四川低山丘陵区香樟和马尾松凋落叶分解进程中土壤节肢动物多样性

doi:10.11707/j.1001-7488.20211112

摘要/Abstract

摘要：

目的: 研究四川低山丘陵区香樟和马尾松凋落叶分解进程中土壤节肢动物群落结构及其动态变化，为深入认识森林土壤动物与凋落物分解的相互作用和联系提供参考。方法: 2010年11月，在香樟和马尾松林样地安装15个凋落叶收集器，一年内每月收集2种树种凋落叶，带回实验室自然风干；2011年10月，将风干凋落叶分树种装入20 cm×20 cm分解袋中，分解袋上下孔径分别为3 mm和0.04 mm；2011年11月，将凋落叶分解袋对应埋置于样地内5个5 m×5 m样方中；2011年12月-2015年10月，分11次采集凋落叶分解袋，采用Tullgren漏斗分离收集土壤节肢动物，利用体式解剖镜和生物显微镜镜检计数并分类。结果: 整个分解期间，2种凋落叶中共捕获土壤节肢动物4 648只，隶属7纲17目58科，香樟凋落叶中以等节跳科、棘跳科、丽甲螨科和懒甲螨科为优势类群，马尾松凋落叶中多为常见类群。香樟和马尾松2种凋落叶中土壤节肢动物的个体密度和类群数量具有相似的动态变化过程，个体密度分别在分解的746和398天最高，类群数量均在分解的746天最大；土壤节肢动物总体以菌食性数量比例最高，分别占总捕获量的39.48%和38.78%，腐食性数量比例最低，分别占总捕获量的8.02%和8.05%，且菌食性土壤节肢动物占比随凋落叶分解进程逐渐上升，而捕食性和植食性土壤节肢动物占比则不断下降。非度量多维尺度（NMDS）分析显示，2种凋落叶中土壤节肢动物群落组成相似性较高；聚类分析表明，2种凋落叶中土壤节肢动物群落相似性随凋落叶分解进程不断降低。结论: 四川低山丘陵区香樟和马尾松凋落叶分解进程中，凋落物类型和分解时间对土壤节肢动物群落结构影响显著。与香樟（阔叶树）凋落叶相比，马尾松（针叶树）凋落叶对土壤节肢动物的特定类群依赖度更高。不同凋落物类型下，土壤节肢动物群落结构和多样性在分解中期（第2~3年）差异最明显，有必要长期监测凋落物分解进程中的土壤动物响应。

关键词: 土壤节肢动物, 功能类群, 凋落物类型, 凋落物分解, 亚热带森林

Abstract:

Objective: We conducted a 4-year (2011-2015) litterbag experiment with Cinnamomum camphora and Pinus massoniana litter to explore the dynamics of soil arthropod diversity during the litter decomposition in low mountains and hilly areas of Sichuan, and the changes of the composition and diversity of the soil arthropods community were determined. The objectives of the study are to understand the differences between the short-term and long-term changes of the composition distribution in the soil arthropod community during the long-term litter decomposition, and the changes of soil arthropod diversity during the decomposition of different types of litters. Method: In November 2010, 15 litter collectors were installed in the study site of C. camphora and P. massoniana forests. The fallen leaves of two tree species were collected monthly, and then brought back to the lab for air drying. In October 2011, the air-dried leaves (10±0.05 g) were put into prepared littebags (20 cm×20 cm, the upper and lower apertures were 3 mm and 0.04 mm, respectively). In November 2011, all the litterbags were buried in five (5 m×5 m) plots on the forest floor and the buried litter decomposition bags were retrieved 11 times from December 2011 to October 2015. The soil arthropods were separated and collected by Tullgren funnel, and then counted and classified under anatomical microscope and biological microscope. Result: During the 4-year study period, a total of 4 648 soil arthropods were captured in the litterbags of C. camphora and P. massoniana, belonging to 17 orders and 58 families. The dominant groups of C. camphora were Isotomidae, Onychiuriclae, Liacaridae and Nothridae. The individual density and group number of soil arthropods in the two litter leaves had similar dynamics. The individual density of soil arthropod community in C. camphora reached the highest at 746 d and fell rapidly afterwards, while the that in P. massoniana reached the highest at 398 d. The two communities reached maximum at 746 d, with similar dynamics of group number in the litter decomposing processes. Fungivore arthropods had the highest proportion in the both litters, accounting for 39.48% and 38.78% of the total catch, respectively. The saprophagous arthropods presented the lowest, accounting for 8.02% and 8.05%, respectively. With the decomposition process of litter, phytophagy and predatory arthropods reduced, while the fungivore arthropods increased. Non-metric multidimensional scale (NMDS) results showed a high similarity in the composition of the two soil arthropod communities. Cluster analysis showed that the similarity of soil arthropod community structure diminished with the decomposition process. Conclusion: The soil arthropod community structure is significantly affected by litter types and decomposition phase during litter decomposition in low mountains and hills area. Compared with broadleaf litter, coniferous litter is more dependent on specific groups of soil arthropods. The differences in soil arthropod community composition and diversity in disparate litter types are most obvious in the mid-decomposition phase (2-3 years), indicating the importance of long-term monitoring of soil fauna response to litter decomposition.

Key words: soil arthropods, functional group, litter types, litter decomposition, subtropical forest

中图分类号:

S718.6

刘静如,曹艺,李晗,张丽,游成铭,徐振锋,谭波. 四川低山丘陵区香樟和马尾松凋落叶分解进程中土壤节肢动物多样性[J]. 林业科学, 2021, 57(11): 119-133.

Jingru Liu,Yi Cao,Han Li,Li Zhang,Chengming You,Zhenfeng Xu,Bo Tan. Diversity of Soil Arthropods during Cinnamomum camphora and Pinus massoniana Litter Decomposition in Low Mountainous and Hilly Areas of Sichuan[J]. Scientia Silvae Sinicae, 2021, 57(11): 119-133.

图/表 10

表1

图1

图2

表附表 1

香樟凋落叶中土壤节肢动物组成及动态变化①"

土壤动物类群Soil fauna groups			平均密度(每袋个体数) Mean densities (number per litterbag)														功能类群 Functional groups
土壤动物类群Soil fauna groups			35 d	156 d	277 d	398 d	516 d	628 d	746 d	896 d	1 079 d	1 261 d	1 444 d	合计 Total	占比 Percentage(%)	优势度 Abundance	功能类群 Functional groups
弹尾纲Collembola	弹尾目Collembola	鳞跳科Tomoceridae	2.4	0	0	0.8	0.4	0.4	0.6	0.2	1.0	0.4	0.2	6.4	1.6	++	F
		疣跳科Neanuridae	0	0	0.6	0.4	0	2.2	0.4	0	0.6	0.2	1.0	5.4	1.3	++	F
		等节跳科Isotomidae	16.8	1.6	3.2	8.0	8.6	1.8	5.6	2.4	5.4	4.4	5.4	63.2	15.7	+++	F
		棘跳科Onychiuriclae	11.2	1.6	2.2	6.8	1.2	6.0	8.8	4.4	6.6	5.4	4.6	58.8	14.6	+++	F
		短脚圆跳科Neelidae	0	0	0	0	0	0	0.4	0.6	0	0	0	1.0	0.2	+	F
		长脚跳科Entomobryidae	1.6	0	0	0	0	0	0.2	0	0	0	0	1.8	0.4	+	F
		驼跳科Cyphoderidae	4.0	0	2.8	0	0.4	0	2.0	0.8	1.0	0.8	0	11.8	2.9	++	F
		球角跳科Hypogastruridae	3.2	0	0	2.0	0	2.2	2.6	1.0	1.0	1.0	0.4	13.4	3.3	++	H
		圆跳科Sminthuridae	4.8	0	1.6	3.2	2.2	0	0.4	0.4	0	0.6	0	13.2	3.3	++	H
蛛形纲Arachnoidea	蜱螨目Acarina	土革螨科Ologamasidae	4.0	1.6	2.4	0	0.8	6.2	1.6	1.0	1.4	0	0	19.0	4.7	++	P
		绥螨科Sejidae	0	2.4	2.6	0	0	0.8	0.6	0.2	0.6	0	0	7.2	1.8	++	P
		植绥螨科Phytoseiidae	1.6	0	0	0	0.4	0.4	0	0	0	0	0	2.4	0.6	+	P
		厚厉蟎科Pachylaelapidae	0.8	0	0.8	0	0	0	0.2	0	0	0	0	1.8	0.4	+	P
		厉螨科Laeclapidae	1.6	1.6	1.6	0.4	2.2	0.4	0	0	0	0	0	7.8	1.9	++	P
		隐颚螨科Cryptognathidae	0	0	8.0	0	0	0	0.4	0.6	0.6	0.2	0	9.8	2.4	++	F
		赤螨科Erythraeidae	0	0	4.8	0	0	0	0.6	0.8	1.4	1.8	0.4	9.8	2.4	++	P
		象螨科Bdellidae	0	0	0	0	0	0	0.2	0	0	0	0	0.2	0	+	F
		长须螨科Stigmaeidae	1.6	0	2.6	0	0	0.8	1.2	1.2	1.2	0	0	8.6	2.1	++	P
		大翼甲螨科Galumnidae	1.6	0	0	0.8	0	2.2	1.2	0.4	0.6	0	0	6.8	1.7	++	P
		若甲螨科Oribatulidae	3.2	0.8	4.0	2.2	1.2	0.8	3.6	3.6	2.4	0.8	0.4	23.0	5.7	++	F
		丽甲螨科Liacaridae	4.8	4.8	1.8	3.8	0.4	12.0	9.2	6.0	3.0	1.8	0.2	47.8	11.8	+++	H
		三甲螨科Oribotritiidae	0	0	1.6	0	0	0	1.8	0.8	0.6	0	0	4.8	1.2	++	F
		奥甲螨科Oppiidae	1.6	0	1.2	0	0	0.6	3.2	4.4	3.0	1.8	0.8	16.6	4.1	++	P
		懒甲螨科Nothridae	7.2	6.4	3.2	9	0	4.2	6.6	4.0	5.6	2.8	0.6	49.6	12.3	+++	P
		洛甲螨科Lohmanniidae	0	0	0	0	0	0	0	0	0.2	0.2	0	0.4	0.1	+	F
		珠甲螨科Damaeidae	0	0	0	0	0	0	0	0.4	0	0	0	0.4	0.1	+	F
		缝甲螨科Hypochthoniidae	0	0	0	0	0	0	0.2	0	0	0	0	0.2	0	+	P
		直卷甲螨科Archoplophoridae	0.8	0	0	0.4	2.2	0	1.6	3.0	1.8	2.0	0	11.8	2.9	++	F
		四奥甲螨科Quadroppiidae	0	0	0	0	0	0.4	1.2	0.6	0	0.4	0.4	3.0	0.7	+	P
	蜘蛛目Araneae	球体蛛科Theridiosomatidae	0	0	0	0	0	0.4	0.2	0	0	0	0	0.6	0.1	+	P
		跳蛛科Salticidae	0	0	0	0	0	1.2	0.8	0.8	0.6	0.2	0	3.6	0.9	+	P
		狼蛛科Lycosidae	0	0.8	0	0	0	0.8	0.4	0	0	0.2	0	2.2	0.5	+	P
		圆蛛科Araneidae	0.8	0	0	0	0	0.6	0	0	0	0	0	1.4	0.3	+	P
		草蛛科Agelenidae	0.8	0	0	0	0	1.2	0	0	0	0	0	2.0	0.5	+	P
	伪蝎目 Pseudoscorpiones	土伪蝎科Chthonidae	0.8	0	0.8	4.0	0	0	0	0	0	0	0	5.6	1.4	++	P
	盲蛛目Opiliones	长奇盲蛛科Phalangiidae	0	0	1.4	0	0	0	0.4	0	0	0.4	0	2.2	0.5	+	P
双尾纲Diplura	双尾目Diplura	原铗叭科Projapygidae	1.6	0	0.6	0	0	0	0.8	1.6	0.4	0.4	0.2	5.6	1.4	++	H
真缓步纲Eutardigrada	离爪目Apochela	小斑科Milnesiidae	0	0	0	0	0	0.4	0	0	0	0	0	0.4	0.1	+	H
唇足纲Chilopoda	石蜈蚣目 Lithobiomorpha	石蜈蚣科Lithobiidae	0	0.8	0.6	0	0	0	1.0	1.2	0	0.4	0.4	4.4	1.1	++	P
昆虫纲Insecta	鞘翅目Coleoptera	隐翅甲科Staphylinidae	1.6	0	0	0	1.2	0	1.6	1.2	1	0.4	0.4	7.4	1.8	++	P
		叶甲科Chrysomelidae	0.8	0	0	0	0	0	0.2	0	0	0	0	1.0	0.2	+	H
	双翅目Diptera (larva)	长角蚊科幼虫Hesperinidae	0.8	0	0.4	0.4	0	0	1.6	0.4	0	0	0	3.6	0.9	+	S
		蝇科Muscidae	1.6	0	0.6	1.2	0	0	0.2	0	0	0	0	3.6	0.9	+	S
		毛蚊科Bibionidae	0	0	0.6	0.8	0	0	0	0	0	0	0	1.4	0.3	+	S
		摇蚊科Chironomidae	2.4	1.6	1.6	0	0	0	0.8	1.8	0	0.6	0.6	9.4	2.3	++	S
		长足蚊科幼虫Dolichopdidae	0	0	0	0	0	0	0.2	0	0	0	0	0.2	0	+	P
	啮虫目Psocoptera	球啮科Sphaeropsocidae	2.4	0	0	0	0	2.8	3.4	1.2	0	0	0	9.8	2.4	++	S
	缨翅目Thysanoptera	管蓟马科Phlaeothripidae	0	0	1.2	0	0	0	0.4	0.6	0	0	0	2.2	0.5	+	H
	蜚蠊目Blattoptera	光蠊科Epilampridae	0	0.8	0	0	0	0.4	1.6	3.2	3.0	2.0	1.2	12.2	3.0	++	S
	膜翅目Hymenoptera	蚁科Formicidae	0.8	3.2	2.8	0	0	4.2	3.2	1.6	0.6	0	1.2	17.6	4.4	++	H
		切叶蚁亚科Myrmicinae	0	0	0.2	0	0	0	0	0	0	0	0	0.2	0	+	H
	鳞翅目Lepidoptera	毒蛾科Lymantriidae	0	0	0	0	0.4	0	0	0	0	0	0	0.4	0.1	+	H
		尺蛾科Geometridae	0	0	0	0	0.4	0	0	0	0	0	0	0.4	0.1	+	S
	直翅目Orthoptera	蟋蟀科Gryllidae	0	0	0	0	0	0	0.2	0	0	0	0	0.2	0	+	H
	合计Total		87.2	28.0	56.4	44.2	22.4	53.4	71.6	50.4	43.6	29.2	18.4	503.6	100.0	―	―
	类群数 Number of groups		29	13	29	16	15	25	42	31	22	24	16	54	―	―	―

表附表 1

表附表 2

马尾松凋落叶中土壤节肢动物组成及动态变化"

土壤动物类群Soil fauna groups			平均密度(每袋个体数) Mean densities (number per litterbag)														功能类群 Functional groups
土壤动物类群Soil fauna groups			35 d	156 d	277 d	398 d	516 d	628 d	746 d	896 d	1 079 d	1 261 d	1 444 d	合计 Total	占比 Percentage(%)	优势度 Abundance	功能类群 Functional groups
弹尾纲Collembola	弹尾目Collembola	鳞跳科Tomoceridae	1.6	0	0	0.4	0	0	0	0	0	0	0	2.0	0.5	+	F
		疣跳科Neanuridae	2.4	0	3.0	2.8	1.8	0	1.8	1.0	0.6	0	0	13.4	3.2	+	F
		等节跳科Isotomidae	3.2	1.6	8.4	12.8	2.4	1.2	3.8	1.8	2.0	1.8	2.2	41.2	9.7	++	F
		棘跳科Onychiuriclae	4.8	1.6	8	6.2	0.8	2.4	7.2	3.6	2.0	3.6	1.6	41.8	9.8	++	F
		短脚圆跳科Neelidae	2.4	0	0	3.4	0	0	0.4	0	0	0	0	6.2	1.5	++	F
		长脚跳科Entomobryidae	0	0.8	0.4	0.8	0	0	0	0	0	0	0	2.0	0.5	+	F
		驼跳科Cyphoderidae	1.6	0	1.6	0	0	0	0	0	0.2	0	0	3.4	0.8	+	F
		爪跳科Paronellidae	0	0	0.6	0	0	0	0	0	0	0	0	0.6	0.1	+	F
		球角跳科Hypogastruridae	1.6	0	0.4	2.2	0.4	0	0	0.2	0.2	0.4	0.4	5.8	1.4	++	H
		圆跳科Sminthuridae	2.4	0.8	1.2	1.4	2.0	0.4	1.6	0.4	0.8	0.2	0	11.2	2.6	++	H
蛛形纲Arachnoidea	蜱螨目Acarina	土革螨科Ologamasidae	4.0	0.8	1.6	3.0	2.4	0.8	0.2	0	0	0	0	12.8	3.0	++	P
		绥螨科Sejidae	0.8	0	3.2	1.2	0	0	0	0	0	0	0	5.2	1.2	++	P
		植绥螨科Phytoseiidae	1.6	0	0	3.4	0.8	0	0	0	0	0	0	5.8	1.4	++	P
		厚厉蟎科Pachylaelapidae	1.6	0.8	1.6	3.2	2.2	0.4	0	0	0	0	0	9.8	2.3	++	P
		厉螨科Laeclapidae	1.6	0	1.0	0	0	0	0	0	0	0	0	2.6	0.6	+	P
		隐颚螨科Cryptognathidae	0	0	1.4	0.4	0	0	0.4	0	0	0	0	2.2	0.5	+	F
		长须螨科Stigmaeidae	0	0	0	0	0	0	0.4	0.2	0.4	0.6	0	1.6	0.4	+	P
		赤螨科Erythraeidae	0	0	2.6	2.2	0	0	1.2	1.0	0.4	1.0	1.0	9.4	2.2	++	P
		象螨科Bdellidae	0	0	0	0	0	0	0.2	0.2	0	0	0	0.4	0.1	+	F
		大翼甲螨科Galumnidae	8.8	1.6	0	6.6	4.6	1.2	0.8	0.6	1.0	0.2	0.4	25.8	6.1	++	P
		若甲螨科Oribatulidae	8.0	0.8	1.6	6.8	2.2	2.2	3.6	1.2	2.2	0.8	1.2	30.6	7.2	++	F
		丽甲螨科Liacaridae	4.0	0.8	1.8	5.6	1.6	0	4.4	3.6	1.6	1.8	0	25.2	5.9	++	H
		三甲螨科Oribotritiidae	0.8	0	0	0	0	0	0.6	1.8	0.6	1.2	1.2	6.2	1.5	++	F
		奥甲螨科Oppiidae	4.0	0	0	12.0	0	1.2	5.0	2.6	0.8	1.8	0.2	27.6	6.5	++	P
		懒甲螨科Nothridae	6.4	1.6	2.8	4.6	6.2	0	6.4	2.0	1.8	0.8	0	32.6	7.7	++	P
		洛甲螨科Lohmanniidae	0	0	0	0	0	0	0.4	0.6	0	0	0	1.0	0.2	+	F
		珠甲螨科Damaeidae	0	0	0	0	0	0	0.4	0.2	0	0	0	0.6	0.1	+	F
		缝甲螨科Hypochthoniidae	2.4	0	0	2.2	2.2	1.0	0	0	0	0	0	7.8	1.8	++	P
		直卷甲螨科Archoplophoridae	3.2	0	0	2.2	3.6	0	1.8	1.2	0.4	0.6	0.2	13.2	3.1	++	F
		四奥甲螨科Quadroppiidae	0	0	0.8	0.8	0	1.2	1.6	0.6	0.6	1.0	0	6.6	1.6	++	P
	蜘蛛目Araneae	球体蛛科Theridiosomatidae	0.8	0	0	0	0	0	0	0	0	0	0	0.8	0.2	+	P
		跳蛛科Salticidae	0.8	0.8	0.4	0	0	0	0.4	0.4	0	0	0	2.8	0.7	+	P
		圆蛛科Araneidae	0.8	0	0	0.4	0	0	0.4	0	0	0	0	1.6	0.4	+	P
	伪蝎目 Pseudoscorpiones	土伪蝎科Chthonidae	0	0	0.6	0	0	0	0	0	0	0	0	0.6	0.1	+	P
	盲蛛目Opiliones	长奇盲蛛科Phalangiidae	0	0	0	0	0	0.8	0.2	0	0	0	0	1.0	0.2	+	P
双尾纲Diplura	双尾目Diplura	原铗叭科Projapygidae	0.8	0	0.6	0	0	0	0.4	0.6	0	0	0	2.4	0.6	+	H
唇足纲Chilopoda	石蜈蚣目 Lithobiomorpha	石蜈蚣科Lithobiidae	0	0	0.2	0	0	0	0.6	0.4	0.6	0.2	0.2	2.2	0.5	+	P
综合纲Symphyla		幺蚰科Scutigerellidae	2.4	0	0	0	0	0	0	0	0	0	0	2.4	0.6	+	H
昆虫纲Insecta	鞘翅目Coleoptera	隐翅甲科Staphylinidae	0	0	0.2	0	0.8	0	0.4	0.4	0.2	0.2	0.2	2.4	0.6	+	P
		蚁甲科Pselaphidae	0	0	0	0	0.4	0	0	0	0.2	0	0	0.6	0.1	+	S
		叶甲科Chrysomelidae	0	0	0	0	0	0	0.2	0.2	0	0	0	0.4	0.1	+	H
	双翅目Diptera	长角蚊科幼虫Hesperinidae	0	0	0	0.4	0	0	0.2	0.2	0.4	0	0	1.2	0.3	+	S
		蝇科Muscidae	0	0	0	0.8	0	0	0	0	0.2	0	0	1.0	0.2	+	S
		摇蚊科Chironomidae	0.8	1.6	2.0	2.2	8.6	0	1.8	2.2	0.4	0.4	0.4	20.4	4.8	++	S
		长足蚊科幼虫Dolichopdidae	0	0	0	2.0	2.2	0	0.2	0	0	0	0	4.4	1.0	++	P
	啮虫目Psocoptera	球啮科Sphaeropsocidae	3.2	0	0	2.2	0	0	0	1.2	0.4	0.4	0.6	8.0	1.9	++	S
	缨翅目Thysanoptera	管蓟马科Phlaeothripidae	0	0	0	0	0	0	0.4	0	0.2	0.2	0	0.8	0.2	+	H
	蜚蠊目Blattoptera	光蠊科Epilampridae	0	0	0	0	0	0	0.4	0.2	0.6	1.0	0.6	2.8	0.7	+	S
	膜翅目Hymenoptera	蚁科Formicidae	1.6	1.6	1.0	2.4	0	0	0.6	1.2	1.2	0.6	1.6	11.8	2.8	++	H
	鳞翅目Lepidoptera	毒蛾科Lymantriidae	0	0	0	0	0.8	0	0	0.4	0	0	0	1.2	0.3	+	H
		尺蛾科Geometridae	0	0	0	0	0	0	0.2	0	0	0	0	0.2	0	+	S
	半翅目Hemiptera	土蝽科Cydnidae	0	0	0	0	0	0	0.4	0.2	0	0	0	0.6	0.1	+	H
	直翅目Orthoptera	蟋蟀科Gryllidae	0	0.8	0	0	0	0	0	0	0	0	0	0.8	0.2	+	H
	合计Total		78.4	16.0	47.0	105.6	46.0	12.8	49.0	30.4	20.0	18.8	12.0	425.0	100.0	―	―
	类群数Number of groups		29	14	25	29	19	11	35	30	26	21	15	53	―	―	―

表附表 2

表2

图3

图4

图5

图6

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树种 Tree species	C/ (g·kg^-1)	N/ (g·kg^-1)	P/ (g·kg^-1)	L (%)	CL (%)	C/N	C/P	N/P	L/N	L/CL
香樟 Cinnamomum camphora	461.22 (17.70)	11.03 (0.56)	0.66 (0.05)	27.02 (0.55)	16.92 (0.49)	41.86 (1.19)	703.68 (28.99)	16.81 (0.61)	2.45 (0.10)	1.60 (0.03)
马尾松 Pinus massoniana	496.99^* (14.64)	12.18^* (0.37)	0.87^* (0.06)	24.80^* (0.72)	16.78 (0.57)	40.80^* (0.46)	576.09^* (29.12)	14.12^* (0.66)	2.04^* (0.03)	1.48 (0.01)

树种 Tree species	指示物种 Indicator species	IndVal	P
香樟C. camphora	原铗叭科Projapygidae	0.703 3	0.003
马尾松P. massoniana	叶甲科Chrysomelidae	0.719 5	0.032

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