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Occasional Publication Series No. 2
BIOLOGICAL CONTROL OF OPIS/NA
ARE.NOSE.LLA W ALKER / LEPIDOPTERA,
OECOPHORIDAE
by
M. J.
w. Cock
(CAB International Institute of Biological Control,
Kenya Station, P.O. Box 30148, Nairobi, Kenya)
and
P. A. C. R. Perera
(Coconut Research Institute, Lunuwila, Sri Lanka)
Editor: R. Mahindapala Ph D.
Coyer photograph: A coconut palm heavily ïnfested with the Coconut Caterpillar.
A rare instance where the nut is also infested. (Credit - R. Mahindapala)
This article was first published in Biocontrol News and Information (Volume 8 No. 4,
1987). It is reproduced with the kind courtesy of the Commonwealth Agricultural
Bureaux International, England.
Published by the
Coconut Research Institute of Sri Lanka
Joly, 1988
CONTENTS
1.
Introduction
ll.
Natural enemies
1.
2.
3.
Page
01
02
Pathogens
02
(a)
(b)
(c)
(d)
02
03
03
03
Viruses
Bacteria
Protozoa
Fungi
Predators
03
(a)
(b)
(c)
03
03
04
Anthocoridae
Reduviidae
Carabidae
Parasitoids
Anastatoidea brachartonae Gahan (Eupelmidae)
Apanteles taragamae Viereck (Braconidae)
Brachymeria excarinata Gahan (Chalcididae)
Brachymeria hime attevae Joseph, Narendran & Joy (Chalcididae)
Brachymeria lasus (Walker) (Chalcididae)
(f)
Brachymeria nephantidis Gahan (Chalcididae)
(g) Brachymeria nosatoi Habu (Chalcididae)
(h) Bracon brevicornis Wesmael (Braconidae)
(i)
Bracon serinopae (Cherian) (Braconidae)
(j)
Elasmus nephantidis Rohwer (Elasmidae)
(k) Eriborus trochanteratus (Morley) Ichneumonidae)
(1)
Eurytoma spp. (Eurytomidae)
(m) Fornicia sp. (Braconidae)
(n) Goniozus nephantidis (Muesebeck) Bethylidae)
(0) Goryphus sp. (Ichneumonidae)
(p) Megaselia sp. (Phoridae)
(q) Meteoridea hutsoni (Nixon) (Braconidae)
(r) Pyemotes ventricosus (Newport) (Acarina)
(s) Stomatoceras sulcatiscutellum Girault (Chalcididae)
(t) Stomatomyia bezziana Baranoff (Tachinidae)
(u) Tetrastichus israeli (Mani & Kurian) (Eulophidae)
(v) Thelairodrino gracilis Mesnil (Tachinidae)
(w) Trichospilus pupivorus Ferriere (Eulophidae)
(a)
(b)
(c)
(d)
(e)
04
04
05
05
05
05
06
06
07
07
07
08
08
08
10
10
10
10
10
11
Il
12
12
(x)
Xanthopimpla spp. (Ichneumonidae)
13
(y)
(z)
Other pupal parasitoids
Other probable hyperparasitoids.
13
13
Ill.
IV.
Classical biological control - historical
13
(a)
Antrocephalus pandens (Walker) releases in Sri Lanka
14
(b)
Bessa remota (Aldrich) releases in India and Sri Lanka
14
(e)
(cl)
(e)
(f)
(g)
(h)
(i)
Bracon brevicornis releases in Sri Lanka
E/asmus nephantidis releases in Sri Lanka
Eriborus trochanteratus releases in India
Stomatomyia bezziana releases in India
Tetrastichus israeli releases in Sri Lanka
Trichogramma brasi/iensis (Ashmead) releases in Sri Lanka
Trichogramma minutum Riley releases in India and Sri Lanka
15
15
15
16
16
16
17
Classical biological control - possibilities
17
1.
Natural enernies of Opisina arenosel/a
17
2.
Other hosts as sources for natural enemies
18
(a)
(b)
(c)
(d)
(e)
(f)
(g)
(h)
(i)
U)
(k)
18
22
22
24
24
24
25
26
27
27
27
27
(1)
Agonoxena spp. (Agonoxenidae)
Antaeotricha spp. (Stenomidae)
Artona catoxantha (Hampson) Zygaenidae)
Durrantia arcane//a (Busek) (Oeeophoridae)
Furcivena rhodoneurialis Hampson (Pyralidae)
Hedy/epta b/ackburni (Butler) (Pyralidae)
Herculia nigrivitta (Walker) (Pyralidae)
Homophy/otis catori Jordan (Zygaenidae)
Loxotoma e/egans Zeller (Stenomidae)
?Odites sp. (Xylorietidae)
Stenoma impressella (Busek) (= S. cecropia Meyrick) (Stenomidae)
Struthocelis semiotarsa Meyriek (Oeeophoridae)
V.
Discussion
27
VI.
Recommendations
30
Acknowledgements
31
References
31
ABSTRACT
The natural enemies of Opisina arenosella Walker are reviewed and their use in classical
biological control summarized. Further use of the known natural enemies of O. arenosella
is not considered promising. The following species of small Lepidoptera whose larvae feed
upon palm leafl.ets in concealed manner are considered as possible sources for natural
enemies to be used against O. arenosella: Agonoxena spp., Antaeotricha spp., Artona
catoxantha (Hampson), Durrantia arcanella (Busck), Furcivena rhodoneurialis Hampson,
Hedylepta blackburni (Butler), Herculia nigrivitta (Walker), Homophylotis catori Jordon
(Loxotoma elegans Zeller. Odïtessp., Stenoma impressella Busck and Struthocelis semiotarsa)
Meyrick. Their natural enemies are reviewed in light of what is known of the population
dynamics and causes of outbreaks of Opisina arenosella and climates of the source and
target areas. It is concluded that Argyrophylax fumipennis (Townsend), a tachinid
parasitoid of Artona catoxantha in Java, particularly merits investigation as a possible
biological control agent of O. arenosella.
J. INTRODUCTION
Opisina arenosella Walker, the black headed caterpillar of coconut, is the most damaging
defoliator of coconut in India and Sri Lanka. The report of Davis & Sudasrip (1982)
of this species in Indonesia is erroneous; it is found in Sri Lanka, South India, Bangladesh
and Burma (CIE 1966).
Until 1981 this pest was known as Nephantis serinopa Meyrick, and there is a very
extensive literature under that name (see reviews by Jayaratnam 1941a; Mohammed et al.
1982b; Nirula 1956a, 1956b; Ramachandran et al. 1979, etc.). In 1981, however, Becker
(1981) examined some F. Walker types from 'unknown lands' and found N. serinopa to
be synonymous with the older Opisina arenosella Walker. In view of the many published
works on 'Nephantis serinopa' there would seem to have been a good case for suppressing
the name 'Opisina arenosella' in the interests of stabilizing the literature. However, the
opportunity to do so has passed, and the name O. arenosella is now in use in the literature
and should be taken as correct.
O. arenosella has only been prominent as a pest in Sri Lanka and India since about
1920. In severe outbreaks thousands of palms are afiected and all the fronds, except the
3 or 4 youngest are killed. In the following year palm production may be halved or
worse, due to Ieduced production of flower spikes, increase in premature nutfalI,
constriction of the trunk and retardation of growth (Lever 1969). Other studies also
clearly show the marked effect of defoliation - whether by O. arenosella, Artona
catoxantha (Hampson), limacodids or artificial simulation - upon subsequent performance
by the palm (Wood 1987).
Coconut is the normal foodplant of O. arenosella, although Butani (1975) records
it as a minor pest of date palm. Manjunath (1985) records that during an outbreak in
Andhra Pradesh, larvae fed on banana. From laboratory studies, Talati & Kapadia
(1984) conclude that fan palm (Livistona chinensis), wild date palin (Phoenix sylvestris)
and date palm (P. dactylifera) are suitable foodplants, but banana is not. Dharmaraju
(1963) lists palm hosts in Sri Lanka, which include palmyra palms (Borassus) (Hutson 1922),
and Lever (1969) lists Borassus, Corypha, Hyphaene, Phoenix and Roystonea.
Lever (1969) has summarized the bio10gy of O. arenosella. The eggs are laid on
the distal undersurface of the leaflets of the older fronds, and hatch in 3-5 days. The
larvae make galleries or runways on the leaflet undersurface, comprising silk and
incorporating the larval frass, in which they feed. The five larval instars are completed
in 5-8 weeks and the moth emerges from the pupa after a further 9-14 days. On average
140 eggs are laid. The complete life cycle takes 2-2.5 months.
O. arenosella is an outbreak pest, which sporadically occurs in large numbers locally
to cause severe defoliation to coconut palms. The mechanism of outbreak initiation may
weIl be mediated through the effects of external factors upon the natural enemies of O.
arenosella (cf. Section II) or even through the intrinsic dynamjç interaction of the pest
and its natural enemies, which acts to synchronize the pest population and render the
natural enemies ineffective (Godfray & Hassell 1987). There have been attempts to
correlate meteorological factors with O. arenosella outbreaks. Thus, in 1974-75 Nadarajan
& Channa Basavanna (1980) studied O. arenosella at Bangalore and Mangalore, India.
Although oviposition peaked from November to March no climatic, effects were noted
in the equable climate at Bangalore, although high summer temperatures at Mangalore
ｾｲ･ｷ
thought to affect natural enemies adversely, 1eading to pest population increases.
Sathiamma et al. (1974) monitored O. arenosella in Kerala from 1967 to 1969. They
correlated population increase with high humidity, and decrease with high temperatures,
much sunshine and heavy rain.
Attempts to control this pest have been mostly cultural or biological. One early
method was to remove infested leaves and leave them on the ground for three weeks to
kill the larvae white preserving the parasitoids (Anon. 1918). Subsequently, growers were
recommended to bum infected fronds in Sri Lanka (Anon. 1920; Hutson 1920, 1921a,
1922, 1933; Jardine et al. 1925; Jebaratnam 1926), although this was qualified by De Mel
(1927) to early in an outbreak (parasitoids being considered effective later in an outbreak),
in India (Anstead 1926; Madhvan Pillai, 1919; Pillai 1921, 1922, 1923, 1924, 1926 and
in Burma (Ghosh 1924) Peris (1926) suggested that the removal offronds causes too much
damage to the palms. He also reported that placing the fronds in gauze cage, allows the
parasitoids to escape whi1e retaining the moths and that this technique has been successfully
used. Eally recommendations for the use oflight traps (Anon. 1920; Hutson 1920, 1921a,
1922) were proclaimed under the Plant Pest Ord nanee in Sri Lanka (Hutson 1921b), but
De Mel (1927) stated that they are of little value.
Augmentative releases of laboratory cultured parasitoids have played a very
substantial role in attempts to control O. arenosella in both Sri Lanka and India. Species
released include Bracon brevicornis Wesmael, Elasmus nephantidis Rohwer, Eriborus
trochanteratus (Morley), Goniozu'i nephantidis(Muesebeck), Stomatomyia bezziana Baranoff,
Tetrastichus israeli (Mani & Kurian) and Trichospilus pupivorus Ferriere.
Some of these species; when released in Sri Lanka, were effectively classical
biological control releases and are treated in Section III below. As part of the continuing
programme of collaboration between the Silwood Centre for Pest Management (SCPM)
and the Coconut Research Institute of Sri Lanka (CRI), it is intended that a critical
evaluation of the augmentative releases will be made. In this review, however, the
emphasis is plaeed upon the natural enemies and the possibilities for classical biological
control.
II. NATURAL ENEMIES
There is a large literature on the natural enemies of O. arenosella. However,
much of this comprises short notes, with little depth of investigation, so that there are still
many gaps in our knowledge of the biology, distribution, and more especially frequency
and importance of these natural enemies. Dharmaraju (1962) pIovides a check list of
natural enemies, but we are aware of no attempt to bring together the scattered literature
on this subject. We try to do so below.
1.
Pathogens
(a)
Viruses
There are very few reports of virus epizootics of O. arenosella. However,
Philip et al. (1982 recorded widespread larval mortality due to a nuclear
polyhedrosis virus in April 1981 at VaUayani, Kerala. This virus merits more
investigation, and could prove an effective means of terminating outbreaks
if applied in the field.
2
(b)
Bacteria
In laboratory trials, Serratia marcescens Bizio and Bacillus thuringiensis
Berliner produced 80% and 20% larval mortality of O. arenosella respectively
(Muthukrishnan & Rangarajan 1974).
(c)
Protozoa
Eugregarines occur mostly as commensals in the guts of insects, and there is
no reason to believe that the?Leidyana sp. reported by Rabindra (1981) is
acting otherwise.
(d)
Fungi
Paecilomyces farinosus (Fries) Brown & Smith is known to have a wide range
of insect hosts. In laboratory tests, Kuruvilla & Jacob (1980) demonstrated
that it would infect O. arenosella. Again in laboratory trials Muthukrishnan
& Rangarajan (1974 and Oblisami et al. (1969) showed Aspèrgillusflavus Link
could cause 90% mortality of O. arenosella larvae. Nirula (I956a, 1956b)
reports that fiooding of the larval galleries during the southwest monsoon
encourages mortality due to fungi.
2.
Predators
Records of predators of O. arenosella are relatively infrequent in the literature. Perera
(unpublished) notes that crows (Corvus splendens) are common predators in Sri
Lanka, and doubtless other common insect-eating birds play a role, Spiders, which
are often important predators of insect pests in the tropics (Reichert & Lockley
1984), are scalcely mentioned (e.g. Dharmaraju 1963), but Perera (unpublished)
found Rhene sp., Hyllus sp. and Plexippus sp. to he common predators of
O. arenosella in Sri Lanka. A handful of predatory bugs and beetles have been
reported to attack the larvae and pupae. General predators, e.g. the asilid
Astochia sp. (Perera unpublished), attack the adults. No attempt has been made
to assess the field incidence or field predation rates of predators. Sorne preliminary
work would be useful to try and assess the importance of these polyphagous
p;edators of O. arenose/la.
(a)
Anthocoridae
Cardiostethus sp. is a predator of the eggs and early instars of O. arenosella
(Abdurahiman et al. 1982). It will also feed on the exoparasitoids Bracon
brevicornis and Goniozus nephantidis It has been recorded in this role from
Kerala. The Triphleps (= Orius) sp. recorded by Lever (1969) may refer to
the same or another species.
(b)
Reduviidae
Nirula (l956a) records that Sphedanolestes aurescens Distant feeds upon the
eggs and larvae of O. arenosella in India.
3
(c)
Carabidae
Two species of Carabidae are recorded as predators of O. arenosella: Parena
nigrolineata (Chaudoir) (=latecincta Hates) and Calleida splendidula CF.)
Details of the biology of the former are given by David et al. (1975) and
Mohamed et al. (1982), and of the latter by Mohamed et al. (1982). Both
are recorded from India, but Dharmaraju (1963) records only P. . nigrolineata
from Sri Lanka.
3.
Parasitoids
We attempt to summarize here the literature on the large and varied fauna
of parasitoids associated with O. arenosella. Because O. arenosella has been so
extensively and intensively studied in India and Sri Lanka, there are several records
of parasitoids which are best regarded as generalist species attacking O. arenosella
opportunistically when and where it is commonly available. However, there remains
a core of specialized species which do seem to be closely associated with O. arenosella,
and emphasis is given to these below.
Given this association of common, specialized parasitoids, one might expect
an effective natural control to prevail (Cock 1986), and on the whole this is so .This
implies that the periodic, often seasonal, outbreaks of O. arenosella oceur when
this natural control breaks down for some reason, and this is diseussed further below.
(a)
Anastatoidea brachartonae Gahan (Eupelmidae).
There is just a single record of this species reared from a field collected pupa
of O. arenosella in Kayamkulam, Kerala State, India (Joy & Joseph 1976).
In West Malaysia this species is a hyperparasitoid of Artona catoxanthae
(Hampson), through Hymenoptera and Tachinidae (Ferriere 1940; Table 7
below). Most probably it was aiso acting as a hyperparasitoid in India,
perhaps through a Xanthopimpla sp.
(b)
Apanteles taragamae Viereck (Braconidae)
A. taragamae was originally described from Bangalore, India, but occurs from
India through Southern Asia to Japan, the Philippines and Indonesia. It
was first recorded from O. arenosella by Wilkinson (1929), but is known from
larvae of several other families of Lepidoptera, including Lasiocamdidae,
Noctuidae, Pyralidae and Tortricidae (Herting 1971-1982; Thompson 19431965). It is conunonly reared from O. arenosella in India and Sri Lanka.
This is a solitary, internaI parasitoid which attacks first and second instar
larvae of O. arenosella. The larva completes development in about 7 days
and the life cycle is about 18 days (Perera observations). Nirula (1956a)
reports a life cycle of 10-14 days on second instar larvae and 16-17 on first
instar larvae in India, Parasitism under laboratory conditions is not very
efficient. Dharmaraju (1963) obtained only 5-10% parasitized larvae, while
the second author's trials were less successful.
4
This is a moderately common parasitoid of O. arenosella in southeast Sri
Lanka, but scarce elsewhere on the island. Under field conditions, 15-20%
hyperparasitism by Aphanogmus manilae (Ashmead) (Ceraphronidae) and
Eurytoma albotibialis Ashmead (Eurytomidae) occurs in Sri Lanka (Dharmaraju
1963; Perera unpublished). In India, Apanteles taragamae seems to be lesll
common than in Sri Lanka, although Nadarajan & Channa Basavanna
(1980) note it as one of the most effective around Bangalore and Mangalore.
Hyperparasitism by Pediobius imbreus (Walker) and Eurytoma braconidis
Ferriere has been reported (Ghosh & Abdurahiman 1985). Ferriere (1930)
described Perilampus microgastris from braconid hosts, including 'a braconid
parasite of Nephantis serinopa, Meyr. (India)' which is probably referable
to A. taragamae, or perhaps a Bracon sp.
(c)
Brachymeria excarinata Gahan (Chalcididae)
The association ofthis species with O. arenosella is based upon a single specimen
reared from 600 pupae collected in Kerala (Joy & Joseph 1972; Joy et al. 1974).
(d)
Brachymeria hime attevae Joseph, Narendran & Joy (Chalcididae)
This species is included among the six Brachymeria spp. listed from O. arenosella
in India by Joy & Joseph (1973); it is not an important parasitoid of
O. arenosella.
(e)
Brachymeria lasus (Walker) (Chalcididae)
Joy et al. (1974) recorded five B. lasus from a sample of 600 pupae of O.
arenosella in Kerala. This very polyphagous species is clearly an opportunistic
parasitoid of O. arenosella, if not a hyperparasitoid (Cock 1987). Joy et al.
(1978) described its biology on this hosto The listing of B. euploeae (Westwood)
from O. arenosella by Joy & Joseph (1973) is almost certainly in error for B.
lasus (cf. Cock 1987).
(f)
Brachymeria nephantidis Gahan (Chalcididae)
This species was originally described from O. arenosella in India (Gahan 1930)
but it also occurs in Sri Lanka. Early reports suggested that it is the commonest
Brachymeria sp. attacking O. arenosella in India. However, once Joy &
Joseph (1972) recognised B. nosatoi from India, it became apparent that the
latter is the commonest Brachymeria sp., and B. nephantidis, although usually
the second commonest, is rather less frequent (Joy & Joseph 1972, 1973; Pillai
& Nair 1981, 1982d).
The biology of this solitary pupal parasitoid is described by Satpathy & Rao
(1972) and Joy et al. (1978). Adult females do not readily parasitize naked
pupae. It can be bred conveniently in the laboratory on one day old
unexcised pupae of Corcyra cephalonica (Stainton) (Pyralidae) (Perera,
unpublished).
Dharmaraju (1963) recorded B. nephantidis as a hyperparasitoid of Eriborus
trochanteratus (as Nythobia sp.)
5
(g)
Brachymeria nosatoi Habu (Chalcididae)
Joy & Joseph (1972) recorded this pupal parasitoid from O. arenosella as 'a
new geographical race'. It differs slightly from B. nosatoi described from
Japan. The biology is described by Joy & Joseph (1973). They note that it
can be fOl;lnd searching around O. arenosel/a galleries. In the laboratory,
although it readily parasitizes O. arenosel/a pupae in their galleries by inserting
its ovipositor through the wall of the gallery, it is reluctant to oviposit on
excised pupae. Similarly, Pillai & Nair (1982b) recorded that the use ofpupae
in cocoons inside galleries was nec;lssary to develop a breeding technique for
B. nosatoi. This evident specialization contrasts with other species of Brachymeria recorded from this host. However, B, nosatoi was described from
pyralid and tortricid hosts in Japan (Habu 1966) and Joy & Joseph (1973)
also record it from pyralid pests of castor and teak.
Studying parasitoid emergence ho1es in field collected pupae from Kerala,
Pillai & Nair (1982d) conc1ude that B. nosatoi is the commonest pupa
parasitoid (up to 30% parasitism) followed by B. nephantidis (16% parasitism)
Pillai & Nair (1982d) recorded levels of pupal parasitism of 19 % and 44 %
in Kerala in October-December 1980 and January 1981 respectively, in an
area where Xanthopimpla nana Schulz was also commoo. Joy & Joseph (1972,
1973) also record that this is the commonest species of Brachymeria attacking
O. arenosel/a and suggest that where it is present the pest is under natural
control, and outbreaks are linked to its absence. They suggest that where it is
absent B. nosatoi should be introduced.
B. nosatoi was not identified as attacking O. arenosella in Sri Lanka untÏl
1982, although there are specimens from Sri Lanka in the British Museum
(Natural History). The Brachymeria spp. on O. arenosella had not been
studied in detail and all parasitism had previously been recorded as due to
B. nephantidis. B. nosatoi is now recorded as widely distributed in Sri Lanka
and has occasionally been recorded as giving a higher percent parasitism than
B. nephantidis (Perera PhD MS).
Joy & Joseph (1977) discuss the use of Brachymeria spp. for release against
O. arenosel/a. They conc1ude that B. nosatoi would be the most effective,
but that problems in developing a mass culture method need to be overcome
first.
(h)
Bracon brevicornis Wesmael (Braconidae)
Two very closely related species, forms or races are involved under this name:
B. hebetor Say and B. brevicornis (e.g. Puttarudriah & Channa Basavanna
1956). We are not aware that the question of whether one or two species are
involved has been satisfactorily resolved, but on the advice of the CIE we treat
both names as referring to one morphologically defined species. Bracon
hebetor is the senior name, but the name B. brevicornis has been consistently
applied in the literature to the species attacking O. arenosella so we use it here.
This is a gregarious larval ectoparasitoid, very widely distributed and with
a wide host range. It is recorded as aparasitoid of O. arenosella from India,
but not Sri Lanka, where it was the subject of an introduction programme.
6
Sudheendrakumar et al. (1982) describe the biology and immature stages.
Development takes 6-7 days only. Narendran et al. (1980) record observations
on oviposition behaviour Adult females paralyze the host by stinging; they
can lay 25 eggs per day, and oviposition is inhibited below 14-15°C. Jacob
et al. (1980) and Mathew et al. (1980) studied the effects of ambient conditions,
host size, parasitoid: host ratios and male: female exposure ratios to optimize
progeny and sex ratio when reared on Corcyra cephalonica. Ghosh & Abdurahiman (1985) record Pediobius imbreus as a hyperparasitoid in India.
(i)
Bracon serinopae (Cherian) (Braconidae)
Cherian (1929) described this species in the genus Microbracon, which is here
treated as below generic rank. His account of its biology remains the definative
one.
The adult females lay 8-12 eggs per host which they first paralyse. The
maximum observed fecundity was 440 eggs in 55 days. Development is
ectoparasitic and takes OnlY 6-9 days. No other hosts have been recorded
in the field, although alternative hosts were attacked in the laboratory. In
the 1920s it was produced and released in large numbers on the west coast
of Tamil Nadu (Ramachandra Rao 1929). Cherian notes that a Pediobius
(=Pleurotropis) sp. has been reared as a hyperparasitoid in Kerala.
G)
Elasmus nephantidis Rohwer (Elasmidae)
As the specifie name implies, this species was first described as a parasitoid
of O. arenosella (Rohwer 1921) from Coimbatore, India. Ferriere (1929)
recorded a specimen from West Malaysia, and Thomas (1960) recorded it
from a sesiid pest of cocoa there. As it does not occur in Sri Lanka, introductions have been attempted.
Ramachandra Rao & Cherian (1927) published notes on the life history
and Pillai & Nair (1982a) give details on the mating behaviour and biology.
Eggs are laid singly on prepupae of O. arenosella after the host has been stung
and paralysed. The larvae develop as ectoparasitoids of the prepupa.
Development from egg to adult takes 10-16 days, the sex ratio is female biased,
and fecundity is 14-57.
Nirula (1956b) states that this parasitoid accounts for 5-10% parasitism
of o. arenosella throughout India. In a three year survey in Kerala, George
et al. (1977) found 3.25 %'
(k)
Eriborus trochanteratus (Morley) (Ichneumonidae)
In the literature on O. arenosella, this species has also appeared in the genera
Dioctes and Angitia, and sometimes as Nythobia sp., Dioctes sp. or Angitia sp.
It was originally described from Sri Lanka (Morley 1913), where it is a common
parasitoid of O. arenosella. Although it has been recorded from 'Dichocrocis
sp.' (Pyralidae) in India (Ramakrishnan Ayyar & Margabandu 1934) and
there are specimens in the British Museum (Natural History) reared in India
from another pyralid, Antigastra catalaunalis (Duponche1) (I.D. Gauld, pers.
comm. 1987), it has never been reported from O. arenosella in India. It seems
7
likely that E. trochanteratus has adapted either to O. arenose/la or the coconut
habitat in Sri Lanka. Certainly there is evidence of sorne plasticity in its
biology. Cultures at CRI maintained on Corcyra cephalonica as factitious
host became unable to parasitize O. arenosella (H.C,J. Godfray & M.P. Hassell,
pers. comm. 1986). Renee, a Sri Lanka strain adapted to O. arellose/la may
weIl be worth introducing into India.
Petera (1977) has described the laboratory culture of this species. It is a
solitary larval endoparasitoid, Fifth instar C. cephalonica larvae are preferred
for parasitization, and the parasitoid emerges from the prepupa and pupates
inside the host cocoon. Ad.ult females live about ten days, during which they
can parasitize about 120 larvae; the life cycle takes about 16 days.
This is the commonest larval parasitoid throughout Sri Lanka and Dharmaraju (1963) recorded parasitism rates of 1-37%. Perera (1977) gives ilIustrative
levels of field parasitism from diverse 10calities varying from 5 to 54 %' This
species has been used for augmentative releases in Sri Lanka.
The incidence of hyperparasitism is not very high, seldom exceeding 6 %
(Perera 1977). The hyperparasitoids include Aphanogmus manilae (Ceraphron
sp. of Dharmaraju 1963 and Perera 1977) which is the commonest, Eurytoma
albotibialis, Brachymeria nephantidis, which normally acts as a primary
parasitoid (Perera 1977), Pediobius imbreus (Perera unpublished) and Goniozus
nephantidis (Dharmaraju 1963).
(1)
Eurytoma spp. (Eurytomidae)
Eu/'ytoma sp., recorded as a pupal parasitoid of O. arenose/la in India
(Seshagirirao & Dharmaraju 1967), could weIl be hyperparasitic. Similarly
Eurytoma sp., recorded from larvae in Sri Lanka (Dharmaraju 1963), is a
hyperparasitoid of Goniozus nephantidis and Apanteles taragamae. Dharmaraju
(1963) records Eurytoma albotibialis as a pupal parasitoid in Sri Lanka, but
Perera (unpublished) finds this to he a hyperparasitoid of A. taragamae.
(m) Fomicia sp. (Braconidae)
Jayaratnam (1941a) lists Fomicia sp. nr. ceylonica Wilkinson as a parasitoid
of O. arenose/la in Sri Lanka, This seems unlikely as Fomicia spp. are otherwise known only as parasitoids of Limacod.idae (Austin in press).
(n)
Goniozus nephantidis (Muesebeck) (Bethylidae)
Muesebeck (1934) described this species as Perisierola nephantidis from O.
arenosella in South India and it has been recorded from Sri Lanka, but from
no other hosts. In the literature it has also appeared as Goniozus, into which
genus it has recently been transferred.
Antony & Kurian (1960), Jayaratnam (1941b) and Ramachandra Rao &
Cherian (1927) describe the biology and Dharmaraju & Pradhan (1977) and
Remadevi et al. (1978) discuss its culture. It is a gregarious larvalor prepupal
ectoparasitoid, and the female parent practices brood care. The host larvae
are paralysed and the parasitoid may host feed; 10-15 eggs are laid on a mature
8
larva, increasing to 15-18 for a prepupa, but only one or two for a third instar
larva. The life cycle takes 12-16 days including a preoviposition period of
three days. The sex ratio is 0.84 females and the fecundity 20 (maximum 90).
The adult female has up to three cycles of oviposition and brood care, each
lasting 12-13 days and separated by 4-5 days (Sundaramurthy & Santhanakrishnan 1978). In the field it seems unlikely that fcmales survive to parasitize
more than one or occasionally two hosts. In the laboratory, G. nephantidis
can be cultured on larvae of Corcyra cephalonica (Remadevi et al. 1981),
although Paul et al. (1979 report that it is not as suitable a host as O. arenosella.
Field observations on rates o' larval parasitism include those of George
et al. (1977), who monitored parasitism from 1967 to 1970 at Kayangulam,
Kerala. They found 8.5 % larval parasitism, out of a total of 12.6 %, was
due to G. nephantidis, and that larval parasitism was directly proportional
to host density. Manjunath (1985) reports that G. Ilephantidis was the
principal parasitoid in an outbreak of O. arenosella in Andra Pradesh,
accounting for 28 % parasitism, Nadarajan & Channa Basavanna (1980)
report it as one of the most effective parasitoids around Bangalore and Mangalore. It is said to be favoured by warm, dry weather (Ramakrishlla Ayyar
& Analltanarayanan 1935). In Sri Lanka, Dharmaraju (1963) recorded rates
of parasitism up to 19 %'
This is one of the parasitoids relcased against O. arenosella in India. Sundaramurthy & Santhanakrishnan (1978) discuss the usefulll(f;s of releasing unmated
females which sometimes predominate in mass culture. Sincc unmated fcmaJes
parasitize hosts as effectively as mated ones, it would make no difference
whether females were rnated or not for an innundative release. However,
because each G. nephantidis will only parasitize 1-3 O. arellosella larvae it i5
likely to be more effective as an augmentative release, when unmated material
will be of little use.
Hyperparasitism is common in Sri Lanka, a eulophid, Pediobius imbreus
(? P. detrimentosus (Gahan) of Dharmaraju 1963), being the commonest species,
accounting for 6-8 % hyperparasitism. Gahan (1930) described Pediobius
detrimentosus from this host in India. Ramachandra Rao & Cherian (1928)
record this (as Pleurotropis sp.) as well as ?Aphanogmus mani/ae ('a small
black species, probably a proctupoid'), Eurytoma sp., and the mite Pyemotes
ventricosus (Newport). Between them these secondary parasitoids can
severely linùt the effectivcness of Goniozus nephantidis towards the end of the
season. In turn, G. nephantidis has been recorded as a hyperparasitoid of
Eriborus trochanteratus, although this seems unlikcly.
Sundaramurthy & Santhanakrishnan (1979) carried out field cage studies
which demonstrated that mortality of O. arenosella larvae due to parasitism
is directly proportional to the density of G. lIephantidis and inversely proportional to the density of hosts. Givcn its biology, such a rcsult is to be expected
and any density dependent effect could only be expressed in a delayed density
depcndent manner.
9
(0)
Goryphus sp. (Ichneumonidae)
Goryphus sp. nr. (Brachycoryuphus) nursei (Cameron) was recOIdec:l. by Nirula
et al. (1955) as a solitary pupal parasitoid of O. arenosella from coastal and
centra.l Kerala (Travancore) at a rate of 6 % only in July to December, when
the relative humidity was high and the tempcrature moderate. The lack of
subsequent records suggests this is an atypical association. Eggs are laid on
or near the prepupa and the mature larva emerges frorn the pupa to pupate
in its own cocoon. Development takes 12-16 days.
(p)
Megaselia sp. (Phoridae)
This phorid has been reported once from field collected pupae of O. arenosella
(Rernadevi et al. 1980). The authors were uncertain as to whether it was
parasitic or saprophytic, but based on the biology which they recOI ded,
concluded that it is at least potentiaUy parasitic.
(q)
Meteoridea hutsoni (Nixon) (Braconidae)
This species was described by Nixon (1941) from a series reared in Sri Lanka
from Syllepte derogata (F.). Although it has not been recorded parasitizing
O. arenosella in Sri Lanka, it does so in India. Sudheendra Kumar et al.
(1979) first recorded it from O. arenosella and Ghosh & Abdurahiman (1984)
provide details of its incidence and biology.
M. hutsoni has a limited distribution in the coastal area of Kerala, where
Ghosh & Abdurahiman (1984) recorded 10% parasitism. This larval-pupal
parasitoid attacks third, fourth and fifth instar larvae and development takes
17-23 days. This is not an important natural enemy of O. arenosella and
probably normally develops on other hosts.
(r)
Pyemotes ventricosus (Newport) (Acarina)
This parasitic mite is infamous as the agent which caused synchronization of
the population of a hispine, Promecotheca coeruleipennis Blanchard, and hence
massive outbreaks in Fiji (Taylor 1937). It has been recorded attacking O.
arenosella in Kerala state, India (Mathen et al. 1970) and Sri Lanka
(Dharmaraju 1963). Taylor (1937) describes its biology in detail. Although
synchronization may play a role in the outbreaks of O. arenosella, there are
insufficient records of P. ventricosus from this host to suggest it plays a role in
this process.
ln mass rearing programmes, P. ventricosus can cause problems as a contaminant, killing larvae of both O. arenosella and its parasitoids (T.M.
Manjunath, pers. comm. 1984).
(8)
Stomatoceras sulcati:>cuteJ/um Girault (Chalcididae)
Joy & Joseph (1974) review this species as a parasitoid of Opisina arenosella
in lndia. Although there are reliable records fJOrn O. arenosella, reports of
it being cornmon and widespread (e.g. Jayaratnarn 1941a; Ramachandra
Rao et al. 1948) most probably refer to Brachymeria spp.
10
(t)
Stomatomyia bezziana Baranoff (Tachinidae)
Although described in the genus Stomatomyia (Baranoff 1934), S. bezziana
also appears in the literature in the genus Spoggosia, of which Stomatomyia
has been considered a subgenus. However, in recent catalogues (Croskeys
1976; Delfinado & Hardy 1977) Stomatomyia is treated as a distinct genus,
hence the combination Stomatomyia bezziana is usee!. here. At present S.
bezziana is known on1y from Sri Lanka, but Crosskey (1976) has suggested
that S. bezziana, S. innocens (Wiedemann) described from Macao, S. acuminata
(Rondani) described from Italy and wid.espreacl in the 01d. world tropics and
S. approximata Villeneuve may ail be synonyms, in which case S. innocens
would be the senior name. The fact that no Stomatomyia s;:>. has been reared
from O. arenosel/a in India, while this is the normal host in Sri Lanka, leads
us to believe that the Sri Lankan species is a disfnct entity to which the name
S. bezziana can be applied.
Rao & Rao (1964) describe culture techniques for S. bezziana and describe
and illustrate the life history. Freshly emerged females were mated with
1-2 day old males, and oviposition commenced after a 2 day pre-oviposition
period. Females survived 17 days and the fecundity varied from 25-127
(average 61) macrotype eggs laid preferentially on full grown larvae of O.
arenosel/a, which are parasitized in their galleries. A culture method on an
alternative host has been developed in Sri Lanka by Perera, using larvae of
Corcyra cephalonica which have first bcen paralysed by the stings of Bracon
breJ'icornis ＬＮ＾［ｲｾ･ｐＨ
lInpublished PhD thcsis). The larval stage lasts 4-5 days
and the pupal stage about 9, giving a minimum life cycle of 17 days from egg
to egg-laying adult, and a maximum of about 27 days if it is assumed that
feroale flies can continue to oviposit for 10 days. Superparasitism is frequent
but only one or, with ｾｲｵｴ｡ｭ
host larvae, sometimes two parasitoids reach
maturity.
This species has a patchy distribution within Sri Lanka, being most frequent
in the Eastern Province and the south-east, but is not always apparent in the
field.. At times it is common, particularly at the end. of outbreaks and
Dharmaraju (1963) gives parasitism rates up to 80 %' Sorne redistribution
within Sri Lanka has been carried out and establishment and rapid
multiplication reported (Dharmaraju 1963), but this work has not been critically
assessed.. 1t suffers hyperparasitism by Brachymeria nephantidis, Eurytoma
albotibialis, Exoristobia philippinensis Ashmead, Neasteropaeus sp., Pediobius
imbreus and Trichospilus pupil'orus (Dharmaraju 1963; Jayaratnam 1941a;
Perera unpublished PhD thesis).
(u)
Tetrastichus israeli (Mani & Kurian) (Eulophidae)
This gIegarioliS pupal endoparasitoid has a wide host range, but is most
commonly associated, at least in the literature, with pyralid stem borers of
rice, sugarcane, etc. (Abraham et al. 1974; Varma et al. 1981). Culture
methods have been studied (Nadarajan & Jayaraj 1975, 1977) and it has been
mass reared on a v?.riety of hosts (Tuhan et al. 1977). Releases have been
made in India for the control of graminaceous stemborers (Varma et al. 1980)
Il
and it has been exported to other countries for release against a variety of pests
(Bordat et al. 1977; Etienne 1973; Maafo 1975). It has been recorded as a
regular parasitoid of O. arenosella in Tamil Nadu, India (Ali & Subramanian
1972) and so introductions were made in Sri Lanka (Section III(g).)
(v)
Thelairodrino graciUs (Mesnil) (Tachinidae)
During surveys carried out in the states of Kerala and Andhra Pradesh, Rao
(1961) reported T. graciUs attacking 2-5 % of O. arenosella pupae in one area
(Ernakulam). T. graciUs has also been reported from a noctuid host (Thompson
1943-1965). According to Nagarkatti (1973 this is the species referred to as
Winthemia sp. by Ramachandra Rao et al. (1948) and, she suggests T. graciUs
merits investigation for augmentative releases, as oruy easily cultured species
can be used.
(w)
Trichospilus pupivorus Ferriere (Eulophidae)
Boucek (1976) reviewed the hosts and distribution of this species. It appears
throughout the literature as T. pupivora, as named by Ferriere (1930), but
Z. Boucek (pers. comm. 1987) advises us that 'pupivora' is an adjective and
as such should agree with the generic name, hence we use T. pupivorus. Boucek
(1976) recorded it from Sri Lanka, southern India, West Malaysia, Java, New
Guinea and New Britain. It was introduced into Mauritius against sugarcane
stemborers and became established upon other hosts (Greathead 1971).
Studies on the biology and culture (Anantanarayanan 1934; Dharmaraju &
Pradhan 1977; Hutson 1939; Jayaratnam 1941a; Pillai & Nair 1982c) show
that it has a wide field and laboratory host range, although O. arenosella
is a principal hosto It can be readily produced in large nwnbers. It is a
gregarious pupal endoparasitoid, with a life cycle of 16-19 days. An average
of 133 adults is obtained from pupae of O. arenosella (less or more From other
hosts depending mostty upon size) The small number of males produced in
each brood mate with the females within the host pupa. Each female can
only par:lsitize one or two host pupae. The pupae aestivate between March
and May in India (Pillai & Nair 1982c). In the only critical assessment of
the biological characters of this speeies, Pillai & Nair (1982c) conc1ude that
poor searching and dispersive ability, low non-optimal temperature tolerance
inability to recognize parasitized pupae (which may not be true - see
Jayaratnam (l941a) and inability to compete with other pupal parasitoids
render it an ineffective species for the natural control of O. arenosella.
In Sri Lanka, T. pupivorus is found most frequently in the west and northwest, occasionally in the south. Dharmaraju (1963) records parasitism rates
of up ta 38 %. Its absence From the east is generally attributed to the hotter
climate, as intensive releases have been made in that area. Rodrigo (1943)
and Seneviratne (1945) suggest that releases in the dry (hot) season are
ineffective. Levels of parasitism are variable, but generally low. Thus, in
Kerala, George et al. (1977) record 6.8 % pupal parasitism by T. pupivorus
compared to 41.5 % by Brachymeria spp. Again in Kerala, Pillai & Nair (1981)
cite pupal parasitism as causing more than 50 % mortality but T. pupivorus
is responsible for only 1.6% of this. On the other hand Venkitasubban (1938)
12
reported T. pupivorus as the most important parasitoid in Kerala. T. pupivorus
is occasionally reared as a secondary parasitoid from the tachinids Stomatomyia
bezziana, Bessa remota (Aldrich) and Argyrophylax fumipennis Townsend.
(x)
Xanthopimpla spp. (Ichneumonidae)
X. nana Schulz is a replacement name for X. parva Cameron which was
described from Sri Lanka. It is a generalist para'litoid of Lepidoptera pupae,
and since it is only sporadically recorded from Opisina arenosella this,
association is consideœd opportunistic. Pillai & Nair (1983) described its
biology in Kerala and recorded levels of pupal para'litism of 27.5 % for 1980
and Ｙｾ
for 1981.
X. punctata (F.) is another species of tbis genus œcorded from O. arenosella
(Ramakrishna Ayyar & Margabandu 1934) and again is an example of a
generalist pupal parasitoid using an abundant host when available. Perera
(unpublished) has occasionally reared a Xanthopimpla sp. in Sri Lanka which
could refer to either of the above species.
(y)
Other pupal parasitoids
Subba Rao & Hayat (1986) list the chalcidids A. Izakonensis (Ashrnead), A.
phaeospi/us Waterston and Brachymeria podagrica (F.) as parasitoids of O.
arenosella in India, in addition to those we have tœated above. We have not
traced the references they cite, but anticipate that aIl these records are of
generalist species wbich occasion1.lly parasitize O. arenosella when it is cornmon
Perera (unpublished) has also recorded Trathala sp. (Ichneumonidae) as a
rare pupai parasitoid in Sri Lanka. It needs to be clarified whether this is
T. flavoorbitalis (Cameron) discussed in Section IV 2(e), which has been
recorded frorn Sri Lanka, but not from O. arenosella.
(z)
Other probable hyperparasitoids
The records of Tetrastichus coorgensis Kurian (Eulophidae) and Invreia opisinae
Narendran (Cha1cididae) listed by Subba Rao & Hayat (1986) from O.
arenosella in India need clarification as to the biologicai roles of the species
concerned. We consider them most likcly to be hyperparasitoids until proven
otherwise.
III. CLASSICAL BIOLOGICAL CONTROL - HISTORICAL
Biologicai control of Opisina arenosella by the introduction of exotic natural enemies
can fall into one of two categories. Either natural enemies can be sought
which attack O. arenosella elsewhere in its range, or natural enemies of species related
to O. arenosella -- whether taxonomically or biologically - may be suitable for use.
Because O. arenosella is indigenous where it is a pest, the possibilities for
introducing natural enemies which attack it elsewhere are limited. Nagarkatti
(1973) has suggested that a survey of natural enemies throughout India may reveal
gaps in the distribution of so:ne of the more important parasitoids, which could
then be moved around within the country. Similarly there are a few differences
between the fauna of Sri Lanka and India which have been xeploited. These
introductions are discussed below.
13
Carl (1982) reviewed the biological control of native pests and showed that there
are sufficient examples of successful biological control introductions for them to be
considered suitable targets for c1assical biological control. The attempted
Bessa remota, a tachinid parasitoid of a coconut feeding zygaenid
introduction
from South-East Asia, for the control of O. arenosella falls into this category.
of
(a)
Antrocephalus pandens (Walker) (Chalcididae) releases in Sri Lanka.
As discussed in Section II, Brachymeria nosatoi is a common pupal parasitoid
of Opisina arenosella in India, but it was not recognised from Sri Lanka until
1982. In the same year CRI obtained a nucleus culture, purportedly of B.
nosatoi, from India, Specimens from this culture were submitted to CIE
and determined by Z. Boucek as Antrocephalus pandens, a species described
from Sri Lanka, i.e. somewhere along the line the culture became
contaminated.. Since O. arenosella was a suitable host in the laboratory,
re1eases : 25,000 were made over two years. However, since no recoveries
were made, it seems that it is not a suitable field hosto
(b)
Bessa remota (Aldrich) releases in India and Sri Lanka
This tachinid is famous as the successful biological control agent of a zygaenid,
Levuana iridescens Bethune-Baker, in Fiji (Tothill et al. 1930). It is indigenous
to South-East Asia, where it is best known as a parasitoid. of the coconut
defoliating zygaenid Artona catoxantha (Hampson) (cf. Section IV.2(c). It
has been leared from several other Lepidoptera and in the laboratory has a
wide host range, including the predatory c1erid beetle Callimerus arcufer
Chapin. The macrotype eggs are laid. on medium to full grown larvae, and
the maggot completes development in the prepupa, from which it emerges
to form its pupariurn within the cocoon of the hosto The life cycle from egg to
ad.ult can take from 13 to 29 days (18 in India). There is a minimum preoviposition phase of 2 days and females can survive more than 3 weeks, but
oviposition is insignificant in the third week. The potential fecundity is 25-100
(Tothill et al. 1930; Jayanth & Nagarkatti 1984).
Bessa remota was imported from Fiji into Sri Lanka in 1975 and cultured
and released (Mahindapala 1977). Just under 8,000 were released in 1976,
mostly in the Eastern Province, but sorne in the north west and some in the
south. No recoveries were made. B. remota was first introduced into India
from Burma in 1938 for the control of the teak defoliators Eutectona machaeralis
(Walker) and Hyblaea puera (Cramer) and released near Madras (Beeson
1940), but no establishment has been reported. A subsequent record from
an Ailanthus defoliator, Atteva fabriciella Swederus (Mathur et al. 1970)
needs confirmation.
ln 1980 CIBC arranged. the supply of B. remota from West Malaysia to
India (CIBC 1981) and again it was successfuUy cultured (Jayanth & Nagarkatti
1984) and released (India 1983) (Table 1). No recoveries were made. Unpublished reports suggest that t\e flies were unable to reach the target larvae
in their galleries (M.J. Ch'lcko, pers. comm. 1986). Observations in field
cages would be needed to confirm this. However, given the wide host range
of B. remota, it could. weIl have become established in either country on other
hosts.
14
Table 1.
Releases of Bessa remota in Southern India, 1981 - 1982 (India 1983: G. B.
Pillai, pers. comm. 1986)
No. released
Locality
Release period
Quilon (Ponmana), coastal Kerala
v-ix.81.iii.82
AUepey (Thottappally), coastal Kerala
iv-vi.82
76
Cochin (Ernakulum), coastal Kerala
iv-vi.82
188
Salem (Kendampatti) (inland.>3500m. Tamil Nadu)
viii-xii.81
448
(c)
1757
Bracon brevicornis releases in Sri Lanka
Bracon brevicornis is known from Sri Lanka, where it principaUy acts as a
parasitoid of lepidopteran stored product pests. It had not been recorded.
from O. arenosella in Sri Lanka, but was known to attack this host in India.
Accordingly CIBC Indian station provided a culture to CRI in 1959
(Dharmaraju 1963). It was mass reared on Corcyra cephalonica and starting
in 1960 sorne 100,000 were released annually. B. brevicornis was subsequently
recovered from O. arenosella in 1975 in North-West province, and now occurs
sporad.ically at 0.8 % parasitism in most areas. Whether these recoveries are
due ta an introduced strain from India becoming established on its normal
host, or the local strain beeoming adapted to a new host is open to speculation,
and no firm evidence is available either way. It is hyperparasitized by Pediobius
?erionotae Kerrieh (Perera unpublished).
(d)
Elasmus nephantidis releases in Sri Lanka
This gregarious prepupal parasitoid has been released against O. arenosella
in Sri Lanka. Its introduction was first attempted in 1962 when a culture
was obtained from the Central Coeonut Research Station, Kerala, and about
5000 released in Eastern Province (Dharmaraju 1963). In 1971, it was supplied.
by the CIBC Indian Station and about 50,000 released in aU areas between
1971 and 1976 (Kanagaratnam 1974, 1975; Mahindapala 1972, 1973), when
culture was stopped (Mahindapala 1977). No recoveries were made.
(e)
Eriborus trochanteratlls releases in India
G. B. Pillai (in press) obtained E. trochanteratus from Coimbatore in 1978.
It was cultured on larvae of O. arenosella and Corcyra cephalonica and 1049
adults (62 % female) were released in 1979-80 at Thottappally, Alleppey District
Kerala. Fifteen cocoons of E. trochanteratlls were subsequently recovered from
the field, of whieh three were hyperparasitized by Brachymeria nephantidis.
If this parasitoid persists in India, it should be a useful addition to the natural
cnemy complex.
15
(f)
Stomatomyia bezziana releases in India
Rao et al. (1971) record the introduction of S. bezziana from Eastern Province,
Sri Lanka, to India during 1962·64. This tachinid was cultured at the CIBC
Indian Station from 1960-1963, using the methods described by Rao & Rao
(1964). It was supplied to Coimbatore fOl culture and release in 1962 and
to Orissa in 1963. Releases on the Kerala coast led to recoveries at a low
rate of parasitism, and Trichospilus pupivorus and Brachymeria nephantidis
wae recorded as hyperparasitoids (G.B. Pillai, pers, comm. 1986). The
colony did not persist.
A reported recovery at Sirpur (Orissa State) from the rice pest, Pelopidas
mathias (F.) (Hesperiidae) by Rao et al. (1969) in 1963 may refer to the
indigenous S. innocens (cf. Section II (t) and needs confirmation Nagarkatti
(1973) subsequently suggested further trials with this species and in 1978 ICAR
obtained a nucleus culture from CRI, Sri Lanka. Although it was cultured
for several generations at the Central Plantation Crops Research Institute,
Kerala, the fecundity, survival and sex ratio deteriorated and tlle culture was
lost (G.B. Pillai, comm. pers. 1986). There is no indication that releases were
made.)
(g)
Tetrastichus israeli releases in Sri Lanka
In 1959 this polyphagous pupal parasitoid was sent from the CIBC Indian
Station to CRI (Dharmaraju 1963), cultured on pupae of Spodoptera litura
(F.), and from 1960 to 1973 about 5.7 millions were released in western,
southern, eastern and north-western provinces before culture was discontinued
in 1974 (CRI records). The fact that no recoveries were made in spite of these
large numbers released suggests that either O. arenosella is not a normal host,
or that host specialization occurs in this species and the material used was
not appropliate. In view of the paucity of records from this host in India,
the former explanation seems more likely.
(h)
Trichogramma brasiliensis (Ashmead) releases in Sri Lanka
This Neotropical species has bcen recorded as an egg parasitoid of Heliothis
zea (Boddie) on cotton in Peru (Nag3.rb.tti & Nag3.raja 1977), Diatraea
saccharalis (F.) on sugarcane in Peru, Plutella xylostella (L.) and Trichoplusia
ni (Hubner) on cabbages in Trinid'ld (Cock 1985) and Keiferia lyc:Jpersicella
(Walsingham) on tomato in Trinidad (Barrow et al. 1979). It was in culture
at the CIBC Indian Station from 1967 to 1980, and quite widely distributed
in Indla. Recoveries were made following releases against Heliothis armigera
Hübner in tomato and okra crops in Hessarghatta, Bangaloœ, during 1973
(CIBC 1974). Fortnightly releases in Punjab cotton field infested by bollworms
during 1977-78 are reported to have reduced the infestation from 70% to 30%
(CIBC 1979).
ln 1973 a culture obtained ｦｲｶｾｮ
California was supplied by the CIBC Indian
Station to the CRI, Sri Lanka. ft was mass reared on Corcyra cephalonica
and about 19.5 million released between 1973 and 1977 in aIl coconut areas
of Sri Lanka. There is no indication that it has become established,
although Kanagaratnam (1975) recorded initial recoveries.
16
The known host records discussed above give no indication that T. brasiliensis
is suited to a coconut palIn habitat, indeed the preferred habitat would seem
to be field crops. Rence, there is no reason to be surpnsed at its failure against
o. arenosella, although it could well be established on other hosts in Sri Lanka.
(i)
Trichogramma minutum Riley releases in India and Sri Lanka.
The CRI, Sri Lanka obtained a stock of Trichogramma minutum from the
CIRC Indian Station in 1960 (CIRC 1961), cultured it and released 50,00060,000 in aIl coconut areas of Sri Lanka (unpublished reports). There is no
evidence that it became established.
Nagarkatti (1973) comments on the release of 730,000 Trichogramma sp.
at each of four sites in Andra Pradesh. Although the species released was
supposed to be T. minutum, a sample from the culture proved to be T.
australicum Girault, and it was probably obtained from Karnataka, where it
is used against sugarcane stemborers. Pointing out that there was no evidence
that this species was adapted to an arboreal, coconut habitat or even that
it could attack eggs of O. arenosella, she suggests that O. arenosella should be
surveyed for egg parasitoids.
IV. CLASSICAL BIOLOGICAL CONTROL - POSSIBILITIES
As pointed out in the introduction to section III, there are two approaches to the
classical biological control of O. arenosella.
1.
Natural enemies of Opisina arenosella
This has been the principal approach to classical biological control of O. arenosella
in the past, and is reviewed in Section III. It has failed so far, and in this section
it remains only to highlight the species with which further trials seem worthwhile.
Eriborus trochanteratus is an important larval parasitoid in Sri Lanka which is
absent from India, so its introduction into India must have some potential. The
recoveries reported in Section III (e) need to be followed up. If establishment has
occurred, redistribution will now be necessary; if not, releases should continue.
Also, further attempts to establish Stomatomyia bezziana in India should be made.
The other introductions summarized in Section III can be considered failures (or
completed in the case of Bracon brevicornis released in Sri Lanka), and further effort
is not justified.
The role of natural enemies in the population dynamics of O. arenosella in Bangladesh and Burma has not been investigated, nor are there significant reports of o.
arenosella as a pest in these countries. This is probabiy a refiection of the fact that
coconut is not an important crop, rather than that O. arenosella is under effective
natural control. The natural enemies of O. arenosella in these areas need to be
surveyed for potentially useful biological control agents, but there is no particular
reason to expect anyth.ing of significance to be found, bearing in mind the land is
contiguous with India.
17
2.
Other hosts as sources for natural enemies
The second approaeh is to use parasitoids or predators which attack species similar
to the target, either taxonomically or ecologically. Hence, natural enemies of
Oecophoridae in general might be eonsidered, or natural enemies of small lepidopterous larvae which feed in a concealed manner on leaflets of palms (preferably
coconut). We feel that the latter is more likely to yield effective natural enemies,
if only because of the difficulty of seleeting subjects for study from the whole gamut
of natural enemies of Oecophoridae.
Scattered through the tropics there are probably many small Lepidoptera whose
larvae fced in a conccaled manner on coconut and other palms. Unless they cause
significant damage, they are unlikely to attract much attention in the literature.
Below, we discuss 14 species whose classification and distribution are sununarized
in Table 2. The rcason that they do not cause significant damage is most likely
to be that normally they are kept under control by their natural enemies, although
the occasional flare-up may occur due to misuse of pesticides. While studies have
been made on the natural enemies of sorne of these, most are unknown and there is,
little to record.
(a)
Agonoxena spp. (Agonoxenidae)
Agonoxella is the only genus in the family Agonoxenidae, wltose taxonomie
affinity is to the family Oecophoridae (Bradley 1965). There are four spccics
in tlte genus (Table 3). The coconut flat moth, A. argaula Meyrick is an
economic pest in the Pacifie and its biology (Singh 1952) and natural eneIllies
(Hinckley 1963) have been studied. The larvae fecd only on the lower
epidermis and parenchyma, and spin a fine web over the surface under which
they feed. The larvae are active and readily leave their shelters. Pupation
IS under a white web, either on a leaflet or on nearby undergrowth. In the
course of attempts at biological control in Fiji the indigenous natural enemies
were surveyed (Hinekley 1963) and are included in Table 4. The braconid
larval parasitoids (Agathis sp., Apanteles agonoxenae Fulœ.way and Bracon
sp.) are eommon and potentially effective control agents, except tltat they are
heavily hyperparasitized by a range of indigenous species (Table 5).
While working on the biological control of other pests for Fiji, R. W. Paine
was able to investigate the natura) enemies of Agolloxena phoenicia Bradley
in Queensland (Paine 1961), and of A. pyrograrnrna Meyrick in Papua New
Guinea and New Britain (O'Connor 1960). He found both species under
effective natural control by a range of parasitoids (Table 4) including Chelonus
sp. (braconid egg-Iarval). Ooellcyrtus sp. (encyrtid pupal), and Actia painei
Crosskey (an ovolarviparous tachinid). Introductions of several parasitoids
were attempted (some in very small numbers). but only Brachymeria
agonoxenae Fullaway and perhaps Macrocentrus sp. became established,
although O'Connor (1960) expresses doubt about the latter. The natural
enemies of Agonoxena miniana (Meyrick), which is not a pest in Java, have
not been investigated adequately and may weIl include additional gui Ids.
18
Table 2.
Swnmary of Lepidoptera which feed upon leaflets of palm in a concealed
manner.
Species
Agonoxenidae
Agonoxena argaula M eyrick
A. pyrogramma Meyrick
A. miniani (Meyrick)
A. phoenicea (Bradley)
Oecophoridae
Durrantia arcanella (Busck)
Struthocelis semiotarsa (Meyrick)
Pyralidae
Furcivena rhodoneurialis
Hampson
Hedylepta blackburni (Butler!
Herculia nigrivitta (Walker)
Stenomidae
Antaeotricha spp.
Loxotoma elegans Zeller
Stenoma impressella (Busck)
Xylorictidae
?Odites sp.
Zygaenidae
Homophylotis catori (Jordan)
Artona catoxantha (Hampson)
Table 3.
Distribution
Status of research
on natural enemies
South-west Pacifie
Melanesia
Java
Queensland
well known
Surveyed
CasuaI observations
SUlveyed
Neotropics
Northern South America
Casual observations
Casuai observations
West Africa
Unknown
Hawaii
South and South East Asia
WeB known
Casual observations
Northern South America
South America
Northern South America
Casuai observations
Unknown
Casual observations
Malaysia
Unknown
West Africa
Burma, Malaysia and
West Indonesia
Casual observations
Well known
The distribution and hostplants of Agonoxena spp.
Species
Distributioil
Hostplants
A. argaula Meyriek
South - west Pacifie: Fiji,
Kiribati, New
CaIed.onia,
Niue, American & Western
Samoa, Tokelau,
Tonga,
Tuvalu, Vanuatu, Wallis &
Futuna Is.
Introdueed
Hawaii, Palmyra
Coconut and other palms:
(Chrysalidoareca
palm
Wendl.)
carpus
lutescens
bottle palm (Hyoporbe sp.)
Kentia sp., Pritchardia pacifica
Seem. & H. Wend1., Veitchia
filifera (H. Wend.l.) H. E.
MOOle
Coconut. ?others
A. pyrogramma (Meyriek) Guam, Marianas, New Britain, Papua New Guinea.
Solomon Is.
Java, ?Papua New Guinea
A. miniana (Meyrick)
A. phoenicea <Bradley)
Queensland
Coconut, (?Calamus sp. lU
PNG)
Northern bungalow palm·
1Archontophoenix alexandrae
Muellel)
19
One attempt has been made to use parasitoids of Opisina arenosella to control
Agonoxena argula in Fiji (Kamath 1979). The CIBC Indian Station sent
cultures of Goniozus nephantidis. Elasmus nephantidis, Bracon brevicornis
and Trichospilus pupivorus ta Fiji in 1971. In Labaratory tests (Rao 1972)
E. nephantidis would not breed on A. argaula, while G. nephantidis and B.
brevicornis would, Releases were made of G. nephantidis (550), E. nephantidis
(45), B. brevicornis (4,600) and T. pupivorus (13,000), but none became
established. The fact that some of the parasitoids of O. arenosella would
attack A. argaula, at least in the laboratory, suggests that the converse may
also be true.
Table 4 Natural enemies of Agonoxena spp.
Specil!s
Braconidae
Agathis sp.
Hosts
Distribution
A. argaula
Fiji
A. phoenicia
A. pyrogramma
A. argaula
Queensland
New Britain
Fiji. Tonga
A. stantoni (Ashmead) A. pyrogramma
Bracon sp.
A. argaula
New Britain
Fiji
Apanteles sp.
Apanteles sp.
A. agonoxenae
Fullaway
Comments
Gregarious larval-prepupal
parasitoid: occasional (6 %):
hyperparasitized (21 %)
Larval parasitoid
Parthenogenetic: solitary: cornmon
(37 %): heavily hyperparasitized
(76%)
Larval parasitoid
2-3 per host: common (28%):
heavily hyperparasitized (42 %)
Bracon sp.
Bracon sp.
Chelonus sp.
A. pyrogramma
A. argaula
A. phoenicia
ｑｵ･ ｮｾｬ｡ｮ､
Macrocentrus sp.
A. miniana
A. argaula
Java
Fiji
J.f. pallidus Fullaway
A. pyrogramm r
Guam
Egg-larval parasitoid:
hyperparasitized
Solitary ｜｡ｲｶｬＭｰ･ｾｵ｡
parasitoid
Introduced from Java, may not be
established
Larval ｰ｡ｲｾｩｴｯ､
Chalcididae
Brachymeria
agonoxenae Fullaway
A. miniana
Java
Pupal parasitoid
A. argaula
A. argaula
Tonga, American Samoa
Fiji
New Britain
American Samoa
Introduced from Java: common
(35 %): facultative hyperparasitoid
Introduced from American Samoa
Pupal parasitoid
Adapted from resident host
A. argaula
B. hammari(Crawford) A. pyrogramma
B.polynesialisCameron A. argaula
Brachymeria sp.
A. argaula
Hawaii
Guam
Hawaii
Fiji, Tuvalu, Kiribati
Encyrtidae
OOi!f/.cyrtus sp.
Ooencyrtus sp.
4. miniana
4. pyrogramma
Java
New Britain
Pupal parasitoid (?)
Pupal parasitoid (?)
Eulophidae
Elachertus
agonoxenae Kerrich
A. pyrogramma
A. phoenicia
New Guinea
Queensland
Larval parasitoid
Larval parasitoid
A. argaula
Hawaü
Adapted from Hedylepta
blackburni'
A. pyrogramma
A. argaula
Solomon Is. New Britain
Pupates within hast larval skin:
Fiji
localised
A. argaula
FiJi
Ichneumonidae
Trathala fial'oorbitalis
(Cameron)
Tachinidae
Actia painei Crosskey
Tongamyia cinerella
Mesnil
Fungi
Indet.
a Already known from Opisina arenosella in Sri Lanka.
20
Common on pupae (16 %)
In terms of gaps within the complex of natural enemies attacking Opisina
arenosella, there are some ､ｩｦ･ｲｾｮｴ
guilds of parasitoids attacking Agonoxena
spp. Chelonus sp. (Braconidae) is an egg-larval para5itoidattacking A. phoenicea
on an ornamental in Queemland. It is by no means certain that it would
adapt to the coconut ecosystem, and it is likely to suifer a degree of hyperparasitism. Hinckley (1963) records that Agathis sp. (Braconidae) i'> a gregariou.
larval-prepupal parasitoid of Agonoxena argaula in Fiji, where it is not very
common \6% parasitism) and sulfers some hyperpara5itism (21 %).
One o' morè Ooencyrtus spp. (Encyrtidae) are œported as pupal parasitoids
of A. miniana in Java and A. pyrogramma in New Britain. Ooencyrtus spp.
are usually (but not exclusively) egg parasitoids, 50 this identification needs
verification. Paine (in Hinckley 1963) has suggested this species can act as
a hyperparasitoid through Macrocentrus sp. and other larval parasitoids and
its role need5 clarification. If it reany is a primary pupal patasitoid it merits
trial for the control of both A. argaula and O. arenosella.
Table 5.
Hyperparasitoids of Agonoxena spp.
Species
Distribution
Ceraphronidae
Aphanogmus fijiellsis
(Ferriere)
Fiji
Chalcididae
Brachymeria agonoxenae Fiji
Fullaway
Eulophidae
Tetrastichus sp.
Eupelmidae
Eupelmus sp.
Fiji
Hosts
Apanteles agonoxenae
Aphanoglllus sp. occurs in India
and Sri Lanka
?Agathis sp., Macrocentrus sp.,
Tongalllyia cinere/la
Facultative hyperparasitoid
Agathis sp., Apanteles sp.,
Bracon sp., Macrocentrus sp.
Fiji
Agathis sp., Apante/es sp.,
BrachYllleria agonoxenae
Macrocentrus sp.
Hawaii
?BrachYllleria sp(p.)
lchneumonidae
Stictopisthus sp.
Gelis teneflus (Say)
Fiji
Hawaii
Apanteles sp.
? Brachymeria sp(p).
Pteromalidae
b"upteromaJus sp.
Fiji
Apanteles sp.
Eupe/mus cushmani
(Crawford) .
Comments
Actia painei Crosskey (Tachinidae) has been ;'eated from larvae of A.
pyrogramma in New Blitain by O'Connor in 1952 and Paine in 1957-58 and
1960 and in Bouganville, Solomon Islands, by Paine in 1960 (Crosskey 1962).
Siphoruni, including Actia spp., are ovolarviparous (Herting 1960) and newly
hatched latvae are able to locate Agonoxena pyrogramma larvae undet their
webs and may weIl be able to find those of O. arenosella in their galleries. As
no culture technique could be deveJoped, this species was never teleased in
Fiji. Paine (pers. comm. 1985) still believes this tachinid is the mos! likely
introduction to succeed against A. argaula and it mcrits trial for control of
O. arenosella.
21
For the biological control of O. arenosella laboratory trials and, if
appropriate field trials should be carried out to test the suitability of these
parasitoids. The natural enemies of A. miniana in Java merit investigation
both for the biological control of A. argaula and as a potential source of
natural enemies for the control of o. arenosella. However, like O.arenose/la
A. argaula is a sporadic outbreak pest. It has been suggested that outbreaks
are linked to periods of hot, dry weather and one possible mechanism is the
climatic effects upon the natural enemies, as has been suggested for O. arenosella
in Sri Lanka. If this were so, there is little reason to think that the species
indigenous to Fiji would be effective in Sri Lanka, while those from other
climates can only b.;; an unknown quantity, testable by trial.
(b)
Antaeotricha spp. (Stenomidae)
In northern South America, three species of this genus, including A. euthrinca
Meyrick and A. phaeoneura (Meyrick) attack oil palm (Genty et al. 1978).
The larva makes a shelter between two leaflets, in which it deve10ps and pupates.
The only significant natural enemy is reported to be a fungus, Beauvaria
tene/la (Delacroix) Siemaszko (Genty et al. 1978).
(c)
Artona catoxantha (Hampson) (Zygaenidae)
This ｳｰ･｣ｩｾ
was described from BUI'ma and occurs as an outbreak pest in
Malaysia, Borneo, Sumatra, Java, Palawan Island (Philippines) and West
Irian (CIE 1955). Nagarkatti (1973) has already suggested that parasitoids
of A. catoxantha should be investigated for the biological control of O.
arenosella, and went on to carry out trials with Bessa remota. A. catoxantha
and its natural enemies have been extensively studied (Betrem 1941; Corbett
1932; Feniere 1940; Gater 1925, 1926 Kalshoven & van de Laan 1981;
Lecfmans 1928; Lever 1953, 1964, 1969; Simmonds 1930; Tothill et al. 1930;
Van der Vecht 1947, 1950, 1954), and the following account is based upon
thcsc sources.
The normalfoodplants are Metroxylon sp. <,.nd coconut, but during outbreaks
a range of other palms can be used. The eggs are laid in small batches and
the larvae, which are well endowed with bristles, feed on the leaflet undersurface
making parallel sided scars. The white cocoon is formed on the leaflet undersurface. Sporadic outbreaks occur, and several times this has been attributed
to the action of parasitoids synchronizing the pest population, with the result
that the unsynchronized parasitoids become rare. The natural enemies are
summarized in Table 6 and their hyperparasitoids, etc. in Table 7. Three
species are reported to be most important: Apanteles artonae Rohwer, Bessa
remota and Argyrophylax jumipennis. The first two of these have been implicated in the synchronization of A. catoxantha populations in West Malaysia
and Java. Bessa remota is discussed in Section III (a), as its use for biological
control of O. arenosella has already been attempted.
Tothill et al. (1930) studied the natural enemies of other zygaenids
in Melanesia. They found that Balataea (Artona) trisignata Snellen, which
feeds on species of Zingiberaceae in Java, i5 attacked by Apanteles artonae
(rare), Bessa remota (rare), Mesostenus sp. (conunon), Trichogrammatoidea
common, but host specific) and an
nana (Zehntner), two chalcidoids ｾｮｯＨ
ichneumonid (common\
22
Table 6.
Natural enemies of Artona catoxantha (Hampson)
Species
Distribution
Role
Parasitoids
Braconidae
Apanteles artonae Rohwer
Java, W. Malaysia
Solitary, young larval parasitoid: common
under normal conditions
Java
Java, Sri Lanka
Young larval parasitoid: common
Larval parasitoid; common
Eulophidae
Neoplectrus bicarinatus
Ferriere
N. maculatus Ferriere
Euplectromorpha artonae
Ferriere
E. viridiceps Ferriere
Eupelmidae
Eupelmus catoxanthae
Ferriere
(?=E. javae auct.)
Anastoidea brachartonae
Gahan
Java
Java, W. Malaysia, India
Gregarious larval and prepupal parasitoid:
common under normal conditions
Java, W. Malaysia
Facultative hyperparasitoid
Java, W. Malaysia.
Recorded ex pupae, but could have becn
hyperparasitic
Java, W. Malaysia
Facultative hyperparasitoid: pupal or
prepupal parasitoid
Java
Java
Java
Java
W. Malaysia
Pupal!prepupal parasitoid: scarce
Pupal/prepupal parasitoid: scarce
Pupal/prepupal parasitoid: common
Pupal/prepupal parasitoid: scarce
Betrem (1941) questions this record of Gater
W. Malaysia
Java
Belrem (1941) questions this record of Gater
Pupal/prepupal parasitoid: scarce
Java, W. Malaysia
Microtype egg-laycr: important: cornrnon in
outbreaks
Fiji, Java, W Malaysia,
Burma, India (?)
Macrotype egg-layer:
larval
prepupal
parasitoid: cornmon in outbreaks
Predators
Cleridae
Callimerus arcufer Chapin
W. Malaysia
Predator of larvae etc.
Pentatomidae
?Cantheconidea furcel/ata
(Wolff)
E. Malaysia (Sabah)
Predator of adults
Pathogens
Fungi: Hyphomycetes
Beuveria bassiana (Balsamo)
Vuillemin
W. Malaysia
Kills mature larvae
Eurytomidae
Eurytoma albotibialis
Ashmead
Ichneumonidae
Goryphus bituberculatus
Szepligeli
G. fasciatipcnnis Szepligeti
G. infcrus Szepligeti
G. javanicus Roman
G. macu/iceps Cameron
G. mesoxanthus maculipennis
(Cameron)
G. rufobasalis Betrem
Tachinidae
Argyrophylax fumipennis
(Townsend)
(= Cadurcia leefinansi
Baranov)
(=Degeeria albiceps allct.)
Bessa remola (Aldrich)
aAlso recorded from india ex Opisina arenosella pupa (Joy & Joseph 1976)
23
We suggest that the tachinid Argyrophylax fumipennis is potentially a useful
control agent for O. arenosella. As it is a microtype egg-layer, it has a very
high potential fecundity and should he capable of curtailing outbreaks.
Microtype egg-layers of this sort have been used successful1y for biological
control in the past, e.g. A. basifulva Bezzi for the control of coconut spike
moth, Tirathaba complexa Butlet (Pyralidae), in Fiji (Paine 1935) and Cyzenis
albicans (Fallen) for thc control of winter moth, Operophtera brumata (L,)
(Geometridae), in Canada (Embree & Otvos 1984). Although difficulties
in culture have becn encountered for A. basifulva (Paine 1935; O'Connor 1950),
use of electric fans to stimulate mating
these were recently overwme by ｴｨｾ
(Godfray 1985), and a method sim lar to that used for A. basifulva should
suffice for A. filmipennis.
(d)
Dl/rrantia (Peleopoda) arcanella (Busck) (Oecophoridae)
This species occurs in Central America and northern Sou'h America, where
it is polyphagous, its host plants including oil palm, Several other Peleopoda
spp. attack palms in the region The larva lives under a white silk shelter and
rests a1.ong the main vein. It is kept under control by its natural enemies,
of which a polyembrionic encyrtid is the most conspicuous (Genty et al. 1978).
(e)
Furcivena rhodoneurialis Hampson (Pyralidae)
Mariau et al. (1981) record that the larvae of this smaU SpeCi0S from West
Africa feed beneath a fine web on the underside of leaves of oil pa1m. ft i s
normaUy scarce, and the authors report no natural enemies.
(f)
Hedylepta blackburni (Butler) (Pyralidae)
This moth. which at different times has been placed in the genera Lamprosema,
Nacoleia, Omiodes and Phryganodes. is restricted to Hawaii, early reports from
Papua New Guinea heiu5 erroneous. The native host plants are Pritchardia
spp., but it has adapted to coconut, on which it is a sporad.ic pest, particularly
in situations exposed to strong winds, which it is thought hinder the action of
its natura1 enemies. The young larvae feed gIegariously on the leaflet undersurface without perforating it, under a fine webbing of silk, Subsequently
they scatter and form individual tubes by spinning the edges of a leaflet
together.
The recorded natural enemies are summarized in Table 8, and these normally
keep H. blackburni under adequate control. Trathalaflavoorbitalis (Cameron)
is stated by severa1 authors to be the conunonest and most important parasitoid
attacking H. blackburni in Hawaii Jt has a wide host range (Bradley & Burgess
1934; Swezey 1929) and would seem a.potentially useful parasitoid against
Opisina arenosella. Yet T. flavoorbitalis is already reported to occur in Sri
Lanka (Chu & Hsia 1937), although it is not known to attack O. arenosella.
The possibility of biological races deserves exploration and trials with T.
flavoorbitalis from Hawaii could he worthwhile. The second. most frequ·:nt
species is Bracon omiodil'orum (Terry), hut it could be consideted a homologue
of B. brel'icornis ｶ､｡Ｇｾｲｬ
present in India and Sri Lanka.
24
(g)
Herculia nigrivitta (Walker) (Pyralidae)
The larvae of this moth fccd on the old or dead leayes of palms, in galleries
lined with silk and frass (Hutson 1925). In the field the damage can he quite
stliking, but as it is Icstrictcd to the oldest leaves it is of no significance. Palm
leaves, hO\.veyer, are also used for thatch and as such are still suitabb food
material for H. nigrivitta, which 1S occasionally reported as a pest feeding on
nipa (Nipa fruticans), sago (Metroxylon sagu) and areca (Chrysalidococarpus
lutescens) thatch in the Philippines (Morril 1953), Sabah (Cheng 19631 and
Indonesia (Kalshoyen & van der Laan 1981). !ts range extends to Sri Lanka
(Hut:;on 1925).
Table 7.
Hyperparasitoids, etc. of Artona catoxantha (Hampson)
Species
Distribution
Hosts
Hyperparasitoids
Ceraphronidae
Aphanogmus sp.
Ja.va, West Malaysia
Apanteles artonae
Chalcididae
Brachymeria apicicornis
Cameron
B. lasus (Walker)
B. lugubris (Walker)
B. punctiventris Cameron
Dirhinus banksi Rohwer
Hockeria sp.
Java
W. Malaysia
Java, W. Malaysia
Java, W. Malaysia
Java
W. Malaysia
Unspecified (Ferriere
Bessa remota
Unspecified (Ferriere
Unspecified (Ferriere
Unspecified (Ferriere
Unspecified (Ferriere
Java, Sumatra,
W.Malaysia,
Java
W. Malaysia
Java, Sumatra, W.
Malaysia
Argyrophylax fumipennis, Bessa remota
Eulophidae
J.felittobia hawaiiensis
Perkins
Pediobi/ls detrimentos/ls
(Gahan)'
P. ptychomyiae Ferriere
(=P. parl'ltlus auct.)
Nesolynx zygaenarum
(Ferriere)
N. thymus (Girault)
(=Syntomosphyrllln
ob"curiceps (Ferriere)
Tetrastichus sp.
1940)
1940)
1940)
1940)
1940)
Neopiec/rus bicarinatus
Bessa remota
Apanteles artonae, Bessa remota
Solomon Is. W. Malaysia Apallteles artonae, Goryphus sp, Bessa remota
Java, W.Malaysia, Burma, Apanteles artonae, Argyrophylax fumipennis,
Sri Lanka
Bessa remota
W. Malaysia
Unspecuied
Java, W. Malaysia
Java, W. Malaysia
Apanteies artonae, Bessa remota
Apanteles artonae, Argyrophylax fumipennis,
BeSStl remo1P, Goryphus sp.
Eurytomidae
Eurytoma albotibialis
Ashmead'
Java, W. Malaysia
Apanteles artonae
Ichneumonidae
Microtoridea sp.
W. Malaysia
Apanteles artonae
Predators
Clcridae
Callimerus arcufer Cha pin
W. Malaysia
Bessa remola
Pathogens
Fungi
Indet. sp.
W. Malaysia
Bessa remota
Eupelmidae
Euplemus catoxanthae
Ferriere
Anastatoidea brachartollae
Gahan
'Also present in Sri Lanka
25
Table 8. Natural enemies of Hedylepta blackburni (Butler) in Hawaii
Species
Status
Sources
Braconidae
Bracon (Microbracon)
omiodivorum (ferry)
Chelonus sp.
Gregarious larval ectoparasitoid;
introduced from Japan
Egg-larval parasitoid
Simmonds 1929; Swezey
1915
Lever 1969
Chalcididae
Brachymeria lasus (Walker)
(=B. obscurata (Walker)
Polyphagous pupal parasitoid;
introduced from Japan
Swezey 1915
lchneumonidae
Trathala fial'oorbitalis (Cameron) Polyphagous pupal parasitoid:
(0:;; Cremastus hymeniae Yiereck) accidentally introduced; commonest
species
Polyphagous pupal parasitoid
Ecthropomorpha ji/scatar (F.)
Eulimneria blackburni (Cameron)
Pimpla sanguineiceps (Cresson)
(=- Pimpla hawaiicnsis Cameron)
Bradley & Burgess 1934;
Simmonds 1929; Swezey
1929
Swezey 1915
Swezey 1915
Swezey 1915
Trichogrammatidae
Trichogramma fiavull1 Perkins
Egg parasitoid
Swezey 1915
Tachinidae
Frontina sp.
Chaetogaedia sp.
Introduced from USA
lntroduced from USA
Lever 1969
Lever 1969
Cheng (1963) repor ed that pupae in thatch arc parasitized by an Antro'
cephalus sp. (Cha1cid.idae). 1n a small outbœak on rec 'ntly pla'lIed. coconut
palms near Butuan, north Mind.anao Island, Philippines, Cock (1983) fomd
the pupae parasitized by Brachymeria lasus. As d.iscussed above (Section Il
B(e», B. lasus is a polyphagous species already known to attack Opisina
arenosella in India and Sri Lanka. There has been no systematic investigation
of the natural enemies of H. nigril'itta repOl ted in the literature.
One question of immediate interest is to what extent the natural enemies
recorded from O. arenosella in lndia and. Sri Lanka utilize H. nigrivitta as an
alternative host. Furthermore, since H. nigril'itta also occurs in South-East
Asia, where it is of no significancc on growing palms, it may weil have effective
natural enemies there. As any such natural enemies would be attacking a
smaU lepidopteran larva feeding in galleries on coconut leaves, they are
potentially capable of attacking O. arenosella if introduced into Sri Lanka or
India. T 0 asses> this potentiaI, surveys need. to be carried. out to establish
the importance of natural enemies in South-East Asia, which should be linked
that the e
to a simila.. exercise in Sri Lanka and In<'.ia which may ､Ｎｾｭｯｮｳｴｲ｡･
natural enemies are already p 'esent, but attacking H. nigrivitta only, and not
O. arenosel/a.
h)
Homophylotis catori Jordan (Zygaenidae)
The larvae of this species feed. openly on the leaves of COCOIlut and oil palm
in West Africa (Mariau et al. 1981). Although occasional outbreaks occur,
it is usually kept und.er con roI by parasitoids (several Hymenoptera and one
t3Ghinid) and a common fungus. Outbr':2.k: a' e terminated by dry weather.
26
(i)
Loxotoma elegans Zeller (Stenomidae)
This species is similar in biology to Stenoma impressella (Busck) (Section (k)
below), and is widespread in South America. No natural enemies are recorded,
but it is of little pest importance (Genty et al. 1978).
U)
?Odites sp. (Xylorictid.ae)
According to Wood (1968) the small (lOmm) larva of this moth constructs
an open ended. tube about 20 mm long on oil palm. It occurs in both East
(Sabah) and West Malaysia (J. D. Bradley, pers, comm. 1986), but is not at
present known from elsewhere. As Wood (1968) states that it is never
common (atthough damage can usually be found), it seems likely that it is kept
under control by natural enemies. Unfortunately, nothing is known of these.
Since ?Odites sp. feeds in a concealed. manner on palm leaves, it is a possible
source of natural enemies for Opisina arenosella. A survey would have to be
made to find out what (if any) natural enemies keep it under control, before
trials could be carried out to see if Opisina arenosella is a suitable alternate
host, or coconut a suitable habitat.
(k)
Stenoma impressella (Busck) (= S. cecropia Meyrick) ',Stenomidae)
This species feeds on numerous tlees in northern South America, but has only
adapted. to oil palm (Genty 1978). The larvae hollow out a leaflet and
construct a protective sheath of silk, leaf fragments and. frJ,ss, in which pupation
occurs. A bra.conid larval parasitoid, Rhysipolis sp., is common and chalcidoids
and ichneumonids attack the pupae, but these do not prcvcnt outbreaks which
are terminated by hot dry, weather. Bccausc S. impressella has only recently
adapted to oil palm as a host, the p3Iasitoid<; must be associated either with
palms, or with S. impressella on other types of plant.
(\)
Struthocelis semiotarsa Meyrick (Oecophoridae)
This is one of two species of the genus which :.l.ttack oil palm, in northernSouth
America. The larvae live under a silk shetter on the und.erside of the leaflet and
it is controllcd by its natuul cnemies, of which hymenopterous pupat parasitoids
are mentioned. (Genty et al. 1978).
V. DISCUSSION
In considering Opisina arenosella as a target for classical biological control by the
introduction of exotic natural enemies, there are two critical factors to bear in mind:
firstly, O. arenosella is an indigenous pest and secondty, it is an outbreak pest.
The fact that it is an indigenous pest means that natural enemies for introduction
must be sought either from ehewhere within its range, or from species related to the
target. The natural cnemies of O. arenosella hwc been summarized above for Sri
Lanka and Indja (Section) II. There are sorne diffe;'cnces in the llatural enemy
complex between Sri Lanka and India, which are reviewed in Sections III and IV.
With the possible exceptions of Stomatomyia bezziana and Eriborus trochanteratus,
which are not known from O. arenosella in India, and Brachymeria nasatoi, which
27
does not attack O. arenosella in Sri Lanka, there seems little scope for the exchange
of natural enemies between these two countries. As nothing is known of the natural
enemy complex of O. arenosella in Burma, this merits investigation, but we are not
,
optimistic.
The second strategy, that of introducing natural enemies of related species, we
base upon a review of the natural cnemies of smalliarvae of Lepidoptera which feed
in a concealed manuer on leaflcts of palms (Section IV and Table 2). The possible
introductions of species representing guilds djfferent to those already present in
lndia and Sri Lanka are summarized in Table 9. For most of these, it is not known
whether O. arenosella is a suitable alternative host. Because its larva hides within
a gallery it may not be accessible to aU types of parasitoid, e.g. the trials with
Bessa remota describec1. in Section III (a) showcd this 110st to be unsuitable as the
tachinid was unable to oviposit directly onto it. Hence, any species considered
for introduction will have to be first screencd in laboratory and field cage trials to
demonstrate that O. arenosella is an appropriatc host. However, to speculate about
which of these natural enemies might provide effective control of O. arenosella, we
must now consider the fact that it is an outbreak pest.
Table 9. Possible introduction of parasitoids of a guild different from those which already occur in India
and Sri Lanka.
Species
Natural host
Bi%gy
Braconidae
Agathis sp.
Agonoxena phoenicea
Agolloxella argal/la
Solitary larval
endoparasitoid
Egg-lar\ial parasitoid
Agonoxena pyrogral1ll1la
Pupal; ?hyperparasitic role
Chelonus sp.
Encrytidae
Ooencyrtus sp.
Ichneumonidae
Tratha/a fiavoorbitalis (Cameron) Hedylepta blackburni, Ostrinia
nl/bi/alis, etc.
Tachinidae
Actia painei Crosskey
Argyrophylax fumipennis
Townsend
Solitary pupal
endoparasitoid
Agonoxena pyrogral1lma
O\iolarviparous
Zeuxippa catoxamha, etc.'
Microtype egg-Iayer
The outbreaks have been linked to periods of hot dry weather, which it has been
suggested have an adverse effect on the natural enemies. Research presently underway is investigating the possible role of synchronizatioll of host populations, perhaps
due to the action of parasitoids, in the initiation of outbreaks (H. C. J. Godfray &
M. P. Hassell, pers. comm. (1986). In such a system, the pest is nonnally under
satisfactory control by a range of natural enemies, but periodically occurs in outbreaks when the natural enemies are incffective.
For an introduced natural enemy to be useful in such a system it can act either
to prevent an outbreak starting, or to bring an outbreak under control. Without
knowing the precise mechanism of the initiation of outbreaks, one cannot predict
appropriate biological characteristics for an introduced natural enemy to act in the
former manner. If the cause is pureJy climatic factors affecting the indigenous natural
cnemies, then a species preadapted. to the wcather extremes would ben ecessary.
A selection of climate diagrams (Fig. 1.) show that the climates in SouthEast Asia
28
BlOcontro! News and lnformation 1987 Vol. 8 No.
COLOMBO
(7m) 26.6° 2370
CALICUT
(9m) 26.8°
3080
KUALA LUMPUR
(39m) 27.4° 2365
TRINCOMALEE
(7m) 27.6° 1641
HAMBANTOTA
(18m) 26.8° 1097
BANGALORE
(92Om) 23.00 864
RANGOON
(Sm) 27.2° 2633
PASURUAN
26.7° 1316
LAUTOKA
25.5° 1771"
Fig.l Climate diagrams for areas where Opisina arenosella Walker oceurs, and of sorne potential
source areas for natural enemies. Colombo represents the areas of western Sri Lanka where outbreaks
seldom occur. Trincomalee in the East and Hambantota in the Soutjl of Sri Lanka have seasonal
dry periods and periodic outbreaks. In India the distribution of O. aIenoseJla centres on Kerala State
exemplified by Calicut on the west coast.' Both Calicut and Bangalore (central southern India) show
a very pronounced dry season. as does Rangoon (Burma). In contras! West Malaysia and western
Java exemplified by Kuala Lumpur are much more equable, alt\1ough parts of eastem Java (e.g.
Pasuruan) show a pronounced dry season. Lautoka is in western Viti Levu, Fiji Is., one of the drier
parts of the archipelago and has a moderate dry season.
The figure in brackets following the locality is the altitude of the recording station in metres;
this is followed by the average temperature in oC, and the average annual rainfall in mm; the
horizontal axis represents the months of the year. the leCt-hand axis is the monthly iRean temperature
(thin line) graduated (from) O"C in 1000C units; the right-hand axis is the rainfall (thick tine) graduated
from 0 in 20 mm units up to 100 mm, above which (solid black area) the scale is reduced to one
tenth of this; dotted areas represent arid periods while the hatched areas represent humid periods.
Redrawn from Walter, Hamickell and Mueller-Dombois (1975)
to Fiji, which is one potential source area, are mostly more equable than those of
the problem areas in Sri Lanka. However, parts of East Java are subject to a dry
season rather more pronounced than that of the outbreak areas of Sri Lanka (Fig. 1,
Pasoeroen). Natural enemies from this part of Java should be able to adapt to
Sri Lankan conditions, although material from other parts of the region could have
problems adapting to the dry areas of Sri Lanka. Without more detail on the
distribution of the palm feeding species reviewed from West Africa and the Neotropical region, we cannot comment on climatic match with these sources. It
seems likely that a reasonable climatic match could be found in the Neotropics, and
there may weB be effective natural enemies of the palm defoliators there. A wide
ranging survey would be necessary to develop this potential source.
If synchronization plays a key role in the initiation of outbleaks, then a natural
enemy which will not be affected by this condition would need to be either synchronized with the host, with the same length of life cycle or, as in the case of Pediobius
parvulus (Ferriere) used for the control of Promecotheca coeruleipennis in Fiji, able
to attack the host throughout most of its life cycle and survive the brief period when
no host material is available (Taylor 1937). The life cycle of O. arenosella takes
about ten weeks, but there are no detailed studies of the biology of any of
the parasitoids considered for introduction in Table 9. Based upon the known life
history of the parasitoids we can predict that the life cycles of the Hymenoptera are
likely to be significantly sholter than that of the host, but if the. tachinids
can parasitize sufficiently young larvae, their life cycle may not be that much shorter
than that of the host (due to a preoviposition period). Thus, to introduce natural
enemies to prevent outbreaks, without knowing the precise cause of those outbreaks,
will be a tlial and error ex rcise, which in light of present knowledge of the available
natural enemies is unlikely to be effective.
On the other hand to cUItail an outbleak once it has started, rather different
biological charactelistics seem appropriate for an intlOduœd natural enemy. These
should maxirnize the reproductive potential and the rapidity with which it can
respond to the available hosts. Further, if the outbreak is synchronized, the control
agent must be able to survive any periods when no hosts are available. For these
reasons we suggest that a parasitoid with a very high fecundity, the ability
to maintain itself on alternative hosts and an appropriate life cycle would he ideal.
chalacters:
One natural enemy amongst those reviewed stands out as having ｾｳ･ｨｴ
Argyrophylax jumipennis. This species is a common parasitoid of Artona catoxantha
on coconut in Java; it is a microtype eggIeayer with a likely fecundity of around
1000; it is known to use other hosts by laying Hs eggs heside feeding damage, the
ploblem of access to the O. arenosella larva in its gallery is resolved, of the species
considered for inrtoduction in Table 9, it probably bas the longest potential life
cycle and it bas been noted as very common in synchronized outbreaks of A.
catoxantha in Java, when parasitoids play the princiP'B1 role in curtailing the
outbreaks.
VI. RECOMMENDATIONS
1.
Releases of Eriborus trochanteratu5 and Stomatomyia in India merit further
effort.
2.
A survey of natura! enemies of O. arenosella hould be made in Bangladesh
and Burma to assess this potential source of biological control ｡ｧｾｮｴｳＮ
3\
3.
Argyrophylax fumipennis should be obtained [tom East Java and tested in
the laboratOlY against O. arenos lla. If suitable, A. fumipennis should ｢ｾ
released for the control of O. arenosella.
ACKNOWLEDGEMENTS
Some years ago Dr. Sudha Nagarkatti, then of the Indian Centre for Agricultural Research,
asked the first author about possible exotic hosts for potential biological control agents
for Opisina arenosella. At the time, she got a rather negative answer, but the seed was
sown and it has grown into this review. It is a pleasure to acknowledge this early stimulus.
This review was prepared as part of a collaborative programme between the Silwood
Centre for Pest Management (SCPM) and the Coconut Research Institute (CRI) of Sri
Lanka, funded by EEC contract no. TSD-A-286 under the direction of Professor M. J.
Way of SCPM and Dr. P. Kanagaratnam of CRI.
The authOl-'s would like to thank Dr. H. C. J. Godfrayand Professor M. P. Hasselt
ofImpetial Coltege for helpful discussions which contributed to this review. Taxononists
of the CAB International Institute of Entomology provided advice on taxonomic useage
and information sources, including Dr. Z. Boucek (Cha1cidoidea), Dr J. D. Holloway
(Lepidoptera), Dr. A. Walker (Braconidae) and Dr. 1. M. White (Tachinidae).
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36
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37
Joy, P.J.; Narendran, T.C.; Joseph, K.J. (1974) New tecOlds of Brachymeria lasus (Walker)
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38
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39
Narendran, T.C.; Mohamed, V.V.K; Abdurahiman, V.C.; Kumar, V.V.S. (1980)
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40
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41
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42
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44
Matthew Cock