Monoculture,polyculture, and polyvariety in tropical forest Swidden cultivation

A number of researchers have suggested that polyculture is characteristic of native tropical forest swiddens and have adduced theory from community ecology to account for its adaptiveness. Ye'kwana and Yanomamö swidden cultivation is examined, and it is shown that polyculture is not practiced to any significant degree. Instead, the concept of polyvariety is introduced along with a number of other cultivation practices that more simply account for the adaptiveness of Ye'kwana and Yanomamö gardening. In addition, comparative data from other parts of the tropical world indicate that polyculture is no more common than monoculture and recent advances in ecological research indicate that the diversity-stability hypothesis that underpins adaptive arguments of polyculture is in need of drastic revision.     

Tropical forest mimicry in Swiddens: A reassessment of Geertz's Model with Amazonian data

Geertz's famous hypothesis that horticulturalists practicing shifting cultivation intercrop their plots to mimic the protective structure of the tropical forest is evaluated in view of data on the structural ecology of the tropical forest and swidden plots in Amazonian Ecuador. The cultivation practices of the Siona-Secoya Indians reveal a three-fold typology of cropping patterns (high-diversity intercropping, low-diversity intercropping, and monocropping), with variation among the types in terms of plot size and distribution, cultigen inventory, structural complexity, and function. These gardens and the tropical forest are compared and analyzed in terms of morphology, ecological characteristics, and human manipulation and utilization. Gardens with high-diversity intercropping show certain similarities to the tropical forest, as Geertz's model predicts, but their highly transient structure does not function as a mature ecosystem. Furthermore, plots with low-diversity intercropping and monocropping show few similarities to the forest. Garden structure is best understood in terms of the economic utilization of tropical cultigens displaying specific habits, rather than by analogy to tropical forest physiognomy and function.     

海南岛霸王岭热带低地雨林植被恢复动态

热带次生林具有重要的物种保育和固碳功能, 然而高强度的干扰会导致次生林早期出现类似季雨林的阶段, 因而群落恢复速度和方向是当前热带生态学研究中最为关注的议题之一。该文以海南岛在刀耕火种弃耕地形成的不同演替阶段的次生林为研究对象, 比较森林不同恢复时间(12年、25年、55年)群落中的不同年龄(幼树、小树、成年树)个体与低地雨林老龄林的物种组成、多样性和群落结构差异, 探讨刀耕火种弃耕地恢复过程中的群落组配过程。首先, 在海南岛霸王岭林区内建立7个1 hm²(100 m × 100 m)的样地, 并调查样地内所有胸径≥ 1 cm的木本植物个体(不包括木质藤本)的种类、胸径大小和树高。无度量多维标定法(NMS)排序结果表明, 刀耕火种弃耕地恢复群落与老龄林的物种组成存在明显差异, 并且其物种组成差异随着径级增加而逐渐减小。刀耕火种弃耕地群落物种累积速度缓慢, 25年和55年恢复群落的种面积、种个体和种多度曲线无差异, 存在一个明显的停滞阶段。与物种组成相比, 群落结构恢复相对较迅速, 但仍没有形成老龄林阶段中的复杂结构。萌生个体在早期恢复群落中占有较高比例, 其个体密度和胸高断面积分别占总数的39.9%和55.9%, 但在恢复中后期迅速降低。刀耕火种弃耕地恢复群落中以先锋种和非先锋喜光种为主。虽然耐阴种随演替而逐渐增加, 但恢复中后期群落中的耐阴种重要值仅为老龄林的27.7%。这些结果表明, 虽然刀耕火种弃耕地恢复群落缓慢地逐渐接近最终恢复目标, 但仍然存在 很大的不确定性。刀耕火种弃耕地恢复过程中的异速恢复和停滞阶段需要纳入今后群落演替模型构建和森林固碳效益核甘共苦算中。     

The Potential of Tree Rings for the Study of Forest Succession in Southern Mexico

Studies of tropical secondary forest succession face strong limitations due to the slow pace of succession and the time-consuming task of monitoring processes. The occurrence of tree rings in secondary forest trees may help expand our knowledge on succession in these systems and may be useful for fallow dating in chronosequence studies. We examine here the potential of tree rings to study forest succession by sampling 70 species along chronosequences of dry and wet forests in southern Mexico. Based on wood anatomical features, we estimated that about 37 percent of the species presented distinct growth rings useful for ring studies. Overall, maximum number of rings matched well the interview-based fallow ages but, at some sites, trees had consistently higher numbers of rings, probably due to errors in fallow ages derived from interviews. Best fallow age estimations were obtained by examining rings in both pioneer and nonpioneer species. Reconstruction of species' establishment dates revealed that pioneer and nonpioneer species establish early during succession, and that species of both groups continue to recruit after many years. Our study clearly shows that tree ring analysis is a promising tool for studies on secondary forest succession in the tropics.     

Farmers’ Perspectives on Vital Soil-related Ecosystem Services in Intensive Swidden Farming Systems in the Peruvian Amazon

A growing dilemma is how to conserve Amazonian forest while allowing local people to secure their livelihoods. Small-scale swidden farming in Amazonia is entirely dependent on the continued provision of ecosystem services (ES) that generate the conditions for agriculture. This study identified soil-related ES needed for, and enhanced by, productive swidden systems from the farmer’s perspective. Workshops in six farming communities in northeastern Peru discussed various land uses, swidden systems that continue to be productive, and swidden systems on degraded land. The participating farmers noted changes in their production systems and described the ES (or lack thereof) in terms of soil quality, crop production quantity and quality, burning practices, forest regeneration, and farming skill. The central elements described in farmers’ own strategies for managing soil-related ES were fallow management for biomass production and crop diversity, factors identified as central to future ES management work in established agricultural areas in Amazonia.     

Secondary Succession and Indigenous Management in Semideciduous Forest Fallows of the Amazon Basin1

To the discussion on secondary succession in tropical forests, we bring data on three under‐addressed issues: understory as well as overstory changes, continuous as opposed to phase changes, and integration of forest succession with indigenous fallow management and plant uses. Changes in vegetation structure and species composition were analyzed in secondary forests following swidden agriculture in a semideciduous forest of Bolivian lowlands. Twenty‐eight fallows, stratified by four successional stages (early = 1–5 yr, intermediate = 6–10 yr, advanced = 12–20 yr, and older = 22–36 yr), and ten stands of mature forests were sampled. The overstory (plants ≥5 cm diameter at breast height [DBH]) was sampled using a 20 × 50 m plot and the understory (plants <5 cm DBH) in three nested 2 × 5 m subplots. Semistructured interviews provided information on fallow management. Canopy height, basal area, and liana density of the overstory increased with secondary forest age. The early stage had the lowest species density and diversity in the overstory, but the highest diversity in the understory. Species composition and abundance differentiated mature forests and early successional stage from other successional stages; however, species showed individualistic responses across the temporal gradient. A total of 123 of 280 species were useful with edible, medicinal, and construction plants being the most abundant for both over‐ and understories. Most of Los Gwarayo preferred mature forests for making new swidden, while fallows were valuable for crops, useful species, and regenerating timber species.     

Shifting Cultivation and Fire Policy: Insights from the Brazilian Amazon

Fires in humid tropical forests are increasingly frequent due to severe dry seasons, forest degradation and agricultural expansion. One agent implicated in current discourse surrounding tropical forest fires is the small-scale farming peasantry who rely on fire in swidden (shifting cultivation, slash-and-burn) agriculture. The environmental degradation associated with fire has led to government responses at multiple scales (international, national, state, regional) via policies aimed mainly at managing ignition sources. However, continued increase in forest fires suggests that these policies may be having limited impact and a fresh evaluation of current policy approaches to fire management is needed. We review fire policy measures with insights of caboclo farming practices and perspectives from Eastern Amazonia and examine the congruence between policy and practice. We demonstrate a significant disparity between policy requirements such as firebreaks and actual fire management practices, in which measures rarely meet requirements outlined in legislation. We explore the origins and the impacts of these disparities, focussing on smallholder farm-level management measures and local capacity. Incomplete knowledge coupled with marginal awareness of legal requirements served to propagate widespread erroneous beliefs in what these are. This analysis at multiple scales (international, national, state, regional) will contribute to developing greater congruence between fire policies and smallholder farming practices.     

Structure,diversity, and regeneration of tropical dry deciduous forest of northern India

This study examined the impact of disturbance on the pattern of diversity, forest structure and regeneration of tree species in the Vindhyan dry tropical forest of India. A total of 1500 quadrats distributed over five, 3-ha permanent plots in five sites, differing in degree of disturbance, were used to enumerate and measure the tree species. A total of 65 species with 136,983 individuals were enumerated in the total 15-ha area for stems 30 cm height. The number of species and number of stems ranged from 12 to 50 and 8063–65331 per 3-ha area. The number of species and stems for trees 10 cm dbh ranged from 3 to 28 species, with a mean value of 16 species ha^–1, and from 16 to 477 stems, with a mean value of 256 stems ha^–1, respectively. The adult based PCA ordination indicated uniqueness of sites in terms of species composition and habitat characteristics. PCA ordination also showed uniqueness of sites in terms of seedling composition, but the seedling and adult distributions were not spatially associated. The distinct species composition at the different sites and at the two life-cycle stages on the same site is indicative of marked spatio-temporal dynamics of the dry tropical forest. The density–diameter semi-logarithmic curves ranged from a near linear to an overall concave appearance with a limited plateau in the mid-diameter ranges. The -diversity and its components decreased with increasing disturbance intensity, reflecting enhanced utilization pressure with increasing disturbance. The site-wise and species-wise regression analyses of the number of individuals in different stages of the species revealed that both the level of disturbance and the nature of species strongly affect the regeneration. In conclusion, although the forest is relatively species-poor, the differential species composition on different sites and the temporal dynamics lend a unique level of diversity to the tropical dry deciduous forest.     

An Assessment of Trends in the Extent of Swidden in Southeast Asia

Swidden systems consisting of temporarily cultivated land and associated fallows often do not appear on land use maps or in statistical records. This is partly due to the fact that swidden is a diverse and dynamic land use system that is difficult to map and partly because of the practice of grouping land covers associated with swidden systems into land use or land cover categories that are not self-evidently linked to swiddening. Additionally, in many parts of Southeast Asia swidden systems have changed or are in the process of changing into other land use systems. This paper assesses the extent of swidden on the basis of regional and national sources for nine countries, and determines the pattern of changes of swidden on the basis of 151 cases culled from 67 articles. Findings include (1) a majority of the cases document swidden being replaced by other forms of agriculture or by other livelihood systems; (2) in cases where swiddening is still practiced, fallow lengths are usually, but not always, shorter; and (3) shortened fallow length does not necessarily indicate a trend away from swidden since it is observed that short fallow swidden is sometimes maintained along with other more intensive farming practices and not completely abandoned. The paper concludes that there is a surprising lack of conclusive data on the extent of swidden in Southeast Asia. In order to remedy this, methods are reviewed that may lead to more precise future assessments.     

Impacts of Selective Logging and Agricultural Clearing on Forest Structure, Floristic Composition and Diversity, and Timber Tree Regeneration in the Ituri Forest, Democratic Republic of Congo

Mature tropical forests at agricultural frontiers are of global conservation concern as the leading edge of global deforestation. In the Ituri Forest of DRC, as in other tropical forest areas, road creation associated with selective logging results in spontaneous human colonization, leading to the clearing of mature forest for agricultural purposes. Following 1-3 years of cultivation, farmlands are left fallow for periods that may exceed 20 years, resulting in extensive secondary forest areas impacted by both selective logging and swidden agriculture. In this study, we assessed forest structure, tree species composition and diversity and the regeneration of timber trees in secondary forest stands (5-10 and ~40 years old), selectively logged forest stands, and undisturbed forests at two sites in the Ituri region. Stem density was lower in old secondary forests (~40 years old) than in either young secondary or mature forests. Overall tree diversity did not significantly differ between forest types, but the diversity of trees ≥10 cm dbh was substantially lower in young secondary forest stands than in old secondary or mature forests. The species composition of secondary forests differed from that of mature forests, with the dominant Caesalpinoid legume species of mature forests poorly represented in secondary forests. However, in spite of prior logging, the regeneration of high value timber trees such as African mahoganies (Khaya anthotheca and Entandrophragma spp.) was at least 10 times greater in young secondary forests than in mature forests. We argue that, if properly managed and protected, secondary forests, even those impacted by both selective logging and small-scale shifting agriculture, may have high potential conservation and economic value.     

Some ecophysiological features in leaves of plants in an oak-hornbeam forest

Six different leaf indexes, indicating the anatomical and ecophysiological features in leaves, were used for characterizing the three principal layers (tree, shrub and herb) of an oak-hornbeam forest in southwest Slovakia, Czechoslovakia. The most pronounced differences in the leaf indexes followed were found between those at the active surface of the forest canopy (sun leaves of tall trees) and those in the herb layer. Differences exist between individual layers as well as between species within each layer. They should be taken into consideration in ecological and ecophysio-logical studies and in modelling forest ecosystems.     

Drivers of aboveground wood production in a lowland tropical forest of West Africa: teasing apart the roles of tree density,tree diversity,soil phosphorus,and historical logging

Tropical forests currently play a key role in regulating the terrestrial carbon cycle and abating climate change by storing carbon in wood. However, there remains considerable uncertainty as to whether tropical forests will continue to act as carbon sinks in the face of increased pressure from expanding human activities. Consequently, understanding what drives productivity in tropical forests is critical. We used permanent forest plot data from the Gola Rainforest National Park (Sierra Leone) – one of the largest tracts of intact tropical moist forest in West Africa – to explore how (1) stand basal area and tree diversity, (2) past disturbance associated with past logging, and (3) underlying soil nutrient gradients interact to determine rates of aboveground wood production (AWP). We started by statistically modeling the diameter growth of individual trees and used these models to estimate AWP for 142 permanent forest plots. We then used structural equation modeling to explore the direct and indirect pathways which shape rates of AWP. Across the plot network, stand basal area emerged as the strongest determinant of AWP, with densely packed stands exhibiting the fastest rates of AWP. In addition to stand packing density, both tree diversity and soil phosphorus content were also positively related to productivity. By contrast, historical logging activities negatively impacted AWP through the removal of large trees, which contributed disproportionately to productivity. Understanding what determines variation in wood production across tropical forest landscapes requires accounting for multiple interacting drivers – with stand structure, tree diversity, and soil nutrients all playing a key role. Importantly, our results also indicate that logging activities can have a long‐lasting impact on a forest's ability to sequester and store carbon, emphasizing the importance of safeguarding old‐growth tropical forests.     

Changes in tree phenology along natural regeneration in a seasonally dry tropical forest

The applicability of succession models from temperate and tropical wet forests to threatened seasonally dry tropical forests (SDTFs) is questioned. Plant phenology affects ecosystem functions and changes along forest regeneration gradient. To investigate the recovery of ecological functions after disturbances in a SDTF, we recorded the vegetative and reproductive phenologies for trees (DBH >5 cm) for 17 months in southeast Brazil in three successional stages: early (10–15 years after clearing), intermediate (25–30) and late (>50). The vegetative phenology of the 523 individuals was strongly seasonal, with 3% of individuals presenting green leaves in a deciduous dry season. Besides structural and floristic differences, phenological trends were similar between the later stages. Reproduction occurred with higher intensities in the early stage and in the advanced stages only in the dry season, providing key resources to local fauna. The studied SDTF is resilient to ecological functions, rapidly recovering functional processes. The integration of structural and functional knowledge of succession of STDFs may lead to better management of its secondary remnants. Our study suggests that classical forest succession theory developed for other ecosystems may not fully reflect the pattern of SDTF succession, an ecosystem that originally covered 42% of the earth's tropical and subtropical landmass.     

Relocating plants from swidden fallows to gardens in Southwestern China

As upland farmers in Southeast Asia change from shifting cultivation to permanent agriculture and lose access to swidden-fallow forests and their resources, they are introducing economically important forest and fallow plants into their house gardens. We describe this process in Daka, a village of Hani ethnicity in Yunnan Province, China. Daka smallholders collect both seeds and seedlings from fallow forests and transfer the plants to house gardens, initiating the transformation of wild species to a cultivated or semi-domesticated one. Two kinds of species are commonly transferred from swidden fallows to house gardens. Some are particularly rare, others are plants in great demand. Between 1998 and 2000 Daka households earned an average of US$68.20 annually from the products of fallow forests. We also found that villagers harvest and use 76 plant species from fallow forests and 126 species from house gardens. Twenty-two species found in house gardens that had been transferred from fallow forests. Households vary widely in the frequency with which they engage in this pattern. We believe that local knowledge of these practices is a potentially important resource in the development of other areas of smallholder farming.     

Spatial distribution of the gypsy moth [Lepidoptera: Lymantriidae] and some of its parasitoids within a forest environment

Aspects of the microhabitat distributions of the gypsy moth,Porthetria dispar (L.) (Lepidoptera: Lymantriidae), and some of its parasitoids were investigated in the field by means of sticky panels and gypsy moth egg masses exposed at different heights in trees, by egg masses exposed within forested and cleared areas, and by gypsy moth pupal collections from different heights in trees.Ooencyrtus kuwanai (Howard)(Hymenoptera: Encyrtidae), Apanteles melanoscelus Ratzeburg (Hymenoptera: Braconidae), and gypsy moths were caught most frequently on stickly panels placed in upper portions of trees. In contrast,Apanteles laeviceps Ashmead, a parasitoid of cutworms, was most often caught near the forest floor.O. kuwanai attacked equally egg masses exposed at different heights in trees, but parasitized those in a clearing less often than those within the forest prosper.Brachymeria intermedia (Nees) (Hymenoptera: Chalcididae) emerged mostly from pupae collected near the tops of trees and not at all from those collected below 5 m. The results are discussed as they relate to field sampling procedures, behavioral activities of gypsy moth and parasitoid adults, and integrated control possibilities for the gypsy moth.     

Plant functional traits under cattle grazing and fallow age scenarios in a tropical dry forest of Northwestern Mexico

Evaluating plant functional traits helps to understand how plants respond to changing environmental conditions and resource availability associated with disturbance events. Livestock production is one of the primary drivers of tropical forest loss and degradation. Livestock alter environmental conditions within the forest by grazing, trampling and nutrient inputs, which in turn can influence species composition and functional traits of species. Understanding how livestock influence functional traits along a successional gradient is poorly understood. Here, we studied the effect of cattle grazing and fallow age on plant functional traits and soil nutrients in secondary and old-growth tropical dry forests. We analyzed plant functional traits of the most important species in successional and old-growth forest communities in both cattle present and cattle excluded plots. Our results showed the effects of cattle grazing and fallow age on plant functional traits, with fallow age explaining more variation than cattle grazing. In early succession, functional traits were associated with water conservation (thicker leaves, lower specific leaf area), and in later successional they were linked with sunlight conservation (larger height, higher specific leaf area). The presence of large fruits and seeds in advanced successional sites suggests high resource availability, which may help plants to successfully reproduce. Moreover, under cattle grazing some functional traits are associated with herbivory defense (high foliar dry weight and thick leaves). Even though N and C increased as succession advanced, the sites with cattle grazing had higher NH4 and NO3 concentrations as a result of fecal deposition. Plant functional traits responded to fallow age than to cattle grazing. Our study showed that cattle exclusion, as a management and biodiversity conservation strategy, contributes positively to soil nutrition. Thus, fallow age and cattle exclusion facilitate soil recovery and allows establishing species with suitable functional attributes for overcoming environmental filters in abandoned cattle fields.     

Land use types influenced avian assemblage structure in a forest–agriculture landscape in Ghana

The conservation of biodiversity within tropical forest regions does not lie only in the maintenance of natural forest areas, but on conservation strategies directed toward agricultural land types within which they are embedded. This study investigated variations in bird assemblages of different functional groups of forest‐dependent birds in three agricultural land types, relative to distance from the interior of 34 tropical forest patches of varying sizes. Point counts were used to sample birds at each study site visited. Data from counts were used to estimate species richness, species evenness, and Simpson's diversity of birds. Mean species richness, evenness, and diversity were modeled as responses and as a function of agricultural land type, distance from the forest interior and three site‐scale vegetation covariates (density of large trees, fruiting trees, and patch size) using generalized linear mixed‐effect models. Mean observed species richness of birds varied significantly within habitat types. Mean observed species richness was highest in forest interior sites while sites located in farm centers recorded the lowest mean species richness. Species richness of forest specialists was strongly influenced by the type of agricultural land use. Fallow lands, density of large trees, and patch size strongly positively influenced forest specialists. Insectivorous and frugivorous birds were more species‐rich in fallow lands while monoculture plantations favored nectarivorous birds. Our results suggest that poor agricultural practices can lead to population declines of forest‐dependent birds particularly specialist species. Conservation actions should include proper land use management that ensures heterogeneity through retention of native tree species on farms in tropical forest‐agriculture landscapes.     

Effect of forest fragmentation on the woody flora of the highlands of Chiapas, Mexico

This study was conducted in the Chiapas Highlands, a tropical mountain region where traditional agricultural practices have resulted in a mosaic landscape of forest fragments embedded in a matrix of secondary vegetation and crop fields. The question addressed was how may woody species richness be affected by forest fragment attributes derived from traditional land-use patterns. Species inventories of total woody species, canopy and understorey trees, and shrubs were obtained in 22 forest fragments (5 ha). Multiple regression analyses were applied to examine the effects of size, matrix, isolation and shape of the forest fragments on richness of these species guilds. Fragment size was correlated with shape (r = 0.75) and isolation (r = –0.69), and isolation was correlated with shape (r = –0.75). Total species richness, and number of shrubs and understorey trees in fragments were related to isolation; moreover, additive effects of fragment shape were found for shrubs. The number of canopy species was not related to any fragment variable. Matrix did not help to explain species richness, possibly due to the landscape structure created by the traditional land-use patterns. In addition to size and isolation, we point out the need of considering shape and matrix as additional fragmentation attributes, along with social and economic factors, if we are ever going to be successful in our management and conservation actions.     

The role of tree sprouts in the restoration of stand structure and species diversity in tropical moist forest after slash-and-burn agriculture in Eastern Paraguay

The contribution of tree sprouts to the recovery of tropical moist forest in Eastern Paraguay after swidden agriculture was examined in 2–15 yr old forest fallows and compared with sprouting in mature forest. The proportion of stems of sprout origin, as individuals arising from stumps or lower parts of live stems ( 1 m), in the stem density declined from 59.5% (stems 1–4.9 cm DBH) and 21.0% (stems 5 cm DBH) in the young regrowth stands (2–5 yr old) to 32.9% and 19.6%, respectively in the older regrowth stands (10 and 15 yr old). Sprouts were absent in the mature forest. Out of 58 species sampled in the regrowth stands, 28 occured both as resprouts and seed regeneration, 7 were only found as resprouts, and 23 were only present as seed-established individuals. No significant relationship was found between the successional status or the growth form of species and apparent resprouting capacity. Seed-established individuals of Trema micrantha were predominant in the two and three-year old regrowth stands. In the more advanced successional stages, T. micrantha was replaced by Cecropia pachystachia and other secondary species. Species richness increased during succession. Species-abundance distribution in the successional stands followed a log series pattern, whereas the mature forest showed a log normal distribution. Floristic similarity to the mature forest, calculated with the qualitative Soerensen index, increased from 0.45 (1–4.9 cm DBH) in the young regrowth stands to 0.52 in the older regrowth stands. In the tree stratum ( 5 cm DBH), however, floristic composition approached only 0.28 in the younger regrowth stands and 0.44 in the older regrowth stands, respectively that of the mature forest.     

Insect attraction to herbivore-induced beech volatiles under different forest management regimes

Insect herbivore enemies such as parasitoids and predators are important in controlling herbivore pests. From agricultural systems we know that land-use intensification can negatively impact biological control as an important ecosystem service. The aim of our study was to investigate the importance of management regime for natural enemy pressure and biological control possibilities in forests dominated by European beech. We hypothesize that the volatile blend released from herbivore-infested beech trees functions as a signal, attracting parasitoids and herbivore enemies. Furthermore, we hypothesize that forest management regime influences the composition of species attracted by these herbivore-induced beech volatiles. We installed flight-interception traps next to Lymantria dispar caterpillar-infested young beech trees releasing herbivore-induced volatiles and next to non-infested control trees. Significantly more parasitoids were captured next to caterpillar-infested trees compared to non-infested controls, irrespective of forest type. However, the composition of the trophic guilds in the traps did vary in response to forest management regime. While the proportion of chewing insects was highest in non-managed forests, the proportion of sucking insects peaked in forests with low management and of parasitoids in young, highly managed, forest stands. Neither the number of naturally occurring beech saplings nor herbivory levels in the proximity of our experiment affected the abundance and diversity of parasitoids caught. Our data show that herbivore-induced beech volatiles attract herbivore enemies under field conditions. They further suggest that differences in the structural complexity of forests as a consequence of management regime only play a minor role in parasitoid activity and thus in indirect tree defense.