Spatial hints of forest ecotone indicating forest succession, a case of larch forests in Baihuashan Reserve, north China

In cold or alpine areas of northern China, birch forests and larch forests are the two primary forest types. These forests are also characteristic of a south branch of boreal forests in Asia. Some ecologists argue that larch forests can replace birch forests, but this still remains a question due to fragmentary or short observations. The ecotone between a larch forest patch and a birch forest patch is the arena in which the two species interplay and compete with each other, and studies of these areas are meaningful to understanding forest succession. In the alpine area of the Baihuashan Reserve, northern China, we sampled a larch-birch forest ecotone with eight plots in four transects and then analyzed population structures of larches and birches. The results show that the edges of the larch forest patch are composed of many larch saplings or young trees, but the edges of the birch forest patch are mainly composed of old birches. Across the ecotone, the larches, on average, are taller than the birches. These facts suggest that larch saplings can permeate into birch forest patches, probably by seed dispersal, germination, success-ful competition and growth, but birch saplings cannot permeate into larch forest patches. Therefore, on the ecotone, larch forest patches can steadily expand by unceasing permeation into birch forest patches, whereas birch forest patches progressively recede due to ultimate death of the old and poor recruitment of the young. Larch forest patches replace birch forest patches in a stepwise manner, causing succession from birch forests to larch forests. This study not only confirms that larch forests can naturally replace birch forests, but also introduces a simple and reliable method, employing spatial hints, to study forest succession. Additionally, the findings are of benefit to cultivation or development of larch forests in cold or alpine areas of the North Temperate Zone, which can be a huge carbon sink.     

大兴安岭森林火烧后不同演替阶段土壤细菌多样性动态

【目的】研究我国最大的林区之一——大兴安岭森林火烧后不同演替阶段土壤细菌多样性动态,为天然林保护工程对于生物多样性的影响增添新的认识。【方法】以空间替代时间的方法分析大兴安岭森林演替对于土壤细菌多样性动态的影响。大兴安岭森林火烧后典型的自然演替序列为火烧迹地(LG-BA)、灌丛(SHR)、白桦林(BP)、白桦落叶松混交林(BP-LG)、落叶松林(LG-OM)。在演替序列的每个典型样地上采集0-10 cm土样,采用Illumina Mi Seq高通量测序技术测定土壤细菌群落组成及其多样性。【结果】细菌操作分类单位(OTU)数量从少到多的顺序为火烧迹地落叶松白桦混交林灌丛落叶松林白桦林。随着森林演替,多样性指数Simpson先增高后降低;Shannon指数先增加后减少再增加;OTU的丰度变化比较平缓,表明物种变化较小。在各演替阶段中,土壤细菌种类主要有变形菌门(Proteobacteria)、酸杆菌门(Acidobacteria)、放线菌门(Actinobacteria)和浮霉菌门(Planctomycetes),4个门的种类含量随演替顺序都呈现先增加后减少的趋势。主成分分析表明不同演替阶段土壤细菌群落存在一定的差异性。冗余分析表明有机质(SOM)、全氮(TN)、全磷(TP)和p H对于土壤细菌群落变化有影响。【结论】随着森林演替,大兴安岭地区土壤细菌种类和生物多样性会发生变化,其变化与土壤化学成分和p H有关。     

Recruitment patterns and northward tree migration through gap dynamics in an old-growth white pine forest in northern Ontario

Decreases in abundances and declines in growth of eastern white pine over the past century due mainly to human activities have resulted in few large intact old-growth white pine forests in Ontario. These stands may be vulnerable to replacement by deciduous species from temperate forests further south, where recruitment in canopy gap disturbances can greatly define the regeneration process. We investigated recruitment dynamics in canopy gaps of an old-growth white pine forest of Temagami, northern Ontario, Canada, the northern limit of the temperate?Cboreal ecotone. White pine, red pine, black spruce and eastern white cedar represented 85?% of the mature canopy abundance, where trees and saplings established equally in gaps and the closed canopy. Balsam fir and paper birch were more abundant in gaps, showing increases of abundance and basal area with increases in gap size representing canopy self-replacement (balsam fir) and autogenic succession (paper birch). Red maple, at its northernmost range limit, was the only species to show linear increases of abundance and basal area with increases in gap size and gap age. This result, along with adult red maples present in gaps but absent from the closed canopy, identifies the establishment of a northward migrating species in gaps as hypothesized for pine forests at the northern limit of this broad ecotone. We discuss how migration pressures, coupled with pine recruitment limitation through reduced fire frequency by regional fire suppression and predicted future increased warming of 2?C4?°C over the next century, threatens replacement of old-growth white pine forests at this latitude with northward migrating tree species found further south.     

Woody plant assemblages in isolated forest patches in a semiarid agricultural matrix

Spatial and temporal isolation and environmental variability are important factors explaining variation in plant species composition. The effect of fragmentation and disturbance on woody plant species composition was studied using data from 32 remnant church forest patches in northern Ethiopia. The church forests are remnants of dry Afromontane forest, embedded in a matrix of intensively used crop and grazing lands. We used canonical correspondence analysis and partial canonical correspondence analysis to analyze the effects of fragmented and isolated forest-patch identity, environmental and spatial variables on woody plant species composition in different growth stages. The dominance of late successional species was higher at the adult growth stage than seedlings and saplings growth stages. In the adult stages, late successional species like Olea europaea subsp. cuspidate had high frequency of occurrence. Forest patch identity was more important in explaining woody plant assemblages than environmental and spatial variables. For all growth stages combined, environmental variables explained more of the explained total fraction of variation in species composition than spatial variables. Topographic variables best explained variations in species composition for saplings, adults and all growth stages combined, whereas the management regime was most important for seedlings species composition. Our results show that in a matrix of cultivated and grazing land, fragmented and isolated forest patches differ in woody plant species assemblages. Some species are widely distributed and occurred in many patches while other occurred only in one or a few forest patches. Thus, our results indicate that remnant forest patches are important for preserving rare plant species and therefore management practices should focus on minimizing disturbance to the church forests and if possible increase church forest patch size.     

No general edge effects for invertebrates at Afromontane forest/grassland ecotones

The Afromontane region of South Africa is characterised by numerous small, remnant forests in a grassland matrix. The edges, or ecotones between forests and grasslands are usually sharp (typically just over a few metres) and are mainly maintained by both natural and, more recently, anthropogenic fires. We investigated epigaeic amphipod, carabid and ant distribution patterns across Afromontane forest/grassland ecotones and found little evidence to support the biological edge effect. Five of the fifty-two sampled species however, did increase significantly in abundance at the ecotone. Among these was a very distinct edge species, the amphipod Talistroides africana. Overall, carabids were more abundant and species rich in forests while for ants it was in the grasslands. Ants and carabids were both more abundant and species rich in spring and summer than in autumn and winter. More interestingly, the abundance and species richness patterns across the ecotone did not change with the passing of the seasons. We argue that a conservation strategy for the Afromontane forest patches must also incorporate the surrounding grassland. The grassland habitat is often perceived as less valuable than forest and, as a consequence, is subject to many anthropogenic disturbances such as fragmentation, cattle grazing and afforestation. Protecting grasslands around forest patches not only conserves the rich ant diversity, but also conserves the biota in the forests and at the edges, and would therefore be more meaningful in terms of the overall conservation of Afromontane biodiversity.     

The effect of altitude,patch size and disturbance on species richness and density of lianas in montane forest patches

The species richness and density of lianas (woody vines) in tropical forests is determined by various abiotic and biotic factors. Factors such as altitude, forest patch size and the degree of forest disturbance are known to exert strong influences on liana species richness and density. We investigated how liana species richness and density were concurrently influenced by altitude (1700–2360 m), forest patch size, forest patch location (edge or interior) and disturbance intensity in the tropical montane evergreen forests, of the Nilgiri and Palni hills, Western Ghats, southern India. All woody lianas (≥1 cm dbh) were enumerated in plots of 30 × 30 m in small, medium and large forest patches, which were located along an altitudinal gradient ranging from 1700 to 2360 m. A total of 1980 individual lianas were recorded, belonging to 45 species, 32 genera and 21 families, from a total sampling area of 13.86 ha (across 154 plots). Liana species richness and density decreased significantly with increasing altitude and increased with increasing forest patch size. Within forest patches, the proportion of forest edge or interior habitat influenced liana distribution and succession especially when compared across the patch size categories. Liana species richness and density also varied along the altitudinal gradient when examined using eco-physiological guilds (i.e. shade tolerance, dispersal mode and climbing mechanism). The species richness and density of lianas within these ecological guilds responded negatively to increasing altitude and positively to increasing patch size and additionally displayed differing sensitivities to forest disturbance. Importantly, the degree of forest disturbance significantly altered the relationship between liana species richness and density to increasing altitude and patches size, and as such is likely the primary influence on liana response to montane forest succession. Our findings suggest that managing forest disturbance in the examined montane forests would assist in conserving local liana diversity across the examined altitudinal range.     

Quercus and Pinus cover are determined by landscape structure and dynamics in peri-urban Mediterranean forest patches

Successional dynamics in Mediterranean forests have been modulated by anthropogenic disturbances during thousands of years, especially in areas densely populated since ancient times. Our objective is to determine whether pine tree cover (early-successional species) and oak tree cover (late-successional species), used as a surrogate of successional stage of peri-urban fragmented forests in the Vallès lowlands (Catalonia, NE, Spain), are primarily determined by (1) climate and topography; (2) anthropogenic disturbances; (3) patch structure; or (4) patch dynamics from 1956 to 1993. Quercus spp. and Pinus spp. tree cover were separately recorded on 252 randomly selected plots of 100 m², within forest patches ranging in size from 0.25 to 218 ha. Multiple linear regressions indicated that forest patch history is the most important variable determining oak and pine tree cover: new forest patches showed higher pine and lower oak tree cover than recently split patches (i.e. those that became fragmented from large forest areas after 1956). Patches already existing as such in 1956 (pre-existent patches) showed higher pine cover than recently split patches. Oak cover increased and pine cover decreased with increasing forest connectivity of the patch. Finally, highly frequented forests were related to high cover of pines. Climatic and topographic variables were not significant. We conclude that pine and oak cover in these peri-urban forests are mainly determined by recent patch dynamics, but also by the spatial pattern of patches. However, human-induced disturbance can modulate this as there is some evidence for pine being associated with a high human frequentation.     

Cropland edge,forest succession,and landscape affect shrubland bird nest predation

The effects of habitat edges on nest survival of shrubland birds, many of which have experienced significant declines in the eastern United States, have not been thoroughly studied. In 2007 and 2008, we collected data on nests of 5 shrubland passerine species in 12 early successional forest patches in North Carolina, USA. We used model selection methods to assess the effect of distance to cropland and mature forest edge on nest predation rates and additionally accounted for temporal trends, nest stage, vegetation structure, and landscape context. For nests of all species combined, nest predation decreased with increasing distance to cropland edge, by nearly 50% at 250 m from the cropland edge. Nest predation of all species combined also was higher in patches with taller saplings and less understory vegetation, especially in the second year of our study when trees were 4–6 m tall. Predation of field sparrow (Spizella pusilla) nests was lower in landscapes with higher agricultural landcover. Nest predation risk for shrubland birds appears to be greater near agricultural edges than mature forest edges, and natural forest succession may drive patterns of local extirpation of shrubland birds in early successional forest patches. Thus, we suggest that habitat patches managed for shrubland bird populations should be considerably large or wide (>250 m) when adjacent to crop fields and maintained in structurally diverse early seral stages. © 2011 The Wildlife Society.     

次生林区斑块形状动态与森林恢复过程分析

以1959年以来4期航片为基础资料,在ARC／INFO支持下,分析丁关帝山天然次生林区森林恢复过程中各类景观要素斑块异圆指数和类斑边界分维数及其动态。研究表明,采用斑块异圆指数即可揭示景观要素斑块形状特征及其变化规律;研究地区景观恢复过程中,各类非林地斑块的异圆指数总体上高于林地斑块,但呈持续下降趋势,形状趋于规则;林地斑块的异圆指数呈持续上升趋势,景观处于活跃的森林恢复和斑块演替中期,演替后期森林斑块的扩展和合并过程始终占主导地位,斑块形状更趋复杂多变,相邻斑块间的联系密切,先锋性森林群落的斑块形状较为简单,斑块逐渐消退。为研究景观结构与功能的关系和变化规律提供了基础。     

Edge effects as the principal cause of area effects on birds in fragmented secondary forest

Bird communities in tropical forests are strongly affected by both patch area and habitat edges. The fact that both effects are intrinsically confounded in space raises questions about how these two widely reported ecological patterns interact, and whether they are independent or simply different spatial manifestations of the same phenomenon. Moreover, do small patches of secondary forest, in landscapes where the most sensitive species have gone locally extinct, exhibit similar patterns to those previously observed in fragmented and continuous primary forests? We addressed these questions by testing edge‐related differences in vegetation structure and bird community composition at 31 sites in fragmented and continuous landscapes in the imperilled Atlantic forest of Brazil. Over a two‐year period, birds were captured with mist nets to a standardized effort of 680 net‐hours at each site (～22 000 net‐hours resulting in 3381 captures from 114 species). We found that the bird community in patches of secondary forest was degraded in species composition compared to primary continuous forest, but still exhibited a strong response to edge effects. In fragmented secondary forests, edge and area effects also interacted, such that the magnitude of edge to interior differences on bird community composition declined markedly with patch size. The change in bird species composition between forest interiors and edges was similar to the change in community composition between large and small patches (because species had congruent responses to edge and area), but after controlling for edge effects community composition was no longer affected by patch area. Our results show that although secondary forests hold an impoverished bird community, ecological patterns such as area and edge effects are similar to those reported for primary forests. Our data provide further evidence that edge effects are the main drivers of area effects in fragmented landscapes.     

长江上游峨眉冷杉林结构特征的研究

通过对长江上游贡嘎山不同演替阶段峨眉冷杉（Abies fabri）林的野外调查和对比分析,探讨了峨眉冷杉林的结构特征。结果表明,成熟峨眉冷杉林下有丰富的峨眉冷杉种源,小于20年的峨眉冷杉更新幼树幼苗数量少,但却为唯一的优势更新树种,在冬瓜杨、峨眉冷杉混交林中,冬瓜杨、桦木等树种在最初30年高生长非常迅速并能很快占据优势地位。峨眉冷杉是一耐荫性很强的树种,幼龄期生长缓慢,但将更替其它树种形成稳定群落。贡嘎山海拔2900－3400m地区大规模泥石流为100多年发生一次,在泥石流发生地,川滇柳、冬瓜杨、桦树等树种首先出现在迹地并占据优势,在自然条件下恢复峨眉冷杉林将需要较长时间,可通过人为干预措施加以促进和调控。     

季节性雪被对高山森林凋落物分解的影响

季节性雪被可能对高山森林凋落物分解产生重要影响, 但一直没有深入的研究。该文采用凋落物分解袋法, 于2010-2012年雪被覆盖下几个关键时期(冻结初期、深冻期和融化期)以及生长季节, 研究了川西高山森林代表性树种岷江冷杉(Abies faxoniana)、红桦(Betula albosinensis)、四川红杉(Larix mastersiana)和方枝柏(Sabina saltuaria)凋落叶在不同厚度冬季雪被下的分解动态。经过两年的分解, 不同雪被覆盖下岷江冷杉凋落物分解率为33.98%-39.55%, 红桦为46.49%-48.22%, 四川红杉为42.30%-44.93%, 方枝柏为40.34%-43.84%。相对于无雪被覆盖环境, 厚型雪被覆盖均小幅提高了4种凋落物两年的失重率(1.57%-5.57%)。3个针叶树种(岷江冷杉、四川红杉和方枝柏) Olson凋落物分解系数k均以厚型雪被覆盖最大, 薄型雪被覆盖最小, 而阔叶树种红桦分解系数k则表现为无雪被>薄型雪被>较厚型雪被>厚型雪被>中型雪被。尽管在第二年生长季中雪被对红桦凋落物分解的促进作用不明显, 但雪被覆盖明显促进了两年各个关键时期岷江冷杉、四川红杉和方枝柏凋落物的分解。第一年雪被期凋落物分解对当年分解总量的贡献达42.5%-65.5%, 季节性雪被变化明显改变了凋落物冬季分解格局, 对深冻期凋落物分解过程影响尤为显著。综上所述, 当前气候变化情景下冬季雪被的减少可能减缓该区森林凋落物分解过程, 但相对于易分解的阔叶凋落物, 针叶凋落物的响应特征可能更为强烈。     

海南岛热带低地雨林刀耕火种弃耕地自然恢复过程中的群落构建

研究群落构建机制是群落生态学的一个重要目标, 群落动态过程中的构建规律对于了解群落演替机理有重要的作用。该文以海南岛刀耕火种干扰后自然恢复的10 hm ²热带低地雨林为研究对象, 通过比较不同恢复阶段的次生林(15年、30年和60年)和老龄林在幼苗、幼树和成年树群落的物种组成, 揭示次生演替过程中的群落构建规律。研究结果表明, 老龄林中不同径级群落的物种多样性及不同径级间的物种相似度显著高于各恢复阶段的次生林, 但优势种在群落中的比例低于各恢复阶段的次生林。随着自然恢复过程的进行, 次生林群落物种组成与老龄林的相似性也逐渐增大, 支持演替平衡理论。所有恢复阶段样地中幼苗的个体、物种丰富度和基于多度涵盖估计量(ACE)都低于幼树和成年树群落, 幼苗层物种组成与幼树、成年树也有较大差异, 说明新增到幼苗群落可能是一个难于预测的过程。研究结果说明了确定过程和随机过程共同决定了次生演替的群落构建。     

长白山阔叶红松林不同演替阶段土壤团聚体粒径组成及有机碳含量变化

阔叶红松林是我国东北重要的原生群落,其土壤团聚体在森林生态系统碳固定中具有重要作用.本研究采用空间代替时间的方法,选取白桦幼龄林、白桦中龄林、白桦成熟林、阔叶红松成熟林和阔叶红松过熟林5个不同演替序列,通过湿筛法研究长白山天然针阔混交林群落恢复演替中土壤团聚体粒径组成及有机碳含量的变化.结果表明: 土壤团聚体粒径组成受演替过程影响较大,不同演替阶段下土壤团聚体各粒级所占比例差异显著.团聚体平均质量直径随演替的进行表现为先升高再降低的单峰形式,且最高点出现在白桦成熟林阶段.土壤中不同粒级的团聚体内有机碳含量随着演替的进行呈先增加后略有下降的趋势,且团聚体内有机碳含量最大值出现在阔叶红松成熟林阶段.在同一演替阶段下,0～5和5～10 cm土层(除演替末期的阔叶红松过熟林外)中的各粒径团聚体内有机碳含量都随着粒径的减小而增加,而10～20 cm土层中的各粒径团聚体内有机碳含量都随着粒径的减小而减小.从演替初期的白桦幼龄林到演替末期的阔叶红松过熟林,每个样地内的同一粒径团聚体内有机碳含量均具有明显的垂直分布特性,均随着土层深度的增加而显著降低.     

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。     

Viability of ecological processes in small Afromontane forest patches in South Africa

Abstract The conservation of biodiversity is dependent on protecting ecosystem‐level processes. We investigated the effects of fragment size and habitat edge on the relative functioning of three ecological processes – decomposition, predation and regeneration of trees – in small Afromontane forests in KwaZulu‐Natal, South Africa. Ten sampling stations were placed in each of four forest categories: the interior of three large indigenous forest fragments (100 m from the edge), the edges of these large fragments, 10 small indigenous fragments (<1 ha) and 10 small exotic woodlands (<0.5 ha). Fragment size and edge effects did not affect the abundance of the amphipod Talitriator africana, a litter decomposer, and overall dung beetle abundance and species richness significantly. Bird egg predation was marginally greater at large patch edges compared with the other forest categories, while seed predation did not differ among forest categories. Tree seedling assemblage composition did not differ significantly among large patch interiors and edges, and small indigenous fragments. Sapling and canopy assemblage composition each differed significantly among these three indigenous forest categories. Thus, while tree recruitment was not negatively affected by patch size or distance from the edge, conditions in small fragments and at edges appear to affect the composition of advanced tree regeneration. These ecological processes in Afromontane forests appear to be resilient to fragmentation effects. We speculate that this is because the organisms in these forests have evolved under fragmented conditions. Repeated extreme changes in climate and vegetation over the Pleistocene have acted as significant distribution and ecological extinction filters on these southern hemisphere forest biota, resulting in fauna and flora that are potentially resilient to contemporary fragmentation effects. We argue that because small patches and habitat edges appear to be ecologically viable they should be included in future conservation decisions.     

受干扰长白山阔叶红松林林分组成及冠层结构特征

通过样地调查对不同干扰方式产生的过伐天然林、次生白桦林和人工落叶松林等群落的结构组成进行分析和分类探讨 ,并选取了林窗片断和叶面积指数两个能表示群落冠层结构的指标进行分析。结果表明 ,林窗片断值分别为 :原始阔叶红松林 0 194、原始阔叶类 0 185、结构转换型 0 315、结构保留型 0 36 3、结构破坏型 0 2 35、严重干扰类型 0 5 5 0、次生白桦林0 2 13和人工落叶松林 0 2 2 7;叶面积指数分别为 :原始阔叶红松林 1 76 6、原始阔叶类 1 6 80、结构转换型 1 2 5 0、结构保留型 1 0 2 8、结构破坏型 1 5 5 0、严重干扰类型 0 6 35、次生白桦林1 731和人工落叶松林 1 4 73。     

Effects of fragmentation of secondary broadleaf deciduous forests on populations of the near-threatened butterfly, Sasakia charonda (Lepidoptera, Nymphalidae), in central Japan

We examined the effects of fragmentation of secondary broadleaf deciduous forests (secondary forests) on populations of the near-threatened butterfly, Sasakia charonda, in central Japan. Regression analyses revealed that the number of overwintering larvae per host tree significantly increased when the area of secondary forest patches and the Isolation Index of the forest patch increased and the distance from secondary forest patches containing the focal host trees to the nearest secondary forest patch decreased. There was a significantly positive correlation between the number of overwintering larvae and the number of host trees in the neighborhood. The host trees were primarily distributed at the edges of secondary forests. From the results of the backward elimination method of multiple linear regression analysis, independent variables other than patch area were eliminated, and the standardized partial regression coefficient of the patch area was significant. This result suggested that a contiguous distribution of large secondary forest patches with many host trees is very important to conserving this butterfly species.     

Higher climate warming sensitivity of Siberian larch in small than large forest islands in the fragmented Mongolian forest steppe

Forest fragmentation has been found to affect biodiversity and ecosystem functioning in multiple ways. We asked whether forest size and isolation in fragmented woodlands influences the climate warming sensitivity of tree growth in the southern boreal forest of the Mongolian Larix sibirica forest steppe, a naturally fragmented woodland embedded in grassland, which is highly affected by warming, drought, and increasing anthropogenic forest destruction in recent time. We examined the influence of stand size and stand isolation on the growth performance of larch in forests of four different size classes located in a woodland‐dominated forest‐steppe area and small forest patches in a grassland‐dominated area. We found increasing climate sensitivity and decreasing first‐order autocorrelation of annual stemwood increment with decreasing stand size. Stemwood increment increased with previous year's June and August precipitation in the three smallest forest size classes, but not in the largest forests. In the grassland‐dominated area, the tree growth dependence on summer rainfall was highest. Missing ring frequency has strongly increased since the 1970s in small, but not in large forests. In the grassland‐dominated area, the increase was much greater than in the forest‐dominated landscape. Forest regeneration decreased with decreasing stand size and was scarce or absent in the smallest forests. Our results suggest that the larch trees in small and isolated forest patches are far more susceptible to climate warming than in large continuous forests pointing to a grim future for the forests in this strongly warming region of the boreal forest that is also under high land use pressure.