北京野鸭湖湿地地表甲虫群落组成与空间分布格局

doi:10.3724/SP.J.1003.2009.08161

摘要/Abstract

摘要：

2007年4-10月在北京野鸭湖湿地两种保存较好的湿地植被类型和3种主要的退化植被类型中设立了32个样地, 采用陷阱法调查地表甲虫群落的组成, 并在科级水平上探讨了湿地退化及植被类型变化对地表甲虫群落组成的影响。整个采样周期共采集甲虫标本42科, 其中步甲科和隐翅虫科为优势科, 蚁形甲科、肖叶甲科和薪甲科为亚优势科。在所研究的5种植被类型中, 湿地景观保存较好的芦苇(Phragmites communis)带与球穗莎草(Cyperus glomeratus)带的甲虫群落活动密度、科丰富度和Shannon-Wiener多样性指数(H')均无显著差异, 而上述两种植被类型的甲虫活动密度以及球穗莎草带的甲虫科丰富度均显著高于3种退化的植被类型。对地表甲虫群落组成与9个环境因子进行的典范对应分析(CCA)表明, 32个样地在CCA排序图中的分布与植被类型之间存在明显的对应关系, 土壤含水量、植物盖度、植物生物量和枯落物盖度是影响地表甲虫群落组成及空间分布的主要环境因子。相关和回归分析结果也显示, 甲虫群落的活动密度与土壤含水量、植物生物量和植物盖度均极显著或显著正相关, 科丰富度与植物生物量显著正相关, 多样性指数(H')与植物盖度极显著负相关; 其中土壤含水量的变化能够解释甲虫群落活动密度总方差的57%。此外, 通过主成分分析获得了反映土壤含水量、植物生物量和植物盖度综合作用的环境变量WBC (Water-Biomass-Coverage)。依据地表甲虫活动密度与WBC的关系, 可将5种植被类型分为彼此差异极显著的3组。研究结果表明保持良好的湿地景观对于保护湿地甲虫具有重要意义。

关键词: 植被类型, 湿地, 典范对应分析, 主成分分析

Abstract

In this paper, ground-dwelling beetle communities (GDBCs) from 32 sampling sites with five main vegetation types, two well-conserved ones and three degradated ones, were investigated in Yeyahu Wetland of Beijing by pitfall traps from April to December, 2007, and the influence of wetland degradation and vegetation variation on the composition of GDBCs was studied at the family level. A total of 42 families were collected in the whole sampling period. Among these, Carabidae and Staphylinidae were dominant, while Anthicidae, Eumolpidae and Lathridiidae were subdominant. In the five types of vegetation investigated, the activity density, family level richness and the Shannon-Wiener diversity index (H') of GDBCs were not significantly different between the well-conserved Phragmites communis and Cyperus glomeratus ones. However, the beetle activity density in these two well-conserved vegetations, and the family richness in the Cyperus glomeratus one were significantly higher than those of the three degradated ones. It was showed by canonical correspondence analysis (CCA) to the relationship between the composition of GDBCs and nine environmental factors, that there was an obviously correspondance between the distribution of the 32 sampling sites in CCA ordination diagram and the vegetation types, and soil water content, plant coverage, plant biomass, and litter coverage were the major factors affecting on the composition and spatial distribution of GDBCs in the wetland. According to the correlation analysis, we also found that the correlation among beetle activity density and the soil water content, plant biomass and coverage was all significantly positive (P<0.05 or P<0.01), and it was between beetle family richness and plant biomass (P<0.05); while that between the beetle diversity index (H') and plant coverage was significantly negative (P<0.01). Moreover, in a nonlinear regression analysis, the change of soil water content could explain about 57% of the variation in beetle activity density. A synthetic environmental variable, WBC (Water-Biomass-Coverage), which reflects the situation of soil water content, plant biomass and coverage, was obtained by principal component analysis (PCA) of the nine environmental factors. On the basis of relationship between beetle activity density and WBC, the five vegetation types could be divided into three extremely different groups. Our results showed that maintaining appropriate wetland landscapes is vital for beetle protection.

Key words: vegetation type, wetland, canonical correspondence analysis, principal component analysis

王玉, 高光彩, 付必谦, 吴专 (2009) 北京野鸭湖湿地地表甲虫群落组成与空间分布格局. 生物多样性, 17, 30-42. DOI: 10.3724/SP.J.1003.2009.08161.

Yu Wang, Guangcai Gao, Biqian Fu, Zhuan Wu (2009) Composition and spatial distribution pattern of ground-dwelling beetle communities in Yeyahu Wetland, Beijing. Biodiversity Science, 17, 30-42. DOI: 10.3724/SP.J.1003.2009.08161.

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链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2009.08161

https://www.biodiversity-science.net/CN/Y2009/V17/I1/30

图/表 10

参考文献 38

[1]	Bates AJ, Sadler JP, Fowles AP (2006) Condition-dependent dispersal of a patchily distributed riparian ground beetle in response to disturbance. Oecologia, 150, 50-60.
[2]	Boscaini A, Franceschini A, Maiolini B (2000) River ecotones: carabid beetles as a tool for quality assessment. Hydrobiologia, 422, 173-181.
[3]	Brose U (2003) Bottom-up control of carabid beetle communities in early successional wetlands: mediated by vegetation structure or plant diversity? Oecologia, 135, 407-413. DOI URL
[4]	Butterfield J (1997) Carabid community succession during the forestry cycle in conifer plantations. Ecography, 20, 614-625.
[5]	Dennis P, Young MR, Gordon IJ (1998) Distribution and abundance of small insects and arachnids in relation to structural heterogeneity of grazed, indigenous grasslands. Ecological Entomology, 23, 253-264.
[6]	Du RQ (杜荣骞) (1999) Biostatistics (生物统计学). China Higher Education Press, Beijing and Springer-Verlag Heidelberg, Beijing. (in Chinese)
[7]	Epstein ME, Kulman HM (1990) Habitat distribution and seasonal occurrence of carabid beetles in East-central Minnesota. American Midland Naturalist, 123, 209-225.
[8]	Evans WG (1983) Habitat selection in the Carabidae. The Coleopterists Bulletin, 37, 164-167.
[9]	Gormley LHL, Furley PA, Watt AD (2007) Distribution of ground-dwelling beetles in fragmented tropical habitats. Journal of Insect Conservation, 11, 131-139.
[10]	Irmler U (2000) Environmental characteristics of ground beetle assemblages in northern German forests as basis for an expert system. Zeitschrift fuer Oekologie und Naturschutz, 8, 227-237.
[11]	Irmler U, Hoernes U (2003) Assignment and evaluation of ground beetle (Coleoptera: Carabidae) assemblages to sites on different scales in a grassland landscape. Biodiversity and Conservation, 12, 1405-1419.
[12]	Jennings DT, Housewear MW, Dunn GA (1986) Carabid beetles (Coleoptera: Carabidae) associated with strip clearcut and dense spruce-fir forests of Maine. The Coleopterists Bulletin, 40, 251-263.
[13]	Koivula M, Punttila P, Haila Y, Niemela J (1999) Leaf litter and the small-scale distribution of carabid beetles (Coleoptera, Carabidae) in the boreal forest. Ecography, 22, 424-435.
[14]	Larsen KJ, Work TW (2003) Differences in ground beetles (Coleoptera: Carabidae) of original and reconstructed tallgrass prairies in northeastern Iowa, USA, and impact of 3-year spring burn cycles. Journal of Insect Conservation, 7, 153-166.
[15]	Li HX (李鸿兴), Sui JZ (隋敬之), Zhou SX (周士秀), Zhou Q (周勤), Sun HG (孙洪国) (1987) Key to Insects (昆虫分类检索). China Agriculture Press, Beijing. (in Chinese)
[16]	Li Q (李巧), Chen YQ (陈又清), Guo X (郭萧), Chen YL (陈彦林) (2006) Arthropod used as bio-indicator to assess the success of ecological restoration. Journal of Central South Forestry University (中南林学院学报), 26(3), 117-122. (in Chinese with English abstract)
[17]	Liu YH (刘云慧), Yu ZR (宇振荣), Liu Y (刘云) (2004) Temporal and spatial structure of carabid community in agricultural landscape of Dongbeiwang, Beijing. Chinese Journal of Applied Ecology (应用生态学报), 15, 85-90. (in Chinese with English abstract)
[18]	Liu YJ (刘玉金) (2007) Wild Duck Wetland (北京野鸭湖湿地). China Forestry Publishing House, Beijing. (in Chinese)
[19]	Lövei GL, Sunderland KD (1996) Ecology and behavior of ground beetles (Coleoptera: Carabidae). Annual Review of Entomology, 41, 231-256.
[20]	Luff ML, Eyre MD, Rushton SP (1989) Classification and ordination of habitats of ground beetles (Coleoptera, Carabidae) in north-east England. Journal of Biogeography, 16, 121-130.
[21]	Magagula CN (2003) Changes in carabid beetle diversity within a fragmented agricultural landscape. African Journal of Ecology, 41, 23-30.
[22]	Magura T, Tóthmérész B, Molnár T (2004) Changes in carabid beetle assemblages along an urbanisation gradient in the city of Debrecen, Hungary. Landscape Ecology, 19, 747-759.
[23]	Negro M, Casale A, Migliore L, Palestrini C, Rolando A (2007) The effect of local anthropogenic habitat heterogeneity on assemblages of carabids (Coleoptera, Caraboidea) endemic to the Alps. Biodiversity and Conservation, 16, 3919-3932.
[24]	Rainio J, Niemelä J (2003) Ground beetles (Coleoptera: Carabidae) as bioindicators. Biodiversity and Conservation, 12, 487-506.
[25]	Rieske LK, Buss LJ (2001) Influence of site on diversity and abundance of ground- and litter-dwelling coleoptera in Appalachian oak-hickory forests. Environmental Entomology, 30, 484-494.
[26]	Sanways MJ (1993) Insects in biodiversity conservation: some perspectives and directives. Biodiversity and Conservation, 2, 258-282.
[27]	Siemann E, Tilman D, Haarstad J, Ritchie M (1998) Experimental tests of the dependence of arthropod diversity on plant diversity. The American Naturalist, 152, 738-750. DOI URL PMID
[28]	Topp W, Kappes H, Kulfan J, Zach P (2006) Litter-dwelling beetles in primeval forests of Central Europe: does deadwood matter? Journal of Insect Conservation, 10, 229-239.
[29]	Tumur A (艾尼瓦尔∙吐米尔), Halik M (马合木提∙哈力克), Frank T (2005) Beetles communities structure differences between farmland and ecological compensation area. Acta Ecologica Sinica (生态学报), 25, 2284-2290. (in Chinese with English abstract)
[30]	Yaacobi G, Ziv Y, Rosenzweig ML (2007) Effects of interactive scale-dependent variables on beetle diversity patterns in a semi-arid agricultural landscape. Landscape Ecology, 22, 687-703.
[31]	Yu XD (于晓东), Luo TH (罗天宏), Yang J (杨建), Zhou HZ (周红章) (2006) Diversity of ground-dwelling beetles (Coleoptera) in larch plantation with different stages of reforestation in Wolong Natural Reserve, Southwestern China. Zoological Research (动物学研究), 27, 1-11. (in Chinese with English abstract)
[32]	Yu XD (于晓东), Luo TH (罗天宏), Zhou HZ (周红章) (2002) Composition and seasonal dynamics of litter-layer beetle community in the Dongling Mountain region, North China. Acta Entomologica Sinica (昆虫学报), 45, 785-793. (in Chinese with English abstract)
[33]	Yu XD (于晓东), Zhou HZ (周红章), Luo TH (罗天宏) (2001) Species diversity of insects on the Ordos Plateau, Northwest China. Biodiversity Science (生物多样性), 9, 329-335. (in Chinese with English abstract)
[34]	Zhang JT (张金屯) (2004) Quantitative Ecology (数量生态学). Science Press, Beijing. (in Chinese)
[35]	Zhang Q (张倩), Zheng FK (郑发科), Zhong ZY (钟志宇) (2008) Diversity of litter-layer beetles in Wuyishan Nature Reserve, Jiangxi Province. Sichuan Journal of Zoology (四川动物), 27, 3-6. (in Chinese with English abstract)
[36]	Zhao ZM (赵志模), Guo YQ (郭依泉) (1990) Principle and Methods of Community Ecology (群落生态学原理与方法). Publishing House of Scientific and Technical Documentation, Chongqing Branch, Chongqing. (in Chinese)
[37]	Zheng LY (郑乐怡), Gui H (归鸿) (1999) Insect Classification (昆虫分类). Nanjing Normal University Press, Nanjing. (in Chinese)
[38]	Zhou HZ (周红章), Yu XD (于晓东), Luo TH (罗天宏), He JJ (何君舰), Zhou HS (周海生), Ye CJ (叶婵娟) (2000) Insect abundance and environmental effects in Shennongjia Natural Reserve, Hubei Province. Chinese Biodiversity (生物多样性), 8, 262-270. (in Chinese with English abstract)

科 Family	甲虫个体数 Individual number					总个体数 Total	百分比 (%)
科 Family	球穗莎草带 (CGT)	猪毛菜带 (SCT)	退化芦苇带 (DPCT)	牛鞭草带 (HAT)	保存较好的芦苇带 (WPCT)	总个体数 Total	百分比 (%)
步甲科 Carabidae	943	282	50	134	1,132	2,541	21.50
隐翅虫科 Staphylinidae	554	533	183	546	585	2,401	20.31
蚁形甲科 Anthicidae	71	680	17	530	66	1,364	11.54
肖叶甲科 Eumolpidae	61	46	1	15	977	1,100	9.31
薪甲科 Lathridiidae	191	40	49	86	595	961	8.13
象甲科 Curculionidae	240	119	18	27	73	477	4.04
蜉金龟科 Aphodiidae	50	48	6	82	197	383	3.24
叩甲科 Elateridae	50	44	2	0	280	376	3.18
拟步甲科 Tenebrionidae	5	247	15	2	48	317	2.68
金龟科 Scarabaeidae	8	201	20	53	10	292	2.47
叶甲科 Chrysomelidae	62	37	2	4	159	264	2.23
埋葬甲科 Silphidae	67	64	20	31	18	200	1.69
隐食甲科 Cryptophagidae	28	108	7	19	28	190	1.61
丽金龟科 Rutelidae	2	100	49	3	2	156	1.32
虎甲科 Cicindelidae	14	88	0	0	33	135	1.14
阎甲科 Histeridae	4	77	12	26	5	124	1.05
粪金龟科 Geotrupidae	1	25	14	57	9	106	0.90
小蕈甲科 Mycetophagidae	46	11	1	7	14	79	0.67
鳃金龟科 Melolonthidae	12	22	1	12	8	55	0.47
瓢甲科 Coccinellidae	6	14	9	16	4	49	0.41
苔甲科 Scydmaenidae	0	0	13	0	25	38	0.32
犀金龟科 Dynastidae	11	2	3	11	11	38	0.32
蕈坚甲科 Merycidae	1	2	6	21	0	30	0.25
花萤科 Cantharidae	2	0	0	0	23	25	0.21
露尾甲科 Nitidulidae	4	2	2	4	10	22	0.19
缨甲科 Ptiliidae	18	0	0	0	3	21	0.18
平唇水龟科 Hydraenidae	12	0	0	0	0	12	0.10
蚁甲科 Pselaphidae	2	2	0	1	6	11	0.09
圆泥甲科 Georyssidae	1	0	0	0	7	8	0.07
锹甲科 Lucanidae	4	0	0	0	4	8	0.07
花蚤科 Mordellidae	0	3	0	3	1	7	0.06
长蠹科 Bostrychidae	0	3	1	1	1	6	0.05
球蕈甲科 Leiodidae	0	1	1	0	3	5	0.04
水龟甲科 Hydrophilidae	3	0	0	0	1	4	0.03
小丸甲科 Nosodendridae	0	0	1	0	3	4	0.03
铁甲科 Hispidae	2	1	0	0	0	3	0.03
水缨甲科 Hydroscaphidae	0	0	0	0	2	2	0.02
花金龟科 Cetoniidae	0	2	0	0	0	2	0.02
天牛科 Cerambycidae	0	1	0	0	0	1	0.01
皮蠹科 Dermestidae	1	0	0	0	0	1	0.01
拟花萤科 Melyridae	0	0	0	0	1	1	0.01
窃蠹科 Anobiidae	0	0	1	0	0	1	0.01
总计 Total	2,476	2,805	504	1,691	4,344	11,820

科 Family	甲虫个体数 Individual number					总个体数 Total	百分比 (%)
科 Family	球穗莎草带 (CGT)	猪毛菜带 (SCT)	退化芦苇带 (DPCT)	牛鞭草带 (HAT)	保存较好的芦苇带 (WPCT)	总个体数 Total	百分比 (%)
步甲科 Carabidae	943	282	50	134	1,132	2,541	21.50
隐翅虫科 Staphylinidae	554	533	183	546	585	2,401	20.31
蚁形甲科 Anthicidae	71	680	17	530	66	1,364	11.54
肖叶甲科 Eumolpidae	61	46	1	15	977	1,100	9.31
薪甲科 Lathridiidae	191	40	49	86	595	961	8.13
象甲科 Curculionidae	240	119	18	27	73	477	4.04
蜉金龟科 Aphodiidae	50	48	6	82	197	383	3.24
叩甲科 Elateridae	50	44	2	0	280	376	3.18
拟步甲科 Tenebrionidae	5	247	15	2	48	317	2.68
金龟科 Scarabaeidae	8	201	20	53	10	292	2.47
叶甲科 Chrysomelidae	62	37	2	4	159	264	2.23
埋葬甲科 Silphidae	67	64	20	31	18	200	1.69
隐食甲科 Cryptophagidae	28	108	7	19	28	190	1.61
丽金龟科 Rutelidae	2	100	49	3	2	156	1.32
虎甲科 Cicindelidae	14	88	0	0	33	135	1.14
阎甲科 Histeridae	4	77	12	26	5	124	1.05
粪金龟科 Geotrupidae	1	25	14	57	9	106	0.90
小蕈甲科 Mycetophagidae	46	11	1	7	14	79	0.67
鳃金龟科 Melolonthidae	12	22	1	12	8	55	0.47
瓢甲科 Coccinellidae	6	14	9	16	4	49	0.41
苔甲科 Scydmaenidae	0	0	13	0	25	38	0.32
犀金龟科 Dynastidae	11	2	3	11	11	38	0.32
蕈坚甲科 Merycidae	1	2	6	21	0	30	0.25
花萤科 Cantharidae	2	0	0	0	23	25	0.21
露尾甲科 Nitidulidae	4	2	2	4	10	22	0.19
缨甲科 Ptiliidae	18	0	0	0	3	21	0.18
平唇水龟科 Hydraenidae	12	0	0	0	0	12	0.10
蚁甲科 Pselaphidae	2	2	0	1	6	11	0.09
圆泥甲科 Georyssidae	1	0	0	0	7	8	0.07
锹甲科 Lucanidae	4	0	0	0	4	8	0.07
花蚤科 Mordellidae	0	3	0	3	1	7	0.06
长蠹科 Bostrychidae	0	3	1	1	1	6	0.05
球蕈甲科 Leiodidae	0	1	1	0	3	5	0.04
水龟甲科 Hydrophilidae	3	0	0	0	1	4	0.03
小丸甲科 Nosodendridae	0	0	1	0	3	4	0.03
铁甲科 Hispidae	2	1	0	0	0	3	0.03
水缨甲科 Hydroscaphidae	0	0	0	0	2	2	0.02
花金龟科 Cetoniidae	0	2	0	0	0	2	0.02
天牛科 Cerambycidae	0	1	0	0	0	1	0.01
皮蠹科 Dermestidae	1	0	0	0	0	1	0.01
拟花萤科 Melyridae	0	0	0	0	1	1	0.01
窃蠹科 Anobiidae	0	0	1	0	0	1	0.01
总计 Total	2,476	2,805	504	1,691	4,344	11,820

	球穗莎草带 (CGT)	猪毛菜带 (SCT)	退化芦苇带 (DPCT)	牛鞭草带 (HAT)	保存较好的芦苇带 (WPCT)
活动密度 Activity density (AD)	619.0 ± 50.2^d	350.6 ± 26.4^c	126.0 ± 8.7^a	211.4 ± 20.1^b	543.0 ± 67.1^d
科丰富度 Family richness (R)	23.5 ± 1.0^c	20.8 ± 0.9^b	19.0 ± 0.4^b	15.8 ± 0.7^a	21.1 ± 0.7^bc
丰富度指数 Margalef richness index (D)	3.51 ± 0.19^bc	3.38 ± 0.14^bc	3.73 ± 0.13^c	2.77 ± 0.11^a	3.23 ± 0.10^b
多样性指数 Shannon-Wiener diversity index (H')	2.01 ± 0.11^ab	2.35 ± 0.08^c	2.22 ± 0.02^bc	1.82 ± 0.09^a	2.02 ± 0.06^ab
均匀度指数 Pielou evenness index (E)	0.64 ± 0.03^a	0.78 ± 0.02^b	0.75 ± 0.01^b	0.66 ± 0.03^a	0.66 ± 0.02^a
优势度指数 Simpson dominance index (C)	0.22 ± 0.03^ab	0.15 ± 0.02^a	0.18 ± 0.01^ab	0.26 ± 0.03^b	0.20 ± 0.02^ab

	球穗莎草带 (CGT)	猪毛菜带 (SCT)	退化芦苇带 (DPCT)	牛鞭草带 (HAT)	保存较好的芦苇带 (WPCT)
活动密度 Activity density (AD)	619.0 ± 50.2^d	350.6 ± 26.4^c	126.0 ± 8.7^a	211.4 ± 20.1^b	543.0 ± 67.1^d
科丰富度 Family richness (R)	23.5 ± 1.0^c	20.8 ± 0.9^b	19.0 ± 0.4^b	15.8 ± 0.7^a	21.1 ± 0.7^bc
丰富度指数 Margalef richness index (D)	3.51 ± 0.19^bc	3.38 ± 0.14^bc	3.73 ± 0.13^c	2.77 ± 0.11^a	3.23 ± 0.10^b
多样性指数 Shannon-Wiener diversity index (H')	2.01 ± 0.11^ab	2.35 ± 0.08^c	2.22 ± 0.02^bc	1.82 ± 0.09^a	2.02 ± 0.06^ab
均匀度指数 Pielou evenness index (E)	0.64 ± 0.03^a	0.78 ± 0.02^b	0.75 ± 0.01^b	0.66 ± 0.03^a	0.66 ± 0.02^a
优势度指数 Simpson dominance index (C)	0.22 ± 0.03^ab	0.15 ± 0.02^a	0.18 ± 0.01^ab	0.26 ± 0.03^b	0.20 ± 0.02^ab

类群 Family	球穗莎草带 (CGT)	猪毛菜带 (SCT)	退化芦苇带 (DPCT)	牛鞭草带 (HAT)	保存较好的芦苇带 (WPCT)
步甲科 Carabidae	235.8 ± 53.1**^{, b}	35.3 ± 3.4*^{, a}	12.5 ± 4.2*^{, a}	16.8 ± 5.8*^{, a}	141.5 ± 25.6**^{, b}
隐翅虫科 Staphylinidae	138.5 ± 9.9**^{, b}	66.6 ± 2.6*^{, a}	45.8 ± 3.8**^{, a}	68.3 ± 11.8**^{, a}	73.1 ± 13.8*^{, a}
蚁形甲科 Anthicidae	17.8 ± 4.5^b	85.0 ± 24.5**^{, c}	4.3 ± 0.8^a	66.3 ± 17.1**^,c	8.3 ± 4.0^a
肖叶甲科 Eumolpidae	15.3 ± 7.2^c	5.8 ± 1.1^bc	0.3 ± 0.3^a	1.9 ± 1.1^ab	122.1 ± 41.5**^{, d}
薪甲科 Lathridiidae	47.8 ± 4.2*^{, b}	5.0 ± 1.3^a	12.3 ± 4.0*^{, a}	10.8 ± 2.7*^{, a}	74.4 ± 16.7*^{, b}
象甲科 Curculionidae	60.0 ± 8.6*^{, d}	14.9 ± 1.9^c	4.5 ± 0.9^ab	3.4 ± 1.3^a	9.1 ± 1.9^bc
叩甲科 Elateridae	12.5 ± 1.7^c	5.5 ± 1.9^bc	0.5 ± 0.3^b	0.0 ± 0.0^a	35.0 ± 4.8*^{, d}
拟步甲科 Tenebrionidae	1.3 ± 0.6^ab	30.9 ± 5.8*^{, d}	3.8 ± 0.6^c	0.3 ± 0.3^a	6.0 ± 2.1^bc
金龟科 Scarabaeidae	2.0 ± 1.1^ab	25.1 ± 2.3*^{, c}	5.0 ± 1.2^b	6.6 ± 2.3^b	1.3 ± 0.8^a
丽金龟科 Rutelidae	0.5 ± 0.3^a	12.5 ± 4.3^b	12.3 ± 3.5*^b	0.4 ± 0.3^a	0.3 ± 0.3^a