natural enemies of bagworms on oil palms - Hawaii Biological Survey

Pacific Insects 14 (1): 57-71 20 March 1972 

THE NATURAL ENEMIES OF BAGWORMS ON OIL 

PALMS IN SABAH, EAST MALAYSIA 

By T. Sankaran 1 and R. A. Syed 2 

Abstract: Three species <strong>of</strong> <strong>bagworms</strong>, Mahasena corbetti Tams, Metisa plana Wlk. 

and Cremastopsyche pendula Joannis, are major pests <strong>of</strong> oil palms in Sabah. Parasites 

recorded from these and other Psychids during 1969 and 1970 were: Apanteles sp., Aulosaphes 

sp., Echthromorpha agrestoria Swed., Xanthopimpla sp., Sympiesis sp., Pediobius 

sp., Eozenillia equatorialis Towns., Palexorista solennis (Wlk.) and Sarcophaga sp. from 

M. corbetti; Apanteles metesae Nixon, Eupelmus catoxanthae Ferr. and Eozenillia psychidarum 

Bar. from M. plana ; Aulosaphes psychidivorus Muesebeck and A. metesae from 

C. pendula; Xanthopimpla pedator Fabr., E. equatorialis and Brachymeria sp. from Clania 

tertia Tempi.; Sericopimpla flavobalteata Cam. from Cryptothelea cardiophaga Westw.; 

and Parasierola sp., Brachymeria sp. and Pediobius lividiscutum (Gahan) from? Pteroma 

sp. C. tertia and C. cardiophaga were also attacked by A. psychidivorus. Paraphylax 

rufipes Cam. was a common secondary parasite <strong>of</strong> <strong>bagworms</strong>. Brachymeria sp., B. deesensis 

Cam. and B. lugubris Wlk. were reared from E. equatorialis ; E. catoxanthae, P. 

lividiscutum and Brachymeria sp. from A. metesae; and Brachymeria sp. from E. psychidarum. 

Apanteles angaleti Muesebeck and Anastatoidea brachartonae Gahan emerged in cages 

with M. corbetti. A. psychidivorus, P. lividiscutum and E. sp. ? equatorialis were reared 

from M. corbetti bags formed <strong>of</strong> leaves from an unknown host plant. A Fislistina sp. 

was obtained from Eumeta sp. on cocoa. Two predators, Callimerus bellus Gorh. associated 

with M. corbetti, and Sycanus sp. with C. pendula, were very scarce. 

Pathogens isolated from dead <strong>bagworms</strong> were Paecilomyces fumoso-roseus (Wize) Brown 

and Smith, from larvae <strong>of</strong> M. plana, C. pendula, C. tertia and C. cardiophaga, and a 

possible nonoccluded virus and an Aspergillus sp. from those <strong>of</strong> M. corbetti. 

Although several species <strong>of</strong> <strong>bagworms</strong> (Lep., Psychidae) occur on oil palms (Elaeis 

guineensis Jacq.) in Sabah, East Malaysia, only three <strong>of</strong> them, Mahasena corbetti Tams, 

Metisa plana Walker and Cremastopsyche pendula Joannis are, at the present time, <strong>of</strong> 

major economic importance as pests (Sankaran, 1970), In W. Malaysia M. corbetti has 

long been known to be a pest <strong>of</strong> coconut palms, while the other 2 species have caused 

severe outbreaks on oil palms. Native parasites play an important role in checking 

these <strong>bagworms</strong> but experience in W. Malaysia has shown that the use <strong>of</strong> broad-spectrum, 

residual, contact insecticides has hampered the beneficial work <strong>of</strong> these parasites, 

leading to fresh pest outbreaks (Wood, 1968). Our knowledge <strong>of</strong> the parasites <strong>of</strong> these 

<strong>bagworms</strong> was more or less confined to Indonesia and W. Malaysia. With the introduction 

and rapid extension <strong>of</strong> oil palm cultivation in E. Malaysia <strong>bagworms</strong> have 

become serious since 1966. Their indigenous parasites and other <strong>natural</strong> <strong>enemies</strong> were 

investigated in Sabah by the authors during 1969 and 1970 with a view to the introduc- 

1. Indian Station, Commonwealth Institute <strong>of</strong> Biological Control, Bangalore, India. 

2. Sabah Sub-station, Commonwealth Institute <strong>of</strong> Biological Control, Tuaran, Sabah (Malaysia).

58 Pacific Insects Vol. 14, no. 1 

tion <strong>of</strong> additional species from other areas as suggested by Simmonds (1967). A 

search for promising bagworm parasites has been taken up by the C.I.B.C. W. Indian 

Station in Trinidad. 

This paper deals with the <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> oil palm <strong>bagworms</strong> recorded in Sabah 

during 1969 and 1970. The same or other species <strong>of</strong> <strong>bagworms</strong> were collected on various 

other economic plants in order to rear out parasites, if any, to ascertain the hostrange 

and host-plant associations <strong>of</strong> the species concerned. 

PARASITES AND PREDATORS OF OIL PALM BAGWORMS 

No egg-parasite <strong>of</strong> any species <strong>of</strong> bagworm has so far been obtained in Sabah. It 

seems unlikely that these would occur because the eggs remain well protected within 

the mother's hardened pupal skin, which itself is enclosed in a bag. 

Several species <strong>of</strong> larval and pupal parasites obtained are listed under their hosts. 

Mahasena corbetti Tams 

Several species <strong>of</strong> larval parasites were reared from this bagworm but with most 

levels <strong>of</strong> parasitism were low. A Tachinid larval parasite, Eozenillia equatorialis Townsend, 

and an Ichneumonid, Echthromorpha agrestoria Swed., which emerged from the 

pupal cases, were by far the most important species recorded but appeared to be incapable 

<strong>of</strong> building up their populations to bring about any substantial reduction <strong>of</strong> the 

pest in an outbreak. 

HYMENOPTERA 

Braconidae 

Apanteles sp.: Cocoons <strong>of</strong> this solitary parasite were found in 4 <strong>of</strong> 45 small larval 

bags collected at Sg. Manila near Sandakan in September 1969. In January 1970 a 

single parasite emerged from one bag in a lot <strong>of</strong> 469 young larvae collected at Mostyn 

Estate. There has been no previous record <strong>of</strong> any Apanteles sp. parasitising M. corbetti. 

Aulosaphes sp.: Twelve cocoons <strong>of</strong> an Aulosaphes sp. were obtained from a larval 

bag in a lot <strong>of</strong> 47 specimens collected at Giram Estate in June 1969. Many <strong>of</strong> these 

cocoons were empty or had imperfectly developed adults. In November a single adult 

emerged from a bag and a group <strong>of</strong> 6 cocoons was found in another bag collected at 

the same estate. This species is different from Aulosaphes psychidivorus Muesebeck, 

which is a common parasite <strong>of</strong> several genera <strong>of</strong> Psychids in Sabah. 

Ichneumonidae 

Echthromorpha agrestoria Sw Qd.: In December 1969 this large Ichneumonid was reared 

for the first time from female pupal cases <strong>of</strong> M. corbetti collected at Mostyn Estate. 

In one field ll specimens in a collection <strong>of</strong> 95 were found parasitised. The parasite 

adults emerged from the hardened pupal skins <strong>of</strong> the hosts. In unparasitised individuals 

such pupal skins contain female moths. Therefore, it seems that parasitism actually 

occurs in the late larval or pupal stage but the parasite completes its development only 

after the host reaches the imaginal stage but is still enclosed in the pupal skin. The 

hardened pupal skin <strong>of</strong> the host is lined internally by a thin, parchment-like membrane. 

Xanthopimpla sp.: A single specimen was reared from a female pupal case <strong>of</strong> M.

1972 Sankaran and Syed : The <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> bag worms on oil palms 59 

corbetti in a lot <strong>of</strong> 71 bags collected at Giram Estate in August 1969. A pupa <strong>of</strong> the 

same parasite was found in another specimen which was dissected in the field. This 

is also a new record <strong>of</strong> an Ichneumonid from M. corbetti. 

Eulophidae 

Pediobius sp.: A single specimen was obtained from a larval bag among 404 specimens 

<strong>of</strong> various stages <strong>of</strong> M. corbetti collected at Mostyn Estate in April 1969. The 

Commonwealth Institute <strong>of</strong> Entomology, London, has retained the specimen, commenting 

that this is a peculiar species, not represented in the British Museum collection. 

Sympiesis sp.: Eighteen parasites emerged from three larval bags <strong>of</strong> M. corbetti 

collected at Giram Estate in November 1969. On opening the bags, empty pupal skins 

<strong>of</strong> the parasite were found in them. Species <strong>of</strong> Sympiesis are generally parasitic on 

coleopterous, dipterous and lepidopterous leafminers. The only other record <strong>of</strong> a 

Sympiesis sp. parasitising a Psychid (Cryptothelea crameri West.) is from Ceylon (Thompson, 

1954). 

DIPTERA 

Tachinidae 

Eozenillia equatorialis Townsend: This was the only important dipterous parasite <strong>of</strong> 

M. corbetti in Sabah. It was frequently reared from fully grown larvae collected in 

all the plantations visited in eastern Sabah but was scarce in the Beaufort area on the 

west coast. Rarely two or three parasites but generally only one emerged per host. 

In two instances, final stage larvae <strong>of</strong> this Tachinid passed through a pupal period <strong>of</strong> 9 

and ll days, respectively. In the laboratory, adult parasites fed on dilute honey lived 

for a maximum <strong>of</strong> 5 days. Puparia have been found in larval bags collected almost 

throughout the year. However, parasitism never increased along with the host's population 

when outbreaks occurred at Mostyn Estate. This Tachinid appears to exercise 

some measure <strong>of</strong> control <strong>of</strong> Mahasena only when the bagworm population remains at 

low levels on oil palms and other alternative host plants. At the time <strong>of</strong> an outbreak 

at Mostyn Estate in November and December 1969 parasitism was hardly 1-2 %. In 

the same estate, E. equatorialis and another Tachinid, Palexorista solennis (Walker), together 

accounted for only 12.6 % parasitism in a collection <strong>of</strong> 400 bags in April 1969 

although the bagworm population here had increased earlier during an outbreak. In 

the Lormalong Scheme, 200 bags were collected in the same month, out <strong>of</strong> which only 

7 had parasite puparia. 

In April 1970 up to 55 percent parasitism <strong>of</strong> Mahasena corbetti on Albizzia sp. was 

recorded in areas around Kota Kinabalu, and at Pamol Estate 80 % <strong>of</strong> the <strong>bagworms</strong> 

in a small sample was found parasitised by E. equatorialis in September. In both areas 

the bagworm population was low. Some specimens <strong>of</strong> E. equatorialis in the insect 

collection <strong>of</strong> the Agricultural Research Center, Tuaran, were earlier determined as Exorista 

psychidarum Bar. Several specimens reared from M. corbetti by the authors 

and submitted to the Commonwealth Institute <strong>of</strong> Entomology, London, were identified 

as Eozenillia equatorialis Towns. E. equatorialis and E. psychidarum are very closely 

allied but distinct species and have been confused in the past. The true E. psychidarum, 

<strong>of</strong> which the holotype is in the British Museum, has the eyes completely bare, and E. 

equatorialis has the eyes densely haired. There are also differences in the male genitalia 

(R. W. Crosskey, pers. com.).


Exorista (=Zenillia) quadrimaculata Bar. is a parasite <strong>of</strong> Cryptothelea variegata Sn. in 

India and Malaya (Thompson, 1945), M. corbetti in Malaya and Thailand (Thompson, 

1946a; Meksongsee, 1963) and unidentified Psychids in Ceylon, India, Malaya and 

Sumatra (Thompson, 1946b). Crosskey (1967a) has synonymised this species with 

Nealsomyia rufella (Bezzi). 

Eozenillia ? equatorialis Towns.: A single specimen <strong>of</strong> this doubtful species was reared 

from a larval bag collected at B. A. L. Estates in June 1969. 

Palexorista solennis (Walker) : This was obtained in small numbers from larval bags 

<strong>of</strong> M. corbetti collected at Mostyn and Giram Estates on several occasions in both years. 

The puparium can be easily distinguished from that <strong>of</strong> E. equatorialis by its hairy 

surface, brown color, more or less parallel sides and by the very prominent posterior 

spiracles, which have three raised slits, resembling a group <strong>of</strong> three mussels. The puparium 

<strong>of</strong> E. equatorialis is dark brown with a smooth and shining surface, and with an 

asymmetrical swelling on one side. P. solennis is the commonest Oriental species <strong>of</strong> 

Palexorista, distributed from India through the Far East and New Guinea to Samoa and 

is polyphagous, parasitising several Lepidoptera (Crosskey, 1966; 1967a and b). The 

species has been recorded from the toon shoot-borer Hypsipyla robusta Moore in India 

(CIBC, unpublished record). 

Sarcophagidae 

Sarcophaga sp.: A Sarcophaga sp. was occasionally obtained from larval bags collected 

at Mostyn and Giram Estates. 

Sarcophaga antilope Boettch. and S. peregrina R.-D. have been recorded as parasites <strong>of</strong> 

M. corbetti in Malaya (Thompson, 1946a). 

HYPERPARASITES 

Three species <strong>of</strong> hyperparasites, Brachymeria sp. (not in British Museum), B. deesensis 

Cam. and B. lugubris Walk, were reared from the puparia <strong>of</strong> E. equatorialis. These 

emerged by cutting circular holes in the puparia while the Tachinids from unparasitised 

puparia emerged by splitting open their anterior ends. Nine <strong>of</strong> 25 puparia (36 %) obtained 

from Mahasena bags at Ranau in November 1969 were found to have been 

attacked by Brachymeria. 

In Indonesia, B. lugubris is a secondary parasite <strong>of</strong> the coconut leaf-moth Artona catoxantha 

Hmps. through the Tachinid Ptychomyia remota Aldr, (van der Vecht, 1950). 

PARASITES OF UNCERTAIN STATUS 

Apanteles angaleti Muesebeck: A single specimen <strong>of</strong> A. angaleti emerged in a laboratory 

cage with 47 larvae <strong>of</strong> M. corbetti collected at Giram Estate in June 1969. At 

that time the host relationship <strong>of</strong> Apanteles with M. corbetti was uncertain. Since another 

Apanteles sp. was later found to be parasitic on young larvae <strong>of</strong> M. corbetti, A. 

angaleti may also prove to be a primary parasite <strong>of</strong> this bagworm. Nixon (1965) mentions 

Sylepta derogata F. and Pectinophora gossypieUa Saunders as hosts, and Pakistan, 

India and Sumatra as the distribution <strong>of</strong> A. angaleti. Like M. corbetti these two hosts 

also live in protected situations. 

Anastatoidea brachartonae Gahan: A single specimen <strong>of</strong> this Eupelmid emerged in a

1972 Sankaran and Syed: The <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> <strong>bagworms</strong> on oil palms 61 

laboratory cage with 24 larvae <strong>of</strong> M. corbetti collected at Sg. Manila in June 1969. A. 

brachartonae has been recorded as a parasite <strong>of</strong> Psyche Vitrea Hmps. in Ceylon (King, 

1933). In Indonesia it occasionally attacks the prepupae and pupae <strong>of</strong> Artona catoxantha 

but is more <strong>of</strong>ten a hyperparasite <strong>of</strong> the Tachinids Ptychomyia remota and Cadurcia 

leefmansi Bar. and <strong>of</strong> Apanteles anonae Rohw. and Goryphus spp. (van der Vecht, 

1950). 

PREDATORS 

In a larval bag <strong>of</strong> M. corbetti collected at Mostyn Estate in August 1969 a freshly 

emerged adult and a larva (in a cocoon) <strong>of</strong> a predator, Callimerus bellus Gorh. (Coleoptera, 

Cleridae), were found. The presence <strong>of</strong> a second, empty, cocoon within the 

same bag indicated that the adult beetle might have emerged from it. Callimerus arcufer 

Chpm. is a predator <strong>of</strong> <strong>bagworms</strong> in West Malaysia (Wood, 1968). It has also been 

recorded feeding on Artona catoxantha and on the parasite Ptychomyia remota in Malaya 

(Corbett and Miller, 1933). There is no record <strong>of</strong> C. arcufer from Sabah. 

Once an unidentified Chrysophid larva was found within a bag collected at Sg. Manila. 

This carried on its back small bits <strong>of</strong> dried, oil palm leaves, apparently derived 

from the Mahasena bag, and is suspected to be a predator <strong>of</strong> the bagworm. In the 

laboratory it attacked the small larvae <strong>of</strong> M. corbetti and pupated but did not complete 

its development. 

OTHER INSECTS FOUND ASSOCIATED WITH M. CORBETTI 

On several occasions maggots <strong>of</strong> a phorid fly, Megaselia sp., were found feeding on 

dead larvae <strong>of</strong> M. corbetti at B.A.L. and Giram Estates. These were probably saprophytic. 

Two adults <strong>of</strong> the Hemerobiid Micromus timidus Hagen emerged in a cage 

with larvae <strong>of</strong> M. corbetti collected at Giram Estate. A full-grown nymph <strong>of</strong> an Asopine 

predator, Cantheconidea sp., was collected on an oil palm infested by M. corbetti 

and the nettle grub Setora nitens Walk, at Giram Estate. It fed on a larva <strong>of</strong> S. nitens 

in the laboratory and developed into an adult. The bug later attacked the larvae <strong>of</strong> 

M. corbetti and C. pendula which were provided for it. It fed on a larva <strong>of</strong> M. corbetti 

continuously for about 5 hours until the prey was sucked dry. 

Specimens <strong>of</strong> a bagworm species collected on oil palms were much smaller than and 

different in appearance from those <strong>of</strong> M. corbetti. They lacked the large pieces <strong>of</strong> oil 

palm leaves characteristic <strong>of</strong> the bags <strong>of</strong> M. corbetti. However, a single male moth 

which emerged from one <strong>of</strong> such bags collected at B.A.L. Estates, Tawau, was identified 

by the CLE. as Mahasena corbetti. Since M. corbetti has been reported to feed 

on other alternative host plants it seems likely that these unusual specimens had actually 

fed on some other (possibly wild) plants in the oil palm fields and then migrated 

to the palms. It would be interesting to obtain a series <strong>of</strong> specimens <strong>of</strong> male moths 

from such bags and compare them with M. corbetti to confirm their identification. The 

three species <strong>of</strong> hymenopterous parasites and a Tachinid obtained from larvae <strong>of</strong> this 

species are listed:


HYMENOPTERA 


A. psychidivorus: Empty and inhabited cocoons <strong>of</strong> this parasite were obtained from 

larval bags collected at Beaufort in April, May and July, at B.A.L. Estates and at 

Merotai in May and at Pamol Estate in June 1969. Parasitism was 17 % in a collection 

<strong>of</strong> 155 bags from Beaufort in May. In July 6 larvae in a small sample <strong>of</strong> 18 received 

from the same area were parasitised. Twelve females and 2 males <strong>of</strong> A. psychidivorus 

emerged from one <strong>of</strong> the bags and these adults fed on dilute honey lived up to 6 days 

in the laboratory. 

It is note-worthy that A. psychidivorus has not been obtained from the larval bags <strong>of</strong> 

M. corbetti which were covered by pieces <strong>of</strong> oil palm leaves as described earlier. 

Ichneumonidae 

Paraphylax sp.: A single specimen emerged from a larva in a collection <strong>of</strong> 42 bags 

from B.A.L. Estate. The Commonwealth Institute <strong>of</strong> Entomology has commented that 

this was "assuredly secondary." 

Eulophidae 

Pediobius lividiscutum (Gahan) : Two specimens were obtained from larvae collected 

at Beaufort. According to the CLE. this species has been reared from Artona catoxantha 

in the Malay Peninsula in 1924 and 1934 and the specimens were then misdetermined 

as P. detrimentosus Gahan in 1940, under the generic name Pleurotropis. 

DIPTERA 

Tachinidae 

Eozenillia ? equatorialis Townsend : A single specimen emerged from a larva collected 

at B.A.L. Estate in May 1969. 

The following parasites were recorded: 

HYMENOPTERA 


Metisa plana Walker 

Apanteles metesae Nixon: This was the only important parasite obtained during the 

period. In April 1969 a sample <strong>of</strong> 260 M. plana was collected at Mostyn Estate. About 

4 % <strong>of</strong> the bags contained Apanteles cocoons, most <strong>of</strong> which were empty. When the 

area was visited in August, the Metisa larval population had greatly increased during 

the next generation. The numbers <strong>of</strong> specimens collected in four fields and the percentages 

<strong>of</strong> parasitism (in brackets) were 168(25), 307 (18.6), 73 (71) and 110 (60), 

respectively. In September 13 larvae <strong>of</strong> 31 from B.A.L. Estate were parasitised. On 

a third visit to Mostyn Estate in November 18 larval bags <strong>of</strong> 73 from a different field 

contained cocoons <strong>of</strong> A. metesae. At Papar parasitism by A. metesae was 66 % in June 

1970. 

Eupelmidae 

Eupelmus catoxanthae Ferriere: This was reared from male pupae <strong>of</strong> M. plana but 

parasitism was negligible. E. catoxanthae has been recorded from the Limacodid Parasa 

lepida (Cram.) in Malaya (Lever, 1964).


DIPTERA 

Tachinidae 

Eozenillia psychidarum Bar.: In May and July 1969 puparia <strong>of</strong> this Tachinid were found 

in a few larval bags <strong>of</strong> M. plana collected at B.A.L. Estate. In June a puparium was 

found in a pupal cocoon <strong>of</strong> M. plana. Most <strong>of</strong> these puparia were empty or aborted. 

A parasite which did emerge was identified as E. psychidarum, which has been recorded 

from an unidentified Psychid in Sumatra (Thompson, 1946b). This is the first record 

<strong>of</strong> a Tachinid parasitising M. plana. 

HYPERPARASITES 

Three species <strong>of</strong> hyperparasites, E. catoxanthae, Pediobius lividiscutum and Brachymeria 

sp., were reared from cocoons <strong>of</strong> A. metesae collected at Mostyn Estate. Some 50 

Apanteles cocoons which were isolated from Metisa bags in August 1969 were kept under 

observation in the laboratory. Since they failed to produce any emergents they 

were dissected in November. They contained dead larvae and pupae and imperfect 

adults <strong>of</strong> Apanteles and <strong>of</strong> the first two species <strong>of</strong> hyperparasites. 

According to van der Vecht (1950) E. catoxanthae has been reared from pupae <strong>of</strong> 

Artona catoxantha, puparia <strong>of</strong> Ptychomyia remota and Cadurcia leefmansi and from pupae 

<strong>of</strong> Apanteles artonae in Indonesia. 

In April 1970 about 80 % <strong>of</strong> the Apanteles cocoons obtained at B.A.L. Estates were 

found to be attacked by the three hyperparasites. 

A Brachymeria sp. was obtained, on one occasion, from a puparium <strong>of</strong> E. psychidarum 

in a bag <strong>of</strong> M. plana. 

Cremastopsyche pendula Joannis 

This bagworm infests oil palms as well as Albizzia sp. and Delonix regia. 

PARASITES 

Two Braconids, A. psychidivorus, and A. metesae, were obtained from larval bags <strong>of</strong> 

C. pendula. The former occurred both as a solitary and as a gregarious parasite (up to 

4 cocoons per host were seen) while A. metesae was invariably a solitary species, its 

cocoon occupying the whole bag <strong>of</strong> the host larva. With both, the parasitised hosts 

were always young and medium-sized larvae. 

In May 1969 parasitism by A. psychidivorus was 12 % and that by A. metesae 2 % in 

a lot <strong>of</strong> 650 bags collected at Pamol Estate. At Papar Apanteles had parasitised 16 % 

<strong>of</strong> the <strong>bagworms</strong> on oil palms and 4 % on D. regia in June 1970. A. psychidivorus has 

been reared from C. pendula at Tuaran, while A. metesae has been reared from the same 

bagworm collected at Beaufort, B.A.L. Estates and Merotai. According to Nixon 

(1967) A. metesae has been recorded from Vietnam "ex. C. pendula ?" 

E. catoxanthae and Paraphylax rufipes Cam. (Ichneumonidae) were reared from C. 

pendula on D. regia at Papar. These together with Apanteles were obtained from 33.5 

% <strong>of</strong> the <strong>bagworms</strong> on oil palms in the same area in May 1970. It is possible that 

these two species are secondary parasites.


PREDATOR 

In June 1969 a female <strong>of</strong> a Sycanus sp. was found on an oil palm heavily infested 

by C. pendula at Pamol Estate. It readily attacked the larvae <strong>of</strong> C. pendula (but not 

those <strong>of</strong> M. corbetti) in the laboratory and lived for 12 days. Stationary bags <strong>of</strong> C. 

pendula (those containing moulting larvae and pupae which were immobile) were ignored 

by the predator and only moving larvae were attacked. A cluster <strong>of</strong> 48 eggs was laid 

by the bug on the lid <strong>of</strong> plastic rearing cage. Forty-one eggs hatched after 20 days. 

The tiny nymphs fed on freshly hatched larvae <strong>of</strong> another bagworm {? Pteroma sp.) but 

not on full-grown larvae <strong>of</strong> C. pendula which were <strong>of</strong>fered to them in glass tubes. 

Two Scelionid parasites, Gryon sp. and Telenomus sp., emerged from two clusters <strong>of</strong> 

eggs closely resembling those <strong>of</strong> Sycanus collected on oil palm leaves at Mostyn and 

Giram Estates, respectively. 

Clania tertia Tempi. 

An Ichneumonid larval parasite, Xanthopimpla pedator Fabr, and the Braconid A. 

psychidivorus were obtained from this bagworm in small numbers, the former only from 

material collected at Beaufort and the latter from other plantations as well. On one 

occasion a single specimen <strong>of</strong> a Xanthopimpla sp. (which appears to be different from 

pedator) emerged from a larva collected at Beaufort. This is similar to the species 

listed under M. corbetti. 

This bagworm species was found on guava at Tuaran, where E. equatorialis and a 

Brachymeria sp. were reared from 19 % <strong>of</strong> the larvae collected in July 1970. 

X. pedator is a solitary parasite. Parasitised larvae become dark in colour, their skin 

shrinking and gradually drying up from the hind end. Parasitism was about 6% in 

two samples <strong>of</strong> 124 and 67 bags collected in May and September 1969 respectively, and 

nearly ll % in another <strong>of</strong> 55 specimens collected in November. The adults emerged 

by severing the head capsules <strong>of</strong> the host larvae, which remained adhering to the hosts' 

bodies inside their bags. 

A. psychidivorus also attacks C. tertia infesting other host plants. In July 1969 a total 

<strong>of</strong> 142 individuals emerged from a single bag collected on a c<strong>of</strong>fee plant at Tuaran. 

E. equatorialis was reared from C. tertia. In June 1970 parasitism was 41 % on oil 

palms at Papar and ll % at Beaufort. 

Cryptothelea cardiophaga Westw. 

This bagworm was very common on oil palms in most <strong>of</strong> the localities surveyed. 

It also infested various other host plants, such as citrus, c<strong>of</strong>fee, cocoa, guava, mango, 

pepper, an unidentified ornamental palm, Annona muricata, Araucaria cunninghamii, Cassia 

sp., Grevillea sp., Hibiscus sp., Lagerstroemia indica, Nephelium lappaceum, Peltophorum 

pterocarpum, Samanea saman and Thuja orientalis. 

A. psychidivorus, Sericopimpla flavobalteata Cam. and P. rufipes (most probably secondary) 

were reared from C. cardiophaga collected on oil palms. Parasitism by the last 

two species was 30.5 % at B.A.L. Estates in August 1970.


? Pteroma sp. 

Larvae <strong>of</strong> this species collected on oil palms at Tuaran were found parasitised by 

a Bethylid, Parasierola ( = Perisierola) sp. and a Brachymeria sp. The former is gregarious ; 

up to 7 parasites per host have been recorded. It has also been reared from larvae collected 

at Beaufort, Pamol and Sabah Palm Estates. Parasitism was \A% in a collection <strong>of</strong> 

166 bags taken at the last two estates in June 1969. On one occasion Pediobius lividiscutum 

was reared from this bagworm at Tuaran. 

Stephens (1962) has recorded a Perisierola sp. attacking small larvae <strong>of</strong> Oiketicus kirbyi 

Guild, in Costa Rica. 

A Tachinid puparium was obtained once from a bag collected at Tuaran in May but 

this actually harboured a Brachymeria sp. 

UNIDENTIFIED BAGWORM 

E. equatorialis and a Palexorista sp. were reared from two specimens <strong>of</strong> this species 

collected on oil palms at Ulu Dusun and B.A.L. Estate in May and June 1969 respectively. 

C. tenia 

PARASITES OF BAGWORMS COLLECTED ON 

OTHER HOST PLANTS 

On Annona muricata: A. psychidivorus; on c<strong>of</strong>fee: A. psychidivorus and Paraphylax (= 

Aphadnus) rufipes (Hym., Ichn.), all from Tuaran. P. rufipes appears to be a secondary 

parasite. 

C. cardiophaga 

On citrus: Sericopimpla flavobalteata, Goryphus sp. and Paraphylax sp. Once, a Brachymeria 

sp. emerged in a cage with this bagworm. On cocoa and c<strong>of</strong>fee: P. rufipes 

(most probably secondary), all from Tuaran. 

? Pteroma sp. 

On paddy: Parasierola sp.; on c<strong>of</strong>fee: Goryphus sp., both from Tuaran. 

? Acanthopsyche sp. 

On tea: Paraphylax sp., Kundasang. 

Eumeta sp. 

On cocoa: FishStina sp.. Tuaran 

? Eurukuttarus sp. 

On cocoa: Pediobius lividiscutum, Tuaran. 

Unidentified <strong>bagworms</strong> 

On Acacia auriculiformis: Eupelmus catoxanthae and Brachymeria sp., B.A.L. Estates. 

On cashew-nut tree: A. psychidivorus, Tuaran. 

On guava: A. psychidivorus, Tuaran.


PATHOGENS ASSOCIATED WITH OIL 

PALM BAGWORMS 

In March 1969 about 120 bags <strong>of</strong> M. corbetti were collected at Sg. Manila. The majority 

<strong>of</strong> these contained dead larvae. Since these were suspected <strong>of</strong> having died <strong>of</strong> 

some disease, specimens preserved in distilled water and Dubosq Brasil Bouin's fluid 

were sent to the Insect Pathology Unit, Oxford. The preliminary diagnosis made by 

IPU indicated that there was no evidence <strong>of</strong> any virus infection but that there were 

many microorganisms in the specimens. 

In April specimens <strong>of</strong> M. plana collected at Mostyn Estate developed a fungal infection 

and died in the laboratory at Tuaran. Some specimens <strong>of</strong> C. pendula collected 

at B.A.L. Estate also showed the same fungus. In June several specimens <strong>of</strong> C. pendula 

were found to be attacked by this fungus at Pamol and Sabah Palm Estates near 

Sandakan. Laboratory colonies <strong>of</strong> two other bagworm species, C. tenia and C. cardiophaga, 

originating from the Klias Scheme at Beaufort and B.A.L. Estates, respectively, 

showed symptoms <strong>of</strong> the fungal disease and eventually died. Specimens <strong>of</strong> the three 

last species were sent to Dr. E. Mtiller-Kogler, Institut fiir Biologische Schadlingsbekampfung, 

Darmstadt, Germany. He placed the infected larvae in moist chambers, allowed 

the fungus to develop and then successfully isolated it and built up a culture. It was 

reported to grow well on malt extract (2-3^), peptone (0.5^), agar (2-3%). The 

fungus was identified by him to be a Paecilomyces sp. Later, Dr Agnes H. S. 

Onions, Commonwealth Mycological Institute, Kew, examined the fungus and identified 

it specifically as Paecilomyces fumoso-roseus (Wize) Brown and Smith. A culture <strong>of</strong> this 

fungus is kept at Kew under the designation "IMI 143085" (E. Mtiller-Kogler, personal 

communication). 

P. fumoso-roseus was originally found by Wize on larvae <strong>of</strong> Cleonus punctiventris Germ. 

in the Ukraine and its other known insect hosts include Hylemyia floralis Fall., H. 

brassicae Beh. and Melolontha melolontha L. This fungus is <strong>of</strong> widespread occurrence, 

chiefly as a parasite <strong>of</strong> insects but also capable <strong>of</strong> growing on other substrata. Isaria 

fumoso-rosea Wize and Spicaria fumoso-rosea (Wize) Vassiljevski are synonyms <strong>of</strong> P. 

fumoso-roseus (Wize) Brown and Smith (Brown and Smith, 1957). The fungus appears 

to exist in different strains. 

Investigations by Sekiguchi (1959) in Japan have shown that P. fumoso-roseus is suitable 

for use in the biological control <strong>of</strong> the peach fruit moth Carposina sasakii Mats. 

(niponensis Wl sm.) 

In November a large number <strong>of</strong> dead larvae and also some dead pupae <strong>of</strong> M. corbetti 

were examined at Mostyn Estate. Some <strong>of</strong> them had a blackened skin and showed 

signs <strong>of</strong> liquefying internally, emitting a foul smell. These were sent to the Insect 

Pathology Unit which later reported in its preliminary diagnosis that the specimens 

showed a "possible non-occluded virus." At the time <strong>of</strong> collection the bags had become 

very wet owing to heavy rains in the Tawau area. 

In laboratory cages young larvae <strong>of</strong> M. corbetti started making their bags with small 

pieces <strong>of</strong> dried oil palm leaves from their mother's bag. Most <strong>of</strong> the larvae which 

used the green leaves for bag construction were attacked by a fungus, Aspergillus sp.

1972 Sanakran and Syed: The <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> <strong>bagworms</strong> on oil palms 67 

DISCUSSION 

The present investigations have shown the existence <strong>of</strong> several <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> 

<strong>bagworms</strong> infesting oil palms and certain other economic plants in Sabah. However, 

the native parasites <strong>of</strong> the three major oil palm bagworm species, M. corbetti, M. plana 

and C. pendula, are not able to prevent the serious outbreaks <strong>of</strong> these three pests which 

have occurred since 1966 following a rapid increase in the area under oil palms. While 

Apanteles metesae parasitises a high percentage <strong>of</strong> the larvae <strong>of</strong> M. plana in some fields 

it is attacked by at least two important species <strong>of</strong> hyperparasites which reduce its 

efficiency. 

There is a striking similarity between the <strong>natural</strong> enemy complexes associated with 

these <strong>bagworms</strong> in West Malaysia and East Malaysia. Although the two areas are 

separated by a vast stretch <strong>of</strong> the South China Sea most <strong>of</strong> the oil palm pests and 

their <strong>natural</strong> <strong>enemies</strong> are common to them. This is because the insect pests <strong>of</strong> oil 

palms in the entire Malay Archipelago are all native species which already existed on 

other palms and several other alternative food-plants prior to the introduction <strong>of</strong> the 

oil palm. They have gradually transferred to oil palms over the last three decades 

or so. The <strong>bagworms</strong> have also attracted to their new host plant (oil palm) a number 

<strong>of</strong> their own parasites and predators. 

A. metesae is the commonest parasite <strong>of</strong> M. plana on oil palms in West Malaysia. 

Other, less important parasites include: Brachymeria sp., Elasmus sp. nr. hutsoni Ferr., 

Eupelmus catoxanthae, Fislistina sp., Goryphus inferus Szepl., Lienella varius Walk., Pediobius 

anomalus Gah., P. detrimentosus Gah. and Syntomosphyrum zygaenarum var. nigricoxis 

Ferr. Parasites <strong>of</strong> C. pendula include: Apanteles sp. nr. hasorae Wilk., Aulosaphes psychidivorus, 

Brachymeria sp., E. catoxanthae, Eurytoma sp., Goryphus sp. nr. inferus Szep., 

G. sp. ? varibalteatus Cam., Genus near Paraphylax, Paraphylax rufipes, P. anomalus, P. 

detrimentosus and Tetrastichus sp. (Wood, 1966 and 1968; C.I.B.C, unpublished records). 

In January 1962 Rao (1962) recorded 14 % parasitism <strong>of</strong> C. pendula by Apanteles sp. (later 

identified as metesae by Nixon) at Layang Layang, Johore. 

In West Malaysia, M. corbetti is parasitised by Brachymeria euploeae Westw., Sarcophaga 

antilope, S. peregrina, Palexorista inconspicuoides (Bar.) (^Sturmia inconspicuoides 

Bar.), Thrycolyga aberrans Strobl., T. psychidarum Bar., T. sorbillans Wied, and Nealsomyia 

rufella (Bezzi) (=Zenillia quadrimaculata Bar.) (Thompson, 1946a). There is no 

information as to the extent to which these parasites are responsible for keeping the 

<strong>bagworms</strong> under <strong>natural</strong> control. 

Tricholyga (=Thrycolygd) psychidarum Bar. is now treated as a synonym <strong>of</strong> Eozenillia 

psychidarum (Bar.) (Crosskey, 1967a) which occurs in Sabah. In addition, a Sarcophaga 

sp. and the Tachinids E. equatorialis and P. solennis occur here. Three Brachymeria 

spp. are hyperparasitic on Eozenillia and limit the capacity <strong>of</strong> this Tachinid to increase 

its population. B. euploeae in West Malaysia may also behave as a secondary parasite. 

Although this species has not been bred from M. corbetti it has been recorded from 

Erionota thrax L. in Sabah. Therefore, none <strong>of</strong> the parasites <strong>of</strong> M. corbetti known at 

present from West Malaysia is considered suitable for introduction here. 

A. metesae is an important parasite <strong>of</strong> M. plana in both Sabah and West Malaysia. 

Similarly, both A. metesae and A. psychidivorus attack C. pendula in the two areas. Many 

<strong>of</strong> the other West Malaysian parasites <strong>of</strong> these two <strong>bagworms</strong> are known to occur as


hyperparasites, not only <strong>of</strong> these <strong>bagworms</strong> but also <strong>of</strong> other lepidopterous pests. These 

are: Brachymeria sp., E. albotibialis, E. catoxanthae, G. infer us, G. varibalteatus, P. rufipes, 

P. detrimentosus and S. zygaenarum. It would, therefore, be undesirable to introduce 

them into Sabah even if some <strong>of</strong> them definitely do not occur here. E. catoxanthae 

and P. rufipes have been recorded from Sabah during the present study. Further work 

may show that the other species are also present here. 

The Reduviid predator S. dichotomus and the Clerid C. arcufer are important general 

predators <strong>of</strong> <strong>bagworms</strong> in West Malaysia. Eggs <strong>of</strong> C. arcufer are parasitised by a Mymarid, 

Cleruchus sp. (Wood, 1966). S. dichotomus is already present in Sabah, according 

to information obtained from the Commonwealth Institute <strong>of</strong> Entomology, London. It 

was not seen in any <strong>of</strong> the plantations surveyed. However, another Sycanus sp. attacks 

C. pendula. Two egg-parasites <strong>of</strong> Reduviids Gryon sp. and Telenomus sp., occur in Sabah 

and these and possibly other parasites appear to be, at least partly, responsible for the 

scaricity <strong>of</strong> Reduviids in the oil palm plantations here. Sycanus spp. are indiscriminate 

feeders, attacking parasitised as well as healthy larvae (Vayssiere, 1958). The Clerid 

predator C. arcufer is known to attack the Tachinid Ptychomyia remota in West 

Malaysia (Corbett and Miller, 1933). Its introduction into Sabah is not recommended 

since it may attack the Tachinid parasites <strong>of</strong> M. corbetti. 

Since M. corbetti, M. plana and C. pendula are not known to occur outside the Southeast 

Asian region any additional parasites which are specific to them and intended for 

introduction into Sabah have to come from other areas in this region. However, a 

comparison <strong>of</strong> the known <strong>natural</strong> <strong>enemies</strong> <strong>of</strong> these <strong>bagworms</strong> in different parts <strong>of</strong> the 

Malay Archipelago reveals a more or less common complex. Therefore, it would be 

desirable to screen the parasites <strong>of</strong> other <strong>bagworms</strong> elsewhere in the world, choosing 

the species, as far as possible, from areas where the climate is predominantly tropical 

and comparable to that <strong>of</strong> Sabah. 

Since there are a very large number <strong>of</strong> hymenopterous and some dipterous parasites 

<strong>of</strong> Oiketicus spp. in South America (Thompson, 1946b and C.I.B.C, card files) it will 

be worthwhile testing the more readily available species on M. corbetti, M. plana and 

C. pendula. The hymenopterous parasites should receive prior attention since no effective 

hymenopterous larval parasites <strong>of</strong> M. corbetti are known from Sabah. Parasites <strong>of</strong> the 

ever-green bagworm Thyridopteryx ephemeraeformis (Haw.) in the U. S. A. might also 

be tested. Since Oiketicus spp. and T. ephemeraeformis are polyphagous the possibilities 

<strong>of</strong> some <strong>of</strong> their parasites exhibiting a "host-plant relationship" have to be borne in 

mind. In this connection it may be noted that many <strong>of</strong> the native parasites <strong>of</strong> M. 

corbetti, M. plana and C. pendula are themselves polyphagous, attacking other <strong>bagworms</strong> 

and various unrelated Lepidoptera. For example, Sarcophaga antilope and Eupelmus 

catoxanthae attack Parasa lepida (Cram.) in Malaysia (Lever, 1964). Goryphus varibalteatus 

Cam. attacks the nettle-grub Natada nararia Moore in Ceylon (Austin 1932) and 

G. inferus Szep. is a common parasite <strong>of</strong> Artona catoxantha in Java (van der Vecht, 

1950). Sarcophaga peregrina R.-D. is a larval parasite <strong>of</strong> Dendrolimus punctatus Walk. 

in China (Sun and Liu, 1958). S. peregrina and Exorista sorbillans have been recorded 

from Mocis frugalis F. in Australia (Mungomery, 1947). E. sorbillans parasitises the bagworm 

Deborrea malagassa Heyl. in Malagasy (Breniere and Dubois, 1965). Echthromorpha 

agrestoria attacks Clania sp., Cryptothelea minuscula Btlr., Hyblaea puera Cramer 

and D. punctatus (Townes, Townes and Gupta, 1961).


Most <strong>of</strong> the hymenopterous parasites <strong>of</strong> Oiketicus spp. in South America have no 

other known hosts. These parasites include Apanteles oeceticola Blanch., Balcarcia 

brethesi Blanch., Cryptus borsani Blanch., Cryptopteryx oeceticola Blanch., Iphiaulax spp., 

Pimpla (Itoplectis) psychidophagus Blanch, and Psychidosmicra spp. Pimpla tomyris Blanch. 

is polyphagous, Pimpla (Itoplectis) oeceticola (Brethes) has been recorded from the pine 

shoot borer Rhyacionia (Evetria) buoliana (Schiff.) in Uruguay (Millan & De Santis, 

1958). Spilochalcis spp. and Tetrastichus pseudoeceticola Blanch, are known to be 

hyperparasites though they may also be primary. Only those species that are entirely 

primary should be tested on the oil palm <strong>bagworms</strong>. 

Oiketicus kirbyi Guild, is a minor pest <strong>of</strong> cacao in Trinidad (Kirkpatrick, 1953). Work 

at the C.I.B.C. West Indian Station during 1970 has already shown that eight species <strong>of</strong> 

parasites and a nuclear polyhedrosis virus disease <strong>of</strong> <strong>bagworms</strong> occur in Trinidad 

(Cruttwell, 1970). 

Many <strong>of</strong> the hymenopterous primary parasites <strong>of</strong> the evergreen bagworm T. ephemeraeformis 

in the U. S. A. are also polyphagous. In a study conducted in October I960 the 

polyphagous Itoplectis conquisitor (Say) was found to be the most important parasite 

<strong>of</strong> the ever-green bagworm. Dipterous parasites were only <strong>of</strong> minor importance (Kulman, 

1965). However more recent work by Pr<strong>of</strong>. Kulman has led him to suspect that 

/. conquisitor may be a hyperparasite more <strong>of</strong>ten than is indicated in the literature (H. 

M. Kulman, pers. com.). Therefore, it would not be desirable to introduce it into 

Malaysia. Work on another Ichneumonid, Sericopimpla sericata (Kchrb.), in South Africa 

has shown that the initial attraction <strong>of</strong> the parasite to its bagworm host Kotochalia 

junodi (Heyl.) seems to be the bag rather than the host larva. The adults, particularly 

the females, are not restricted to wattle trees or other Acacia spp. on which the hosts 

occur (Smithers, 1956). It is worth investigating whether S. sericata would parasitise 

the <strong>bagworms</strong> on oil palms. 

The size <strong>of</strong> the host larva (enclosed in the bag, which itself varies in size and 

structure according to the food-plant) is a critical factor in host selection with some 

<strong>of</strong> the bagworm parasites. A. metesae parasitises only the young larvae <strong>of</strong> M. plana. 

According to Stephens (1962), Iphiaulax sp. attacks all sizes <strong>of</strong> larvae <strong>of</strong> O. kirbyi 

while Psychidosmicra sp. attacks only small larvae. Sarcophaga (Sarcodexia) lambens 

Wied, and Achaetoneura sp. attack medium and large larvae. Griot and Icart (1947) 

have studied the bionomics <strong>of</strong> Psychidosmicra brethesi Blanch., P. brasiliensis (Brethes) 

and P. australis Blanch. These parasites do not attack larvae in bags which are larger 

than 1 cm. These and possibly other parasites <strong>of</strong> Oiketicus which prefer the young 

larval stages <strong>of</strong> the host may prove useful against the early stages <strong>of</strong> M. corbetti, M. 

plana and C. pendula in Sabah. In central Argentina P. brasiliensis has been reported 

to give as much as 80 % parasitism <strong>of</strong> the larvae <strong>of</strong> O. kirbyi in some places (Griot 

and Icart, 1947). 

Miller (1970) has stated that many species <strong>of</strong> the Eulophid Sympiesis are "nichespecific" 

and that any trends in their habits towards host-specificity are attributable to 

the fact that only certain species <strong>of</strong> insects occupy the niches sought by the parasites. 

S. stigmata Girault seeks a niche usually occupied by lepidopterous case-bearing insects. 

A similar habit may account for the occurrence <strong>of</strong> a Sympiesis sp. as a parasite <strong>of</strong> M. 

corbetti in Sabah.


Mahasena theivora (Dudg.) and Metisa plana are minor pests <strong>of</strong> tea in north-east India 

(Das, 1956). Their <strong>natural</strong> <strong>enemies</strong>, if any, have not been investigated. Mahasena 

graminivora Hmps. is an unimportant pest <strong>of</strong> paddy in north-east India. A Paraphylax sp. 

and a Lienella sp. have been reared from this bagworm. 

M. corbetti, C. pendula, C. terita and Pteroma spp. have been recorded from New 

Guinea. As suggested by Simmonds (1971) a survey for parasites <strong>of</strong> these and other 

<strong>bagworms</strong> in this area will be <strong>of</strong> value both to Malaysia and to New Guinea. 

The entomogenous fungus /. psychidae and the virus (Borrelinavirus kotochaliae Ossowski) 

attacking the wattle bagworm in South Africa are also worth testing for their 

host specificity. If they prove to be pathogenic to M. corbetti, M. plana and C. pendula 

(or to any <strong>of</strong> these species) they could be introduced into both East and West Malaysia. 

Acknowledgments: The authors are grateful to the Director <strong>of</strong> Agriculture, Sabah, 

for the various facilities provided to them at the Agricultural Research Centre, Tuaran, 

for carrying out these investigations. Their thanks are due to the Director, Commonwealth 

Institute <strong>of</strong> Entomology, London for identifying the insects mentioned in 

this paper, and to the Deputy Head, Insect Pathology Unit, Commonwealth Forestry 

Institute, Oxford and to Dr E. Muller-Kogler, Institut fiir Biologische Schadlingsbekampfung, 

Darmstadt, Germany for identifying the insect pathogens. The Managers 

<strong>of</strong> the different oil palm plantations which were visited by the authors were very helpful 

and hospitable to them. Their co-operation in this work is greatly appreciated. 

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