林业科学 ›› 2019, Vol. 55 ›› Issue (11): 153-162.doi: 10.11707/j.1001-7488.20191117
扈梦梅1,田龙1,吴亚楠1,杨晋宇1,2,3,*,吕小翠1,黄选瑞1,2,3
收稿日期:
2019-03-10
出版日期:
2019-11-25
发布日期:
2019-12-21
通讯作者:
杨晋宇
基金资助:
Mengmei Hu1,Long Tian1,Yanan Wu1,Jinyu Yang1,2,3,*,Xiaocui Lü1,Xuanrui Huang1,2,3
Received:
2019-03-10
Online:
2019-11-25
Published:
2019-12-21
Contact:
Jinyu Yang
Supported by:
摘要:
目的: 比较研究塞罕坝地区华北落叶松人工林间伐和混交改造对大型土壤动物群落组成和多样性的影响,为实现华北落叶松人工林质量精准提升和可持续利用提供理论依据和技术支撑。方法: 以塞罕坝机械林场未经营的华北落叶松人工林(CK)、抚育间伐后的华北落叶松人工林(TH)和引入阔叶树的华北落叶松白桦混交林(M)为对象,于2014年5-9月,利用手拣法调查大型土壤动物群落,研究不同经营措施林分内土壤动物的结构组成、空间分布、多样性和功能群组成的差异及其影响因素。结果: 共获得大型土壤动物55类1 405头,隶属2门4纲12目,优势类群为正蚓目、叶甲科幼虫、象甲科幼虫。总体上,经间伐和混交改造后华北落叶松人工林内大型土壤动物的类群数显著高于未经营的对照林分(P < 0.05),TH内土壤动物密度亦显著高于CK(P < 0.05)。不同经营措施林分大型土壤动物类群数均表现为沿土壤剖面位置上升逐渐增加的趋势,且M和TH凋落物层土壤动物类群数及0~10 cm土层土壤动物密度显著高于CK(P < 0.05)。间伐和混交改造后,土壤动物群落各多样性指数均有提高,其中M的Shannon-Wiener多样性指数(H)显著高于CK(P < 0.05)。各功能群中,植食性土壤动物所占比例(47.19%)最高,其次为腐食性土壤动物(33.31%);捕食性(19类)和植食性(18类)土壤动物的类群数居多;多因素方差分析显示,经营措施、土层和时间均对大型土壤动物功能群组成和结构影响显著(P < 0.05),其中间伐和混交改造均可显著提高植食性土壤动物的类群数。冗余度分析(RDA)显示,凋落物全碳含量(P=0.03)和C/N(P=0.08)对凋落物层大型土壤动物分布影响显著;而土壤含水量(P=0.008)、pH(P=0.012)和土壤密度(P=0.062)对土壤层大型土壤动物分布影响显著。结论: 塞罕坝华北落叶松人工林经间伐和混交改造后,林分结构和微环境的改善可提高大型土壤动物的种类、数量和多样性水平,尤其是部分功能群种类增加可能更有利于凋落物早期的粉碎和分解。
中图分类号:
扈梦梅,田龙,吴亚楠,杨晋宇,吕小翠,黄选瑞. 塞罕坝华北落叶松人工林间伐和混交改造对大型土壤动物群落结构的影响[J]. 林业科学, 2019, 55(11): 153-162.
Mengmei Hu,Long Tian,Yanan Wu,Jinyu Yang,Xiaocui Lü,Xuanrui Huang. Influences of Thinning and Mixed Transformation of Larix principis-rupprechtii Plantations on the Community Structure of Soil Macro Faunal in Saihanba Area[J]. Scientia Silvae Sinicae, 2019, 55(11): 153-162.
表1
塞罕坝华北落叶松人工林研究样地概况①"
林分特征Forest characteristics | 未经营CK | 间伐TH | 混交M |
林龄Forest age/a | 28 | 28 | 28 |
乔木密度Forest density/(tree·hm-2) | 2 082±142.03a | 1 015±55.67b | 1 170±75.50b |
平均树高Mean tree height/m | 11.00±0.23b | 13.80±0.31a | 10.53±1.12b |
平均胸径Mean DBH/cm | 14.80±0.06b | 19.76±1.31a | 13.94±0.38b |
郁闭度Canopy closure(%) | 71.67±2.33a | 59.00±3.01b | 65.33±2.03ab |
土壤pH Soil pH | 6.09±0.02a | 6.12±0.02a | 6.06±0.03a |
土壤全氮含量Soil total nitrogen/(g·kg-1) | 1.80±0.01a | 2.11±0.02a | 1.24±0.02b |
土壤全碳含量Soil total carbon/(g·kg-1) | 25.31±1.54a | 22.49±1.15a | 16.65±1.39b |
土壤含水量Soil water content(%) | 17.50±1.21a | 17.11±1.53a | 17.65±0.01a |
土壤密度Soil density/(g·cm-3) | 0.70±0.07a | 0.73±0.07a | 0.75±0.06a |
凋落物含水量Litter water content(%) | 39.00±4.04a | 33.33±5.24a | 29.00±1.16a |
凋落物厚度Litter depth/cm | 6.00±0.41a | 6.13±0.21a | 5.27±0.29a |
凋落物生物量Litter biomass/(kg·m-2) | 5.90±0.72a | 5.98±0.59a | 4.97±0.51a |
凋落物全氮含量Litter total nitrogen/(g·kg-1) | 7.91±0.05b | 7.82±0.02b | 12.24±0.08a |
凋落物全碳含量Litter total carbon/(g·kg-1) | 168.70±4.13a | 171.40±13.19a | 150.90±1.52a |
附表1
大型土壤动物组成①"
类群Group | 未经营CK | 间伐TH | 混交M | 总个体数Total | 百分比Percentage(%) | |||||
个体数Individuals | 百分比Percentage(%) | 个体数Individuals | 百分比Percentage(%) | 个体数Individuals | 百分比Percentage(%) | |||||
正蚓目Lumbricida | 63 | 20 | 57 | 9.56 | 26 | 5.26 | 146 | 10.39 | ||
颤蚓目Tubificida | 9 | 2.85 | 1 | 0.17 | 18 | 3.64 | 28 | 1.99 | ||
石蜈蚣目Lithobiomorpha | 13 | 4.13 | 28 | 4.69 | 12 | 2.43 | 53 | 3.77 | ||
地蜈蚣目Geophilomorpha | 4 | 1.27 | 5 | 0.84 | 9 | 1.82 | 18 | 1.28 | ||
盲蛛目Opiliones | 5 | 1.59 | 2 | 0.34 | 3 | 0.61 | 10 | 0.71 | ||
球蛛科Theridiidae | — | — | 4 | 0.67 | 5 | 1.01 | 9 | 0.64 | ||
漏斗蛛科Agelenidae | 2 | 0.63 | 1 | 0.17 | — | — | 3 | 0.21 | ||
肖蛸科Tetragnathidae | — | — | 2 | 0.34 | 1 | 0.20 | 3 | 0.21 | ||
圆颚蛛科Corinnidae | — | — | 1 | 0.17 | — | 1 | 0.07 | |||
平腹蛛科Gnaphosidae | — | — | — | — | 2 | 0.41 | 2 | 0.14 | ||
狼蛛科Lycosidae | — | — | 7 | 1.17 | 7 | 1.42 | 14 | 1.00 | ||
蟹蛛科Thomisidae | — | — | 3 | 0.50 | 4 | 0.81 | 7 | 0.50 | ||
跳蛛科Salticidae | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
圆珠科Araneidae | 5 | 1.59 | 2 | 0.34 | 4 | 0.81 | 11 | 0.78 | ||
伪蝎目Pseudoscorpiones | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
啮虫目Psocoptera | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
叶蝉科Cicadellidae | 2 | 0.63 | 8 | 1.34 | 2 | 0.41 | 12 | 0.86 | ||
同蝽科Acanthosomatidae | — | — | 1 | 0.17 | 14 | 2.83 | 15 | 1.07 | ||
蚜科Aphididae | 1 | 0.32 | 1 | 0.17 | 1 | 0.20 | 3 | 0.21 | ||
长蝽科Lygaeidae | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
奇蝽科Enicocephalidae | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
盲蝽科Miridae | 1 | 0.32 | 2 | 0.34 | 8 | 1.62 | 11 | 0.78 | ||
土蝽科Cydnidae | — | — | 2 | 0.34 | 1 | 0.20 | 3 | 0.21 | ||
蝽科Pentatomidae | — | — | 3 | 0.50 | 3 | 0.61 | 6 | 0.43 | ||
步甲科/成Carabidae/A | 8 | 2.54 | 18 | 3.02 | 34 | 6.88 | 60 | 4.27 | ||
阎甲科/成Histeridae/A | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
象甲科/成Curculionidae/A | — | — | 2 | 0.34 | — | — | 2 | 0.14 | ||
瓢甲科/成Coccinellidae/A | — | — | 4 | 0.67 | 1 | 0.20 | 5 | 0.36 | ||
金龟科/成Scarabaeidae/A | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
叶甲科/成Chrysomelidae/A | 4 | 1.27 | 27 | 4.53 | 14 | 2.84 | 45 | 3.21 | ||
拟花蚤科/成Scraptiidae/A | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
隐翅甲科/成Staphylinidae/A | 1 | 0.32 | 5 | 0.84 | 2 | 0.41 | 8 | 0.57 | ||
埋葬甲科/成Silphidae/A | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
水龟甲科/成Hydrophilidae/A | — | 2 | 0.34 | 2 | 0.14 | |||||
水龟甲科/幼Hydrophilidae/L | 2 | 0.63 | — | — | — | — | 2 | 0.14 | ||
阎甲科/幼Histeridae/L | 1 | 0.32 | — | — | — | — | 1 | 0.07 | ||
隐翅甲科/幼Staphylinidae/L | 1 | 0.32 | 10 | 1.68 | 10 | 2.02 | 21 | 1.50 | ||
叶甲科/幼Chrysomelidae/L | 68 | 21.59 | 167 | 28.02 | 61 | 12.35 | 296 | 21.07 | ||
金龟科/幼Scarabaeidae/L | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
歩甲科/幼Carabidae/L | 1 | 0.32 | — | — | — | — | 1 | 0.07 | ||
象甲科/幼Curculionidae/L | 33 | 10.47 | 110 | 18.45 | 126 | 25.51 | 269 | 19.15 | ||
长角亚目/幼Nematocera/L | 16 | 5.08 | 44 | 7.38 | 35 | 7.09 | 95 | 6.76 | ||
短角亚目/幼Brachycera/L | 35 | 11.11 | 39 | 6.54 | 57 | 11.54 | 131 | 9.33 | ||
芒角亚目/幼Aristocera/L | 18 | 5.71 | 9 | 1.51 | 7 | 1.42 | 34 | 2.42 | ||
短角亚目Brachycera | 1 | 0.32 | — | — | — | — | 1 | 0.07 | ||
长角亚目Nematocera | 1 | 0.32 | — | — | 1 | 0.20 | 2 | 0.14 | ||
蚁科Formicidae | 1 | 0.32 | 6 | 1.00 | 7 | 1.42 | 14 | 1.00 | ||
胡蜂科Vespidae | — | — | — | — | 1 | 0.20 | 1 | 0.07 | ||
叶蜂科Tenthredinidae | — | — | 2 | 0.34 | 3 | 0.61 | 5 | 0.36 | ||
姬蜂科Lchneumonidae | 3 | 0.95 | 3 | 0.50 | 1 | 0.20 | 7 | 0.50 | ||
小卷叶蛾科/幼Olethreutidae/L | 1 | -0.32 | — | — | — | — | 1 | 0.07 | ||
卷叶蛾科/幼Tortricidae/L | 1 | 0.32 | — | — | — | — | 1 | 0.07 | ||
夜蛾科/幼Noctuidae/L | — | — | 2 | 0.34 | 4 | 0.81 | 6 | 0.43 | ||
尺蛾科/幼Geometridae/L | 14 | 4.44 | 10 | 1.68 | 5 | 1.01 | 29 | 2.07 | ||
舟蛾科/幼Notodontidae/L | — | — | 1 | 0.17 | — | — | 1 | 0.07 | ||
总个体数Total individuals | 315 | 100 | 596 | 100 | 494 | 100 | 1 405 | 100 | ||
总类群数Total group number | 29 | — | 41 | — | 39 | — | 55 | — |
表2
土壤动物群落的垂直分布及动态变化(均值±标准误)①"
经营样地 Plot | 月份 Month | 类群数Group numbers | 平均密度Mean density/(ind·m-2) | |||||
凋落物层 Litter layer | 0~10 cm土层 0-10 cm layer | 10~20 cm土层 0-20 cm layer | 凋落物层 Litter layer | 0~10 cm土层 0-10 cm layer | 10~20 cm土层 10-20 cm layer | |||
未经营CK | 5月May | 6.00±1.15Ba | 6.00±0.00Aa | 2.00±0.58Bb | 14.08±3.65Aa | 17.04±2.97Aa | 11.85±2.60Aa | |
7月July | 6.00±1.00Aa | 5.00±0.58Aab | 2.00±0.58Bb | 13.71±5.46Aa | 19.26±5.46Ba | 13.71±1.62Ba | ||
9月Sep. | 4.33±1.76Ba | 3.33±0.67Aa | 3.33±0.33Aa | 16.67±8.98Aa | 6.30±1.85Aa | 4.08±0.74Aa | ||
间伐TH | 5月May | 12.00±2.89Aa | 7.67±1.76Ab | 6.00±1.00Ab | 22.22±7.70Aa | 24.82±3.65Aa | 16.30±0.74Aa | |
7月July | 9.00±1.15Aa | 7.00±1.15Aab | 4.67±0.88ABb | 27.41±6.46Ab | 52.97±10.23Aa | 53.71±28.51Aa | ||
9月Sep. | 6.33±0.33ABa | 3.00±1.15Aa | 4.00±0.58Aa | 10.37±2.67Aa | 5.56±2.57Aa | 7.41±2.06Aa | ||
混交M | 5月May | 7.33±1.76Ba | 6.33±2.33Aa | 4.33±1.45ABa | 12.97±2.97Aa | 31.85±7.76Aa | 7.04±2.60Aa | |
7月July | 9.00±2.65Aa | 8.33±0.88Aa | 6.33±0.88Aa | 19.26±4.18Ab | 60.00±4.01Aa | 32.97±10.82ABb | ||
9月Sep. | 9.67±0.88Aa | 4.67±1.20bAa | 2.00±0.58Bb | 18.89±2.57Aa | 12.22±1.29Aa | 3.71±1.48Aa |
表3
大型土壤动物群落多样性指数(平均值±标准误)"
经营样地Plot | H | Jd | H | Jd | |||||
5月May | 7月July | 9月Sep. | 5月May | 7月July | 9月Sep. | ||||
未经营CK | 2.21±0.01B | 0.90±0.01A A | 1.66±0.18Ba | 1.66±0.20Aa | 1.79±0.170Aa | 0.82±0.04Aa | 0.80±0.08Aa | 0.90±0.06Aa | |
间伐TH | 2.37±0.07AB | 0.87±0.03A | 2.36±0.22Aa | 1.75±0.13Ab | 2.13±0.07Aab | 0.91±0.03Aa | 0.83±0.06Aa | 0.93±0.02Aa | |
混交M | 2.52±0.09A | 0.91±0.01A | 2.32±0.12ABa | 1.89±0.15Aa | 2.22±0.16Aa | 0.95±0.02Aab | 0.83±0.01Ab | 0.95±0.01Aa |
表4
大型土壤动物功能群多因素方差分析①"
项目Source | 腐食性Saprophagous | 捕食性Predatism | 植食性Phytophagy | 杂食性Omnivory | |||||||
平均密度 Density | 类群 Group | 平均密度 Density | 类群 Group | 平均密度 Density | 类群 Group | 平均密度 Density | 类群 Group | ||||
经营措施Man. | 0.804 | 0.067 | 0.040* | 0.161 | 0.067 | 0.003** | 0.005** | 0.141 | |||
土层Lay. | 0.000** | 0.000** | 0.018* | 0.000** | 0.021** | 0.150 | 0.000** | 0.000** | |||
月份Mon. | 0.004** | 0.021* | 0.036* | 0.052 | 0.000** | 0.169 | 0.025* | 0.029* | |||
经营措施×取样层次Man.×Lay. | 0.065 | 0.194 | 0.690 | 0.539 | 0.489 | 0.331 | 0.095 | 0.578 | |||
月份×经营措施Mon.×Man. | 0.092 | 0.001** | 0.295 | 0.392 | 0.064 | 0.105 | 0.044* | 0.103 | |||
月份×取样层次Mon.×Lay. | 0.555 | 0.489 | 0.835 | 0.880 | 0.002** | 0.003* | 0.043* | 0.126 | |||
月份×经营措施×取样层次Mon.×Man.×Lay. | 0.331 | 0.127 | 0.942 | 0.571 | 0.529 | 0.573 | 0.005** | 0.845 |
图2
赛罕坝华北落叶松人工林大型土壤动物群落结构与土壤环境因子的关系 OC:有机碳Organic carbon; TN:全氮Total nitrogen; LD:凋落物厚度Litter thickness; WC:含水量Water content; LB:凋落物生物量Litter biomass; pH: pH值pH value; VW:土壤密度Soil bulk density; C/N:碳氮比Carbon nitrogen ratio; Lum.:正蚓目Lumbricina; Tub.:颤蚓目Tubificida; Lytho.:石蜈蚣目Lithobiomorpha; Geo.:地蜈蚣目Geophilomorpha; Pha.:盲蛛目Phalangida; The.:球蛛科Theridiidae; Age.:漏斗蛛科Agelenidae; Tet.:肖蛸科Tetragnathidae; Gna.:平腹蛛科Gnaphosidae; Lyco.:狼蛛科Lycosidae; Tho.:蟹蛛科Thomisidae; Ara.:圆珠科Araneidae; Cic.:叶蝉科Cicadellidae; Aca.:同蝽科Acanthosomatidae; Aph.:蚜科Aphididae; Mir.:盲蝽科Miridae; Cyd.:土蝽科Cydnidae; Pen.:蝽科Pentatomidae; Cara.(A):步甲科Carabidae adult; Cocc.(A):瓢甲科(成)Coccinellidae; Chry.(A):叶甲科(成)Chrysomelidae adult; Sta.(A):隐翅甲科(成)Staphylinidae adult; Sta.(L):隐翅甲科(幼)Staphylinidae larvae; Chry.(L):叶甲科(幼)Chrysomelidae larvae; Cur(L):象甲科(幼)Curculionidae larvae; Nema.(L):长角亚目(幼)Nematocera larvae; Bra.(L):短角亚目(幼)Brachycera larvae; Ari.(L):芒角亚目(幼)Aristocera larvae; Nema.(A):长角亚目(成)Nematocera adult; For.:蚁科Formicidae; Cim.(A):叶蜂科(成)Cimbicidae adult; Lch.:姬蜂科Lchneumonidae; Noc.(L):夜蛾科(幼)Noctuidae larvae; Goc.(L):尺蛾科(幼)Geometridae larvae. "
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