282 March - April 2007

ACAROLOGY Life History of Amblyseius herbicolus (Chant) (: Phytoseiidae) on Coffee Plants

PAULO R. REIS1,ADENIR V. T EODORO2, MARÇAL PEDRO NETO2 AND ESTER A. DA SILVA3

1EPAMIG-CTSM/EcoCentro, C. postal 176, 37200-000, Lavras, MG, CNPq Researcher SDXORUHEHOOHV#HSDPLJXÀDEU 2UFLA, C. postal 3037, 37200-000 Lavras, MG. Fapemig and CBP&D/Café Research Assistantship Recipients 3 Univ. Estadual do Maranhão, 65055-150, São Luís, MA

Neotropical Entomology 36(2):282-287 (2007)

História de Vida de Amblyseius herbicolus (Chant) (Acari: Phytoseiidae) em Cafeeiro

RESUMO - O ácaro predador Amblyseius herbicolus &KDQW pRVHJXQGR¿WRVHtGHRPDLVDEXQGDQWH em cafeeiro (Coffea arabica L.), após Euseius alatus DeLeon, na região de Lavras, MG, associado DRYHWRUGRYtUXVGDPDQFKDDQXODU phoenicis (Geijskes) (Acari: ). Sua KLVWyULDGHYLGDIRLHVWXGDGDOHYDQGRHPFRQWDDVSHFWRVELROyJLFRVWDEHODGHYLGDDWLYLGDGHSUHGDWyULD e respostas funcional e numérica em função da densidade de presa. Foi constatada longevidade de 38 dias para fêmeas adultas quando alimentadas com B. phoenicis. A estimativa da capacidade inata de

crescimento da população (rm) foi 0,150 e a duração média de uma geração (T) 25,3 dias. A população GREURXDFDGDGLDV7ULQWDiFDURVB. phoenicis /arena de folha de cafeeiro (3 cm de diâmetro) foram oferecidos separadamente para um espécime de cada fase do ácaro predador. Fêmeas adultas foram PDLVH¿FLHQWHVQDSUHGDomRGHWRGDVDVIDVHVGRGHVHQYROYLPHQWRGRiFDURSUHVDVHJXLGDVGDVIDVHV QLQIDLV3DUDRHVWXGRGDVUHVSRVWDVIXQFLRQDOHQXPpULFDDSUHVDIRLRIHUHFLGDQDVGHQVLGDGHVGH DIRUPDVLPDWXUDVGHB. phoenicis /cm2 de arena, por serem preferidas para predação. A predação e oviposição de A. herbicolus aumentam com o aumento da densidade de presa, com uma correlação SRVLWLYDHDOWDPHQWHVLJQL¿FDWLYD$DQiOLVHGHUHJUHVVmRUHDOL]DGDVXJHUHXPDUHVSRVWDIXQFLRQDOGR tipo II, com uma predação máxima diária de aproximadamente 35 B. phoenicis /cm2 /fêmea adulta.

3$/$95$6&+$9(%LRORJLD¿WRVHtGHRUHVSRVWDIXQFLRQDOUHVSRVWDQXPpULFDCoffea arabica

ABSTRACT - The predaceous Amblyseius herbicolus &KDQW LVWKHVHFRQGPRVWDEXQGDQW phytoseiid on coffee plants (Coffea arabica L), after Euseius alatus'H/HRQLQ/DYUDV0*%UD]LO associated to the vector of the coffee ring spot virus, Brevipalpus phoenicis (Geijskes) (Acari: 7HQXLSDOSLGDH ,WVOLIHKLVWRU\ZDVVWXGLHGWDNLQJLQWRDFFRXQWELRORJLFDODVSHFWVOLIHWDEOHSUHGDWRU\ activity and functional and numerical responses in relation to the density of the prey. The adult female

has longevity of 38 days when supplied with B. phoenicis. The intrinsic rate of population increase (rm) ZDVDQGWKHPHDQJHQHUDWLRQWLPH 7 GD\V7KHSRSXODWLRQGRXEOHVHYHU\GD\V7KLUW\ B. phoenicis /3-cm diameter coffee leaf arenas were separately offered to one specimen of each SUHGDWRUSKDVH$GXOWIHPDOHVZHUHPRUHHI¿FLHQWLQNLOOLQJDOOGHYHORSPHQWDOSKDVHVRIB. phoenicis, IROORZHGE\WKHQ\PSKVWDJHV)RUWKHIXQFWLRQDODQGQXPHULFDOUHVSRQVHVVWXGLHVIURPWR immature specimens of the prey /cm2RIDUHQDZHUHVXEPLWWHGWRWKHSUHGDWRUWKHSUHIHUUHGSKDVHIRU predation. Predation and the oviposition of A. herbicolus increased with increasing prey density, with a SRVLWLYHDQGKLJKO\VLJQL¿FDQWFRUUHODWLRQ5HJUHVVLRQDQDO\VLVVXJJHVWVDIXQFWLRQDOW\SH,,UHVSRQVH with a maximum daily predation near 35 B. phoenicis /cm2 /one adult female.

KEY WORDS: Biology, phytoseiid, functional response, numerical response, Coffea arabica

Phytoseiid mites are the most important and one of the %UD]LOLDQFRIIHHFURSV Coffea arabica L.) associated to most studied natural enemies of pest mites (McMurtry et al. the pest mites Brevipalpus phoenicis (Geijskes) (Acari: 0RUDHV0F0XUWU\ &URIW IDFWRUWREH Tenuipalpidae) and Oligonychus ilicis (McGregor) (Acari: considered in integrated pest management. The predatory Tetranychidae) (Pallini et al. 1992, Reis 2002). mite Amblyseius herbicolus (Chant) (Acari: Phytoseiidae) Moraes et al. (1991) reported 55 (A. herbicolus LVRQHRIWKHPRUHIUHTXHQWDQGDEXQGDQWSK\WRVHLLGLQ among them) of phytoseiids in Latin America, and Lin et March - April 2007 Neotropical Entomology 36(2) 283

al. (2000) cited its presence in China. This species was phoenicisWKURXJKVWXGLHVRILWVELRORJ\IHUWLOLW\OLIHWDEOH DOVRIRXQGLQ%UD]LODVVRFLDWHGWR$UHFDFHDH 6DQWDQD  predatory capacity and study of functional and numerical )OHFKWPDQQ*RQGLP 0RUDHV (XSKRUELDFHDH responses. (Zacarias & Moraes 2001), and native plants of the State of Rio Grande do Sul (Ferla & Moraes 2002). The predaceous A. herbicolus was cited previously as Material and Methods Typhlodromus (Amblyseius) herbicolus Chant, Amblyseius deleoni Muma & Denmark and Amblyseius impactus $OOELRDVVD\VDQGUHDULQJRIPLWHVZHUHFDUULHGRXW Chaudhri, according to Gondim & Moraes (2001), Zacarias XQGHU“ž&“5+DQGKSKRWRSKDVHLQ & Moraes (2001) and Moraes et al   WKH/DERUDWyULRGH$FDURORJLDRIWKH&HQWURGH3HVTXLVD A question raised is if phytoseiids can reduce high em Manejo Ecológico de Pragas e Doenças de Plantas population densities of phytophagous mites, and the - EcoCentro, of the Empresa de Pesquisa Agropecuária de studies that answer such question are those of functional Minas Gerais - EPAMIG, located at the Universidade Federal QXPEHURISUH\FRQVXPHGSHUWLPHXQLW DQGQXPHULFDO de Lavras - UFLA campus. SURJHQ\SURGXFHGE\WLPHXQLWRURWKHUFKDQJHLQSUHGDWRU GHQVLW\ UHVSRQVHV 6RORPRQ0RUL &KDQW Stock rearing. All mites, predator and prey, used in the /DLQJ 2VERUQ 7KHODVWWZRDXWKRUVIRXQGWKDW IXQFWLRQDODQGQXPHULFDOUHVSRQVHVELRDVVD\VZHUHFROOHFWHG WKHWZRUHVSRQVHVDUHFRUUHODWHGDQGVKRXOGEHFRQVLGHUHG LQWKH¿HOGLQDFURSRIWKH&DWXDtFXOWLYDURIFRIIHH C. together. arabica), which had received no phytossanitary treatments +ROOLQJ  UHSRUWHGWKDWWKUHHEDVLFW\SHV IRUWKHSDVWWKUHH\HDUVDQGNHSWLQWKHODERUDWRU\DV RIIXQFWLRQDOUHVSRQVHWRSUH\GHQVLW\FRXOGEHLGHQWL¿HG maintenance rearing. in general: linear, convex and sigmoid. In type I (linear) Stock rearing of A. herbicolusWREHXVHGLQWKH WKHQXPEHURISUH\FRQVXPHGLQFUHDVHVOLQHDUO\XSWRD H[SHULPHQWVZDVPDGHLQDUHQDVZKLFKFRQVLVWHGRIFP PD[LPXPLQW\SH,, FRQYH[ WKHQXPEHURIFRQVXPHG FRIIHHOHDIGLVNVÀRDWLQJLQGLVWLOOHGZDWHULQFPGLDPHWHU SUH\LQFUHDVHVZLWKWKHQXPEHURISUH\RIIHUHGEXWEHJLQV petri dishes. In the middle of each arena there was a small to decrease when a maximum point is reached, that is, hole to allow the passage of a pin that was glued with the EHJLQVWRVKRZDUHGXFWLRQLQFRQVXPHGSUH\UDWHZLWKWKH KHDGRQWKHERWWRPRIWKHGLVKZLWKDGKHVLYHRIVLOLFRQDQG increase of its density, and in type III (sigmoid) predation with the point up. This allowed the arenas to move inside of result in a sigmoid form with the increase of prey density WKHSHWULGLVKÀXFWXDWLQJXSDQGGRZQDFFRUGLQJWRWKHZDWHU XSWRDPD[LPXP7KH¿UVWW\SHRIUHVSRQVHLVVXSSRVHGO\ OHYHOSUHYHQWLQJWKHPRIWRXFKLQJWKHERUGHURIWKHGLVKHV W\SLFDOLQDTXDWLFLQYHUWHEUDWHVWKHVHFRQGLQSUHGDWRUDQG Each arena contained a 20 x 20-mm microscope slide cover SDUDVLWRLGVDUWKURSRGVDQGWKHWKLUGRIYHUWHEUDWHSUHGDWRUV JODVVRYHUFRWWRQ¿EHUVZKLFKVHUYHGDVVKHOWHURYLSRVLWLRQ (Hassel 1978). VLWHDQGORFDOIRUSODFLQJSROOHQRIFDVWRUEHDQSODQW Ricinus $FFRUGLQJWR+ROOLQJ  WKHEDVLFIDFWRUVWKDWDIIHFW communis L.) (Reis & Alves 1997). Pollen and immature functional response are: the time of exposure to predator stages of B. phoenicis were supplied to the mites. and prey, search rate, identification, capture, and prey The stock rearing of B. phoenicisLQWKHODERUDWRU\ZDV consumption. Other factors like predator stimulation due to PDGHRQSDUDI¿QFRDWHGFLWUXVIUXLWV Citrus sinensis2VEHFN  prey interference or the kind of learning response (Sandness except in approximately 3-cm diameter areas. These areas & McMurtry 1970), the confusion effect and hunger are ZHUHFOHDQHGXSE\XVLQJDVPDOOEUXVKDQGLPPHGLDWHO\ VXEVLGLDU\EXWLPSRUWDQWEHFDXVHWKH\FDQWULJJHUSDUWRUDOO VXUURXQGHGE\D%LR6WLFN®EDUULHUWRSUHYHQWPLWHVIURP EDVLFIDFWRUV HVFDSLQJ &KLDYHJDWR  $VIRUWKHOLIHWDEOHGHWHUPLQDWLRQLW¶VNQRZQWKDWWKH\KHOS ERWKDVLQWKHG\QDPLFVSRSXODWLRQVSHFLHVFRPSUHKHQVLRQ Biological aspects. 'DWDUHIHUULQJWRWKHELRORJ\RIA. and in the natural enemies impact evaluation causes over herbicolusZHUHREWDLQHGLQFPGLDPHWHUDUHQDVPDGHZLWK population (Lenteren & Woets 1988, Bellows Jr et al. 1992). FRIIHHOHDIIURPWKHWKLUGDSLFDOSDLUDQGEUDQFKHVIURPWKH

WKHPLWHVWRWKHSURGXFWXVHGLQDGGLWLRQWRWKHELRORJLFDO PLWHZDVEXLOWXVLQJVXUYLYDOGDWD lx VSHFL¿FIHUWLOLW\ mx) disruption caused to the environment. DQGVH[UDWLR %LUFK'LQKet al. 1988), after female This work was carried out to evaluate the predatory deutonymphs. Each deutonymph was placed on a 3-cm in potential of A. herbicolusDVELRORJLFDOFRQWURODJHQWRIB. diameter arena of coffee leaf (Reis et al DQGREVHUYHG Reis et al. - Life History of Amblyseius herbicolus (Chant) (Acari: Phytoseiidae) on Coffee Plants

DWKLQWHUYDOVUHPRYLQJWKHHJJVODLGDQGUHJLVWHULQJWKH 7KHQXPEHURIGHDGSUH\DQGHJJVODLGE\WKHSUHGDWRU\ dead mites. Twenty-two females originated from the stock PLWHZHUHHYDOXDWHGHYHU\KZLWKUHPRYDORIGHDGSUH\ UHDULQJZHUHREVHUYHG7KLUW\LPPDWXUHVRIB. phoenicisE\ QRWFRQVXPHGDQGHJJV7KHQXPEHURISUH\ZDVEURXJKWWR GD\ERWKODUYDHDQGQ\PSKV were supplied to the predaceous the original one every day during eight days. mite.

The following parameters were calculated: Ro = ¦ mx.lx QHWUHSURGXFWLYHUDWHRUWKHQXPEHURIWLPHVWRWKHSRSXODWLRQ Results and Discussion increases in each generation); T = ¦ mx.lx.x /¦ mx.lx (mean generation time); rm (approximate) = ln Ro /T (estimate of the Biological aspects. The average duration of egg stage was intrinsic rate of population increase) and O = erm ¿QLWHUDWHRI “GD\VZLWKYLDELOLW\2WKHUGHYHORSPHQWDO LQFUHDVHZKLFKUHSUHVHQWVWKHQXPEHURLQGLYLGXDOVDGGHG stage average durations were 1.2 ± 0.31 days for the larvae, to the population /time unit /female which will originate a “GD\IRUWKHSURWRQ\PSKDQG“GD\VIRU female). the deutonymph. Development from egg to adult was lasted

The true value of rm was calculated interactively through “GD\VDQGIHPDOHORQJHYLW\“GD\V 7DEOH Lotka equation (Carey 1993): 1). The sex ratio was equal to 1 (100% of females) and no males was found in A. herbicolusFRORQLHV7KHDEVHQFHRI T A. herbicolusPDOHVZDVUHODWHGLQRWKHUSDSHUVFRQ¿UPLQJ -rm (x+1) WKHUHIRUHWKHUHSURGXFWLRQRIWKLVVSHFLHVE\WHOLWRN\ 0RUDHV ™ lxmxe = 1 x = 0 & Mesa 1988).

Fertility life table. The intrinsic rate of population increase $IWHUREWDLQWLRQRIWKHWUXHYDOXHRIU , the mean duration m of the predator (r ) was 0.150 female /female /day; the mean of one generation was recalculated, T = ln R /r . m o m generation time (T) 25.3 days; the net reproductive rate (R ) 7KHQXPEHURIGD\VQHHGHGIRUWKHSRSXODWLRQWRGXSOLFDWH o IHPDOHVIHPDOHDQGWKH¿QLWHUDWHRILQFUHDVH Ȝ  was also calculated, and according to Tanigoshi et al. (1975) 7KHSRSXODWLRQLVHVWLPDWHGWRGRXEOHHYHU\GD\V7KHVH is equal to ln 2 /r . m results demonstrate that this species is well adapted to B. phoenicis, presenting higher predation than I. zuluagai (Reis Predatory activity. The experiments were conducted in et al 8QGHUODERUDWRU\FRQGLWLRQVFRQVLGHUHGLGHDO FPGLDPHWHUDUHQDVRIFRIIHHOHDIÀXFWXDWLQJLQGLVWLOOHG and where there is appropriate nourishment, the predator can water in 15-cm diameter petri dishes (Reis et al. 2000). show high reproduction capacity. On the other hand, under Four experiments were performed, one for each phase of ¿HOGFRQGLWLRQVRQHFDQVWDWHWKDWVHYHUDOHFRORJLFDOIDFWRUV development of B. phoenicis - egg, larva, nymph (protonymph can interfere with its reproductive capacity. It is known that, as SOXVGHXWRQ\PSK DQGDGXOWZLWKDFRPSOHWHO\UDQGRPL]HG KLJKHULVWKH¿QLWHLQFUHDVHUDWHKLJKHUZLOOEHWKHGDLO\JURZWK GHVLJQDQG¿YHWUHDWPHQWVFRQWUROODUYDQ\PSKDQGA. RIWKHSRSXODWLRQDQGWKHRQHVKRZHGE\A. herbicolus Ȝ  herbicolus adult female. Thirty B. phoenicis were placed  LVVOLJKWHUKLJKHUWKDQIRUI. zuluagai Ȝ   5HLV /arena, for one predaceous mite specimen according to the et al. 1998), species also frequently founded on coffee crops SKDVHWREHWHVWHGZLWKUHSOLFDWHV$IWHUKIROORZLQJ (Pallini et al. 1992, Reis 2002). the introduction of the mites in the arenas, predated, partially predated, naturally dead, dead in water and live B. phoenicis Predatory activity. Only nymphs and adults of A. herbicolus ZHUHFRXQWHG%RWKSUHGDWRUDQGSUH\VSHFLHVZHUHREWDLQHG predate adults of B. phoenicisEXWWKH\FRQVXPHGOHVVWKDQ IURPWKHVWRFNUHDULQJZKLFKDOORZVWRREWDLQPLWHVRIWKH 30% of the 30 mites offered. Adult female was the most same age. HI¿FLHQWLQSUHGDWLQJDOOSKDVHVRIGHYHORSPHQWRIWKHSUH\ Functional and numerical responses. An adult female of A. herbicolusZDVFRQ¿QHGGXULQJHLJKWGD\VLQDFPGLDPHWHU coffee leaf arena (7.1 cm2 ÀXFWXDWLQJLQGLVWLOOHGZDWHULQD 7DEOH'XUDWLRQLQGD\V PHDQ“VWDQGDUGHUURU RIDOO 15-cm diameter and 2-cm deep petri dish. Eight arenas were developmental phases of A. herbicolus fed on B. phoenicis, placed in each dish at equal distances from each other and XQGHU“ž&“5+DQGKSKRWRIDVH NHSWLQSODFHE\DVPDOOKROHLQWKHPLGGOHWRDFFRPPRGDWH DQXSVLGHGRZQSLQZLWKWKHKHDGJOXHGLQWKHERWWRP Phases of life cycle n Mean ± SE (days) GHVFULEHGSUHYLRXVO\,PPDWXUHVRIB. phoenicis were placed Egg stage 68 1.50 ± 0.31 LQWKHIROORZLQJGHQVLWLHV Larva 67 1.16 ± 0.31 ZLWKVHYHQUHSOLFDWHV  ZLWKIRXUUHSOLFDWHV  ZLWK WKUHHUHSOLFDWHV DQGFP2 (with Protonymph 67 1.00 ± 0.26 two replicates). Larvae and nymphs were used due to the Deutonymph 64 1.26 ± 0.32 SUHIHUHQFHIRULPPDWXUHVWDJHVRIWKHSUH\IRUSUHGDWLRQE\ phytoseiids (Gravena et al5HLVet al. 2000). The same Egg - adult 64 6.88 ± 0.64 QXPEHURIB. phoenicis was used as control treatment in the Longevity (female) 64 37.9 ± 5.47 DUHQDVEXWLQDEVHQFHRIWKHSUHGDWRUIRUQDWXUDOPRUWDOLW\ REVHUYDWLRQ Q 1XPEHURIVWXGLHGPLWHV March - April 2007 Neotropical Entomology 36(2) 285

although nymphs showed also high predation (Fig. 1), similar WRWKHUHVXOWVREWDLQHGZLWKRWKHUVSHFLHVRISK\WRVHLLGV (Gravena et al5HLVet al. 2000). 100 Functional and numerical responses. Results show that 80 predation and oviposition of A. herbicolus increases according 60 to the increase in prey population density, with a positive 40 Female DQGKLJKO\VLJQL¿FDQWFRUUHODWLRQ ZLWKWKHGHWHUPLQDWLRQ Nymph 20 Larva FRHI¿FLHQWHTXDOVWR DQG UHVSHFWLYHO\ ZLWK Predation (%) 0 Control almost 100% consumption in the population density of 7.7 B. Egg Larva Nymph Adult 2 phoenicis /cm DQGHJJOD\LQJRIHJJVGD\IURPWR Phases of development of B. phoenicis SUH\FP2 7DEOH)LJVDQG $OWKRXJKWKHQXPEHU of B. phoenicis killed increase with the increase of density Fig. 1. Percent predation of B. phoenicis, on its different RIWKHPLWHVRIIHUHGWKHSHUFHQWRIDWWDFNUHGXFHV )LJ  GHYHORSPHQWVWDJHVE\ODUYDQ\PSKDQGDGXOWIHPDOHRIWKH VLPLODUWRWKHUHVXOWVVKRZHGE\5HLVet al. (2003) with other predatory mite A. herbicolus. two species of phytoseiids. Such fact may result from their satiety or interference on their predation capacity related to SUH\GHQVLW\ 0RUL &KDQW6DQGQHVV 0F0XWU\ when compared to Euseius concordis (Chant) and 1970), suggesting that A. herbicolusZLOOEHPRUHHI¿FLHQW *DOHQGURPXVÀRULGDQXV (Muma) (Acari: Phytoseiidae). under low and medium prey densities. Predator-prey models A. herbicolusZDVVKRZQWREHDQHI¿FLHQWSUHGDWRU\ LQJHQHUDOVXSSRUWWKHLGHDWKDWFRQWUROE\SUHGDWLRQLVYHU\ mite at several densities of the coffee ring spot mite, B. HI¿FLHQWSDUWLFXODUO\XQGHUORZSUH\GHQVLW\DQGWKHPLWHV phoenicis,WVLPSRUWDQFHDVDQHI¿FLHQWQDWXUDOHQHP\ZDV of the group Gamasina, important group of predatory mites, evidenced and, therefore, among other predatory mites, it are included in this class (Koehler 1999). PXVWEHSUHVHUYHGWKURXJKLQWHJUDWHGDQGDJURHFRORJLFDO The regression analysis suggests a functional type II management programs. response, with a maximum daily predation near to 35 B. phoenicis /cm2 /one female (Fig. 2) which, according to Hassel (1978), is typical in predatory . Acknowledgments 6DQGQHVV 0F0XUWU\  REVHUYHGWKDWAmblyseius largoensis (Muma) (Acari: Phytoseiidae), species To the Consórcio Brasileiro de Pesquisas e taxonomically close to A. herbicolusZDVPRUHHI¿FLHQWLQ 'HVHQYROYLPHQWRGR&DIp&%3 '&DIpIRUWKH¿QDQFLDO killing Oligonychus punicae (Hirst) (Acari: Tetranychidae) support and scholarship, and to the Fundação de Amparo à

7DEOH)XQFWLRQDODQGQXPHULFDOUHVSRQVHVRIWKHSUHGDWRU\PLWHA. herbicolus fed on the phytophagous mite B. phoenicis,XQGHU“ž&“GH5+DQGKSKRWRSKDVH Mean ± SE Range Prey density 2 Consumption Predation (%) Natural mortality (%) (mites /cm ) Eggs laid /day Preys Eggs (preys /day) 0.14 0.86 ± 0.06 0.14 ± 0.01 0-1 0-1 86.00 0.00 0.28 1.93 ± 0.10 0.27 ± 0.03 1-2 0-1 96.50 0.00 0.7 4.92 ± 0.08 0.25 ± 0.06 4-5 0-2 98.40 0.00 1.4 9.87 ± 0.50 0.23 ± 0.06 7-10 0-2 98.70 0.00 2.8 19.37 ± 0.03 0.60 ± 0.09 14-20 0-2 96.85 0.00 4.2 28.14 ± 1.20 0.75 ± 0.09 17-30 0-2 93.80 4.17 4.9 33.07 ± 1.20 1.09 ± 0.10 15-35 0-3 94.48 1.07 6.3 43.78 ± 1.15 1.22 ± 0.09 38-45 0-2 97.29 0.28 7.7 54.50 ± 2.69 1.08 ± 0.12 53-55 0-2 99.09 1.59 9.8 61.68 ± 2.50 1.43 ± 0.15 50-70 1-2 88.11 0.71 14.1 87.37 ± 2.35 1.68 ± 0.18 70-96 1-2 87.37 2.00 17.6 82.99 ± 2.65 1.56 ± 0.17 68-96 1-3 66.39 1.80 28.2 101.50 ± 3.05 1.62 ± 0.17 88-120 1-3 50.75 0.19 42.3 127.81 ± 6.01 1.68 ± 0.18 82-184 1-3 42.60 3.46 286 Reis et al. - Life History of Amblyseius herbicolus (Chant) (Acari: Phytoseiidae) on Coffee Plants

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dead / day r = 0.98** 20 232. 0 &KLDYHJDWR/*%LRORJLDGRiFDURBrevipalpus phoenicis Number of 0 5 10 15 20 25 30 35 40 45 HPFLWURV3HVTX$JURSHF%UDV 2 Number of B. phoenicis /cm /day &KLDYHJDWR/*000LVFKDQ 0$6LOYD3UHMXt]RV Fig. 2. Number of B. phoenicis preyed upon by one female HWUDQVPLVVLELOLGDGHGHVLQWRPDVGHOHSURVHSHORiFDUR of A. herbicolus according to the density offered. Brevipalpus phoenicis *HLMVNHV 6D\HG $FDUL 7HQXLSDOSLGDH HPFLWURV&LHQWt¿FD 'LQK1YDQ$-DQVVHQ 0:6DEHOLV5HSURGXFWLYH 1.8 success of Amblyseius idaeus and A. anonymus on a diet of 1.6 WZRVSRWWHGVSLGHUPLWHV([S$SSO$FDURO 1.4 )HUOD1- *-GH0RUDHVÈFDURVSUHGDGRUHV $FDUL HP laid / day 1.2 1 SODQWDVQDWLYDVHFXOWLYDGDVGR(VWDGRGR5LR*UDQGHGR6XO 0.8 %UDVLO5HY%UDV=RRO 0.6 y = -2E-06x4 + 0.0002x3 - 0.0108x + 0.2194x + 0.0987 *RQGLP-U0*& *-GH0RUDHV3K\WRVHLLGPLWHV 0.4 r2 =0.97**

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B. phoenicis 40 (QWRPRO Percent of y = -1.4247x + 99.741 .RHKOHU++3UHGDWRU\PLWHV *DPDVLQD0HVRVWLJPDWD  dead 20 r2 = 0.87** $JULF(FRV\VW(QYLURQ 0 0 5 10 15 20 25 30 35 40 45 /DLQJ-( -$/2VERUQ7KHHIIHFWRISUH\GHQVLW\RQWKH Number of B. phoenicis /cm2 /day functional and numerical response of three species of predatory PLWHV(QWRPRSKDJD Fig. 4. Percent of B. phoenicis preyed upon by one female of A. herbicolus according to the density offered. /HQWHUHQ-&YDQ -:RHWV%LRORJLFDODQGLQWHJUDWHGSHVW FRQWUROLQJUHHQKRXVHV$QQX5HY(QWRPRO /LQ-=4=KDQJ<;=KDQJ4</LX --L&KHFNOLVWRI Pesquisa do Estado de Minas Gerais - Fapemig and Conselho PLWHVIURPPRVREDPERRLQ)XMLDQ&KLQD6\VW$SSO$FDURO 1DFLRQDOGH'HVHQYROYLPHQWR&LHQWt¿FRH7HFQROyJLFR 6SHF3XEO CNPq for the scholarships granted. 0F0XUWU\-$ %$&URIW/LIHVW\OHVRISK\WRVHLLGPLWHV DQGWKHLUUROHVLQELRORJLFDOFRQWURO$QQX5HY(QWRPRO References  0F0XUWU\-$&%+XIIDNHU 0YDQGH9ULH(FRORJ\ %HOORZV-U765*YDQ'ULHVFKH -6(ONLQWRQ/LIHWDEOH RIWHWUDQ\FKLGPLWHVDQGWKHLUQDWXUDOHQHPLHV$UHYLHZ, construction and analysis in the evaluation of natural enemies. 7HWUDQ\FKLGDHHQHPLHV7KHLUELRORJLFDOFKDUDFWHUVDQGWKH $QQX5HY(QWRPRO LPSDFWRIVSUD\SUDFWLFHV+LOJDUGLD March - April 2007 Neotropical Entomology 36(2) 287

0RUDHV*-GH&RQWUROHELROyJLFRGHiFDURV¿WyIDJRV,QI Reis, P.R., E.O. Sousa, A.V. Teodoro & M. Pedro Neto. 2003. Effect $JURSHF of prey density on the functional and numerical response of two species of predaceous mites (Acari: Phytoseiidae). Neotrop. Moraes, G.J. de, J.A. McMurtry, H.A. Denmark & C.B. Campos. (QWRPRO $UHYLVHGFDWDORJRIWKHPLWHIDPLO\3K\WRVHLLGDH $XFNODQG0DJQROLD3UHVVS =RRWD[D  Reis, P.R., L.G. Chiavegato & E.B. Alves. 1998. Biologia de Iphiseiodes zuluagai Denmark & Muma (Acari: Phytoseiidae). Moraes, G.J. de & N.C. Mesa. 1988. Mites of the family Phytoseiidae An. Soc. Entomol. Bras. 27: 185-191. $FDUL LQ&RORPELDZLWKGHVFULSWLRQVRIWKUHHQHZVSHFLHV ,QW-$FDURO Sandness, J.N. & J.A. McMurtry. 1970. Functional response of three species of Phytoseiidae (Acarina) to prey density. Can. Moraes, G.J. de, N.C. Mesa & A. Braun. 1991. Some phytoseiid (QWRPRO mites of Latin America (Acari: Phytoseiidae). Int. J. Acarol. 17: 117-139. Santana, D.L.Q & C.H.W. Flechtmann. 1998. Mite (Arthropoda, $FDUL DVVRFLDWHVRISDOPV $UHFDFHDH LQ%UD]LO,3UHVHQW 0RUL+ '$&KDQW7KHLQÀXHQFHRISUH\GHQVLW\ VWDWXVDQGQHZUHFRUGV5HY%UDV=RRO UHODWLYHKXPLGLW\DQGVWDUYDWLRQRQWKHSUHGDFLRXVEHKDYLRURI Phytoseiulus persimilis Athias-Henriot (Acarina: Phytoseiidae). 6RORPRQ0(7KHQDWXUDOFRQWURORIDQLPDOSRSXODWLRQV- &DQ-=RRO Anim. Ecol. 18: 1-35. Pallini Filho, A., G.J. de Moraes & V.H.P. Bueno. 1992. Ácaros Tanigoshi, L.K., S.C. Hoyt, R.W. Browne & J.A. Logan. 1975. associados ao cafeeiro (Coffea arabica L.) no Sul de Minas ,QÀXHQFHRIWHPSHUDWXUHRQSRSXODWLRQLQFUHDVHRIMetaseiulus *HUDLV&LrQF$JURWHFQRO occidentalis (Acarina: Phytoseiidae). Ann. Entomol. Soc. Am.  Reis, P.R. 2002. Brevipalpus phoenicis, ácaro vetor da mancha- anular em cafeeiro: Bioecologia, dano e controle, p.257-280. 7KRQJWDE7$&KDQGUDSDW\D *7%DNHU%LRORJ\DQG ,Q5GH2(QFDUQDomR3&$IRQVR-U -/GRV65X¿QR HI¿FDF\RIWKHSUHGDWRU\PLWHAmblyseius longispinosus (coords.), Palestras do I Simpósio de Pesquisa dos Cafés do (YDQV  $FDUL3K\WRVHLLGDH DVDELRORJLFDOFRQWURODJHQWRI %UDVLO%UDVtOLD(PEUDSD&DIpS Eotetranychus cendanai Rimando (Acari, Tetranychidae). J. $SSO(QWRPRO Reis, P.R., A.V. Teodoro & M. Pedro Neto. 2000. Predatory activity of phytoseiid mites on the developmental stages of coffee Zacarias, M.S. & G.J. de Moraes. 2001. Phytoseiid mites (Acari) ring spot mite (Acari: Phytoseiidae: Tenuipalpidae). An. Soc. DVVRFLDWHGZLWKUXEEHUWUHHVDQGRWKHUHXSKRUELDFHRXVSODQWV (QWRPRO%UDV LQVRXWKHDVWHUQ%UD]LO1HRWURS(QWRPRO Reis, P.R. & E.B. Alves. 1997. Criação do ácaro predador Iphiseiodes zuluagai Denmark & Muma (Acari: Phytoseiidae) HPODERUDWyULR$Q6RF(QWRPRO%UDV Received 12/IX/05. Accepted 27/VII/06.