火后恢复时间影响大兴安岭寒温带森林内部与边缘鸟类多样性
How forest fires affect bird diversity over time in boreal forest interiors and edges in the Greater Khingan Mountains
通讯作者: *E-mail:qiaoqi.sun@nefu.edu.cn
编委: 王彦平
责任编辑: 闫文杰
收稿日期: 2022-11-30 接受日期: 2023-03-29
基金资助: |
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Corresponding authors: *E-mail:qiaoqi.sun@nefu.edu.cn
Received: 2022-11-30 Accepted: 2023-03-29
中重度火干扰通常会显著改变森林结构与功能, 甚至引发森林演替, 严重影响鸟类群落组成与多样性。大兴安岭是我国林火高发热点地区之一, 但林火对鸟类多样性影响的研究较少, 特别是对功能多样性的影响尚不清楚。本研究于2021年7-8月在大兴安岭地区的黑龙江南瓮河国家级自然保护区, 采用样点法对森林鸟类展开调查, 量化不同林火干扰时间下森林内部和边缘鸟类功能多样性的差异。同时, 通过系统性文献检索, 进一步比较了大兴安岭地区不同林分组成与鸟类功能多样性的关系。结果发现: (1)不同林火事件后, 森林内部鸟类群落的组成差异较小, 但边缘鸟类组成差异较大; (2)森林内部的鸟类功能丰富度和均匀度随火后恢复时间增加逐渐升高, 并且在火后早期阶段森林边缘的功能丰富度更高; (3)随着火后恢复时间的增加, 森林边缘的留鸟比例增加, 夏候鸟比例减少; (4)在大兴安岭地区, 针阔混交林中的鸟类功能丰富度显著高于兴安落叶松(Larix gmelinii)林。本研究建议在大兴安岭寒温带森林中, 保持森林内部结构稳定性, 同时提升火后早期阶段森林边缘的植被多样性, 这有利于维持鸟类较高的功能丰富度和功能均匀度。
关键词:
Aim: Moderate/high-severity fire is highly likely to induce changes in forest structure, function, and forest succession, which further threaten bird composition and diversity. Greater Khingan Mountains is a fire-prone region, however, little is known about how forest fires influence bird diversity, particularly its functional diversity.
Methods We used the point counts approach to survey birds and quantified changes in bird community structure and functional diversity after several fires had occurred at both forest interiors and edges in the Heilongjiang Nanwenghe National Nature Reserve from July to August 2021. Furthermore, a systematic literature review was conducted to investigate the relationship between forest composition and bird functional diversity in the Greater Khingan Mountains.
Results: The results showed (1) the difference in bird community composition was small in the forest interiors, but was great at forest edges, with different fire severity histories; (2) the functional richness and evenness of birds in the forest interiors gradually increased with the increased recovery time since the fire, and bird functional richness was greatest at the forest edges in the early stages of the post-fire disturbance; (3) over time since the post-fire recovery, the proportion of resident birds increased, but the proportion of summer migratory birds decreased at the forest edges; (4) bird functional richness was significantly higher in the mixed coniferous forests compared to the larch dominant forests in the Greater Khingan Mountains region.
Conclusions: This study suggests that maintaining the structural stability in forest interiors and enhancing plant diversity at the forest edges in the early stages of the post-fire are beneficial for maintaining high bird functional richness and evenness in the boreal forests.
Keywords:
本文引用格式
李发扬, 李滢钰, 蒋文妮, 刘曙光, 霍超, 孙巧奇, 邹红菲 (2023)
Fayang Li, Yingyu Li, Wenni Jiang, Shuguang Liu, Chao Huo, Qiaoqi Sun, Hongfei Zou (2023)
人类活动导致的气候变化增加了全球林火发生的频次与严重程度(Parks & Abatzoglou, 2020; Clark-Wolf et al, 2022)。林火是影响北方针叶林结构与功能的重要因子之一, 可以直接改变乔木层、灌木层和草本层的组成与比例, 同时也会提高生境的复杂性(Johnson et al, 1998; Kremen et al, 2007; Waito et al, 2018)。不同林火事件发生以后, 由于恢复时间上的差异, 森林植被组成与结构上的差异也越发明显①(①杨扬 (2019) 火灾对大兴安岭地区森林碳储量、植物组成多样性和群落结构影响研究. 硕士学位论文, 东北林业大学, 哈尔滨.)。与此同时, 林火不仅影响初级生产者, 也间接影响较高营养级的消费者。
鸟类在维持生态系统动态平衡、物质循环与能量流动等方面具有重要作用, 同时对环境变化极为敏感, 容易受到林火干扰的影响(Florenzano, 2004; Freeman et al, 2021)。根据“环境过滤假说” (environmental filtering hypothesis), 林火干扰后形成的新生境通常会偏好具备某一类相似特征的鸟类, 而“过滤”其他特征的鸟类物种(Barnagaud et al, 2014; Le Bagousse-Pinguet et al, 2017)。譬如, 在中重度火干扰后, 相较于地面觅食鸟类, 在森林的中上层觅食的鸟类受到的影响更为严重(Connell et al, 2022); 另外, 部分在灌木营巢的鸟类则可能优先选择火后开阔的栖息地, 从而在林火干扰中受益(Rainsford et al, 2022)。因此, 林火后森林植被组成与结构的变化影响着鸟类群落组成与多样性。
林火对鸟类群落组成及多样性的影响复杂, 受到诸多因素的作用, 包括林火干扰后的恢复时间以及鸟类自身对觅食与休憩生境的需求差异等(Schieck & Song, 2006; Taillie et al, 2018)。特别是重度火干扰后, 不同恢复阶段的森林组分存在差异, 进而影响鸟类群落结构(Schieck & Hobson, 2000)。“景观分化假说” (landscape-divergence hypothesis)认为: 同一森林中的鸟类在物种组成上较为趋同, 不同森林中的鸟类组成会有所变化(Laurance et al, 2007)。在很大程度上, 火后生境对鸟类功能特征(如筑巢位置)具有筛选作用, 导致鸟类群落物种组成发生变化, 但其功能丰富度可能保持不变(Lindenmayer et al, 2015)。因此, 仅基于物种多样性难以全面地评估并掌握鸟类群落对林火干扰的响应。
近年来, 作为度量生物多样性的重要指标之一, 功能多样性的研究受到更多关注(Petchey & Gaston, 2002; Schleuter et al, 2010; Spasojevic et al, 2014)。通过量化物种特征值和范围(比如觅食位置、食物类型、生物量等), 可以全面反映群落在多维特征空间中所占据的体积以及物种间的功能差异(Petchey & Gaston, 2002; Mason et al, 2005; Villéger et al, 2008)。量化功能多样性将有助于我们理解不同时空变化下生物群落的构建机制, 也有助于评估生态系统的过程和功能稳定性(Münkemüller et al, 2012; Rainsford et al, 2022)。
受全球林火发生频次与影响程度升高的影响, 国外越来越多的研究开始关注林火对鸟类功能多样性的影响。有研究指出鸟类功能丰富度在很大程度上受到火后生境质量的影响, 同时功能均匀度与火后植被结构的复杂性也有重要的联系(Barbaro et al, 2014; Sitters et al, 2016)。但在我国林火对鸟类群落影响的研究有限, 相关研究仅围绕火后不同生境下鸟类的物种组成和数量变化展开(李健和王文, 2009; 于洋, 2010② (②于洋 (2010) 大兴安岭北坡火烧迹地夏季鸟类群落及多样性研究. 硕士学位论文, 东北林业大学, 哈尔滨.)), 对火后鸟类功能多样性的研究尚属空白。基于此, 本文选择林火频发的大兴安岭寒温带森林湿地生态系统为研究地, 通过野外实地调查, 量化不同林火干扰时间下森林内部和边缘鸟类物种组成和功能多样性的差异, 揭示鸟类功能特征在不同生境下对林火干扰的响应模式。同时, 通过文献综述与数据再分析, 比较大兴安岭地区典型植被结构下鸟类功能多样性的差异, 进一步揭示寒温带森林不同演替阶段(从阔叶林到针阔混交林再到顶级植被群落)森林鸟类群落组成变化, 旨在为火灾后森林资源管理与鸟类多样性的保护提供科学依据和数据支撑。
1 研究区概况
研究地位于黑龙江省大兴安岭地区的黑龙江南瓮河国家级自然保护区(图1), 地理坐标51°05′- 51°39′ N, 125°07′-125°50′ E, 海拔370-1,044 m, 总面积2,290 km2。属于寒温带大陆性季风气候, 年平均气温‒3℃, 年平均降水量500 mm, 降水集中分布于每年7-8月。研究区属于较为独特的寒温带森林湿地生态系统, 林冠层以兴安落叶松(Larix gmelinii)和白桦(Betula platyphylla)为主, 土壤类型主要是表层砂壤土(0-20 cm)和深层的砾砂土(20-40 cm)。该保护区处于完全的冻土带, 湿地、森林、草甸、沼泽和水域等多种生境类型组成的镶嵌景观使得该地区具有高度的复杂性和异质性。同时, 这里林火干扰频繁, 火灾发生后以自然恢复为主, 未进行人工干预, 为我们研究鸟类对火后恢复时间的响应提供了理想环境。
图1
图1
黑龙江南瓮河国家级自然保护区鸟类野外调查样地分布图及其对应的林火历史。图中“FI”和“FE”分别是“forest interior”和“forest edge”的缩写。小图表示自然保护区边界。
Fig. 1
Bird sampling locations with different fire histories in the study area of the Heilongjiang Nanwenghe National Nature Reserve. “FI” and “FE” are abbreviations of “forest interior” and “forest edge” respectively. Inset map shows the boundary of the nature reserve.
由于无法从当地保护区管理局获得精准的林火历史信息, 调查地范围内林火事件发生时间的回溯主要依赖于欧洲航天局(European Space Agency, ESA)提供的MODIS FireCCI51 Burn area遥感数据(空间分辨率为250 m, 覆盖时间2001-2020年)。而对超出MODIS FireCCI51遥感数据时间范围的林火事件(比如早于2001年), 我们则通过树木年轮年代学方法去判定最近一次火灾发生的时间(胡海清等, 2010)。在每个调查样地随机采集20棵大小相对均匀的树木个体, 利用生长锥在每棵树距地面1.3 m处获取树芯样本, 再经过实验室内的交叉定年, 以此判断研究区树木平均年龄, 最终推断样地林火历史。
遥感数据显示, 野外调查样地内分别于2006年(火后15年)、2015年(火后6年)、2017年(火后4年)发生过3次较大规模的森林火灾。由于我们关注的是不同火后恢复时间下鸟类多样性的变化, 同时为了使数据具有可比性, 因此将2015年(火后6年)以及2017年(火后4年)林火历史平均确定为火后5年。最终, 结合林火历史与森林内不同位置, 即森林内部(forest interior, FI)和森林边缘(forest edge, FE), 我们累计调查了8块样地, 包括火后5年(3块)、火后15年(2块)以及火后50年(3块)。每块样地大小在10-30 ha, 样地之间相距1 km以上。样地内的林火历史地图采用QGIS 3.20.0制作(图1)。
2 研究方法
2.1 鸟类调查
2021年7-8月, 在上述8块样地内各设置样点15个, 总计120个样点。森林内部样点之间相距100 m, 而森林边缘由于鸟类可探测性较高, 样点间保持200 m的距离。同时, 对于森林植被和鸟类, 50 m是一个接近边缘效应的阈值(Harper et al, 2005; Barbaro et al, 2014), 故森林内部样点距离边缘至少50 m以上。我们在到达每个样点后, 静待3 min后开始记录看到或听到的鸟类, 并且估计鸟类到样点的距离。每个样点计数15 min, 每个点进行3次重复调查(Gregory et al, 2004)。鸟类调查时间都在6:00-9:00和15:00-18:00进行, 并且在整个调查期间无极端天气(Ehlers Smith et al, 2015)。对于上空飞过的鸟类也进行记录, 但不纳入后续分析; 猫头鹰、猛禽类以及燕类也被排除在分析之外, 因为点计数方法不适用于这些鸟类(Bouvet et al, 2016)。
2.2 鸟类功能特征
鸟类对干扰的响应与鸟类相应的功能特征有关(Hansen & Urban, 1992; Haddad et al, 2008)。目前, 功能丰富度(functional richness, FRic)、功能均匀度(functional evenness, FEve)和功能离散度(functional divergence, FDiv) 3个指数已被广泛证实可以有效评估群落在多维特征空间中所占据的体积和物种间的功能差异(Villéger et al, 2008)。因此, 我们选取了6个主要功能特征计算上述功能多样性指数, 其中包括5个分类变量(觅食位置、食物类型、筑巢类型、筑巢位置和迁徙状态)和1个连续变量(体重) (附录1)。相关的功能特征数据参考王彦平等(2021), 并以Birds of the World (
2.3 林分组成与鸟类功能多样性的关系
通过检索文献资料, 整理已发表的鸟类分布记录信息, 重新计算了大兴安岭地区不同林分组成下的鸟类功能多样性指数。主要分析步骤如下: 首先, 通过中国知网(CNKI,
2.4 数据分析
式中, j为两个鸟类群落之间共有的物种数, a、b分别为两个群落的鸟类物种数。
鸟类群落相似度Jaccard指数的测算在Microsoft Excel 2016中完成。我们统一通过“存在” (1)或“不存在” (0)来标识鸟类物种信息, 并通过R软件“FD”程序包中的“dbFD”函数计算鸟类的FRic、FEve和FDiv, 具体计算公式见Mason等(2005)。同时计算群落加权均值(the community-level weighted means of trait values, CWM)来反映鸟类群落功能特征的变化规律(Laliberté & Legendre, 2010)。随后采用方差分析对不同林火干扰时间下森林内部与边缘的鸟类功能多样性指数进行差异显著性检验。同时, 我们在保持物种丰富度恒定的条件下, 从群落中随机选择物种(1,000次)构建随机群落(零模型, null model), 并计算随机群落的FRic、FEve、FDiv的均值。再对实际观测值与零模型的均值进行比较, 计算两者的标准化效应值(standardized effect sizes, SES), 分别表示为SES.FRic、SES.FEve、SES.FDiv, 如公式(2)所示(丁志锋等, 2020)。我们进一步对SES采用单样本t检验, 判断其是否显著区别于0。SES被认为是生物多样性度量中控制物种丰富度的指标, 可以用于推测群落的构建机制, 如物种特征的相似性限制、环境过滤等(de Bello et al, 2012; Andrikou-Charitidou et al, 2020)。其正、负值分别代表群落在功能上过度分散或聚集(Montaño-Centellas et al, 2020; Jarzyna et al, 2021)。
3 结果
3.1 不同火后恢复时间森林内部与边缘植被组成的差异
在森林边缘, 植被均以小叶章(Calamagrostis angustifolia)、薹草(Carex spp.)、问荆(Equisetum arvense)、木贼(Equisetum hyemale)、地榆(Sanguisorba officinalis)、泽芹(Sium suave)、菖蒲(Acorus calamus)等草本植物为主。但由于森林边缘林火发生后恢复时间的不同, 灌木、乔木等树种的组成存在差异。在火后15年, 细叶沼柳(Salix rosmarinifolia)、笃斯越桔(Vaccinium uliginosum)等灌木开始增加; 火后50年, 除了灌木以外, 兴安落叶松、白桦、东北桤木(Alnus mandshurica)等树种也逐渐增加(图2a-c)。
图2
图2
大兴安岭地区不同火后恢复时间森林内部和边缘样地示意图(大图), 小图为微生境图。FE-5: 火后5年的森林边缘; FE-15: 火后15年的森林边缘; FE-50: 火后50年的森林边缘; FI-5: 火后5年的森林内部; FI-15: 火后15年的森林内部; FI-50: 火后50年的森林内部。
Fig. 2
Photos of study areas at the forest interior and edge since post-fire recovery in the Greater Khingan Mountains. Inset maps show details of micro-habitats. FE-5, Forest edge 5 years post-fire; FE-15, Forest edge 15 years post-fire; FE-50, Forest edge 50 years post-fire; FI-5, Forest interior 5 years post-fire; FI-15, Forest interior 15 years post-fire; FI-50, Forest interior 50 years post-fire.
在森林内部, 林冠层均主要是由兴安落叶松和白桦组成, 但林下植被结构大不相同: 火后5年的灌木层优势种为珍珠梅(Sorbaria sorbifolia)、越桔(Vaccinium vitis-idaea)、笃斯越桔, 草本层以小叶章、地榆、蚊子草(Filipendula palmata)、薹草为主; 火后15年的灌木层以兴安杜鹃(Rhododendron dauricum)、细叶沼柳、绣线菊(Spiraea salicifolia)为主, 草本层以蕨类(Pteridium spp.)、兴安老鹳草(Geranium maximowiczii)、铃兰(Convallaria majalis)、山尖子(Parasenecio hastatus)、毛百合(Lilium dauricum)为主; 火后50年的灌木层以越桔、兴安杜鹃、珍珠梅、绣线菊为主, 草本层以悬钩子(Rubus arcticus)、兴安鹿药(Smilacina dahurica)、红花鹿蹄草(Pyrola incarnata)、假升麻(Aruncus sylvester)、蹄叶橐吾(Ligularia fischeri)为主(图2d-f)。
3.2 火后恢复时间对森林内部与边缘鸟类群落组成的影响
我们在8个样地共记录到鸟类13目27科54种524只(附录2)。鸟类以雀形目为主(36种), 占鸟类种数的66.7%。在排除猛禽类(7种)、燕类(2种)和空中飞过的鸟类(4种)后, 共有41种420只鸟类纳入分析。结果显示, 除了火后15年森林边缘的鸟类多度(68)高于森林内部(43)以外, 在其他林火干扰条件下, 森林内部相比森林边缘均有着更多的物种丰富度和多度(表1)。抽样强度的估计结果显示, 随着鸟类多度的增加, 样本覆盖逐渐趋于1, 此时森林内部物种数估计值为31种, 森林边缘为45种(图3)。在森林内部与边缘实际调查的物种丰富度分别占估计值的93.5%、91.6%, 表明抽样调查足够充分。
表1 不同火后恢复时间森林内部和边缘鸟类物种丰富度与多度
Table 1
火后时间 Time since fire (yr) | 生境类型 Habitat type | 物种丰富度 Species richness | 多度 Abundance | 总计 Total | |
---|---|---|---|---|---|
物种丰富度 Species richness | 多度 Abundance | ||||
5 | 森林边缘 Forest edge | 18 | 69 | 33 (80.5%) | 173 (41.2%) |
森林内部 Forest interior | 23 | 104 | |||
15 | 森林边缘 Forest edge | 14 | 68 | 20 (48.8%) | 111 (26.4%) |
森林内部 Forest interior | 16 | 43 | |||
50 | 森林边缘 Forest edge | 16 | 41 | 31 (75.6%) | 136 (32.4%) |
森林内部 Forest interior | 23 | 95 |
括号内的百分比代表物种丰富度与多度占各自总数的比例
The percentage in parentheses represents the proportion of species richness and abundance to their respective total counts
图3
图3
基于鸟类多度的样本覆盖度与物种丰富度估计。a: 基于鸟类多度对样本覆盖度的估计; b: 以图a估计的样本覆盖度为基础, 对物种丰富度进一步估计。
Fig. 3
Estimation of sample coverage and species richness based on bird abundance. a, Estimation of sample coverage based on bird abundance; b, Further estimation of species richness based on the sample coverage estimated in figure a.
森林内部与边缘鸟类群落相似度较低(表2, Jaccard指数 < 0.5)。比如, 白鹡鸰(Motacilla alba)、普通朱雀(Carpodacus erythrinus)、白腰草鹬(Tringa ochropus)等物种只出现在森林边缘; 而花尾榛鸡(Tetrastes bonasia)、极北柳莺(Phylloscopus borealis)、黑啄木鸟(Dryocopus martius)、白腹鸫(Turdus pallidus)等只在森林内部观察到(附录2)。不同火后恢复时间森林内部的鸟类群落相似度较高(表2, Jaccard指数 ≥ 0.5)。如白眉鹀(Emberiza tristrami)、普通䴓(Sitta europaea)、树鹨(Anthus hodgsoni)等都在森林内部的样地被记录到。
表2 火后不同恢复时间森林内部和边缘的Jaccard指数(C)
Table 2
5年 5 years | 15年 15 years | 50年 50 years | ||||
---|---|---|---|---|---|---|
森林边缘 Forest edge | 森林内部 Forest interior | 森林边缘 Forest edge | 森林内部 Forest interior | 森林边缘 Forest edge | ||
5年 5 years | 森林内部 Forest interior | 0.24 | ||||
15年 15 years | 森林边缘 Forest edge | 0.33 | 0.28 | |||
森林内部 Forest interior | 0.26 | 0.50 | 0.38 | |||
50年 50 years | 森林边缘 Forest edge | 0.36 | 0.34 | 0.15 | 0.28 | |
森林内部 Forest interior | 0.21 | 0.59 | 0.32 | 0.50 | 0.26 |
3.3 火后恢复时间对森林内部与边缘鸟类功能多样性的影响
图4
图4
火后不同恢复时间森林内部和边缘的鸟类功能丰富度(FRic)、功能均匀度(FEve)和功能离散度(FDiv)。不同字母表示组间差异显著, 大小写分别代表P < 0.01和P < 0.05。
Fig. 4
Functional richness (FRic), functional evenness (FEve) and functional divergence (FDiv) of birds at the forest interior and edge since post-fire recovery. Different letters indicate significant differences among groups, with upper and lower case letters representing P < 0.01 and P < 0.05, respectively.
图5
图5
火后不同恢复时间森林内部和边缘鸟类功能组成所占比例。FE-5: 火后5年的森林边缘; FE-15: 火后15年的森林边缘; FE-50: 火后50年的森林边缘; FI-5: 火后5年的森林内部; FI-15: 火后15年的森林内部; FI-50: 火后50年的森林内部。
Fig. 5
Proportion of functional composition of birds at the forest interior and edge since post-fire recovery. FE-5, Forest edge 5 years post-fire; FE-15, Forest edge 15 years post-fire; FE-50, Forest edge 50 years post-fire; FI-5, Forest interior 5 years post-fire; FI-15, Forest interior 15 years post-fire; FI-50, Forest interior 50 years post-fire.
SES的单样本均值t检验结果显示, 在控制物种丰富度后, 不同林火干扰时间下森林内部与边缘的SES.FRic、SES.FEve、SES.FDiv的增减大小并不一致。火后5年森林边缘SES.FRic大于0 (P < 0.05), 但大部分SES.FRic小于0 (图6), 表明火后5年森林边缘的鸟类群落功能特征呈现发散的趋势。
图6
图6
火后不同恢复时间森林内部和边缘鸟类功能丰富度(FRic)、功能均匀度(FEve)、功能离散度(FDiv)的标准化效应值(SES)。FE-5: 火后5年的森林边缘; FE-15: 火后15年的森林边缘; FE-50: 火后50年的森林边缘; FI-5: 火后5年的森林内部; FI-15: 火后15年的森林内部; FI-50: 火后50年的森林内部。
Fig. 6
Standardized effect size (SES) of functional richness (SES.FRic), functional evenness (SES.FEve) and functional divergence (SES.FDiv) at the forest interior and edge since post-fire recovery. FE-5, Forest edge 5 years post-fire; FE-15, Forest edge 15 years post-fire; FE-50, Forest edge 50 years post-fire; FI-5, Forest interior 5 years post-fire; FI-15, Forest interior 15 years post-fire; FI-50, Forest interior 50 years post-fire.
3.4 不同林分组成对鸟类多样性的影响
图7
图7
大兴安岭地区不同林分组成下(落叶松、樟子松与混交林)鸟类丰富度、功能丰富度(FRic)、功能均匀度(FEve)、功能离散度(FDiv)的变化。* P < 0.05; ** P < 0.01。
Fig. 7
Changes in bird richness, functional richness (FRic), functional evenness (FEve), functional divergence (FDiv) under different forest compositions (Larix gmelinii, Pinus sylvestris and mixed forest) in the Greater Khingan Mountains. * P < 0.05; ** P < 0.01.
4 讨论
4.1 林火对森林内部鸟类多样性的影响
通过对森林内部鸟类功能多样性的分析可知,FRic、FEve随着火后恢复时间的增加逐渐上升, 但FDiv之间的差异并不明显, 表明林火改变了鸟类在多维特征空间中的分布模式, 进而导致了FRic、FEve的差异(图4)。这种差异可能是受林火干扰早期与中后期不同阶段的生境差异所致。
在林火早期, 林下层恢复较慢, 部分鸟类会主动规避这类早期生境, 因为简单的森林结构难以满足它们对食物、隐蔽条件等资源的需求(Lindenmayer et al, 2006)。但也有一些鸟类, 如在地面筑巢的树鹨、白眉鹀以及在洞穴筑巢的大斑啄木鸟(Dendrocopos major)、普通䴓等在该生境下有着较高的比例, 这与前人的研究结果相似(Kotliar et al, 2002; Newbold et al, 2013; Lindenmayer et al, 2019)。因此, 我们的研究部分支持了“环境过滤假说”, 因为对某一类型资源来说, 特定鸟类或者具备相似功能特征的物种才能对其展开利用(Venn et al, 2011; Le Bagousse-Pinguet et al, 2017)。与此同时, 在这种资源配置下, 鸟类在功能特征上会较为接近, 进而形成特定的功能群, 最终导致FEve的变化。
在林火干扰中后期, 林下植被包括兴安杜鹃、细叶沼柳等灌木的生长, 增加了林分结构的复杂性, 而偏好在灌丛中筑巢的鸟类如灰头鹀(Emberiza spodocephala)、白眉地鸫(Geokichla sibirica)、巨嘴柳莺(Phylloscopus schwarzi)也开始增加。同时, 林火干扰后期留鸟的比例增加, 一部分原因可能是由于火烧迹地的植被垂直结构异质性增加和森林之间的连通性增强, 为鸟类提供了丰富的食源选择、隐蔽与筑巢条件, 更有利于鸟类生存和繁殖(Tews et al, 2004; Burton et al, 2006; White et al, 2016); 另外的原因可能是由于调查地温暖季节短暂, 夏候鸟和旅鸟需要尽快完成迁徙前的能量补给, 会主动避免与留鸟的资源竞争, 从而减少自身能量消耗(李家兴等, 2022), 因此出现留鸟比例增加的情况。
4.2 林火对森林边缘鸟类功能多样性的影响
森林边缘鸟类的FRic在火后5年最高, 并随火后恢复时间的增加表现出先降低后升高的趋势, 然而FEve和FDiv并没有显著的变化(图4), 表明林火主要影响了森林边缘鸟类群落在特征空间中所占据的体积。相比于15年后的火烧迹地, 火后早期阶段在森林边缘灌丛地带营巢的鸟类占比较少, 这与Taillie等(2018)发现与灌木相关的鸟类随火后时间的恢复呈现出先减后增的趋势一致。过去的研究也证实, 具有不同功能特征的鸟类对火后时间的响应存在差异, 从而造成FRic的不同, 因为林火本身以及火后新的生境均对鸟类产生了持续深远的影响(Smucker et al, 2005; Hart & Chen, 2006)。火后5年森林边缘的SES.FRic > 0, 表明此时群落功能丰富度出现发散(Jarzyna et al, 2021), 即鸟类功能特征在多维空间中占据了更大的范围, 这一结果可能是受环境过滤和物种特征相似性限制的双重影响(de Bello et al, 2012; Andrikou-Charitidou et al, 2020)。在火后早期阶段, 森林边缘植被结构较为简单的草本层“过滤”了其他特征如依赖树洞筑巢的鸟类, 而具有相似功能特征(如筑巢类型、觅食位置、筑巢位置)的鸟类对有限资源的竞争增加, 导致了鸟类群落在功能上呈现发散的趋势。
随着火后演替的推进, 灌木物种开始增加, 使得群落形成更新和演替层(杨健等, 2013)。这种栖息生境令偏好灌木的鸟类如灰头鹀、巨嘴柳莺等的占比相对更高(图5e)。这与前人研究结果一致, 即在火烧迹地灌木较多的区域, 从灌木中受益的物种其密度会更高(Fontaine et al, 2009; Pons & Clavero, 2010)。在火后5年和15年的森林边缘有更多的迁徙鸟类, 比如北红尾鸲(Phoenicurus auroreus)、褐柳莺(Phylloscopus fuscatus)等。但在火后50年, 小嘴乌鸦(Corvus corone)、松鸦(Garrulus glandarius)、褐头山雀(Poecile montanus)等留鸟有着更高的比例。我们推测这种现象可能跟鸟类的生态位分化有关。在火后50年的森林边缘, 鸟类可以较大程度上利用相对稳定的植被群落资源, 迁徙鸟类会倾向选择火后5年和15年有空余生态位的生境, 避免与当地留鸟的竞争(Martin, 1987)。这与Pons和Clavero (2010)发现干扰后的生境相比未干扰的生境有着更多迁徙物种的结论是一致的。
4.3 北方寒温带森林类型对鸟类功能多样性的影响
我们的结果表明, 落叶松林中的鸟类物种数显著高于樟子松林, 但和混交林之间没有显著差异(图7a), 一部分混交林常见的鸟类如山斑鸠(Streptopelia orientalis)、大杜鹃(Cuculus canorus)、树鹨等也频繁出现在落叶松林。同时, 星鸦(Nucifraga caryocatactes)、北噪鸦(Perisoreus infaustus)、白腰朱顶雀(Acanthis flammea)等偏好密闭冠层的鸟类, 几乎只出现在落叶松林。这表明在大兴安岭地区, 以白桦和落叶松为主的混交林作为一种过渡林型, 其鸟类群落的物种多样性水平与以落叶松为主的顶级群落比较一致。森林作为一种关键资源, 强烈影响鸟类的物种组成(Venn et al, 2011; Ibarra & Martin, 2015), 这也符合“景观分化”假说。
我们的结果也发现3种林型之间的FEve、FDiv没有显著差异, 但混交林中FRic显著高于落叶松林和樟子松林(图7b)。这可能是由于混交林相对复杂的林分组成可以为鸟类提供更多的觅食生境、食物资源以及筑巢位置等必要的生存条件(Schieck & Hobson, 2000; Barton et al, 2014; White et al, 2016)。譬如, 混交林中的白桦会招引更多以树皮或木质纤维为食的甲虫, 这将为银喉长尾山雀(Aegithalos glaucogularis)、沼泽山雀(Poecile palustris)、燕雀(Fringilla montifringilla)等食虫性鸟类创造独特的觅食资源, 提供更多生存和繁殖机会(Novoa et al, 2021)。因此, 混交林中的鸟类群落在功能特征空间中占据了更大的体积, 对环境变化的抵抗能力也会更强(Mason et al, 2005)。我们的研究再次证实了探讨环境变化与鸟类多样性关系的时候, 不仅需要从物种多样性角度考虑, 还需要结合鸟类功能多样性的研究, 才能全面掌握鸟类对林火干扰的响应。
5 结论
本研究发现林火对森林内部和边缘鸟类群落结构组成的影响程度不同, 鸟类功能丰富度受到火后恢复时间的显著影响, 并且证实了功能多样性对于理解鸟类与生境之间内在联系的重要性。通过量化寒温带森林生态系统林火对鸟类功能多样性的影响, 有助于火后森林资源的管理以及鸟类多样性保护。同时, 我们也注意到本研究存在一定的局限性。首先, 受林火发生的随机性、复杂性的影响, 我们仅在有限的火烧迹地开展鸟类调查, 在统计分析中有一定的局限性, 因此未来的研究亟待加强对不同火疤地、森林类型、森林相对位置的鸟类群落组成与多样性的长期监测。其次, 现阶段我国北方寒温带森林生态系统鸟类功能多样性与林火干扰研究非常有限, 限制了与其他调查地相似研究的横向比较, 但也凸显了本项研究的基础性与重要性。最后, 在未来的森林管理框架下, 可以适当地将鸟类功能特征的动态变化纳入考虑范围, 更好地实现生物多样性保护与自然保护地科学管理目标。
附录 Supplementary Material
附录1 鸟类功能特征及其对应的生态学意义
Appendix 1 Bird functional traits and their related ecological significance
附录2 不同火后恢复时间森林边缘和森林内部观察到的鸟类物种及数量
Appendix 2 Bird species and numbers observed in forest edge (FE) and forest interior (FI) under different recovery time post-fire
致谢
感谢黑龙江南瓮河国家级自然保护区管理局对我们野外工作的支持, 感谢协助野外调查的金殊宇同学。
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Functional species pool framework to test for biotic effects on community assembly
Functional trait differences among species are increasingly used to infer the effects of biotic and abiotic processes on species coexistence. Commonly, the trait diversity observed within communities is compared to patterns simulated in randomly generated communities based on sampling within a region. The resulting patterns of trait convergence and divergence are assumed to reveal abiotic and biotic processes, respectively. However, biotic processes such as competition can produce both trait divergence and convergence, through either excluding similar species (niche differences, divergence) or excluding dissimilar species (weaker competitor exclusion, convergence). Hence, separating biotic and abiotic processes that can produce identical patterns of trait diversity, or even patterns that neutralize each other, is not feasible with previous methods. We propose an operational framework in which the functional trait dissimilarity within communities (FDcomm) is compared to the corresponding trait dissimilarity expected from the species pool (i.e., functional species pool diversity, FDpool). FDpool includes the set of potential species for a site delimited by the operating environmental and dispersal limitation filters. By applying these filters, the resulting pattern of trait diversity is consistent with biotic processes, i.e., trait divergence (FDcomm > FDpool) indicates niche differentiation, while trait convergence (FDcomm < FDpool) indicates weaker competitor exclusion. To illustrate this framework, with its potential application and constraints, we analyzed both simulated and field data. The functional species pool framework more consistently detected the simulated trait diversity patterns than previous approaches. In the field, using data from plant communities of typical Northern European habitats in Estonia, we found that both niche-based and weaker competitor exclusion influenced community assembly, depending on the traits and community considered. In both simulated and field data, we demonstrated that only by estimating the species pool of a site is it possible to differentiate the patterns of trait dissimilarity produced by operating biotic processes. The framework, which can be applied with both functional and phylogenetic diversity, enables a reinterpretation of community assembly processes. Solving the challenge of defining an appropriate reference species pool for a site can provide a better understanding of community assembly.
Functional and phylogenetic diversity of birds in urban habitat patches in Macao, China
Understanding how communities assemble is a central and fundamental question inecology, which is critical for explaining species coexistence and maintenance of species diversity, and for improving biodiversity conservation practice and policy. Compared with species diversity (species richness and other species diversity indices), functional and phylogenetic diversity consider the differences of species in traits and evolutionary history that closely correlated with ecosystem services and functioning, providing insights into the mechanisms driving community clustering or overdispersion. We used line transects to collect data on bird species richness andabundance in five urban habitat patches in Macao, China (Ecological Zone I: STQ;Lu’niaolin: LNL; tidal flat wetland nearby Saimachang: SMC; tidal flat wetland nearby port: GZ; mangrove forest under Lianhuaqiao: LHQ) from May 2014 to January 2017. Surveys were conducted during the breeding season in 2014, 2015, and 2016, and during the wintering season in 2014, 2015, 2017, and were repeated for 14 times. Species diversity was estimated as species richness and abundance, while functional and phylogenetic diversity were estimated using functional richness (FRic) and Faith’s PD, respectively. Abundance weighted mean pairwise functional distance (MFD) and mean pairwise phylogenetic distance (MPD) were calculated to evaluate changes in the functional and phylogenetic structure of bird community, respectively. The results were as follows: (1) A total of 98 bird species were recorded, belonging to 14 orders and 32 families. Passerines and Ardeidae had the highest species richness (15 and 11 species, respectively). In addition,<em>Egretta garzetta</em> was the most abundant species (849 individuals). (2) Bird species richness (PD and MPD) was the highest in STQ (species richness was 68), whereas abundance was the highest in LHQ (2940 individuals), FRic and MFD was the highest in SMC. (3) Patch area was important factor in explaining the richness, abundance, FRic and PD of birds, whereas vegetation area and tidal flat area had important effects on MFD and MPD, respectively. (4) SES.MFD were significantly clustered in GZ and LHQ, and SES.MPD were significantly clustered in LNL and LHQ, indicating that functional clustering occurred in GZ and LHQ might be a result of environmental filtering. Taken together, our findings suggest that the differences in habitat characters lead to changes in community assembly mechanisms, which is difficult to provide by just considering species diversity. Our results highlight that the necessity of integrating multiple dimension of biodiversity.
澳门城市栖息地斑块中鸟类群落功能和谱系多样性
了解群落构建过程是生态学的核心和基本问题,这对于解释物种共存和物种多样性的维持,完善生物多样性保护政策至关重要。与传统的物种多样性相比,功能和谱系多样性能够提供更多的信息,特别是在群落构建过程方面提供深刻见解。2014年5月—2017年1月,采用样线法对澳门地区5个城市栖息地斑块中(生态一区、鹭鸟林、赛马场滩涂、关闸口岸滩涂和莲花桥侧红树林)的鸟类进行繁殖季和越冬季的调查,共调查14次。利用鸟类丰富度和多度来表征群落的物种多样性,其中多度以最大值保留法计;功能和谱系多样性分别采用功能丰富度(functional richness: FRic)和Faith’PD。同时,利用基于多度加权的平均成对功能距离(mean pairwise functional distance, MFD)和平均成对谱系距离(mean pairwise phylogenetic distance, MPD)来表征群落的功能和谱系结构。结果表明:共记录98种鸟类,隶属于14目32科,目、科、种中数量最多的依次为,雀形目鸟类最多为47种,鹭科物种数为11种,白鹭多度为849只;鸟类物种丰富度(68种)和PD在生态一区最高,多度在莲花桥侧红树林最高为2940只,MFD和FRic在赛马场滩涂最高,MPD在生态一区最高;斑块面积在鸟类丰富度、多度、FRic和PD中具有普遍重要性,植被面积和滩涂面积则分别强烈影响着MFD和MPD;关闸口岸滩涂和莲花桥侧红树林的鸟类群落SES.MFD显著低于随机值(P<0.05),鹭鸟林和赛马场滩涂的鸟类群落SES.MPD显著低于随机值(P<0.05),表明关闸口岸滩涂和莲花桥侧红树林鸟类群落功能聚集(functional clustering),可能与生境过滤驱动群落构建有关。栖息地斑块的特征差异导致了鸟类群落构建机制的不同,这是仅仅考虑物种多样性难以提供的信息,说明了研究群落构建机制时考虑多维度多样性的必要性。
Response of avian diversity to habitat modification can be predicted from life-history traits and ecological attributes
DOI:10.1007/s10980-015-0172-x URL [本文引用: 1]
Functional richness and relative resilience of bird communities in regions with different land use intensities
Birds as indicators of recent environmental changes in the Apennines (Foreste Casentinesi National Park, central Italy)
DOI:10.1080/11250000409356589 URL [本文引用: 1]
Bird communities following high-severity fire: Response to single and repeat fires in a mixed-evergreen forest
Enhancing bird-mediated seed dispersal to increase rainforest regeneration in disused pasture—A restoration experiment
DOI:10.1016/j.foreco.2020.118536 URL [本文引用: 1]
Bird census and survey techniques
In: Bird Ecology and Conservation (eds Sutherland WJ, Newton I, Green R), pp.
Species' traits predict the effects of disturbance and productivity on diversity
DOI:10.1111/j.1461-0248.2007.01149.x
PMID:18201199
[本文引用: 1]
Disturbance is an important factor influencing diversity patterns. Ecological theory predicts that diversity peaks at intermediate levels of disturbance, but this pattern is not present in a majority of empirical tests and can be influenced by the level of ecosystem productivity. We experimentally tested the effects of disturbance on diversity and show that species' autecological traits and community relations predicted species loss. We found that - alone or in concert - increasing disturbance intensity or frequency, or decreasing productivity, reduced diversity. Our species did not exhibit a clear competition-colonization trade-off, and intrinsic growth rate was a more important predictor of response to disturbance and productivity than measures of competitive ability. Furthermore, competitive ability was more important in predicting responses when, in addition to killing individuals, disturbance returned nutrients to the ecosystem. Our results demonstrate that species' traits can help resolve conflicting patterns in the response of diversity to disturbance and productivity.
Avian response to landscape pattern: The role of species' life histories
DOI:10.1007/BF00133308 URL [本文引用: 1]
Edge influence on forest structure and composition in fragmented landscapes
DOI:10.1111/cbi.2005.19.issue-3 URL [本文引用: 1]
Understory vegetation dynamics of North American boreal forests
DOI:10.1080/07352680600819286 URL [本文引用: 1]
Fire history of Mongolian pine (Pinus sylvestris var. mongolica) forests in Mengkeshan of Tahe, China
基于树轮火疤塔河蒙克山樟子松林火灾的频度分析
Beyond species richness: An empirical test of top predators as surrogates for functional diversity and endemism
Global functional and phylogenetic structure of avian assemblages across elevation and latitude
DOI:10.1111/ele.13631
PMID:33124188
[本文引用: 2]
Mountain systems are exceptionally species rich, yet the associated elevational gradients in functional and phylogenetic diversity and their consistency across latitude remain little understood. Here, we document how avian functional and phylogenetic diversity and structure vary along all major elevational gradients worldwide and uncover strong latitudinal differences. Assemblages in warm tropical lowlands and cold temperate highlands are marked by high functional overdispersion and distinctiveness, whereas tropical highlands and temperate lowlands appear strongly functionally clustered and redundant. We additionally find strong geographic variation in the interplay of phylogenetic and functional structure, with strongest deviations between the two in temperate highlands. This latitudinal and elevational variation in assemblage functional structure is underpinned by nuanced shifts in the position, shape and composition of multivariate trait space. We find that, independent of latitude, high-elevation assemblages emerge as exceptionally susceptible to functional change.© 2020 John Wiley & Sons Ltd.
Wildfires in the western Canadian boreal forest: Landscape patterns and ecosystem management
DOI:10.2307/3237276 URL [本文引用: 1]
Effects of fire and post-fire salvage logging on avian communities in conifer-dominated forests of the western United States
Pollination and other ecosystem services produced by mobile organisms: A conceptual framework for the effects of land-use change
Many ecosystem services are delivered by organisms that depend on habitats that are segregated spatially or temporally from the location where services are provided. Management of mobile organisms contributing to ecosystem services requires consideration not only of the local scale where services are delivered, but also the distribution of resources at the landscape scale, and the foraging ranges and dispersal movements of the mobile agents. We develop a conceptual model for exploring how one such mobile-agent-based ecosystem service (MABES), pollination, is affected by land-use change, and then generalize the model to other MABES. The model includes interactions and feedbacks among policies affecting land use, market forces and the biology of the organisms involved. Animal-mediated pollination contributes to the production of goods of value to humans such as crops; it also bolsters reproduction of wild plants on which other services or service-providing organisms depend. About one-third of crop production depends on animal pollinators, while 60-90% of plant species require an animal pollinator. The sensitivity of mobile organisms to ecological factors that operate across spatial scales makes the services provided by a given community of mobile agents highly contextual. Services vary, depending on the spatial and temporal distribution of resources surrounding the site, and on biotic interactions occurring locally, such as competition among pollinators for resources, and among plants for pollinators. The value of the resulting goods or services may feed back via market-based forces to influence land-use policies, which in turn influence land management practices that alter local habitat conditions and landscape structure. Developing conceptual models for MABES aids in identifying knowledge gaps, determining research priorities, and targeting interventions that can be applied in an adaptive management context.
A distance-based framework for measuring functional diversity from multiple traits
A new framework for measuring functional diversity (FD) from multiple traits has recently been proposed. This framework was mostly limited to quantitative traits without missing values and to situations in which there are more species than traits, although the authors had suggested a way to extend their framework to other trait types. The main purpose of this note is to further develop this suggestion. We describe a highly flexible distance-based framework to measure different facets of FD in multidimensional trait space from any distance or dissimilarity measure, any number of traits, and from different trait types (i.e., quantitative, semi-quantitative, and qualitative). This new approach allows for missing trait values and the weighting of individual traits. We also present a new multidimensional FD index, called functional dispersion (FDis), which is closely related to Rao's quadratic entropy. FDis is the multivariate analogue of the weighted mean absolute deviation (MAD), in which the weights are species relative abundances. For unweighted presence-absence data, FDis can be used for a formal statistical test of differences in FD. We provide the "FD" R language package to easily implement our distance-based FD framework.
Habitat fragmentation, variable edge effects, and the landscape-divergence hypothesis
DOI:10.1371/journal.pone.0001017
PMID:17925865
[本文引用: 1]
Edge effects are major drivers of change in many fragmented landscapes, but are often highly variable in space and time. Here we assess variability in edge effects altering Amazon forest dynamics, plant community composition, invading species, and carbon storage, in the world's largest and longest-running experimental study of habitat fragmentation. Despite detailed knowledge of local landscape conditions, spatial variability in edge effects was only partially foreseeable: relatively predictable effects were caused by the differing proximity of plots to forest edge and varying matrix vegetation, but windstorms generated much random variability. Temporal variability in edge phenomena was also only partially predictable: forest dynamics varied somewhat with fragment age, but also fluctuated markedly over time, evidently because of sporadic droughts and windstorms. Given the acute sensitivity of habitat fragments to local landscape and weather dynamics, we predict that fragments within the same landscape will tend to converge in species composition, whereas those in different landscapes will diverge in composition. This 'landscape-divergence hypothesis', if generally valid, will have key implications for biodiversity-conservation strategies and for understanding the dynamics of fragmented ecosystems.
Testing the environmental filtering concept in global drylands
DOI:10.1111/1365-2745.12735
PMID:28642625
1. The environmental filtering hypothesis predicts that the abiotic environment selects species with similar trait values within communities. Testing this hypothesis along multiple - and interacting - gradients of climate and soil variables constitutes a great opportunity to better understand and predict the responses of plant communities to ongoing environmental changes. 2. Based on two key plant traits, maximum plant height and specific leaf area (SLA), we assessed the filtering effects of climate (mean annual temperature and precipitation, precipitation seasonality), soil characteristics (soil pH, sand content and total phosphorus) and all potential interactions on the functional structure and diversity of 124 dryland communities spread over the globe. The functional structure and diversity of dryland communities were quantified using the mean, variance, skewness and kurtosis of plant trait distributions. 3. The models accurately explained the observed variations in functional trait diversity across the 124 communities studied. All models included interactions among factors, i.e. climate - climate (9% of explanatory power), climate - soil (24% of explanatory power) and soil - soil interactions (5% of explanatory power). Precipitation seasonality was the main driver of maximum plant height, and interacted with mean annual temperature and precipitation. Soil pH mediated the filtering effects of climate and sand content on SLA. Our results also revealed that communities characterized by a low variance can also exhibit low kurtosis values, indicating that functionally contrasting species can co-occur even in communities with narrow ranges of trait values. 4. We identified the particular set of conditions under which the environmental filtering hypothesis operates in drylands worldwide. Our findings also indicate that species with functionally contrasting strategies can still co-occur locally, even under prevailing environmental filtering. Interactions between sources of environmental stress should be therefore included in global trait-based studies, as this will help to further anticipate where the effects of environmental filtering will impact plant trait diversity under climate change.
Avian diversity of different habitats of grassland-forest ecosystem in Honghuaerji, Inner Mongolia
内蒙古红花尔基地区草原-森林生态系统不同生境鸟类多样性
Bird diversity in different habitats under agriculturalization in Guangxi, China
DOI:10.17520/biods.2021515
[本文引用: 2]
<p id="p00010"><strong>Aims:</strong> Due to the increasing demand for agricultural land, biodiversity in China has faced increased losses. Our goal in this study is to evaluate how agriculturalization influences bird communities.</p> <p id="p00015"><strong>Methods</strong> We conducted field surveys from October 2020 to October 2021 in three different habitats: (1) forest, (2) agricultural land close to the forest (0‒3 km), and (3) agricultural land far from the forest (6‒9 km). In total, we surveyed bird communities in 180 plots and then repeated the survey for each plot in each season (spring, summer, autumn and winter). All plots were one of three different national nature reserves in Guangxi: Dayaoshan, Damingshan, and Shiwandashan.</p> <p id="p00020"><strong>Results:</strong> (1) The species accumulation curve of each habitat increased at a decreasing rate. The ratio of each habitat’s recorded species richness to the estimated species richness was larger than 60%, indicating that sufficient sampling was conducted; (2) We recorded a total of 196 species belonging to 14 genera and 54 families; passerine birds accounted for the highest proportion of recorded species (71.4%). The total number of species in each habitat type from highest to lowest were: forest (103), agricultural land close to the forest (101), and agricultural land far from the forest (94); (3) Similarity analysis implied that species composition was more similar between the two agricultural habitats, and both were different from forests; (4) Agricultural plots far from the forest had on average the highest Shannon-Wiener diversity index, while forest plots had the lowest in different seasons; (5) The results of the generalized linear mixed model indicated that the bird diversity in each plot per survey in both agricultural land far and close was significantly higher than in forest habitat, but there was no difference between the two agricultural habitats.</p> <p id="p00025"><strong>Conclusion:</strong> Our results suggest that although forests have the lowest species diversity, some species in the forest have very strict habitat requirements. Therefore, protecting forest species and their habitat is vital. Although many species have strong preferences for agricultural habitat, human disturbance in agricultural areas is often extensive and detrimental, so clear protection measures for birds in those areas are necessary as well.</p>
广西山地农业化背景下鸟类多样性比较
DOI:10.17520/biods.2021515
[本文引用: 2]
随着农业用地需求增加, 生物多样性受到严重威胁并急剧下降。为探讨农业化对鸟类多样性的影响, 本文在2020年10月至2021年10月期间, 对广西大瑶山、大明山、十万大山3个国家级自然保护区以及周边农田区域内鸟类进行了调查。研究选取森林、近地农田(距离森林较近的农田)、远地农田(距离森林较远的农田) 3种生境, 布设样点共计180个, 并在不同季节(春、夏、秋、冬)对鸟类多样性进行了调查。结果如下: (1) 3种生境物种累积曲线呈先快速上升, 后变为渐近线或增速放缓趋势, 各生境实际调查鸟类物种数与预测值比例均大于60%, 表明鸟类调查充分; (2)共记录到鸟类196种, 隶属于14目54科, 其中雀形目鸟类占比最高(71.4%)。不同生境记录到鸟类种数从多到少依次为: 森林(103)、近地(101)、远地(94); (3)相似性分析结果表明, 森林与近地农田和远地农田间鸟类相似性差异都较大, 近地与远地间鸟类群落更为相似; (4) 3种生境鸟类群落Shannon-Wiener多样性指数在不同季节里均表现为: 远地 > 近地 > 森林; (5)广义线性混合模型结果显示, 两种农田生境鸟类多样性显著高于森林, 而近地和远地农田生境间鸟类多样性无显著差异。研究结果表明, 虽然森林鸟类多样性较低, 但对其特有种具有较高的保护作用; 尽管鸟类对农田生境表现出更高的喜好, 但农田内人为干扰较为频繁, 缺乏相关的保护措施, 应加强对农田鸟类的保护。
Richness is not all: How changes in avian functional diversity reflect major landscape modification caused by pine plantations
DOI:10.1111/ddi.2015.21.issue-7 URL [本文引用: 1]
Spatiotemporal effects of logging and fire on tall, wet temperate eucalypt forest birds
General management principles and a checklist of strategies to guide forest biodiversity conservation
Functional richness, functional evenness and functional divergence: The primary components of functional diversity
DOI:10.1111/oik.2005.111.issue-1 URL [本文引用: 3]
Using functional and phylogenetic diversity to infer avian community assembly along elevational gradients
DOI:10.1111/geb.v29.2 URL [本文引用: 1]
From diversity indices to community assembly processes: A test with simulated data
Fire regimes shape biodiversity: Responses of avian guilds to burned forests in Andean temperate ecosystems of southern Chile
Warmer and drier fire seasons contribute to increases in area burned at high severity in western US forests from 1985 to 2017
Functional diversity (FD), species richness and community composition
DOI:10.1046/j.1461-0248.2002.00339.x URL [本文引用: 2]
Bird responses to fire severity and time since fire in managed mountain rangelands
DOI:10.1111/j.1469-1795.2009.00337.x URL [本文引用: 2]
R: A Language and Environment for Statistical Computing
Fire and functional traits: Using functional groups of birds and plants to guide management in a fire-prone, heathy woodland ecosystem
DOI:10.1111/ddi.v28.3 URL [本文引用: 2]
Bird communities associated with live residual tree patches within cut blocks and burned habitat in mixedwood boreal forests
DOI:10.1139/x00-061
URL
[本文引用: 2]
By retaining patches of trees in cut blocks, managers expect to retain some forest birds and create bird communities more similar to those found after fires. We surveyed birds from a range of patch sizes (1 to >3000 live residual trees), at four ages (2, 15, 30, 60 years), following two disturbance types (harvest, fire) in mixedwood boreal forest. Bird communities varied among patch sizes, forest ages, and disturbance types. Immediately post-disturbance, bird communities from large patches (>100 residual trees) were more similar to those from old forest than were communities from small patches ([Formula: see text]10 residual trees). Birds that nest or forage in open or riparian habitats had highest densities in small patches 2 years post-harvest, whereas birds that nest in large snags had highest densities in small patches 2 years post-fire. Throughout the chronosequence following both disturbance types, birds from small patches became more similar to those from old forest. Birds that nest and forage in small trees and shrubs were common 15 and 30 years post-disturbance, and generalist forest birds were common 60 years post-disturbance. Birds associated with old forest were present in all patch sizes 15, 30, and 60 years post-disturbance, although larger patches had higher densities of these species.
Changes in bird communities throughout succession following fire and harvest in boreal forests of western North America: Literature review and meta-analyses
DOI:10.1139/x06-017
URL
[本文引用: 1]
Within boreal forests of western North America, the dominant large-scale natural disturbance is wildfire. Thus, harvesting that is as similar as possible to fire is expected to best maintain natural bird communities. We reviewed studies of birds (mainly grouse, woodpeckers, and songbirds) from boreal forests of western North America and conducted meta-analyses to compare the similarity of bird communities occurring postfire versus postharvest. We compared the bird communities at five seral stages and also summarized the effects of retaining large live trees at harvest. Bird communities immediately postharvest differed greatly from those postfire. Differences between disturbance types disappeared as the forest regenerated, and both disturbance types became dominated by relatively few bird species in 31- to 75-year-old forests. During the period 76\u0096125 years postdisturbance, old-forest birds became present and bird species richness increased. However, the trajectory of forest succession during this period influenced bird communities; old aspen (Populus tremuloides Michx.), old mixedwood, and old white spruce (Picea glauca Moench (Voss)) forests all had different bird communities. Retention of large live trees in cutblocks resulted in their use by many old-forest birds, but results were not consistent among studies. Although most bird species had clear peaks in abundance in a specific forest type, no species with more than five detections was limited to a single forest type.
A user's guide to functional diversity indices
DOI:10.1890/08-2225.1 URL [本文引用: 1]
Bird functional diversity decreases with time since disturbance: Does patchy prescribed fire enhance ecosystem function?
Animal species diversity is often associated with time since disturbance, but the effects of disturbances such as fire on functional diversity are unknown. Functional diversity measures the range, abundance, and distribution of trait values in a community, and links changes in species composition with the consequences for ecosystem function. Improved understanding of the relationship between time since fire (TSF) and functional diversity is critical given that the frequency of both prescribed fire and wildfire is expected to increase. To address this knowledge gap, we examined responses of avian functional diversity to TSF and two direct measures of environmental heterogeneity, plant diversity, and structural heterogeneity. We surveyed birds across a 70-year chronosequence spanning four vegetation types in southeast Australia. Six bird functional traits were used to derive four functional diversity indices (richness, evenness, divergence, and dispersion) and the effects of TSF, plant diversity and structural heterogeneity on species richness and the functional diversity indices were examined using mixed models. We used a regression tree method to identify traits associated with species more common in young vegetation. Functional richness and dispersion were negatively associated with TSF in all vegetation types, suggesting that recent prescribed fire generates heterogeneous vegetation and provides greater opportunities for resource partitioning. Species richness was not significantly associated with TSF, and is probably an unreliable surrogate for functional diversity in fire-prone systems. A positive, relationship between functional evenness and structural heterogeneity was comnon to all vegetation types, suggesting that fine-scale (tens of meters) structural variation can enhance ecosystem function. Species more common in young vegetation were primarily linked by their specialist diets, indicating that ecosystem services such as seed dispersal and insect control are enhanced in more recently burnt vegetation. We suggest that patchy prescribed fire sustains functional diversity, and that controlled use of patchy fire to break up large expanses of mature vegetation will enhance ecosystem function.
Changes in bird abundance after wildfire: Importance of fire severity and time since fire
DOI:10.1890/04-1353 URL [本文引用: 1]
Using functional diversity patterns to explore metacommunity dynamics: A framework for understanding local and regional influences on community structure
DOI:10.1111/ecog.00711 URL [本文引用: 1]
Interacting and non-linear avian responses to mixed-severity wildfire and time since fire
Animal species diversity driven by habitat heterogeneity/diversity: The importance of keystone structures
DOI:10.1046/j.0305-0270.2003.00994.x URL [本文引用: 1]
Using plant functional traits to explain community composition across a strong environmental filter in Australian alpine snowpatches
DOI:10.1007/s11258-011-9923-1 URL [本文引用: 2]
New multidimensional functional diversity indices for a multifaceted framework in functional ecology
DOI:10.1890/07-1206.1
PMID:18724739
[本文引用: 2]
Functional diversity is increasingly identified as an important driver of ecosystem functioning. Various indices have been proposed to measure the functional diversity of a community, but there is still no consensus on which are most suitable. Indeed, none of the existing indices meets all the criteria required for general use. The main criteria are that they must be designed to deal with several traits, take into account abundances, and measure all the facets of functional diversity. Here we propose three indices to quantify each facet of functional diversity for a community with species distributed in a multidimensional functional space: functional richness (volume of the functional space occupied by the community), functional evenness (regularity of the distribution of abundance in this volume), and functional divergence (divergence in the distribution of abundance in this volume). Functional richness is estimated using the existing convex hull volume index. The new functional evenness index is based on the minimum spanning tree which links all the species in the multidimensional functional space. Then this new index quantifies the regularity with which species abundances are distributed along the spanning tree. Functional divergence is measured using a novel index which quantifies how species diverge in their distances (weighted by their abundance) from the center of gravity in the functional space. We show that none of the indices meets all the criteria required for a functional diversity index, but instead we show that the set of three complementary indices meets these criteria. Through simulations of artificial data sets, we demonstrate that functional divergence and functional evenness are independent of species richness and that the three functional diversity indices are independent of each other. Overall, our study suggests that decomposition of functional diversity into its three primary components provides a meaningful framework for its quantification and for the classification of existing functional diversity indices. This decomposition has the potential to shed light on the role of biodiversity on ecosystem functioning and on the influence of biotic and abiotic filters on the structure of species communities. Finally, we propose a general framework for applying these three functional diversity indices.
Recent fire activity in the boreal eastern interior of North America is below that of the past 2000 yr
A dataset on the life-history and ecological traits of Chinese birds
DOI:10.17520/biods.2021201 URL [本文引用: 1]
中国鸟类的生活史和生态学特征数据集
DOI:10.17520/biods.2021201
[本文引用: 1]
中国现有1,445种鸟类, 是世界上鸟类物种数最多的国家之一。物种特征反应了生物有机体的功能和适合度, 在生态学、进化生物学和保护生物学研究中具有重要作用。但是, 目前还没有关于我国鸟类生活史、生态学和地理分布等物种特征的完整数据库。通过系统查阅文献和各种数据资料, 本文共收集整理出了中国1,445种鸟类17个功能特征数据: 体重、体长、嘴峰长、翅长、尾长、跗蹠长、食性、窝卵数、卵大小、卵体积、巢址、巢的类型、集群状况、迁徙状况、是否特有种、地理分布范围和分布省份等。在这些特征中, 除迁徙状况、是否特有种、地理分布范围和分布省份外, 其余特征数据均存在不同程度的缺失, 数据的完整度为60.83%‒100%。本数据库是目前关于中国鸟类最新和最全的物种特征数据库, 期望能为我国鸟类生态学、进化生物学、生物地理学、保护生物学等研究提供支持。 数据库(集)基本信息简介 数据库(集)名称 中国鸟类的生活史和生态学特征数据集 作者 王彦平, 宋云枫, 钟雨茜, 陈传武, 赵郁豪, 曾頔, 吴亦如, 丁平 通讯作者 王彦平(wangyanping@njnu.edu.cn) 时间范围 截止到2021年5月 地理区域 地理区域为全中国, 包括香港、台湾和澳门等地区 文件大小 1.0 MB 数据格式 *.xlsx 数据链接 http://dataopen.info/home/datafile/index/id/222 http://doi.org/10.24899/do.202109003 https://www.biodiversity-science.net/fileup/1005-0094/DATA/2021201.zip 数据库(集)组成 数据集共包括1个数据文件, 包括中国1,445种鸟类的17个物种特征
Avian community responses to post-fire forest structure: Implications for fire management in mixed conifer forests
DOI:10.1111/acv.2016.19.issue-3 URL [本文引用: 2]
A review of effects of fire disturbance on understory vegetation in boreal coniferous forest
DOI:10.3724/SP.J.1258.2013.00049
[本文引用: 1]
Understory vegetation in boreal forests is the most diverse and least understood component of boreal plant com- munities. In addition, the aboveground net primary productivity of understory vegetation is almost comparable to that of the trees, and the biomass turns over more rapidly than does that of the trees. Fire disturbance is an important landscape process, driving understory structure and function in boreal forests. We review recent research about how fire, interacting with topography and overstory canopy, affects environmental resources (e.g., light and soil nutrients) and understory vegetation. Recent studies have demonstrated that understory vegetation in boreal forests can also influence post-fire tree seedling regeneration, rate of recovery and successional pathways. It can also affect the quantity and heterogeneity of environmental resources by influencing biogeochemical processes such as decomposition and nutrient flow. Understanding understory vegetation dynamics after fire, especially during early succession, is very important for biodiversity conservation and management of boreal forest.
林火干扰对北方针叶林林下植被的影响
DOI:10.3724/SP.J.1258.2013.00049
[本文引用: 1]
林下植被在北方针叶林植被群落中的物种多样性最高, 且具有较高的生物量周转率和地上部分净初级生产力, 对北方针叶林生态系统功能起着重要作用。火干扰是决定北方针叶林林下植被结构与功能的一个重要景观过程。该文综述了火干扰是如何通过与地形、火前林冠组成的交互作用而影响环境资源和林下植被的。最近的研究表明: 林下植被能够影响火后树木更新苗的定植、重建速率及森林演替轨迹; 林下植被还会通过影响元素的生物地球化学过程(凋落物降解和养分循环)影响林下环境资源的数量与异质性。因此, 研究火后初期北方针叶林林下植被的动态变化, 对于物种多样性保护和森林管理具有重要意义。
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