Folivory with leaf folding by giant flying squirrels: its patterns and possible function

Japanese giant flying squirrels, Petaurista leucogenys, are entirely arboreal folivores. From spring to summer, their principal food source consists of leaves of the deciduous oak Quercus acutissima in western Tokyo, Japan. Before consuming the central part of each leaf, the squirrels fold the leaf two or more times with their forelegs. In the wild, leaf folding is a very rare behavior, even in primates. In our study, this peculiar feeding behavior was not observed across all study sites or even in some local populations. Herbivores generally try to maximize their intake of nutritious foods (e.g., sugars) containing low levels of plant defensive chemicals (e.g., phenols). We found that total phenolic contents in individual leaves of Q. acutissima were lower in the center than at the margins, whereas sugar (glucose) was homogenously distributed. Consequently, consumption of only leaf centers appears to be an adaptation by Japanese giant flying squirrels to avoid leaf margin defensive chemicals and structures. These leaf margin characteristics, which may have developed as defenses against herbivorous insects that generally feed from the margins, have contributed to the skillful and complex feeding behavior of this generalist folivore.     

麻栎和闽楠幼苗叶功能性状及生物量对光照和施肥的响应

光照和养分条件是影响植物生长的重要环境因子,不同生活型植物对环境异质性的响应机制不同。以落叶阔叶树种麻栎和常绿阔叶树种闽楠幼苗为研究对象,设置2个光照梯度（全光照和45%全光照）和4个施肥梯度（不施肥、氮磷供应比为5、15和45）共8种处理,研究光照和施肥及其交互作用对麻栎和闽楠生物量和叶形态、生理及化学性状的影响,并探讨了叶功能性状和生物量的关系。结果表明：（1）光照、施肥及其交互作用对光合气体交换参数（除水分利用效率外）、叶绿素荧光参数、叶形态指标（除比叶面积外）、单位质量叶氮含量和根冠比影响显著（P<0.05）。此外,光照和施肥对地上生物量和总生物量影响显著（P<0.05）。（2）全光照显著增加了麻栎和闽楠单株总叶面积和地上、地下生物量及总生物量（P<0.05）,而遮荫降低了非光化学猝灭系数、光合氮利用率和根冠比,增加了单位质量叶氮含量。（3）在全光照处理中,施肥显著增加了麻栎和闽楠水分利用效率（P<0.05）;在遮荫处理中,氮磷供应比45显著增加了麻栎和闽楠净光合速率和水分利用效率（P<0.05）。（4）麻栎和闽楠在全光照中倾向于资源获取策略,在遮荫中偏向于资源保守策略。在光照和施肥处理中,麻栎和闽楠单株总叶面积与地上生物量均显著正相关（P<0.05）。总之,单株总叶面积是预测麻栎和闽楠幼苗地上生物量变化的稳定指标,施肥有助于增加低光环境下麻栎和闽楠幼苗的生态适应能力。     

Relationships between leaf age and the food quality of cottonwood foliage for the gypsy moth,Lymantria dispar

Summary The cottonwood tree, Populus deltoides, continues to produce leaves late into the growing season, exposing midseason herbivores to leaves of a wide range of maturity. Gypsy moth larvae preferred and grew best on the oldest cottonwood leaves and suffered higher mortality and 85% less growth when fed young, expanding leaves. Concentration of phenolics in the youngest leaves was 3 times that in the oldest leaves and was negatively correlated with caterpillar growth rate. The active phenolics were not identified; tannin was present but its concentration changed more with season than leaf age.     

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.     

Strong influence of leaf tie formation and corresponding weak effect of leaf quality on herbivory in eight species of Quercus

1. Leaf shelter construction by herbivorous insects can improve leaf quality, sometimes changing resultant herbivory. In two experiments in a Missouri (USA) deciduous forest we quantified the impact of leaf tie construction and changes to leaf quality on subsequent leaf damage.
2. First, using eight Quercus species, we compared damage to single leaves versus experimental leaf ties that had been stocked with either Pseudotelphusa quercinigracella (Gelechiidae) or Psilocorsis cryptolechiella (Depressariidae) to determine how initial leaf quality (total phenolics) influenced damage caused by shelter inhabitants. Skeletonization by leaf tying caterpillars and leaf edge chewing by free feeding species were 12.2× and 1.3× greater on tied than on non-tied leaves, respectively. July and September leaf phenolic content had a slight positive effect on the probability of skeletonization, none on the probability of edge damage, and a weakly positive or negative effect on the intensity of skeletonization and edge damage, depending on leaf position.
3. Second, we created experimental leaf ties, protected from herbivores, on the same Quercus species to determine whether tie formation changes leaf quality (total phenolics, nitrogen, water, toughness). Tie formation decreased phenolics, but this change was predicted to add only 0.8% leaf area loss.
4. Synthesis. Herbivory increased dramatically when leaves were in ties, with the effect mostly due to the tie itself rather than a change in leaf quality. We predict that the advantages of building and using leaf ties in this system are more likely to be escape from natural enemies and changes in abiotic factors.

     

Comparison of Herbivores and Herbivory in the Canopy and Understory for Two Tropical Tree Species1

The Janzen–Connell model of tropical forest tree diversity predicts that seedlings and young trees growing close to conspecific adults should experience higher levels of damage and mortality from herbivorous insects, with the adult trees acting as either an attractant or source of the herbivores. Previous research in a seasonal forest showed that this pattern of distance‐dependent herbivory occurred in the early wet season during the peak of new leaf production. I hypothesized that distance‐dependent herbivory may occur at this time because the new foliage in the canopy attracts high numbers of herbivores that are limited to feeding on young leaves. As a consequence, seedlings and saplings growing close to these adults are more likely to be discovered and damaged by these herbivores. In the late wet season, when there is little leaf production in the canopy, leaf damage is spread more evenly throughout the forest and distance dependence disappears. I tested three predictions based on this hypothesis: (1) the same species of insect herbivores attack young and adult trees of a given plant species; (2) herbivore densities increase on adult trees during leaf production; and (3) herbivore densities in the understory rise during the course of the wet season. Censuses were conducted on adults and saplings of two tree species, raribea asterolepis and Alseis blackiana. Adults and saplings of both species had largely the same suite of chewing herbivore species. On adults of Q. asterolepis, the density of chewing herbivores increased 6–10 times during leaf production, but there was no increase in herbivore density on adults of A. blackiana. Herbivore densities increased 4.5 times on A. blackiana saplings and 8.9 times on Q. asterolepis saplings during the wet season, but there were no clear trends on the adults of either species. These results suggest that the potential of adult trees as a source of herbivores on saplings depends on the value of new leaves to a tree species' herbivores, which may differ across tree species.     

Leaf morphology and leaf chemical composition in three Quercus (Fagaceae) species along a rainfall gradient in NE Spain

 Leaf features were examined in three Quercus species (Q. coccifera, Q. ilex and Q. faginea) along a steep rainfall gradient in NE Spain. The analyzed leaf traits were area, thickness, density, specific mass, leaf concentration of nitrogen, phosphorous, lignin, cellulose and hemicellulose, both on a dry weight basis (N_w, P_w, L_w, C_w, H_w) and on an area basis (N_a, P_a, L_a, C_a, H_a). These traits were regressed against annual precipitation and correlated with each other, revealing different response patterns in the three species. Q. faginea, a deciduous tree, did not show any significant correlation with rainfall. In Q. coccifera, an evergreen shrub, N_w, N_a, L_w, L_a and C_a increased with higher annual rainfall, while H_w decreased. In Q. ilex, an evergreen tree, leaf area, P_w and L_w increased with precipitation, whereas specific leaf mass, thickness and H_a showed the reverse response. Correlations between the leaf features revealed that specific mass variation in Q. faginea and Q. coccifera could be explained by changes in leaf density, while in Q. ilex specific leaf mass was correlated with thickness. Specific leaf mass in the three species appeared positively correlated with all the chemical components on a leaf area basis except with lignin in Q. ilex and with P in Q. ilex and Q. faginea. In these two tree species P_w showed a negative correlation with specific leaf mass. It is suggested that each species has a different mechanism to cope with water shortage which is to a great extent related to its structure as a whole, and to its habit. Received: 18 December 1995 / Accepted: 8 March 1996     

Strategies of leaf expansion in Ficus carica under semiarid conditions

Leaf area expansion, thickness and inclination, gas exchange parameters and relative chlorophyll content were analysed in field‐grown fig (Ficus carica L.) leaves over time, from emergence until after full leaf expansion (FLE). Ficus carica leaves showed a subtle change in shape during the early stages of development, and FLE was reached within ca. 30 days after emergence. Changes in leaf thickness and inclination after FLE demonstrated good adaptation to environmental conditions during summer in areas with a Mediterranean climate. Changes in gas exchange parameters and relative chlorophyll content showed that F. carica is a delayed‐greening species, reaching maximum values 20 days after FLE. Correlation analysis of datasets collected during leaf expansion, confirmed dependence among structural and functional traits in F. carica. Pn was directly correlated with stomatal conductance (Gs), transpiration (E), leaf area (LA) and relative chlorophyll content up to FLE. The effect of pruning on leaf expansion, a cultural technique commonly applied in this fruit tree, was also evaluated. Although leaf development in pruned branches gave a significantly higher relative leaf area growth rate (RGR_l) and higher LA than non‐pruned branches, no significant differences were found in other morphological and physiological traits, indicating no pruning effect on leaf development. All studied morphological and physiological characteristics indicate that F. carica is well adapted to semiarid conditions. The delayed greening strategy of this species is discussed.     

桥山栎林群落结构特征与物种多样性相关关系分析

为了解森林群落结构特征与物种多样性之间的相关关系,以黄土高原桥山林区典型麻栎纯林、麻栎阔叶混交林和麻栎油松混交林为研究对象,调查分析了群落结构特征,计算物种重要值及物种多样性,并进行了冗余分析,结果表明:(1)麻栎油松混交林主要以乔木层胸径、树高、枝下高等最高;麻栎阔叶混交林以灌木层盖度、地径、冠幅、高度等最高;麻栎纯林以草本地径、草本盖度及草本冠幅最高。(2) 3种类型林分乔木层重要值最高的均为麻栎(Quercus acutissima),灌木层为狼牙刺(Sophora viciifolia)、南蛇藤(Celastrus orbiculatus)、胡枝子(Lespedeza bicolor),草本层为苔草(Carex tristachya)。(3)麻栎油松混交林乔、灌层物种多样性较高,麻栎阔叶混交林以草本层物种多样性最高。(4)不同类型麻栎林群落结构特征与物种多样性关系有差异。松栎混交林中,对物种多样性影响最大的为乔木胸径、新稍长及灌木高度;麻栎阔叶混交林对物种多样性影响最大的为灌木层高度及冠幅;麻栎纯林中,对物种多样性影响最大的为乔木胸径。(5)麻栎油松混交林、麻栎阔叶混交林的胸径、树高、物种多样较麻栎纯林高,具有较高木材生产能力和生态防护功能,是未来森林经营培育的方向。     

Perfect is best: low leaf fluctuating asymmetry reduces herbivory by leaf miners

Fluctuating asymmetry (FA) represents small, random variation from symmetry and can be used as an indicator of plant susceptibility to herbivory. We investigated the effects of FA of two oak species, Quercus laevis and Q. geminata, and the responses of three herbivore guilds: leaf miners, gallers, and chewers. To examine differences in FA and herbivory between individuals, 40 leaves from each tree were collected, and FA indices were calculated. To examine differences in FA and herbivory within-individuals, we sampled pairs of mined and unmined leaves for asymmetry measurements. Differences in growth of leaf miners between leaf types were determined by tracing 50 mines of each species on symmetric leaves and asymmetric leaves. Asymmetric leaves contained significantly lower concentrations of tannins and higher concentrations of nitrogen than symmetric leaves for both plant species. Both frequency of asymmetric leaves on plants and levels of asymmetry positively influenced the abundance of Brachys, Stilbosis and other leaf miners, but no significant relationship between asymmetry and herbivory was observed for Acrocercops. Brachys and Stilbosis mines were smaller on asymmetric leaves, but differences in mine survivorship between symmetric and asymmetric leaves were observed only for Stilbosis mines. This study indicated that leaf miners might use leaf FA as a cue to plant quality, although differential survivorship among leaf types was not observed for all species studied. Reasons for the different results between guilds are discussed.     

Use of leaf resources by howling monkeys (Alouatta palliata) and leaf-cutting ants (Atta cephalotes) in the tropical rain forest of Los Tuxtlas,Mexico

Use of leaf resources by a troop of howling monkeys and two colonies of leaf cutting ants was studied for an annual cycle in the rain forest of Los Tuxtlas, Mexico. Howling monkeys spent half their annual foraging time feeding on leaves; leaf-cutting ants spent at least 80% of their recorded foraging time harvesting leaves. Both herbivores preferred young leaves over nature ones, and chemical analysis showed that the protein: fibre ratio of the leaves used was correlated with these preferences. Howling monkeys used 34 tree species as leaf sources. Leaf-cutting ants used 40 plant species of which 38 were trees. Eighteen species used by Alouatta were also used by Atta; species of Moraceae and Lauraceae were among the most important in their foraging preferences. The plant species used by monkeys and ants occurred at low densities (? 4.0 ind/ha). The seasonal production of leaves, the high density of leaf-cutting ant colonies at the study site, and the high amounts of young foliage harvested by the ants from tree species, and individual trees used by howling monkeys as sources of young leaves suggest that the foraging activities of Atta may represent a significant pressure upon leaf resources available to Alouatta.     

Outdoor infrared imaging for spatial and temporal thermography: A case study of necrotic versus healthy leaf areas on woody plants

The temperature of plants can be measured using infrared (IR) thermography. Despite the extensive use of IR imaging indoors, outdoor IR imaging is uncommon. We used IR imaging to compare leaf temperatures between necrotic spots and healthy areas of oriental cherry (Prunus serrulata var. spontanea), Japanese cornel (Cornus officinalis) and sawtooth oak (Quercus acutissima) in the field. There was a significant difference in the mean leaf temperatures between the necrotic spots (26.4°C) and healthy areas (25.6°C) of oriental cherry (p = .01). The mean temperatures in the necrotic spots of Japanese cornel and sawtooth oak leaves were 22.3°C and 29.6°C, respectively, which were not significantly different from the mean temperatures of the healthy areas. A consecutive, 2‐day temporal leaf analysis in October 2018 revealed that the temperatures in the necrotic spots were generally higher than those in the healthy areas of the three species. The temperature difference between the spots and healthy areas (up to 1.4°C) was more pronounced at 13:00 hr in all three species on both days. These results reveal differences in the spatial and temporal thermal state across the necrotic spotted leaves. There is potential for use of outdoor IR imaging to visualize the response of trees to pathogen infection and abiotic stress.     

Plastic leaf morphology in three species of Quercus: The more exposed leaves are smaller,more lobated and denser

Phenotypic plasticity and developmental instability in leaf traits are common in oak species but the role of environmental factors is not well understood. To decipher possible correlations between different leaf traits and effects of the position of leaves within the tree canopy, we quantified the plasticity of three leaf traits of 30 trees of Quercus alba L., Quercus palustris Muench and Quercus velutina Lam. We hypothesized that trees could modify the shape of their leaves for better adaptation to the variable microclimate within the canopy. Our results demonstrated that the south and north outer leaves were significantly smaller, more lobed and denser than those situated in the inner canopy. The order of leaves on the branch accounted for the plasticity of leaf traits in Q. alba only. Plasticity of lobing in Q. alba and Q. velutina depended on the height of the trees. We detected fluctuating asymmetry (FA) in all three species, but the source of variation depended on branch position in Q. velutina only. FA was more pronounced in north-facing leaves. Plasticity of the leaf traits ranged from small to medium. Plasticity of leaf area and leaf mass per area (LMA) depended on the branch position. However, the plasticity of lobation was not affected by the location of a branch within the tree canopy. Quercus alba and Q. palustris had similar plastic responses but the plasticity of Q. velutina was significantly smaller. We concluded that individual plants detect and cope with environmental stress through vegetative organ modification.     

Effect of leaf litter characteristics on leaf conditioning and on consumption by Gammarus pulex

相似文献   

Leaf physiology does not predict leaf habit; examples from tropical dry forest

Leaf structure and physiology are thought to be closely linked to leaf longevity and leaf habit. Here we compare the seasonal variation in leaf hydraulic conductance (k_leaf) and water potential of two evergreen tree species with contrasting leaf life spans, and two species with similar leaf longevity but contrasting leaf habit, one being deciduous and the other evergreen. One of the evergreen species, Simarouba glauca, produced relatively short-lived leaves that maintained high hydraulic conductance year round by periodic flushing. The other evergreen species, Quercus oleoides, produced longer-lived leaves with lower k_leaf and as a result minimum leaf water potential was much lower than in S. glauca (–2.8 MPa vs –1.6 MPa). Associated with exposure to lower water potentials, Q. oleoides leaves were harder, had a higher modulus of elasticity, and were less vulnerable to cavitation than S. glauca leaves. Both species operate at water potentials capable of inducing 20 (S. glauca) to 50% (Q. oleoides) loss of k_leaf during the dry season although no evidence of cumulative losses in k_leaf were observed in either species suggesting regular repair of embolisms. Leaf longevity in the deciduous species Rhedera trinervis is similar to that of S. glauca, although maximum k_leaf was lower. Furthermore, a decline in leaf water potential at the onset of the dry season led to cumulative losses in k_leaf in R. trinervis that culminated in leaf shedding.     

Evaluating community effects of a Keystone Ant,Azteca sericeasur,on Inga micheliana leaf litter decomposition in a shaded coffee agro-ecosystem

Our research examined the effect of Azteca sericeasur, a keystone arboreal ant, on the decomposition of leaf litter of the shade tree, Inga micheliana, in coffee agro-ecosystems. This interaction is important in understanding spatial heterogeneity in decomposition. We hypothesized that A. sericeasur could affect leaf litter decomposition by excluding other ants, which could release decomposers, like collembolans, from predation pressure. Determining the relative strengths of these interactions can illuminate the importance of A. sericeasur in decomposition and nutrient cycling processes. We assessed the ant and arthropod communities surrounding 10 pairs of trees, where each pair included one shade tree with an established A. sericeasur nest. Tuna baits were used in conjunction with pitfall traps to assess the ant and arthropod community, and litterbags with I. micheliana leaf litter were used to assess rates of decomposition. The species richness of ants did not change in proximity to A. sericeasur nests, though the ant communities were distinct. Abundance of Collembola and community composition of other invertebrates did not change in the presence of A. sericeasur nests, and there were no differences in leaf litter decomposition rates. This contradicts past studies that suggest A. sericeasur reduces ant species richness in its territory. We suggest that other ants may avoid A. sericeasur by moving within and beneath the leaf litter. Our results indicate that there is no net effect of A. sericeasur on leaf litter decomposition.     

Influence of leaf traits on the spatial distribution of arboreal arthropods within an overstorey rainforest tree

Abstract.

• 1 Several attributes of foliage were measured from the Australian rainforest tree Argyrodendron actinophyllum Edlin (Sterculiaceae). These were related to estimates of abundance per leaf area of the most common arthropod guilds and families sampled with restricted canopy fogging.
• 2 When all these arthropod groups were considered, much of the overall variance in arthropod spatial distribution could be attributed to leaf age characteristics, arthropod aggregation patterns, arthropod activity and distance to tree trunk.
• 3 The fraction of variance which could be specifically explained by foliage attributes such as nitrogen-, water- and fibre-content, specific leaf weight, and epiphyll load was small for most arthropod groups (usually <30%). However, an index of food quality explained a higher proportion of variance (50%) in the abundance of phloem-feeders. Leaf size and foliage compactness did not influence significantly the abundance of any arthropod group.
• 4 Most herbivores were more abundant on young foliage than on mature leaves. With the exception of Corylophidae and Chrysomelidae, which were more abundant in the lower and upper canopy respectively, arthropod stratification was not conspicuous within the inner core of tree crowns.
• 5 The results firstly emphasize the distribution of young foliage as a key factor affecting the abundance of many herbivores and, secondly, the importance of the local illumination regime for host leaf production and its indirect effects on the spatial distribution of arboreal arthropods.

     

Quantitative and qualitative shifts in defensive metabolites define chemical defense investment during leaf development in Inga,a genus of tropical trees

Selective pressures imposed by herbivores are often positively correlated with investments that plants make in defense. Research based on the framework of an evolutionary arms race has improved our understanding of why the amount and types of defenses differ between plant species. However, plant species are exposed to different selective pressures during the life of a leaf, such that expanding leaves suffer more damage from herbivores and pathogens than mature leaves. We hypothesize that this differential selective pressure may result in contrasting quantitative and qualitative defense investment in plants exposed to natural selective pressures in the field. To characterize shifts in chemical defenses, we chose six species of Inga, a speciose Neotropical tree genus. Focal species represent diverse chemical, morphological, and developmental defense traits and were collected from a single site in the Amazonian rainforest. Chemical defenses were measured gravimetrically and by characterizing the metabolome of expanding and mature leaves. Quantitative investment in phenolics plus saponins, the major classes of chemical defenses identified in Inga, was greater for expanding than mature leaves (46% and 24% of dry weight, respectively). This supports the theory that, because expanding leaves are under greater selective pressure from herbivores, they rely more upon chemical defense as an antiherbivore strategy than do mature leaves. Qualitatively, mature and expanding leaves were distinct and mature leaves contained more total and unique metabolites. Intraspecific variation was greater for mature leaves than expanding leaves, suggesting that leaf development is canalized. This study provides a snapshot of chemical defense investment in a speciose genus of tropical trees during the short, few‐week period of leaf development. Exploring the metabolome through quantitative and qualitative profiling enables a more comprehensive examination of foliar chemical defense investment.     

Differential effects of host plant hybridization on herbivore community structure and grazing pressure on forest canopies

Interspecific hybridization in plants is known to have ecological effects on associated organisms. We examined the differences in insect herbivore community structure and grazing pressure on tree canopy leaves among natural hybrids and their parental oak species. We measured leaf traits, herbivore community structure, and grazing pressure on leaves of two oak species, Quercus crispula and Q. dentata, and their hybrids. The concentration of nitrogen in canopy leaves was greater in hybrids and in Q. dentata than in Q. crispula. The concentration of total phenolics was lower in hybrids than in Q. crispula. The concentration of condensed tannin was greater in hybrids than in Q. crispula. Relative herbivore abundance and species richness were greater on oak hybrids than on either parental species; herbivore species diversity and composition on hybrids were close to those on Q. crispula. Herbivore grazing pressure was lower on hybrids and Q. dentata than on Q. crispula. There was a negative correlation between herbivore grazing pressure and leaf nitrogen, suggesting that interspecific variation among oak taxa in herbivore pressure may be explained by leaf nitrogen; variation in herbivore community structure among oak taxa is likely to be controlled by polygenic leaf traits. Differing responses of (1) herbivore community structure and (2) herbivore grazing pressure to host plant hybridization may play important roles in regulating herbivore biodiversity in cool‐temperate forest canopies.     

Home-field advantage in decomposition of leaf litter and insect frass

Home-field advantage (HFA) hypothesis regarding litter decomposition states that litter is decomposed more rapidly in the habitat from which it is derived (i.e., home) than in other habitat (i.e., away) due to local adaptation of soil decomposers. We tested the HFA hypothesis regarding decomposition of leaf litter, insect frass, and their mixtures, using laboratory incubation of leaf litter from an evergreen (Pinus densiflora) and a deciduous (Quercus acutissima) tree species, frass excreted by two insect herbivores (Dendrolimus spectabilis and Lymantria dispar) fed on one of the two trees, and soil collected underneath the two trees. We found evidence that decomposers in each soil were specialized to decompose the litter derived from the tree species above them, indicating that the HFA occurred in litter decomposition. In contrast, the HFA was not detected in the decomposition of insect frass or litter-frass mixtures. Mixing with D. spectabilis frass non-additively decelerated, while mixing with L. dispar frass non-additively accelerated, decomposition of the mixtures, independent of soil and litter types. These indicate that the presence of insect herbivores may make it difficult to form and maintain a decomposer community specialized to a certain leaf litter, and that it may consequently cancel or weaken HFA in litter decomposition.