Factors driving mortality and growth at treeline: a 30-year experiment of 92 000 conifers

Understanding the interplay between environmental factors contributing to treeline formation and how these factors influence different life stages remains a major research challenge. We used an afforestation experiment including 92 000 trees to investigate the spatial and temporal dynamics of tree mortality and growth at treeline in the Swiss Alps. Seedlings of three high-elevation conifer species (Larix decidua, Pinus mugo ssp. uncinata, and Pinus cembra) were systematically planted along an altitudinal gradient at and above the current treeline (2075 to 2230 m above sea level [a.s.l.]) in 1975 and closely monitored during the following 30 years. We used decision-tree models and generalized additive models to identify patterns in mortality and growth along gradients in elevation, snow duration, wind speed, and solar radiation, and to quantify interactions between the different variables. For all three species, snowmelt date was always the most important environmental factor influencing mortality, and elevation was always the most important factor for growth over the entire period studied. Individuals of all species survived at the highest point of the afforestation for more than 30 years, although mortality was greater above 2160 m a.s.l., 50-100 m above the current treeline. Optimal conditions for height growth differed from those for survival in all three species: early snowmelt (ca. day of year 125-140 [where day 1 is 1 January]) yielded lowest mortality rates, but relatively later snowmelt (ca. day 145-150) yielded highest growth rates. Although snowmelt and elevation were important throughout all life stages of the trees, the importance of radiation decreased over time and that of wind speed increased. Our findings provide experimental evidence that tree survival and height growth require different environmental conditions and that even small changes in the duration of snow cover, in addition to changes in temperature, can strongly impact tree survival and growth patterns at treeline. Further, our results show that the relative importance of different environmental variables for tree seedlings changes during the juvenile phase as they grow taller.     

Foliar absorption of intercepted rainfall improves woody plant water status most during drought

A large proportion of rainfall in dryland ecosystems is intercepted by plant foliage and is generally assumed to evaporate to the atmosphere or drip onto the soil surface without being absorbed. We demonstrate foliar absorption of intercepted rainfall in a widely distributed, continental dryland, woody-plant genus: Juniperus. We observed substantial improvement in plant water status, exceeding 1.0 MPa water potential for drought-stressed plants, following precipitation on an experimental plot that excluded soil water infiltration. Experiments that wetted shoots with unlabeled and with isotopically labeled water confirmed that water potential responded substantially to foliar wetting, that these responses were not attributable to re-equilibration with other portions of the xylem, and that magnitude of response increased with water stress. Foliar absorption is not included in most ecological, hydrological, and atmospheric models; has implications for interpreting plant isotopic signatures; and not only supplements water acquisition associated with increases in soil moisture that follow large or repeated precipitation events, but also enables plants to bypass soil water uptake and benefit from the majority of precipitation events, which wet foliage but do not increase soil moisture substantially. Foliar absorption of intercepted water could be more important than previously appreciated, especially during drought when water stress is greatest.     

Tree species effects on decomposition and forest floor dynamics in a common garden

We studied the effects of tree species on leaf litter decomposition and forest floor dynamics in a common garden experiment of 14 tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Betula pendula, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pinus nigra, Pinus sylvestris, Pseudotsuga menziesii, Quercus robur, Quercus rubra, and Tilia cordata) in southwestern Poland. We used three simultaneous litter bag experiments to tease apart species effects on decomposition via leaf litter chemistry vs. effects on the decomposition environment. Decomposition rates of litter in its plot of origin were negatively correlated with litter lignin and positively correlated with mean annual soil temperature (MAT(soil)) across species. Likewise, decomposition of a common litter type across all plots was positively associated with MAT(soil), and decomposition of litter from all plots in a common plot was negatively related to litter lignin but positively related to litter Ca. Taken together, these results indicate that tree species influenced microbial decomposition primarily via differences in litter lignin (and secondarily, via differences in litter Ca), with high-lignin (and low-Ca) species decomposing most slowly, and by affecting MAT(soil), with warmer plots exhibiting more rapid decomposition. In addition to litter bag experiments, we examined forest floor dynamics in each plot by mass balance, since earthworms were a known component of these forest stands and their access to litter in litter bags was limited. Forest floor removal rates estimated from mass balance were positively related to leaf litter Ca (and unrelated to decay rates obtained using litter bags). Litter Ca, in turn, was positively related to the abundance of earthworms, particularly Lumbricus terrestris. Thus, while species influence microbially mediated decomposition primarily through differences in litter lignin, differences among species in litter Ca are most important in determining species effects on forest floor leaf litter dynamics among these 14 tree species, apparently because of the influence of litter Ca on earthworm activity. The overall influence of these tree species on leaf litter decomposition via effects on both microbial and faunal processing will only become clear when we can quantify the decay dynamics of litter that is translocated belowground by earthworms.     

Herbivores and variation in the composition of specific phenolics of boreal coniferous trees: a search for patterns

Coniferous trees of different species, or of the same species growing at different locations, vary in the extent to which they are attacked by various herbivores and pathogens. Plant secondary metabolites might be a key to understanding some of this variation. At the site level, we investigated if there was an intra- or interspecies pattern for individual compounds (or for groups of compounds) and their relationship to indices of plant nitrogen and plant productivity. For example, do plants exhibit similar covariance in defence compounds when evaluated across a number of sites varying in productivity? Here, we concentrated on the phenolic profile of Pinus sylvestris, Picea abies, Juniperus communis and Pinus contorta. Our results indicate striking differences in secondary chemistry profiles of the twigs including needles of the trees and in the inter-relationships amongst individual compounds and groups of compounds. Flavonols occurred in high variety in P. sylvestris and were highly correlated with each other, differing from P. contorta. But the results of the factor analyses indicate an underlying pattern for flavonols of the coumaroyl type for P. contorta. In contrast, the compounds of the other tree species showed a low degree of inter-correlation. Co-occurring phenolics of different tree species were not correlated. Overall, our analysis of site indices indicated that plant productivity was not a useful predictor for the concentration of specific phenolics. The relationship amongst plant nitrogen and specific phenolics might be the result of two defence strategies (one related and the other not related to nitrogen content). This might enable the plant to shift its defences against attacks with a high degree of flexibility.     

川西亚高山、高山森林土壤微生物生物量和酶活性动态特征

为深入了解川西亚高山/高山森林冬季生态学过程,于2008年11月─2009年10月,在土壤冻结初期、冻结期和融化期及植被生长季节,研究了不同海拔岷江冷杉林（Abies faxoniana）土壤微生物生物量和酶活性动态。各海拔森林土壤在冬季维持着较高的微生物生物量含量和酶活性,并随土壤冻融过程不断变化。土壤有机层和矿质土壤层冬季微生物生物量碳和氮含量及转化酶和尿酶活性均表现出受冻结初期土壤冻融循环影响显著降低,在冻结期变化不明显,在融化期急剧增加至融化后显著降低的趋势,且土壤有机层微生物生物量含量和酶活性在融化期具有一个明显的年高峰值。海拔变化显著影响了土壤酶活性,但对土壤微生物生物量不显著。土壤温度与土壤微生物生物量含量相关显著。这表明季节性冻融期是土壤生态过程的重要时期,土壤冻融格局显著影响川西亚高山/高山森林土壤微生物生物量和酶活性动态。     

Role of multidecadal climate variability in a range extension of pinyon pine

Evidence from woodrat middens and tree rings at Dutch John Mountain (DJM) in northeastern Utah reveal spatiotemporal patterns of pinyon pine (Pinus edulis Engelm.) colonization and expansion in the past millennium. The DJM population, a northern outpost of pinyon, was established by long-distance dispersal (approximately 40 km). Growth of this isolate was markedly episodic and tracked multidecadal variability in precipitation. Initial colonization occurred by AD 1246, but expansion was forestalled by catastrophic drought (1250-1288), which we speculate produced extensive mortality of Utah Juniper (Juniperus osteosperma (Torr.) Little), the dominant tree at DJM for the previous approximately 8700 years. Pinyon then quickly replaced juniper across DJM during a few wet decades (1330-1339 and 1368-1377). Such alternating decadal-scale droughts and pluvial events play a key role in structuring plant communities at the landscape to regional level. These decadal-length precipitation anomalies tend to be regionally coherent and can synchronize physical and biological processes across large areas. Vegetation forecast models must incorporate these temporal and geographic aspects of climate variability to accurately predict the effects of future climate change.     

Relationships between simulated water stress and mortality and growth rates in underplanted Toona ciliata Roem. in subtropical Argentinean plantations

Toona ciliata Roem. (Australian red cedar) requires a nurse-tree overstory to prevent damage from drought and irradiation in some regions of north-eastern Argentina. T. ciliata was planted in the understory of Pinus taeda L. (625 stems/ha), Pinus elliottii Engelm. × Pinus caribaea Morelet (625 stems/ha), and Grevillea robusta A. Cunn. (833 stems/ha) nurse trees, which were thinned to 0, 25, 50, 75 and 100% of the initial densities. We measured initial T. ciliata mortality and growth as well as Leaf Area Index (LAI) based on light transmission. T. ciliata soil water availability and its effect on early growth and mortality were examined by modelling drought stress using the two-dimensional forest hydrology model ForWaDy. Simulated patterns in T. ciliata water stress for the different overstory treatments were consistent with observed patterns of mortality. Early mortality was lowest with a G. robusta overstory, with corresponding lowest drought stress values and high modelled soil water contents in the top soil layer in intermediate and high overstory densities. Mortality was highest with a P. elliottii × P. caribaea overstory in treatments with the highest modelled drought stress values in the most open treatments. The model supported our field observations by indicating that water stress was an important limitation to T. ciliata survival and growth on our study sites. The linkage between T. ciliata establishment success, early growth and soil water availability as indicated by ForWaDy, leads us to conclude that the model is a suitable stand management tool for guiding establishment of T. ciliata plantations.     

Perennial-like adaptation mechanisms of annual legumes to limited irrigation

The hydrodynamic behavior of two annual legumes (Trifolium angustifolium L. and Onobrychis caput-galli (L.) Lam.) under water shortage was studied in a rain shelter experiment. Seeds were collected from natural grasslands of northern Greece and were sown in pots. Two months after seedlings' emergence, full irrigation (up to field capacity) and limited irrigation (40% of field capacity) were applied. During the vegetative period the leaf water potential and the relative water content were measured at seven day intervals in both treatments. T. angustifolium retained the lowest values of psi both under full (-0.11 to -1.78 MPa) and limited irrigation (-0.16 to -2.90 MPa), while the highest values in both cases were those of O. caput-galli (-0.05 to -0.5 MPa). The results suggested that T. angustifolium was the species mostly affected by limited water supply. T. angustifolium seemed to display adaptation mechanisms to drought similar to those of perennial plants. O. caput-galli displayed a more isohydric behavior by not altering its water potential under limited irrigation.     

Seed germination of three provenances of Pinus brutia (Ten.) as influenced by stratification, temperature and water stress

Seeds from three provenances of Pinus brutia were stratified for 0 or 45 d (days) at 4 +/- 1 degrees C and then germinated at 15 degrees C or 20 degrees C on filter paper saturated with polyethylene glycol solutions to provide water potentials of 0, -0.2 and -0.4 MPa (mega Pascal). Regardless of stratification, germination was lower at 15 degrees C than 20 degrees C for seeds of all provenances. Stratification significantly increased germination percentage at all water potentials regardless of the germination temperatures. Lowering the water potential to -0.4 MPa reduced germination for all three provenances in unstratified and stratified seeds when averaged for two temperatures, but reaction to the increased water stress was different among the provenances. Combined over two temperatures, stratified or unstratified seeds from the highest elevation (Cehennemdere) had the lowest germination performance at all water potentials, and stratified and nonstratified seeds from a coastal elevation (Silifke) had the highest germination parameters at the lowest water potential (-0.4 MPa). It might be concluded that seed germination and resistance to water stress vary according to provenance and stratification.     

宁夏六盘山8种木本植被的叶片水分利用效率

为了解宁夏六盘山地区主要树种的叶片水分利用效率（WUE）的种间和水分生境差异及季节变化,2010年6-9月份在宁夏六盘山具有半湿润气候和较多树种的香水河小流域和具有半干旱气候及较少树种的叠叠沟小流域,选择8个主要树种测定其叶片的 WUE。结果表明,（1）在香水河小流域,树种叶片 WUE 大小排序为油松 Pinus tablaeformis＞山桃 Prunus davidiana＞沙棘Hippophae rhamnoides＞辽东栎Quercus liaotungensis＞华北落叶松Larix principris-rupprechtii＞华山松Pinus armandii＞少脉椴Tilia paucicostata＞白桦Betula platyphylla;在不同生活型之间,叶片WUE的大小排序基本上呈现出灌木＞小乔木＞常绿针叶树＞落叶针叶树＞落叶阔叶树的变化规律;所有8个测试树种的叶片WUE的季节变化均为生长季初期较高,在中后期持续降低,但季节变幅随树种而异。（2）对2个小流域共有的测试树种（华北落叶松、山桃和沙棘）的叶片WUE进行了比较,结果表明存在着明显的水分生境差异,即在半干旱区极显著（p＜0.01）大于半湿润区,前者为后者的2.03倍以上。（3）对于沙棘、山桃、油松和华北落叶松,它们在较干旱气候类型立地和较干旱季节条件下都具有较高的叶片WUE,属抗旱能力较强的生态型节水树种,可选为干旱缺水区主要造林树种。此外,树种叶片WUE同时具有保守性和变异性,说明可采取适度抗旱锻炼来在一定范围内提高造林树种的WUE。     

Transpiration and hydraulic strategies in a pi?on-juniper woodland

Anthropogenic climate change is likely to alter the patterns of moisture availability globally. The consequences of these changes on species distributions and ecosystem function are largely unknown, but possibly predictable based on key ecophysiological differences among currently coexisting species. In this study, we examined the environmental and biological controls on transpiration from a pi?on-juniper (Pinus edulis-Juniperus osteosperma) woodland in southern Utah, USA. The potential for climate-change-associated shifts in moisture inputs could play a critical role in influencing the relative vulnerabilities of pi?ons and junipers to drought and affecting management decisions regarding the persistence of this dominant landscape type in the Intermountain West. We aimed to assess the sensitivity of this woodland to seasonal variations in moisture and to mechanistically explain the hydraulic strategies of P. edulis and J. osteosperma through the use of a hydraulic transport model. Transpiration from the woodland was highly sensitive to variations in seasonal moisture inputs. There were two distinct seasonal pulses of transpiration: a reliable spring pulse supplied by winter-derived precipitation, and a highly variable summer pulse supplied by monsoonal precipitation. Transpiration of P. edulis and J. osteosperma was well predicted by a mechanistic hydraulic transport model (R2 = 0.83 and 0.92, respectively). Our hydraulic model indicated that isohydric regulation of water potential in P. edulis minimized xylem cavitation during drought, which facilitated drought recovery (94% of pre-drought water uptake) but came at the cost of cessation of gas exchange for potentially extended periods. In contrast, the anisohydric J. osteosperma was able to maintain gas exchange at lower water potentials than P. edulis but experienced greater cavitation over the drought and showed a lesser degree of post-drought recovery (55% of pre-drought uptake). As a result, these species should be differentially affected by shifts in the frequency, duration, and intensity of drought. Our results highlight the sensitivity of this woodland type to potential climate-change-associated shifts in seasonal moisture patterns and demonstrate the utility of mechanistic hydraulic models in explaining differential responses of coexisting species to drought.     

西藏生态整体性水平测度

为了探讨资源-环境双重约束下地处我国生态环境脆弱区的西藏的生态整体性特征,利用生态环境质量指数和生态现代化指数（EMI）对地处我国西南边陲的西藏地区的生态现代化水平进行了分析。结果表明：全球生态现代化水平在区域之间存在比较明显的空间分异。2004年西藏生态现代化指数为50,在全国排名为3位,较2000年的排名16位有了明显的提高。生态进步指数、经济生态化指数好于其他4个少数民族地区,但社会生态化指数落后于其他4个少数民族地区,人均SO2排放与新疆、广西相同,但高于内蒙古和宁夏;1980—2007年西藏现代化程度虽然有所提升,但远低于世界平均水平和中等发达国家以及我国平均水平和其他4个少数民族地区;1996—2007年西藏区域环境水平呈＂W＂型变动态势。资源转化率和水污染指数呈＂N＂型不稳定变动,生态保护指数和环境治理指数呈剧烈变动,生态脆弱性和环境支持系统的不稳定性没有明显改观。因此,西藏应立足地缘和资源优势,充分考虑西藏生态地域、生态系统服务功能、生态资产、生态敏感性以及人类活动对生态环境胁迫等要素的综合影响,依据青藏高原高寒生态大区和高寒草甸生态区、青藏高原高山草原、高寒草甸生态地区生态区划,发展人工设计生态方案的生态重建途径。建立严格的生态补偿制度,逐步提升生态现代化水平。     

Pastoral Politics: Gaddi Grazing, Degradation, and Biodiversity Conservation in Himachal Pradesh, India

Two assumptions underlie the current conservation focus worldwide. The first is that democratic governments can restrict human resource use within protected areas, and the second is that human land use for subsistence leads to degradation and is incompatible with the maintenance of high levels of biological diversity. An examination of official policy documents over the past century indicates that Gaddi herders of Himachal Pradesh, northwestern Indian Himalaya, have used political influence to circumvent bureaucratic policies of exclusion and that there is an absence of scientific evidence to support the notion that Gaddi grazing leads to land degradation. Although grazing intensity has profoundly shaped the structure and composition of the Siwalik forests (the Gaddi winter grazing grounds), as demonstrated by transect-based data presented here, deviations from a supposed "climax" community need not constitute degradation. A growing rather than declining cattle population attests to the regenerative capacities of these forests. Within the alpine meadows grazed by the Gaddi in summer, mean plant species richness increased along transects originating at herder camps and extending 250 m north of border camp sites. Intense grazing pressure or heavy manuring by livestock bedded at night are likely to be responsible for the observed low species diversity adjacent to the campsite, but the effect is insignificant at the level of the overall landscape. Interviews with borders also suggest the presence of a sizable, though hunted, mammalian fauna in these high altitude meadows. Recognition of the difficulties associated with implementing restrictive policies, and the fact that human land-use practices need not lead to degradation or to a decline in biological diversity, should lead to more inclusive conservation policies within protected areas as well as an expansion of the conservation focus beyond protected-area boundaries.     

Microbial stress-response physiology and its implications for ecosystem function

Microorganisms have a variety of evolutionary adaptations and physiological acclimation mechanisms that allow them to survive and remain active in the face of environmental stress. Physiological responses to stress have costs at the organismal level that can result in altered ecosystem-level C, energy, and nutrient flows. These large-scale impacts result from direct effects on active microbes' physiology and by controlling the composition of the active microbial community. We first consider some general aspects of how microbes experience environmental stresses and how they respond to them. We then discuss the impacts of two important ecosystem-level stressors, drought and freezing, on microbial physiology and community composition. Even when microbial community response to stress is limited, the physiological costs imposed on soil microbes are large enough that they may cause large shifts in the allocation and fate of C and N. For example, for microbes to synthesize the osmolytes they need to survive a single drought episode they may consume up to 5% of total annual net primary production in grassland ecosystems, while acclimating to freezing conditions switches Arctic tundra soils from immobilizing N during the growing season to mineralizing it during the winter. We suggest that more effectively integrating microbial ecology into ecosystem ecology will require a more complete integration of microbial physiological ecology, population biology, and process ecology.     

长白山不同海拔岳桦(Betula ermanii)的光合生理

通过研究不同海拔(1 750 m,1 900 m,2 000 m)岳桦的光合生理,探讨岳桦叶片对高山环境的适应特征.结果表明:与海拔1 750 m相比,1 900 m和2 000 m处植物叶片具有较低的暗呼吸速率(Rd)、光补偿点(LCP)和CO2补偿点(CCP),较高的最大光合速率(Amax)、光近饱和点(LK)、表观羧化速率(CE)、最大羧化速率(Vcmax)和最大电子传递速率(Jmax).1 900 m处光化学淬灭系数(qP)和作用光存在时PSⅡ实际光化学量子效率(ФpsⅡ)最高,2 000 m处非光化学淬灭系数(NPQ)则最高.随海拔升高,岳桦叶片水分利用效率(WUE)先升高再降低.与1 750 m相比,1 900和2 000 m处植物叶片具有较高的气孔导度(gs)和CO2吸收速率(Pn/Ci斜率).分析表明:1 900 m是岳桦的最适生长区,具有较高的光合能力,且所受胁迫较小.图6表3参26     

Impact of low dose of organophosphate, monocrotophos on the epithelial cells of gills of Cyprinus carpio communis Linn.--SEM study

The paper deals with the deleterious changes at ultrastructural level of the epithelial cells of gills of Cyprinus carpio communis Linn. upon exposure to 1/10th of LC50 of monocrotophos which is considered to be insignificant concentration from toxicological point of view. The gills of the fish are the primary corridor formolecularexchange between the internal milieu of a fish and its environment. Gills perform numerous functions such as oxygen uptake and CO2 excretion, osmoregulation, acid-basic balance, excretion of nitrogenous compounds and taste. Hazardous chemicals present in water may alter the morphology of the epithelial cells of gills of the fish, which may affect the process of diffusion of gases and ultimately the overall health of the fish. To prove this fact Cyprinus carpio communis Linn. was kept in water for 30 days having low concentration of 0.038 ppm (1\10th of LC50) of monocrotophos and an attempt was made to study the different types of degenerations produced in the epithelial cells of gills as compared to the normal epithelial cells of gills of this culturable fish using Scanning Electron Microscope (SEM) technique. The ultrastructural changes due to the toxic exposure at finer scale were thinning of microridges, upliftment of epithelial cells, development of hyperplasia, decrease in the density of mucous cells which are considered to be the first line of defence and total dystrophy of epithelial tissue. Thus, it is opined that a low concentration of monocrotophos has the potential to bring different type of degenerations at finer scale hence affecting the fish's health drastically and altering the fitness of the fish in water even having insignificant amount of this toxicant in the ambient water     

Constraints on tree seedling establishment in montane grasslands of the Valles Caldera, New Mexico

Montane and subalpine grasslands are prominent, but poorly understood, features of the Rocky Mountains. These communities frequently occur below reversed tree lines on valley floors, where nightly cold air accumulation is spatially coupled with fine soil texture. We used field experiments to assess the roles of minimum temperature, soil texture, grass competition, and ungulate browsing on the growth, photosynthetic performance, and survival of transplanted ponderosa pine (Pinus ponderosa) seedlings at 32 sites straddling such reversed tree lines in the Valles Caldera National Preserve (VCNP) of the Jemez Mountains, New Mexico (USA). Seedling growth increased most strongly with increasing nighttime minimum temperatures away from the valley bottoms; seedlings experiencing the coldest temperatures on the caldera floor exhibited stunted needles and often no measurable height growth. Based on the chlorophyll fluorescence ratios PhiPSII and Fv/Fm, we found that low minimum temperatures, low soil moisture, and fine soil texture all contributed to photoinhibition. Neighboring herbs had only minor negative effects on seedlings. We found no effect of ungulates, but golden-mantled ground squirrels (Spermophilus lateralis) caused substantial seedling mortality. Second-year seedling survival was highest on sandy soils, and third-year survival was highest at sites with higher minimum temperatures. We conclude that differential tree seedling establishment driven by low minimum temperatures in the valley bottoms is the primary factor maintaining montane grasslands of the VCNP, although this process probably operated historically in combination with frequent surface fire to set the position of the tree line ecotone. As at alpine tree lines, reversed tree lines bordering montane and subalpine grasslands can represent temperature-sensitive boundaries of the tree life form.     

塔克拉玛干绿洲外围胡杨林的水分特征研究

塔克拉玛干沙漠南缘地区年均降水量不足50mm,胡杨在这一地区绿洲外围自然植被中占有重要地位.对塔克拉玛干沙漠绿洲外围胡杨林水分特征的研究表明,在生境地下水含量较稳定的条件下,胡杨林受到的水分胁迫程度不大,干旱胁迫未对胡杨林的持续存在构成主要威胁.胡杨的水分生理特点显示植物更加依靠稳定的水分供应来应对极端干旱的环境条件,并且在蒸腾作用中形成了细胞水平上的生理适应策略.因此,保持地下水的稳定是维持绿洲前沿天然胡杨林持续存在的关键因素.     

Monoterpene chemodiversity of ponderosa pine in relation to herbivory and bark beetle colonization

Summary. Ponderosa pine, Pinus ponderosa Laws. (Pinaceae), forests in Arizona have suffered from a nine-year period of drought and bark beetle, Ips lecontei Swaine (Coleoptera: Scolytidae), outbreaks. Abiotic and biotic stress in ponderosa pine results in the induced synthesis of certain monoterpenes that may in turn affect bark beetle behavior and survival. In this study, we investigate whether induced monoterpene production could result in a different monoterpene composition that remains stored in the needles or the trunk resin of the tree. Needle and resin samples in addition to trunk cores were collected from ponderosa pines at three locations in Arizona. Ungulate browsing induced a significant increase in limonene (P=0.010) and in chemodiversity (P=0.009), a measure of the evenness of distribution among the monoterpenes present in needles. We compared the level of ‘stress’ of the trees by measuring the thickness of annual rings in living trees and those that were killed by bark beetles. Where drought occurred, the spacing of annual rings from the last 10 years of trees killed by bark beetles was significantly smaller (P=0.020) compared to living trees. There was no difference in the monoterpene composition between the core sections of closest spacing of annual rings (stressed years) compared to the sections of widest spacing, which indicates that monoterpenes are distributed evenly throughout the extended resin system. In the area where the degree of drought was less overall, none of the individual monoterpenes present in the resin was related to bark beetle killed trees. However, about half the living pines had resin in which one of the major monoterpenes (α-pinene, Δ³-carene, and limonene) was absent, and these trees had a lower monoterpene chemodiversity compared to trees killed by bark beetles. Trees with these three major monoterpenes, corresponding to the average relative proportion in living pines at that location, may sustain higher selection and colonization by bark beetles.     

Shifting prey selection generates contrasting herbivore dynamics within a large-mammal predator-prey web

Shifting prey selection has been identified as a mechanism potentially regulating predator-prey interactions, but it may also lead to different outcomes, especially in more complex systems with multiple prey species available. We assessed changing prey selection by lions, the major predator for 12 large herbivore species in South Africa's Kruger National Park. The database was provided by records of found carcasses ascribed to kills by lions assembled over 70 years, coupled with counts of changing prey abundance extending over 30 years. Wildebeest and zebra constituted the most favored prey species during the early portion of the study period, while selection for buffalo rose in the south of the park after a severe drought increased their vulnerability. Rainfall had a negative influence on the proportional representation of buffalo in lion kills, but wildebeest and zebra appeared less susceptible to being killed under conditions of low rainfall. Selection by lions for alternative prey species, including giraffe, kudu, waterbuck, and warthog, was influenced by the changing relative abundance and vulnerability of the three principal prey species. Simultaneous declines in the abundance of rarer antelope species were associated with a sharp increase in selection for these species at a time when all three principal prey species were less available. Hence shifting prey selection by lions affected the dynamics of herbivore populations in different ways: promoting contrasting responses by principal prey species to rainfall variation, while apparently being the main cause of sharp declines by alternative prey species under certain conditions. Accordingly, adaptive responses by predators, to both the changing relative abundance of the principal prey species, and other conditions affecting the relative vulnerability of various species, should be taken into account to understand the interactive dynamics of multispecies predator-prey webs.