Cascading effects of moose (<Emphasis Type="Italic">Alces alces</Emphasis>) management on birds

Large herbivores often have key functions in their ecosystems, and may affect ecosystem processes with cascading effects on other animals. The mechanisms often involve relocations of resources of various kinds, including reduction in resource availability following foraging and increase in resources from animal excreta. As large herbivore populations in Europe generally are intensely managed, management activities may interact with the activities of the herbivores themselves in the effect on other ecosystem components. We investigated the effects of moose (Alces alces) winter browsing, together with the effect of net nutrient input via supplementary winter feeding of moose on functional composition and species richness of birds in a boreal forest. Supplementary feeding stations for moose had a net zero effect on bird species richness and abundance, because negative effects of moose browsing were balanced by positive effects of nutrient input. Sites with a similar browsing intensity as at feeding stations but without nutrient input had lower abundance and species richness than feeding stations. Functional groups of bird species showed differing responses: birds nesting at or below browsing height were negatively affected by moose browsing, whereas species nesting above the browsing zone were positively affected by moose browsing. Insect-eating species responded negatively to moose browsing on birch but positively to nutrient input at feeding stations, whereas seed-eating species responded positively to birch browsing and negatively to feeding stations. This study showed that both high levels of cervid activity and human management interventions influence bird communities.     

Deciduous sapling responses to season and large herbivores in a semi‐arid African savanna

Impacts of large herbivores (>5 kg) on woody plants in African savannas are potentially most severe among plants shorter than 1.6 m. It is well established that severe browsing leads to longer shoots, yet prevents saplings from recruiting into adult size‐classes in African savannas. Increased shoot length, indicating faster shoot growth, is often associated with reduced concentrations of tannins and increased nutrient concentrations, suggesting carbon limitation. We hypothesized that, on average, large herbivores suppress stem height or circumference, but increase shoot length. We also hypothesized that if there were concomitant positive effects on nutrients, or negative effects on tannin concentrations, they would be greatest early in the wet season. We sampled saplings of four deciduous woody species (Acacia grandicornuta, Dichrostachys cinerea, Combretum apiculatum and Grewia flavescens) at different stages of the wet season in a large‐scale, long‐term herbivore exclusion experiment in Kruger National Park, South Africa. Plant height, shoot length and stem circumference were generally not adversely affected by large herbivores, suggesting C limitation is rarely present among deciduous saplings in semi‐arid African savannas, allowing them to tolerate browsing. Time since first rainfall emerged as a predominant factor consistently affecting nutrient and tannin concentrations, rather than large herbivores. Nitrogen and phosphorus generally decreased (by 20–50%), while condensed tannin concentration increased (150–350%) during the wet season, except for one species. We postulate that A. grandicornuta is less prone than other species to accumulating tannins during the wet season because of high investment of C in spines. Although nutrient and tannin concentrations were generally not affected by large herbivores, species‐specific responses were evident very early in the wet season, which is when herbivore populations are most likely to be affected by differential forage quality among plants.     

Disentangling the causes of damage variation by deer browsing on young Thuja plicata

Long‐lived trees experience different levels of damage due to mammalian herbivores. To untangle the mechanisms that underlie this variation, we combined chemical with dendrochronological analyses to study variation in browsing on Western redcedars (Thuja plicata) on Haida Gwaii (British Columbia, Canada). Since the last glaciation, Haida Gwaii forests had lacked large herbivore browser until Sitka black‐tailed deer (Odocoileus hemionus sitkensis) were introduced at the beginning of the 20th century. Dendrochronology yielded information on radial growth and plant annual responses to environmental stresses including herbivory. Secondary metabolite content and plant nutritional quality provided insights into proximate causes of food choices made by herbivores. We sampled lightly‐ and heavily‐browsed young trees at four sites: three clear‐cut sites with high browsing pressure and one old‐growth forest site where browsing pressure had, until recently, been lower. Heavily‐browsed young trees had lower concentrations of secondary metabolites and were of lower nutritive value than lightly‐browsed trees at all sites. Under high browsing pressure, tree growth patterns suggested that all young trees were initially severely browsed until some trees, currently scored as lightly‐browsed, started to escape deer. At the old‐growth site, both lightly‐ and heavily‐browsed trees tended to have lower overall average secondary metabolite concentrations than those of all other sites, a trend possibly related to greater canopy closure. Lightly‐browsed trees were older than heavily‐browsed ones which resulted, during the period of lower browsing pressure, in higher growth rate and a same pattern of change in growth from one year to the next year. This suggests that, under low browsing pressure, selection of young trees related to chemical defense was weak and that growth differences due to other factors than browsing could be expressed. Under strong browsing pressure, however, all young trees had equally low growth rates until trees with better genetic potential to produce effective defenses were able to escape deer. This suggests that selection by deer could occur on a long‐lived tree.     

Food preference and performance of the larvae of a specialist herbivore: variation among and within host-plant populations

Specialist herbivores are suggested to be unaffected by or attracted to the defense compounds of their host-plants, and can even prefer higher levels of certain chemicals. Abrostola asclepiadis is a specialist herbivore whose larvae feed on the leaves of Vincetoxicum hirundinaria, which contains toxic alkaloids and is unpalatable to most generalist herbivores. The food choice, leaf consumption and growth of A. asclepiadis larvae were studied to determine whether there is variation among and within host-plant populations in their suitability for this specialist herbivore. There was significant variation in food preference and leaf consumption among host-plant populations, but no differences were found in larval growth and feeding on different host-plant populations. A. asclepiadis larvae preferred host-plant populations with higher alkaloid concentrations, but did not consume more leaf material from plants originating from such populations in a no-choice experiment. There was also some variation in food preference of larvae among host-plant individuals belonging to the same population, suggesting that there was variability in leaf chemistry also within populations. Such variation in larval preference among host-plant genotypes and populations may create potential for coevolutionary dynamics in a spatial mosaic.     

Generalized plant defense: effects on multiple species

Summary Two species of lepidopteran herbivores, Manduca sexta (Sphingidae) and Trichoplusia ni (Noctuidae), were reared on synthetic diet containing either the alkaloid nicotine or the flavonoid rutin. Survival and pupal weight of the specialist M. sexta did not differ when larvae were reared on diet containing nicotine or rutin. In contrast, the generalist T. ni did not survive on diet containing 0.125% nicotine or greater, whereas larvae survived on all concentrations of rutin. These data demonstrate that the alkaloid nicotine is inhibitory toward generalist, but not specialist herbivores, whereas the flavonoid rutin has no effect on specialist herbivores and limited effects on generalist herbivores. Five species of Pseudomonas bacterial pathogens: P. syringae, P. syringae pv. angulata, P. syringae pv. tabaci, P. fluorescens, and P. solanacearum were grown on nutrient agar containing nicotine or rutin at concentrations ranging from 0.0 to 1.0% wet weight in 0.1% intervals. No species of Pseudomonas grew at concentrations greater than 0.5% nicotine when 10⁶ colony forming units (cfu) were used, but growth occurred at all concentrations of rutin when 10² cfu were used. These data indicate that nicotine was inhibitory to growth of both herbivores and pathogens, suggesting that certain plant secondary chemicals with high toxicity are of a generalized nature and affect multiple species. Differences in the sensitivity of organisms to allelochemicals such as generalist or specialist can make it appear that specific allelochemicals affect specific organisms, when in fact it is the tolerance of the organism to the plant chemical that is responsible. In four separate studies, the growth of M. sexta, T. ni and Helicoverpa zea was significantly lower on plants inoculated with P. solanacearum. Alteration in leaf quality by P. solanacearum was due to either reductions in leaf nutrients or increases in allelochemicals. We speculate that localized or systemic induction by both herbivores and pathogens can cause changes in leaf quality, effecting each other's subsequent colonization. The generalized nature of plant secondary compounds and potential reciprocal effects on induction by both species suggests that herbivores and pathogens may affect plant quality through induction and diffuse interactions of disparate species can alter the community of organisms colonizing a plant.     

Indirect effects of domestic and wild herbivores on butterflies in an African savanna

Indirect interactions driven by livestock and wild herbivores are increasingly recognized as important aspects of community dynamics in savannas and rangelands. Large ungulate herbivores can both directly and indirectly impact the reproductive structures of plants, which in turn can affect the pollinators of those plants. We examined how wild herbivores and cattle each indirectly affect the abundance of a common pollinator butterfly taxon, Colotis spp., at a set of long‐term, large herbivore exclosure plots in a semiarid savanna in central Kenya. We also examined effects of herbivore exclusion on the main food plant of Colotis spp., which was also the most common flowering species in our plots: the shrub Cadaba farinosa. The study was conducted in four types of experimental plots: cattle‐only, wildlife‐only, cattle and wildlife (all large herbivores), and no large herbivores. Across all plots, Colotis spp. abundances were positively correlated with both Cadaba flower numbers (adult food resources) and total Cadaba canopy area (larval food resources). Structural equation modeling (SEM) revealed that floral resources drove the abundance of Colotis butterflies. Excluding browsing wildlife increased the abundances of both Cadaba flowers and Colotis butterflies. However, flower numbers and Colotis spp. abundances were greater in plots with cattle herbivory than in plots that excluded all large herbivores. Our results suggest that wild browsing herbivores can suppress pollinator species whereas well‐managed cattle use may benefit important pollinators and the plants that depend on them. This study documents a novel set of ecological interactions that demonstrate how both conservation and livelihood goals can be met in a working landscape with abundant wildlife and livestock.     

Delayed induced responses of birch glandular trichomes and leaf surface lipophilic compounds to mechanical defoliation and simulated winter browsing

Changes in morphology and chemistry of leaf surface in response to herbivore damage may increase plant resistance to subsequent herbivore attack; however, there is lack of studies on induced responses of glandular trichomes and their exudates in woody plants and on effects of these changes on herbivores. We studied delayed induced responses in leaf surface traits of five clones of silver birch (Betula pendula Roth) subjected to various types of mechanical defoliation and simulated winter browsing. Glandular trichome density and concentrations of the majority of surface lipophilic compounds increased in trees defoliated during the previous summer. This induced response was systemic, since control branches in branch defoliated trees responded to the treatments similarly to defoliated branches, but differently from control trees. In contrast to defoliation treatments, simulated winter browsing reduced glandular trichome density on the following summer and had fewer effects on individual surface lipophilic compounds. Moreover, constitutive density of glandular trichomes was negatively correlated with induced total amount of lipophilic compounds per trichome, indicating a trade-off between constitutive and induced resistance in silver birch. Induced changes in leaf surface traits had no significant effect on leaf damage by chewers, miners and gall mites, but increased susceptibility of birch trees to aphids. However, leaf damage by chewers, miners and gall mites in defoliated (but not in control) trees was correlated with concentrations of some fatty acids and triterpenoids, although the direction of relationships varied among herbivore species. This indicates that induction of surface lipophilic compounds may influence birch resistance to herbivores. Our study thus demonstrated both specificity of elicitation of induced responses of birch leaf surface traits by different types of damage and specificity of the effects of these responses on different types of herbivores.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Regrowth and tannin production in woody and succulent karoo shrubs in response to simulated browsing

Allocation of carbon to chemical defences has often been suggested to be a direct response to browsing or grazing by herbivores. This study quantitatively compares total polyphenol and tannin production in response to simulated browsing of three karoo shrubs in order to test this induced defence hypothesis. The three species studied responded to browsing either by rapid regrowth or by increasing polyphenol production in the remaining tissues. The patterns did not follow any phylogenetic relationships but were weakly associated with the palatability of each species. The highly palatable deciduous species Osteospermum sinuatum, which is capable of rapid regrowth, showed no or very low levels of constitutive and browsing-induced total polyphenols, condensed tannins and protein-precipitating tannins. The evergreen sclerophyllous species Pteronia pallens showed a limited regrowth capacity and had intermediate levels of polyphenols, while the evergreen succulent species Ruschia spinosa showed almost no regrowth over the study period. R. spinosa contained the highest constitutive and browsing-induced levels of polyphenols, condensed tannins and protein-precipitating tannins. In two of the species more than one anti-herbivore defence feature co-occur. P. pallens foliage contains both hepatotoxins and polyphenols while R. spinosa has both structural (spines) and chemical defences. Responses of karoo shrubs to simulated browsing are interpreted as the result of passive alterations in plant chemistry rather than as an active defence response to herbivores.     

福建省马尾松和杉木针叶中7种营养元素含量特征

为了解福建省马尾松和杉木人工林养分需求特征,选取25个马尾松(Pinus massoniana)林和31个杉木(Cunninghamia lanceolata)林,测定了针叶中7种主要营养元素(C、N、P、S、K、Ca、Mg)含量。结果表明,马尾松针叶K、Ca、Mg含量显著低于杉木,分别是杉木的68%、14%、50%,表明杉木对矿质养分尤其Ca的需求远高于马尾松,提示马尾松比杉木更能适应矿质养分尤其Ca较贫瘠的立地。两树种针叶C、N、P、S含量及其化学计量比均表现出相似的内稳态特征,而Ca、Mg、K含量及其化学计量比的变异大,表明这3种矿质元素受立地条件的影响较大。两树种对N、P、K需求表现为协同正相关关系,N与Ca含量则为负相关关系,暗示持续N沉降或营林实践中长期施N肥可能抑制Ca的吸收。     

The impact of simulated moose densities on conifer aphids along a productivity gradient

Large herbivores have a significant impact on boreal forest ecosystems. The modification of resources through their feeding induces changes in trophic dynamics and affects the direction of interactions in a community. Moose Alces alces may decrease the available plant biomass for herbivorous insects on one of their main winter forage species in Fennoscandia, Scots pine Pinus sylvestris, and indirectly alter the abundance of invertebrates through exploitative competition. Moose browsing can also induce chemical, morphological, and phenological changes in plants, changing their nutritive value to insect herbivores such as aphids. Habitat productivity may further modify the responses of aphids to moose browsing. We studied the responses of the gray pine aphid Schizolachnus pineti to different moose densities, and their relations to habitat productivity by sampling pine branches and measuring the number of aphids on pine needles. The experimental setup consisted of 8 exclosures along a productivity gradient, where the feeding, defecation, and urination of 4 densities of moose had been simulated for 7 yr. We here show that high levels of simulated browsing decrease the amount of gray pine aphids in areas with high productivity. In areas with low productivity, however, simulated browsing had no such effect. Habitat productivity should therefore be considered as an important factor that may determine the strength of an areas buffering capacity to high moose densities. Low resource environments appear to be favourable to specialist conifer aphids on pines under high browsing pressure, but the performance of generalist insect herbivores might be lowered.     

Aphid genotypes vary in their response to the presence of fungal endosymbionts in host plants

Genetic variation for fitness‐relevant traits may be maintained in natural populations by fitness differences that depend on environmental conditions. For herbivores, plant quality and variation in chemical plant defences can maintain genetic variation in performance. Apart from plant secondary compounds, symbiosis between plants and endosymbiotic fungi (endophytes) can produce herbivore‐toxic compounds. We show that there is significant variation among aphid genotypes in response to endophytes by comparing life‐history traits of 37 clones of the bird cherry‐oat aphid Rhopalosiphum padi feeding on endophyte‐free and endophyte‐infected tall fescue Lolium arundinaceum. Clonal variation for life‐history traits was large, and most clones performed better on endophyte‐free plants. However, the clones differed in the relative performance across the two environments, resulting in significant genotype × environment interactions for all reproductive traits. These findings suggest that natural variation in prevalence of endophyte infection can contribute to the maintenance of genetic diversity in aphid populations.     

Elemental and isotopic analyses of mammalian fauna from Southern Africa and their implications for paleodietary research

Elemental analyses of mammalian bone (e.g., strontium-calcium ratios, or Sr/Ca) distinguish between herbivores and carnivores; however, the relationships among herbivores are unclear. To study this question, a modern faunal sample from the Nagupande Tsetse Control Area (Zambezi drainage, Northwestern Zimbabwe) was used. This collection has the advantage of well-established geographical controls in addition to a varied fauna, which includes both bovids and suids. The grazing/browsing dietary status of each species was ascertained by means of isotopic analysis of carbon. Clear differences were seen in the δ¹³C of grazing and browsing animals; a specialized grazer was found to have significantly lower Sr/Ca than less specialized grazers and browsers. In this study it was also possible to examine differences in Sr/Ca by sex; female warthogs were found to have significantly lower Sr/Ca than males. The variation in certain animal groups was found to be abnormal. Implications for reconstruction of prehistoric human diets using trace-element techniques are discussed.     

Fast and efficient DNA-based method for winter diet analysis from stools of three cervids: moose, red deer, and roe deer

Effects of cervid browsing on timber production, especially during winter, lead to economic losses in forest management. The aim of this study was to present an efficient DNA-based method which allows qualitative assessment of the winter diet from stools of moose (Alces alces), roe deer (Capreolus capreolus), and red deer (Cervus elaphus). The preliminary results of the diet composition of the three cervids from Poland were also presented with a special emphasis on moose. The electropherograms of the chloroplast intron trnL (UAA) P6 loop amplification products using g (fluorescence-labeled) and h primers revealed differences in the length of PCR products among various plant species eaten by these herbivores. In addition, the usage of species-specific primers allowed unambiguous identification of different gymnosperms and angiosperms. The preliminary moose diet analysis, based on winter fecal samples from the entire range of moose occurrence in Poland, revealed the presence of 15 to 24 tree, shrub, and herbaceous species. This fast, cost-efficient, and simple method proved also to be reliable for the diet analysis of red deer and roe deer. It may be a valuable tool in forest and conservation management, as well as a way of enhancing ecological studies focusing on the impact of herbivores on the ecosystems and their possible food niche overlap.     

Density- and growth stage-dependent responses to defoliation in two rhizomatous grasses

Summary Responses to defoliation were studied in two tallgrass prairie perennials (Andropogon gerardii and Panicum virgatum) established from seed at three densities. P. virgatum was also grown from transplanted rhizomes of established clones. Plants of both species displayed a continuum of responses to defoliation, from large reductions in biomass, tillering and seed production to significant increases in one or more performance measures. In crowded populations, defoliation shifted plants into subordinate positions within the competitive hierarchy. Plants competing intraspecifically and those that were initially small suffered more from defoliation than either plants grown at low density or those that were larger than their neighbors. At the highest plant density, the effects of defoliation or initial plant size were overshadowed by the effects of crowding. When defoliated and grown at similar densities, P. virgatum and A. gerardii grown from seed showed large reductions in biomass, seed production, and new rhizome production, but established P. virgatum ramets grown from rhizomes showed increases in these performance measures. Thus, herbivory may be particularly detrimental to P. virgatum during juvenile stages before perennating organs have developed. Overcompensation of P. virgatum clones in response to defoliation only occurred if all ramets within the clone were defoliated. In clones containing both defoliated and undamaged ramets, there were no differences in their performance, suggesting that genets are capable of integrating the effects of differential defoliation among shoots. Defoliated P. virgatum clones allocated a smaller fraction of their total biomass to new rhizomes, indicating that the short-term regrowth response following defoliation may incur a longer-term cost associated with gradual reduction in biomass of the perennating organs and reduced genet success.     

Aspects of dietary quality,nutrient assimilation and water balance in wild howler monkeys (Alouatta palliata)

Summary Wild howler monkeys were fed natural fruit and leaf diets in order to determine their ability to assimilate minerals (Ca, P, Mg, Si, Sr, Ba, Fe, Al, Mn, B, Cu, Li, V), and to assess their water and electrolyte budgets by the balance method. Results were extrapolated to the field situation so that the nutritional status of free-living monkeys could be examined in relation to their diet. The figs and young leaves of Ficus insipida and F. yoponensis trees (the most important foods in the dry season) provide adequate amounts of most required minerals, and an abundance of water. However, the copper requirement would not be met by this diet, which also provides only marginal amounts of sodium and phosphorus. These deficiencies may be corrected by consuming other available food items that are rich in Cu, Na or P. These results help explain the diversity that has been observed in the diet of wild howler monkeys, and suggest that these herbivores should feed selectively in order to obtain a nutritionally adequate diet.     

Multiple herbivores and coevolutionary interactions in an Ipomopsis hybrid swarm

Studies focusing on pairwise interactions between plants and herbivores may not give an accurate picture of the overall selective effect of herbivory, given that plants are often eaten by a diverse array of herbivore species. The outcome of such interactions may be further complicated by the effects of plant hybridization. Hybridization can lead to changes in morphological, phenological and chemical traits that could in turn alter plant–herbivore interactions. Here we present results from manipulative field experiments investigating the interactive effects of multiple herbivores and plant hybridization on the reproductive success of Ipomopsis aggregata formosissima X I. tenuituba. Results showed that ungulate herbivores alone had a net positive effect on plant relative fitness, increasing seed production approximately 2-fold. Caterpillars had no effect on plant relative fitness when acting alone, with caterpillar-attacked plants producing the same number of flowers, fruits and seeds as the uneaten controls. Caterpillars, however, significantly reduced flower production of ungulate browsed plants. Flower production in these plants, however, was still significantly greater (approximately 1.7-fold greater) than uneaten controls, likely leading to an increase in reproductive success through the paternal component of fitness given that fruit and seed production was not significantly different from that of herbivore-free controls. Although results suggest that herbivore imposed selection is pairwise, ungulates likely have a large influence on the abundance of, and hence the amount of damage caused by, caterpillar herbivores. Thus, because of the ecological interactions between ungulates and caterpillars, selection on Ipomopsis may be diffuse rather than pairwise, assuming such interactions translate into differential effects on plant fitness as herbivore densities vary. Plant hybridization had no significant effect on patterns of ungulate or caterpillar herbivory; i.e., no significant interactions were detected between herbivory and plant hybridization for any of the fitness traits measured in this study nor did plant hybridization have any significant effect on host preference. These results may be due to patterns of introgression or the lack of species-specific differences between I. aggregate formosissima and I. tenuituba. Plant hybridization per se resulted in lowered reproductive success of white colored morphs due in part to the effects of pollination. Although it appears that there would be strong directional selection favoring darker flower colors due to the lower reproductive success of the white colored morphs in the short run, the natural distribution of hybrids suggest that over the long run selection either tends to average out or there are no fitness differences among morphs in most years due to the additive fitness effects of hawkmoth and hummingbird pollinators.     

Genetic variation in defensive chemistry in Plantago lanceolata (Plantaginaceae) and its effect on the specialist herbivore Junonia coenia (Nymphalidae)

To examine genetic variation in defensive chemistry within and between natural populations of Plantago lanceolata, we performed a greenhouse experiment using clonal replicates of 15 genotypes from each of two populations, from a mowed lawn and an abandoned hayfield. Replicates of each genotype were harvested for determinations of aboveground biomass and leaf chemical content either at the beginning of the experiment (initial controls), after exposure to herbivory by larvae of Junonia coenia, a specialist on P. lanceolata (herbivory treatment), or at the end of the experiment without exposure to herbivory (final controls). Allocation to the iridoid glycosides aucubin and catalpol and the phenylpropanoid glycoside verbascoside displayed significant genetic variation within and between populations, and differed with leaf age. Significant genotypextreatment interactions indicated genetic variation in response of leaf chemistry to the treatments. There was no evidence for a cost of allocation to chemical defense: genetic correlations within and between chemical pathways and between defensive chemicals and aboveground growth were positive or nonsignificant. Although iridoid glycosides are known to be qualitative feeding stimulants for J. coenia, multiple regression of larval survivorship on leaf chemical content and shoot biomass indicated that larvae had lower survivorship on P. lanceolata ge-notypes with higher concentrations of aucubin in the leaves. Larval survivorship was unaffected by levels of catalpol and verbascoside. Thus, although specialist herbivores may respond to defensive chemicals as qualitative feeding stimulants, they do not necessarily have higher fitness on plant genotypes containing higher concentrations of these chemicals.     

Short-term effects of thinning and liming on forest soils of pitch pine and Japanese larch plantations in central Korea

The influences of thinning (50% of standing density) and liming (Ca+Mg, 2 Mg ha⁻¹) on soil chemical properties were investigated for 2 years (2001, 2002) in 40-year-old pitch pine (Pinus rigida Mill.) and 44-year-old Japanese larch (Larix leptolepis Gord.) plantations established on similar soils. In general, soil properties varied significantly among plantations and treatments. For both plantations, thinning significantly increased soil organic C (SOC) concentrations whereas there were no significant changes in soil pH and Ca and Mg concentrations. In addition, thinning increased total soil N and Na concentrations for the pitch pine plantation and available P concentration for the Japanese larch plantation in the second year after the treatment. Liming did not affect soil chemical characteristics for the pitch pine plantation except for Na concentration. However, for the Japanese larch plantation, liming significantly increased soil pH and K, Ca and Mg concentrations and decreased SOC and total soil N concentrations. For both plantations, soil Al concentration did not change after thinning and liming and decreased exponentially with increased pH values. The increases in SOC and total soil N concentrations after thinning were possibly due to increases in decomposition of organic matter and root death. Although differences were not statistically significant, soil available P concentration tended to increase at early stages of liming for both plantations. These results suggested that thinning and liming seemed to regulate soil chemical properties for pitch pine and Japanese larch plantations established on similar soils.     

Soil type, microsite, and herbivory influence growth and survival of Schinus molle (Peruvian pepper tree) invading semi-arid African savanna

Naturalization of Schinus molle (Anacardiaceae) has been observed in semi arid savanna of the Northern Cape Province of South Africa. However, with high dispersal ability, the species is expected to achieve greater densities and invade more widely. The study involved a field manipulation experiment over 14 months using a factorial block design to examine transplanted seedlings in different savanna environments. The experiments examine the effects of soil type (sandy and clay), microsite, and herbivores on seedling performance (establishment, growth and survival). Seedlings were grown in a greenhouse and individually transplanted into four treatment groups: in open grassland, under tree canopies, and with and without cages to exclude large herbivores (cattle and game). The same experiment was repeated in two different soil types: coarse sand and fine-textured clay soil. Results suggest that protection provided by canopies of large indigenous Acacia trees facilitates S. molle invasion into semi-arid savanna. In the field, S. molle seedlings performed considerably better beneath canopies of indigenous Acacia trees than in open areas regardless of soil type. Whether exposed or protected from large herbivores, no seedlings planted in open grassland survived the first winter. Although, seedlings grew better and had higher survival rates beneath tree canopies than in the open sites, exposure to large herbivores significantly decreased heights and canopy areas of seedlings compared with those protected from large herbivores. The effect was greater on clay soil than on sandy soil. The results suggest that low temperature (frost), and possibly inter-specific competition with grasses, may limit S. molle seedling establishment, survival and growth away from tree canopies in semi arid savannas. Low soil nutrient status and browsing may also delay growth and development of this species. The invasive potential of S. molle is thus greatest on fertile soils where sub-canopy microsites are present and browsing mammals are absent.     

Elk browsing increases aboveground growth of water-stressed willows by modifying plant architecture

In the northern elk wintering range of Yellowstone National Park, USA, wolf (Canis lupus) removal allowed elk (Cervus elaphus) to overbrowse riparian woody plants, leading to the exclusion of beaver (Castor canadensis) and a subsequent water table decline in many small stream valleys. Reduced elk browsing following wolf reintroduction may or may not facilitate willow (Salix sp.) recovery in these areas. To determine if the effect of elk browsing on willow interacts with that of beaver abandonment, we manipulated elk browsing and the water table in a factorial experiment. Under the condition of an ambient (low) water table, elk browsing increased shoot water potential (Ψ _s), photosynthesis per unit leaf area (A), stomatal conductance per unit leaf area (g _s), and aboveground current annual growth (CAG) by 50%. Elk browsing occurred entirely during dormancy and did not affect total plant leaf area (L). Improved water balance, photosynthetic rate, and annual aboveground productivity in browsed willows appeared to be due to morphological changes, such as increased shoot diameter and decreased branching, which typically increase plant hydraulic conductivity. An elevated water table increased Ψ _s, A, g _s, CAG, and L, and eliminated or lessened the positive effect of browsing on CAG for most species. Because low water tables create conditions whereby high willow productivity depends on the morphological effects of annual elk browsing, removing elk browsing in areas of water table decline is unlikely to result in vigorous willow stands. As large willow standing crops are required by beaver, a positive feedback between water-stressed willow and beaver absence may preclude the reestablishment of historical conditions. In areas with low water table, willow restoration may depend on actions to promote the re-establishment of beaver in addition to reducing elk browsing.