Ecological outcomes of agroforests and restoration 15 years after planting

Large‐scale forest restoration relies on approaches that are cost‐effective and economically attractive to farmers, and in this context agroforestry systems may be a valuable option. Here, we compared ecological outcomes among (1) 12–15‐year‐old coffee agroforests established with several native shade trees, (2) 12–15‐year‐old high‐diversity restoration plantations, and (3) reference old‐growth forests, within a landscape restoration project in the Pontal do Paranapanema region, in the Atlantic Forest of southeastern Brazil. We compared the aboveground biomass, canopy cover, and abundance, richness, and composition of trees, and the regenerating saplings in the three forest types. In addition, we investigated the landscape drivers of natural regeneration in the restoration plantations and coffee agroforests. Reference forests had a higher abundance of trees and regenerating saplings, but had similar levels of species richness compared to coffee agroforests. High‐diversity agroforests and restoration plantations did not differ in tree abundance. However, compared to restoration plantations, agroforests showed higher abundance and species richness of regenerating saplings, a higher proportion of animal‐dispersed species, and higher canopy cover. The abundance of regenerating saplings declined with increasing density of coffee plants, thus indicating a potential trade‐off between productivity and ecological benefits. High‐diversity coffee agroforests provide a cost‐effective and ecologically viable alternative to high‐diversity native tree plantations for large‐scale forest restoration within agricultural landscapes managed by local communities, and should be included as part of the portfolio of reforestation options used to promote the global agenda on forest and landscape restoration.     

天然林重建过程中单优箬叶竹灌丛对树木更新的影响

利用1996年在湖南省会同林区布置的20个1 hm~2的实验样地,以剔除或林下不含箬叶竹样地作为对照(C),保留箬叶竹作为处理(I),评估了天然林重建过程中林下单优箬叶竹灌丛形成对树种更新过程的影响.处理10a以后结果显示,箬叶竹灌丛盖度上升到95%.进一步通过2007年两种样地乔木多度、多样性、生物量以及物种组成等指标的测量表明,单优箬叶竹灌丛显著抑制了树种的存活与生长,18～390 cm高度级别的树种多度和多样性显著降低,尽管高度18 cm以下级别的幼苗以及成年树的多度和多样性没有显著下降,但是生物量从幼苗到成年树,处理样地普遍下降.幼苗、幼树、成年树地上部分生物量分别下降47.45%、39.87%、59.64%,地下部分生物量分别下降55.70%、46.08%、63.62%.NMS(非计量多元尺度法)排序进一步表明两种样地具有显著不同的树种组成,揭示两种样地处于不同的演替更新阶段.结果表明单优箬叶竹灌丛的形成能延缓树种更新的进程,因此天然林重建过程中应注意避免该类林下植被的出现.     

Tree recovery and seed dispersal by birds: Comparing forest, agroforestry and abandoned agroforestry in coastal Ecuador

We used a highly replicated study to examine vegetation characteristics between patches of intervened forest, abandoned agroforestry systems with coffee and actively managed agroforestry systems with coffee in a tropical landscape. In all habitats, plant structural characteristics, individual abundance, species richness and composition were recorded for the three plant size classes: adult trees, saplings and seedlings. Furthermore, bird species richness and composition, and seeds dispersed by birds were recorded. Tree abundance was higher in forest habitats while saplings and seedlings were more abundant in abandoned coffee sites. Although species richness of adult trees was similar in the three habitats, species richness of saplings and seedlings was much higher in forest and abandoned coffee than in managed coffee sites. However, in spite of their relatively low species richness, managed coffee sites are an important refuge for tree species common to the almost disappeared mature forest in the area. Floristic similarity for adult trees was relatively low between land use types, but clearly higher for seedlings, indicating homogenizing processes at the landscape level. More than half of the saplings and seedling were not represented by adults in the canopy layer, suggesting the importance of seed dispersal by birds between habitats. Our results show that each of the studied ecosystems plays a unique and complementary role as seed source and as habitat for tree recovery and tree diversity.     

Phylogenetic Structure of Tree Species across Different Life Stages from Seedlings to Canopy Trees in a Subtropical Evergreen Broad-Leaved Forest

Investigating patterns of phylogenetic structure across different life stages of tree species in forests is crucial to understanding forest community assembly, and investigating forest gap influence on the phylogenetic structure of forest regeneration is necessary for understanding forest community assembly. Here, we examine the phylogenetic structure of tree species across life stages from seedlings to canopy trees, as well as forest gap influence on the phylogenetic structure of forest regeneration in a forest of the subtropical region in China. We investigate changes in phylogenetic relatedness (measured as NRI) of tree species from seedlings, saplings, treelets to canopy trees; we compare the phylogenetic turnover (measured as β_NRI) between canopy trees and seedlings in forest understory with that between canopy trees and seedlings in forest gaps. We found that phylogenetic relatedness generally increases from seedlings through saplings and treelets up to canopy trees, and that phylogenetic relatedness does not differ between seedlings in forest understory and those in forest gaps, but phylogenetic turnover between canopy trees and seedlings in forest understory is lower than that between canopy trees and seedlings in forest gaps. We conclude that tree species tend to be more closely related from seedling to canopy layers, and that forest gaps alter the seedling phylogenetic turnover of the studied forest. It is likely that the increasing trend of phylogenetic clustering as tree stem size increases observed in this subtropical forest is primarily driven by abiotic filtering processes, which select a set of closely related evergreen broad-leaved tree species whose regeneration has adapted to the closed canopy environments of the subtropical forest developed under the regional monsoon climate.     

Composition and Dynamics of Functional Groups of Trees During Tropical Forest Succession in Northeastern Costa Rica

We compared the functional type composition of trees ≥10 cm dbh in eight secondary forest monitoring plots with logged and unlogged mature forest plots in lowland wet forests of Northeastern Costa Rica. Five plant functional types were delimited based on diameter growth rates and canopy height of 293 tree species. Mature forests had significantly higher relative abundance of understory trees and slow-growing canopy/emergent trees, but lower relative abundance of fast-growing canopy/emergent trees than secondary forests. Fast-growing subcanopy and canopy trees reached peak densities early in succession. Density of fast-growing canopy/emergent trees increased during the first 20 yr of succession, whereas basal area continued to increase beyond 40 yr. We also assigned canopy tree species to one of three colonization groups, based on the presence of seedlings, saplings, and trees in four secondary forest plots. Among 93 species evaluated, 68 percent were classified as regenerating pioneers (both trees and regeneration present), whereas only 6 percent were classified as nonregenerating pioneers (trees only) and 26 percent as forest colonizers (regeneration only). Slow-growing trees composed 72 percent of the seedling and sapling regeneration for forest colonizers, whereas fast-growing trees composed 63 percent of the seedlings and saplings of regenerating pioneers. Tree stature and growth rates capture much of the functional variation that appears to drive successional dynamics. Results further suggest strong linkages between functional types defined based on adult height and growth rates of large trees and abundance of seedling and sapling regeneration during secondary succession.
Abstract in Spanish is available at http://www.blackwell-synergy.com/loi/btp     

The potential of small exclosures in assisting regeneration of coffee shade trees in South‐Western Ethiopian coffee forests

Ethiopian Afromontane moist forests where coffee grows as understorey shrub are traditionally managed by the local communities for coffee production through thinning of the shade tree canopy and slashing of competing undergrowth. This management practice has a negative impact on the coffee shrubs, because the removal of shade tree saplings and seedlings reduces the succession potential of the shade tree canopy, which threatens the very existence of the shade coffee production system. We assessed the functionality of small exclosures to initiate coffee shade tree canopy restoration through natural regeneration. Our results show that small exclosures have a strong restoration potential for the coffee shade trees preferred by farmers (Albizia schimperiana, A. gummifera and Millettia ferruginea), as evidenced from their seedling abundance, survival and growth. The regeneration of late‐successional tree species of the moist Afromontane forest was not successful in the small exclosures, most probably due to the low abundance or absence of adult trees as seed sources for regeneration. Therefore, temporary establishment of small exclosures in degraded coffee forest fragments where shade trees are getting old or dying is recommended for sustainable shade coffee production.     

Gap regeneration of major tree species in different forest types of Japan

Gap regeneration of major tree species was examined, based on the pattern of gap phase replacement, in primary old-growth stands of warm-temperate, cool-temperate and subalpine forests, Japan. Using principal component analysis, the gap-regeneration behavior of major tree species could be divided into three guilds and that of Fagus crenata (monodominant species of cool-temperate forests). The criteria used for this division were total abundance of canopy trees and regenerations and relative abundance of regenerations to canopy trees. The gap-regeneration behavior of species in the first guild was that canopy trees regenerate in gaps from seedlings or saplings recruited before gap formation; they had higher total abundance and more abundant regenerations relative to their canopy trees. The gap-regeneration behavior of F. crenata was same as species in the first guild, but F. crenata had less abundant regenerations relative to its canopy trees. Species in the second guild had lower total abundance and less abundant regenerations to their canopy trees. The guild contained species whose canopy trees regenerate in gaps from seedlings or saplings recruited after gap formation or regenerate following largescale disturbance. The third guild consisted of species with lower total aboundance and more abundant regenerations relative to their canopy trees. The gap-regeneration behavior of some species in this guild was that trees regenerate in gaps from seedlings or saplings recruited before gap formation, and grow, mature, and die without reaching the canopy layer, while the gap-regeneration behavior of other species was same as that of species in the first guild or F. crenata. Major tree species of subalpine forests were not present in the third guild.     

The influence of tree canopies and elephants on sub-canopy vegetation in a savannah

The apparent influence of elephants on the structure of savannahs in Africa may be enhanced by management activities, fire and other herbivores. We separated the effect elephants have on grasses, woody seedlings (<0.5 m) and saplings (0.5–2 m) from the effect of tree canopies (canopy effect), and herbivory (park effect). We defined the canopy effect as the differences between plant abundances and diversity indices under tree canopies and 20 m away from these. Our testing of the park effect relied on the differences in the sub-canopy plant indices inside and outside a protected area that supported a range of herbivores. We based our assessment of the elephant effect on sub-canopy vegetation indices associated with elephant induced reductions in tree canopies. The park and canopy effects were more pronounced than the elephant effect. The park effect suppressed the development of woody seedlings into saplings. Conditions associated with tree canopies benefited woody plants, but not the grasses, as their indices were lower under trees. Elephants reducing canopies facilitated grass species tolerant of direct solar radiation. We concluded that management should consider other agents operating in the system when deciding on reducing the impact that elephants may have on vegetation.     

Cultivation of Non‐timber Forest Products Alters Understory Light Availability in a Humid Tropical Forest in Mexico1

The planting of non‐timber forest products (NTFPs) in the understory of tropical forests is promoted in many regions as a strategy to conserve forested lands and meet the economic needs of rural communities. While the forest canopy is left intact in most understory plantations, much of the midstory and understory vegetation is removed in order to increase light availability for cultivated species. We assessed the extent to which the removal of vegetation in understory plantations of Chamaedorea hooperiana Hodel (Arecaceae) alters understory light conditions. We also examined how any changes in light availability may be reflected by changes in the composition of canopy tree seedlings regenerating in understory plantations. We employed a blocked design consisting of four C. hooperiana plantation sites; each site was paired with an adjacent, unmanaged forest site. Hemispherical canopy photographs were taken and canopy tree seedlings were identified and measured within 12 3 × 2 m randomly placed plots in each site for a total of 96 plots (4 blocks × 2 sites × 12 plots). Plantation management did not affect canopy openness or direct light availability but understory plantations had a higher frequency of plots with greater total and diffuse light availability than unmanaged forest. Comparisons of canopy tree seedling composition between understory plantations and unmanaged forest sites were less conclusive but suggest that management practices have the potential to increase the proportion of shade‐intolerant species of tree seedlings establishing in plantations. Given the importance of advanced regeneration in gap‐phase forest dynamics, these changes may have implications for future patterns of succession in the areas of forest where NTFPs are cultivated.     

大明山常绿阔叶林冠层垂直结构与林下植物更新

2009—2011年,在广西大明山常绿阔叶林一个典型坡面的上坡、中坡和下坡分别建立24个20 m×20 m的固定样地,测定不同林冠层次(上层、中层和下层冠高分别为:>8、4～8和0～4 m)的覆盖度并监测林下植物的更新,研究林冠垂直结构的动态变化及其对林下植物更新的影响.结果表明: 随着2008年特大冰冻灾害后森林的恢复,林冠总覆盖度显著增加,从2009年的54.0%提高到2011年的67.4%,不同冠层覆盖度和恢复增长存在显著差异,上层林冠覆盖度显著高于中、下层,而中、下层林冠的恢复增长显著优于上层林冠.林下更新的木本植物共55种,优势科和优势种与现有群落的基本一致.同一年份不同坡位林下更新植物的物种多样性指数差异不显著,但同一坡位不同年份间存在显著差异.不同冠层的覆盖度与更新植物的物种丰富度和多度相关性不显著;中层、下层和林冠总覆盖度与林下更新植物的Shannon指数、Simpson指数和Pielou均匀度指数呈显著负相关,表明冠层覆盖度显著影响着林下更新,中、下层林冠对林下更新植物多样性的作用更明显.     

Density-dependent shoot-borer herbivory increases the age of first reproduction and mortality of neotropical tree saplings

Shoot herbivory by the sapling specialist shoot-borer Cromarcha stroudagnesia (Lepidoptera, Pyralidae, Chrysauginae) is shown to have large direct and indirect effects on the rates of height increment and mortality of saplings of its host tree, Tabebuia ochracea (Bignoniaceae), in the secondary successional tropical dry forests of the Area de Conservación Guanacaste, Costa Rica. Experiments and field observations over 3-4 years show a substantial reduction in sapling height increments due to C. stroudagnesia herbivory, of equivalent magnitude to the difference in height increments between undamaged saplings in canopy gaps and full understorey shade. Extrapolating this data at average amounts of C. stroudagnesia herbivory increases the duration of the pre-reproductive sapling life stage by about 40% relative to undamaged plants. This is an underestimate, as top shoot herbivory by C. stroudagnesia also increased the probability of canopy gap saplings being overtopped and shaded by surrounding vegetation. Sapling mortality was increased by C. stroudagnesia herbivory, with 11.8% of the most heavily damaged young saplings dying in 3 years while no undamaged saplings died. Cromarcha stroudagnesia herbivory strongly increases with the number of conspecific T. ochracea saplings and the number of conspecific shoots within 50 m of focal saplings. It is therefore likely to disproportionately decrease the number of saplings and rate of recruitment to reproductive age in areas of high conspecific sapling density. These results suggest that sapling herbivory, especially herbivory of terminal meristems, has an important but largely unexplored influence on the population dynamics of tropical tree species. They further demonstrate that sapling herbivory by insects, in addition to the well-studied insect predation and herbivory of seedlings, is likely to influence tree species coexistence in tropical forests.     

Seedling and Sapling Dynamics of Treefall Pits in Puerto Rico1

Seedling and sapling dynamics in a Puerto Rican rain forest were compared between forest understory and soil pits created by the uprooting of 27 trees during Hurricane Hugo. Soil N and P, organic matter, and soil moisture were lower and bulk densities were higher in the disturbed mineral soils of the pits than in undisturbed forest soils ten months after the hurricane. No differences in N and P levels were found in pit or forest soils under two trees with N–fixing symbionts (Inga laurina and Ormosia krugii) compared to soils under a tree species without N–fixing sym–bionts (Casearia arborea), but other soil variables (Al, Fe, K) did vary by tree species. Forest plots had greater species richness of seedlings (<10 cm tall) and saplings (10–100 cm tall) than plots in the soil pits (and greater sapling densities), but seedling densities were similar between plot types. Species richness and seedling densities did not vary among plots associated with the three tree species, but some saplings were more abundant under trees of the same species. Pit size did not affect species richness or seedling and sapling densities. Recruitment of young Cecropia schreberiana trees (>5 m tall) 45 months after the hurricane was entirely from the soil pits, with no tree recruitment from forest plots. Larger soil pits had more tree recruitment than smaller pits. Defoliation of the forest by the hurricane created a large but temporary increase in light availability. Recruitment of C. schreberiana to the canopy occurred in gaps created by the treefall pits that had lower soil nutrients but provided a longer–term increase in light availability. Treefall pits also significantly altered the recruitment and mortality of many understory species in the Puerto Rican rain forest but did not alter species richness.     

阔叶红松林林冠斑块特征及其与林木更新的关系

调查了长白山北坡海拔740m处阔叶红松林主要树种林冠斑块的特征,及其对林分结构与更新的影响,结果表明,林冠斑块主要由红松,椴树,色木,蒙古栎4个树种组成,其林冠斑块覆盖面积占林地面积的57．1％,占样地内林冠斑块总面积的87％,其中以红松和紫椴林冠为主,小林冠斑块数量多,分布广,大斑块则很少,林冠斑块面积基本在100m^2以上,只有个别的斑块超过100m^2,林冠斑块形状变异较大,呈现不规则的多边形,林冠树种对幼龄个体的分布存在一定的影响,色木槭幼龄个体几乎分布在所有的紫椴和红松林冠斑块内,最大密度出现32m^2的紫椴林冠斑块和28m^2的红松林冠斑块内,紫椴的幼龄个体只集中在少数林冠斑块内。     

Spatial structure and regeneration of Tetramerista glabra in peat swamp rain forest in Indonesian Borneo

We examined the spatial structure and regeneration of Tetramerista glabra, a dominant canopy tree in peat swamp rain forest in Borneo (West Kalimantan, Indonesia). T. glabra has strong spatial structure that changes dramatically during the life cycle; seedlings were highly aggregated, saplings were random and trees were evenly distributed. Germination and seedling relative growth were highest within canopy gaps, but seedling densities were highest at gap edges. Concentration of seedlings in gap edges, rather than localized seed dispersal, was responsible for seedling patchiness. The slope of the relationship between relative growth rate and seedling height decreased from gap, to gap edge to understory habitats, suggesting that competition within the seedling layer is more important in gaps than in the understory. The processes that break down seedling aggregation, leading to over-dispersion of trees, must be density dependent, but remain unknown.     

Large-Scale Patterns of Quercus ilex, Quercus suber, and Quercus pyrenaica Regeneration in Central-Western Spain

In Central-Western Spain, forests and woodlands composed of Quercus sp. support outstanding levels of biodiversity, but there is increasing concern about their long-term persistence due to a lack of regeneration. We hypothesize that this regenerative lack is operating on a large geographic scale; that there are differences in the abundance of regeneration between three oak species; that oak regeneration is governed mainly by forest management and structure; and that shrubs act as important physical protectors of seedlings and saplings. We analyzed whether densities of oak seedlings and saplings in several size classes were related to stand-structure, understory, and physiographic variables potentially affecting regeneration. Data collected at a regional level (1 km × 1 km grid) by the Spanish Forest Inventory were evaluated from 2,816 plots. Results revealed that regeneration failure was common for all size categories, from small seedlings to large saplings, and for the three oak species studied, especially the evergreens. Of the Quercus ilex, Q. suber, and Q. pyrenaica plots studied, 49%, 62%, and 20% were lacking any small seedlings, and 82%, 96%, and 56% did not have any large saplings, respectively. Regeneration was positively correlated with tree cover and density, especially of small and medium-sized trees, and negatively correlated with the presence of large trees, indicating that regeneration failure is mostly associated with more open, uniform, and/or aged woodlands. Regeneration densities of Q. ilex and Q. suber were positively correlated with all understory variables, suggesting that the presence of pioneer shrubs represent a major safe site for early tree recruitment, independent from specific shrub species.     

Relationship between tree size and insect assemblages associated with Anadenanthera macrocarpa

This study analyzed the effects of tree size, and correlated architectural tree characteristics, on the assemblages of ants and insect herbivores associated with Anadenanthera macrocarpa (Mimosaceae). The latter is a myrmecophilous tree species from the Atlantic rainforest in south-eastern Brazil. Ants and insect herbivores were collected in 30 individuals of A. macrocarpa , ranging from young individuals (>3 m in height) to emergent trees (up to 40 m). Tree height was a strong indicator of other tree characteristics, including trunk diameter, crown height, crown volume, and number of bifurcations. Ants were collected using arboreal pitfall traps and beating, while insect herbivores with beating only. There was a significant increase in both abundance and species richness of ants and insect herbivores with an increase in tree height. In addition, tree height had a significant effect on the species composition of ants and insect herbivores. Assemblages of both taxa showed a nested organization pattern. The species found in small- and medium-sized trees, in general, consisted of a subset of the species found in the crowns and branches of larger, canopy or emergent trees. Thus, in A. macrocarpa , there was not a replacement of insect species with plant ontogeny. This finding is at variance with those conducted in tropical evergreen forests and which show a clear stratification between the understory and canopy insect faunas. Additional studies are needed to explain these contrasting patterns, but it is possible that differences in microclimate are involved. As the forest we studied is semi-deciduous, microclimatic gradients between the understory and the canopy habitat are probably less severe than in an evergreen forest, thus resulting in a lower turnover of species.     

Foraging Behavior and Coexistence of Two Sunbird Species in a Kenyan Woodland

We investigated the mechanism of coexistence of the rare Amani Sunbird (Hedydipna pallidigastra) and the widespread Collared Sunbird (H. collaris), within Brachystegia woodland in the Arabuko‐Sokoke Forest, Kenya. We compared how prey abundance and search strategies affect resource exploitation by the two species. We used foraging theory to direct our measures of feeding activities as influenced by sunbird species, tree species and foraging height. We evaluated invertebrate abundance among tree species at different heights within trees. The Collared Sunbird primarily used the understory, and the Amani Sunbird primarily used the upper‐canopy. Overall, the rate of prey attacks per flight of the Amani Sunbird was 2.8 times greater than that of the Collared Sunbird. The Amani Sunbird, however, used increased search and attack rates in the understory compared with the mid‐ and upper‐canopies, but the Collared Sunbird foraged similarly throughout all strata. We hypothesize that the increased foraging rate of the Amani in the understory reflects increased foraging costs due to interference from the Collared Sunbird in that stratum. Furthermore, the Collared Sunbird exploits rich patches by moving frequently from place to place. The Amani Sunbird forages slowly, with reduced travel rates, and with a greater number of prey captures within a patch. Arthropod density did not differ among the vegetative strata, but was higher in Brachystegia spiciformis and Hymenaea verrucosa than in six other tree species. We hypothesize that the Amani Sunbird appears dependent upon continued tall B. spiciformis trees within the canopy of the Arabuko‐Sokoke Forest.     

秦岭山地锐齿栎次生林幼苗更新特征

分析了秦岭山地锐齿栎次生林群落乔木及木本幼苗物种特征值、更新生态位宽度及不同影响因子下（包括林分密度、坡向、海拔等）乔木更新动态。结果表明,乔木层物种有25种,锐齿栎占有明显地优势,重要值为149.18%。其次为华山松(Pinus armandii)和千斤榆(Carpinus cordata)等;更新层共有木本植物41种,其中乔木种有28种,占所有木本植物总数量的68%,优势种有青蛙皮槭(Acer grosseri)、木姜子(Litsea pungens)等。林下有丰富的幼苗库,径级及龄级比较小。更新方式以实生为主;分析了10种更新优势乔木种群的生态位宽度,同种幼树种群生态位宽度均大于幼苗种群生态位宽度。幼苗种群生态位宽度从大到小依次为榛子(Corylus heterophylla)、锐齿栎、青蛙皮槭等。幼树种群生态位宽度从大到小依次为榛子、青蛙皮槭、锐齿栎等;林分密度对林下乔木幼苗和幼树数量影响不同。当林分密度（株?hm-2）从720增加到1460时,幼苗密度逐渐增大。随后随着林分密度的增加,又呈现出减少的趋势。而幼树密度随着林分密度增加一直呈现出减少的趋势。坡向对锐齿栎次生林林下乔木幼苗和幼树的影响各异,阳坡（南偏西3）均利于幼苗和幼树的更新。南偏西45林分内,幼苗密度急剧减少。随着坡向转为阴坡,幼苗密度又逐渐增加。而幼树密度逐渐减少。当海拔从1083 m增加到1547 m时,幼树密度逐渐增加。随后,随着海拔增加到1882 m,幼树密度则逐渐减小。而幼苗密度随着海拔的增高一直呈现减小的趋势。     

伏牛山自然保护区森林冠层结构对林下植被特征的影响

在伏牛山自然保护区典型地段设立样方,测定了森林生态系统内几种典型群落类型的冠层结构、光环境特征,调查了林下植被的特征,分析了它们之间的相互关系.结果显示:各群落的冠层结构和光环境有一定的差异,单因素方差分析表明,部分群落间的差异性达到显著水平;各群落灌木层物种丰富度、多样性和均匀度均高于草本层,而优势度正相反;线性拟合的结果表明,草本层的物种丰富度、多样性与冠下光合量子通量密度间呈极显著负相关,优势度与冠下光合量子通量密度间呈显著正相关,灌木层各参数与冠层结构特征间相关性不显著.研究表明,冠层结构的变化对草本层(包括更新幼苗)的影响显著高于灌木层.林隙/林窗或林中空地的出现可能对草本物种或其他阳性及先锋物种具有促进作用,而对优势种幼苗的萌发和定植产生负效应.推测在典型的落叶阔叶林生态系统演替进程中,林下光照强度可能不是最主要的限制因素,优势种种子的扩散、萌发和定植限制可能更重要.     

Introduced deer reduce native plant cover and facilitate invasion of non-native tree species: evidence for invasional meltdown

Invasive species are a major threat to native communities and ecosystems worldwide. One factor frequently invoked to explain the invasiveness of exotic species is their release in the new habitat from control by natural enemies (enemy-release hypothesis). More recently, interactions between exotic species have been proposed as a potential mechanism to facilitate invasions (invasional meltdown hypothesis). We studied the effects of introduced deer on native plant communities and exotic plant species on an island in Patagonia, Argentina using five 400 m² exclosures paired with control areas in an Austrocedrus chilensis native forest stand. We hypothesized that introduced deer modify native understory composition and abundance and facilitate invasion of introduced tree species that have been widely planted in the region. After 4 years of deer exclusion, native Austrocedrus and exotic Pseudotsuga menziesii tree sapling abundances are not different inside and outside exclosures. However, deer browsing has strongly inhibited growth of native tree saplings (relative height growth is 77% lower with deer present), while exotic tree sapling growth is less affected (relative height growth is 3.3% lower). Deer significantly change abundance and composition of native understory plants. Cover of native plants in exclosures increased while cover in controls remained constant. Understory composition in exclosures after only 4 years differs greatly from that in controls, mainly owing to the abundance of highly-browsed native species. This study shows that introduced deer can aid the invasion of non-native tree species through negatively affecting native plant species.