浙江天童木本植物小枝的“大小-数量”权衡

枝条大小和数量关系反映了植物适应环境胁迫的构型和生物量分配策略。该研究以浙江天童木本植物为对象, 通过对小枝大小(横截面积)与数量(稠密度)关系的研究发现: 1)小枝稠密度与枝截面积显著负相关(斜率为-1.32, CI = -1.48- -1.17; p < 0.05); 2)在相同曝光度, 在II级曝光环境(植株40%-80%暴露在直射光中)中常绿植物比落叶植物单位小枝截面积的小枝稠密度高, 而在I级(植株<40%暴露在直射光中)和III级(>80%暴露在直射光中)曝光环境中, 小枝稠密度在两种生活型间无显著差异; 3)在不同曝光度下, 常绿植物单位枝条在I和II级比III级曝光水平具有更高的小枝稠密度; 但落叶植物单位枝条的小枝稠密度在3个曝光水平相同; 4)相同枝条大小下, 4 m以下灌木比4 m以上的亚乔木和乔木具有更高的小枝稠密度。总之, 天童地区木本植物的小枝“大小-数量”关系符合Corner法则(描述枝叶“大小-数量”关系的法则), 且在不同生活型间存在差异, 常绿植物相对于落叶植物, 灌木相对于乔木具有较高的小枝稠密度, 从而有利于它们适应光资源的限制。     

Predators,fire or resources: What drives the distribution of herbivores in fragmented mesic forests?

Trophic interactions and disturbance events can shape the structure and function of ecosystems. However, the effects of drivers such as predation, fire and climatic variables on species distributions are rarely considered concurrently. We used a replicated landscape‐scale predator management experiment to compare the effects of red fox Vulpes vulpes control, time‐since‐fire, vegetation type and other environmental variables on native herbivore distributions. Occurrence data for four native herbivores and an invasive predator – the red fox – were collected from 240 sites across three baited (for lethal fox control) and three unbaited forest blocks (4659–9750 ha) in south‐western Victoria, Australia, and used to build species distribution models. The herbivore taxa were as follows: red‐necked wallaby Macropus rufogriseus, black wallaby Wallabia bicolour, grey kangaroo Macropus fuligenosus and Macropus giganteus and common brushtail possum Trichosurus vulpecula. Fox control and fire had little effect on herbivore occurrence, despite the literature suggesting it can influence abundance, while climate, proximity to farmland and topography were more influential. This may be because the region’s high productivity and agricultural pastures subsidise food resources for both predators and prey within the forest blocks and so dampen trophic interactions. Alternatively, these drivers may affect herbivore abundance, but not herbivore occurrence. Understanding the drivers of herbivore distributions is an important step in predicting the effects of herbivory on other species, particularly after management interventions such as predator control and prescribed burns.     

Exploitation ecosystems and trophic cascades in non‐equilibrium systems: pasture – red kangaroo – dingo interactions in arid Australia

The exploitation ecosystems hypothesis (EEH) proposes that 1) plant biomass reflects the primary productivity of an ecosystem modified by the regulating effect of herbivory, and 2) herbivore abundance reflects the productivity of plants modified by the regulating effect of predation. Primary productivity thus determines the number of trophic levels in an ecosystem and the extent to which bottom–up and top–down regulation influence the biomass ratios of adjacent and non‐adjacent trophic levels (i.e. trophic cascading). We constructed an interactive model of plant (pasture), herbivore (red kangaroo Macropus rufus) and predator (dingo Canis lupus dingo), a system in which trophic cascades have been suggested to occur, and used it to test the effects of increasing stochastic variation in primary productivity and dingo culling on predictions of the EEH. The model contained four feedback loops: the predator–herbivore and herbivore–plant feedback loops, and the predator and plant density‐dependent feedback loops. The equilibrium conditions along the primary productivity gradient reproduced the three zones of trophic dynamics predicted by the EEH, plus an additional zone at productivities above which the maximum density of a predator is achieved due to social regulation: that zone is characterized by increasing herbivore density and decreasing plant biomass. Culling dingoes produced trophic cascades that were strongly attenuated at primary productivities below which the maximum density of dingoes was attained. Results were robust to uncertainty in kangaroo off‐take by dingoes and to the efficacy of dingo culling, but prey switching by dingoes from red kangaroos to reptiles would weaken trophic cascades. We conclude that social regulation of carnivores has important implications for expression of the EEH and trophic cascades, and that attenuation of trophic cascades increases with increasing stochasticity in primary productivity. Our model also provides a framework for understanding the conditions in which dingo‐mediated trophic cascades might be expected to occur, and generates testable predictions about the effects of higher dingo densities (e.g. by stopping culling or reintroduction to former range) on kangaroo and pasture dynamics.     

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.     

Spine production is induced by fire: a natural experiment with three Berberis species

Earlier studies indicate that some plant species allocate more mass to produce longer spines in shoots resprouting after browsing. Here we present, for the first time, evidence that fire induces a similar response. Many terrestrial herbivores may benefit from fire through the enhanced availability of fast growing species colonizing or re-sprouting in burned areas. It is less clear whether post-fire plant growth responds to the enhanced risk of herbivory by an increased investment in defensive traits. In this study, we tested whether the production of spines is influenced by the set of environmental conditions that result from fire events. We compared the resource allocation pattern of resprouting shoots from three Berberis species growing in two areas that burned 1999 with samples collected from unburned areas within the same plant communities. We divided the shoot into three main components: supporting tissue (twigs), assimilating tissue (leaves) and defensive structures (spines). We found that plants resprouting after fire allocated more mass to spines and leaves but not twigs. This resulted in a higher density of both spines and leaves. Spines were significantly longer in plants resprouting after fire. Leaves were shorter at the apical end of the shoot, but did not show any significant change in size following fire. We suggest that this type of post-fire response may be a general adaptation to pruning and leaf picking by browsing herbivores in arid and semi-arid regions. Changes in the browsing pressure following fire will determine the fitness value of this response.     

Comparative finite element analysis of the cranial performance of four herbivorous marsupials

Marsupial herbivores exhibit a wide variety of skull shapes and sizes to exploit different ecological niches. Several studies on teeth, dentaries, and jaw adductor muscles indicate that marsupial herbivores exhibit different specializations for grazing and browsing. No studies, however, have examined the skulls of marsupial herbivores to determine the relationship between stress and strain, and the evolution of skull shape. The relationship between skull morphology, biomechanical performance, and diet was tested by applying the finite element method to the skulls of four marsupial herbivores: the common wombat (Vombatus ursinus), koala (Phascolarctos cinereus), swamp wallaby (Wallabia bicolor), and red kangaroo (Macropus rufus). It was hypothesized that grazers, requiring stronger skulls to process tougher food, would have higher biomechanical performance than browsers. This was true when comparing the koala and wallaby (browsers) to the wombat (a grazer). The cranial model of the wombat resulted in low stress and high mechanical efficiency in relation to a robust skull capable of generating high bite forces. However, the kangaroo, also a grazer, has evolved a very different strategy to process tough food. The cranium is much more gracile and has higher stress and lower mechanical efficiency, but they adopt a different method of processing food by having a curved tooth row to concentrate force in a smaller area and molar progression to remove worn teeth from the tooth row. Therefore, the position of the bite is crucial for the structural performance of the kangaroo skull, while it is not for the wombat which process food along the entire tooth row. In accordance with previous studies, the results from this study show the mammalian skull is optimized to resist forces generated during feeding. However, other factors, including the lifestyle of the animal and its environment, also affect selection for skull morphology to meet multiple functional demands. J. Morphol. 276:1230–1243, 2015. © 2015 Wiley Periodicals, Inc.     

Patterns in the abundance of kangaroo populations in arid Australia are consistent with the exploitation ecosystems hypothesis

The exploitation ecosystems hypothesis (EEH) makes predictions about trophic interactions along gradients of primary productivity. The EEH has been shown to apply to a wide range of terrestrial environments but its applicability to arid environments has received little attention. One reason for this is that arid environments may not satisfy the assumptions of the EEH because dearth of water may limit biological activity in both temporal and spatial contexts. The EEH predicts that herbivore biomass should increase linearly with primary productivity in the absence of predators; but when predators are present herbivore biomass will remain relatively constant due to top down regulation. We tested this prediction in an arid environment using rainfall as a proxy of primary productivity and an index of the abundance of the dominant herbivores (kangaroos Macropus spp.). We compared an index of kangaroo abundance at 18 areas situated along a gradient of mean annual rainfall in areas where a top predator (the dingo Canis lupus dingo) was rare and common. We also explored the relationship between the density of artificial water points (AWPs) and kangaroo abundance to investigate if the resource subsidy provided by AWPs allows kangaroos to persist in high numbers. Consistent with the EEH, kangaroo abundance showed a weak relationship with mean annual rainfall in the presence of dingoes but increased with increasing annual rainfall in the absence of dingoes. The density of AWPs was a poor predictor of kangaroo abundance. Our analysis of macro‐ecological patterns suggests that kangaroo populations are primarily top down regulated in the presence of dingoes, but are bottom up regulated in the absence of dingoes. Our findings provide evidence that top down regulation can prevail over bottom up regulation of herbivore populations in arid ecosystems and highlights the usefulness of the EEH as a predictor of macro‐ecological patterns of species abundance.     

Evolutionary irony: evidence that ‘defensive’ plant spines act as a proximate cue to attract a mammalian herbivore

Many plants produce structural defenses to deter feeding by herbivores. However, many previous studies testing whether spines are effective at defending against mammalian herbivores have produced equivocal results. These ambiguous results are hypothesized to be due to herbivore counter‐adaptations. We investigated potential counter‐adaptations in a population of white‐throated woodrats Neotoma albigua that specialize on cactus by investigating feeding behavior and preference for cacti varying in spinescence. Neotoma albigula exhibited a unique behavior of clipping cactus spines, which renders these defenses ineffective. Strikingly, these woodrats chose to collect spiny cacti over experimentally de‐spined cacti, demonstrating that spines act as a proximal cue that attracts woodrats. This attraction is likely due to the higher protein and lower fiber content of spiny cacti compared to naturally non‐spiny cacti. Thus, the ‘defensive’ spines of cacti are ineffective against a specialist herbivore and instead serve as an indicator of nutritional quality that promotes herbivory. Our results support the ‘rule‐of‐thumb’ hypothesis of foraging, which states that herbivores forage according to obvious visual cues that are indicative of nutritional content, rather than sampling nutrient composition of plants. We propose that specialist herbivores are unique systems in which to study other counter‐adaptations to structural defenses and ‘rule‐of‐thumb’ foraging strategies.     

亚热带常绿林不同冠层小枝叶面积-叶生物量关系研究

叶面积与叶生物量的关系对于理解植物叶片的碳收益和投资权衡策略具有重要意义。收益递减假说认为植物的叶面积与叶生物量成显著异速生长关系，其异速生长指数<1.0，但该假说是否适用于不同生活型（常绿与落叶）亚热带木本植物不同冠层高度（上下冠层）当年生小枝的叶片仍不清楚。以江西亚热带常绿阔叶林的69种常绿与落叶木本植物当年生小枝上的叶为研究对象，采用标准化主轴回归估计（standardized major axis estimation，SMA）方法检验不同冠层高度和生活型叶面积与叶生物量的异速生长关系。结果显示：（1）当年生小枝叶生物量在不同冠层高度和生活型的植物中无显著差异（P>0.05），叶面积在常绿和落叶植物中有显著差异（P<0.05），常绿和落叶植物的比叶重存在显著差异（P<0.05），而落叶植物的比叶重在不同冠层高度存在显著差异（P<0.05），同一冠层，常绿植物比叶重显著高于落叶植物（P<0.05）；（2）69种植物的叶面积与叶生物量异速生长指数具有物种特异性，60.9%的物种叶面积与叶生物量呈等速生长关系；（3）不同冠层和生活型植物的叶面积与叶生物量呈等速生长关系，但其异速生长常数在不同冠层高度与生活型间存在差异。这些结果表明冠层高度和生活型未改变叶面积-生物量之间的等速生长关系，不支持"收益递减"假说。     

Effects of Snow on Sitka Black-Tailed Deer Browse Availability and Nutritional Carrying Capacity in Southeastern Alaska

ABSTRACT Snow affects the nutritional ecology of northern ungulates during winter through burial of important winter forages. We used nonlinear regression analyses to model snow-burial dynamics of blueberry (Vaccinium spp.) browse biomass, a key winter food item of Sitka black-tailed deer (Odocoileus hemionus sitchensis) in southeastern Alaska, USA. During November 2003—March 2004 we collected data from 546 individually marked twigs located on 100 plants of differing sizes and architectures across a range of snow depths. In general, browse biomass became buried and unavailable to deer at snow depths substantially lower than prewinter twig heights. Plant architecture and plant height were related to the probability of a twig being buried. Probability of twig burial was higher on plants with lateral than on those with erect architectures. Twig height also affected the probability of burial by snow but the relationship was complex. For twigs located on erect plants, probability of burial was greatest for twigs near the bottom and top of the plant due to ground-up burial and bending of flexible apex stems, respectively. We used estimated nonlinear equations to model blueberry browse availability in a simulated upland old-growth habitat patch subject to a range of snow depths. We then compared subsequent estimates of deer winter nutritional carrying capacity for this habitat patch to findings derived using an alternative, simple linear (ground-up) model of winter-browse burial by snow. Comparisons indicated that ground-up models of browse burial overestimated browse availability and nutritional carrying capacity for most snow depths. Our findings demonstrate the importance of applying detailed snow-burial models when characterizing nutritional landscape of northern ungulates during winter.     

Plant biological warfare: thorns inject pathogenic bacteria into herbivores

Thorns, spines and prickles are among the rich arsenal of antiherbivore defence mechanisms that plants have evolved. Many of these thorns are aposematic, that is, marked by various types of warning coloration. This coloration was recently proposed to deter large herbivores. Yet, the mechanical defence provided by thorns against large herbivores might be only the tip of the iceberg in a much more complicated story. Here we present evidence that thorns harbour an array of pathogenic bacteria that are much more dangerous to herbivores than the painful mechanical wounding by the thorns. Pathogenic bacteria like Clostridium perfringens, the causative agent of the life-threatening gas gangrene, and others, were isolated and identified from date palm (with green-yellow-black aposematic spines) and common hawthorn (with red aposematic thorns). These thorn-inhabiting bacteria have a considerable potential role in antiherbivory, and may have uniquely contributed to the common evolution of aposematism (warning coloration) in thorny plants.     

Responses of woody vegetation to exclusion of large herbivores in semi‐arid savannas

The Nkuhlu large‐scale long‐term exclusion experiment in Kruger National Park was designed to study the long‐term effects of large herbivores on vegetation. One treatment excludes elephants, another excludes all herbivores larger than hares and another one comprises an open, control area. Vegetation monitoring was implemented in 2002 when a baseline survey was conducted prior to exclusion. Monitoring was repeated 5 years after exclusion. Data from the surveys were analysed to establish how structure and composition of woody vegetation had changed 5 years after herbivore exclusion. The analysis showed that neither plant assemblage nor mean vegetation height had changed significantly since exclusion. However, both species richness and density of woody plants increased 5 years after exclusion of all large herbivores, but not after the exclusion of elephants alone. One already common species, Dichrostachys cinerea, became more common after excluding all large herbivores compared with either no exclusion or elephant exclusion, possibly leading to competitive suppression of other species. Species other than D. cinerea tended to either increase or decrease in density, but the changes were insufficient to induce significant shifts in the overall assemblage of woody plants. The results indicate that after 5 years of exclusion, the combined assemblage of large herbivores, and not elephants alone, could induce changes in species richness and abundances of woody plants, but the effect was so far insufficient to induce measureable shifts in the assemblages of woody plants. It is possible that assemblages will change with time and increasing elephant numbers may amplify future changes.     

Modeling the dynamics of woody plant–herbivore interactions with age-dependent toxicity

In this paper we study the effects that woody plant chemical defenses may have on interactions between boreal hares that in winter feed almost entirely on twigs. We focus particularly on the fact that toxin concentration often varies with the age of twig segments. The model incorporates the fact that the woody internodes of the youngest segments of the twigs of the deciduous angiosperm species that these hares prefer to eat are more defended by toxins than the woody internodes of the older segments that subtend and support the younger segments. Thus, the per capita daily intake of the biomass of the older segments of twigs by hares is much higher than their intake of the biomass of the younger segments of twigs. This age-dependent toxicity of twig segments is modeled using age-structured model equations which are reduced to a system of delay differential equations involving multiple delays in the woody plant-hare dynamics. A novel aspect of the modeling was that it had to account for mortality of non-consumed younger twig segment biomass when older twig biomass was bitten off and consumed. Basic mathematical properties of the model are established together with upper and lower bounds on the solutions. Necessary and sufficient conditions are found for the linear stability of the equilibrium in which the hare is extinct, and sufficient conditions are found for the global stability of this equilibrium. Numerical simulations confirmed the analytical results and demonstrated the existence of limit cycles over ranges of parameters reasonable for hares browsing on woody vegetation in boreal ecosystems. This showed that age dependence in plant chemical defenses has the capacity to cause hare-plant population cycles, a new result.     

The impact of elevated CO2 on plant-herbivore interactions: experimental evidence of moderating effects at the community level

Surprisingly little research has been published on the responses to elevated [CO₂] at the community level, where herbivores can select their preferred food. We investigated the combined effects of atmospheric [CO₂] and herbivory on synthesised plant communities growing on soils of different fertility. Factorial combinations of two [CO₂] (350 or 700 l l⁻¹), two fertility (fertilised or non-fertilised), and two herbivory (herbivores present or absent) treatments were applied to a standard mixture of seven fast- and eight slow-growing plants in outdoor microcosms. The herbivores used were the grain aphid (Sitobion avenae) and the garden snail (Helix aspersa). We measured plant biomass, foliar nitrogen and soluble tannin concentration, aphid fecundity, and snail growth, fecundity, and feeding preferences over one growing season. Elevated [CO₂] did not have a significant impact on (1) the combined biomass of fast-growing or slow-growing plants, (2) herbivore feeding preferences, or (3) herbivore fitness. There was, however, a significant biomass increase of Carex flacca (which represented in all cases less than 5% of total live biomass), and some chemical changes in unpalatable plants under elevated [CO₂]. The herbivory treatment significantly increased the biomass of slow-growing plants over fast-growing plants, whereas fertilisation significantly increased the abundance of fast-growing plants over slow-growing plants. Predictions on the effects of elevated [CO₂] based on published single-species experiments were not supported by the results of this microcosm study. Received: 30 November 1997 / Accepted: 24 July 1998     

Root herbivore performance suppressed when feeding on a jasmonate‐induced pasture grass

1. Plants defend themselves from insect herbivore attack using a range of physical and chemical defences which are in many cases regulated by phytohormones such as jasmonates. While much more is known about how jasmonates regulate defence against above‐ground herbivores (e.g. herbivores of leaves), there is increasing interest in how they influence below‐ground defences. 2. For the Poaceae, most below‐ground studies focus on highly domesticated cereals. Here it is demonstrated how exogenous application of methyl jasmonate (MeJA) to the leaf blades of a non‐domesticated pasture grass (Microlaena stipoides) caused a more than two‐fold decrease in relative growth rate (RGR) of a root‐feeding chafer (Dermolepida albohirtum). MeJA treatment did not affect root consumption rates, but substantially reduced the efficiency of conversion of ingested food to body mass. 3. Non‐targeted metabolomics identified significant changes in the metabolome of MeJA‐induced plants, with three compounds (a galactolipid, a trihydroxy fatty acid and a lysophospholipid) found to be correlated with herbivore RGR, although their roles in herbivore defence remain uncertain. 4. This study suggests that an important Australian pasture grass can become better defended against root herbivores via enhanced jasmonate activity.     

Exotic herbivores and fire energy drive standing herbaceous biomass but do not alter compositional patterns in a semiarid savanna ecosystem

Questions

Fire regime alterations are pushing open ecosystems worldwide past tipping points where alternative steady states characterized by woody dominance prevail. This reduces the frequency and intensity of surface fires, further limiting their effectiveness for controlling cover of woody plants. In addition, grazing pressure (exotic or native grazers) can reinforce woody encroachment by potentially reducing fine-fuel loads. We investigated the effects of different fire energies on the herbaceous plant community, together with mammalian wildlife herbivory (exotic and native combined) exclusion, to inform best management practices.

Location

Texas semi-arid savanna, southern Great Plains, USA.

Methods

We conducted an experiment in which we manipulated fire intensity and herbivore access to herbaceous biomass in a split-plot design. We altered fire energy via fuel addition rather than applying fire under different environmental conditions to control for differences in standing biomass and composition attributable to differential plant physiological status and fire season.

Results

High-energy fire did not reduce herbaceous biomass or alter plant community composition, although it did increase among-plot variability in composition and forb biomass relative to low-energy fire and non-burned controls. Grazing pressure from native and non-native mammalian herbivores reduced above-ground herbaceous biomass regardless of fire treatments, but did not alter community composition.

Conclusions

Managers seeking to apply high-intensity prescribed fire to reduce woody encroachment will not negatively impact herbaceous plant productivity or alter community composition. However, they should be cognizant that repeated fires necessary for greatly reducing woody plants in heavily invaded areas might be difficult to accomplish due to fine-fuel reduction from wild herbivores. High fencing to restrict access by wildlife herbivores or culling might be necessary to build fuels sufficient to conduct high-intensity burns for woody-plant reduction.     

Response of Brassica oleracea to temporal variation in attack by two herbivores affects preference and performance of a third herbivore

1. Plants are frequently under attack by multiple insect herbivores, which may interact indirectly through herbivore‐induced changes in the plant's phenotype. The identity, order, and timing of herbivore arrivals may influence the outcome of interactions between two herbivores. How these aspects affect, in turn, subsequently arriving herbivores that feed on double herbivore‐induced plants has not been widely investigated. 2. This study tested whether the order and timing of arrival of two inducing herbivores from different feeding guilds affected the preference and performance of a subsequently arriving third herbivore, caterpillars of Mamestra brassicae L. (Lepidoptera: Noctuidae). Aphids [Brevicoryne brassicae L. (Hemiptera: Aphididae)] and caterpillars [Plutella xylostella L. (Lepidoptera: Yponomeutidae)] were introduced onto wild Brassica oleracea L. (Brassicaceae) plants in different sequences and with different arrival times. The effects of these plant treatments on M. brassicae caterpillars were assessed in pairwise preference tests and no‐choice performance tests. 3. The caterpillars of M. brassicae preferred to feed from undamaged plants rather than double herbivore‐induced plants. Compared with undamaged plants, they preferred plant material on which aphids had arrived first followed by caterpillars, whereas they avoided plant material with the reverse order of herbivore arrival. Performance of the caterpillars increased with increasing arrival time between herbivore infestations in double herbivore‐induced plants. Although M. brassicae grew faster on plants induced by aphids than on those induced by caterpillars alone, its performance was not affected by the order of previous herbivore arrival. 4. These results imply that the timing of colonisation by multiple herbivores determines the outcome of plant‐mediated herbivore–herbivore interactions.     

Behavioral archives link the chemistry and clonal structure of trembling aspen to the food choice of North American porcupine

Understanding the links among plant genotype, plant chemistry, and food selection by vertebrate herbivores is critical to assess the role of herbivores in the evolution of plant secondary chemistry. Some specialized vertebrate herbivores have been shown to select plants differentially according to plant genotype, but examples from generalists, which constitute the vast majority of vertebrate herbivores, are few, especially in natural conditions. We examined the relationship between the North American porcupine (Erethizon dorsatum), a generalist mammalian herbivore, and clonal trembling aspen (Populus tremuloides), a preferred food source of porcupines. We determined preference for certain aspen trees through visual examination of porcupine climbing scars left on tree bark, and through a controlled feeding experiment. We used genetic and biochemical analyses to link the behavioral archives (climbing scars) left by porcupines on aspen trunks to the clonal structure and chemical composition of trees. We show that two phenolic glycosides (tremulacin and salicortin), which are under a high degree of genetic control and thus vary in concentration across clones, are the chemical variables that most influence (deter) feeding choices by porcupines. Using behavioral archives left by a wild herbivore on a natural stand of plants thus allowed us to demonstrate that a generalist vertebrate herbivore can choose plants according to their clonal structure and genetically based chemical composition. Our results contribute to extending previous findings obtained with generalist herbivores studied in controlled conditions, and with specialist herbivores studied in the field. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

观赏灌木小枝和叶性状在林下庇荫环境中的权衡关系

小枝与叶片间的关系能够很好地反映植物对环境的适应策略。目前,从枝叶间权衡关系的角度研究树木对庇荫环境适应性的报道尚少,其中主要包括"生长-生长"权衡和"生长-生存"权衡两种假说。因此,为了探讨灌木在庇荫环境中,是否会在小枝和叶性状间采取权衡策略,来提高光照资源的利用能力或增强生存和防御能力,在城市绿地的林隙和林冠下两种光环境中,选择了没有人工修枝、整形等经营措施的金银木(Lonicera maackii)、小花溲疏(Deutzia parviflora)和连翘(Forsythia suspensa)等11种、79株观赏灌木作为研究对象,采集其当年生小枝,观测并计算了每棵灌木所处环境的有效光合辐射(PAR)相对累积光量和红光/远红光(R/FR),以及灌木的小枝干重(TDW)、单位小枝叶干重(LDW)、出叶强度(LN/TDW)、叶面积支持效率(LA/TDW)、比枝长(TL/TDW)和单叶面积(ILA) 6个性状。对数据进行标准化和数据转换后,运用Pearson相关检验分析光环境指标和灌木枝叶性状间的相关性;采用标准化主轴估计法进行异速生长方程的参数估计;使用多元回归分析不同光强下各性状的相互关系。结果表明:(1) PAR相对累积光量和R/FR与TDW和LDW呈极显著(P0.01或P0.001)正相关,与TL/TDW、LA/TDW、LN/TDW呈极显著(P0.001)负相关,与ILA相关性不显著(P 0.05)。整体上,R/FR与各灌木性状的相关性大于PAR相对累积光量;(2) TDW和LDW存在极显著(P0.001)的异速生长正相关关系。随着光强的减小,灌木当年生小枝和叶片的生物量都趋于减小,但是表现出相对偏向于叶片生物量的投资偏好;(3) LA/TDW和TL/TDW存在极显著(P0.001)的等速生长正相关关系。但随着光强的减小,比枝长随叶面积支持效率增加而增加的速率却减小,说明灌木在庇荫条件下,更倾向于采取忍耐型的光资源利用策略;(4) ILA和LN/TDW呈极显著(P0.001)异速生长负相关关系。随着光强的减小,灌木趋向于表现出单位小枝上着生大量小叶的现象。所以总体上,庇荫环境下的观赏灌木存在投资偏好和权衡,倾向于通过枝叶生长提高光截获能力来适应弱光环境,与"生长-生长"的权衡假说相符,但整体上,观赏灌木的耐阴性较差,故不建议种植在庇荫环境中。     

Interactions between large African browsers and thorny Acacia on a wildlife ranch in Kenya

Some plants respond to browsing with compensatory regrowth of plant tissues and with increased thorn growth. Associations between browsers and their preferred forage were examined through wandering quarter vegetation sampling and observational studies in an effort to understand how some plants respond to browsing by large African herbivores. Acacia seyal (n = 2680) A. drepanolobium (n = 1850), and Balanites glabra (n = 960) were three species of frequently browsed indigenous plants examined on Game Ranching Ltd. in Kenya. There were several statistically significant associations revealed. Individual A. seyal exposed to intensive browser utilization were observed to lose shoot tips, produce long thorns, and have relatively few flowers and fruits. Browser utilization was associated with increased lateral branching in A. drepanolobium and with an increased occurrence of short, thickened spines in B. glabra. Thorns, spines and flowers were measurable indicators of relative browser utilization, and may be useful features to monitor in the management of large African mammals and their prickly forage