Assessment of woody species encroachment in the grasslands of Nechisar National Park,Ethiopia

Ecological survey was executed to assess woody species encroachment into the grassland plain of Nechisar National Park (NNP). Forty‐one woody species were recorded. Dichrostachys cinerea Wight & Arn., Acacia mellifera (Vahl) Benth., Acacia nilotica (L) Willd., Acacia senegal (L.) Willd., Acacia seyal Del. and Acacia tortilis (Forssk.) Hayne were among the major encroaching woody species. The majority of the woody species were found to be highly aggregated in their pattern of distribution, while only few species showed some degree of randomness. The mean woody species density was ca. 1995 woody plants ha^?1. Mean cover of woody, grass, unpalatable forbs and total herbaceous species were 31%, 58%, 68% and 121%, respectively. The woody species density and cover, unpalatable forbs and bare land cover were significantly higher in the highly grazed and fire‐suppressed part of the grassland plain. Pearson correlation coefficient matrix indicated that woody species cover and density were negatively correlated with total herbaceous and grass cover. The high woody, unpalatable forbs and bare land cover indicated the progressively increasing perennial grass species diversity deterioration in the grass plain of the Park. Decline in the grassland condition, unless reversed, will jeopardize the biological diversity as well as the aesthetic value of the NNP.     

The role of phenolic compounds and other plant constituents in feeding by Canada geese in a coastal marsh

Summary We have assessed the relative importance of phenolic compounds, other secondary metabolites, and gross nutrient levels as feeding cues to Canada geese. Phenolic content was the most significant constituent influencing feeding selection by geese. Nutrient content had little or no effect on feeding selection.Correlative data showing the negative influence of plant phenolics on food choices by wild geese were supported by feeding preference tests. Extracts of unpalatable plants inhibited feeding by captive geese relative to extracts of palatable plants. In high phenolic plants, the phenolic containing methanol extract was more inhibitory than extractions made with petroleum ether. In a relatively low phenolic, unpalatable plant, an inhibitory factor was extractable in petroleum ether, indicating that for this species, another class of deterrents was involved. Preference tests with individual secondary metabolites showed that tannic acid and quebracho tannin were very effective in inhibiting feeding by geese and phenolic acids were slightly inhibitory, but a sesquiterpene lactone was not deterrent. These results point out the primacy of some secondary metabolites in determining food choices by geese.     

Phenological influences on the utilization of woody plants by eland in semi-arid shrubland

Our study monitored the acceptability, shoot phenology and chemical content of karoid woody species over the seasonal cycle. Species favoured by eland as long as they retained leaves, were classified as palatable, while those neglected for part of the seasonal cycle, even though they retained leaves, were classified as unpalatable. Changes in the acceptability of unpalatable species over the seasonal cycle appeared to be related to changes in the proportion of young shoots on plants, as influenced by rainfall and temperature. These species were typically only favoured when they offered relatively high proportions of young shoots. Our study identified total fibre as the most important chemical factor influencing the acceptability of woody species to eland. The shoots of palatable species contained low fibre concentrations, while the shoots of unpalatable species were more variable in fibre content. When favoured, the shoots of unpalatable species typically had similar levels of fibre to palatable species, but when neglected, the shoots of unpalatable species generally had increased fibre levels. Eland have thrived since being introduced into our study area, which is dominated by microphyllous and leptophyllous woody species. We suggest that this is because eland are able to select a diet sufficiently low in fibre content.     

Differences in construction costs and chemical composition between deciduous and evergreen woody species are small as compared to differences among families

We tested to what extent differences in construction costs (CC) and chemical composition of woody species are attributed to leaf habit. Eight evergreen and eight deciduous species belonging to six families were selected to form eight phylogenetic independent contrasts (PICs). The plants were grown from seed in a glasshouse. Differences in leaf, stem and root CC between evergreen and deciduous species were minor, the proportion of variance explained by leaf habit generally being less than 6%. Surprisingly, differences in leaf chemical composition between deciduous and evergreen species were small as well. Variation in CC and chemical composition among families was substantial, the factor 'family' explaining 50-85% of variance. We therefore conclude that in this case, phylogeny is a more important factor than functional group. Leaves of the fast-growing species in this experiment showed high levels of minerals, organic acids, proteins and lipids, whereas leaves of the slow-growing species had higher concentrations of soluble phenolics, lignin as well as higher carbon/nitrogen (C/N) ratio. These relationships suggest a trade-off between growth and defence. In contrast, CC of leaves, stems, roots or whole plants showed no or only a weak correlation with relative growth rate (RGR). The C/N ratio of the leaves is an easily measured parameter that correlated strongly in a negative way with the RGR of the plants and reflected better the balance between investment in structure and physiological functioning than CC.     

Chemical and morphological variation of Mediterranean woody evergreen species: Do plants respond to ungulate browsing?

Abstract. Plant defense theories suggest that chemical or structural defences should be maximized when and where browsing is most likely to occur. We tested this hypothesis on four evergreen woody species growing in a Mediterranean area with a high density of ungulates. In this system, levels of browsing are more intense in the winter (due to the lack of annual plants) and young foliage is often preferred. Therefore we predicted that the chemical defences of these species, namely their phenolic content, would vary with leaf age, season and damage intensity. In addition, we tested whether ungulates preferentially selected species containing lower phenolic levels, and also whether browsing induced either chemical or morphological changes in damaged plants. Phenolic levels varied greatly between plant species; ungulates browsed preferentially on the species with the lowest phenolic levels. No difference in phenolic content was found between browsed and unbrowsed trees. Morphological changes in heavily browsed trees included an increase in shoot and leaf density and a net decrease in leaf size. We suggest that for Mediterranean plants, which have evolved under high browsing pressure from large mammals, the production of small leaves and dense shoots in response to browsing might decrease ungulate foraging efficiency and hence reduce the rate of further damage as effectively as high levels of chemical defence.     

The effects of fire on woody plant encroachment are exacerbated by succession of trees of decreased palatability

The ingression of woody plants into the grassy layer of savannas and grasslands has become a global concern. The increase of woody plants has been primarily attributed over grazing, fire and more recently to the increase of atmospheric CO₂. We used long-term observations and analyses to assess changes in woody vegetation in Ithala Game Reserve (IGR), South Africa. Textural analysis of aerial photographs was used to detect changes in woody vegetation, from 1943 to 2007 in Ithala Game Reserve (IGR), South Africa. Daily rainfall data from 1905 to 2009 were used in a time-series analysis to determine if rainfall patterns have changed. The time-series analysis showed that the low magnitude (0–10 mm) rainfall events decreased from 1916 to 2009 and high magnitude rainfall events increased (10–20 and >20 mm). The mean annual rainfall increased from ～700 to ～850 mm from the 1930s to the 2000s. This change in rainfall was a key factor in the increase in woody vegetation from 1943 to 2009. We also used field data from the same reserve collected over 30 years to assess the increases in tree cover. Tree cover and density increased significantly by 32.5% and 657.9 indiv ha^?1 respectively, over 64 years. Before the proclamation of IGR in 1972, increases in woody vegetation from 1943 were non-significant. After the proclamation of IGR, herbivore population numbers and spatial distribution influenced the accumulation of grassy biomass required to fuel fires. In areas with reduced fuel loads, the consequential suppression of fire accelerated the rate of woody plant invasion into savannas. The increase in woody vegetation coincided with a decrease in palatable (e.g. Acacia gerrardii and Acacia davyi) and an increase in unpalatable woody plants. The avoidance of the unpalatable trees (e.g. Euclea and Searsia species) by large mammalian herbivores has allowed these trees to increase in density relatively unhindered.     

Secondary succession in Acacia nilotica (L.) savanna in the Hluhluwe Game Reserve,South Africa

Analysis of aerial photographs indicates that woody plant biomass has rapidly increased in Hluhluwe Game Reserve over the last 40 years. Open Acacia nilotica savanna is being replaced by broadleaf species, especially Euclea spp. We were interested in whether this secondary successional shift was due to high numbers of seedlings establishing and growing to maturity under acacias (facilitation) or due to the release of already established, but suppressed individuals (gullivers) of the resprouting broadleaf species. We examined the recruitment patterns and size-class distributions (height, basal diameter) of important species in this savanna.Densities of euclea seedlings (<0.6 cm basal diameter) under Acacia nilotica were low (median of 0 and mean of 0.06 m²) below adult canopies and effectively zero in adjacent interspaces. No differences in numbers of other broadleaf species were found between open sites and under Acacia nilotica sites. Few large eucleas or other broadleaf species occurred under Acacia nilotica. Few Acacia nilotica recruits were found either under adults or in the open whereas Acacia karroo recruits were more common.The overall size class distribution for eucleas was dominated by individuals in the intermediate size class, suggesting that recruitment is not the dominant demographic process. We propose that the escape of intermediate sized eucleas from the fire trap has caused the increase in woody plants.Analysis of a time sequence of aerial photographs shows that invasion occurred rapidly between 1954 and 1975. A common feature for sites where woody plant invasion has taken place, was the presence of barriers to fire (especially roads). We suggest that the recent and rapid increase in woody vegetation is due to a decrease in the frequency of intense fires, rather than the recent absence of megaherbivores which allowed Acacia nilotica establishment. Few intense fires allow suppressed tree or shrub individuals, ('gullivers'), to escape the grass/fire layer and thereby become tall and fire-resistant. This release may explain the rapid rate of invasion by inherently slow growing broad-leaf species.     

Comparison of plant secondary metabolites and digestibility of three different boreal coniferous trees

Moose (Alces alces L.) prefer to browse Pinus sylvestris over other coniferous trees. One reason for this preference might be due to differences in the chemical composition of various coniferous plants, and/or differences in digestibility. We evaluated the in-sacco digestibility of P. sylvestris, Pinus contorta and Picea abies using a fistulated cow (Bos taurus). Additionally, we determined the chemical profile of the different plant species (with emphasis on specific phenolics) and we modified the concentration of the individual phenolic (+)catechin in plant material of P. sylvestris to test the influence of this particular phenolic on digestibility in vitro (IVDMD). The in-sacco experiment revealed the highest digestibility of dry matter, and neutral detergent fibre for P. sylvestris. In addition, the concentration of nitrogen was significantly higher, and the concentrations of total phenolics and condensed tannins were significantly lower in P. sylvestris than in the other species. Individual phenolics showed high variation but were low in P. sylvestris, whereas P. contorta and P. abies had at least one species-specific phenolic compound of high concentration each. We found no changes in the digestibility of samples in which we experimentally elevated (+)catechin content.     

Chemical defenses and the susceptibility of tropical marine brown algae to herbivores

Summary I assayed phenolic and tannin concentrations in a number of species of temperate and tropical brown algae of the genera Sargassum and Turbinaria. Tropical species in both genera contained consistently low levels of phenolics and tannins (species means ranged between 0 and 1.6% [measured as % dry weight of the thallus]). Levels of phenolics in temperate species of Sargassum were variable and consistently much higher than in tropical species (species means ranged between 3 and 12% by dry weight). This pattern of latitudinal variation in phenolic levels in Sargassum conflicts with previous predictions for latitudinal variation in the chemical defenses of marine organisms. The low levels of phenolics present in the tropical species that I analyzed may also explain recent results (Hay 1984; Lewis 1985) demonstrating that tropical Sargassum and Turbinaria are often preferentially consumed by herbivorous fishes and echinoids.     

Foliage acceptability to browsing ruminants in relation to seasonal changes in the leaf chemistry of woody plants in a South African savanna

Summary We investigated seasonal changes in food selection by hand-reared kudus and impalas in savanna vegetation in northern Transvaal, South Africa. The acceptability of the leaves of woody plants to these animals was compared with leaf concentrations of nutrients, fibre components and old leaf phenophases. No consistently significant correlation was found between acceptability and any single chemical factor. Based on an a priori palatability classification, discriminant function analysis separated relatively palatable species from unpalatable species in terms of a linear combination of protein and condensed tannin concentrations. The high acceptability of certain otherwise unpalatable species during the new leaf phenophase was related to elevation of protein levels relative to condensed tannin contents. Species were added to the diet during the dry season approximately in the order of their relative protein-condensed tannin difference.     

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。     

Impacts of a mineral lick‐centred land use system on woody vegetation cover in an East African Savannah

Evaluation of woody vegetation changes with distance from a salt crater was conducted in the semi‐arid rangelands of southern Ethiopia. Data on live woody plants were collected over three seasons at 0, 1, 4, 6, 9 and 12 km from the salt crater. The density and diversity of woody plants differed significantly (p < .01) along the distance gradient. Six woody plant families were identified of which Fabaceae and Burseraceae were the dominant families. Acacia drepanolobium, Acacia nilotica, Commiphora africana and Acacia mellifera were among the severely encroaching woody species. There were high proportions of seedlings and saplings recorded closer to the salt crater showing a vigorous recruitment by woody plants. Woody plant encroachment along the 12‐km transect ranged from a low to severe encroachment, which could be translated into poor rangeland condition. Changes in soil characteristics increased grazing pressure and sedentary settlement around the salt crater, and the breakdown of traditional institutions seems to be major contributing factors to these vegetation changes. We suggest that severely encroached areas could be improved through a combination of methods such as bush clearing, prescribed fire, browsing animals and proper grazing management.     

Secondary metabolites and nutrients of woody plants in relation to browsing intensity in African savannas

Carbon-based secondary metabolites (CBSMs) are assumed to function as defences that contribute to herbivore-avoidance strategies of woody plants. Severe browsing has been reported to reduce concentrations of CBSMs and increase N concentrations in individual plants, causing heavily browsed plants to be characterised by N-rich/C-poor tissues. We hypothesised that concentrations of condensed tannins (CT) and total polyphenols (TP) should decrease, or N increase, in relation to increasing intensity of browsing, rendering severely browsed plants potentially more palatable (increased N:CT) and less N-limited (increased N:P) than lightly browsed ones. We sampled naturally browsed trees (taller than 2 m) of four abundant species in southern Kruger National Park, South Africa. Species-specific relationships between N:CT, CT, TP and P concentrations and increasing browsing intensity were detected, but N and N:P were consistently invariable. We developed a conceptual post-hoc model to explain diverse species-specific CBSM responses on the basis of relative allocation of C to total C-based defence traits (e.g. spines/thorns, tough/evergreen leaves, phenolic compounds). The model suggests that species with low allocation of C to C-based defence traits become C-limited (potentially more palatable) at higher browsing intensity than species with high allocation of C to C-based defences. The model also suggests that when N availability is high, plants become C-limited at higher browsing intensity than when N availability is low.     

Leafing patterns and leaf traits of four evergreen shrubs in the Patagonian Monte, Argentina

We assessed leafing patterns (rate, timing, and duration of leafing) and leaf traits (leaf longevity, leaf mass per area and leaf-chemistry) in four co-occurring evergreen shrubs of the genus Larrea and Chuquiraga (each having two species) in the arid Patagonian Monte of Argentina. We asked whether species with leaves well-defended against water shortage (high LMA, leaf longevity, and lignin concentration, and low N concentration) have lower leaf production, duration of the leafing period, and inter-annual variation of leafing than species with the opposite traits. We observed two distinctive leafing patterns each related to one genus. Chuquiraga species produced new leaves concentrated in a massive short leafing event (5–48 days) while new leaves of Larrea species emerged gradually (128–258 days). Observed leafing patterns were consistent with simultaneous and successive leafing types previously described for woody plants. The peak of leaf production occurred earlier in Chuquiraga species (mid September) than in Larrea species (mid October–late November). Moreover, Chuquiraga species displayed leaves with the longest leaf lifespan, while leaves of Larrea species had the lowest LMA and the highest N and soluble phenolics concentrations. We also observed that only the leaf production of Larrea species increased in humid years. We concluded that co-occurring evergreen species in the Patagonian Monte displayed different leafing patterns, which were associated with some relevant leaf traits acting as plant defenses against water stress and herbivores. Differences in leafing patterns could provide evidence of ecological differentiation among coexisting species of the same life form.     

重庆石灰岩地区主要木本植物叶片性状及养分再吸收特征

以重庆石灰岩地区15种常绿木本植物和14种落叶木本植物为研究对象,对两种生活型植物叶片衰老前后叶干物质含量(LDMC)、比叶面积(SLA)和叶片厚度(LT)进行了比较,并采用不同的计算方法(单位质量叶片养分含量、单位面积叶片养分含量)分析了两类植物叶片衰老前后养分含量及再吸收特征,最后对养分再吸收效率与其他叶性状因子之间的关系进行了相关分析。结果表明:常绿植物成熟叶LDMC、LT及衰老叶LT显著低于落叶植物,落叶植物成熟叶和衰老叶SLA均显著高于常绿植物(P0.05);基于单位质量叶片计算的养分含量,常绿植物成熟和衰老叶N、P量均低于落叶植物,而基于单位面积叶片计算的N、P含量则表现出相反的趋势;基于不同方法计算的N、P再吸收效率差异不明显,其中常绿植物基于单位质量叶片养分含量计算的N、P平均再吸收效率为39.42%、43.79%,落叶植物的为24.08%、33.59%;常绿和落叶植物N、P再吸收效率与LDMC、SLA、LT和成熟叶N、P含量之间没有显著相关性,但与衰老叶养分含量存在显著负相关(P0.05)。研究发现,无论是常绿植物还是落叶植物,衰老叶N、P含量均较低,表明石灰岩地区植物具有较高的养分再吸收程度。     

Tree species diversity alters plant defense investment in an experimental forest plantation in southern Mexico

How plant species diversity affects traits conferring herbivore resistance (e.g., chemical defenses), as well as the mechanisms underlying such effects, has received little attention. One potential mechanism for the effect of diversity on plant defenses is that increased plant growth at high diversity could lead to reduced investment in defenses via growth–defense trade‐offs. We measured tree growth (diameter at breast height) and collected leaves to quantify total phenolics in 2.5‐year‐old plants of six tropical tree species (N = 597 plants) in a young experimental plantation in southern Mexico. Selected plants were classified as monocultures or as polycultures represented by mixtures of four of the six species examined. Tree species diversity had a significant negative effect on total phenolics, where polycultures exhibited a 13 percent lower mean concentration than monocultures. However, there was marked variation in the effects of diversity on defenses among tree species, with some species exhibiting strong reductions in phenolic levels in mixtures, whereas others were unresponsive. In addition, tree species diversity had no effect on growth, nor was the negative effect of diversity on chemical defenses mediated by a growth–defense trade‐off. These results demonstrate that tree diversity can alter investment in chemical defenses in long‐lived tree species but that such effect may not always be under strong control by plant endogenous resource allocation trade‐offs. Regardless of the underlying mechanism, these findings have important implications for predicting effects on consumers and ecosystem function.     

海南岛霸王岭热带季雨林树木的死亡率

树木死亡是调节群落组成和结构的一种方式, 在森林生态系统动态中发挥着重要作用。该文在对海南岛霸王岭热带季雨林群落大量调查和树种功能群划分研究的基础上, 探索了热带季雨林群落及其不同功能群树木的死亡率及其随径级和环境条件的变化规律。结果表明: 海南岛霸王岭热带季雨林群落内树木死亡率的变化范围为3.42%-18.71%, 平均值为7.60%。按照功能群比较, 乔木死亡率显著高于灌木, 落叶树种死亡率高于常绿树种, 但具刺树种死亡率显著低于无刺树种。按照树木径级划分, 5-30 cm径级范围的死亡率均超过10%, 最高值出现在5-10 cm径级范围, 在相同径级范围内比较时, 落叶树种在 胸径(DBH ) < 5 cm时的死亡率显著高于常绿树种, 而在其他径级范围内, 二者之间无显著差异, 具刺和无刺树种的死亡率在所有径级范围内均无显著差异。乔木死亡率与群落距河流距离具有显著的相关性, 距离河流越远死亡率越高; 落叶树种和无刺树种则与坡位存在显著的相关性, 坡上部的死亡率显著高于坡中部。热带季雨林内树木的死亡率与其所处的群落生境有关, 较干旱的生境是导致树木死亡率较高的重要原因。     

Testing adaptive plasticity to UV: costs and benefits of stem elongation and light-induced phenolics

On exposure to ultraviolet radiation (UV), many plant species both reduce stem elongation and increase production of phenolic compounds that absorb in the UV region of the spectrum. To demonstrate that such developmental plasticity to UV is adaptive, it is necessary to show that the induced phenotype is both beneficial in inductive environments and maladaptive in non-inductive environments. We measured selection on stem elongation and phenolic content of seedlings of Impatiens capensis transplanted into ambient-UV and UV-removal treatments. We extended the range of phenotypes expressed, and thus the opportunity for selection in each UV treatment, by pretreating seedlings with either a low ratio of red:far-red wavelengths (R:FR), which induced stem elongation and reduced phenolic concentrations, or high R:FR, which had the opposite effect on these two phenotypic traits. Reduced stem length relative to biomass was advantageous for elongated plants under ambient UV, whereas increased elongation was favored in the UV-removal treatment. Selection favored an increase in the level of phenolics induced by UV in the ambient-UV treatment, but a decrease in phenolics in the absence of UV. These results are consistent with the hypotheses that reduced elongation and increased phenolic concentrations serve a UV-protective function and provide the first explicit demonstration in a wild species that plasticity of these traits to UV is adaptive. The observed cost to phenolics in the absence of UV may explain why many species plastically upregulate phenolic production when exposed to UV, rather than evolve constitutively high levels of these compounds. Finally, pretreatment with low R:FR simulating foliar shade did not exacerbate the fitness impact of UV exposure when plants had several weeks to acclimate to UV. This observation suggests that the evolution of adaptive shade avoidance responses to low R:FR in crowded stands will not be constrained by increased sensitivity to UV in elongated plants when they overtop their neighbors.     

Seasonality and edge effect determine herbivory risk according to different plant association models

We report evidence of hierarchical resource selection by large herbivores and plant neighbouring effects in a Mediterranean ecosystem. Plant palatability was assessed according to herbivore foraging decisions. We hypothesize that under natural conditions large herbivores follow a hierarchical foraging pattern, starting at the landscape scale, and then selecting patches and individual plants. A between- and within-patch selection study was carried out in an area formed by scrubland and pasture patches, connected by habitat edges. With regard to between-patch selection, quality-dependent resource selection is reported: herbivores mainly consume pasture in spring and woody plants in winter. Within-patch selection was also observed in scrub habitats, influenced by season, relative patch palatability and edge effect. We defined a Proximity Index (PI) between palatable and unpalatable plants, which allowed verification of neighbouring effects. In spring, when the preferred food resource (i.e. herbs) is abundant, we observed that in habitat edges large herbivores basically select the relatively scarce palatable shrubs, whereas inside scrubland, unpalatable shrub consumption was related to increasing PI. In winter, a very different picture was observed; there was low consumption of palatable species surrounded by unpalatable species in habitat edges, where the latter were more abundant. These outcomes could be explained though different plant associations described in the literature. We conclude that optimal foraging theory provides a conceptual framework behind the observed interactions between plants and large herbivores in Mediterranean ecosystems.     

Seasonal changes in leaf chemistry and leaf selection of the Japanese giant flying squirrel upon two tree species

Tree leaves are important food sources for arboreal herbivores, such as primates, rodents, and marsupials. These animals do not eat leaves randomly in habitats with many tree species but rather choose based on the chemical components of leaves, such as sugars, fibers, proteins, and toxins. However, the effects of the microscale distribution of these chemicals within each leaf have not been examined for these animals. The giant flying squirrels Petaurista leucogenys are entirely arboreal, nocturnal herbivores, usually feeding on leaves and dropping leaf debris on the ground after partially consuming them. Therefore, we could easily assess which species of trees and which parts of the individual leaves they preferred to eat. We also examined microscale distributions of phenolics, sugar, and water within individual leaves. Of the two dominant food tree species, the deciduous Quercus acutissima was preferred over the evergreen Q. sessilifolia. The latter tree is only used during winter to early spring when the former had no leaves. Our chemical analyses revealed that Q. acutissima contained much more glucose than Q. sessilifolia in all seasons. Three types of leaf debris, eaten apically, basally, or centrally with a hole, were found. In Q. sessilifolia, which had low phenolic concentrations, apical eating was most common, whereas central eating was rare. In Q. acutissima, which had high phenolics, basal or central eating was common. Central feeding may be caused by avoiding the periphery because of a higher phenolic concentration in the leaf margin. Thus, microscale distributions of phenolics within individual leaves affect which parts P. leucogenys eats, whereas leaf sugar concentration is an important factor affecting which species of leaves they eat.