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海南茶區茶角盲蝽形態發育與生物學特性研究

2025-04-14 00:00:00李文惠廖永林楊太慧吉訓聰姚其
熱帶作物學報 2025年4期

摘""要:茶角盲蝽(Helopeltis"theivora"Waterhouse)屬于半翅目(Hemiptera)盲蝽科(Miridae)角盲蝽屬(Helopeltis),是海南熱區茶園最重要的吸汁類害蟲之一。茶角盲蝽主要以若蟲和成蟲刺吸茶梢幼嫩組織為害,形成黑斑壞死,導致茶葉產量降低、品質下降。針對海南茶區茶角盲蝽的研究報道尚不多見,本研究描述了茶角盲蝽的外部形態,重點記錄卵孵化、若蟲蛻皮、成蟲羽化以及取食、交尾、產卵等生物學特征。結果表明:茶角盲蝽卵呈長橢圓形,頂端具兩根長短不一的呼吸突,卵初產白色,后逐漸變橙色,臨孵化為橙紅色;若蟲分為5齡,初孵若蟲橘紅色,后漸變為淡綠色至綠色;成蟲頭部較小,復眼為黑色,觸角呈絲狀,分為柄節、梗節、鞭節Ⅰ和鞭節Ⅱ,觸角長度約為體長的2倍,背板以淡黃色或黑色為主,腹部呈綠色或藍綠色。茶角盲蝽孵化過程的時長為(53.25±9.00)min,在室內條件下,卵孵化率僅為69.9%。茶角盲蝽若蟲的蛻皮和成蟲的羽化主要在葉片的背面,呈倒掛狀態,時長分別為(6.20±0.97)min和(8.40±1.33)min;其中,臨近蛻皮和羽化時,取食頻率明顯變少、活動變弱。茶角盲蝽喜歡吸取茶梢嫩葉和嫩莖的汁液,取食時間主要集中在晚上;成蟲交尾則集中在9:00—10:00和17:00—18:00,每次交尾持續時間為(117.75±18.70)min,此外,當雌蟲拒絕雄蟲求偶要求時,一般會將腹部向身體內側卷起。茶角盲蝽孕卵期較短,交尾完成24"h后即可產卵。雌蟲產卵時,腹部拱起,頭朝下,整個身體呈彎弓狀態;卵主要產在嫩莖表皮內,少部分在葉柄和葉脈,卵期為6~8"d。本研究結果為海南茶區茶角盲蝽的形態鑒定與行為生態學相關研究提供基礎理論依據。

關鍵詞:茶角盲蝽;茶樹;形態發育;生物學;行為習性中圖分類號:S435.711""""""文獻標志碼:A

Morphological"Development"and"Biological"Characteristics"of"Helopeltis"theivora"Waterhouse"in"Hainan"Tea"Region

LI"Wenhui1,2,"LIAO"Yonglin3,"YANG"Taihui4,"JI"Xuncong1,2*,"YAO"Qi2*

1."School"of"Tropical"Agriculture"and"Forestry,"Hainan"University,"Danzhou,"Hainan"571737,"China;"2."Institute"of"Plant"Protection,"Hainan"Academy"of"Agricultural"Sciences"/"Research"Center"for"Quality,"Safety"and"Standards"of"Agricultural"Products,"Hainan"Academy"of"Agricultural"Sciences"/"Key"Laboratory"of"Plant"Disease"and"Pest"Control,"Haikou,"Hainan"571100,"China;"3."Institute"of"Plant"Protection,"Guangdong"Academy"of"Agricultural"Sciences,"Guangzhou,"Guangdong"510640,"China;"4."College"of"Tropical"Crops,"Yunnan"Agricultural"University,"Pu’er,"Yunnan"665099,"China

Abstract:"Helopeltis"theivora"Waterhouse"belongs"to"the"genus"Helopeltis"of"the"family"Miridae"of"thenbsp;order"Hemiptera,"and"is"one"of"the"most"important"sap-sucking"pests"in"the"tropical"tea"plantations"in"Hainan,"China."The"nymphs"and"adults"of"H."theivora"mainly"damage"by"piercing"and"sucking"in"the"young"tissues"of"tea"shoots,"which"causes"the"black"brown"spots"that"necrosis"on"the"leaf"and"results"in"the"reduction"of"tea"yield"and"quality."The"research"on"the"H."theivora"in"Hainan"tea"region"is"still"rare."In"this"paper,"we"first"investigated"the"external"morphological"characteristics"of"H."theivora,"then,"the"growth"and"development"characteristics"such"as"egg"hatching,"nymph"molting,"and"adult"emergence,"as"well"as"the"behavioral"characteristics"including"feeding,"mating"and"oviposition"were"recorded"and"analyzed,"respectively."The"results"showed"that"the"egg"of"H."theivora"was"in"a"sausage"shape,"with"two"respiratory"processes"on"the"egg"operculum"at"different"length."The"color"of"the"freshly"laid"egg"was"white."It"was"then"gradually"transformed"to"the"orange"and"ultimately"exhibited"the"orange-red"as"it"approached"incubation."The"nymphal"stage"was"divided"into"5"instars,"the"color"of"the"freshly"hatched"nymph"was"orange-red,"and"then"gradually"turned"to"pale"green"and"green."The"adult"H."theivora"possessed"a"small"head,"an"elongated"beak,"and"two"black"compound"eyes."The"antennae"presented"filamentous,"and"the"length"was"twice"more"than"the"body."The"color"of"adult"H."theivora"was"brown"or"yellowish"brown."The"duration"of"incubation"of"H."theivora"was"(53.25±9.00)min."The"hatchability"was"very"low"while"the"rate"of"egg-"hatching"was"only"69.9%"under"the"laboratory"conditions."The"moulting"of"nymphs"and"the"emergence"of"adults"happened"mainly"on"the"back"of"leaves,"hanging"upside"down,"and"the"duration"was"(6.20±0.97)min"and"(8.40±1.33)min,"respectively."Particularly,"as"approached"to"the"molting"and"eclosion,"the"feeding"frequency"became"significantly"less"and"the"activity"became"weaker."The"H."theivora"preferred"to"sucking"the"saps"of"the"young"leaves"and"stems"of"tea"shoots,"and"the"feeding"time"was"mainly"concentrated"in"the"evening."The"mating"behavior"of"adults"H."theivora"mainly"happened"between"9:00"and"10:00"and"between"17:00"and"18:00,"and"the"duration"of"each"mating"was"(117.75±18.70)"min."In"addition,"when"a"female"decided"to"refuse"a"male’s"request"for"courtship,"it"generally"rolled"her"abdomen"inwards."The"stage"of"the"fertilized"egg"of"H."theivora"was"short."During"the"oviposition,"the"female"arched"itsr"abdomen,"its"head"was"downward,"and"the"whole"body"was"in"a"curvedly"arched"state."The"oviposition"site"was"mainly"in"the"epidermis"of"the"tender"stems,"while"a"small"part"of"eggs"was"laid"in"the"petiole"and"leaf"vein."The"egg"stage"was"6-8"days."The"results"of"this"study"would"provide"a"theoretical"basis"for"the"further"study"on"the"morphological"identification"and"behavioral"ecology"of"H."theivora"in"Hainan"tea"region.

Keywords:"Helopeltis"theivora"Waterhouse;"tea"tree;"morphological"development;"biology;"behavioral"habit

DOI:"10.3969/j.issn.1000-2561.2025.04.020

茶角盲蝽(Helopeltis"theivora"Waterhouse)屬于半翅目(Hemiptera)盲蝽科(Miridae)角盲蝽屬(Helopeltis)[1],是海南茶區主要害蟲之一。該蟲以成蟲、若蟲刺吸茶梢幼嫩組織汁液為害,造成嫩芽黑斑壞死,導致茶葉減產,品質和風味下降。已有研究表明,茶角盲蝽可為害多種經濟作物,如黑胡椒(Piper"nigrum)、腰果(Anacardium"occidentale)、金雞納(Cinchona"spp.)、可可(Theobroma"cocoa)等[2]。但是,相比較而言,茶樹是茶角盲蝽的首選和主要寄主[3]。茶角盲蝽在我國主要分布在海南、廣東、廣西和云南等熱帶地區[4-5],在國外則主要分布于印度、孟加拉國、肯尼亞和澳大利亞等國家[6-7]。半翅目盲蝽科昆蟲的危害特點在于種群繁殖速度較快,世代重疊現象嚴重[8]。在茶園中,吸汁類害蟲是破壞力最強、危害性最大的害蟲之一[9]。前人研究顯示,茶角盲蝽的5齡若蟲取食量最大,被稱為最貪婪的取食者[10],每頭成蟲1"d可取食約150個斑點[11]。據報道,茶角盲蝽在印度至少可造成25%~50%的經濟損失[12],而在我國海南,受害嚴重的茶園幾乎絕收[13]。

目前,國內外針對茶角盲蝽的研究主要集中于形態發育特征和防治策略等方面。如賀蓓等[14]通過調查揭陽生態防治茶園顯示,茶角盲蝽在6—7月開始進入大面積爆發期,生態防治茶園比化學防治茶園物種多樣性更加豐富。朱茜[15]探討了可可園茶角盲蝽的發育歷期和生物學特性,并應用浸豆法進行了室內毒力測定,發現高效氯氰菊酯對茶角盲蝽成蟲的毒力最強。王政等[16]通過室內選擇性試驗研究茶角盲蝽對可可種質果實的取食偏好性,發現品種STS-17對該蟲具有抗性;同時利用“Y”型嗅覺儀測定了茶角盲蝽對一些揮發物的行為反應,表明揮發物成分在茶角盲蝽寄主選擇行為過程中具有導向作用。張中潤等[17]研究比較了5個不同腰果品系對茶角盲蝽的抗蟲性,發現品系HL2-21的抗性最強。潘賢麗等[18]對腰果園中茶角盲蝽的田間分布和防治策略進行了研究,結果顯示每年11月至次年1月是茶角盲蝽種群為害擴展的臨界期,也是該蟲化學防治最有利的時機。此外,在對茶角盲蝽發生與環境的關系以及田間分布型調查中發現,該蟲發生數量和危害程度與溫濕度、陰蔽度、風雨強度、地形、種植密度有密切關系,在田間以負二項分布和N-核心分布型為主[19]。

在國外的研究中,1865年,在印度阿薩姆邦首次發現茶角盲蝽為害,最初被認為是一種真菌病害[20]。目前,在印度共記錄了3種茶角盲蝽,即H."antonii"Signoret、H."bradyi"Waterhouse和H."theivora"Waterhouse[6,"21]。MADHU等[22]通過采用可可的豆莢和嫩枝飼養茶角盲蝽來比較生物學參數,發現茶角盲蝽在豆莢上的存活率和孵化率最高,此外在2組飼養中雌蟲的壽命均大于雄蟲。DAS等[23]采集了印度東部茶園茶角盲蝽的10個種群進行殺蟲劑抗藥性測定,發現所有種群對喹硫磷均無抗性。MANASA等[24]分別選取2株球孢白僵菌菌株BKN20和BKN1/14對茶角盲蝽進行殺蟲活性測定,結果表明BKN20分離物可以作為茶角盲蝽的一種替代微生物殺蟲劑。MANASA等[25]也采用白僵菌劑對番石榴茶角盲蝽進行了防治試驗,結果表明在番石榴坐果期每周噴灑白僵菌劑3~4次能有效控制茶角盲蝽。

綜上所述,國內外針對茶角盲蝽已有較為廣泛的研究基礎。然而,目前國內茶角盲蝽的研究主要集中在可可和腰果,對茶園茶角盲蝽的研究仍較為少見。因此,本研究采用錄像技術與超景深三維顯微技術相結合,觀察比較海南茶區茶角盲蝽的外部形態特征,然后分別對該蟲孵化、蛻皮、羽化等生長發育特征以及取食、交尾、產卵等行為學特征進行記錄分析,以期為海南茶角盲蝽種群鑒定及其綠色防控技術研究提供基礎理論。

1""材料與方法

1.1""材料

茶角盲蝽采自海南省海口市瓊山區大坡鎮東昌茶園(20°00′N,"110°21′E),種植品種為海南大葉種。蟲源帶回實驗室后用茶梢飼養,飼養室環境條件:溫度為(26±10)℃,相對濕度為(70±10)%,光周期為12"L∶12"D。

1.2""方法

1.2.1""茶角盲蝽外表形態觀察""(1)卵。茶角盲蝽主要將卵產于茶梢嫩莖表皮內,2根白色呼吸突伸出表皮外,肉眼清晰可見。雌蟲當天產卵后,取出帶卵茶梢單獨培育,用標簽標注產卵日期。選用產后1~2"d的卵,在體式顯微鏡(Nikon"SMZ745)下用解剖刀縱向解刨茶梢,然后用鑷子清除周圍組織后將卵取出,置于超景深三維顯微系統(VHX-7000)中觀察卵的形狀和顏色,并測量卵的長度以及2根呼吸突的長度,重復4次。

(2)若蟲和成蟲。分別隨機選取健康、有活力的1~5齡若蟲、雌雄成蟲,每個齡期和蟲態各取15頭。為方便觀察,在試驗前先將供試茶角盲蝽若蟲和成蟲分別置于透明的果蠅管中,然后置于–20"℃冰箱中使其凍暈,若蟲約需1~2"min,成蟲約需2~3"min。待蟲子凍暈后,用鑷子取出,先在體式顯微鏡下觀察不同齡期若蟲的體色、翅芽與雌雄蟲的體色、斑紋以及其他形態特征,然后利用超景深三維顯微系統(VHX-7000)分別測量不同齡期若蟲和雌雄成蟲的體長。

1.2.2""茶角盲蝽生長發育觀察""(1)孵化過程。據前人報道,茶角盲蝽卵期為5~13"d[26]。將帶卵的茶梢置于泡水的花泥上單獨培育,每天早、中、晚用體式顯微鏡(Nikon"SMZ745)各觀察1次。當發現卵逐漸突出嫩莖表皮、卵殼變黑且有液體出現時,即代表準備開始孵化。隨即將茶梢置于超景深三維顯微系統(VHX-7000)中,觀察記錄完整的孵化過程,并統計孵化時間,重復4次。此外,利用體式顯微鏡(Nikon"SMZ745)觀察記錄成功孵化的若蟲數量,直至所有若蟲孵化完畢,計算茶角盲蝽的卵孵化率。每天早、中、晚各1次,每批調查15粒卵,重復4次。

(2)蛻皮過程。室內以茶角盲蝽3齡若蟲作為調查對象。將供試茶角盲蝽置于茶梢上飼養,以花泥保水且每天及時更換新鮮茶梢。為便于捕捉到蛻皮過程,共設置2組茶梢,每組茶梢放3頭試蟲,白天不間斷觀察,晚上則采用攝像頭輔助觀察。待蛻皮發生時,采用照相機(佳能EOS6D)記錄完整的蛻皮過程,統計每次蛻皮的持續時間,重復5次。

(3)羽化過程。室內以茶角盲蝽5齡若蟲作為調查對象。將供試茶角盲蝽置于茶梢上,飼養方法同1.2.2-(2)。為防止羽化后的成蟲飛走,將帶有5齡若蟲和茶梢的花泥置于透明玻璃圓管(直徑28"cm,高30"cm)內,在圓管頂端套上紗網通風透氣。為便于捕捉到羽化過程,共設置2組茶梢,每組茶梢放3頭5齡若蟲,白天不間斷觀察,晚上采用攝像頭輔助觀察。羽化過程中采用照相機(佳能EOS6D)記錄完整羽化過程,統計羽化的持續時間,重復5次。

1.2.3""茶角盲蝽行為學觀察""(1)取食行為。室內分別將供試茶角盲蝽若蟲和成蟲置于新鮮茶梢上,利用照相機(佳能EOS6D)跟蹤記錄茶角盲蝽的取食過程。其中,取食時長以每斑點為準,即從試蟲口針刺入葉片表皮開始,至口針拔出為止。統計取食形成每個斑點的時間,若蟲與成蟲各重復5次。

(2)交尾行為。室內選擇較強活力、已性成熟但未進行過交配的雌、雄蟲各3頭,置于透明的果蠅管中自由求偶。待有交尾行為發生時,利用照相機(佳能EOS6D)觀察記錄交尾過程,統計每次交尾從開始至結束的時間,重復8次。

(3)產卵行為。將成功交尾的雌蟲單獨置于裝有新鮮茶梢的養蟲籠中飼養,待開始產卵時,使用照相機(佳能EOS6D)跟蹤記錄完整的產卵過程,統計每次產卵的時間、產卵位置及產卵粒數,重復6次。此外,調查茶角盲蝽卵期。每次雌蟲產卵結束后,取出帶卵茶梢單獨培育,并標注日期,最初每隔1"d觀察1次,臨孵化時早、晚各觀察1次,統計從卵產下至若蟲孵出的天數,重復20次。

1.3""數據處理

首先用Excel"2019軟件對試驗數據進行統計整理,然后用IBM"SPSS"Statistics"26.0軟件進行顯著性檢驗和方差分析。其中茶角盲蝽不同齡期體長采用單因素方差分析(One-way"ANOVA),以圖基法(Tukey)進行事后檢驗。

2""結果與分析

2.1""海南茶區茶角盲蝽形態特征

2.1.1""卵""長橢圓形,卵殼質地堅硬,卵頂端分布2根呼吸突,長短不一,一般長的在上面,短的在下面(圖1A)。卵的長度為(997.93±12.50)μm,2根呼吸突長的為(640.63±23.40)μm,短的為(310.00±12.50)μm。卵初產時為白色,后逐漸變為橙色,臨孵化時則變為橙紅色。卵的外表分布著白色硬殼,臨近孵化時硬殼由白色變為黑色。

2.1.2""若蟲""茶角盲蝽若蟲活動靈敏、善于爬行。若蟲一共分為5齡,其中,1齡若蟲體色為橘紅色,無盾片、無翅芽,體表分布有密密麻麻的剛毛(圖1B);2齡若蟲體色同為橘紅色,無盾片和翅芽,體表的剛毛消失(圖1C);3齡若蟲體色由橘紅色逐漸轉變為綠色,開始出現翅芽和盾片(圖1D);4齡若蟲體色為綠色,翅芽約占體長的1/3,盾片突起(圖1E);5齡若蟲體色為綠色,翅芽約占體長的1/2,盾片突起(圖1F)。茶角盲蝽若蟲體長隨著齡期的增長逐漸變長,其中,1~5齡若蟲的體長分別為(1251±97.49)、(1943.99±"139.97)、(2959.98±179.20)、(3548.03±242.43)、(4014.86±133.69)μm,存在顯著性差異。

2.1.3""成蟲""茶角盲蝽成蟲頭部較小、喙細長,復眼呈黑色,并向兩側突出。觸角呈絲狀,分為柄節、梗節、鞭節Ⅰ和鞭節Ⅱ共4節,長度約為體長的兩倍。胸部分布有三對足和一對翅,足上分布著剛毛和黑色的斑點,翅前端革質,后端膜質。在胸和腹部的中間著生小盾片,盾片底端1/3處為黑色,其余為淡黃色,頂端呈圓形凸起,類似棒槌。雌蟲體型較雄蟲大,背板顏色以淡黃色為主,產卵管位于腹溝內側,腹部顏色為綠色,性成熟時腹部尾端的1/3處為灰綠色(圖1G)。

雄蟲體型較小,背板顏色以黑色為主,腹部顏色為藍綠色(圖1H)。雌、雄蟲體長分別為(4489.65±"110.70)、(3961.79±52.09)μm,無顯著性差異。

2.2""茶角盲蝽生長發育特征

2.2.1""孵化過程""茶角盲蝽雌蟲成功產卵后,隨著天數的增加,卵逐漸向莖表皮外突出,呈透

明狀。在臨孵化時,整個卵體開始變色,初為黃色,后轉變為橘紅色,最后初孵若蟲頭部頂破卵殼,身體掙扎而出,同時觸角從腹部伸出(圖2)。茶角盲蝽完成1次孵化過程的時間為(53.25±"9.00)min。此外,在實驗室條件下茶角盲蝽卵孵化率僅為69.9%。

2.2.2""蛻皮過程""茶角盲蝽蛻皮主要集中在茶梢葉片背面。若蟲在臨蛻皮前,取食頻率明顯變少、活動變弱。蛻皮時,蟲體在葉背倒掛,胸部先裂開,新的蟲體逐漸鉆出來,同時觸角從腹部掙扎抽離(圖3)。茶角盲蝽若蟲完成1次蛻皮的時長約為(6.20±0.97)min。剛蛻完皮的若蟲,體色呈淡黃色,體表較為柔軟。

2.2.3""羽化過程""茶角盲蝽羽化也主要集中在茶梢葉片的背面(圖4)。羽化前,5齡若蟲提前找到合適的位置,然后保持靜止不動。羽化過程和蛻皮過程較為相似但時間略長,完成1次羽化的平均時長約為(8.40±1.33)min。剛羽化出來的成蟲體色呈淡黃色,復眼呈紅色。成蟲羽化后1~3"h即可開始取食。

2.3""茶角盲蝽行為學特征

2.3.1""取食行為""茶角盲蝽喜歡吸取茶梢嫩葉和嫩莖的汁液,食量大、頻率高。葉片被取食后,葉面呈現近圓形水漬透明狀斑點,隨著時間推移,邊緣開始變為褐色,最后形成黑褐色的斑點,導致葉片干枯脫落(圖5A)。茶角盲蝽成蟲和若蟲白天不間斷取食,但是取食高峰期主要集中在晚上。從低齡若蟲到高齡若蟲,再到成蟲,取食斑點由小變大,由緊密變為稀疏,顏色由淺變深。若蟲取食1個斑點的時間約為(4.00±0.64)min,成蟲約為(2.33±0.20)min,當取食結束,口針拔出后約(4.80±0.63)min即可見黑褐色的斑點。

2.3.2""交尾行為""茶角盲蝽性成熟后,雄蟲一般比較活躍,來回爬行尋找適合交尾的雌蟲。交尾開始前,雄蟲首先爬上雌蟲背部,用足抱握雌蟲的胸部,調整合適位置,隨即將交配器插入雌蟲腹部(圖5B、圖5C)。交尾過程中,雌雄蟲一般在葉面靜止不動,當受到外界干擾時,由雌蟲牽引雄蟲爬行逃離。此外,當雌蟲拒絕雄蟲求偶交配,會將腹部向身體內側卷起(圖5D)。茶角盲蝽交尾時間多集中在9:00—10:00和17:00—18:00,完成1次交尾的時間為(117.75±18.70)min。

2.3.3""產卵行為""茶角盲蝽孕卵期較短,雌蟲交尾成功后不久即可產卵,產卵位置多數集中在茶梢的嫩莖上,少部分在葉柄和葉脈。在產卵前,雌蟲首先在嫩梢上來回爬動,尋找合適的產卵位點。雌蟲產卵時,頭朝下,腹部拱起,整個身體呈彎弓狀態(圖5E),然后將產卵器刺入嫩莖的表皮內,產下卵粒。茶角盲蝽產卵方式有2種:一種是直接將產卵器刺入茶梢幼莖表皮內完成產卵;前人在樟頸曼盲蝽的研究中發現了另一種產卵方式,即雌蟲先用刺吸式口器在莖上鉆出1個小孔,隨后將產卵器插入孔內進行產卵[27]。茶角盲蝽完成1次產卵的時長為(2.04±0.22)min,每頭雌蟲每次平均產卵1~6粒不等,且每產1粒卵后,中間需要休息1~2"min才能進行下一次產卵。此外,茶角盲蝽卵期為6~8"d,平均為(6.75±0.14)d。

3""討論

過去10"a,茶角盲蝽頻繁爆發,已對我國熱帶地區茶葉生產構成重大威脅[28]。然而,關于該蟲的研究報道仍然較少,可供借鑒的資料不多。因此,本研究系統描述了海南茶園茶角盲蝽不同蟲態的外部形態特征,并且記錄分析卵孵化、若蟲蛻皮與成蟲羽化等生長發育過程,以及成蟲取食、交尾、產卵等行為學特征。半翅目盲蝽科害蟲可為害多種作物,如條赤須盲蝽為害禾本科作物[29];樟頸曼盲蝽為害樟樹[30];薇甘菊頸盲蝽為害薇甘菊[31];紅色拉盲蝽為害桉樹[32];煙盲蝽[33]為害烤煙;棉盲蝽[34]、綠盲蝽[35]和牧草盲蝽[36]為害棉花等。但是,不同種類之間,生物學習性不一,為害特征各異,只有準確掌握海南茶園茶角盲蝽的形態特征、生長發育特征以及行為學特征,才能進行準確地鑒定和識別,并進行科學防控[37]。

在形態特征方面,茶角盲蝽卵殼頂端分布2根長短不一的呼吸突,LAI等[38]將呼吸突切除之后,發現卵不能正常孵化,表明呼吸突在卵發育過程中發揮著不可或缺的作用。茶角盲蝽若蟲分為5齡,不同齡期之間可從體型的大小、體色的差異以及有無翅芽進行劃分。雌蟲和雄蟲的外部形態各異,一是體型大小,雌蟲明顯比雄蟲大;二是前胸背板顏色,雌蟲以淡黃色為主,雄蟲則以黑色為主;三是腹部顏色,雌蟲為綠色,雄蟲則是藍綠色。在生長發育特征方面,本研究室內調查茶角盲蝽卵的孵化率僅為69.9%,王紀文[13]統計茶園茶角盲蝽卵的孵化率為56.0%,朱茜[15]統計腰果園茶角盲蝽的孵化率則為71.6%,表明茶角盲蝽孵化率普遍較低。茶角盲蝽的蛻皮與羽化過程相似,位置大多集中在茶梢葉片的背面,呈倒立狀況,可能有利于觸角的抽離伸出。

在行為學特征方面,本研究通過調查發現茶角盲蝽若蟲和成蟲均喜食茶梢嫩葉,同樣,ROY等[39]報道印度阿薩姆茶區茶角盲蝽3~4齡若蟲和成蟲最喜歡取食茶梢第2片葉,而1~2齡若蟲則更喜歡在第1片葉取食。因此,茶梢嫩度相關內含物可能在該蟲取食定位過程中發揮重要作用。盲蝽類害蟲在求偶過程中,性成熟的雄蟲一般能夠通過感受雌蟲釋放的性信息素進行定位[40],如SACHIN等[41]發現茶角盲蝽雌蟲胸部散發的性信息素對雄蟲最有吸引力。在交尾過程中,雌、雄蟲一般在葉面保持不動,如受到外界干擾,則由雌蟲牽引雄蟲迅速躲至葉片背面或順著莖桿向下逃走。有趣的是,當雌蟲拒絕雄蟲發出的求偶信號時,通常會將腹部向內側卷起。一方面可能該雌蟲已經進行過交尾,交尾后雌蟲體內誘導產生了某種驅避信息素,使其無法再進行正常交尾[42-43],或者先前交尾時雄蟲向雌蟲體內傳遞了某種抑性欲素[44]或分泌性信息素拮抗劑[45]等,最終導致雌蟲拒絕交配;另一方面可能雌蟲未進行過交尾,但拒絕接受該雄蟲傳遞的求偶信號,表明茶角盲蝽雌蟲對雄蟲求偶具有選擇性識別能力。茶角盲蝽雌蟲喜歡將卵產在嫩莖表皮內,而非葉片,這與ZHANG等[45]和SANGITA等[46]的研究結果一致。但是,另一種紅色拉盲蝽的產卵習性與此不同,該蟲主要將卵粒產在植物表皮外面[32]。

除此之外,茶角盲蝽具有避光性,當外界施加強光干擾刺激時,若蟲將迅速轉到葉片的背面躲避,或者順著莖桿直接自然掉落。RAHMAH等[47]通過調查發現,茶角盲蝽更傾向于在茶樹陰涼處活動。WAGIMAN等[48]則研究證實茶園茶角盲蝽的種群密度主要以若蟲為主,傍晚密度最大,中午密度較小,表明茶角盲蝽更喜歡避光的環境。因此,在本研究明確海南茶園茶角盲蝽的外部形態特征、生長發育特征以及行為學特征的基礎上,下一步可通過深入研究茶園光照對茶角盲蝽行為活動的影響,以期開發和應用更具針對性、更為科學合理的茶角盲蝽綠色防控策略。

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