Chance versus determinism in canopy gap regeneration in coastal scarp forest in South Africa

Question: Is tree regeneration in canopy gaps characterized by chance or predictable establishment. Location: Coastal scarp forests, Umzimvubu district, Eastern Cape Province, South Africa. Methods: Estimation of richness of gap‐filling species across canopy gaps of different size. Data are compared with regeneration under the canopy. Probability of self‐replacement of gap forming species is calculated. Results: Forest area under natural gap phase was 7.8%, caused mostly by windthrow (54%). The abundance and average size of gaps (87.8 m²) suggests that species diversity may be maintained by gap dynamics. However, only four of 53 gap‐filler species displayed gap size specialization and these were pioneer species. An additional 13 species were more common in larger gaps but there was no gradient in composition of gap‐filler species across gap size (p= 0.61). Probabilities of self‐replacement in a gap were low (< 0.3) and common canopy species were equally abundant in gaps and the understorey. Species composition in gaps showed no pattern of variation, i.e. was unpredictable, which suggests absence of a successional sequence within tree‐fall gaps. There was also only a slight increase in species richness in gaps at intermediate levels of disturbance. Conclusions: Coastal scarp forest appears not to comprise tightly co‐evolved, niche‐differentiated tree species. Unpredictable species composition in gaps may be a chance effect of recruitment limitation of species from the species pool. Chance establishment slows competitive exclusion and may maintain tree diversity in these forests. These data suggest that current levels (≤ 3 gaps per ha) of selective tree harvesting may not cause a reduction in species richness in this forest.     

Interactive effects of flooding and forest gap formation on tree composition and abundance in the Peruvian Amazon

In order to better understand how flooding and gap formation affect Amazonian rainforests, I set up plots both in three major forest types that differed by flooding duration (referred to here as dry, wet, very wet) and in their respective gaps. Sampling of those plots after 4 years of regeneration showed: (1) common species exist between wet forests and their gaps and between wet and very wet gaps, (2) tree richness is maximum in dry forest and minimum in very wet gaps except in the wet gaps that show the second highest number of species, (3) there were less stems in gaps compared to forests and less stems in forests as flooding increased, except again in the wet gaps, and (4) dominance-diversity curves have more dominance by single species in the dry gap plots compared to other gaps and in dry forest compared to other forests. In general while some aspects of structure such as tree stem density is largely determined by tree-fall gap dynamics, tree composition is determined by flooding regime. Finally a jump in tree richness in wet forests and wet gaps compared to other plots suggests a “mass effects” hypothesis where species from dry and very wet forest and gaps have overlapping ranges in the wet forest and gap. This effect may help explain the high species diversity seen in this part of the Amazon.     

Does restoration help the conservation of the threatened forest of Robinson Crusoe Island? The impact of forest gap attributes on endemic plant species richness and exotic invasions

Invasive plant species are major drivers of biodiversity losses, especially on islands which are prone to invasions and extinctions. In the “endemic montane forest” of Robinson Crusoe Island (Pacific Ocean, Chile) invasive exotic plant species threaten conservation efforts by establishing in gaps and outcompeting native tree species regeneration. We compared gap attributes and ground vegetation cover in three gap types: those dominated by native species (<5 % cover of invasive species), invaded gaps (>30 % cover by invasive species), and treated gaps (invasive species removed). We examined (a) which gap attributes favored native and exotic species, (b) the relationship between gap size and species richness, and (c) species responses to invasion and treatment. Gaps ranged in size from 46 to 777 m² caused mainly by uprooted and snapped trees. Multi response permutation procedures showed a different floristic composition between natural, invaded and treated gaps. The presence of Myrceugenia fernandeziana (native species) and Aristotelia chilensis (invasive species) as gap border trees was positively and negatively correlated with native species richness, respectively. New gaps had more native species than old gaps, and smaller gaps contained relatively more native species than larger ones. An increase in invasive species cover was related to a decline in native species cover and richness. 1–6 years after treatment gaps tended to recover their native floristic composition. Highly effective conservation management programs will concentrate on monitoring gap creation, early control of invasive species, and by treating smaller gaps first.     

Potential causes of arrested succession in Kibale National Park, Uganda: growth and mortality of seedlings

Recent studies suggest that regeneration following large-scale disturbance in Kibale National Park, Uganda, is slow or possibly arrested. Here, data is provided on the growth and mortality of seedlings in the forest understory, treefall gaps, and in large gaps that suggest that this pattern of arrested succession can be attributed partially to the fact that this East African community lacks aggressive colonizing tree species. Growth and mortality rates were contrasted for seedlings of six tree species planted in the understory, small gaps, and large gaps for 36 months. Data suggest that species are adapted to gaps of particular sizes. For example, Uvariopsis congensis grew faster in the understory than in small gaps, whereas Warburgia ugandensis had the lowest mortality rate and highest growth rate in large gaps. Seedlings (n=170) of 15 species were transplanted to assess the response of the tree community to large gap conditions. The limited survival of seedlings in large gaps relative to the understory suggests that only a small proportion of the tree community in this forest regenerates best in gaps larger than those created by the collapse of a single tree. These findings differ from a number of studies conducted in other geographical regions, and suggests that tree communities differ with respect to the proportion of tree species adapted to gaps of particular sizes. This may relate to variation among regions in their history of disturbance and thus frequency of gap formation, size of gaps, and the duration of periods of release. Such variation could imply the existence of a corresponding pattern among tropical forests of differential vulnerability to human disturbance, which tends to create many large gaps.     

Seed Dynamics in Relation to Gaps in a Tropical Montane Rainforest of Hainan Island, South China： （I） Seed Rain

Seed dynamics is an important part of stand dynamics in forest ecosystems. In this paper, 26 gaps were randomly selected to study the influence of gaps on the spatial and temporal patterns of seed rains in a tropical montane rainforest of Hainan Island, South China. Three zones for each gap, including outside gap zone （Non-gap）, transitional gap zone （EG-CG）, and central gap zone （CG）, were designed, and fourseed traps （each lm x lm in size） were placed in each zone. Seed rains were collected by these traps every 10 days from June 2001 to May 2002. Seed rain varied greatly with season and generally exhibited a pattern of unimodal change during the study period： seed abundance and species richness were both greater in the wet season than in the dry season. Gaps significantly influenced the temporal patterns of both species richness and density of seed rains. Gaps had no significant influences on the spatial distribution patterns of seed rain species richness, but significantly affected the spatial distribution pattern of seed rain densities. Among the three different zones of gaps, the outside gap zone generally received more seeds inputs than the two other gap zones.     

Effects of Treefall Gap Disturbances on Ant Assemblages in a Tropical Montane Cloud Forest

The study of gap dynamics and the effects of gaps on diversity has been at the center of tropical ecology for decades. While most studies have focused on the responses of plant species and communities to gap formation, in this study, we consider the effects of treefall gap disturbances on leaf litter ant assemblages in a Neotropical montane cloud forest. We sampled leaf litter ant assemblages and estimated a suite of abiotic parameters in 12 large (>80‐m²) treefall gaps across a chronosequence and in 12 paired adjacent intact forest sites in the Monteverde Cloud Forest Preserve in Costa Rica. No species were more common in gaps than in intact forests, and in fact, species that were common in gaps were also among the most common in forests. The Chao2 estimate of species richness, however, was higher in gap sites than in intact forest sites. In addition, ant assemblages in gap sites did not become more similar to those in adjacent intact sites as gaps aged. In contrast to other studies, our work demonstrates that ant assemblages in the Monteverde Cloud Forest Preserve are weakly affected by the formation of treefall gaps. Together, these results indicate that treefall gap dynamics probably play little role in promoting ant diversity at more regional scales, or coexistence among species at more local scales.     

马尾松人工林不同大小林窗植物多样性及其季节动态

人工林目前存在结构单一、土壤退化、生物多样性降低等人类普遍关注的生态问题。马尾松(Pinus massoniana)是长江上游低山丘陵区退耕还林的主要人工林树种。研究采伐林窗对植物物种组成和更新的影响, 对马尾松低效人工林的改造, 提升其生态服务功能具有重要的意义。该文以采伐39年生的马尾松人工林形成的7种不同大小的林窗为研究对象, 分析了不同季节林窗内的植物生活型组成及多样性变化。结果表明: 1)马尾松人工林林下植物以高位芽植物居多, 其次是地面、地下芽植物, 一年生植物较少而缺少地上芽植物。在林窗形成初期, 林窗的高位芽植物比例明显低于林下, 大林窗的高位芽植物比例稍高于小林窗, 地下芽和一年生植物的比例低于小林窗。2)林下的物种丰富度和物种多样性指数显著低于大林窗。不同林窗下植物的丰富度指数、优势度指数、多样性指数也存在显著差异。3)夏季林窗下植物多样性最高, 其次是秋季, 春季多样性最低。1225-1600 m²的大林窗能够促进马尾松人工林植物多样性恢复和植被更新。     

长白山自然保护区阔叶红松林林隙更新的研究

通过对林隙及非林隙林分组成树种数量特征的对比分析,研究了长白山自然保护区阔叶红松林中主要树种对林隙的更新反应特点,阐述了林隙在阔叶红松林结构与多样性维持中的作用．随着林隙与非林隙的交替变化,红松和阔叶树以及主林层和中下层树种的相对优势（或重要性）亦呈现出交替变化的规律．林隙提高了阔叶红松林的物种丰富度,增加了其多样性,为不同特性物种的共存提供了可能,从而保持了阔叶红松林的整体稳定性．     

Plant diversity and seasonal dynamics in forest gaps of varying sizes in Pinus massoniana plantations

人工林目前存在结构单一、土壤退化、生物多样性降低等人类普遍关注的生态问题。马尾松(Pinus massoniana)是长江上游低山丘陵区退耕还林的主要人工林树种。研究采伐林窗对植物物种组成和更新的影响, 对马尾松低效人工林的改造, 提升其生态服务功能具有重要的意义。该文以采伐39年生的马尾松人工林形成的7种不同大小的林窗为研究对象, 分析了不同季节林窗内的植物生活型组成及多样性变化。结果表明: 1)马尾松人工林林下植物以高位芽植物居多, 其次是地面、地下芽植物, 一年生植物较少而缺少地上芽植物。在林窗形成初期, 林窗的高位芽植物比例明显低于林下, 大林窗的高位芽植物比例稍高于小林窗, 地下芽和一年生植物的比例低于小林窗。2)林下的物种丰富度和物种多样性指数显著低于大林窗。不同林窗下植物的丰富度指数、优势度指数、多样性指数也存在显著差异。3)夏季林窗下植物多样性最高, 其次是秋季, 春季多样性最低。1225-1600 m²的大林窗能够促进马尾松人工林植物多样性恢复和植被更新。     

鼎湖山林窗形成特征及其对幼树组成和多样性的影响

在对鼎湖山自然保护区植物群落动态长期监测的基础上,研究了保护区内各林窗成因、大小和年龄等形成特征,以及地形位置、附近同种的成年乔木等其它相关因子,探讨其对幼树组成和多样性的影响。林窗主要由山体滑坡、暴风雨或雷击大树、虫害等原因形成。结果显示,幼树组成、密度和多样性随林窗形成方式而异,因耐荫能力的差异使出现在林窗的植物不尽相同。林窗大小、年龄和地形位置的变化,导致植物更新生态位的分化,使幼树在林窗内分布形成同源种团。     

长白山红松阔叶林林冠木竞争生长及林冠空隙动态研究

以长白山红松阔叶林林冠木的年龄资料为基础,分析林木冠重叠与年轮径向生长的关系;通过林冠空隙边缘人隙边缘株的干扰释压分析,揭示林冠空隙集中形成的动态规律。红松林相邻林木间理论树冠重叠率与年轮径向生长呈显著负相关,不同树种间存在不同的竞争和协同关系;红松林中除个别小空隙是经过一个集中形成期形成外,大部分空隙是经过２－３个集中形成期形成的,１９００－１９２０和１９１０－１９３０是大部分空隙的集中形成期。     

Flooding × tree fall gap interactive effects on blackwater forest floristics and physical structure in the Peruvian Amazon

To better understand how flooding and tree fall structure forests in the Amazon, I sampled vegetation plots within three blackwater forests (least flooded, medium flooded, highest flooded) and their tree fall gaps in the Peruvian Amazon. I found (1) increased flooding decreased family richness in the closed-canopy forests but increased it in their gaps, with no trends for no. of unique species, (2) flooding decreased stem size everywhere as did the number of stems as size increased especially for larger stems, (3) Green's index showed clumping only for least-flooded forests with the closed-canopy forest showing more than its tree fall gap, and (4) both flooding and tree fall gap creation decreased canopy coverage perhaps as an additive effect. Further among the stem size classes, only the smallest stems were significantly affected by openness and by type of forest, with a significant interaction term where flooding significantly decreased the number of these smaller stems in all forests and their gaps, except those with the highest level of flooding. Also tree fall gaps had significantly more, smaller stems than their forests in the two most-flooded forests, but not in the least-flooded forest. I conclude that flooding is a greater stressor and influence on the structure of these forests than tree fall, and so, in these forests, gradients and disturbances overlap in their traditional roles.     

长白山红松阔叶混交林林冠空隙树种更新动态规律的研究

分析了长白山红松阔叶混交林林冠空隙(Gap)内更新树种的数量特征.结果表明, 在34个林冠空隙中, 更新乔木树种出现频度为7.27%, 更新灌木树种出现频度为21.02%, Gap形成后, 阳性先锋树种首先侵入, 出现频度较高, 占据较宽的生态位, 随着Gap年龄的增加, 树种间的竞争逐渐增大, 阳性树种的生长逐渐受到限制, 阴性树种逐渐增多;Gap年龄与Gap郁闭度和Gap面积呈负相关, Gap面积与树种出现频度呈负相关;Gap年龄主要分布在5～15a和46～55a两段期间, 树种出现频度与Gap年龄存在着密切关系, 当Gap年龄小于25a时, 二者呈正相关;当Gap年龄在25～40a时, 二者呈负相关;当Gap年龄大于40a时, 树种出现频度趋于稳定;34个Gap内4种主要乔木更新树种和4种主要灌木更新树种的胸径均基本上呈正态分布, 乔木更新树种的最高点胸径为4～6cm径级, 灌木更新树种的最高点胸径为1.2cm径级.     

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.     

Seed Dynamics in Relation to Gaps in a Tropical Montane Rainforest of Hainan Island, South China: (Ⅱ) Seed Bank

Spatial and temporal patterns of seed bank dynamics In relation to gaps in an old growth tropical montane rainforast of Hainan Island, South China, were studied over two consecutive years. From June 2001 to June 2003, soil seed bank sampling blocks were taken near each of the four sides of each seed trap and immediately put into a nursery for observation of seedling emergence dynamics in four seasons (each experiment in each season). The abundances of seedlings that emerged from seed banks showed the trend of vine functional group (VFG) > shrub functional group (SFG) > tree functional group (TFG) > herb functional group (HFG), but the trend in species richness of seedlings that emerged from the soil seed banks was TFG > VFG > SFG > HFG. The abundances of seedlings that emerged from seed banks in the three gap zones showed no significant differences, but significant differences did exist for the species richness. The time of sampling or seasons of experiments had significant influences on both the species richness and seedling abundances. The seedling emergence processes of each experiment all revealed the unimodal patterns. Few emergences occurred 1 year after each experiment. Compared with those under closed canopies, the recruitment rates from seed to seedlings and from seedlings to saplings in gaps were higher, but the mortality rates from saplings to big trees were also higher in the gaps.     

The Response of Ground Beetles (Coleoptera: Carabidae) to Selection Cutting in a South Carolina Bottomland Hardwood Forest

We compared the response of ground beetles (Coleoptera: Carabidae) to the creation of canopy gaps of different size (0.13, 0.26, and 0.50 ha) and age (1 and 7 years) in a bottomland hardwood forest (South Carolina, USA). Samples were collected four times in 2001 by malaise and pitfall traps placed at the center and edge of each gap, and 50 m into the surrounding forest. Species richness was higher at the center of young gaps than in old gaps or in the forest, but there was no statistical difference in species richness between old gaps and the forests surrounding them. Carabid abundance followed the same trend, but only with the exclusion of Semiardistomis viridis (Say), a very abundant species that differed in its response to gap age compared to most other species. The carabid assemblage at the gap edge was very similar to that of the forest, and there appeared to be no distinct edge community. Species known to occur in open or disturbed habitats were more abundant at the center of young gaps than at any other location. Generalist species were relatively unaffected by the disturbance, but one species (Dicaelus dilatatus Say) was significantly less abundant at the centers of young gaps. Forest inhabiting species were less abundant at the centers of old gaps than in the forest, but not in the centers of young gaps. Comparison of community similarity at various trapping locations showed that communities at the centers of old and young gaps had the lowest similarity (46.5%). The community similarity between young gap centers and nearby forest (49.1%) and old gap centers and nearby forest (50.0%) was similarly low. These results show that while the abundance and richness of carabids in old gaps was similar to that of the surrounding forest, the species composition between the two sites differed greatly.     

南亚热带常绿阔叶林不同大小和发育阶段林隙的树种多样性研究

分析了南亚热带常绿阔叶林不同大小和发育阶段林隙内树种多样性的变化规律.结果表明,在南亚热带常绿阔叶林中,多样性指数在<400m²的林隙中变化不大,但在400～500m²的林隙中达到最大,而在500～600m²的林隙中最小,在>600m²的林隙中又有所增大.树种多样性指数随林隙年龄的变化趋势是中间高两端低,即在20～50年期间的多样性最大,其次是20年以下的,50年以后的多样性相对最小.林隙更新层中树种多样性指数在500～600m²的林隙中达到最大,在>600m²和200～300m²的林隙中最小.林隙更新层树种多样性指数在林隙形成最初的10年内达到最大值,但随着林隙年龄的增加,总体上表现出下降趋势,在30～40年和50～60年左右又分别形成两个相对的峰值.物种丰富度的变化趋势总体上与树种多样性指数相一致.不同大小和发育阶段的林隙通过其生态因子的改变,对不同树种的更新起到了不同的作用,从而使得不同大小和发育阶段的林隙中树种的多样性特征不同.林隙是维持南亚热带常绿阔叶林树种多样性的一个重要机制.     

Mind the gap: treefalls as drivers of parental trade‐offs

Tree‐fall gaps are small‐scale disturbances whose formation, colonization, and role in forest dynamics are well documented, but whose effects on animal ecology are still greatly overlooked, except for studies comparing species richness of gaps 6+ months old to that in the closed canopy. Other factors associated with the invasion of fresh tree‐fall gaps such as animal breeding adaptations have been largely neglected. I studied the immediate (within hours and days) arrival of the poison frog Dendrobates tinctorius in new tree‐fall gaps to examine the dynamics of their invasion in relation to tadpole rearing. I found that rearing sites are occupied sooner and tadpoles deposited at higher rates in fresh gaps than in the closed forest, but that the rate of cannibalism is also much greater in the former. This suggests that invading new tree‐fall gaps can be the best parental decision when parents arrive early because they get access to fresh, high‐quality resources, but it could be to the detriment of the offspring if parents arrive late, because of overcrowding and cannibalism. These results highlight the importance of studying the earliest stages of invasions in order to have a better understanding of the composition of communities in disturbed ecosystems at later successional stages.     

Effect of Tree‐Fall Gaps on Fruit‐Feeding Nymphalid Butterfly Assemblages in a Peruvian Rain Forest

One of the main natural disturbances that affects the structure of rain forests is treefalls, frequently resulting in gaps. Tree‐fall gaps can bring drastic changes in environmental conditions compared with the undisturbed understory. We investigated the effect of tree‐fall gaps on fruit‐feeding butterfly (Nymphalidae) species assemblages in an undisturbed lowland rain forest in southeastern Peru. We used fruit‐baited traps suspended 2 m above ground in 15 tree‐fall gaps ranging in area from 100 to 1000 m² and in adjacent undisturbed understory. Our data support the hypothesis that tree‐fall gap and understory habitats are utilized by different butterfly species assemblages. There were morphological differences between gap and understory species, where the understory species had a larger wing area to thoracic volume. Vegetation structure and composition were important factors affecting the butterfly assemblages. Most of the butterfly species showed an avoidance of vines and a strong association with the presence of trees and shrubs in gaps. There were also differences among gap assemblages that increased with gap size. Some of the species that were associated with gaps have been considered as canopy species. Other gap species in the present study, however, are known to feed on fruits and/or use host plants mainly, or only, occurring in gaps, implicating that the gap assemblage is a mix of canopy species and those unique to gaps. This indicates that, in an undisturbed Amazon forest, tree‐fall gaps may contribute to maintain species diversity by creating a mosaic of specific habitats and resources that favors different butterfly assemblages.     

海南霸王岭热带山地雨林林隙更新规律的研究

通过对海南岛霸王岭自然保护区原始热带山地雨林中林隙和非林隙林分的调查,分析了海南岛热带山财雨林中主要树种在林隙内外的数量特征、树各对林隙的更新反应和树种多样性的变化规律。结果表明：（１）不同树林隙内和非隙林分中出现的频度、密度和显著度不同,从而表现出其重要值的不同,根据不同树种在林隙内外重要值位序的差异大小,将海南热带山地雨林中树各划分为７种类型：即（１）只出现于林隙中,（２）只出现于非林隙林分中