in Oregon, Washington, and British Columbia. Here, it is considered to be a threat to the native plant community. We tested the hypothesis that broom would increase available soil nitrogen by comparing soil nutrients in contiguous broom-invaded and non-invaded sites. We then looked for changes in patterns of diversity in the herbaceous community that might indicate a role of Scotch broom in changing conditions following its invasion. Finally we carried out greenhouse assays to test whether broom had a greater impact on the growth of a native and an introduced grass compared to that of a native shrub. Broom was associated with only a weak trend in increased soil nitrogen, but a significant decrease in soil phosphorus was observed. Patterns of plant diversity differed between two sites. At one site, 60% of the plants whose abundances increased in the broom-invaded plots were introduced species while native species abundances decreased in the broom-invaded plots compared to broom-free plots. At the other site, 60% of the plants that caused the differences between broom-invaded and un-invaded plots were native species that were less abundant in the broom-invaded plots. Finally, in greenhouse assays grass growth was not affected as a result of being grown with broom; however, grasses appeared to produce more flowers when grown with broom. We conclude that broom does not necessarily modify soil nitrogen availability but may deplete soil phosphorus availability and that broom invasion can be associated with increase of exotic species and/or the decline of native species.     

Effects of simulated browsing on growth and leaf chemical properties in Colophospermum mopane saplings

Browsing intensity influences a plant's response to herbivory. Plants face a trade-off between investment in the production of secondary compounds and investment in growth. To elucidate this trade-off, we simulated four browsing intensities (0%, 50%, 75% and 100%) on mopane saplings, Colophospermum mopane (J.Kirk ex Benth.) J.Léonard, in a greenhouse experiment. This showed that, with increasing defoliation intensity, plants change their investment strategy. At intermediate levels of defoliation (50%), mopane saplings increased the synthesis of condensed tannins, so that tannin concentrations followed a hump-shaped relation with defoliation intensity, with significantly higher tannin concentration at intermediate defoliation levels. When defoliated heavily (75% and 100%), tannin concentrations dropped, and plants were carbon stressed as indicated by a reduced growth rate of the stem diameter, and leaf production and mean individual leaf mass were reduced. This suggests that, at intermediate defoliation intensity, the strategy of the plants is towards induced chemical defences. With increasing defoliation, the relative costs of the secondary metabolite synthesis become too high, and therefore, the plants change their growing strategy. Hence, browsers should be able to benefit from earlier browsing by either adopting a low or a relatively high browsing pressure.