城市是鸟类的重要栖息地, 城市中的鸟类可以作为重要的指示物种(Sandström et al, 2006)。城市绿地作为城市生态系统的重要组成部分, 是鸟类及其他动物的重要生境(庄艳美等, 2012)。其中, 居住绿地在城市绿地中占比最大, 是人类与鸟类共同生存的环境, 是人工环境和自然环境的复合体, 小区中的植物是人工选择的结果, 因此鸟类多样性可能不同于公园等其他城市绿地。近年来, 对城市鸟类多样性的研究主要集中在城市公园(王勇等, 2014; 杨刚, 2015)、森林公园(张皖清和董丽, 2015)、绿化带(隋金玲等, 2007)、湿地(李相逸等, 2018)等城市公共绿地, 而对居住绿地鲜有研究。
北京作为高度城市化地区和鸟类迁徙路线上的重要节点, 公共绿地的植物配置将直接影响到鸟类的食物来源, 而居住绿地又占比最大, 因此有必要探究居住区的鸟类与食物资源的关系。本研究将从鸟类的食物偏好入手, 构建鸟类的食源关系, 探究在居住区中鸟类取食的影响因素, 以期为营建鸟语花香的森林社区、构建鸟类友好型城市提供科学支撑。
1 研究区域与研究方法
1.1 研究地点
北京市(115°24°-117°30° E, 39°38°-41°05° N)地处华北平原的北部, 属暖温带半湿润半干旱季风气候, 四季分明, 夏季高温多雨, 冬季寒冷干燥。本研究在中心城区的五环路范围内沿8个地理方向随机选取40个居住小区(图1), 各小区之间至少间隔1 km, 以减少相互影响。
图1
1.2 食源情况调查
2020年6月至2021年5月, 选择晴朗无风的上午(夏季6:00-9:00、冬季8:00-11:00), 沿小区人行道以1-1.5 km/h的步速无重复遗漏地调查所有小区, 每月调查1次, 累计调查12次。采用直接计数法进行连续观测, 记录鸟类取食行为的数据。由于麻雀(Passer montanus)属于城市适生种(刘娜娜等, 2018), 长期适应人类生活环境(张淑萍等, 2006), 在小区中广泛分布, 较少受到食源的限制, 对小区的环境特征、食源植物等不敏感, 指示作用不明显, 且数量过多属于离群值, 故本研究中不涉及麻雀。对居住小区的乔、灌木进行全面调查, 记录种类、数量、物候等特征。
1.3 鸟类多样性指数计算
本研究分析了鸟类群落的丰富度、多度、密度(D)和Shannon-Wiener多样性指数(H')。其中物种丰富度为记录到取食行为的鸟类物种数, 多度即记录到的个体数。
式中, D为某个小区中记录到取食行为的鸟类密度, N为某个小区中记录到取食行为的鸟类总数量, S为居住小区的总面积(ha)。
式中, R为居住小区中记录到取食行为的鸟类物种数, Pi为某个小区中记录到第i种取食行为的鸟类数量占鸟类总数的比例。
鸟类食源多样性指数: 借鉴蝴蝶蜜源植物网络中的伙伴多样性指数(partner diversity) (韩丹等, 2021), 计算了鸟类食源多样性指数, 即每个物种食物来源的Shannon-Wiener指数, 较高的值表明了该物种的食物来源广泛。具体计算公式如下:
式中, R为记录到取食行为的鸟类物种数, Fi为某鸟种第i种食物来源的数量占该鸟种食物总数的比例。
1.4 环境影响因子获取
结合居住区的特点, 本研究选取了以下10个影响鸟类取食的环境因子(附录1): (1)小区面积: 在影像图上勾勒小区边界并测量面积(ha)。(2)建成时间: 根据实地调查或从网上查阅相关数据。(3)建筑密度: 在影像图上勾勒建筑边界并测量面积(单位: ha)。建筑密度 = 建筑占地面积/小区面积。(4)平均层数: 调查小区中建筑物的层数, 并按照占地面积进行加权平均。(5)容积率(plot ratio): 依据《建筑工程建筑面积计算规范》(GB/T50353-2013)进行计算, 容积率 = 总建筑面积/净用地面积, 用来反映小区开发强度。(6)周长面积比(perimeter-area ratio, PAR): 周长面积比 = 小区周长/小区面积(李相逸等, 2018)。(7)树冠覆盖率: 对居住小区的影像图进行人工目视解译, 勾绘出树冠的覆盖范围, 计算公式为树冠覆盖率 = 树冠覆盖面积/小区面积(李彤等, 2021)。(8)食源树种丰富度: 统计居住小区中食源树种的物种数。(9)食源树种覆盖率: 统计小区中食源树种的总覆盖面积, 食源树种覆盖率 = 食源树种总覆盖面积/小区面积。(10)食源树种Shannon-Wiener多样性指数:
式中, n为居住小区中食源树种的物种数, Ti为某个小区中记录到第i个食源树种数量占食源树种总量的比例。
1.5 数据分析
1.5.1 食源特征分析
利用R软件的sankeyD3程序包绘制桑葚图, 反映鸟类与食物类型和取食地点的关系, 同时对不同季节也构建关系图。利用circlize程序包绘制弦图, 反映鸟类不同季节对不同生活型植物的利用情况。计算不同季节的鸟类和食源的Bray-Curtis相似性系数, 比较不同季节之间的差异。
1.5.2 影响因素分析
为避免各环境因子之间的多重共线性, 首先检验了Pearson相关性系数(r)。结果显示小区面积与周长面积比、平均层数与容积率、树冠覆盖率与食源树种树冠覆盖率、食源树种丰富度与食源树种Shannon-Wiener多样性指数之间的相关系数均大于0.7 (附录2), 考虑到小区面积和周长面积比反映了小区不同的特征, 分析时仅去除了平均层数、树冠覆盖率和食源树种丰富度3个因子。将每个小区的鸟类多样性作为响应变量(附录3), 对计数数据物种丰富度构建符合Poisson分布的广义线性模型, 密度和Shannon-Wiener多样性指数构建一般线性模型, 建模前对所有数据进行标准化处理。采用小样本校正后的赤池信息量准则(Akaike information criterion, corrected, AICc)进行模型评价, 利用MuMIn程序包中的“dredge”函数进行模型排序和选择, ΔAICc ≤ 2的模型被认为是最优模型(Mortelliti et al, 2010)。利用“model.avg”函数对最优模型进行模型平均, 计算每个因子的参数估计值及标准误。
自身食性的影响采用基于Bray-Curtis距离的非度量多维标度排序(non-metric multidimensional scaling, NMDS)进行分析, 应力函数值stress小于0.2说明排序模型合理。进一步对鸟类的食源多样性指数和多度、分布范围进行一般线性回归。分析过程利用vegan和lme4程序包完成。
2 结果
2.1 居住区有取食行为的鸟类群落特征
本研究共记录到14科35种2,242只鸟类有取食行为, 占调查到鸟类种数(50种)的70%, 其中留鸟16种(占种数的45.71%)、旅鸟15种(42.86%)、夏候鸟1种(2.86%)、冬候鸟4种(11.43%)。灰喜鹊(Cyanopica cyanus, 376只)、白头鹎(Pycnonotus sinensis, 281只)、珠颈斑鸠(Spilopelia chinensis, 147只)和喜鹊(Pica pica, 107只)是小区中数量最多的留鸟。燕雀(Fringilla montifringilla)是北京地区典型的集群过冬的冬候鸟, 共记录到414只。黄腰柳莺(Phylloscopus proregulus, 222只)和黄眉柳莺(P. inornatus, 214只)在春秋迁徙季集中过境时记录到的数量也较多。
物种数在迁徙季时最多, 依次为春季28种(占种数的80%)、秋季25种(71.4%)、冬季16种(45.7%)、夏季9种(25.7%)。鸟类数量从多到少依次为春季(999只)、冬季(659只)、秋季(504只)、夏季(80只)。根据所有小区的总体数据计算出的鸟类群落的Shannon-Wiener多样性指数依次为秋季(2.3959) > 春季(2.3732) > 冬季(2.0342) > 夏季(1.8285)。
2.2 居住区鸟类的食源特征
2.2.1 鸟类的食源构成
鸟类的食源主要为昆虫(33.87%)、翅果(18.33%)、浆果(9.77%)、球果(8.16%)、草籽(5.17%)、聚花果(4.37%)等(图2)。昆虫主要依附在国槐(Styphnolobium japonicum)、绦柳(Salix matsudana ʻPendulaʼ)、刺槐(Robinia pseudoacacia)、榆树(Ulmus pumila)等高大乔木上, 为虫食性鸟类提供食物; 翅果主要来源于元宝枫(Acer truncatum)、白蜡(Fraxinus chinensis)和榆树, 前两者主要在秋冬季为燕雀等植食性的冬候鸟提供食物, 榆树则在早春提供食物; 浆果主要来自金银木(Lonicera maackii)、蔷薇科的小浆果和柿树(Diospyros kaki), 为白头鹎、灰喜鹊等杂食性鸟类提供食物; 球果则来自圆柏(Juniperus chinensis)、侧柏(Platycladus orientalis)、油松(Pinus tabuliformis)等松柏类树种, 主要取食者为灰喜鹊等杂食性鸟类; 喜鹊、珠颈斑鸠等地面觅食的鸟类则常在草地觅食草籽; 桑树(Morus alba)和构树(Broussonetia papyrifera)的聚花果富含充足的水分和糖分, 在5月和6月深受灰喜鹊、白头鹎的青睐。此外, 投喂物也是鸟类的重要食源。
图2
图2
北京居住区鸟类食源关系图。图中的数字表示取食行为频次。
Fig. 2
Relationship between birds and food sources in residential areas of Beijing. The number indicates the feeding frequency.
2.2.2 鸟类不同季节的食源构成
图3
图3
北京居住区不同季节鸟类食源关系图。图中的数字表示取食行为频次。
Fig. 3
Relationship between birds and food sources in residential areas of Beijing in different seasons. The number indicates the feeding frequency.
图4
图4
不同季节不同类型食物的取食频次
Fig. 4
Feeding frequency of different food types in different seasons
不同季节的食源植物存在一定差异, 春季和秋季的Bray-Curtis相似性系数最高(0.4725), 其次为冬春季(0.4655)和秋冬季(0.4291), 夏季与其他季节的相似性系数均低于0.2。这与鸟类相似性存在一致性, 春季和秋季旅鸟过境, 相似性系数高达0.5756 (表1)。
表1 北京居住区不同季节食源植物、鸟类的Bray-Curtis相似性系数
Table 1
| 季节 Season | 食源植物 Food source plant | 鸟类 Bird | ||||
|---|---|---|---|---|---|---|
| 夏季 Summer | 秋季 Autumn | 冬季 Winter | 夏季 Summer | 秋季 Autumn | 冬季 Winter | |
| 春季 Spring | 0.0738 | 0.4725 | 0.4655 | 0.1389 | 0.5756 | 0.4789 |
| 夏季 Summer | 0.1524 | 0.1115 | 0.2641 | 0.2027 | ||
| 秋季 Autumn | 0.4291 | 0.4789 | ||||
2.2.3 鸟类对不同生活型植物的利用
40个小区共有乔木22,628株, 主要的食源树种共计14,674株, 使用量高达64.85%, 其中使用较多的为国槐、圆柏、绦柳、白蜡、元宝枫等北京常见乡土树种。直接取食植物器官的占总取食频次的60.4%, 依托于植物的间接利用占比39.6%。根据不同的植物生活型, 乔木贡献占比高达84.77%, 各利用方式依次为: 乔木直接利用(51.12%)、乔木间接利用(33.65%)、草本直接利用(4.95%)、灌木直接利用(3.90%)、草本间接利用(3.09%)、竹类间接利用(2.62%)、藤本直接利用(0.48%)、灌木间接利用(0.19%), 藤本植物未记录到间接利用方式。
在各季节中均是利用乔木最多, 春季直接利用略多于间接利用, 秋季间接利用略多于直接利用, 而冬季间接利用明显减少; 灌木主要为秋冬季节的直接利用; 而草本直接利用主要在秋冬季节, 间接利用主要在春夏季节(图5)。基于所有小区的数据计算食源植物Shannon-Wiener多样性指数, 依次为秋季(3.1612) > 冬季(2.9651) > 春季(2.9203) > 夏季(2.1763), 在食物匮乏的秋冬季, 鸟类的食源更加丰富。
图5
图5
不同季节鸟类对不同生活型植物的利用方式关系图。图中的数字刻度表示取食行为的频次。
Fig. 5
Plant utilization of different life forms by birds in different seasons. The numerical scale in the figure indicates the feeding frequency.
2.3 鸟类在居住区中取食的影响因素
2.3.1 外界环境因素的影响
通过模型选择, 包含食源树种覆盖率和食源树种Shannon-Wiener多样性指数的模型是预测取食行为鸟类的丰富度(ΔAICc = 0, R2 = 0.5729)和Shannon-Wiener多样性指数(ΔAICc = 0, R2 = 0.2516)的最佳模型, 而密度的最佳模型仅包括食源树种覆盖率(ΔAICc = 0, R2 = 0.2352) (表2), 小区面积、周长面积比、建成时间和建筑密度仅被包括在部分模型中。对上述模型进行模型平均, 食源树种覆盖率(P < 0.001)和食源树种Shannon-Wiener多样性指数(P < 0.001)是影响取食行为鸟类丰富度的关键环境因子, 食源树种覆盖率(P < 0.001)是影响取食行为鸟类密度的关键环境因子, 食源树种Shannon-Wiener多样性指数(P = 0.0023)是影响取食行为鸟类Shannon-Wiener多样性指数的关键环境因子(表3)。综上, 食源树种是影响鸟类在居住区中取食最关键的外界环境因素, 而小区特征的影响不显著。
表2 基于取食行为的鸟类多样性与环境因子的模型选择
Table 2
| 响应变量 Response variables | 模型 Model | 自由度 df | ΔAICc | 权重 Weight | R2 |
|---|---|---|---|---|---|
| 物种丰富度 Species richness | CFST + SFST | 3 | 0 | 0.1518 | 0.5729 |
| CFST + SFST + PAR | 4 | 0.0188 | 0.1504 | 0.5936 | |
| CFST + SFST + CY | 4 | 1.2920 | 0.0796 | 0.5786 | |
| CFST + SFST + AREA | 4 | 1.3209 | 0.0784 | 0.5692 | |
| 物种密度 Species density | CFST | 3 | 0 | 0.1447 | 0.2352 |
| CFST + SFST | 4 | 0.5466 | 0.1101 | 0.2516 | |
| CFST + SFST + AREA | 5 | 1.7703 | 0.0597 | 0.2572 | |
| Shannon-Wiener多样性指数 Shannon-Wiener diversity index | CFST + SFST | 4 | 0 | 0.1129 | 0.2516 |
| CFST + SFST + CY | 5 | 0.9647 | 0.0697 | 0.2516 | |
| SFST | 3 | 1.1036 | 0.0650 | 0.0358 | |
| CFST + SFST + PAR | 5 | 1.2596 | 0.0601 | 0.2431 | |
| CFST + SFST + CY | 5 | 1.9831 | 0.0419 | 0.2431 | |
| CFST + SFST + BD | 6 | 1.9843 | 0.0419 | 0.2253 |
CFST: 食源树种覆盖率; SFST: 食源树种Shannon-Wiener多样性指数; PAR: 周长面积比; CY: 建成时间; AREA: 小区面积; BD: 建筑密度。
CFST, Coverage of food source tree; SFST, Shannon-Wiener diversity index of food source tree; PAR, Perimeter-area ratio; CY, Completed year; AREA, Residential area; BD, Building density.
表3 基于取食行为的鸟类多样性与环境因子平均模型的参数估计
Table 3
| 响应变量 Response variables | 环境因子 Environmental factors | 参数估计 Estimate | 标准误 SE | P |
|---|---|---|---|---|
| 物种丰富度 Species richness | CFST | 0.3885 | 0.1141 | < 0.001*** |
| SFST | 0.5979 | 0.1749 | < 0.001*** | |
| PAR | -0.0918 | 0.1684 | 0.5909 | |
| CY | 0.0227 | 0.0707 | 0.7528 | |
| AREA | 0.0248 | 0.0776 | 0.7538 | |
| 物种密度 Species density | CFST | 8.9247 | 2.5447 | < 0.001*** |
| SFST | 2.1968 | 2.9115 | 0.4577 | |
| AREA | -0.6399 | 1.8525 | 0.7342 | |
| Shannon-Wiener多样性指数 Shannon-Wiener diversity index | CFST | 0.2340 | 0.1696 | 0.1768 |
| SFST | 0.5013 | 0.1594 | 0.0023** | |
| CY | 0.0322 | 0.0928 | 0.7326 | |
| PAR | -0.0665 | 0.1527 | 0.6672 | |
| AREA | 0.0138 | 0.0681 | 0.8434 | |
| BD | 0.0282 | 0.1034 | 0.7877 |
缩写含义见
The meaning of the abbreviation is shown in
2.3.2 自身食性因素的影响
NMDS排序图表明, 虫食性和杂食性的鸟种较为聚集且存在部分重叠, 而植食性的鸟种较为分散(stress = 0.1247) (图6)。相似食性的鸟种排序聚集说明食性是影响鸟类取食重要的内在因素。白头鹎、灰喜鹊、大斑啄木鸟(Dendrocopos major)、喜鹊等食源丰富度较高, 取食多种食源植物, 其中白头鹎、灰喜鹊、大斑啄木鸟食源多样性指数也较高即食性广泛, 而乌鹟(Muscicapa sibirica)等10种鸟类仅观测到1种食源(附录4)。线性回归分析表明, 鸟类多度和所分布的小区数量均与鸟类食源多样性指数呈显著的正相关且解释率较高(R2 = 0.3455, P < 0.001; R2 = 0.7044, P < 0.001) (图7)。表明食源越丰富、食性越广泛的鸟种, 其种群数量越大、分布范围越广泛, 更能够适应城市居住区的复杂环境, 而食源越单一的鸟种, 数量越少、分布越集中, 对特定的食物依赖度越高。
图6
图6
基于取食行为的北京居住区鸟种排序图。图中数字表示前10位的取食地点, 椭圆表示95%的置信区间。1: 元宝枫; 2: 国槐; 3: 草地; 4: 榆树; 5: 绦柳; 6: 白蜡; 7: 裸土地; 8: 侧柏; 9: 重瓣粉海棠; 10: 栾树。
Fig. 6
Ranking map of bird species based on feeding behavior in residential areas of Beijing. Number indicates top 10 feeding sites, and ovals indicate 95% confidence intervals. 1, Acer truncatum; 2, Styphnolobium japonicum; 3, Lawn; 4, Ulmus pumila; 5, Salix matsudana ʻPendulaʼ; 6, Fraxinus chinensis; 7, Bare land; 8, Platycladus orientalis; 9, Malus spectabilis var. riversii; 10, Koelreuteria paniculata.
图7
图7
北京居住区鸟类多度(A)、分布小区数(B)与食源多样性指数的线性回归。灰色区域表示拟合线的置信区间。
Fig. 7
Relationship between bird abundance (A), number of residential areas (B) and Shannon-Wiener diversity index of food source in residential areas of Beijing. Gray areas indicate confidence intervals for the fitted lines.
3 讨论
3.1 在居住区中取食的鸟类群落特征
北京共有鸟类503种(
与徐诗等(2021)在某社区中的研究结果相类似, 鸟种在春季和秋季迁徙期较多。夏季记录到的鸟种数少于其他季节, 推测原因是: (1)夏季繁殖期鸟类以取食昆虫为主(隋金玲等, 2006), 例如家燕(Hirundo rustica)等夏候鸟为空中捕食, 对植物的依赖度较低; (2)冬季城市环境是重要的越冬场所, 物种丰富度在冬季显著升高(Tzortzakaki et al, 2018), 例如大山雀(Parus major)等许多留鸟在夏季会到近郊山地或森林里进行繁殖育雏, 而秋冬季食物匮乏时则会到城区游荡觅食; (3)夏季鸟类更易寻找到食物, 取食活动时间短且常隐蔽在树冠中不易观测, 因此记录到的物种数和数量较少。
3.2 鸟类食源特征及人为活动的影响
鸟类的食物具有种间特异性和时空特异性(李翔等, 2018), 其食物来源与动植物的物候相关联, 对植物的利用在11月至次年3月达到高峰。食物可获得性决定了鸟类分布(Ferger et al, 2014), 各季节的鸟类种类和数量与盛花期和盛果期的乔木种类变化趋势相同, 即使同一季节, 盛花盛果乔木上的鸟类较多(史慧灵等, 2016), 植物的结果期不同也使得鸟类对果实的利用存在差异(Gallinat et al, 2020), 当鸟类无法获得成熟的果实时, 则会取食平时不吃的未成熟的果实, 来确保营养和能量的获取(Mercedes, 1977)。植物的果肉能够吸引众多的食果鸟(Zhu et al, 2022), 它们的体重与可获得的果实丰度正相关(González-Varo et al, 2021)。此外, 初春植物体内各种组织开始活跃, 燕雀、啄木鸟类、山雀类等鸟类会通过吸食元宝枫树干分泌出的树液来补充养分。
鸟类对昆虫的取食集中在4-10月的昆虫活跃期, 繁殖期鸟类靠捕捉昆虫育雏, 如, 迁徙过境的柳莺类、鹟类等虫食性鸟类, 春季在槐树和各类柳树上取食, 秋季在紫叶李(Prunus cerasifera ʻPissardiiʼ)、榆树、栾树(Koelreuteria paniculata)等易受虫害的树种上取食。大斑啄木鸟、星头啄木鸟(Dendrocopos canicapillus)等攀禽也在刺槐、泡桐(Paulownia tomentosa)和榆树等高大乔木上取食天牛、小蠹、吉丁虫、象甲等各类蛀干害虫。研究表明, 食虫鸟的体型大小与其所食节肢动物的体型大小正相关, 鞘翅目是被鸟类利用最多的节肢动物, 其次是蛛形目、膜翅目和鳞翅目(Katerina et al, 2017), 甚至枯死木上大量的鳞翅目昆虫幼虫亦是鸟类的食物来源(范喜顺等, 2005)。今后有待将鸟类的食性与具体的昆虫种类进行关联。
受人为活动影响, 人工投喂在维持鸟类多样性上具有重要作用, 例如小米、玉米等谷物为食谷鸟珠颈斑鸠提供了全年的口粮。居民的友好行为为鸟类提供了食源, 在食物匮乏期实现与植物食源的互补, 灰喜鹊等鸟在深冬也会取食流浪猫的猫粮。
3.3 鸟类对不同生活型植物的利用
居住区的食源植物具有类型丰富、种类繁多、从早春到晚秋覆盖全物候、供给期长的特点。木本植物的丰富度和多样性显著影响鸟类多样性(Yang et al, 2015), 乔木是利用率最高的树种, 由于利用期长, 花芽、叶芽、果实等全物候均可作为食物。同时多数鸟类喜欢在冠大荫浓的树上采食(隋金玲等, 2006), 隐蔽性强增加了取食安全性。利用灌木或草本的较少, 这与绿化配置方式和养护管理方式有关, 灌木多为高度修剪的绿篱, 金银木等丛生型灌木也常过度修剪, 规则式或频繁修剪不易形成花与果实(冼丽铧等, 2020)。自生草本地被是维持生物多样性的重要因素(李晓鹏等, 2018; 李晓鹏和董丽, 2020; 陈颖等, 2021), 可全年为鸟类提供丰富的节肢动物、植物和种子(Cabodevilla et al, 2021), 枯枝落叶层也有利于维持较高的鸟类多样性(Shwartz et al, 2008)。而小区中多是人工种植的草或草坪, 自生草本也作为杂草被清理, 调查中发现苦荬菜(Ixeris polycephala)就吸引了成群的灰椋鸟(Spodiopsar cineraceus)前来取食花瓣。
3.4 鸟类在居住区中取食的影响因素
食源树种是影响鸟类取食最关键的外界环境因素, 虽然研究表明高度城市化使得鸟类食源更窄(Peneaux et al, 2021), 但居住区绿地碎片化、异质性较高(Jiao et al, 2021), 人工种植的食源植物为鸟类提供了丰富的食源。鸟类多样性与食源树种物种数之间存在正相关(谭丽凤和杨昌尚, 2010; 史慧灵等, 2016), 但密度仅受食源树种量的影响, 与种类无关, 即使种类单一但量多时也会吸引鸟类, 例如灰喜鹊常集群在松柏林中觅食。以往的研究中, 鸟类多样性与绿地面积常呈正相关(Huang et al, 2015; Callaghan et al, 2018), 鸟类更倾向于占据城市公园(Tiwary & Urfi, 2016), 但本研究中面积效应不显著, 可能是由于小区与大型城市公园绿地相比规模较小, 相较于鸟类的活动范围, 小区只是其中一部分或仅作为取食的餐厅。此外当拥有充分的食物供给时, 面积就不再是影响鸟类丰富度的主要因素(李相逸等, 2018)。本研究中用建筑密度反映城市化程度和人类干扰强度, 多数研究表明鸟类多样性与建筑物比例呈现负相关或不显著(郭诗怡等, 2022), 随着建筑密度的增加, 鸟类的丰富度以及虫食性鸟的丰度下降, 而城市适应者和杂食性物种却增加(Amaya-Espinel et al, 2019)。小区中鸟类多为城市适应者和开拓者(Geschke et al, 2018), 且以杂食性为主, 但其取食不存在明显的空间异质性, 可能是由于研究区是用地类型均质的高度城市化地区。
3.5 居住区绿化植物配置与养护管理的建议
居住区植物配置时, 要做到乔灌草科学配置、优化时空布局、乡土树种协调发展。配置杨、柳、榆、槐等乡土树种以及金银木等观果植物作为引鸟树种, 适当减少观赏性高而生态价值较低的绿篱、观花灌木和小乔木。增加野化灌木层, 为在不同生态位生活的多种鸟类提供适合的栖息条件(彭子嘉等, 2020)。这与北京市新一轮百万亩造林绿化工程 (
本研究探究了北京居住区鸟类的食源特征, 揭示了绿化植物为鸟类直接或间接地提供食物, 人类的投喂行为也是鸟类度过食物匮乏期的重要补充。小区在绿化建设时配置食源树种、减少绿篱使用, 采取低干扰的近自然园林养护管理措施, 有利于营造鸟语花香的森林社区, 建设鸟类友好型城市。本研究是基于鸟类取食行为开展的, 而实际鸟类对生境的选择还依赖于是否能提供安全的筑巢场所、停息的庇护所等, 今后有待进一步在鸟类植物生境偏好、夜栖地选择等方面开展相关研究。
附录 Supplementary Material
附录1 北京市40个居住小区环境参数
Appendix 1 Environmental parameters of 40 residential areas in Beijing
附录2 外界环境因子的Pearson相关性(r)分析结果
Appendix 2 External Pearson correlations coefficients (r) among environmental factors
附录3 北京市居住小区基于取食行为的鸟类多样性参数
Appendix 3 Bird diversity parameters based on feeding behavior in residential areas of Beijing
附录4 北京市居住小区有取食行为的鸟类名录
Appendix 4 Bird list of feeding behavior in residential areas of Beijing
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北京城市绿地冬季鸟类群落特征
Study on bird-feed trees in parks of Liuzhou urban area in winter
柳州城市公园冬季鸟类食源树种调查研究
Spatial variations of bird occupancy in Delhi: The significance of woodland habitat patches in urban centres
Seasonal patterns of urban bird diversity in a Mediterranean coastal city: The positive role of open green spaces
DOI:10.1007/s11252-017-0695-9 URL [本文引用: 1]
Bird preference to food source trees in urban green space, Guangzhou, China
广州城市绿地中鸟类对食源树种的偏好
Biodiversity characteristics in near-natural community parks: A case study in the central area of Beijing
近自然社区公园的生物多样性特征研究——以北京市中心城区为例
The composition of common woody plant species and their influence on bird communities in urban green areas
DOI:10.3724/SP.J.1003.2014.13168
[本文引用: 2]
Urban green areas are important habitats for birds in rapidly urbanized areas. The composition of woody plant species has a significant influence on the structure of bird communities in urban green areas. Therefore, to enhance the function of public green areas as wild bird habitats, we studied how woody plants influenced the composition of the bird community in Binjiang Forest Park, a typical urban green area in Shanghai city. A total of 5,368 individual birds belonging to 64 species from 25 families and 10 orders were recorded during line transect and point-count surveys conducted between 2009 and 2010. Data on bird abundance showed significant differences among months, with the two highest abundances recorded in April and November. A plot of the data on bird species richness against time (months) showed a curve with two peaks, with the highest abundance recorded in April and December. The values of the Simpson index showed significant variation in bird diversity over the year, with the highest value recorded in October and the lowest in August. With square-plot (400 m2) sampling analysis, a total of 77 woody plant species in the park was identified. Only 14 of these species were recorded in at least five plots (nine trees and five shrubs). These species were considered common woody species. The fruiting seasons of 12 of these species occurred during periods between September and the following February. The principal outbreaks of insects with significance to forestry occurred from May to November. A principal component analysis was used to group all birds observed on the 14 woody plant species into 8 guilds according to their diets, foraging behaviors, and distributions in various vegetation layers. A Spearman correlation analysis indicated that the autumn-winter fruiting season was significantly positively correlated with 5 bird guilds, including vegetarians, omnivores, raptors, and insectivores. The abundant food supply coincided with the highest bird abundance and richness in autumn and winter. An analysis of the importance of each common woody plant species for the bird community was also performed based on the evaluation of the relative richness, relative abundance, and relative number of bird guilds. The results showed that during autumn and winter, fruiting plants provided rich food resources for terrestrial migratory birds; trees such as Populus lasiocarpa, Pterocarya stenoptera, and Cinnamomum camphora had a greater importance than shrubs. The relatively lower importance of the common shrub species to birds implies that the shrub community of the park needs to be reviewed. Comments and recommendations are provided based on our research results.
城市公共绿地常见木本植物组成对鸟类群落的影响
DOI:10.3724/SP.J.1003.2014.13168
[本文引用: 2]
快速城市化背景下, 城市公共绿地已经成为重要的鸟类栖息地, 其中的木本植物群落构成对鸟类群落结构有显著影响, 研究木本植物配置与鸟类多样性的关系对提升城市公共绿地作为鸟类栖息地的生态服务功能有重要的理论和应用价值。我们于2009–2010年间在上海市滨江森林公园就木本植物和鸟类群落的相互关系展开研究。样线法结合样点调查共记录到鸟类10目25科64种5,368只(次), 鸟类多度全年变化显著, 峰值分别出现在4月和11月。丰富度全年也呈现双峰型变化, 峰值出现在4月和12月。全年鸟类多样性(Simpson指数)差异显著, 10月最高, 8月最低。样方调查共记录到77种木本植物, 其中有14种(乔木9种、灌木5种)出现率超过5个样方, 定义为常见种, 其果期主要集中在9月到翌年2月, 其上常见林业致病害虫的发生盛期集中在5–11月。主成分分析显示, 常见木本植物上观察到的鸟类可划分为8个鸟类集团, Spearman秩相关检验显示秋冬季常见木本植物果期与植食性、肉食性、食虫性、杂食性等多个鸟类集团的多度均呈显著正相关。说明在秋冬季鸟类迁徙高峰期不同鸟类集团均能获得丰富的食物资源。就常见木本植物对鸟类群落的重要值进行排序, 结果显示, 大叶杨(Populus lasiocarpa)、枫杨(Pterocarya stenoptera)、香樟(Cinnamomum camphora)等乔木对鸟类重要值较高, 而灌木层对鸟类的重要性整体偏低, 说明滨江森林公园的灌木层作为鸟类栖息地的功能建设尚需加强。根据以上研究结果, 我们提出了提高上海城市公园鸟类多样性的植被配置建议。
Application of food source tree species and the construction of ecological landscape in urban green space of Guangzhou
广州城市园林绿地食源树种应用及其生态景观营造
Bird community composition and its seasonal variation in comprehensive communities in Beijing: A case study of Chinese Academy of Forestry
北京综合性社区鸟类群落构成及其季节变化——以中国林业科学研究院社区为例
The influence of habitat types on bird community in urban parks
城市公园生境类型对鸟类群落的影响
Evaluation of microhabitats for wild birds in a Shanghai urban area park
Habitat use of urban tree sparrows in the process of urbanization: Beijing as a case study
DOI:10.1360/biodiv.060013
[本文引用: 1]
The tree sparrow (<i>Passer montanus</i>) is a dominant species of urban bird communities. With the development of urbanization, the habitats and food resources of tree sparrows are likely to decrease. Can the urban tree sparrow adapt to changes in the urban environment? To answer this question, we studied the habitat use of tree sparrows in eight types of urban areas in Beijing. The results showed that the number of both breeding and wintering tree sparrows decreases with increasing urbanization. The habitat use of tree sparrows, analyzed by discriminant analysis, was positively correlated with the number of brick bungalows, coniferous and broad-leaved trees, and air conditioners, whereas it was negatively correlated with the area of high buildings and hardened roads, and people and automobile flux. This indicates that the tree sparrow is not adaptable to rapid urbanization, although in general it is an adaptable species. Urban planning should take into consideration birds such as tree sparrows.
城市化对城市麻雀栖息地利用的影响: 以北京市为例
DOI:10.1360/biodiv.060013
[本文引用: 1]
本文以北京市为例研究了城市化水平不同的8个区域中麻雀(Passer montanus的数量分布现状以及影响麻雀分布的栖息地因子。结果表明, 越冬期和繁殖期的麻雀数量均与城市化程度呈显著的负相关关系; 城市化程度高的城市中心商业区、高层居民区和城市主干道中的麻雀数量均很少; 城市化程度较低的城乡结合区、公园、城市的平房区及古建筑区域中麻雀数量均较多; 高校校园和低层楼房居民区, 虽然城市化程度相对较高, 但由于植被较丰富, 麻雀数量也较多。平房面积、针叶树数量、阔叶树数量、空调数量、高层楼房的面积、硬化地面的面积、人流量及车流量是影响麻雀栖息地利用的重要因素。其中平房面积、针叶树、阔叶树、空调的数量增加, 可为麻雀提供栖息条件而有利于麻雀的利用; 硬化地面的面积、高层楼房的面积、人流量及车流量的增加, 由于减少了杂草等麻雀的食物来源并增加了干扰, 不利于麻雀的利用。结果表明, 虽然麻雀是一个适应人类生活环境的物种, 但在快速的城市化变迁中,它已表现出对高度城市化环境的不适应。在城市的规划与建设中, 应考虑到为以麻雀为代表的城市鸟类提供生存必要的植被和繁殖场所, 构建人鸟和谐共存的生态城市。
Study of bird preference to plant habitat and species in Beijing urban park
北京城市公园中鸟类对植物生境及种类的偏好研究
Arboreal camera trapping: A reliable tool to monitor plant-frugivore interactions in the trees on large scales
DOI:10.1002/rse2.v8.1 URL [本文引用: 1]
A review on the urban green space pattern affecting avian community
城市绿地空间格局对鸟类群落影响的研究进展
