Predation on artificial avian nests is higher in forests bordering small anthropogenic openings

Habitat edges alter the diversity of avian communities and are often associated with higher rates of nest predation. However, most previous studies on habitat edges have been conducted along long linear corridors or at the transition between large field and forest patches in agricultural systems. Less is known about predation rates when the habitat edge is the result of a small interior forest opening. We assessed predation rates on artificial nests mimicking ground and shrub nesters in Northern Michigan forests perforated by small clearings used previously for oil and gas extraction. Nests were placed at varying distances from the edges of these clearings, and in similar spatial arrangements within unfragmented interior forest plots. Predation rates increased in forests near edges, but significant impacts were limited to shrub nests. Markings on predated clay eggs indicated that the type of predation also differed. Scratch marks were the most prevalent egg indentation, but eggs with poked holes were twice as common near the forest edge. The increase in the number of poked eggs suggests that a higher density of avian predators occurred in forests near an edge. Predation rates at forest edges did not vary by distance from the forest edge. Surveys of the avian community revealed differences between edge and interior forests: American Crows Corvus brachyrhynchos and Blue Jays Cyanocitta cristata, two species known to predate bird nests, were more common near edges. Our results suggest that small forest openings alter the avian community and may adversely impact reproductive output in some species. If the alteration of these processes results in population‐level impacts, small forest perforations should be avoided when possible and reforestation of abandoned well‐pads should be encouraged.     

Relationships between flowering phenology,plant size and reproductive success in shape Lotus corniculatus (Fabaceae)

Over three years the flowering phenology of individuals of Lotus corniculatus has been studied in relation to fruit set and seed predation to determine the relationships between four components of flowering time, plant size and reproductive success. Timings of first and peak flowering, and duration and synchrony of flowering differed between individuals in the same years. Between years, timing of first flowering was highly correlated for the same individuals, and was closely correlated with plant size and duration of flowering–larger plants flowered earlier and for a longer period. Peak flowering and synchrony were not correlated between-years for individuals.Fruit production and seed predation were correlated with some of the components of flowering phenology in some years, but not in others. The inconstancy of these relationships suggests that directional or stabilising selection is not acting consistently on the aspects of reproductive success studied in this work. The inconstancy of selection may result in the rather asynchronous flowering phenologies of individuals of L. corniculatus observed.We emphasize the importance of studying different components of flowering phenology in relation to individual plant size over several seasons. This work has shown that plant size not only has a direct effect on individual plant fecundity but also can influence flowering time and hence indirectly affect reproductive output.     

Dissecting components of flowering pattern: size effects on female fitness

Flowering synchrony is essential for plant reproductive success, especially in the case of small‐sized populations of self‐incompatible species. Closely related to synchrony, flowering intensity influences pollinator attraction and pollinator movements. Thus, a high flowering intensity may increase pollinator attraction but, at the same time, may also increase the probability of geitonogamous pollinations. Depending on the mating system, the female fitness of plants in small populations may be affected by both the positive effects of higher flowering synchrony and pollinator attraction and the negative effects of geitonogamous pollinations induced by a high flowering intensity. It was hypothesized that different‐sized plants in a population would show contrasting flowering patterns, resulting in differences in pollinator behaviour. These influences could result in differences in mating and female reproductive success. This hypothesis was tested by studying the flowering pattern of Erodium paularense (Geraniaceae), a rare and endangered endemic of central Spain. The temporal distribution of flower production was explored throughout the reproductive season, and the probability of xenogamy and geitonogamy and their relationship to plant size and fitness components were calculated. The analysis of this partially self‐compatible species showed diverse flowering patterns related to different plant sizes. Small plants produced a larger number of seeds per fruit in spite of having lower values of flowering synchrony. By contrast, large plants produced a larger number of seeds from geitonogamous pollinations. The effect of different flower displays and outcrossing rates on seed set varied throughout the season in the different groups. Our findings highlight the relevance of individual plant size‐dependent phenology on female reproductive success and, in particular, on the relationship between flowering synchrony and fitness. © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society, 2008, 156 , 227–236.     

Consequences of variation in flowering phenology for seed head herbivory and reproductive success in Erigeron glaucus (Compositae)

Summary We examined the relationship between flowering phenology, reproductive success (seed production only), and seed head herbivory for 20 similarly sized clones of Erigeron glaucus growing at Bodega Bay Reserve, northern California, USA. Although clones tended to reach peak flowering on the same date, they differed in the proportion of their total flowers produced around that date (flowering synchrony). Clones also differed in the number and density of flower heads presented at any one time to pollinators and herbivores (floral display). Both of these characteristics had consequences for herbivory and plant reproductive success. The proportion of flower heads damaged by insect herbivores was greater for clones that concentrated flowering activity during the main flowering period for the population as a whole (high synchrony) compared to clones that spread flowering out temporally. The primary reason for this result was that clones with low flowering synchrony produced a significant proportion of their flower heads during the fall and therefore, escaped attack by the tephritid fly, Tephritis ovatipennis. Clones with intermediate synchrony had lower seed success (total number of viable seeds produced over the year) than clones with either low or high synchrony. The proportion of flower heads damaged by insect herbivores and number of tephritid flies reared from flower heads were both negatively correlated to floral display while seed head mass and germination rates were positively related to display. Thus, clones which produced dense floral displays were favored both in terms of reduced herbivory and increased successful seed production.     

濒危植物长柄双花木自然种群生殖构件的时空动态

根据野外观测数据,研究了濒危植物长柄双花木(Disanthus cercidifolius var.longipes)在不同生境条件下自然种群开花物候变异以及生殖构件的时空动态.结果表明,不同群落类型中长柄双花木开花物候存在一定差异,在纯林中个体的平均花期最长,达75d,而个体间开花同步性最低,仅为0.717;竹林中的个体花期最短,只有47d,而同步指数最高,达0.968.生殖枝数量及花序数均随个体年龄增长而增加,在年龄为30～35年时达最高,此后又呈下降趋势;在海拔为810m左右处,个体生殖枝数及着生的花序数均达最大值,且冠层间差异明显.纯林中个体着生的生殖枝数及每生殖枝上着生的花序数均最高,分别为411.39和7.857;竹林中个体每生殖枝着生的花序数显著低于其它群落.年龄小的个体,各时期生殖构件败育率均高于年龄大的个体;另外,生殖构件的败育与海拔高度间无相关性,而与构件发育时间、分布的树冠层次以及所处的群落类型相关.影响生殖构件败育的主要因子是光照条件.     

Gender specialization and factors affecting fruit set of the wind-pollinated heterodichogamous Juglans regia

Gender expression, flowering phenology, reproductive performance and factors affecting fruit set (i.e., flowering synchrony, size and distance to the nearest pollen donor) were investigated in a cultivated population of a wind-pollinated self-compatible heterodichogamous Juglans regia (Juglandaceae). Four flowering morphs, (i.e., protandrous, protogynous, male and female) were observed. The sexual functions of the protandrous and protogynous morphs were almost synchronous; however, they were not reciprocal, and the separation of male and female flowering within most monoecious individuals was not complete. Thus, within-morph mating and geitonogamous pollination may be common. The ratio of protandrous versus protogynous morphs was biased towards the protandrous morph, but the fruit set did not differ between the morphs, suggesting that the fruit set of the protandrous morph could be partly compensated by within-morph pollination. The ratio of the female flower number or fruit number to the total male catkin length was higher in the protogynous morph than in the protandrous morph and did not vary with plant size, suggesting that gender variation was not size dependent and that the sexual function of protandrous morphs was more male biased. Fruit set depended on plant size only for protandrous morphs. The fruit set of individual plants decreased with increasing distance to the nearest pollen donor regardless of morph, possibly because of pollen limitation. The fruit set of individual plants increased with flowering synchrony, indicating that flowering synchrony could affect reproductive success.     

Differential effects of anthropogenic edges and gaps on the reproduction of a forest‐dwelling plant: The role of plant reproductive effort and nectar robbing by bumblebees

Forest fragmentation produces sharp changes in the spatial configuration of remnant forest patches, which include the increasing influence of patch edges on the interior of forest patches (‘edge effects’). Human activities responsible for forest fragmentation tend also to change the internal characteristics of remnant patches, for example, through the creation of gaps by selective logging. While edges and gaps can be expected to cause comparable changes in the micro‐environmental conditions of the forest, their effects on forest‐dwelling species and their interactions are not necessarily comparable. This study compares the effect of forest edges and anthropogenic gaps on the reproductive success of a self‐incompatible epiphytic plant (Mitraria coccinea), mediated by changes in its relationship with mutualists (pollinators) and antagonists (flower larcenists). Mitraria coccinea's flowers are pollinated by hummingbirds and robbed by the bumblebee Bombus dahlbomii. Edges and gaps had comparable positive effects on flower production and fruit set; however, nectar robbing was up to sevenfold higher in patches with numerous gaps and resulted in lower reproductive success (fruit set and total fruit crop). Forest fragmentation studies should therefore avoid treating forest remnants as homogeneous units (i.e. focusing exclusively on their characteristics and connectivity), without taking into account the internal heterogeneity caused by anthropogenic pressures (e.g. gaps and edge effects).     

Rainfall and flowering synchrony in a tropical shrub: Variable selection on the flowering time ofErythroxylum havanense

Summary We tested the adaptive significance of flowering synchrony by means of a quantitative analysis of selection and by flowering induction experiments with the deciduous shrubErythroxylum havanense. Temporal schedules of flower and fruit production were determined for a local population (in three sites) in a Mexican seasonal forest for 2 years (1987–1988). The consequences of natural variation in flowering time (flowering initiation day) on maternal reproductive success (fecundity) were evaluated. We observed high levels of inter- and intraindividual flowering synchrony in 1987, but not in 1988 and this contrast was related to differences in rainfall patterns between the two years. A significant proportion (15.4%) of the phenotypic variation in flowering initiation day was accounted for by environmental variance. The expression of phenotypic variance of flowering time and, consequently, the opportunity for selection to act, are controlled by annual variation in rainfall. Despite the between-year difference in flowering synchrony, we detected a relatively intense directional selection on flowering initiation day in both years, but selection coefficients were of opposite sign (standardized directional gradients were –0.326 and 0.333 for 1987 and 1988, respectively). For both years there was a significant relationship between individual relative fitness and the number of neighbouring flowering plants in a given day, suggesting positive frequency-dependent selection.     

Temperature effects on forest herbs assessed by warming and transplant experiments along a latitudinal gradient

Slow‐colonizing forest understorey plants are probably not able to rapidly adjust their distribution range following large‐scale climate change. Therefore, the acclimation potential to climate change within their actual occupied habitats will likely be key for their short‐ and long‐term persistence. We combined transplant experiments along a latitudinal gradient with open‐top chambers to assess the effects of temperature on phenology, growth and reproductive performance of multiple populations of slow‐colonizing understorey plants, using the spring flowering geophytic forb Anemone nemorosa and the early summer flowering grass Milium effusum as study species. In both species, emergence time and start of flowering clearly advanced with increasing temperatures. Vegetative growth (plant height, aboveground biomass) and reproductive success (seed mass, seed germination and germinable seed output) of A. nemorosa benefited from higher temperatures. Climate warming may thus increase future competitive ability and colonization rates of this species. Apart from the effects on phenology, growth and reproductive performance of M. effusum generally decreased when transplanted southwards (e.g., plant size and number of individuals decreased towards the south) and was probably more limited by light availability in the south. Specific leaf area of both species increased when transplanted southwards, but decreased with open‐top chamber installation in A. nemorosa. In general, individuals of both species transplanted at the home site performed best, suggesting local adaptation. We conclude that contrasting understorey plants may display divergent plasticity in response to changing temperatures which may alter future understorey community dynamics.     

Phenology, sex ratio, and spatial distribution among dioecious species of Trichilia (Meliaceae)

The flowering, sex ratio, and spatial distribution of four dioecious species of Trichilia (Meliaceae) were studied in a semi-deciduous forest in southeastern Brazil. All reproductive trees (T. clausseni, T. pallida and T. catigua) with dbh > or = 5 cm within a 1-ha plot were collected, sexed, mapped and, for individuals of each species, the distances to the nearest neighbour of the same and opposite sex were measured. For the shrub species T. elegans (dbh < 5 cm), all reproductive individuals were sampled randomly in 10 samples of 10 x 10 m. The reproductive phenology was observed at weekly to monthly intervals from May 1988 to January 1990. The species are strictly dioecious, did not present any sex-mixed trees or sex switching during the study, and sex ratio did not differ significantly from 1 : 1. The size distributions and the relative size variation were not significantly different between sexes. There was no significant segregation or clumping between individuals of either sex and no fruit production without pollination. Onset of flowering and flowering peak were synchronous between male and female plants for all species studied. Flower synchrony was related to outcrossing and pollinator attraction rather than climatic factors.     

Vulnerability and determinants of reproductive success in the narrow endemic Antirrhinum microphyllum (Scrophulariaceae)

The breeding system and flowering phenology of the narrow endemic Antirrhinum microphyllum (Scrophulariaceae) were studied in order to assess the main factors affecting female reproductive success and to identify existing or potential threats to the viability of its populations. Hand-pollination experiments showed that A. microphyllum is an allogamous self-incompatible species. In both populations studied, the flowering season was 4 mo long and mean flowering duration per plant was about 1 mo. Peak flower production took place between mid-April and mid-May and overlapped with the period of activity of the main pollinator, Rhodanthidium sticticum (Megachilidae). Estimated mean number of seeds produced per plant was 9391, showing that population viability is not presently limited by seed output. The study of the direct and indirect effects of plant size, phenological traits (first flowering date, flowering duration, and flowering synchrony), and distance to neighbor plants on reproductive success was performed using structural equation modeling (SEM). In both populations, number of flowers and plant size were the main factors that determined the total number of fruits produced by a plant. First flowering date and flowering synchrony also affected fruit production. Multisample comparison of path coefficients for the two populations rejected the possibility that reproductive patterns could be described by one single model.     

准噶尔无叶豆的开花物候与生殖特征

 为了研究沙漠稀有植物准噶尔无叶豆(Eremosparton songoricum)的开花物候特征及其对生殖成功的影响, 2005和2006年连续两年对其自然种群的开花物候和开花过程中的花部表型变化进行了观察, 并运用相对开花振幅、开花强度和开花同步性等开花物候指数研究了开花物候特征。结果表明: 准噶尔无叶豆在5月下旬至6月中下旬开花, 其种群、个体、花序和单花的花期分别历时26~29 d、8~10 d、5~7 d和2~3 d。单花开花进程依其形态和散粉特征可分为散粉前期、散粉初期、散粉盛期和凋谢期4个时期。其个体水平的开花物候进程(开花振幅曲线)呈渐进式单峰曲线, 具有很高的开花同步指数, 表现出一种集中开花的模式。开花物候指数与座果数之间的相关分析结果表明, 始花日期与花期持续时间存在负相关关系, 而与开花数和座果数存在正相关关系; 花期持续时间与开花数和座果数存在显著正相关关系。准噶尔无叶豆个体开花物候在很大程度上是由其遗传因子决定的, 而开花物候在年度间的变异, 可能是由于荒漠气候的差异(主要是水分和温度的差异)所引起的。作为沙漠窄域分布特有种, 准噶尔无叶豆在环境和人为干扰的双重选择压力下, 为了吸引更多的传粉者访问而达到生殖成功, 形成了大量集中开花的模式。     

Synchronized expression of FLOWERING LOCUS T between branches underlies mass flowering in Fagus crenata

Masting is the intermittent and synchronized production of a large amount of flower and seed in plant populations. This population-level phenomenon is caused by individual-level variability in reproduction and its synchrony between individuals. The variability at the individual level is induced by synchronized reproduction between branches within an individual because a tree is an assemblage of branches that are considered as semiautonomous units. However, there have been no empirical studies that quantify the degree of reproductive synchrony at the branch level within the same tree in masting species. Here, we evaluated the reproductive synchrony within individuals by monitoring flowering dynamics and expression level of a flowering-time gene at the branch-level in a typical masting species, Fagus crenata Blume. The 4-year census showed that the branch-level gene expression was highly variable between years and was strongly synchronized between branches. The branch-level synchrony in flowering-time gene expression was followed by coherent flowering cycle at the whole individual. To examine the causal relationship between gene expression and climatic factors, we performed a nonlinear statistical analysis called convergent cross-mapping using the time course data of gene expression and environmental variables. Our results indicated that the observed gene expression pattern was well cross-mapped by temperature or precipitation. However, this cross-mapping skill was lower than that of randomly generated seasonal dynamics, implying a combination of internal and external environmental signals is more likely to regulate gene expression dynamics in F. crenata. Our results provide the first empirical evidence that synchronized expression of a flowering-time gene between branches underlies integrated flowering behavior at the individual level.     

Sexual vs. vegetative reproduction in relation to forest dynamics in the understorey shrub,Hydrangea paniculata (Saxifragaceae)

In the forest understorey, shrubs usually reproduce vegetatively rather than sexually, but the relative contribution of these two reproductive modes may vary with temporal changes in environmental conditions (e.g., light, substrates of forest floor) that are closely related to forest dynamics. To evaluate the occasional changes in reproductive modes of the clonal understorey shrub Hydrangea paniculata, the reproductive characteristics (i.e., flowering, seedling establishment, vegetative propagation, reproductive size and age) were investigated in four different developmental stages in a Japanese beech forest (early gap, EG; late gap, LG; building, BU; mature, MA). Flowering individuals occurred only in EG and LG, and a much greater number of seedlings was observed in both EG and LG than in BU and MA, mainly because canopy gaps provided sufficient light for flowering and suitable substrates (fallen trees and exposed mineral-soil) for seedling establishment. The flowering individuals originated from plants that had persisted in the shaded understorey until gap formation. In contrast to sexual reproduction, clonal fragmentation increased with forest development in the order LG, EG, BU, MA. This is because, in later stages of forest dynamics (BU and MA), large individuals, some of which had previously flowered in gap stages, were layered on the forest floor and subsequently produced a substantial number of clonal fragments by separation of the buried branches from the main stems which had decayed. This resulted in a large number of clonal fragments originating from a single mother plant. These results suggest that the reproductive modes of H. paniculata is strongly influenced by the changes in environmental conditions with respect to the dynamics of canopy trees. Reproduction from seed in gap stages, despite their short period, provide new genets in established populations, whereas vegetative propagation via fragmentation in closed-canopy stages enhance the stability of the population until the next disturbance.     

Edge influence on plant litter biomass in forest and savanna in the Brazilian cerrado

Edge influence, characterized by differences in ecosystem characteristics between the edge and the interior of remnants in fragmented landscapes, affects a variety of organisms and ecosystem processes. An important feature that may be affected by edges is the amount of plant litter, which provides important habitat for a large variety of organisms and influences ecological processes such as fire dynamics. We studied edge influence on plant litter and fine woody debris in the cerrado of São Paulo state, south‐eastern Brazil. We collected, sorted, dried and weighed plant litter along 180 m‐long transects perpendicular to three savanna and eleven forest edges adjacent to different anthropogenic land uses, with four to five transect per edge. There tended to be less biomass of the finer portions of fine woody debris at both savanna and forest edges. Graminoid litter at savanna edges was greater than in the corresponding interior areas, whereas other litter portions were either unaffected by edges or did not show consistent patterns in either savanna or forest. Edge influence was usually restricted to the first 20 m from the edge, was not influenced by edge characteristics and exhibited no clear differences between savanna and forest areas. Several mechanisms may have led to the variable patterns observed including variation in the plant community, plant architecture, and invasive species. The edge‐related variation in plant litter may putatively lead to, for example, increased fire frequency and intensity at the savanna edges and altered trophic dynamics at forest edges; the mechanisms and consequences of this edge influence should be addressed in future studies.     

The role of individual variation in flowering and pollination in the reproductive success of a crepuscular buzz-pollinated plant

Background and AimsPlant individuals within a population differ in their phenology and interactions with pollinators. However, it is still unknown how individual differences affect the reproductive success of plants that have functionally specialized pollination systems. Here, we evaluated whether plant individual specialization in phenology (temporal specialization) and in pollination (pollinator specialization) affect the reproductive success of the crepuscular-bee-pollinated plant Trembleya laniflora (Melastomataceae).MethodsWe quantified flowering activity (amplitude, duration and overlap), plant–pollinator interactions (number of flowers visited by pollinators) and reproductive success (fruit set) of T. laniflora individuals from three distinct locations in rupestrian grasslands of southeastern Brazil. We estimated the degree of individual temporal specialization in flowering phenology and of individual specialization in plant–pollinator interactions, and tested their relationship with plant reproductive success.Key Results Trembleya laniflora presented overlapping flowering, a temporal generalization and specialized pollinator interactions. Flowering overlap among individuals and populations was higher than expected by chance but did not affect the individual interactions with pollinators and nor their reproductive success. In contrast, higher individual generalization in the interactions with pollinators was related to higher individual reproductive success.ConclusionsOur findings suggest that individual generalization in plant–pollinator interaction reduces the potential costs of specialization at the species level, ensuring reproductive success. Altogether, our results highlight the complexity of specialization/generalization of plant–pollinator interactions at distinct levels of organization, from individuals to populations, to species.     

The Seedling Community of Tropical Rain Forest Edges and Its Interaction with Herbivores and Pathogens1

Edge effects alter biotic interactions and forest regeneration. We investigated whether edge creation affected the seedling community and its interactions with herbivores and leaf‐fungal pathogens. In forest edges and interior sites in Chiapas, Mexico, we counted all woody seedlings and species (10–100 cm tall) present in 1‐m² plots, measured their size (height and leaf number), and examined them for the occurrence of herbivory and pathogen damage. We investigated relationships between levels of damage and size, species richness and density. Species composition and biotic damage varied greatly among sites and habitats (edge vs interior). Late‐successional species dominated the community, but richness was lower in interior sites and species similarity was greater among edges than among interiors. Nearly all species (95%) present at edges and interiors showed herbivory damage, whereas 76 percent of the species in edge plots and 68 percent in interior plots showed pathogen damage. Although leaf area damaged by herbivores was similar between habitats (average 9.2%), pathogen damage was three times greater in edge plots (1.85%) than in interior plots (0.57%). Size was positively related to biotic damage at both habitats. Relationships between herbivory and pathogen damage and between pathogen damage and leaf number were significant only for edges. Biotic damage was not related to density or species richness. Overall, plant community structure was similar between habitats, but biotic damage was enhanced at edges. Thus, disease spread at edges may arise as a threat to tropical rain forest vegetation.     

Edge Structure Determines the Magnitude of Changes in Microclimate and Vegetation Structure in Tropical Forest Fragments1

Edge structure is one of the principal determinants of the extent and magnitude of edge effects in forest fragments. In central Amazonia, natural succession at forest edges typically produces a dense wall of vegetation dominated by Cecropia spp. that buffers the forest interior. Fire encroachment into forest edges, however, eliminates the soil seed bank, enhances plant mortality, and promotes succession to an open, Vismia–dominated edge that does not buffer the forest interior. Contrasting open, fire–encroached forest edges and closed, non–fire–encroached edges were examined in central Amazonia to assess the effects of edge structure on microclimate and vegetation structure in tropical forest fragments. Edge penetration distances for most microclimate and vegetation structure variables were as much as two to five times greater at open edges than at closed edges. The magnitude of these differences suggests that edge structure is one of the main determinants of microclimate and vegetation structure within tropical forest fragments. Edge effects also varied systematically with fragment area. For a given edge type, 100–ha fragments had consistently lower canopy height, higher foliage density, higher temperature, a higher rate of evaporative drying, lower leaf litter moisture content, and lower litter depth than continuous forest, at all distances from the forest edge. These differences, however, were relatively minor compared to the striking differences in edge penetration between open and closed forest edges. For organisms in small fragments, the difference between open and closed edges may be the difference between total edge encroachment on one hand and an effective nature reserve on the other, relatively independent of absolute fragment area.     

Clidemia hirta Invasion at the Pasoh Forest Reserve: An Unexpected Plant Invasion in an Undisturbed Tropical Forest1

Clidemia hirta is a highly invasive shrub in tropical forests throughout the world, but has had little success invading mainland sites and undisturbed forests. In the early 1990s, this plant was found to have invaded an unexpected site, an undisturbed continental tropical forest at Pasoh, Peninsular Malaysia. In 1997, a study was conducted of the C. hirta population at the Pasoh Forest Reserve. A demographic survey of the 50–ha long–term research plot at Pasoh located 1002 C. hirta individuals, 69 of which were reproductive at the time of the study. All but 8 individuals were located in high light gaps or gap edges. Relative growth rates were significantly higher in gaps and gap edges than in the understory, and no reproductive individuals were found in the understory. Mean plant size and dry biomass density increased steadily over the course of the study, while the observed mortality rate was 0 percent over two months. The biomass density of Clidemia at Pasoh was <500 g/ha at the conclusion of this study, but because the population is confined almost exclusively to high light environments, its density in these sites is much higher. The location of C. hirta plants in gaps was significantly correlated with past disturbance by wild pigs, suggesting that soil disturbance and light availability are essential for their establishment. The implication of this study is that by competing with native species in gaps, C. hirta invasion has the potential to alter forest regeneration at Pasoh. Changing land use practices near the reserve have increased the number of wild pigs, and thus the level of disturbance, which may explain the recent success of C. hirta at Pasoh.