Abstract
Diachasmimorpha longicaudata(Ashmead) has been produced in the laboratory for > 160 generations on the larvae
of oriental fruit fly, Bactrocera dorsalis (Hendel), the propagation hosts raised routinely on a semi-synthetic wheat
diet formulation. Choice tests using modified stinging units were conducted in the laboratory to investigate whether
insectary rearing had altered the host seeking and oviposition behavior of female parasitoids. Results showed
that fruit fly larvae that developed in papaya, Carica papaya L. var. ‘solo’, were less preferred for oviposition
than fruit fly larvae that developed on wheat diet when both were exposed concurrently to naive D. longicaudata
females (= females without prior oviposition experience). The substrates (pureed papaya or wheat diet) in which
treatment larvae were exposed to parasitoids did not affect oviposition preference of gravid D. longicaudata for
wheat diet-reared fruit fly larvae. Our study demonstrated the possibility that rearing in an insectary system may
have modified the parasitization behavior of female D. longicaudata.
Introduction
Diachasmimorpha longicaudata (Ashmead) [= Opius
longicaudatus Ashmead], a braconid parasitoid
of tephritid fruit flies, was introduced into Hawaii
between 1947 and 1952 subsequent to the discovery
and establishment of the oriental fruit fly, Bactrocera
(= Dacus) dorsalis (Hendel) (Zwaluwenberg, 1947;
Clausen et al., 1965). This initial effort was undertaken
primarily to deter spoilage of fruits and vegetables from
fruit fly infestation and reduce the risks of fruit fly
introductions to the continental United States through
shipment of Hawaiian produce. Once very abundant
in 1948, D. longicaudata accounted for only 30% of
total fruit fly parasitization in 1979 and 1985 (Wong &
Ramadan, 1987).
Diachasmimorpha longicaudata is an obligate
endoparasitoid of B. dorsalis larvae. A female lays her
eggs inside fruit fly larvae where they complete development
(Greany et al., 1976). Adult parasitoids emerge
from fruit fly pupae usually a few days after emergence
of adult flies from unparasitized pupae. When hosts
are scarce, it is not uncommon for a female to oviposit
more than 1 egg in a host larva (Lawrence et al., 1978).
Nonetheless, D. longicaudata being solitary, only one
individual from each pupa will complete development
to adult.
From initial field collections of fruit fly pupae in the
islands of Oahu, Maui, and Hawaii, a laboratory colony
of D. longicaudata was started in 1981. Subsequently,
a mass-rearing procedure was developed using larvae
of B. dorsalis as propagation hosts (Wong & Ramadan,
1992). Currently, D. longicaudata is one of six species
of fruit fly parasitoids reared in this laboratory which
produces more than a million wasps each week (Wong
& Ramadan, 1992).
D. longicaudata is commonly used in laboratory
and field assays because it can be produced easily in
large numbers (Wong & Ramadan, 1992). Notwithstanding,
efforts to suppress resident populations of
B. dorsalis in guavas through augmentative releases
were unsuccessful on the island of Kauai, Hawaii
GR: 201003406, Pips nr. 133525 BIO2KAP *133525* ento1508.tex; 13/05/1997; 12:55; v.7; p.1
214
(M.F. Purcell, J.C. Herr, R.H. Messing & T.T.Y. Wong,
unpubl.). Despite releases of 6–800 000 parasitoids
during the first year of the test, parasitization of B. dorsalis
in common guava, Psidium guajava L. hardly
exceeded 6%. However, it is not known whether conditions
during shipment of parasitized B. dorsalis pupae
(Purcell et al., 1994) or errant behavior of adult parasitoids
as they emerged and dispersed from release sites
(Messing et al., 1994) may have affected the efficacy
of D. longicaudata. Recently, in a study on interspecific
competition among 3 parasitoids of B. dorsalis,
we observed that females of D. longicaudata were
less responsive in parasitizing fruit fly larvae infesting
papayas (Bautista & Harris, unpubl.). Again, the
reason why is not understood.
We were curious as to whether insectary rearing
may have modified the parasitization behavior of
D. longicaudata. Therefore, choice tests were undertaken
in the laboratory to compare the host seeking
and oviposition behavior of female parasitoids between
fruit fly larvae that developed in C. papaya and those
that developed on wheat diet.
Materials and methods
Rearing of fruit flies for assays. Approximately 1000
B. dorsalis eggs (18 to 24 h old), obtained from
insectary-colonized fruit flies (Tanaka et al., 1969;
Vargas, 1989), were inoculated in half-ripe whole
C. papaya L. var. ‘Solo’ (weight range = 430–488 g
per fruit) or seeded on 235 g wheat diet. Each of these
rearing media was placed in 21 cm-diameter plastic
containers with mesh screen lids. Five to 6 days later,
following development of fruit fly larvae to 3rd instars
(Wong & Ramadan, 1992), fruit fly larvae were teased
out from deteriorated papaya with a pair of forceps
then collected in a petri dish with tap water. In the
case of the wheat diet, fruit fly larvae were screened
from the wheat substrate with a mesh sieve (1 mm2)
then emptied into another dish with tap water. Fruit fly
larvae were taken at random from each petri dish for
subsequent assays.
Experimental procedure. A modified stinging (= oviposition)
unit developed and described by Wong &
Ramadan (1992) was used in all the assays. A pair of
stinging units, which contained fruit fly larvae, were
exposed concurrently to 25 females of D. longicaudata
(5 to 6-d-old after emergence) inside a cubical cage
(26 26 26 cm). One of the stinging units contained
300 larvae reared in papaya while the other stinging
unit contained 300 larvae reared on wheat diet. During
the tests, adult parasitoids were fed with spun honey
(Sioux Honey, Sioux City, IA) and agar (source of
water).
Fruit fly larvae were not exposed naked to parasitoids
but were mixed with wheat diet substrate before
packing them in the stinging unit (Wong & Ramadan,
1992). In our tests, besides wheat diet, we likewise
utilized pureed papaya as a substrate. Both substrates
were used merely for purposes of confirming whether
odor stimuli other than those that may be associated
with the treatment larvae would influence the choice
made by the female parasitoids. Wheat millfeed (Armstrong,
B.C. Canada), a bulking material routinely used
in B. dorsalis larval diet formulation, was mixed with
pureed papaya or wheat diet substrate in equal parts
(by weight) to make mixture drier. Otherwise, female
parasitoids would shy away from the wet surface of the
stinging unit (Wong & Ramadan, 1992).
All assays were conducted under laboratory conditions
with mean ambient low and high temperatures
of 23 0:3
C and 24 0:2
C, respectively, relative
humidity of 61:5 0:8%, and L10:D14 photoperiod.
Each test was replicated 12 times. Fresh cohorts of
naive D. longicaudata females (= no prior oviposition
experience) were used in each replication.
Test 1. Fruit fly larvae reared in papaya (Pa) or wheat
diet (Wd) were exposed to parasitoids in pureed papaya
substrate (Pus) or wheat diet substrate (Wds), respectively.
Treatments were subsequently referred to as
(Pa-Pus)1 and (Wd-Wds)1, respectively. The numerical
subscript in each treatment indicates the test number.
One half hour after initial exposure of (Pa-Pus)1
and (Wd-Wds)1, and at hourly intervals for a total
exposure time of 6 h, the number of parasitoids on
each stinging unit that exhibited typical host seeking
behavior, i.e., intense antennation on surface of stinging
unit, extension of ovipositor, and insertion of ovipositor
accompanied by pumping movements, were
recorded. The number of females observed on each
stinging unit per unit time was expressed in percentage
based on the original number of females assayed in
each cage.
At the end of the 6 h period (total exposure time),
the pair of stinging units were retrieved from the cage,
then 30 larvae were sampled at random from each stinging
unit. Fruit fly larvae were dissected individually
under a stereo microscope for presence of parasitoid
eggs. Percentage parasitization (pooled counts of paraento1508.tex;
13/05/1997; 12:55; v.7; p.2
215
sitized fruit fly larvae which contained 1 and 2 or more
parasitoid eggs) was computed by dividing the number
parasitized by total number dissected.
The remainder of fruit fly larvae from each stinging
unit was placed in separate holding cups (11 cmdiameter)
provisioned with vermiculite (W. R. Grace,
Cambridge, MA) for fruit fly pupation. After 9–10 d
fruit fly pupae were sifted with a mesh screen (1 mm2).
A cohort of 200 pupae was sampled and then placed
in emergence cups with mesh-screen covers until eclosion
of parasitoids. Pooled counts of emerged parasitoids
and dead parasitoid cadavers in uneclosed pupae
were recorded. Percent yields of parasitoid progeny in
each treatment was expressed as a ratio of total number
of parasitoids that developed per 200 pupae sampled.
Test 2. The same variables were generated according
to procedure described in preceding test except that
fruit fly larvae reared in Pa or Wd were both exposed
to parasitoids in -Wds substrate only. Treatments were
subsequently referred to as (Pa-Wds)2 and (Wd-Wds)2.
Test 3. Fruit fly larvae reared in Pa and Wd were both
exposed to parasitoids in -Pussubstrate only. As in Test
1, similar procedure was followed in the processing
of samples and data collection. Treatments were subsequently
referred to as (Pa-Pus)3 and (Wd-Pus)3.
Data analysis. Data on proportion of responding
D. longicaudata females, rate of parasitization, and
percentage parasitoid progeny yield were compared
between paired treatments by Student’s t-test at P =
0:05. Percentages were transformed to arcsine p
proportion
before analyses. Untransformed means (
SEM) are used in presentation of results.
Results
Test 1. The overall mean percentage of female parasitoids
that responded to fruit fly larvae following a
total exposure time of 6 h was 35% higher in (PaPus)
AbstractDiachasmimorpha longicaudata(Ashmead) has been produced in the laboratory for > 160 generations on the larvaeof oriental fruit fly, Bactrocera dorsalis (Hendel), the propagation hosts raised routinely on a semi-synthetic wheatdiet formulation. Choice tests using modified stinging units were conducted in the laboratory to investigate whetherinsectary rearing had altered the host seeking and oviposition behavior of female parasitoids. Results showedthat fruit fly larvae that developed in papaya, Carica papaya L. var. ‘solo’, were less preferred for ovipositionthan fruit fly larvae that developed on wheat diet when both were exposed concurrently to naive D. longicaudatafemales (= females without prior oviposition experience). The substrates (pureed papaya or wheat diet) in whichtreatment larvae were exposed to parasitoids did not affect oviposition preference of gravid D. longicaudata forwheat diet-reared fruit fly larvae. Our study demonstrated the possibility that rearing in an insectary system mayhave modified the parasitization behavior of female D. longicaudata.IntroductionDiachasmimorpha longicaudata (Ashmead) [= Opiuslongicaudatus Ashmead], a braconid parasitoidof tephritid fruit flies, was introduced into Hawaiibetween 1947 and 1952 subsequent to the discoveryand establishment of the oriental fruit fly, Bactrocera(= Dacus) dorsalis (Hendel) (Zwaluwenberg, 1947;Clausen et al., 1965). This initial effort was undertakenprimarily to deter spoilage of fruits and vegetables fromfruit fly infestation and reduce the risks of fruit flyintroductions to the continental United States throughshipment of Hawaiian produce. Once very abundantin 1948, D. longicaudata accounted for only 30% oftotal fruit fly parasitization in 1979 and 1985 (Wong &Ramadan, 1987).Diachasmimorpha longicaudata is an obligateendoparasitoid of B. dorsalis larvae. A female lays hereggs inside fruit fly larvae where they complete development(Greany et al., 1976). Adult parasitoids emergefrom fruit fly pupae usually a few days after emergenceof adult flies from unparasitized pupae. When hostsare scarce, it is not uncommon for a female to ovipositmore than 1 egg in a host larva (Lawrence et al., 1978).Nonetheless, D. longicaudata being solitary, only oneindividual from each pupa will complete developmentto adult.From initial field collections of fruit fly pupae in theislands of Oahu, Maui, and Hawaii, a laboratory colonyof D. longicaudata was started in 1981. Subsequently,a mass-rearing procedure was developed using larvaeof B. dorsalis as propagation hosts (Wong & Ramadan,1992). Currently, D. longicaudata is one of six speciesof fruit fly parasitoids reared in this laboratory whichproduces more than a million wasps each week (Wong& Ramadan, 1992).D. longicaudata is commonly used in laboratoryและฟิลด์ assays เนื่องจากมันสามารถผลิตได้ในตัวเลขขนาดใหญ่ (วงและเดือนรอมฎอน 1992) อย่างไรก็ตามความพยายามที่จะปราบปรามกลุ่มประชากรที่มีถิ่นที่อยู่ของDorsalis เกิดใน guavas ผ่านรุ่น augmentativeบนเกาะเกาะ ฮาวายประสบความสำเร็จGR: 201003406, Pips nr 133525 BIO2KAP * 133525 * ento1508.tex 13/05/1997 12:55 v.7 p.1214(M.F. กเพอร์เซลล์ J.C. Herr, R.H. Messing และ วงศ์ T.T.Y.unpubl) แม้ มีรุ่น parasitoids 6-800 000ในระหว่างปีแรกของการทดสอบ parasitization dorsalis เกิดในฝรั่งทั่วไป Psidium guajava L. แทบเกิน 6% อย่างไรก็ตาม ไม่ทราบว่าเงื่อนไขในระหว่างการจัดส่งเกิด dorsalis parasitized pupae(กเพอร์เซลล์ et al., 1994) หรือลักษณะการทำงานของผู้ใหญ่ parasitoids พเนจรพวกเขาปรากฏขึ้น และออกจากไซต์ย่อย(Messing et al., 1994) อาจมีผลต่อประสิทธิภาพของ D. longicaudata เมื่อเร็ว ๆ นี้ ในการศึกษา interspecificแข่งขันระหว่าง 3 parasitoids dorsalis เกิดเราสังเกตว่า หญิงของ D. longicaudata มีตอบสนองน้อยในตัวอ่อนแมลงวันผลไม้ infesting parasitizingpapayas (Bautista และแฮร์ริส unpubl) อีกครั้ง การเหตุผลว่าทำไมจึงไม่เป็นที่เข้าใจเราก็อยากรู้อยากเห็นเป็นไปว่าแม่ insectaryอาจมีการปรับเปลี่ยนพฤติกรรม parasitizationD. longicaudata ดังนั้น ทางเลือกทดสอบได้ดำเนินในห้องปฏิบัติการเพื่อเปรียบเทียบหาโฮสต์และ oviposition ลักษณะการทำงานของ parasitoids หญิงระหว่างตัวอ่อนแมลงวันผลไม้ที่พัฒนาในมะละกอ C. และผู้that developed on wheat diet.Materials and methodsRearing of fruit flies for assays. Approximately 1000B. dorsalis eggs (18 to 24 h old), obtained frominsectary-colonized fruit flies (Tanaka et al., 1969;Vargas, 1989), were inoculated in half-ripe wholeC. papaya L. var. ‘Solo’ (weight range = 430–488 gper fruit) or seeded on 235 g wheat diet. Each of theserearing media was placed in 21 cm-diameter plasticcontainers with mesh screen lids. Five to 6 days later,following development of fruit fly larvae to 3rd instars(Wong & Ramadan, 1992), fruit fly larvae were teasedout from deteriorated papaya with a pair of forcepsthen collected in a petri dish with tap water. In thecase of the wheat diet, fruit fly larvae were screenedfrom the wheat substrate with a mesh sieve (1 mm2)then emptied into another dish with tap water. Fruit flylarvae were taken at random from each petri dish forsubsequent assays.Experimental procedure. A modified stinging (= oviposition)unit developed and described by Wong &Ramadan (1992) was used in all the assays. A pair ofstinging units, which contained fruit fly larvae, wereexposed concurrently to 25 females of D. longicaudata(5 to 6-d-old after emergence) inside a cubical cage(26 26 26 cm). One of the stinging units contained300 larvae reared in papaya while the other stingingunit contained 300 larvae reared on wheat diet. Duringthe tests, adult parasitoids were fed with spun honey(Sioux Honey, Sioux City, IA) and agar (source ofwater).Fruit fly larvae were not exposed naked to parasitoidsbut were mixed with wheat diet substrate beforepacking them in the stinging unit (Wong & Ramadan,1992). In our tests, besides wheat diet, we likewiseutilized pureed papaya as a substrate. Both substrateswere used merely for purposes of confirming whetherodor stimuli other than those that may be associatedwith the treatment larvae would influence the choicemade by the female parasitoids. Wheat millfeed (Armstrong,B.C. Canada), a bulking material routinely usedin B. dorsalis larval diet formulation, was mixed withpureed papaya or wheat diet substrate in equal parts(by weight) to make mixture drier. Otherwise, femaleparasitoids would shy away from the wet surface of thestinging unit (Wong & Ramadan, 1992).All assays were conducted under laboratory conditionswith mean ambient low and high temperaturesof 23 0:3 C and 24 0:2C, respectively, relativehumidity of 61:5 0:8%, and L10:D14 photoperiod.Each test was replicated 12 times. Fresh cohorts ofnaive D. longicaudata females (= no prior ovipositionexperience) were used in each replication.Test 1. Fruit fly larvae reared in papaya (Pa) or wheatdiet (Wd) were exposed to parasitoids in pureed papayasubstrate (Pus) or wheat diet substrate (Wds), respectively.Treatments were subsequently referred to as(Pa-Pus)1 and (Wd-Wds)1, respectively. The numericalsubscript in each treatment indicates the test number.
One half hour after initial exposure of (Pa-Pus)1
and (Wd-Wds)1, and at hourly intervals for a total
exposure time of 6 h, the number of parasitoids on
each stinging unit that exhibited typical host seeking
behavior, i.e., intense antennation on surface of stinging
unit, extension of ovipositor, and insertion of ovipositor
accompanied by pumping movements, were
recorded. The number of females observed on each
stinging unit per unit time was expressed in percentage
based on the original number of females assayed in
each cage.
At the end of the 6 h period (total exposure time),
the pair of stinging units were retrieved from the cage,
then 30 larvae were sampled at random from each stinging
unit. Fruit fly larvae were dissected individually
under a stereo microscope for presence of parasitoid
eggs. Percentage parasitization (pooled counts of paraento1508.tex;
13/05/1997; 12:55; v.7; p.2
215
sitized fruit fly larvae which contained 1 and 2 or more
parasitoid eggs) was computed by dividing the number
parasitized by total number dissected.
The remainder of fruit fly larvae from each stinging
unit was placed in separate holding cups (11 cmdiameter)
provisioned with vermiculite (W. R. Grace,
Cambridge, MA) for fruit fly pupation. After 9–10 d
fruit fly pupae were sifted with a mesh screen (1 mm2).
A cohort of 200 pupae was sampled and then placed
in emergence cups with mesh-screen covers until eclosion
of parasitoids. Pooled counts of emerged parasitoids
and dead parasitoid cadavers in uneclosed pupae
were recorded. Percent yields of parasitoid progeny in
each treatment was expressed as a ratio of total number
of parasitoids that developed per 200 pupae sampled.
Test 2. The same variables were generated according
to procedure described in preceding test except that
fruit fly larvae reared in Pa or Wd were both exposed
to parasitoids in -Wds substrate only. Treatments were
subsequently referred to as (Pa-Wds)2 and (Wd-Wds)2.
Test 3. Fruit fly larvae reared in Pa and Wd were both
exposed to parasitoids in -Pussubstrate only. As in Test
1, similar procedure was followed in the processing
of samples and data collection. Treatments were subsequently
referred to as (Pa-Pus)3 and (Wd-Pus)3.
Data analysis. Data on proportion of responding
D. longicaudata females, rate of parasitization, and
percentage parasitoid progeny yield were compared
between paired treatments by Student’s t-test at P =
0:05. Percentages were transformed to arcsine p
proportion
before analyses. Untransformed means (
SEM) are used in presentation of results.
Results
Test 1. The overall mean percentage of female parasitoids
that responded to fruit fly larvae following a
total exposure time of 6 h was 35% higher in (PaPus)
การแปล กรุณารอสักครู่..
diachasmimorpha นามธรรม longicaudata ( ashmead ) ถูกผลิตในห้องปฏิบัติการสำหรับ > 160 รุ่นต่อหนอนของแมลงวันผลไม้
, และแบคโทรเซรา ( Wasmann ) , การขยายพันธุ์โยธาขึ้นมาตรวจบนอาหารสูตรกึ่งสังเคราะห์ข้าวสาลี
. การเลือกการทดสอบที่ใช้ดัดแปลงแสบหน่วยการทดลองในห้องปฏิบัติการ เพื่อศึกษาว่า
insectary เลี้ยงดูมีการเปลี่ยนแปลงพฤติกรรมการโฮสต์และวางไข่ของแมลงตัวเมีย ผลการศึกษาพบว่าตัวอ่อนที่พัฒนา
แมลงวันผลไม้ในมะละกอ มะละกอ L . var . ' เดี่ยว ' , ที่ต้องการน้อยลงสำหรับวางไข่
กว่าหนอนแมลงวันผลไม้ที่พัฒนาบนข้าวสาลีอาหารเมื่อทั้งสองได้สัมผัสทั้งไร้เดียงสาดี longicaudata
หญิง ( = เพศหญิงก่อนวางไข่ประสบการณ์ )สารอาหาร ( หม้อ มะละกอ หรือข้าวสาลีอาหาร ) ซึ่งในการเปิดรับ
ตัวอ่อนแมลงไม่มีผลต่อความชอบของ longicaudata สำหรับวางไข่ตั้งครรภ์ d
ข้าวสาลีอาหารเลี้ยงหนอนแมลงวันผลไม้ การศึกษาของเราแสดงให้เห็นถึงความเป็นไปได้ที่เลี้ยงในระบบ insectary อาจ
ดัดแปลงทำลายพฤติกรรมของหญิง longicaudata แนะนำ
. .diachasmimorpha longicaudata ( ashmead ) [ = opius
longicaudatus ashmead ] ,
braconid ของแมลงเบียนแมลงวันผลไม้ tephritid , เป็นที่รู้จักในฮาวาย
ระหว่าง 1947 และ 1948 ภายหลังการค้นพบ
และสถานประกอบการของแมลงวันผลไม้ตะวันออก , แบคโทรเซรา
( = dacus ) และ ( Wasmann ) ( zwaluwenberg 1947 ;
เคลาเซิ่น et al . 1965 ) ความพยายามครั้งแรกนี้มีวัตถุประสงค์
เป็นหลักเพื่อยับยั้งการเน่าเสียของผักและผลไม้จาก
รบกวนแมลงวันผลไม้และลดความเสี่ยงของการแนะนำบิน
ผลไม้ไปยังประเทศสหรัฐอเมริกาผ่าน
การจัดส่งผลิตฮาวาย เมื่อชุกชุมมาก
ในปี 1948 , D . longicaudata คิดเป็นเพียง 30% ของ
รวมแมลงวันผลไม้ทำลายในปี 1979 และปี 1985 ( วง&
รอมฎอน , 1987 ) longicaudata เป็น Obligate
diachasmimorphaendoparasitoid พ. และตัวอ่อน ตัวเมียวางไข่ของเธอ
ภายในผลไม้ หนอนแมลงวันที่สมบูรณ์การพัฒนา
( greany et al . , 1976 ) แมลงผู้ใหญ่โผล่ออกมาจากดักแด้แมลงวันผลไม้
ปกติไม่กี่วันหลังงอก
ผู้ใหญ่บินจาก unparasitized ดักแด้ . เมื่อโยธา
ขาดแคลน มันไม่แปลกสำหรับผู้หญิงที่จะ oviposit
มากกว่า 1 ไข่เป็นหนอน ( Lawrence et al . , 1978 ) .
กระนั้นD . longicaudata อยู่โดดเดี่ยว , เพียงหนึ่ง
บุคคลจากแต่ละดักแด้จะสมบูรณ์พัฒนา
จากสนามถึงผู้ใหญ่ เริ่มต้นคอลเลกชันของดักแด้แมลงวันผลไม้ใน
เกาะ Oahu Maui และฮาวาย , ห้องปฏิบัติการอาณานิคม
D . longicaudata เริ่มต้นในปี 1981 ต่อมา
มวลเลี้ยงดูกระบวนการพัฒนาโดยใช้ตัวอ่อน
B และเป็นการโยธา ( วง&รอมฎอน ,
1992 ) ขณะนี้ , Dlongicaudata เป็นหนึ่งในหกชนิดของแตนเบียนแมลงวันผลไม้ ?
นี้ห้องปฏิบัติการที่ผลิตมากกว่าล้านตัวต่อแต่ละสัปดาห์ ( วง
&รอมฎอน , 1992 ) .
d longicaudata มักใช้ในห้องปฏิบัติการ
และสนาม ) เพราะสามารถผลิตได้อย่างง่ายดายใน
ตัวเลขขนาดใหญ่ ( วง&รอมฎอน , 1992 ) แต่ความพยายามที่จะปราบปรามกลุ่มถิ่น
Bและในฝรั่งผ่านแต่งเติมออก
สําเร็จบนเกาะ Kauai , GR ฮาวาย
: 201003406 จุด 133525 133525 Nr bio2kap * * ento1508.tex ; 13 / 05 / 1997 ; 12:55 ; v.7 ; ป.
( 214 ) เพอร์เซล , J.C . แฮร์ , R.H . ยุ่ง& t.t.y. วงศ์ ,
unpubl . ) แม้จะมีรุ่น 6 – 800 000 ตัวเบียน
ในระหว่างปีแรกของการทดสอบและทำลายของ B .
ในฝรั่งทั่วไปฝรั่งไม่เกิน L .
6 % แต่มันไม่เป็นที่รู้จักไม่ว่าจะเป็นเงื่อนไข
ในระหว่างการจัดส่งของเป็นปรสิตพ. และดักแด้
( เพอร์เซลล์ et al . , 1994 ) หรือพฤติกรรมที่ประพฤติชั่วของผู้ใหญ่เช่นที่พวกเขาเกิดและตัวเบียน
กระจายจากการปล่อยเว็บไซต์
( ล้อเล่น et al . , 1994 ) อาจมีผลกระทบต่อประสิทธิภาพ
D . longicaudata . เมื่อเร็วๆ นี้ ในการศึกษาเกี่ยวกับแมลงในการแข่งขัน interspecific
3 พ.
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