Leaf litter decomposition rates in degraded and fragmented tropical rain forests of Borneo

Previously extensive tracts of primary rain forest have been degraded by human activities, and we examined how the effects of forest disturbance arising from habitat fragmentation and commercial selective logging affected ecosystem functioning in these habitats by studying leaf litter decomposition rates in litter bags placed on the forest floor. The rain forests of Borneo are dominated by trees from the family Dipterocarpaceae, and we compared leaf litter decomposition rates of three dipterocarp species at eight forest fragment sites (area 3–3529 ha) that had different histories of disturbance pre‐fragmentation: four fragments had been selectively logged prior to fragmentation and four had been formed from previously undisturbed forest. We compared these logged and unlogged forest fragments with sites in continuous forest that had been selectively logged (two sites) and fully protected and undisturbed (two sites). After 120 d, undisturbed continuous forest sites had the fastest rates of decomposition (52% mass loss). Forest fragments formed from unlogged forest (32% mass loss) had faster decomposition rates than logged forest fragments (28% mass loss), but slower rates than continuous logged forest (39% mass loss). Leaves of a light‐demanding species (Parashorea malaanonan) decomposed faster than those of a shade‐tolerant species (Hopea nervosa), but decomposition of all three dipterocarp species that we studied responded similarly to logging and fragmentation effects. Reduced decomposition rates in logged and fragmented forest sites may affect nutrient cycling and thus have detrimental consequences for forest regeneration. Conservation management to improve forest quality should be a priority, particularly in logged forest fragments.     

Low Herbivory among Targeted Reforestation Sites in the Andean Highlands of Southern Ecuador

Insect herbivory constitutes an important constraint in the viability and management of targeted reforestation sites. Focusing on young experimental stands at about 2000 m elevation in southern Ecuador, we examined foliar damage over one season as a function of tree species and habitat. Native tree species (Successional hardwood: Cedrela montana and Tabebuia chrysantha; fast-growing pioneer: Heliocarpus americanus) have been planted among prevailing local landcover types (abandoned pasture, secondary shrub vegetation, and a Pinus patula plantation) in 2003/4. Plantation trees were compared to conspecifics in the spontaneous undergrowth of adjacent undisturbed rainforest matched for height and foliar volume. Specifically, we tested the hypotheses that H. americanus as a pioneer species suffers more herbivory compared to the two successional tree species, and that damage is inversely related to habitat complexity. Overall leaf damage caused by folivorous insects (excluding leafcutter ants) was low. Average leaf loss was highest among T. chrysantha (7.50% ± 0.19 SE of leaf area), followed by H. americanus (4.67% ± 0.18 SE) and C. montana (3.18% ± 0.15 SE). Contrary to expectations, leaf area loss was highest among trees in closed-canopy natural rainforest, followed by pine plantation, pasture, and secondary shrub sites. Harvesting activity of leafcutter ants (Acromyrmex sp.) was strongly biased towards T. chrysantha growing in open habitat (mean pasture: 2.5%; shrub: 10.5%) where it could result in considerable damage (> 90.0%). Insect folivory is unlikely to pose a barrier for reforestation in the tropical Andean mountain forest zone at present, but leafcutter ants may become problematic if local temperatures increase in the wake of global warming.     

Lichen succession on Wilkiea macrophylla leaves

Abstract: Changes in foliicolous lichen populations of the shrub Wilkiea macrophylla in rainforest at Mt Glorious (south-east Queensland) were observed in the field for 3 years. Leaves successively distal from the twig apices were treated as a chronosequence for lichen successional studies. Five species of lichen each colonized more than 50% of the leaves, and leaves successively distal from the apex showed increasing mean species number. Lichen cover rose until the seventh most distal leaf from the first colonized leaf along a twig, reaching 52%. More distal (older) leaves had apparently lower cover. Different lichen species show different instantaneous colonization rates that vary with time. The sequence of lichen colonization and subsequent community composition is best described by a 'Tolerance' model of succession if a single leaf is selected as the unit or with a 'Chronic Disturbance' model if a leaf population is considered.     

A comparative study of aboveground biomass of three Mediterranean species in a post-fire succession

The aboveground biomass of three woody species (Cistus albidus, Quercus coccifera and Pinus halepensis) in two early successional stages (3- and 10-year old) of a post-fire Mediterranean ecosystem was investigated. Among these three species, which belong to the successional series of holm oak (Quercus ilex), C. albidus and Q. coccifera are two dominant shrub species in the garrigue ecosystem and P. halepensis is a pioneer tree species widely represented in the Mediterranean area. The results obtained showed that in monospecific stands, C. albidus and Q. coccifera had a high recovery potential. In the 3-year-old stands, the cover of P. halepensis was only 19.8% for a total biomass of 0.75 ± 0.21 t ha^–1, while the plant cover of C. albidus and Q. coccifera was, respectively, 26% and 85.5% and biomass was 4.72 ± 1.09 and 11.5 ± 0.16 t ha^–1. Only 10 years after fire, the plant cover of C. albidus and Q. coccifera was, respectively, 55% and 100% and total biomass 13.2 ± 1.7 and 35.8 ± 4.7 t ha^–1. The greatest increase in biomass was noted for P. halepensis (29.7 t ha^–1). If mean annual biomass increments are considered, it appears that there was a significant decrease with the stand age for the two shrub species although the tree species showed an increase in productivity. These differing patterns in biomass and productivity of shrub and tree species with stand age provide information on biomass accumulation rates of pioneer species in a Mediterranean succession and their importance in the vegetation dynamics.     

Trade-offs between light interception and leaf water shedding: a comparison of shade- and sun-adapted species in a subtropical rainforest

Species in high-rainfall regions have two major alternative approaches to quickly drain off water, i.e., increasing leaf inclination angles relative to the horizontal plane, or developing long leaf drip tips. We hypothesized that shade-adapted species will have more pronounced leaf drip tips but not greater inclination angles (which can reduce the ability to intercept light) compared to sun-adapted species and that length of leaf drip tips will be negatively correlated with photosynthetic capacity [characterized by light-saturated net photosynthetic rates (A _max), associated light compensation points (LCP), and light saturation points (LSP)]. We tested this hypothesis by measuring morphological and physiological traits that are associated with light-interception and water shedding for seven shade-adapted shrub species, ten sun-adapted understory shrub species, and 15 sun-adapted tree species in a subtropical Chinese rainforest, where mean annual precipitation is around 1,600 mm. Shade-adapted understory species had lower LMA, A _max, LSP, and LCP compared to understory or canopy sun-adapted species; their leaf and twig inclination angles were significantly smaller and leaf drip tips were significantly longer than those in sun-adapted species. This suggests that shade-adapted understory species tend to develop pronounced leaf drip tips but not large leaf inclination angles to shed water. The length of leaf drip tips was negatively correlated with leaf inclination angles and photosynthetic capacity. These relationships were consistent between ordinary regression and phylogenetic generalized least squares analyses. Our study illustrates the trade-offs between light interception and leaf water shedding and indicates that length of leaf drip tips can be used as an indicator of adaptation to shady conditions and overall photosynthetic performance of shrub species in subtropical rainforests.     

Phenology of the shrub strata of successional sub-tropical humid forests of north-eastern India

The phenology of 70 shrub species occurring in 5-, 25- and 60-yr old forest fallows, developed after slash and burn agriculture in north-eastern India, was studied. The early successional shrubs were largely deciduous with predominant leaf fall during the dry winter, while late successional shrubs were evergreen and with a less pronounced leaf fall pattern; the species in the 25-yr old fallow were largely of the leaf-exchanging-evergreen type. Peak leaf production, flowering and fruiting are delayed by one month in the 60-yr old fallow. The phenological separation of the growth types in the three fallows is discussed and related to possible micro-environmental differences.     

Using functional trait diversity to evaluate the contribution of multiple ecological processes to community assembly during succession

Successional chronosequences provide a unique opportunity to study the effects of multiple ecological processes on plant community assembly. Using a series of 0.5 × 0.5 m² plots (n = 30) from five successional sub‐alpine meadow plant communities (ages 3, 5, 9, 12, and undisturbed) in the Qinghai‐Tibetan Plateau, we investigated whether community assembly is stochastic or deterministic for species and functional traits. We tested directional change in species composition, functional trait composition, and then functional trait diversity measured by Rao's quadratic entropy for four traits – plant height, leaf dry matter content, specific leaf area, and seed mass – along two comparable successional chronosequences. We then evaluated the importance of species interactions, habitat filtering and stochasticity by comparing with random communities and partitioning the environmental and spatial components of Rao's quadratic entropy. We found no directional change in species composition, but clear directionality in functional trait composition. None of the abiotic environmental variables (except P) showed linear change with successional age, but soil moisture and nitrogen were positively related to functional diversity within meadows. Functional trait diversity increased significantly with the increase in successional age. Comparison with random communities showed a significant shift from trait divergence in early stages of succession (3‐ and 5‐yr) to convergence in the later stages of succession 9‐, 12‐yr and undisturbed). The relative importance of abiotic variables and spatial structure for functional trait diversity changed in a predictable manner with successional age. Stochasticity at the species level may indicate dispersal limitation, but deterministic effects on functional trait distributions show the role of both habitat effects and biotic interactions.     

Functional traits associated with plant colonizing and competitive ability influence species abundance during secondary succession: Evidence from subalpine meadows of the Qinghai–Tibetan Plateau

It is widely recognized that colonists and competitors dominate early and late succession, respectively, with selected species having different colonizing and competitive abilities. However, it remains unknown whether colonizing and competitive ability can determine species abundance directly over succession. The data for five key functional traits were collected (photosynthesis rate, leaf turgor loss point, leaf proline content, seed mass, and seed germination rate), which are direct indicators of plant competitive and colonizing abilities including growth, drought and cold stress resistance, dispersal, and seed dormancy. Here, we tested the effects of colonizing and competitive abilities on species abundance, by employing a linear mixed‐effects model to examine the shifts in the relationship between species abundance and these five colonization and competition‐related traits in species‐rich subalpine secondary successional meadows (at 4, 6, 10, 13 years of age, and undisturbed, respectively) of the Qinghai–Tibetan Plateau. The abundant species at the early‐successional meadows tend to have high photosynthetic rate, high leaf proline content, low seed mass, and seed germination rate for having high colonizing ability, but low competitive ability. By contrast, late‐successional communities tend to be dominated by species with high competitive ability, but low colonizing ability, indicated by large seeds, high seed germination rate, low photosynthetic rate, and leaf proline content. The observed directional shifts in the relationships between traits (photosynthetic rate, leaf proline content, seed mass, and seed germination rate) and abundance with successional age, bring two new understandings of community assembly during succession of subalpine meadows in the Qinghai–Tibetan Plateau. First, it discloses that the differences in species abundance over succession can be directly attributed to differences in colonizing and competitive abilities of different species. Second, it expands the effects of multiple life historical differences including growth, resource competitive ability, cold stress resistance, dispersal, and seed germination strategy, represented by functional traits on community assembly along succession, that is, from the species to the community level.     

岷江冷杉林皆伐后次生群落结构和物种多样性的演替动态

为阐明岷江冷杉林皆伐后次生群落的演替过程,采用空间代替时间的方法,在川西米亚罗林区海拔3100—3600 m的阴坡选择岷江冷杉林皆伐后次生演替10、20、30、40和50a阶段的次生群落作为研究对象,对其群落结构和物种多样性的动态进行了研究。不同演替阶段的树木均呈显著聚集分布。按群落中优势种的重要值将该演替序列划分为3个类型:悬钩子-蔷薇灌丛、白桦阔叶林和桦木-岷江冷杉针阔混交林。随次生演替,乔木和灌木种的物种丰富度趋于增加,而草本种的物种丰富度趋于减少;乔木和灌木种的Shannon-Wiener多样性指数趋于增大,而草本的Shannon-Wiener多样性指数趋于减小;乔木、灌木和草本层的Pielou均匀度指数均趋于增大;乔木层的Simpson优势度指数趋于减小,灌木和草本层的Simpson优势度指数在演替0—40年阶段趋于增大,而在演替50a阶段趋于减小。在该演替序列中,乔木、灌木和草本层的物种组成均呈耐荫种替代非耐荫种的趋势。     

Current reclamation practices after oil and gas development do not speed up succession or plant community recovery in big sagebrush ecosystems in Wyoming

Reclamation is an application of treatment(s) following disturbance to promote succession and accelerate the return of target conditions. Previous studies have framed reclamation in the context of succession by studying its effectiveness in reestablishing late‐successional plant communities. Reestablishment of plant communities is especially important and challenging in drylands such as shrub steppe ecosystems where succession proceeds slowly. These ecosystems face threats from climate change, invasive species, altered fire regimes, and land‐use change, as well as fossil‐fuel extraction and associated disturbance. As such, the need for effective reclamation after this type of energy development is great. However, past research regarding this type of reclamation has focused on mining rather than oil and gas development. To better understand the effect of reclamation on rates of succession in dryland shrub steppe ecosystems, we sampled oil and gas wellpads and adjacent undisturbed big sagebrush plant communities in Wyoming, U.S.A., and quantified the extent of recovery for forbs, grasses, and shrubs on reclaimed and unreclaimed wellpads relative to undisturbed plant communities. Reclamation increased the recovery rate for early‐successional types, including combined forbs and grasses and perennial grasses, but did not affect recovery rate of late‐successional types, particularly big sagebrush and perennial forbs. Rather, subsequent analyses showed that recovery of late‐successional types was affected by soil texture and time since wellpad abandonment. This is consistent with studies in other ecosystems where reclamation has been implemented, suggesting that reclamation may not help reestablish late‐successional plant communities more quickly than they would reestablish naturally.     

Nocturnal primates of kibale forest: Effects of selective logging on prosimian densities

To evaluate the potential impact of logging on nocturnal primates, densities ofGalago demidovii, G. inustus, andPerodictus potto were determined in two areas of the Kibale Forest Reserve, Uganda. One of these areas had been logged at a moderate level in the late 1960's, while the second area was relatively undisturbed. The density of the nocturnal primates was lower in the logged area than in the neighboring unlogged area. For the most frequently sighted nocturnal primate,G. demidovii, sightings were spatially clustered both within a year and between years, suggesting that clumped resources and/or social factors were influencing space use.     

Phyllostomid bat assemblages in different successional stages of tropical rain forest in Chiapas,Mexico

Due to their role in seed dispersal, changes in the community of phyllostomid bats have direct consequences on ecological succession. The objective of this work was to document changes in the structure of bat assemblages among secondary successional stages of tropical rain forest in Chiapas, Mexico. Bats were mist-netted at ground level during 18 months in 10 sites belonging to 3 successional stages: four sites represented early succession (2–8 years of abandonment), four intermediate succession (10–20 years of abandonment), and two late succession (mature old-growth forest).We captured 1,179 phyllostomids comprising 29 species. Phyllostomid species richness was 17 (58% of all species) in the early stage, 18 (62%) in the intermediate stage and 24 (83%) in the late stage. The late successional mature forest possessed nine species that were exclusively found there, whereas early and intermediate successional stages contained only one exclusive species. Sturnira lilium, Artibeus lituratus, Carollia perpicillata, Artibeus jamaicensis and Glossophaga soricina represented 88% of all captured phyllostomid bats. Frugivores made up more than 90% of the species captured in early and intermediate successional stages and 84% in late successional forest. The Bray–Curtis index of dissimilarity showed a replacement of species through successional stages with the largest dissimilarity between early and late stages, followed by intermediate and late, and the lowest dissimilarity between early and intermediate stages. The number of gleaning insectivore species increased during succession. The carnivorous guild was exclusively found in the late stage (three species). We conclude that the late successional mature forest was the main reservoir for the gleaning insectivore and carnivore guilds; however, early and intermediate successional stages possessed a great diversity of species including many frugivores.     

洮河自然保护区大峪沟林区紫果云杉群落木本植物种群生态位特征

基于群落调查数据,以样地代表多种资源的综合状态,物种重要值作为生态位的计测指标,利用Levins和Shannon生态位宽度,Schoener生态位相似和Pianka生态位重叠对洮河自然保护区大峪沟林区的紫果云杉群落木本植物种群生态位进行定量研究。结果表明：紫果云杉群落乔木层共有树种8种,隶属于4科6属,紫果云杉占绝对优势,岷江冷杉为次优势种;灌木层共有植物50种,隶属于16科25属,忍冬科的蓝靛果、葱皮忍冬和唐古特忍冬等以及蔷薇科的峨眉蔷薇和银露梅等构成了灌木层的主体,紫果云杉幼苗优势不明显,但生态位宽度最大。本研究表明,生态位宽度受物种生态特性及其分布和生境共同影响,重要值大的物种生态位宽度一般较大,生态位宽度大的物种之间生态位相似性和重叠性一般较大,并与其它物种重叠的概率高、重叠程度大,生态位宽度小的物种集中出现时也会出现较大的生态位重叠,同科或同属的物种间常有较大生态位相似和重叠。生态位宽度反映着种群的扩散潜力,生态位相似反映着群落的稳定性,生态位重叠反映着物种间存在或潜在的竞争程度。紫果云杉在乔、灌木层均有最大的生态位宽度,生态适应性最强,群落结构稳定,具有进一步扩散的可能。     

Tropical forests are thermally buffered despite intensive selective logging

Tropical rainforests are subject to extensive degradation by commercial selective logging. Despite pervasive changes to forest structure, selectively logged forests represent vital refugia for global biodiversity. The ability of these forests to buffer temperature‐sensitive species from climate warming will be an important determinant of their future conservation value, although this topic remains largely unexplored. Thermal buffering potential is broadly determined by: (i) the difference between the “macroclimate” (climate at a local scale, m to ha) and the “microclimate” (climate at a fine‐scale, mm to m, that is distinct from the macroclimate); (ii) thermal stability of microclimates (e.g. variation in daily temperatures); and (iii) the availability of microclimates to organisms. We compared these metrics in undisturbed primary forest and intensively logged forest on Borneo, using thermal images to capture cool microclimates on the surface of the forest floor, and information from dataloggers placed inside deadwood, tree holes and leaf litter. Although major differences in forest structure remained 9–12 years after repeated selective logging, we found that logging activity had very little effect on thermal buffering, in terms of macroclimate and microclimate temperatures, and the overall availability of microclimates. For 1°C warming in the macroclimate, temperature inside deadwood, tree holes and leaf litter warmed slightly more in primary forest than in logged forest, but the effect amounted to <0.1°C difference between forest types. We therefore conclude that selectively logged forests are similar to primary forests in their potential for thermal buffering, and subsequent ability to retain temperature‐sensitive species under climate change. Selectively logged forests can play a crucial role in the long‐term maintenance of global biodiversity.     

Tree–Shrub Interactions During Early Secondary Forest Succession in Uganda

Abstract One of the greatest challenges for ecologists this century will be restoring forests on degraded tropical lands. This restoration will require understanding complex processes that shape successional pathways, including interactions between trees and other plants. Shrub species often quickly invade disturbed tropical lands, yet little is known about whether they facilitate or inhibit subsequent tree recruitment and growth. We examined how shrubs and other vegetation (e.g., vines, grasses, herbs) affect tree recruitment, survival, and growth during the first 6 years of forest succession in Kibale National Park, Uganda. The study was undertaken in two recently logged exotic softwood plantations. We studied the successional trajectories in two recently logged areas that varied in their initial densities of trees and shrubs. Analyses suggested tree seedling presence and density were not strongly related to shrub density or height during succession. Tree sapling presence and density were positively significantly related to shrub density and height. We found little response in the tree community to experimental shrub removal, and although removal of all nontree vegetation temporarily enhanced tree growth, the effect disappeared after 2 years. Some early‐successional trees benefited from reduced competition, whereas some mid‐successional trees benefited from the presence of other vegetation. Some specific tree species responded strongly to vegetation removal. We interpret our findings in light of designing manipulations promoting forest restoration for biodiversity conservation and conclude with four tentative guidelines: (1) manage at the species level, not the community level; (2) increase facilitation for seedlings, reduce competition for saplings; (3) be cautious of assumptions about plant interactions; and (4) be adaptable and creative with new strategies when manipulations fail.     

Effects of Historical Selective Logging on Anuran Communities in a Wet Evergreen Forest,South India

Vast areas of tropical evergreen forests have been selectively logged in the past, and many areas continue to be logged. The impacts of such logging on amphibians are poorly understood. I examined the response of anuran communities to historical selective logging in a wet evergreen forest in south India. Anuran assemblages in unlogged forest were compared with assemblages in selectively logged forest. Forty 10 m × 10 m quadrats in forest, riparian zones, and streams of unlogged and selectively logged forests were searched at night for anurans. Species richness did not appear to be affected by logging. However, anuran density varied significantly and was 42 percent lower in selectively logged forests compared to unlogged forests. Anuran densities also varied significantly across microhabitats, with highest densities in streams of both selectively logged and unlogged forests. Patterns of niche overlap varied with selective logging as niche breadth either expanded, contracted, or remained neutral for different species. Ordination analysis explained 95 percent of the variation in species assemblage across selectively logged and unlogged forests. The assemblage in selectively logged forest was nested within unlogged forest. Among the habitat characteristics, litter thickness and water depth had the highest influence on the assemblage. This was followed by litter/water temperature, air temperature, and lastly relative humidity. It appears that species richness and composition of anurans in selectively logged forests is converging with unlogged forests, but the effects of historical logging seem to persist on anuran densities and their niche characteristics even ca 40 yr since logging ceased.     

Identifying Native Vegetation for Reducing Exotic Species during the Restoration of Desert Ecosystems

There is currently much interest in restoration ecology in identifying native vegetation that can decrease the invasibility by exotic species of environments undergoing restoration. However, uncertainty remains about restoration's ability to limit exotic species, particularly in deserts where facilitative interactions between plants are prevalent. Using candidate native species for restoration in the Mojave Desert of the southwestern U.S.A., we experimentally assembled a range of plant communities from early successional forbs to late‐successional shrubs and assessed which vegetation types reduced the establishment of the priority invasive annuals Bromus rubens (red brome) and Schismus spp. (Mediterranean grass) in control and N‐enriched soils. Compared to early successional grass and shrub and late‐successional shrub communities, an early forb community best resisted invasion, reducing exotic species biomass by 88% (N added) and 97% (no N added) relative to controls (no native plants). In native species monocultures, Sphaeralcea ambigua (desert globemallow), an early successional forb, was the least invasible, reducing exotic biomass by 91%. However, the least‐invaded vegetation types did not reduce soil N or P relative to other vegetation types nor was native plant cover linked to invasibility, suggesting that other traits influenced native‐exotic species interactions. This study provides experimental field evidence that native vegetation types exist that may reduce exotic grass establishment in the Mojave Desert, and that these candidates for restoration are not necessarily late‐successional communities. More generally, results indicate the importance of careful native species selection when exotic species invasions must be constrained for restoration to be successful.     

Interspecific variation of plant traits associated with resistance to herbivory among four species of Ficus (moraceae)

• Background and aims To understand the defensive characteristics of interspecies varieties and their responses to herbivory damage, four species of Ficus plants (Ficus altissima, F. auriculata, F. racemosa and F. hispida) were studied. They were similar in life form, but differed in successional stages. Of these, Ficus altissima is a late successional species, F. hispida is a typical pioneer and F. auriculata and F. racemosa are intermediate successional species. We addressed the following questions: (1) What is the difference in plant traits among the four species and are these traits associated with differences in herbivory damage levels? (2) What is the difference in the damage-induced changes among the four species?• Methods Herbivory damage was measured in the field on randomly planted seedlings of the four species of the same age. Defences to herbivory were also tested by feeding leaves of the four species to larvae of Asota caricae in the laboratory. A total of 14 characters such as water content, thickness, toughness, pubescence density on both sides, leaf expansion time, lifetime and the contents of total carbon (C), nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg) and calcium (Ca) were measured. Leaf calcium oxalate crystal (COC) density, total Ca and N content, leaf toughness and height were measured to investigate induced responses to artificial herbivory among the four species.• Key results and conclusions Herbivory damage in the four studied species varied greatly. The pioneer species, F. hispida, suffered the most severe herbivory damage, while the late successional species, F. altissima, showed the least damage. A combination of several characteristics such as high in content of N, Ca and P and low in leaf toughness, lifetime and C : N ratio were associated with increased herbivore damage. The late successional species, F. altissima, might also incorporate induced defence strategies by means of an increase in leaf COC and toughness.Key words: Calcium oxalate crystals, defensive characteristics, Ficus; herbivory, induced defence     

Responses to nitrogen pulses and growth under low nitrogen availability in invasive and native tree species with differing successional status

Invasive species are frequently found in recently disturbed sites. To examine how these disturbance-dependent invasive species exploit resource pulses resulting from disturbance, twelve physiological and morphological traits, including age-dependent responsiveness in leaf traits to nitrogen pulse, were compared between Bischofia javanica, an invasive tree species in Ogasawara islands, and three native Ogasawara species, each having a different successional status. When exposed to a nitrogen pulse, invasive B. javanica showed higher increases in photosynthetic capacity, leaf area, epidermal cell number and cell size in leaves of broad age classes, and root nitrogen absorption ability than two native mid-/late or late-successional species, but showed no particular superiority to a native pioneer species in these responses. Under low nitrogen, however, it showed the largest relative growth rate among the four species, while the native pioneer showed the lowest growth. From these results, we concluded that the combination of moderately high responsiveness to resource pulses and the ability to maintain steady growth under resource limitations may give B. javanica a competitive advantage over a series of native species with different successional status from early to late-successional stages.     

Exotic species as modifiers of ecosystem processes: Litter decomposition in native and invaded secondary forests of NW Argentina

Invasions of exotic tree species can cause profound changes in community composition and structure, and may even cause legacy effect on nutrient cycling via litter production. In this study, we compared leaf litter decomposition of two invasive exotic trees (Ligustrum lucidum and Morus sp.) and two dominant native trees (Cinnamomum porphyria and Cupania vernalis) in native and invaded (Ligustrum-dominated) forest stands in NW Argentina. We measured leaf attributes and environmental characteristics in invaded and native stands to isolate the effects of litter quality and habitat characteristics. Species differed in their decomposition rates and, as predicted by the different species colonization status (pioneer vs. late successional), exotic species decayed more rapidly than native ones. Invasion by L. lucidum modified environmental attributes by reducing soil humidity. Decomposition constants (k) tended to be slightly lower (−5%) for all species in invaded stands. High SLA, low tensile strength, and low C:N of Morus sp. distinguish this species from the native ones and explain its higher decomposition rate. Contrary to our expectations, L. lucidum leaf attributes were similar to those of native species. Decomposition rates also differed between the two exotic species (35% higher in Morus sp.), presumably due to leaf attributes and colonization status. Given the high decomposition rate of L. lucidum litter (more than 6 times that of natives) we expect an acceleration of nutrient circulation at ecosystem level in Ligustrum-dominated stands. This may occur in spite of the modified environmental conditions that are associated with L. lucidum invasion.