RESEARCH PAPER

Endoparasitoids of larval Anomis privata (Lepidoptera:Noctuidae), major pest of Hibiscus syriacus(Columniferae: Malvaceae)Youngjin KIM, Youngho CHO, Yong-Gu HAN and Sang-Ho NAM

Department of Biology, Daejeon University, Daejeon, Korea

Correspondence

Sang-Ho Nam, Department of Biology,Daejeon University, Daejeon 300-716,Korea. Email: shnam@dju.kr

Received 22 August 2011;accepted 16 September 2011.

doi: 10.1111/j.1748-5967.2011.00348.x

Abstract

Endoparasitoids of Anomis privata larvae include five species in three families oftwo orders. In this work, two species of Hymenoptera Braconidae (Cotesia sp.,Microplitis sp.), one species of Ichneumonidae (Mesochorus vittator) and twospecies of Diptera Tachinidae (Exorista (Podotachina) sorbillans, Timavia amoena)were investigated. Of the 261 larvae of A. privata examined, 32 had a parasite, so therate of parasitism was 12.26%. Parasitism by taxon was the highest, at 10.35% (27individuals), in Cotesia sp. in Hymenoptera Braconidae. Parasitoids of Braconidaeand Ichneumonidae were larval parasitoids. A parasitic insect of Tachinidae was alarva–pupal parasitoid. Solitary parasitoids included Microplitis sp. in Braconidaeand E. sorbillans in Tachinidae. Gregarious parasitoids included Cotesia sp. inBraconidae, M. vittator in Ichneumonidae and T. amoena in Tachinidae. There wasalso a multiparasitoid (T. amoena) and two superparasitoids (Cotesia sp., M. vitta-tor). A larva of A. privata sought feed even after it was parasitized every parasitoidinvestigated in this study, so five species of parasitoids were all koinobiont.

Key words: Anomis privata, endoparasitoid, Hibiscus syriacus, insect pest.

Introduction

Anomis privata (Lepidoptera: Noctuidae) is a harmful insectthat occurs two or three times a year between May andOctober. It is a phyllophagous pest that damages leaves ofH. syriacus leaving only a midrib remaining (Fig. 1). Apolyphagous pest damages H. syriacus, Hibiscus mutabilis,Prunus persica, Prunus salicina and Malus pumila etc. (Kimet al. 1982; Lee & Chung 1997; Shin 2001). Anomis privatais distributed in Korea and Japan, China, Southeast Asia,India, Australia and North America (Kononenko et al.1998). Since 61 000 units of H. syriacus were plantedaccording to the dissemination plans of the government from1983 to 2001, pest damage to H. syriacus first occurredlocally, than spread nationally in 1990 (Lee & Chung 1997;Song 2004). For control of this species, pesticides such asDEP, MEP, pyrethroid, trichlorfon and carbaryl have beenused (Lee & Chung 1997; Song 2004). Excessive spraying

with agricultural chemicals inhibits development of a pest,but also decreases the number of other biota in an ecosystem,including natural enemies of the pest, when selective spray-ing is not performed (Debach 1974). Therefore, biologicalcontrol using parasitoids has been the focus of recentstudies. Parasitoids in an ecosystem provide many speciesand individuals that function to control the density of theirhosts (Godfray 1994).

In addition, A. privata is an entomophagous insect whichhas evolved successfully by interaction with a host (Vinson1975; Vinson & Iwantsch 1980). It shows various life cyclesas it is parasitic on each developmental stage of hosts invarious habitats (Strand 2000; Harvey 2005).

Thus, the aim of this study is to confirm the possibility ofusing parasitoid species to control A. privata, which is theworst pest of H. syriacus. The significance is that this plantis a national flower that is often used as a garden tree and totree line public streets.

Entomological Research 41 (2011) 257–263

© 2011 The AuthorsEntomological Research © 2011 The Entomological Society of Korea and Blackwell Publishing Asia Pty Ltd

Materials and methods

Study sites

To confirm the existence of endoparasitoids of A. privata,this investigation was performed in Chu-dong and Jusan-dong, Dong-gu, Daejeon (South Korea), where there is asubstantial H. syriacus community. There larvae of A. pri-vata were collected without regard for which instars werebeing collected (Fig. 2).

Insects

Collected larvae were placed in Petri dishes (diameter100 mm, height 15 mm) with filter paper (90 mm #2, Advan-tec, Toyo, Japan). For constant humidity, 1 mL of primarydistilled water was added every day. Leaves of H. syriacus,previously washed in flowing water, were provided as feedand then larvae were reared in a growth chamber (VS-91G09MN; Vision Scientific Co., Bucheon, Korea) underconditions of 27°C, 70% relative humidity (RH) and 12 000

(a) (b)

Figure 1 (a) Larva of Anomis privata. (b) Damage on Hibiscus syriacus caused by Anomis privata.

Figure 2 Map showing study areas in Chu-dong and Jusan-dong, Daejeon, South Korea.

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258 Entomological Research 41 (2011) 257–263© 2011 The Authors. Entomological Research © 2011 The Entomological Society of Korea and Blackwell Publishing Asia Pty Ltd

� 100 lx during the day, and 27°C, 70% RH and 0 lx duringthe night under conditions of 14 h light : 10 h dark(LD 14:10). Parasitoids were reared under the same condi-tions as A. privata larvae and species with eclosion werecollected and identified.

External morphological characteristics

of parasitoids

The size of its cocoon, as well as the body length, forewinglength, hindwing length, antenna length and head capsulewidth of a dry parasitoid of A. privata larvae were measuredby a BMI v 4.0 image analysis system (BMI, Ansan, Korea).Body length was measured from the head to the end of theabdomen along the midline of the side. Antenna and wingswere measured in a flat state. Head capsule width was mea-sured between compound eyes of the front part, and thecocoon was measured on its major and minor axes.

Results and discussion

Composition of species of parasitoid

Parasitoids of A. privata larvae consisting in total of twoorders, three families and five species, all of which areendoparasitoids, were found in the A. privata larvae. Byorder, Hymenoptera Braconidae (Cotesia sp., Microplitissp.) and Ichneumonidae (M. vittator) were investigated asone order, two families and three species. Diptera Tachinidae(Exorista sorbillans, Timavia amoena) was investigated asone order, one family and two species (Table 1).

Emergence was highest (84.39%) in Cotesia sp.(Hymenoptera: Braconidae). That of Microplitis sp. (Bra-conidae) was 6.25%, while Mesochorus vittator (Ichneu-monidae), and the two species E. sorbillans and T. amoena(Diptera, Tachinidae), were all 3.12%. Park et al. (2004)reported Cotesia sp. as a parasitic natural enemy for H. syri-acus pests, except predatory pests, and four more specieswere confirmed in this investigation.

Cotesia sp. is a typical parasitoid estimated to includeabout 400 species in North America and 1000 species world-wide (Shaw & Huddleston 1991; Whitefield 1995). It isNoctuidae, Sphingidae, Lymantriidae, Arctiidae, Pieridaeand Geometridae (Alice & James 2004) and was first con-firmed as a parasitoid of A. privata in Korea.

There are 140 species of the genus Microplitis in theworld. Members of this genus are used for biological controlof 71 species in 17 families of parasitoids of larval Noctu-idae (Austin & Dangerfield 1993). Microplitis demolitor is aparasitoid of Helicoverpa armigera, a well-known pest ofCapsicum annuum, Nicotiana tabacum, Zea mays and Gos-sypium arboretum (Shepard et al. 1983). In China, Wanget al. (1984) reported that Microplitis mediator was parasiticon 40 species of Lepidoptera and that it makes up 58% oflarva parasitoids. Microplitis plutelle is parasitic on 30–40%of 3–4 instar larvae of Plutella xylostella, one of the mainpests of economic crops in the world (Jianxiang et al. 2001).Thus, M. plutelle is very important for biological control. Inaddition, M. demolitor was reported as a parasitoid, parasiticon eight species of Noctuidae in Australia (Shepard et al.1983; Austin & Dangerfield 1993). In Korea, 16 species ofgenus Microplitis were recorded in the host insect Acronictarumics (Lepidoptera: Noctuidae) (Cho et al. 2006). This wasits first confirmation as a parasitoid of A. privata.

Exorista sorbillans is a major parasitoid of Bombyx mori(Lepidoptera: Bombycidae) (Yoshimi 2005). In Japan, 46species including Mythimna separate, Spodoptera litura andMamestra brassicae were recorded as host insects ofExorista japonica, which is an allied species (Oku & Koba-yashi 1974). It was confirmed as a parasitoid of A. privatafirst in Korea.

Parasitism

Altogether, 261 larvae of A. privata were collected and 32larvae among those had a parasitoid, so the rate of parasitismwas 12.26%. Among those 32 larvae, HymenopteraBraconidae was 10.35%, Ichneumonidae was 0.38% andDiptera Tachinidae was 0.76% (Table 1). Parasitism by

Table 1 Species composition and parasitism rates

Orders Families Species PA/CI

Parasitism (%)

Species Families Total

Hymenoptera Braconidae Cotesia sp. 27/261 10.35 11.11 12.26Microplitis sp. 2/261 0.76

Ichneumonidae Mesochorus vittator 1/261 0.38 0.38Diptera Tachinidae Exorista (Podotachina) sorbillans 1/261 0.37 0.73

Timavia amoena 1/261 0.18

PA, parasitoid appearance on A. privata; CI, collected individuals from larva of A. privata.

Endoparasitoids of larval Anomis privata

259Entomological Research 41 (2011) 257–263© 2011 The Authors. Entomological Research © 2011 The Entomological Society of Korea and Blackwell Publishing Asia Pty Ltd

taxon was 90.625% in Hymenoptera and 6.25% in Diptera.This result means that parasitic bees made up more than 80%of all the parasitoids (Quick 1997) and could be a generalparasitic pattern. Within Hymenoptera, parasitism was93.75% in Hymenoptera Braconidae and 6.25% inIchneumonidae.

Species composition and parasitism by month

Parasitoid species composition in A. privata larvae eachmonth showed Microplitis sp. in May, Cotesia sp.,T. amoena and E. sorbillans in July, Cotesia sp. in Augustand September and Cotesia sp. and M. vittator in October.Anomis privata was observed between May and October.Hibiscus syriacus blooms between July and September, soA. privata appeared in this period because H. syriacus is ahost plant for A. privata. Because of an increase in thenumber of A. privata at this time, species composition ofparasitoids was the most diverse, with three species in July.

Relative to the total number of infestations, parasitism ofA. privata larvae by month is shown in (Table 2). In May,parasitism was the highest because two of three individualshad parasitoids. Hatching of A. privata larvae occur morethan twice a year (Lee & Chung 1997; Shin 2001). Weinvestigated from May through October, but the period ofintensive appearance was between July and September.Parasitism of A. privata in each month was high in July andAugust, except May because of limited collection. Parket al. (2004) reported that parasitism of Anomis mesogona,which is also a pest of H. syriacus, was 11.4% because 5.7individuals of Cotesia sp. were parasitic on 50 larvae inAugust, the peak period. Parasitism in the same period washigher in this study.

External morphology of parasitoids

For 261 males of Cotesia sp., average eclosion time fromcocoon to adult was 10.22 � 6.07 days and the life span ofan adult was 2.57 � 0.90 days. Cocoons were white oryellow. The length was 3.51 � 0.14 mm on the major axisand 1.49 � 0.11 mm on the minor axis. Adult body lengthwas 2.37 � 0.13 mm. Forewing length was 2.74 � 0.15 mm,hindwing length 1.77 � 0.10 mm, antenna length 2.34 �

0.05 mm and head capsule width 0.62 � 0.04 mm (Table 3).For two males of Microplitis sp., eclosion time from

cocoon to adult was 5 � 0 days and life span of adults was

3 � 0 days. Cocoons were dark gray. The length was 3.88 �

0.06 mm for the major axis and 1.69 � 0.05 mm for theminor axis. Body length of an adult was 3.10 � 0.11 mm.Forewing length was 2.97 � 0.04 mm, hindwing length 1.85� 0.04 mm, antenna length 3.17 � 0.04 mm and headcapsule width 0.95 � 0.02 mm.

For three males and four females of M. vittator, eclosiontime from cocoon to adult was 9 � 0 days in both sexes andthe life span of adults was 3 � 0 days. The length of acocoon was 3.68 � 0.01 mm for the major axis and 1.62 �

0.01 mm for the minor axis in males. It was 3.66 � 0.02 mmfor the major axis and 1.63 � 0.01 mm for the minor axis infemales, so the length was similar in both sexes. Body lengthof an adult was 3.90 � 0.01 mm in males and 4.13 �

0.01 mm in females. Forewing length was 3.25 � 0.02 mmin males and 4.12 � 0.01 mm in females, hindwing lengthwas 2.10 � 0.02 mm in males and 1.95 � 0.04 mm infemales, antenna length was 4.10 � 0.02 mm in males and4.22 � 0.01 mm in females and head capsule width was 0.59� 0.01 mm in males and 0.63 � 0.01 mm in females. Exceptfor hindwing length, females were longer than males.

For one male of E. sorbillans, eclosion time from cocoonto adult was 21 days and life span of the adult was 5 days. Thecocoon was dark brown. Cocoon length was 8.02 mm for themajor axis and 3.76 mm for the minor axis. Body length of anadult was 9.43 mm. Forewing length was 7.29 mm, antennalength was 1.49 mm and head capsule width 3.14 mm. Aoki(1969) reported that body length of an adult of this species,which is a parasitoid of B. mori, was 12.0 mm (male),11.4 mm (female) and that pupal duration and life span of anadult were 1–14 days and 14 days, respectively; thus, thisresult was different from our result. It may have been causedby differences in the growth of E. sorbillans larvae accordingto the size and age of the host insect larva.

For three males and one female of T. amoena, eclosiontime from cocoon to adult was 17 � 0 days in both sexes andthe life span of adults was 8 � 0 days. The length of cocoonswas 7.77 � 0.18 mm for the major axis and .33 � 0.06 mmfor the minor axis in males. It was 6.98 mm for the majoraxis and 2.98 mm for the minor axis in females. Body lengthof an adult was 9.49 � 0.06 mm in males and 9.32 mm infemales. Forewing length was 7.80 � 0.03 mm in males and7.69 mm in females. Antenna length was 1.27 � 0.03 mm inmales and 1.17 mm in females. Head capsule width was 2.81� 0.03 mm in males and 2.66 mm in females. Generally,males were larger than females.

Table 2 Parasitism by monthsMay June July August September October Total

I/CI 2/3 0 10/25 5/14 4/33 11/186 32/261Parasitism (%) 66.67 0 40.00 35.70 12.12 5.91 12.26

I/CI, individuals (parasitoid)/collected individuals from larva of A. privata.

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260 Entomological Research 41 (2011) 257–263© 2011 The Authors. Entomological Research © 2011 The Entomological Society of Korea and Blackwell Publishing Asia Pty Ltd

Ecological characteristics of these parasitoid

Among the endoparasitoids of A. privata, Cotesia sp.(Hymenoptera: Braconidae), which showed the highest rateof parasitism and density, is a larval parasitoid that forms anexterior cocoon from which to escape from a host body afteremergence. Larvae of A. privata cast their skins and ingestedregardless of whether it hosted parasitoids, so they are koi-nobiont. In one larva of A. privata, eclosion of dozens ofCotesia sp. was observed, so it is a gregarious parasitoid(Table 4). It showed up every month between July andOctober, so it is also a multivoltine insect. Species of thegenus Cotesia are parasitic bees including in microgastri-nae, the largest classification in North America. In particu-lar, Cotesia is widely used for biological control of pests butit does not control many species. However, it is very impor-tant to control a pest community in the specific area. The lifecycles of some Cotesia species are used as a key model forinteraction between hosts and parasitoids in physiology,molecular biology, behavioral ecology and ecology (Alice &James 2004).

Microplitis sp. is a parasitoid of larval A. privata. With aparasitoid, A. privata could cast its skin and ingest so it iskoinobiont. However, unlike Cotesia sp., eclosion of onlyone parasitic bee occurs in one larva, so it is a solitaryparasitoid. In addition, most species of this genus arepolyphagous and are parasitic on two or more hosts (Kadashet al. 2003). Jiancheng et al. (2006) reported that M. media-tor, which is parasitic on Mythimma separate (Noctuidae),showed high rates of parasitism (60–70%) during 2–3 instarsin the parasitism test, so the possibility for biological controlwas confirmed.

Mesochorus vittator is a superparasitoid of Cotesia sp.,which is was koinobiont because it cast its skin and ingestedwith a parasitoid. Mesochorus vittator is also a gregariousparasitoid. About a hundred hosts for M. vittator have beenrecorded and it is distributed in 17 countries on the conti-nents of Australia, Europe and Asia (Lee & Suh 1991; Suhet al. 1997). The Mesochorus genus is a huge group inMesochorinae. About 500 species re reported worldwide, ofwhich 140 are Palearctic, 12 African, 2 Australian, 96 Nearc-tic and 245 Neotropic (Lee & Suh 1991). In addition, Meso-chorinae is a superparasitoid for other species withinBraconidae or Tachinidae. It is a potential risk factor andimportant for biological control in particular (Whal 1993).

Exorista sorbillans and T. amoena (Diptera: Tachinidae)are parasitic on larvae of a host and they emerge from anexterior cocoon. In addition, a larva cast its skin and ingestedregardless of parasitism, so it was koinobiont. However,E. sorbillans showed eclosion of only one individual in onelarva, so it was a solitary parasitoid. Aoki (1969) reportedthat this species was a multivoltine parasitic insect with 5–6cycles of birth with oviposition on the surface of a host larva,T

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Endoparasitoids of larval Anomis privata

261Entomological Research 41 (2011) 257–263© 2011 The Authors. Entomological Research © 2011 The Entomological Society of Korea and Blackwell Publishing Asia Pty Ltd

and that it oviposited more than 800 eggs during a produc-tion period. Timavia amoena cast its skin and ingestedregardless of parasitism, so it is koinobiont. There was eclo-sion of four individuals of Tachinidae in one larva, so it wasa gregarious parasitoid.

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Table 4 Ecological characteristic of endoparasitoids on A. privata

Parasitizedhost age

Parasitoidescape stage

Parasitizedhost ecdysis

Food consumption ofparasitized host

Solitary (S) orGregarious (G)

Idiobiont (I) orKoinobiont (K)

Hymenoptera: BraconidaeCotesia sp. Larva Larva Yes Yes G KMicroplitis sp. Larva Larva Yes Yes G KIchneumonidaeM. vittator Larva Larva Yes Yes G KDiptera: TachinidaeE. sorbillans Larva Pupa Yes Yes S KT. amoena Larva Pupa Yes Yes G K

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