Invasive nonindigenous species pose a serious threat to native biodiversity and ecosystem functioning. Understanding how species’ performance varies under conditions in the current and invaded range can help to predict the dynamics of the invading species in its new environment. Plants with the ability to alter growth in response to variation in light conditions may be favored in landscapes that experience frequent disturbance, as these species may be able to exploit a wide range of niches. Seedbank persistence may also play a critical role in successful plant invasion, as extended seed viability may increase the chance of outlasting unfavorable conditions, maintain population genetic diversity, and allow reinvasions. This study investigated seed longevity and the effect of light intensity on germination of wavyleaf basketgrass [Oplismenus undulatifolius (Ard.) Roem. & Schult.], a newly established invasive species in U.S. mid-Atlantic forest understories. Oplismenus undulatifolius seeds were collected across 5 yr from the original site of introduction in Maryland, USA, and stored in standard lab conditions, then subjected to germination trials under four light conditions in a controlled growth chamber. Seeds remained viable for at least 9 yr, and light intensity did not significantly impact seed germination. Our study demonstrates the importance of evaluating environmental and temporal effects on germination traits, because the scope of surveillance in the field may need to be expanded based on new information about environmental tolerance. Long-term monitoring may also be necessary to effectively control invasive plant populations capable of forming a persistent seedbank.

Understanding the mechanisms by which an invasive plant species is able to colonize and successfully expand into native plant communities can help in estimating the potential threat posed by a new invader and predict impacts on community diversity, structure, and function. Wavyleaf basketgrass [Oplismenus undulatifolius (Ard.) P. Beauv.] is a perennial, shade-tolerant grass species that has been recently introduced to the mid-Atlantic United States. Areas invaded by O. undulatifolius typically have low species richness, but it is unknown whether O. undulatifoius actively outcompetes other species or simply thrives primarily in species-poor habitats. This study used a greenhouse experiment to quantify interspecific competition in shade and sun among seedlings of O. undulatifolius; Japanese stiltgrass [Microstegium vimineum (Trin.) A. Camus], an invasive annual grass common in the region; and a mix of three native perennial grass species commonly used in restoring areas invaded by M. vimineum. In this experiment, shade did not significantly affect growth or competitive ability. Interspecific competition irrespective of shade had a negative effect on growth of all species, but O. undulatifolius was affected to a much greater degree than either M. vimineum or the native grass mix. These results suggest that, at least under these conditions, O. undulatifolius is a weak interspecific competitor and may be capable of forming dense monotypic stands only in areas that already have low species diversity. In the mid-Atlantic region, postagricultural legacies and overabundant deer populations, which lead to depauperate understories, may be a major facilitator of O. undulatifolius invasion in forests.

Brassicaceae weeds can be problematic in canola varieties that have not been modified to resist specific broad-spectrum herbicides. The overall objective of this study was to evaluate the potential for increased rapeseed seeding rate as a management strategy for flixweed. To accomplish this objective, a field study was conducted to determine crop seeding rate effects on canopy light transmission and rapeseed yield characteristics, as well as a greenhouse study to determine morphological and photosynthetic responses of flixweed to decreasing irradiance levels. Results from the field study indicated that light transmittance through the canopy decreased linearly as crop seeding rate increased from 1.8 to 9.0 kg ha-1. Increasing crop seeding rate did not influence rapeseed aboveground biomass, seed yield, and harvest index, but negatively affected rapeseed seed oil content in one of two site-years. Greenhouse study results indicated that declining irradiance levels caused reductions in flixweed biomass, root allocation, and photosynthetic light compensation point. Flixweed leaf allocation, foliage area ratio, and specific foliage area increased in response to decreasing irradiance levels. Combined results of field and greenhouse studies suggest that increasing rapeseed seeding rate can suppress flixweed growth while not causing yield penalties from increased intraspecific competition. However, increased rapeseed seeding rate might not be an adequate control strategy on its own because flixweed displays characteristics of a shade-tolerant species.

Common cocklebur, jimsonweed, and velvetleaf were grown with soybeans in the field to determine how soybean interference affects weed canopy architecture. Common cocklebur had more leaves within the soybean canopy than jimsonweed or velvetleaf. At the end of the season, common cocklebur leaf area was distributed evenly below and above the top of the soybean canopy, while nearly all the jimsonweed and velvetleaf leaf area was above the soybean canopy. Common cocklebur exhibited more shade tolerance than jimsonweed or velvetleaf by also maintaining leaves in the shade within the soybean canopy. Differences among these weeds in leaf distribution within the soybean canopy were not related to differences in abscission of the lower leaves but to a differential response of lower axillary buds to soybean shading. Growth from lower axillary buds in jimsonweed and velvetleaf was strongly inhibited by soybean interference, but interference had little effect on lower axillary bud growth in common cocklebur. Axillary bud growth in the lower canopies of both common cocklebur and soybeans, and their similarity in height, caused these plants to compete for the same aboveground niche. However, common cocklebur had more extensive axillary growth along the lower stem than soybeans, which may allow it to compete for resources in this niche more aggressively than soybeans. Velvetleaf and jimsonweed did not share the same aboveground niche with soybeans due to the placement of their leaves above rather than within the soybean canopy. The lower branching characteristics and apparent shade tolerance of common cocklebur may be important factors in the superior competitive ability with soybeans compared to jimsonweed and velvetleaf.

The effects of shade on dry-matter production, leaf area, and biomass partitioning in purple nutsedge (Cyperus rotundus L.) and yellow nutsedge (C. esculentus L.) were determined in a controlled-environment greenhouse with a day/night temperature regime of 32/26 C. For comparison with a full-light treatment, screening provided 40, 70, and 85% shade. Shading significantly reduced height of yellow nutsedge and dry-matter production, leaf-area production, and rhizome and tuber formation of both species. Shading decreased the partitioning of plant biomass into tubers and rhizomes and increased partitioning into leaves. Yellow nutsedge had a higher net-assimilation rate than purple nutsedge at all levels of shading, but the greater leaf-area duration or total amount of leaf area present in purple nutsedge resulted in greater dry-matter production by that species. The responses of the two species showed no differences in shade tolerance.

Brachypodium sylvaticum, a shade-tolerant, forest dwelling, and aggressive invasive grass native to Eurasia, is a noxious weed in California, Oregon, and Washington. This species could cause ecosystem collapse by altering forest fire regimes. To examine interactions with fire, we divided two Willamette National Forest sites into eight units and randomly selected half for treatment with prescribed fire in spring 2011. We assessed the effect of B. sylvaticum on fire (severity and intensity) as well as the effect of fire on B. sylvaticum (cover, seedling emergence, and dispersal). We found that B. sylvaticum cover decreased fire severity but had no effect on intensity. Furthermore, fire severity influenced B. sylvaticum cover; in areas receiving low-severity fire, the grass increased from 21 ± 15.05 to 34 ± 15.81%, but in areas of high-severity fire, cover remained consistently around 0% (0 ± 0% cover in yr 1 to 0.2 5± 0.25% in yr 3). In the field, prescribed fire decreased seedling emergence by 32% compared to controls, but not in an associated greenhouse experiment. However, in the greenhouse, severely burned plots had zero emergence, compared to 0.29 ± 0.14 seedlings low-severity m−2 plot. Fire severity also influenced dispersal in the field; we monitored plots with < 0.5% cover B. sylvaticum initially; when these plots experienced low severity fire, they had greater B. sylvaticum cover (increasing 1,200%), suggesting increased dispersal with less severe fires. High-severity dispersal plots did not experience increased cover. High severity fires have the potential to control the grass, but low-severity fires will likely increase its cover.

We analyzed the recent (< 25 yr) spread in New Hampshire, USA, of the exotic tree Kalopanax septemlobus, native to Asia. The invasion was likely initiated by a single tree planted ca. 1972. Our objective was to assess the viability of the invasion, especially in light of the small propagule size. We tallied, mapped, aged, and measured the height and growth of K. septemlobus individuals at two sites, the University of New Hampshire campus (UC) and Thompson Farm (TF), both in Durham. We found over 3,800 plants at UC and 270 at TF in < 120 ha (296 ac) total area. Plant age ranged from 0 to 22 yr, and UC plants were as far as 775 m (2,543 ft) from the purported parent tree. Annual height growth was comparable to midtolerant native trees. Plants occurred in both open and forested habitats, and the mean level of photosynthetically active radiation incident on understory plants was 4 to 6% of full sun. The large population size, shade tolerance, rapid height growth, and ability to sprout from damaged stems suggest potential for K. septemlobus to invade and persist in forests, the most common natural ecosystem in the northeastern United States. We further suggest that small propagule size, likely a single tree, has not prevented K. septemlobus from initiating a spatially extensive and vigorous population. Kalopanax septemlobus has been planted as an ornamental in the northeastern United States, and prevention of region-wide invasion might depend on removal of these trees, even when they occur as single individuals.

Some species-rich secondary forests in New Caledonia have a monodominant canopy. Here we investigate growth and biomass allocation traits that might explain single-species’ dominance of these post-disturbance stands, and their later decline in the absence of large-scale disturbance. Seedlings of 20 rain-forest trees were grown in two light treatments in a nursery house. In the sun treatment, monodominants grew faster (56.7 ± 1.4 mg g−1 wk−1) than subordinates (40.2 ± 2.6 mg g−1 wk−1). However, some episodically regenerating (ER) subordinates had high growth rates similar to those of monodominants. In the shade treatment, monodominants and subordinates had similar growth rates (33.7 ± 2.6 and 34.0 ± 1.9 mg g−1 wk−1 respectively). Notably, monodominants in both sun and shade treatments had lower root mass fraction (0.29 ± 0.02 and 0.27 ± 0.02 g g−1 respectively) than subordinates (0.39 ± 0.02 and 0.37 ± 0.02 g g−1). Fast growth in sunny conditions is probably imperative for these relatively shade-intolerant ER monodominants. In field conditions, high shoot mass fraction combined with efficient root performance may facilitate faster growth in monodominants competing with other ER species in sunlit sites. Slower growth in shade may contribute to loss of dominance over time in undisturbed forests.

Understanding the ecological function of large frugivores in tropical forests is important considering their central position in seed-dispersal networks. This study illustrates the seed-dispersal effectiveness of a western lowland gorilla (Gorilla gorilla gorilla) population in Cameroon by looking at the interaction of seed quantity, species regeneration traits and the environmental characteristics of the deposition sites (sleeping sites vs. feeding trails) in order to discuss the role of gorillas within the ecosystem. The analysis of 1030 faecal samples collected over a 3-y period (September 2009–August 2012) showed that gorillas dispersed a diverse array of species (58 spp. identified), with an average of 289 intact seeds (>5 mm) d−1 per individual. Seed damage during mouth and gut processing occurred for 12 spp., generally accounting for <5% of specific seed load. Germination trials suggested that germination success of passed seeds remained unchanged (N = 6 spp.) or improved (N = 5 spp.) after gut passage compared with hand-cleaned seeds, with one exception. Indicator species identification (IndVal method) and General Discriminant Analysis suggested that sleeping sites (N = 36), as a main type of seed deposition site, correspond to regenerating microhabitats and correspondingly degree of canopy openness is one variable discriminating significantly this type of deposition site with random locations in the forest (N = 36) used as a proxy for non-directed seed deposition on feeding trails. With a mean canopy openness of 19–27%, sleeping sites offer optimal light conditions for the seedling recruitment of non-pioneer light-demanding species, the seeds of which encompass 47.4% of the total seed load dispersed over the 3-y period. From this study, it is hypothesized that the gorilla performed directed-dispersal of some light-demanding plant species at sleeping sites, and hence, by being actively implicated in gap-phase dynamics, fulfils a valuable ecological function within its ecosystem.

We investigate the effect of patch size on liana diversity and distribution in 19 patches of montane evergreen forest in the Nilgiri hills, Western Ghats, southern India. Additionally, we examined how liana species richness and community assemblage in both edge (within 10 m of the forest edge) and interior regions of forest patches respond to patch size, in order to infer the impact of forest expansion or reduction on the liana communities. A total of 1276 woody liana individuals of 15 species were identified, belonging to 10 genera and nine families. Total species richness of lianas was significantly positively related to forest-patch area, both when analysed for the entire patch, in addition to both core and edge regions when examined separately. Species richness of larger lianas also showed a significant positive relationship with increasing forest patch area. Community assemblage varied with respect to forest edge, with shade-dependent species only occurring in interior patch regions, shade-averse species in edge regions, and shade-tolerant species occurring throughout. Disturbance also played a role in determining the response of liana diversity to patch size, with heavily disturbed patches showing no relationship between patch size and diversity, whereas positive relationships exist in low to moderately disturbed patches. The most significant result is the change in liana community composition between small and larger fragments. Many species present in smaller patches are also present in edge zones of larger fragments. This suggests that lianas are important structural components of montane forest ecosystems, and their compositional patterns are possibly driven by succession. Moreover, this study reveals the importance of edge effect and patch size in influencing liana species richness and compositional patterns.

Abundance and physiology of three understorey tree fern species were compared in a Mexican cloud forest. We hypothesized that the distribution of species would be associated with canopy openness and leaf physiological characteristics. In gullies (1–2% full sun), Alsophila firma was abundant, Cyathea divergens was distributed in moderately open places (4–9%), and Lophosoria quadripinnata preferred more open canopy (9–30%). Although 11 leaf traits of five plants of each species growing under closed and open canopies over 1 y did not differ within species, there were significant interspecific differences. Alsophila firma had comparatively low maximum electron transport rate ETRmax (26.8 ± 1.81 μmol m−2 s−1) and ETR light saturation point (ETRLSP: 261 ± 36.1 μmol m−2 s−1), high specific leaf area (SLA), thin leaves and decreased quantum yield during a leaf desiccation experiment. Cyathea divergens had relatively high maximum quantum yield (0.84 ± 0.004), ETRmax (37.3 ± 1.8 μmol m−2 s−1) and ETRLSP (409 ± 40.0 μmol m−2 s−1). Lophosoria quadripinnata had comparatively thick leaves, low SLA, high predawn water potential, high density (606 ± 25.5 mm−2) and small length (0.026 ± 0.002 mm) stomata. The results support the hypothesis that light sensitivity shapes tree fern distribution in the cloud forest.

The responses of plants to shade and foliar herbivory jointly affect growth rates and community assembly. We grew 600 seedlings of ten species of the economically important Dipterocarpaceae in experimental gradients of shading (0.3–47.0% of full sunlight) and defoliation (0, 25%, 50% or 75% of leaf area removed). We assessed stem diameters initially, after 2 and 4 mo, and calculated relative growth rates (RGR) with a linear model. Shading interacted with defoliation, reducing RGR by 21.6% in shaded conditions and 8.9% in well-lit conditions. We tested three hypotheses for interspecific trade-offs in growth responses to shading and defoliation. They could be positively related, because both reduce a plant's access to carbon, or inversely related because of trade-offs between herbivore resistance and tolerance. We observed, however, that species varied in their response to shading, but not defoliation, precluding an interspecific trade-off and suggesting that plants tolerate shade and herbivory with differing strategies. Shading most strongly reduced the growth of species with less-dense wood and larger seeds. The strong and variable growth responses to shade, contrasted with the weak and uniform responses to defoliation, suggest that variation in light availability more strongly affects the growth of tropical tree seedlings, and thus community assembly, than does variation in herbivory.

Sprouting may play a significant role in maintenance of plant diversity where prevailing disturbance frequency and severity allows. When disturbance frequency and severity decrease, strong sprouters may be outcompeted. As a result, species composition and diversity may change. This study was carried out to investigate the relationship between sprouting, succession and species diversity in a coastal dune forest that currently suffers from low-severity, chronic disturbance due to sea winds and loose sand substrate. Historically, the site was occupied by shifting cultivators who left the site about 80 y ago. Data on trees that were at least 1.3 m tall from 42 sample plots measuring 20 × 20 m were used. The plots were ranked in order of advancement of succession using the first axis of Principal Components Analysis of forest structural variables. Regeneration pattern was examined using analysis of stem diameter frequency distribution. Abundance and regeneration of strong basal sprouters, incidence of basal sprouting and species diversity decreased with advancement of succession. Only a few species could regenerate under the canopy of late-successional sites. Basal sprouts decreased with advancement of succession whereas trunk sprouts increased. These results suggest that maintenance of high species diversity may need a level of disturbance that allows regeneration and maintenance of strong basal sprouters.

An index of canopy closure was used to estimate closure above the crowns of all trees ≥ 10 cm dbh in 11.11 ha of undisturbed lowland tropical forest at La Selva, Costa Rica. To correct for the efFects of tree size on canopy closure, we used the residual of the regression of the canopy closure index on tree size. Analyses were carried out for the 104 species which had ≥ individuals; a total of 3224 trees were included. Nine species were found to have their crowns in significantly more open conditions than expected by chance and five species were found to have their crowns in significantly more closed conditions than expected by chance (P < 0.05). The remaining 90 species (86.5% of the assemblage) were distributed at random with respect to canopy closure, occupying the available light conditions indiscriminately. Species occurring under higher light levels did not show a narrower range of tolerance than did other species. Most species were found to occur over a substantial proportion of the canopy closure continuum present in the stand; overlap among the great majority of species in the assemblage is extensive. The results support the view that tropical forests comprise assemblages of generalist tree species, and raise questions about the classic notions of gap-phase dynamics.

Some large-seeded tree species have cotyledonary reserves that persist for months after seedling establishment. We carried out two screened growing-house experiments with seedlings of Gustavia superba (Lecythidaceae) to test hypotheses proposed to explain why cotyledons are retained. We grew seedlings from large and small seeds in sun and shade to determine if cotyledon reserves supplement photosynthetic carbon gain, and in a second experiment applied defoliation and shoot removal treatments to determine if reserves are allocated to resprout tissue. In each experiment we tracked cotyledonary resource use over time and measured the fraction of seedling biomass allocated to roots and shoots. We found no evidence that light environment, seed size or damage treatment affected the rate of cotyledon resource usage; 20% of the cotyledonary mass remained 9 wk after leaves were fully developed in both sun and shade and 25–30% of the cotyledonary mass remained 6 wk after leaf or shoot removal. Instead, cotyledon reserves appear to be slowly translocated to roots regardless of light environment or seedling damage. Once seedlings are established, lost tissue is replaced using reserves stored in roots; in high light, damaged seedlings had a lower root mass fraction (0.42) than undamaged ones (0.56) when considering the mass of tissue removed and resprout tissue combined. We conclude that cotyledon reserves are important for resprouting during early seedling emergence and establishment, but do not directly contribute to seedling growth or biomass recovery from herbivores at the post-establishment stage. Persistence of cotyledons may ultimately depend on the development of sufficient root mass for reserve reallocation.

Light is a key environmental factor limiting growth and survival of trees in the subcanopy of wet tropical forests (Davies 2001, Thomas 1996). Light availability varies both vertically and horizontally and affects tree height, crown shape and tree architecture (Bongers & Sterck 1998, Sterck & Bongers 2001, Sterck et al. 1999) in addition to growth and survival (Clark & Clark 1992, 2001). Although many studies of tree seedlings and saplings have shown that growth varies significantly with light availability in tropical wet forests (Clark et al. 1993, Iriarte & Chazdon 2005, King 1991, Kohyama 1991, Montgomery & Chazdon 2002, Oberbauer et al. 1988, 1993; Poorter & Werger 1999, Sterck et al. 1999, Welden et al. 1991), few studies have examined these relationships in size classes above 5 cm dbh (Sterck 1999). King et al. (2005) found that annual increment growth of trees in the 8–20-cm dbh size class in two Asian forests was positively dependent on an index of crown light interception, but no direct measurements of light availability were taken in this study. Due to logistical challenges, few direct measurements of light environments above tree crowns have been made in tropical forests (Sterck & Bongers 2001). To our knowledge, no measurements have been made in second-growth forests.

Plants often modify microsite conditions important for seedling establishment. In tropical moist forest, advance regeneration in the form of shade-suppressed seedlings is a major component of regrowth in new gaps. Tree seedlings may be filtered by the composition and structure of the forest understorey. In a lowland forest in Central Panama, we examined light availability, litter accumulation and the seedling community (abundance, diversity and composition) under and away from the canopies of a common subcanopy, clonal palm, Oenocarpus mapora, and tested whether seedling abundance varies as a function of changes in understorey light conditions and leaf litter depth. We found evidence that leaf litter was five times deeper and light availability 27% lower under O. mapora canopy than where it was absent. The probability of finding a seedling in plots under O. mapora canopies was 33% lower than in plots without the palm. Plots under O. mapora canopies also had 49% fewer species of seedlings and those seedlings came from significantly larger seeds than seedlings in plots not under the palm. Overall, seedling density was positively correlated with light availability and negatively correlated with leaf litter depth. A transplant experiment confirmed that for at least one large-seeded species of dicot, seedlings are negatively affected by the reduction in light availability and increase in leaf litter. Under O. mapora, seedlings of Gustavia superba (Lecythidaceae) had lower survivorship and biomass accumulation than away from the palm. Thus, site conditions produced by large, clonal, multi-layered palms such as O. mapora may reduce establishment of small-seeded and shade-intolerant species, potentially affecting species composition in the understorey seedling pool.

Current research efforts to understand the relative invasibility of different plant communities have mostly ignored tropical forests. Only a few studies have treated invasive species in tropical forests, and recent worldwide analyses have not provided clear predictions concerning the relative invasibility of tropical forests. In this review, the extent to which exotic species have invaded tropical forests is summarized and four leading hypotheses to explain the apparently low frequency of invading plants in tropical forests are evaluated. In general, it is found that invasibility positively correlates with human disturbance, and that undisturbed tropical forests harbour few exotic species. To date, there is no evidence to attribute the low invasibility of undisturbed tropical forests to either their high species diversity or their high diversity of functional types. Instead, the low occurrence of exotic species in most tropical forests is most likely due to the fact that the great majority of exotic species that are transported to tropical countries lack specific life history traits, most importantly shade tolerance, necessary for successful invasion of undisturbed tropical forests. Unfortunately, this situation could change in the future with the expected increase in the plantation forestry of high-grade timber combined with common forestry practices that favour the cultivation of exotic species.

An appendix is available for this article which was not included in the print edition of the Journal. The appendix may be found at the rear of the full text, PDF version of the article.

A popular conceptual model asserts that shade tolerance is characterized by morphological and physiological traits that enhance the net rate of carbon capture in low light. We tested this model by quantitatively reviewing growth, leaf lifespan, CO2 exchange and morphological data from 76 studies on woody seedlings grown under conditions of low light. Data were placed into three tolerance categories (intolerant, intermediate, tolerant), two light categories (less than 4% and 4–12%) and two leaf phenology categories (broad-leaved evergreen and winter deciduous). For both evergreen and deciduous groups, intolerant species had traits conferring greater growth potential than tolerant species in both light categories. These traits included greater leaf mass ratio, leaf area ratio, specific leaf area and mass-based photosynthetic rates above light compensation. However, in 0–4% light, growth rates were similar for intolerant and tolerant species, because low light together with higher respiration rates for intolerant species limited the expression of their growth potential differences. Deciduous and evergreen intolerant species were similar in many respects. However, both intermediate and tolerant deciduous species had markedly lower leaf mass ratios and higher root mass ratios than intermediate and tolerant evergreen species. In addition, deciduous species and intolerant evergreens must cope with as much as sixfold higher leaf turnover rates than tolerant evergreen species. Thus, rather than maximizing growth rates in low light, tolerant evergreen species minimize biomass loss through long leaf lifespans and low respiration rates. Tolerant deciduous species also minimize biomass losses by minimizing whole-plant respiration rates but they accomplish low biomass turnover though low leaf mass ratio and not low leaf turnover rates. Furthermore, unlike most tropical evergreens, tolerant deciduous species can gain large fractions of their total growing season carbon during short periods when the overstory is leafless and then allocate this carbon to storage (as reflected by high root mass ratios) rather than new leaves. In conclusion, we found no support for the low-light-enhanced carbon capture model of shade tolerance as viewed strictly from the perspective of physiological growth capacity. This can be explained by the disadvantages to net growth and survival of maintaining a high growth potential at low light, because high growth potential results in greater rates of whole-plant respiration, tissue turnover, herbivory and mechanical damage and in decreased storage. Thus, shade tolerance can be characterized by traits that maximize survival and net growth, where net growth includes losses to all agents.