馬艷芳,　張山林,　張永強(qiáng),　常承秀,　馬　慧,　朱惠英,　陳小玲

(甘肅臨夏州森林病蟲(chóng)害防治檢疫站　731100)

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異色瓢蟲(chóng)幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜若蟲(chóng)的捕食效應(yīng)

馬艷芳*,張山林,張永強(qiáng),常承秀,馬慧,朱惠英,陳小玲

(甘肅臨夏州森林病蟲(chóng)害防治檢疫站731100)

在室內(nèi)研究了捕食性天敵異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜若蟲(chóng)的捕食作用。結(jié)果表明,異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜若蟲(chóng)的捕食功能反應(yīng)符合Holling Ⅱ型反應(yīng),擬合的圓盤(pán)方程為Na=1.028 9Nt/(1+0.009 8Nt)。根據(jù)HollingⅢ型功能反應(yīng)新模型Na=a·exp(-b/Nt)計(jì)算出最佳尋找密度43.54頭;在獵物密度一定的情況下,利用其捕食作用率與個(gè)體間相互干擾作用的關(guān)系用Hassell-Varley的干擾反應(yīng)模型擬合為E=0.242 4P-0.398 5;分?jǐn)偢?jìng)爭(zhēng)強(qiáng)度與其密度的關(guān)系式為I=0.005 4 lgP+0.679 6。利用Holling (1959)尋找效應(yīng)(E)與獵物密度(Nt)關(guān)系式計(jì)算了尋找效應(yīng);利用Hassell & Varley(1969)模型和Beddington(1975) 2種模型對(duì)異色瓢蟲(chóng)4齡幼蟲(chóng)尋找效應(yīng)和自身密度之間的關(guān)系進(jìn)行了模擬,模擬方程分別為:E=0.748 7P-1.298 4、E=1.021 3/(2.479 4P-1.479 4)和E=1.021 3/(0.01Na+2.479 4P-1.479 4),異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜的尋找效應(yīng)隨獵物密度及捕食者密度的增加而下降。

異色瓢蟲(chóng);云南云杉長(zhǎng)足大蚜;捕食作用

(Forest Pest and Diseases Management & Quarantine Station of Linxia Hui Autonomous District, Gansu731100, China)

云南云杉長(zhǎng)足大蚜(CinaraalbaZhang, 1982)屬同翅目、大蚜科、長(zhǎng)足大蚜屬。主要為害云杉的一年生主側(cè)梢, 成、若蚜群集主側(cè)嫩梢,吸液為害,單枝蟲(chóng)口常在百頭以上,嫩梢覆蓋膠狀分泌物,嚴(yán)重影響光合作用,使樹(shù)勢(shì)衰弱。主側(cè)梢被害后,輕則變曲, 重則枯死,幾年后,被害植株形成叢狀主梢, 生長(zhǎng)下降, 材質(zhì)降低[1]。云南云杉長(zhǎng)足大蚜主要分布在內(nèi)蒙古(呼和浩特)、云南(麗江)、四川(丹巴、臥龍)、西藏(墨脫、松林口、樟木)、甘肅(古浪縣、岷縣)、青海(互助)等地。筆者在觀察云南云杉長(zhǎng)足大蚜生物學(xué)特性時(shí)發(fā)現(xiàn)異色瓢蟲(chóng)是其主要的捕食性天敵,國(guó)內(nèi)對(duì)云南云杉長(zhǎng)足大蚜的報(bào)道特別少,且尚未見(jiàn)天敵捕食作用的報(bào)道,為了更好地利用異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜進(jìn)行生物防治,我們對(duì)異色瓢蟲(chóng)4齡幼蟲(chóng)捕食云南云杉長(zhǎng)足大蚜若蟲(chóng)的作用進(jìn)行了研究。

1　材料與方法

1.1材料來(lái)源

2014年5月中旬在臨夏縣韓集鎮(zhèn)云杉行道樹(shù)上采集異色瓢蟲(chóng)3齡幼蟲(chóng),以云南云杉長(zhǎng)足大蚜飼養(yǎng),取蛻皮后的4齡幼蟲(chóng)供試,試驗(yàn)前統(tǒng)一取食1 d再饑餓1 d。云南云杉長(zhǎng)足大蚜若蚜采自同一地點(diǎn)云杉行道樹(shù),取齡期基本一致的3齡若蚜供試。試驗(yàn)于室溫下(23±0.5)℃進(jìn)行。

1.2試驗(yàn)設(shè)計(jì)

1.2.1功能反應(yīng)

在培養(yǎng)皿(直徑12 cm)內(nèi)測(cè)定異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)獵物云南云杉長(zhǎng)足大蚜若蟲(chóng)的功能反應(yīng)。獵物密度分別設(shè)每皿50、100、150、200、250頭(獵物密度的設(shè)置依預(yù)試驗(yàn)確定)。先將云杉針葉放入培養(yǎng)皿內(nèi),再分別將1頭異色瓢蟲(chóng)4齡幼蟲(chóng)與不同獵物密度組合放入皿中加蓋。每組合重復(fù)5次。24 h后檢查捕食量。

1.2.2干擾反應(yīng)

在每個(gè)培養(yǎng)皿中放入300頭云南云杉長(zhǎng)足大蚜3齡若蚜,用云杉針葉喂養(yǎng),再相應(yīng)放入1、2、3、4、5頭異色瓢蟲(chóng)4齡幼蟲(chóng),每處理重復(fù)5次。24 h后檢查統(tǒng)計(jì)捕食量。

1.2.3尋找效應(yīng)

在每個(gè)培養(yǎng)皿中放入50、100、150、200、250頭云南云杉長(zhǎng)足大蚜3齡若蚜,用云杉針葉喂養(yǎng),再相應(yīng)放入1、2、3、4、5頭異色瓢蟲(chóng)4齡幼蟲(chóng),每處理重復(fù)5次。24 h后檢查統(tǒng)計(jì)捕食量。

2　結(jié)果與分析

2.1異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜的捕食功能反應(yīng)

圖1　異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜3齡若蚜的 捕食功能反應(yīng)及搜尋效應(yīng)Fig.1　Functional response and searching efficiency of Harmonia axyridis 4th instar larvae to Cinara alba 3rd instar nymphs

2.2異色瓢蟲(chóng)4齡幼蟲(chóng)的最佳尋找密度

表1　不同獵物密度下異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南 云杉長(zhǎng)足大蚜3齡若蚜的捕食量Table 1　Predation of Cinara alba 3rd instar nymphs byHarmonia axyridis 4th instar larvae at different densities

當(dāng)捕食者密度P=1頭時(shí),在1 d內(nèi),1頭異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜3齡若蚜達(dá)到最大捕食量為82.95頭,最佳尋找密度為43.54頭,因此,在利用異色瓢蟲(chóng)4齡幼蟲(chóng)防控云南云杉長(zhǎng)足大蚜3齡若蚜?xí)r,益害比可以1∶43作為參考值[5]。

2.3異色瓢蟲(chóng)4齡幼蟲(chóng)捕食過(guò)程中的干擾反應(yīng)

表2　異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜3齡若蚜捕食作用率(E)和分?jǐn)偢?jìng)爭(zhēng)強(qiáng)度(I)Table 2　Predation ratio (E) and intensity of scrambling competition (I) of Harmoniaaxyridis 4th instar larvae on Cinara alba 3rd instar nymphs

2.4異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜的尋找效應(yīng)

Holling[2]認(rèn)為尋找效應(yīng)必須依賴于獵物(寄主)密度且受到天敵的食欲(或產(chǎn)卵能力)限制,他提出E=a/(1+athNt),Hassell & Varley[7]考慮到天敵數(shù)量增加會(huì)出現(xiàn)相互干擾而使其搜索效率降低,提出了干擾競(jìng)爭(zhēng)模型E=QP-m,Beddington[8]考慮到尋找效應(yīng)與獵物密度(寄主)的關(guān)系,提出E=a/[1 +btw(P-1)];綜合考慮獵物(寄主)密度、天敵密度與尋找效應(yīng)的關(guān)系,提出E=a/[1+athNa+btw(P-1)]。本文用這4種方程估計(jì)尋找效應(yīng),結(jié)果見(jiàn)表3。

表3　異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜3齡若蚜的尋找效應(yīng)Table 3　The searching efficiency of Harmonia axyridis 4th instar larvae on Cinara alba 3rd instar nymphs

將求得的各參數(shù)代入各尋找效應(yīng)方程得:Holling 尋找效應(yīng)與寄主密度的模型E=1.028 9/(1+0.010 1Nt); Hassell & Varley 干擾競(jìng)爭(zhēng)模型E′=0.748 7P-1.298 4;Beddington提出的尋找效應(yīng)與獵物密度(寄主)的模型E=1.021 3/(2.479 4P-1.479 4),獵物(寄主)密度、天敵密度與尋找效應(yīng)的模型E=1.021 3/(0.01Na+2.479 4P-1.479 4)。由圖3可看出Holling 尋找效應(yīng)與寄主密度的模型曲線E=a/(1+athNt)與E=Na/(NtP)較遠(yuǎn),Hassell & Varley 干擾競(jìng)爭(zhēng)模型E=QP-m和Beddington提出的E= a/[1+athNa+ btw(P-1)]、E=a/[1+ btw(P-1)]模型曲線與E=Na/(NtP)接近,但E=QP-m曲線更接近尋找效應(yīng)實(shí)測(cè)值,說(shuō)明E=QP-m模型能更好地描述異色瓢蟲(chóng)4齡幼蟲(chóng)尋找效應(yīng)與自身密度之間的關(guān)系。但無(wú)論用哪種模型估計(jì),結(jié)果表明在一定的空間或生境內(nèi),異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜的尋找效應(yīng)隨獵物密度及捕食者密度的增加而下降,且當(dāng)異色瓢蟲(chóng)密度大于2頭/皿后,尋找效應(yīng)的下降將趨于平緩(表3、圖2)。

圖2　異色瓢蟲(chóng)4齡幼蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜 3齡若蚜尋找效應(yīng)比較Fig.2　Comparison of the searching efficiency curves of Harmonia axyridis 4th instar larvae on Cinara alba 3rd instar nymphs

3　討論

云南云杉長(zhǎng)足大蚜在甘肅臨夏地區(qū)只為害云杉,未見(jiàn)在其他作物上為害。有翅蚜有較強(qiáng)的擴(kuò)散飛行能力,且隨著造林綠化需求,云杉苗木調(diào)運(yùn)頻繁,云南云杉長(zhǎng)足大蚜在臨夏云杉種植區(qū)都有分布,云杉人工林均受到不同程度的為害,受害嚴(yán)重的幼林,單株蟲(chóng)口密度平均在800頭以上, 平均有蟲(chóng)株率達(dá)60%,生長(zhǎng)勢(shì)衰弱,高生長(zhǎng)受到嚴(yán)重影響。云南云杉長(zhǎng)足大蚜天敵主要有異色瓢蟲(chóng)、月斑鼓額食蚜蠅[Scaevaselenitica(Meigen)]和三突花蛛[Misumenopostricuspidata],異色瓢蟲(chóng)數(shù)量居多,是云南云杉長(zhǎng)足大蚜的優(yōu)勢(shì)天敵。異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜功能反應(yīng)表明,該瓢蟲(chóng)具有很強(qiáng)的捕食能力,1頭異色瓢蟲(chóng)4齡幼蟲(chóng)一日最大捕食量達(dá)82.95頭,說(shuō)明異色瓢蟲(chóng)在云南云杉長(zhǎng)足大蚜的生物防治中具有很大的利用潛力[9]。在云南云杉長(zhǎng)足大蚜若蟲(chóng)密度相同條件下,隨著自身密度的增大,異色瓢蟲(chóng)幼蟲(chóng)的日均捕食量逐漸減少,捕食作用率逐漸降低,個(gè)體間存在分?jǐn)偢?jìng)爭(zhēng),故在田間利用異色瓢蟲(chóng)控制云南云杉長(zhǎng)足大蚜?xí)r應(yīng)以益害比1∶43作為參考值。

本試驗(yàn)用Holling關(guān)于尋找效應(yīng)的理論、Hassell & Varley(1969)和Beddington(1975) 兩種模型估計(jì)了異色瓢蟲(chóng)幼蟲(chóng)不同密度下的尋找效應(yīng),從獵物密度和捕食者密度兩方面探討了對(duì)尋找效應(yīng)的影響。當(dāng)獵物密度增大時(shí),捕食者總的處理時(shí)間會(huì)加長(zhǎng),導(dǎo)致尋找效應(yīng)不會(huì)太高,而當(dāng)捕食者密度增大時(shí),則由于相互干擾而限制了尋找效應(yīng)增加。異色瓢蟲(chóng)4齡幼蟲(chóng)在一定的空間和獵物密度相同的情況下,得出搜索常數(shù)Q=0.242 4,干擾系數(shù)m=0.398 5,干擾方程為:E=0.242 4P-0.398 5;在一定的空間下異色瓢蟲(chóng)4齡幼蟲(chóng)和獵物密度同比增加的情況下,得出搜索常數(shù)Q=0.748 7,干擾系數(shù)m=1.298 4,干擾方程為:E′=0.748 7P-1.298 4。說(shuō)明獵物密度和捕食者都增加的情況下干擾強(qiáng)度增強(qiáng),尋找效應(yīng)降低。自然界中捕食者對(duì)獵物的捕食效應(yīng)往往受到多方面的影響,如捕食者不同齡期、性別[10]及自身密度[11],獵物密度及不同齡期個(gè)體[12-13]、溫度[14]、寄主植物類型[15]及大小[16]、空間異質(zhì)性[17]及其他外界條件[18-19],實(shí)際捕食作用必然下降,如何客觀、定量地描述自然界異色瓢蟲(chóng)對(duì)云南云杉長(zhǎng)足大蚜的控制作用,有待于進(jìn)一步研究。

[1]蔡茂全.云南云杉大蚜觀察初報(bào)[J].森林病蟲(chóng)通訊,1990(4):10-11.

[2]Holling C S.Some characteristics of simple types of predation and parasitism [J].Canadian Entomologist,1959,91(7):385-398.

[3]劉愛(ài)萍,王俊清,徐林波,等.食蟲(chóng)齒爪盲蝽對(duì)枸杞木虱的捕食作用研究[J].植物保護(hù),2008,34(4):85-89.

[4]汪世澤,夏楚貴.Holling-Ⅲ型功能反應(yīng)新模型[J].生態(tài)學(xué)雜志,1988,7(1):1-3.

[5]許俊杰,劉鳳蓮,李秀芬,等. 中華嚙粉蛉對(duì)柏小爪螨捕食作用的研究[J].山東農(nóng)業(yè)大學(xué)學(xué)報(bào),2007,38(2):207-212.

[6]張安盛,于毅,門(mén)興元,等. 東亞小花蝽若蟲(chóng)對(duì)西花薊馬若蟲(chóng)的捕食作用[J].植物保護(hù)學(xué)報(bào),2008,35(1):7-11.

[7]Hassell M P, Rogers D J. Insect parasite responses in the development of population model [J]. Journal of Animal Ecology,1972,41:661-667.

[8]Beddington J R. Mutual interference between parasites or predators and its effect on searching efficiency [J]. Journal of Animal Ecology, 1975,44(1):331-340.

[9]陳步波,林光國(guó). 紅基盤(pán)瓢蟲(chóng)對(duì)橄欖星室木虱捕食作用及其模擬模型的研究[J].華東昆蟲(chóng)學(xué)報(bào),1995,4(2):79-83.

[10]金劍雪,李鳳良,程英,等. 七星瓢蟲(chóng)對(duì)豆蚜的功能反應(yīng)[J].植物保護(hù),2011,37(4):68-71.

[11]封紅兵,徐靜,張青文,等. 新疆棉區(qū)主要捕食性天敵對(duì)棉鈴蟲(chóng)捕食功能反應(yīng)的研究[J].應(yīng)用生態(tài)學(xué)報(bào),2003,14(6):1026-1028.

[12]Fantinou A A, Perdikis D C, Labropoulos P D. Preference and consumption ofMacrolophuspygmaeuspreying on mixed instar assemblages ofMyzuspersicae[J]. Biological Control, 2009, 51 (1):76-80.

[13]蔣杰賢,梁廣文. 叉角厲蝽對(duì)斜紋夜蛾不同齡期幼蟲(chóng)的選擇捕食作用[J].生態(tài)學(xué)報(bào),2001,21(4):684-687.

[14]Pakyari H,Fathipour Y, Rezapanah M. Temperature-dependent functional response ofScolothripslongicornis(Thysanoptera:Thripidae) preying onTetranychusurticae[J]. Journal of Asia-pacific Entomology, 2009, 12(1):23-26.

[15]Clercq P D, Mohaghegh J, Tirry L. Effect of host plant on the functional response of the predatorPodisusnigrispinus(Heteroptera: Pentatomidae)[J]. Biological Control, 2000,18(1):65-70.

[16]Rubia E G, Almazan L P, Heong K L. Predation of yellow stem borer (YSB) by wolf spider[J]. International Rice Research Newsletter, 1990,15(4):22.

[17]李桂亭,鄒運(yùn)鼎,周夏芝.干擾作用及空間異質(zhì)性對(duì)大草蛉雄成蟲(chóng)捕食作用的影響[J].應(yīng)用生態(tài)學(xué)報(bào),2002,13(4):433-434.

[18]楊國(guó)慶,吳進(jìn)才,王洪全. 水對(duì)稻田蜘蛛捕食功能的影響[J].生態(tài)學(xué)報(bào),2003,4(4):681-687.

[19]呂仲賢,俞曉平,Heong K L,等. 稻田氮肥施用量對(duì)黑肩綠盲蝽捕食功能的影響[J].昆蟲(chóng)學(xué)報(bào),2005,48(1):48-56.

(責(zé)任編輯：田喆)

Predatory efficiencies of the multicolored Asian lady beetleHarmonia

axyridislarvae on the aphidCinaraalbanymphs

Ma Yanfang,Zhang Shanlin,Zhang Yongqiang,Chang Chengxiu,Ma Hui,Zhu Huiying,Chen Xiaoling

In the laboratory, the predation of the multicolored Asian lady beetleHarmoniaaxyridislarvae to the aphidCinaraalbanymphs was studied. The results showed that the functional response ofH.axyridislarvae toC.albanymphs fitted to Holling type Ⅱmodel and the equation wasNa=1.028 9Nt/(1+0.009 8Nt). The best seeking density was about 43.54. The relationship between searching efficiency andH.axyridislarvae density was also studied by Holling’s modelE=a/(1+athNt). The relationship between predation efficiency and individual interference could be fitted to the Hassell-Varley modelE=0.242 4P-0.398 5．The relationship between intensity of scrambling competition (I) and the density of theH.axyridislarvae (P) wasI=0.005 4 lgP+0.679 6. The relationship between searching efficiency andH.axyridislarval density was also studied by Holling model．The relationship between searching efficiency (E) andH.axyridislarval density (P) could be described by Hassell & Varley model and Beddington (1975) two models asE=0.748 7P-1.298 4,E=1.021 3/(2.479 4P-1.479 4) andE=1.021 3/(0.01Na+2.479 4P-1.479 4)．The searching efficiency declined when the densities ofH.axyridislarvae and the prey increased．

Harmoniaaxyridis;Cinaraalba;predation

2015-05-28

2015-08-09

甘肅省臨夏州科技計(jì)劃項(xiàng)目(2011-N-S-10)

E-mail:lxgymyf@126.com

Q 968

A

10.3969/j.issn.0529-1542.2016.02.018

致謝：承中國(guó)科學(xué)院動(dòng)物研究所姜立云副研究員鑒定云南云杉長(zhǎng)足大蚜標(biāo)本，謹(jǐn)此致謝。