A critical review of the literature on the pest Erionota spp.(Lepidoptera, Hesperiidae): taxonomy, distribution, food plants,early stages, natural enemies and biological control

Matthew J. W. Cock*

Address: CABI Europe – UK, Bakeham Lane, Egham TW20 9TY, UK

*Correspondence: Matthew J. W. Cock. Email: m.cock@cabi.org

Received: 2 December 2014

Accepted: 15 February 2015

doi: 10.1079/PAVSNNR201510007

The electronic version of this article is the definitive one. It is located here: http://www.cabi.org/cabreviews

g CAB International 2015 (Online ISSN 1749-8848)

Abstract

The classification, taxonomy, distribution, spread, food plants, early stages, natural enemies

and biological control of South-East Asian Erionota spp. (Hesperiidae, Hesperiinae, incertae sedis)

are reviewed with specific attention to Erionota thrax (Linnaeus), Erionota torus Evans and Erionota

acroleuca (Wood-Mason and de Niceville) (=E. hiraca (Moore)). Erionota thrax and E. torus are

shown to be pests of Musa spp. but not palms, while E. acroleuca is a pest of various palms. Where

they overlap in distribution in mainland South-East Asia and the Southern Philippines, E. thrax

and E. torus have not been distinguished in the economic literature. The species which became

established in Mauritius around 1968 and subsequently the target of a successful biological control

programme is shown to be E. torus, and not E. thrax as hitherto reported. Research gaps are

identified, including the need to find diagnostic features or tools to separate the immature stages

of the three species in order to confirm these conclusions, and clarify the ecology and pest status

of the pest species.

Keywords: Erionota thrax, Erionota torus, Erionota acroleuca, Banana, Musa, Palm, Arecaceae.

Review Methodology: Because these pests are large butterflies, there is an extensive literature based on regional butterfly books and

lists as well as the applied entomological literature. The author attempted to track down and examine all available information on the

genus Erionota, particularly the applied literature dealing with the biology, pest status and control of the pest species. The taxonomic

work of Evans [1] and the CABI Crop Protection Compendium datasheet for E. thrax were important starting points. This was

supplemented by searching CAB Abstracts, internet searching, and consulting selected reference works dealing with butterflies and

pests in the region. The information sources were traced back to their original publication whenever possible. The author familiarised

himself with the biology of the two banana skippers (E. thrax and E. torus) by examination of field material in West Malaysia, which

helped considerably in interpreting the literature for this review. The Erionota spp. material in the insect collections of the Natural

History Museum, London, the Hope Entomological Collections (Oxford University Museum) and the Forest Research Institute of

Malaysia were examined to check for locality records, voucher specimens and biological information; the genitalia figures of Inoue and

Kawazoe [2] enabled several misidentifications in these collections to be recognised by examination of the externally visible genitalia.

The taxonomy of the natural enemies and plant names were checked against key online databases. As indicated in the

acknowledgements a variety of informed individuals were consulted on specific points.

Introduction

Erionota thrax (Linnaeus) is well known as ‘the banana

skipper’ or ‘banana leaf roller’ in South-East Asia, and

as an introduced pest in various Pacific territories. It

is amongst the largest members of the family Hesperiidae

(Figure 1). A second species, Erionota torus Evans

(Figure 2), has a very similar biology, also occurs in South-

East Asia and as an introduced in Mauritius, Taiwan and

Japan (Okinawa). Both have been the target of successful

biological control programmes where they have been

introduced. A third, Erionota acroleuca (Wood-Mason

and de Niceville) (Figure 3) is similar in appearance but

the larvae feed on palms. All three overlap in much

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CAB Reviews 2015 10, No. 007

of their distribution in South-East Asia and there is an

extensive literature which indicates considerable confu-

sion as to the Erionota spp. concerned. In this review,

I attempt to untangle this literature, correct some errors,

and identify aspects needing further attention.

Classification

Erionota Mabille is a genus in the subfamily Hesperiinae,

family Hesperiidae. In his work on the Hesperiidae of

Europe, Asia and Australia, Evans [1] grouped seven

Figure 1 Pinned adults of E. thrax thrax, reared on Musa sp., Luzon Is, Philippines; male on the left, female on the right;dorsal surface above, ventral surface below; life size, the scale bar is 1 cm.

Figure 2 Pinned adults of E. torus; male on the left, female on the right; dorsal surface above, ventral surface below; lifesize, the scale bar is 1 cm. Male: SE China, Macao, 20 Aug 1904, J. C. Kershaw; female: west Malaysia, Negri Sembilan,Seremban, Feb 1914, V.G Bell and J. W. B. Bell (specimens in Hope Entomological Collections, Oxford UniversityMuseum). The specimens illustrated are less dark than those of the other two Erionota spp. illustrated because they havefaded with age. Erionota torus often cannot be reliably distinguished from E. thrax thrax (Figure 1) without examination of thegenitalia.

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2 CAB Reviews

Oriental genera as the Erionota sub-group within the

Plastingia group of Hesperiinae. In their work on the

genitalia of the Hesperiidae of South Vietnam, Inoue and

Kawazoe [2] suggested affinities between Lotongus Distant

([1] Plastingia subgroup), Gangara Moore, Erionota, Matapa

Moore ([1] Erionota subgroup), Unkana Distant, Hidari

Distant and Pirdana Distant ([1] Unkana subgroup)

based primarily on characters of the male genitalia.

Although Warren [3] treated Erionotini as a provisional

tribe of Hesperiinae in his Ph.D. thesis on the classification

of the Hesperioidea, this was a much enlarged group of

genera, which in his subsequent co-authored publications

[4, 5] were placed incertae sedis.

Of the seven genera that Evans grouped in his Erionota

subgroup, no information has been located on the early

stages of species of Zela De Niceville, Ilma Swinhoe, Ge

De Niceville or Pudicitia De Niceville, but there is infor-

mation on Gangara, Erionota and Matapa. Gangara thyrsis

(Fabricius) feeds on various palms and the early

stages have been illustrated in Igarashi and Fukuda [6] and

Cock et al. [7] among others. Matapa aria (Moore) is

a bamboo-feeder [8], and the early stages are illustrated

by Bascombe et al. [9] and Saji et al. [10]. Erionota spp. feed

on palms (Arecaceae), Musa spp. (Musaceae, bananas and

plantains) and some related Zingiberales. Igarashi and

Fukuda [6] illustrate the early stages of E. torus, E. thrax

and E. grandis (Leech), while Bascombe et al. [9] also

illustrate the early stages of E. torus. The leaf shelters

of at least some members of all three genera are note-

worthy for being tubes made by rolling palm leaflets

or cut sections of Musa leaves into a tube, either in a

spiral (Gangara, Matapa) or in a roll (Erionota). This

behaviour separates them from all other Hesperiinae

familiar to the author, and supports their recognition as a

distinct group.

There is information available on the early stages of

the additional genera that Inoue and Kawazoe [2] sug-

gested could be grouped with Erionota. Lotongus calathus

(Hewitson) has an unusual biology as documented by

Igarashi and Fukuda [11]. The distinctive ova are flattened

with a grooved ring around the top. They were found laid

singly, and the newly hatched larvae make individual

leaf folds on their unidentified palm seedling food plant.

The second instar larvae then aggregate to make a

communal shelter, which is always co-occupied by a small

ant, Dolichoderus bituberculatus Mayr, which does not

attack the larvae. The mature larva has a four-lobed black

marking on the face, reminiscent of those of some African

Artitropa spp. [12, 13]. The larvae pupate in a stout

cocoon within the shelter, and the pupa is pale brown

with no striking features. Igarashi and Fukuda [11] docu-

ment the life history of Unkana ambasa (Moore) and

Unkana mytheca (Hewitson) on Pandanus odorifer (as

P. odoratissimus, Pandanaceae). The ova are smooth and

domed, similar to those of many other Hesperiinae;

the larvae are translucent grey-green with dark, oval

heads, and a dark anal plate in the case of U. ambasa;

the leaf shelters are cylindrical and made by rolling the

leaf along the axis of the mid-rib; the pupae are brown,

the proboscis reaching the cremaster. Rearing revealed

Figure 3 Pinned adults of E. acroleuca apicalis; male on the left, female on the right; dorsal surface above, ventral surfacebelow; life size, the scale bar is 1 cm. Both collected as larvae on Caryota sp. (Arecaceae), Peninsular Malaysia, Selangor,Kedong, Forest Research Institute of Malaysia, 2 Apr 1988, L. G. Kirton, rearing reference B:5 (specimens in ForestResearch Institute of Malaysia). The extent of the white apical forewing apex is variable, especially in the female, and whenit is reduced this species is difficult to distinguish from E. thrax (Figure 1) and E. torus (Figure 2).

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Matthew J. W. Cock 3

that the all-female species Erionota harmachis (Hewitson)

is a synonym of the all-male species U. mytheca [14],

indicating affinities in adult structure characters. Hidari

irava (Moore) makes leaf shelters between leaflets of

its palm hosts where small numbers of larvae cohabit

[11, 15, 16]. Pirdana hyela (Hewitson) and Pirdana distanti

(Staudinger) feed on Dracaena spp. (Asparagaceae)

[11]. The ovum is strongly ribbed; the head and body of

the larva of P. hyela superficially resembles that of an

Artitropa sp. [12, 13], while that of P. distanti has a

similar head, but the body is dark with a yellow dorsal

line and yellow dorsolateral transverse bars; leaf shelters

are made by rolling the food plant leaf longitudinally; the

pupa has a short, blunt, frontal spike. The rather diverse

life histories of representatives of these genera do not

suggest any obvious affinities with Erionota, unlike that

pointed out for Gangara and Matapa above.

Taxonomy

The checklist in Table 1 presents the current position.

The species of immediate concern as pests are E. torus,

E. thrax and E. acroleuca. However, as some or all the

other species also feed on Musacaeae and Arecaceae as

larvae, they are all of concern as potential quarantine

pests.

Linnaeus [22] described Papilio thrax from Java, referring

to an un-named image in Clerck (36, plate 42.2). Honey

and Scoble [37] explain that there are two specimens

of E. thrax in the Linnean Collection, one of which has

been designated the lectotype, but as Mabille [20] also

pointed out, Clerck’s [36] image is of a different, unre-

lated species (a South American species of the genus

Thracides).

Evans [21] described E. torus from Sikkim and gave

characters to separate the adults from E. thrax based

on appearance and male genitalia. In his catalogue of the

Hesperiidae of Europe, Asia and Australia [1] he added his

diagrammatic figures of the male genitalia. Males of E. torus

(Figure 2) can usually but not always be distinguished

from those of E. thrax (Figure 1) by the more rounded

distal margin of the forewing in E. torus compared to

the straighter margin and apically more pointed forewing

of E. thrax. Separation of the two species, particularly

females, is only reliable if based on examination of the

genitalia ([38], H. Chiba, personal communication, 2014;

author’s observations). It is fortunate that the male

and female genitalia of both species have been accurately

documented and drawn by Inoue and Kawazoe [2] and

available on the internet. Corbet et al. [39] include

figures of the male genitalia of both species, and the male

genitalia and female venation of E. torus are illustrated

by Bascombe et al. [9] from Hong Kong. The diagnostic

characters of both male and female genitalia can usually be

seen by brushing off the scales of the abdomen tip using a

short, fine paint brush, obviating the need for traditional

dissection. In particular, the strongly upturned, more

heavily chitinized apical process of the valves of male

E. torus, compared with the simple, blunt apical process

of E. thrax can often be distinguished. A recent student

thesis yet to be published [40] concluded that ‘the banana

skippers, E. thrax and E. torus, are a monophyletic group

which represent a single host shift from Arecaceae or

Zingiberaceae to Musaceae’.

There is some confusion regarding the correct name

for E. acroleuca [19]. It was first described by Wood-

Mason and de Niceville [25] in August 1881 as Telegonus

acroleucus in the abstract of a paper which was not pub-

lished in full until December that year when the combi-

nation Hesperia acroleuca was used [26]. In the interim, in

September, Moore [27] independently described the same

species as Hesperia hiraca. The description in the abstract

[25] gives the new species name, indicates that it is new

species, gives a brief description and comparison with

E. thrax, and indicates many specimens from ‘S. Andaman’,

i.e. it meets the requirements for the publication of the

new name and so has precedence over Moore’s name.

Gender agreement has been applied under the ICZN [41],

i.e. E. acroleuca is correct rather than E. acroleucus fol-

lowing the treatments of De Jong and Treadaway [17] and

Xue and Lo [19]. The male of E. acroleuca has the apex

of the forewing upper side white, which distinguishes it

from the males of E. thrax and E. torus. The female is

generally smaller than those of E. thrax and E. torus, but

the white patch at the forewing apex is variable, and

some specimens are not reliably distinguishable except

by examination of the genitalia. Fortunately, Inoue and

Kawazoe [2] again provide good drawings of both male

and female genitalia of this species (ssp. apicalis).

The authorship of ssp. apicalis also needed clarification.

Xue and Lo [19] showed that when Evans [29] published

the name as var. apicalis, this made it an unavailable

infraspecific name under Article 45.5.1 of the International

Code of Zoological Nomenclature [41], and it should

take the authorship from the first publication to use it as

a valid name, i.e. De Jong and Treadaway [17], as listed in

Table 1.

Astictopterus harmachis Hewitson was described from

Sumatra [42], and was subsequently treated as a species

of Erionota [1, 21]. However, it is now known to be the

female of U. mytheca, which had previously only been

known from the male [11, 14].

Distribution and Spread

There are four subspecies of E. thrax recognized in

South-East Asia: E. thrax thrax widespread from north-east

India to the Philippines, Sulawesi and Java, E. thrax mindana

Evans in the southern Philippines but overlapping with

E. thrax thrax in the northern Philippines, E. thrax alexandra

Semper from upland northern Luzon and E. thrax has-

drubal Fruhstorfer from Kep. Sula (Mangole, Sanana),

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Table 1 Checklist of Erionota spp. based mostly on Evans [1], and in light of De Jong and Treadaway [17, 18] and Xue and Lo [19]

GenusSpeciesSubspeciesSynonym

Original genus of description(or species if describedas subspecies) Type locality1

Erionota Mabille, 1878 [20]torus Evans, 1941 [21] Erionota Kurseong, Sikkim [India]thrax Linnaeus, 1767 [22] Papilio Java [Indonesia]ssp. thrax Linnaeus, 1767 [22]ssp. mindana Evans, 1941 [21] Erionota Mindanao [Philippines]ssp. hasdrubal Fruhstorfer, 1910 [23] Erionota thrax Batjan [= Bacan, Maluku, Indonesia]ssp. alexandra Semper, 1892 [24] Erionota Vigan, Nordwest Luzon, Philippines

acroleuca Wood-Mason & De Niceville, 1881 [25, 26] Telegonus S[outh] Andaman [Islands], Indiassp. acroleuca Wood-Mason & De Niceville, 1881 [25, 26]=acroleucus Wood-Mason & De Niceville, 1881a [25] Hesperia [South Andaman Islands, India]=hiraca Moore, 1881 [27] Hesperia Andaman Is., India=lara Swinhoe, 1890 [28] Teligonus [sic] Nicobar Is., India [locality error]

ssp. apex Semper, 1892 [24] Erionota thrax var. Luzon, Philippinesssp. apicalis De Jong & Treadaway 1992 [17] Erionota acroleuca Bassein, Burma [= Pathein, Myanmar]=apicalis Evans 1932 [29] Erionota thrax thrax var. Bassein, Burma [= Pathein, Myanmar]

ssp. sakita Ribbe, 1889 [30] Casyapa thrax Ost-Celebes [East Sulawesi, Indonesia]=toradja Fruhstorfer, 1911 [23] Paduka Ost-Celebes [East Sulawesi, Indonesia]

surprisa De Jong & Treadaway, 1992 [17] Erionota Leyte, Philippinestribus Evans, 1941 [21] Erionota E. Celebes [East Sulawesi, Indonesia]hislopi Evans, 1956 [31] Erionota L. Tahan, Pahang, Malaya

[Peninsular Malaysia]grandis Leech, 1890 [32] Plesioneura Chang Yang [Yichang, Hubei] Chinassp. grandis Leech, 1890 [32]ssp. alsatia Evans, 1939 [33] Erionota Likiang, Yunnan, China

sybirita Hewitson, 1876 [34] Hesperia Singapore=mabillei Staudinger, 1889 [35] Erionota Palawan, Philippines

1Information in square brackets has been added where it was not included in the original description.

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North Moluku, Indonesia [1, 18, 43]. Of these, only

E. thrax thrax is known to have spread beyond its indi-

genous range.

In the Pacific E. thrax thrax spread to Guam by 1957

[44] and Saipan in the Northern Mariana Islands by

the late 1960s [45], to Oahu, Hawaii by 1973 [46], to

Irian Jaya, east Indonesia some years before 1983 [47],

to mainland Papua New Guinea (PNG) in 1983 [45],

to New Britain, PNG, in 1991 [48], to Duke of York

and New Ireland islands, PNG, by 1994, and possibly to

Bougainville, PNG, about this time [47]. Muniappan [49]

refers to the original introduction into Guam as being

from the Philippines, based on both localities having

American air force and naval bases, and the heavy civilian

traffic between the Philippines and Guam (R. Muniappan,

personal communication, 2014). Within the Hawaiian

Islands it spread from Oahu to Kauai and Maui in 1974 and

Hawaii, Molokoi and Lanai in 1975 [50, 51].

The records from Guam, Saipan and Hawaii have been

referred to incorrectly as E. torus, e.g. Igarashi and Fukuda

[11], Opler and Warren [52] and perhaps others. The

origin of this error and its correction are outlined in

Pelham [53]. Kawazoe and Wakabayashi [54] illustrated

E. thrax as E. torus and stated that it occurs in Hawaii

and Guam. Snyder and Kumon [55] and Fukuda and Nicho

[56] clarified that E. thrax is indeed the species occurring

in Hawaii, not E. torus.

Although Parsons [57] treats the population that

spread into PNG as E. thrax hasdrubal, the specimen that

he illustrates is E. thrax thrax, based on the characters in

Evans [1, 21], and is treated as such by Waterhouse and

Norris [45] and Sands et al. [58]. Apart from the records

from New Guinea, it is unclear to what extent E. thrax of

either subspecies thrax or subspecies hasdrubal has spread

through eastern Indonesia, either naturally, or with acci-

dental human assistance.

Erionota thrax was first reported in Mauritius in 1970

[59], although it was suspected to have been introduced in

1968 [45]. The author has examined the three Mauritius

specimens in the Natural History Museum, London

(NHM), collected by P. M. H. Davis at Black River in 1979,

and reported in Davis and Barnes [60], and finds that all

are E. torus, based on the visible external genitalia. The

original identification reported in Monty [59] was made

at the NHM, but only female specimens seem to have

been provided, and at that time the distinctive female

genitalia of E. thrax and E. torus had not been documented.

Although it is not impossible that both species were

accidentally introduced into Mauritius, in the absence

of any specimens of E. thrax, it seems safe to assume that

only E. torus is present.

Veenakumari and Mohanraj [61] documented the

first record of E. thrax thrax from the Andaman Islands,

although it would be desirable to confirm this identifica-

tion by examination of the genitalia (K. Veenakumari,

personal communication, 2014). The author was unable

to locate any specimens in the NHM to do so. Erionota

thrax was recently reported from Palpa District, central

southern Nepal [62], but this needs confirmation in

view of Smith’s [63] statement that only E. torus oc-

curs in Nepal (R. Muniappan, personal communication,

2014).

Erionota torus was more restricted in distribution, but is

now widely introduced. It is indigenous in northern India:

an early report of E. thrax as common around Dehradun

(=Dehra Dun) at the base of the Western Himalaya [64]

is considered to refer to E. torus since only specimens of

this species are present in the NHM from Dehra Dun [1].

It is also found in Nepal [63, 65] and Northeastern

India to Southern China and Peninsular Malaysia [1], but it

spread to Mauritius around 1968 (reported as E. thrax, but

see above), Southern Japan in 1971 [66], Taiwan in 1986

[67, 68], the southern Philippines probably in the early

1980s (Cebu, Dinagat, Leyte, Mindanao, Negros, Samar;

[18]), and has just been found in the Western Ghats

of India [69]. There is a single male specimen labelled

Sarawak in the NHM, but the absence of any other

records from Borneo suggests its presence there should

be considered unconfirmed. It does not seem to be pre-

sent in Java or the rest of Indonesia.

Erionota acroleuca occurs in the Andaman Islands

as the nominate subspecies. Although Swinhoe [28]

described a synonym from the Nicobar Islands, Evans [1]

treats this as an erroneous locality. Erionota acroleuca apex

is restricted to the Philippines [17, 18]. Erionota acroleuca

apicalis was formerly treated as a synonym of E. acroleuca

apex, but De Jong and Treadaway [17] established that it is

a separate subspecies found from Myanmar and Northern

Vietnam, through the Malaysian peninsula to Borneo,

Sumatra, Java and east to West Nusa Tenggara. Finally,

E. acroleuca sakita is found on Sulawesi, and the islands to

the east of Central Sulawesi [1, 43].

It likely seems that the major translocations of E. thrax

and E. torus were mediated by aircraft flights carrying

infested plants or planting materials, infested leaves or

adult butterflies. Ova occasionally occur on fruit in bun-

ches, and so may also be transported internationally [70],

but the challenge for the hatching larvae to reach Musa

spp. leaves after arrival is very considerable. Military flights

have been implicated or suggested as the route for

the spread to Mauritius and Hawaii: E. torus is ‘thought to

have been introduced in military aircraft based in Malaysia

in 1968’ [45], and E. thrax was discovered in Hawaii at

Hickam Air Force Base, Oahu [46]. The spread of E. thrax

thrax to Guam has been linked to American military bases

on Guam and the Philippines, as well as heavy civilian

traffic between the two (R. Muniappan, personal com-

munication). The fact that Guam and Saipan both had

strong military bases in the 1950s suggests how the sub-

sequent introduction to Saipan may have been facilitated,

rather than to other islands in the region. During its

outbreak phase in PNG, E. thrax spread throughout the

PNG mainland within 6 years, moving several hundred

km per year [58]. This would have been due to a mixture

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of natural dispersal of gravid females, and accidental man-

assisted transport of planting materials, leaves and adult

butterflies.

What is less clear is the mechanism of spread between

relatively nearby islands, e.g. through the Hawaiian Islands

and from New Guinea to New Britain. Waterhouse

and Norris [45] speculate that ‘the water gaps between

Southern PNG and Northern Australia are probably well

within the flight range of adults. . . . Southwest Pacific

islands will be at risk if inter-island movement of fresh

banana leaves is not prohibited. In any case adults may be

able to fly to nearby islands and establish themselves.’ This

speculation by Waterhouse and Norris [45] has subse-

quently been presented in a pest risk assessment as

‘E. thrax has spread from Oahu to the other Hawaiian

Islands flying over water for distances of up to 150 km

[45]’ [71] – 150 km being approximately the distance

between PNG and the northern tip of Australia. No

published evidence to support the possibility of gravid

females flying these distances has been found. The fact

that E. thrax and E. torus have not spread more widely

already, e.g. within the parts of Indonesia where they

do not occur naturally, or within the Northern Mariana

Islands, suggests that such long distance flights over sea do

not occur. Similarly, the evolution of isolated subspecies

of E. thrax (above) indicates a species that is not highly

mobile.

Inter-island movement of boats seems a more likely

pathway of introduction, as Sands et al. [48] discuss:

‘Lights on board boats are considered to be a means for

attracting the crepuscular adult E. thrax and allowing them

to be carried from one island to another. A specimen in

the Natural History Museum, London, was collected

on board a ship in 1929, near Seram (less than 200 km

west of New Guinea), outside of its native range. From

the distinctive pattern of spots beneath the hind wings,

this specimen probably came from Java. A specimen from

Port Moresby, PNG in 1961 and referred to by Parsons

[57] probably travelled to PNG on board a ship since this

specimen pre-dated first establishment of E. thrax in

North-western PNG by 22 years.’

In view of the foregoing discussion and the confusion

that has arisen, it is worth recognizing the possibility that

in the last 65 years or so since Evans [1] completed his

review of the Hesperiidae of the region, E. torus could

have spread to areas where only E. thrax thrax was known

(or vica versa), and easily been overlooked. This possibi-

lity merits checking with recent material, for example

in the island of Borneo where there is already a single

record of E. torus, and Sumatra and Java which are in

relatively easy reach of Peninsular Malaysia.

Food Plants

I have reviewed much of the literature on food plants

of Erionota spp. and find considerable confusion and

indications of errors. Accordingly, I present my analysis

in some detail below. Erionota thrax thrax and E. torus are

similar in adult appearance [1, 21] and life history [6, 39],

and where their ranges overlap, have almost certainly

been confused – with each other and with other Erionota

spp., particularly E. acroleuca. Reports from localities

where only E. thrax or only E. torus occur are more likely

to be reliable, but where both occur there will be

confusion between the two. The situation is rendered

more difficult to interpret by the habit of many authors

of compiling food plant records from multiple sources

without attribution, so that it is not clear whether any

particular observation, such as a food plant record, is an

original observation or repeated from an earlier source,

which may be misidentified, represent a different sub-

species, or be from a different geographical area [72].

Food plants of E. thrax

I have found reports on the food plants and biology of

E. thrax thrax only, and none on the other subspecies of

E. thrax, although it seems likely that they will have a simi-

lar food plant range. In its indigenous range, many refer-

ences (e.g. [15, 39, 73–77]) indicate that E. thrax feeds

primarily on Musa spp. (Musaceae), especially banana

(M. � paradisiaca, = M. sapientum and M. paradisiaca) and

M. textilis (abaca or Manila hemp). Sands et al. [58] sur-

veyed E. thrax in PNG when it first spread through the

island of New Guinea. They report collecting immature

stages of E. thrax from Musa � paradisiaca, M. textilis, other

Musa spp. (including sections Australimusa and Eumusa)

and several varieties. Waterhouse and Norris [45] point

out that in its introduced range, E. thrax thrax is known

only from cultivated and wild species of banana (based on

personal communications of experienced entomologists

who worked on the introduced pest), although their

observation to this effect from Mauritius is actually based

on E. torus. However, records from other recorded food

plants, especially in the older literature, need critical

examination. It is possible that different species of Erionota

are responsible for the records on bamboo and palms.

Probably the oldest record of E. thrax as a pest

of banana (pisang, Musa paradisiaca) is that of Horsfield &

Moore [78] in A Catalogue of the Lepidopterous Insects in the

Museum of the Hon. East-India Company. As these obser-

vations were from Java, where E. torus is not reported,

they should be reliable. Soon afterwards, Distant [79] in

Rhopalocera Malayana, repeats Horsfield & Moore’s [78]

observation on the food plant, although his illustration of

the adult is E. torus according to Evans [1]. In Lepidoptera

Indica Vol. X, Swinhoe [80] gives the food plants as ‘wild or

cultivated Musa’. Although Evans [1] indicates that the

species illustrated is E. thrax, the observations could relate

to either E. thrax or E. torus or both. Duport [81] refers to

E. thrax as a banana pest in Indochina, and this could refer

to both E. thrax and E. torus. Williams [82] describes the

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Matthew J. W. Cock 7

biology on banana on Luzon Island, Philippines, which

should be correctly associated with E. thrax. Beller and

Bhenchitr [83] include E. thrax thrax as a banana pest in

their list of insect pests of Thailand (Siam); this could be E.

thrax or E. torus.

There are many other food plant references compiled

in Robinson et al. [84] with sources, and in Vane-Wright

and De Jong [43] and CABI CPC [85] without sources.

These include various palms (Arecaceae), bamboos

and grasses (Poaceae), Heliconia spp. (Heliconiaceae),

Curcuma angustifolia (Zingiberaceae) and traveller’s palm

(Ravenala madagascariensis, Strelitziaceae), which are

assessed below.

Robinson et al. [84] cite Bhumannavar et al.’s [86]

Insects of Agricultural Importance in Andaman and Nicobar

Islands that E. thrax feeds in rolled leaves on the intro-

duced R. madagascariensis. Veenakumari et al. [87] made

new observations and reported the food plants of E. thrax

thrax in the Andaman Islands as only Musa�paradisiaca

and R. madagascariensis. As Veenakumari et al. [87] sepa-

rate their food plant records of E. thrax thrax from those

of E. acroleuca, and the leaf structure of R. madagascariensis

is suitable for leaf rolling as practiced by E. thrax, this

record is accepted here, even though Sands et al. [58] did

not find immature stages of E. thrax on this food plant in

PNG. Mau and Kessing [88] give Strelitzia (Strelitziaceae)

as a food plant genus in Hawaii, but this does not seem to

have been formally published. Nevertheless, it is a possible

food plant, although perhaps as a spill-over effect (below).

Similarly, Mau and Kessing [88] list the genus Heliconia

as a food plant in Hawaii. In PNG, the ornamental native

species Heliconia papuana is a widespread host for E. thrax

thrax (F. M. Dori and D. P. A. Sands, personal commu-

nication, in [47]), although the introduced Neotropical

Heliconia bihai is not, and is not in the laboratory [58].

Igarashi and Fukuda [6] list C. angustifolia (wild or Indian

arrowroot; Zingiberaceae) as a food plant. No other

source has been located for this record (which is repeated

in 43), but given that C. angustifolia has large leaves, similar

to those of Musa, Ravenala and Heliconia, it seems a pos-

sible food plant. A record of Canna (Cannaceae) as a food

plant genus in Hawaii [88] could be interpreted in this

way, but could also be due to a spill-over effect.

At this point it is worth noting that Musaceae

and Strelitziaceae are closely related families in the

order Zingiberales, while Heliconiaceae, Cannaceae and

Zingiberaceae although also placed in Zingiberales are

less closely related [89, 90]. Oligophagous species are

mostly, but not always, more likely to feed on species

closely related to their main food plant, than on those less

closely related (e.g. [91]). The remaining food plant

records are of species from other plant orders, and

correspondingly less likely.

The original records of E. thrax [thrax] feeding on

Arecaceae seem to be those of Piepers and Snellen [73]

from Java, who in addition to Musa spp., list Cocos nucifera

(coconut), Rhapis excelsa (as R. flabelliformis), Metroxylon

sagu (sago palm), Arenga pinnata (as A. saccharifera) (sugar

palm), and based on a verbal report Elaeis guineensis (West

African oil palm) as food plants. Keuchenius [92] and

Leefmans [93] also report E. thrax as a coconut pest in

Indonesia (Dutch East Indies). However, since Piepers

and Snellen [73] treat E. acroleuca as a synonym of E. thrax,

it is suggested here that the records from palms are

more likely to relate to E. acroleuca, which has since

been documented as a palm pest (below). Specimens from

P. C. T. Snellen’s collection (which includes material from

M. C. Piepers) are in the Naturalis Biodiversity Centre,

Leiden, Netherlands but they cannot be examined

at present during rearrangements of the collection

(R. De Jong, personal communication, 2014); perhaps in

the future examination of this collection will clarify some

of these records. It is furthermore suggested that Piepers

and Snellen’s records are the basis of the continuing

references to palms as food plants of E. thrax in the lit-

erature. Seitz [94] lists sugar-cane, coconut, Rhapis and

Metroxylon in addition to Musa spp.; the similarity to the

species listed in Piepers and Snellen [73] suggests that

their record of A. pinnata (as A. saccharifera) was mistaken

for sugar-cane.

P. B. Richards [95] and R. M. Richards [96] reported

H. irava Moore and E. thrax as hesperiid pests of coconut

in West Malaysia, but Corbett [74] in his Insects of Coco-

nuts in Malaya considers the two have been confused, that

H. irava is the coconut pest and E. thrax is only a banana

pest. Corbet et al. [39] in The Butterflies of the Malay

Peninsula highlight Musa spp. as the food plants of E. thrax

thrax and E. torus, point out that the two do not differ in

life history, and include coconut and sugar palm (i.e.

A. pinnata) as reported food plants.

In Les Insectes des Palmiers, Lepesme [75] considers

E. thrax to be very common on banana, but rarer on Elaeis,

Cocos, Metroxylon, Calamus, Cyrtostachys [Arecaceae],

sugar-cane and bamboo [Poaceae]’, citing De Joannis

(in [81]), Richards [95], Seitz [94], Williams [82],

Dammerman [97], Corbett [74] and Beller and Bhenchitr

[83]. As noted above, Duport [81], Williams [82] and

Beller and Bhenchitr [83] report E. thrax (and/or E. torus)

as a banana pest. Richards [95], Seitz [94] and Corbett

[74] are also discussed above. Dammerman [97] treated

E. thrax as a banana pest, also found on ‘Manila hemp,

bamboo, coconut, oil palm and other palms’ but does not

indicate his sources. None of these sources listed by

Lepesme [75] are the source of records on Calamus and

Cyrtostachys.

Hutacharern and Tubtim [98] listed E. thrax thrax from

Calamus sp. (rattan) in Thailand, and Johari and Che Aziz

[99] listed it as a pest of Calamus trachycoleus in Peninsular

Malaysia. Erionota thrax was listed as a pest of C. manan

in Peninsular Malaysia by Norani et al. [100], although

in a subsequent treatment of rattan pests in Peninsular

Malaysia, Maziah et al. [101] refer to and illustrate

the larva as E. torus, rather than E. thrax. In their survey

of hesperiids associated with C. manan, Steiner and

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8 CAB Reviews

Aminuddin [102] also record E. torus, but later correct

this to E. acroleuca apicalis (as E. hiraca apicalis) [103]. This

correction was based on the advice of L. G. Kirton, who

indicates that E. acroleuca (as E. hiraca) is the Erionota sp.

that feeds on Calamus spp. and other palms. Hence, in his

recent Naturalist’s Guide, Kirton [77] refers to E. thrax

thrax feeding on bananas only.

In ‘Pests of Oil Palms in Malaysia and their Control’ Wood

[104] includes E. thrax as a minor pest, but the illustration

of the adult (plate XIVb above) is of a male E. acroleuca

apicalis with the distinctive white fore wing tips. There is a

relatively recent record of E. thrax hasdrubal as a pest of

oil palm in New Britain [105]. The adult illustrated in the

paper is not E. thrax hasdrubal, but it is not clear whether

it represents E. thrax thrax, E. torus, an individual of

E. acroleuca lacking the pale forewing apex, or some other

taxon of Erionota. Of these, only E. thrax thrax is known

from New Britain, but above we have discounted all

records of palms as food plants for this subspecies. It will

be necessary to examine and dissect voucher specimens

to confirm or clarify this record.

There are further records from Arecaceae catalogued

by Robinson et al. [84]. In their ‘List of economic pests,

host plants, parasites and predators in West Malaysia

(1920–1978)’ Yunus and Ho [106] include Caryota and

Nypa fruticans as food plants of E. thrax, but do not specify

the source for this record, and Pholboon [107] and

Hutacharern and Tubtim [98] in their compilations of

plant pests in Thailand, list Licuala grandis as a food plant of

E. thrax. These records from palms have not been tracked

back to their original source for this review, but below

we document that in West Malaysia E. acroleuca apicalis is

recorded to feed on Caryota spp. and E. sybirita on a Licuala

sp., indicating that these records are likely to be based on

misidentifications. Kitamura [108] reared a single Erionota

sp. from a leaf shelter on coconut on Panay Island,

Philippines, and illustrated the larva, pupa and shelter

(a longitudinally rolled leaflet). He identified the resultant

adult female as E. thrax. Could it have been another

species such as E. acroleuca or E. surprisa De Jong

and Treadaway? Kitamura [109] subsequently reared

E. acroleuca apex from a palm and compares the larva

with his coconut-reared specimen, so clearly considered

them different. Nevertheless, given the concerns raised

here, and the similarity of adult females, this record based

on a single female merits checking by examination of the

genitalia before accepting.

In their survey of E. thrax in New Guinea, Sands

et al. [58] specifically report that no immature stages

were found on Ensete glaucum (Musaceae), H. bihai

(Heliconiaceae), R. madagascariensis (Strelitziaceae),

N. fruticans, Metroxylon sp., Calamus spp., C. nucifera, Areca

catechu or E. guineensis (Arecaceae), even when infested

Musa spp. plants were growing close by. Furthermore,

in the laboratory, groups of ten newly-hatched of E. thrax

died without feeding when placed on fresh foliage of

H. bihai, C. nucifera and E. guineensis, whereas all larvae

placed on M. � paradisiaca (as M. sapientum) as controls,

began feeding. Furthermore, the leaf shelters that E. thrax

and E. torus make on banana (see below) involve beha-

viour patterns that would not be readily transferable to

a pinnate leaved palm. Hence, the evidence indicates

that E. thrax thrax does not use palms as food plants, and it

is anticipated that all the records from Arecaceae, should

be referred to E. acroleuca. It is unfortunate, therefore,

that the records from palms have led to E. thrax being

mentioned repeatedly as a pest of palms in the pest lit-

erature (e.g. [75, 110–115]). It is equally unfortunate that

the common name ‘palm redeye’ has been applied to this

species. The oldest use of this name appears to derive

from Evans [116, 117], and is assumed to be based on the

food plant records referred to above, ignoring the well-

documented damage to bananas. Although Brigadier

W. H. Evans was an excellent morphological taxonomist,

there is almost no reference to food plants and early

stages in any of his works, so it seems likely that he was

not that well-informed on this aspect, hence the use of

this inappropriate name. I recommend that the use of this

common name should be discontinued for all species of

Erionota to avoid future confusion.

There are also records from Poaceae. Robinson et al.

[84] attribute a record of Chrysopogon to Sevastopulo’s

[118] The Food Plants of Indian Rhopalocera, but this is

incorrect, as Sevastopulo does not list this grass.

Saccharum (sugar cane) is listed as food plant by

Sevastopulo [118] based on Seitz [94], which it was sug-

gested above is an error for sugar palm. Robinson et al.

[84] attribute a record of Saccharum to Corbet et al. [39],

but the food plant listed there is sugar palm, i.e. A. pinnata

as originally listed by Piepers and Snellen [73] and dis-

cussed above. Robinson et al. [84] also cite A Manuscript

List of Butterfly Hostplants in the Western Ghats and Southern

India by H. Gaonkar (2000), which does not seem to have

been subsequently published, that Saccharum officinarum

is a food plant of E. thrax thrax. Whether this is a repeat

of one of the earlier errors or a new record, it is anyway

unlikely to be an original record as E. thrax is not reported

from this part of India, and E. torus has only recently been

found there. It is concluded that none of the published

records of Poaceae as food plants for E. thrax should be

accepted.

Finally, Robinson et al. [84] cite Hutacharern and

Tubtim [98] that Butea monosperma (Fabaceae) is a food

plant of E. thrax thrax in Thailand, but this is an error for

the record of Calamus sp. on the same page ([98], p. 41),

and this record should not be perpetuated.

Interpreting these food plant records, it is worth

remembering two facts documented in weed biological

control. Firstly, oligophagous species are mostly, but not

always, more likely to feed on species closely related to

their main food plant, than on those less closely related

[91]. On this basis, Musaceae and Strelitziaceae are part of

one clade of Zingiberales, while Marantaceae, Cannaceae

and Heliconiaceae are in the other [89]. Families from

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Matthew J. W. Cock 9

outside the order Zingiberales, i.e. Arecaceae in this case,

are much less likely to be suitable food plants. Secondly,

when a successful weed biological control agent is first re-

leased, there is often a population explosion, during which

there can be spill-over effects where feeding damage

occurs on other plants, usually because insects have

strayed or fallen from the normal food plant onto adjacent

plants. The same thing may happen with any newly

introduced species, such as E. thrax and E. torus in their

exotic range, so that food plant records during a popu-

lation explosion will include species that are not normally

suitable, on which eggs will not normally be laid, or which

will not sustain the full life cycle. Some of the observations

reported, e.g. from Hawaii [88], may be of this nature.

To summarize, it is concluded that the food plants of

E. thrax are restricted to Musa spp., perhaps some other

Musaceae and a few related, similarly broad-leaved species

in Heliconiaceae, Strelitziaceae and possibly Zingiberaceae

(all Zingiberales). No convincing food plant records have

been found of subspecies of E. thrax apart from the

nominate subspecies.

Food plants of Erionota torus

As has become evident during this analysis of E. thrax,

there has been confusion between E. thrax and E. torus

where the two co-occur in mainland South-East Asia.

Accordingly, food plant records without voucher speci-

mens can only be accepted for E. torus with full confidence

where E. thrax does not occur. It is clear that like E. thrax,

E. torus is primarily a pest of Musa spp. Only E. torus

occurs in Hong Kong and adjacent China, and here it has

been documented to feed on Musa spp. but not on

other plants since the late nineteenth century as E. thrax

[119–121], and more recently as E. torus (e.g. [9, 122]).

Similarly, only E. torus appears to be present in the

Western Himalayas [1], so Mackinnon & de Niceville’s

[64] report of E. thrax as common on cultivated plantains

around Dehradun is assumed to refer to E. torus.

Since the early stages of E. thrax and E. torus cannot

be distinguished, and they are both common pests of

Musa spp., it follows that where both species co-occur in

mainland South-East Asia and the Southern Philippines,

then any studies on the pest status, natural enemies,

control, etc. which do not address the presence of the

two species can be assumed de facto to treat either or

both. This would be the case for the detailed studies by

J. Okolle and colleagues in Peninsular Malaysia on the

population dynamics, within-field and within-plant dis-

tribution of the banana skipper and its parasitoids

[70, 123–129].

There are a few published records of palms as food

plants for E. torus. Robinson et al. [84] list Caryota

and Roystonea regia. As noted above, in their treatment of

pests of rattan palm in Peninsular Malaysia, Maziah et al.

[101] include E. torus, although Steiner [130] and Steiner

and Aminuddin [103] subsequently indicate that this

record should be treated as E. acroleuca (=E. hiraca).In a laboratory study in Taiwan, Tsai et al. [131] fed

cohorts of each of the five instars of larvae of E. torus with

cut portions of leaves of banana, Strelitzia reginae (bird

of paradise flower; Strelitziaceae), C. nucifera (coconut;

Arecaceae), areca nut (A. catechu; Arecaceae), Bambusa

oldhamii (bamboo; Poaceae) and sugar cane (Saccharum

officinarum; Poaceae). Feeding was most extensive on

banana, but almost as heavy on Strelitzia reginae; there was

very little feeding on coconut (none for the cohort of

instar 1), less on areca nut and none on bamboo or sugar

cane. Average survival for all instar cohorts was 50% on S.

reginae, 5% on coconut and 0% on bamboo and sugar

cane. This is a measure of the survival of larvae trans-

ferred to a plant other than banana, and does not reflect

what would happen following oviposition on that plant in

the field. If there was no feeding in instar 1 on coconut, all

larvae would die in the field, while the data does not allow

us to interpret what total mortality would be on S. regi-

nae, it would be more than 50%. In this experiment, the

larvae on banana are reported to have all completed

development, but in a parallel experiment, only 34 and

21% of individuals completed their metamorphosis on

banana at 20 and 30 �C, respectively, most mortality

occurring in instars 1 and 2. It is difficult to know how to

interpret these two apparently conflicting results. How-

ever, we can conclude that sugar cane, bamboo, areca nut

and coconut are unsuitable food plants, whereas S. reginae

is less suitable than banana.

There are internet records of E. torus using Canna sp(p).

(Cannaceae) as food plants (e.g. [132]), but these have not

been traced to any formal publication, and may well

be derived from the records for E. thrax, when the name

E. torus was misapplied to that species.

As noted above and discussed below, the early stages

and leaf shelters of E. thrax and E. torus are very similar.

The shelters can only be built using a leaf with a large flat

lamina, e.g. Musa spp. Heliconia spp., some Strelitziaceae,

Marantaceae, Zingiberaceae, etc. (all in the Zingiberales),

and unlikely to be used on pinnate palms. It is concluded

that all food plant records of E. torus from palms should

be disregarded, and although the possibility of other

Zingiberales being used as food plants cannot be discoun-

ted, none should be considered confirmed at this time.

Food plants of E. acroleuca

Erionota acroleuca acroleuca has been recorded using

palms as food plants in the Andaman Islands: C. nucifera,

Pinanga kuhlii, Calamus sp. [87]. Erionota acroleuca apicalis

from South-East Asia also feeds on palms. In Peninsular

Malaysia, it is recorded from ‘oil palm’ ([104] as E. thrax,

[76]), Caryota sp. and Orania sylvicola (=macrocladus) [106],

Calamus manan, C. ornatus, C. scipionum, Caryota mitis,

Areca triandra and Adonidia merrillii [130, 133]. Recently,

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10 CAB Reviews

Xue and Lo [19] report rearing this subspecies from

Caryota maxima (as C. ochlandra) in Yunnan Province

and Arenga westerhoutii in Guangxi Province, China, and

an unidentified palm in Singapore; they conclude that

E. acroleuca is primarily a palm feeder. Kitamura [109]

reports Saribus rotundifolia (=Livistona rotundifolia) to be a

food plant of E. acroleuca apex on Samar Island, Philippines.

L. G. Kirton (in 103, 130) has suggested that all records

of Erionota spp. from rattan palms (Calamus spp.) should

be treated as E. acroleuca, and here this generalization is

extended to all records from palms (although some of

these could be errors for H. irava). In particular, the

records of E. thrax from C. nucifera, R. excelsa, M. sagu,

A. pinnata and E. guineensis in Java [73] and the various

repetitions of these in the literature (see above) are

considered to be in error for E. acroleuca. L. G. Kirton

(personal communication, 2014) transferred larvae of

E. acroleuca apicalis of various sizes from palms to Musa sp.

and they all died. The record of E. thrax from oil palm in

New Britain, PNG [105] needs clarification by examina-

tion of voucher specimens; it might prove to be E. acro-

leuca. In conclusion, E. acroleuca is the Erionota species that

is commonly found on a wide range of palms, and until

such time as there are authoritative records of E. thrax

or E. torus breeding on palms, all such existing records

should be considered misidentifications, most probably

for E. acroleuca.

Food plants of other Erionota spp.

The Chinese species, E. grandis feeds on palms [6], as does

E. sybirita (Hewitson). The former is a gregarious species

and so unlikely to be mistaken for other Erionota spp.

Steiner [130, 133] records the later feeding on Licuala

kunstleri in Peninsular Malaysia. It is a rare species found

from Southern Myanmar and Thailand to Borneo and

Palawan [1, 18], so is unlikely to account for many of the

records of other Erionota spp. from palms. The food plants

of the remaining Erionota spp. are unknown.

Immature Stages

Erionota thrax, E. torus and E. acroleuca are similar in adult

appearance [1, 21]. The life history and food plants of

E. thrax and E. torus are very similar [6, 39]. However,

while the early stages of E. acroleuca are also similar, the

food plants and shelters are not. Hence, at this time, the

early stages cannot be reliably separated by diagnostic

characters, and E. acroleuca can only be provisionally

separated by its food plants and shelters.

The larvae of E. thrax and E. torus make their leaf

shelters by cutting and rolling the lamina of their food

plant leaves. More detailed observations of shelter for-

mation are needed for both species, with information on

individual variation, feeding habits, etc. At present, the

shelter making behaviour cannot be used to separate

E. thrax and E. torus. In contrast, although hardly docu-

mented, the leaf shelter of E. acroleuca on palm leaves may

be constructed similarly to those of E. thrax and E. torus,

but is also made by longitudinally rolling a leaflet or pulling

together two leaflets or parts of a larger leaf. This latter

type of shelter seems typical for palm-feeding Hesperiinae

[7], and contrasts with the shelters of E. thrax and E. torus

where the leaf is rolled transversely.

Life history of E. thrax

Horsfield and Moore [78] provide the first illustrations of

the larva and pupa. As these observations were from Java,

where E. torus is not reported, they should be reliably

associated with E. thrax. The figure portrays the larva as

white, with long erect setae on the body, and a dark

brown-black head. The pupa is incorrectly shown, in that

it has a short, strong, pointed, slightly upturned frontal

spike. Horsfield’s emerged pupa is preserved in the NHM,

and is accurately drawn, so is here assumed to belong to

some other species. Distant [79] reproduces these

figures of the larva and pupa in Rhopalocera Malayana,

perpetuating this error regarding the pupa. Semper [24]

illustrates the larva, correct pupa and a large leaf shelter

implicitly from Manila, Luzon Is, Philippines, in which case

they are safe to associate with E. thrax.

Piepers and Snellen [73] document the biology: the

larva ‘lives in a part of a leaf that is rolled up into a case

and fastened with threads, it is chalk-white with a black

head overgrown with very short hairs and wholly covered

with a white waxy powder, that also entirely fills its

abode’. Although Piepers and Snellen treat E. acroleuca as

a synonym of E. thrax, it seems likely given the description

of the shelter on Musa in Java that they are dealing with

the immature stages of E. thrax. Their painting of the pupa

also appears to represent this species, having no frontal

spike, and a long proboscis extending beyond the cre-

master.

In Lepidoptera Indica Vol. X, Swinhoe [80] gives the food

plants as ‘wild or cultivated Musa’ and states that the larva

‘lives in a shelter made of a portion of rolled-up leaf. To

make this shelter it has to cut into the edge of the

enormous leaves of the wild or cultivated Musa, or

plantains, to obtain a suitable segment to be rolled up.’

The larva is figured from ‘Grote’s original drawing’.

Arthur Grote prepared the first catalogue of the butterfly

collection of the Asiatic Museum in 1857–1859; he also

took notes on the life histories, making use of a Munshi

Zynulabdin, a local artist [134], so the material for these

figures was likely to have been collected in the vicinity

of Kolkata (Calcutta), and could represent either E. thrax

or E. torus. Although not acknowledged, the figure of a

pupa is copied from [78] and so is incorrect. Evans [1]

indicates that the adult illustrated by Swinhoe [80] is

E. thrax; however, the observations on early stages and

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Matthew J. W. Cock 11

the figure of the larva could refer to either or both

E. thrax or E. torus.

Williams [82] describes the biology on banana on Luzon

Island, Philippines, which should be correctly associated

with E. thrax. ‘The eggs are hemispherical and well ribbed

from the apex and are not quite 2 mm in diameter. The

little larvae cut obliquely inward from the edge of the

banana leaf and form a tube by fastening this rolled strip

with silk; a large larva makes a large roll as illustrated.

Here it also pupates. A full-fed larva is fusiform-cylindrical

with a large head set on a small neck, as is usual among the

Hesperiidae, and is covered with a whitish flocculent

substance.’ Williams [82] includes drawings of a banana

leaf with many small shelters, a leaf with a single large

shelter, and a parasitized pupa. The figure of the large

shelter is not entirely clear, nor is it explained in the text.

It can be interpreted as indicating that the larva makes

two parallel cuts along the leaf and rolls the resultant flap,

leaving the leaf edge intact. An alternative explanation is

that the area of leaf adjacent to the edge of the lamina

reflects the portion of the leaf that is eaten or not eaten

by the larva.

The following account is synthesized from several

sources. Ashari and Eveleens [135] provide drawings of

the life cycle and provide some additional information.

More recent observations are similar, but some have

been made from areas where E. thrax and E. torus occur

together and so cannot be allocated to species with

confidence (e.g. [16, 70]). Igarashi and Fukuda [6] include

the life history of E. thrax from India (in their text they

only mention Sikkim). As noted above, Igarashi and

Fukuda [6] refer to the species that has spread to the

Pacific and Asia as E. torus, rather than E. thrax. As they do

not refer to the diagnostic differences in the genitalia, one

cannot be sure that their identifications were thus con-

firmed. However, although both E. torus and E. thrax occur

in Sikkim, India, and the possibility of confusion cannot

be ignored, the adults that they illustrate are correctly

associated (the difference in male forewing shape is very

clear), so it is assumed that the early stages are also

correctly associated. Although there are photographs and

brief descriptions of the biology of E. thrax in its intro-

duced range [45, 136] no detailed accounts have been

located in this review.

Eggs are laid singly or in groups, mostly on the under-

side of the leaves, but also on the upper side. Eggs are

2.3 mm wide�1.1 mm high, with about 22 fine ribs stop-

ping short of the micropyle.

Detailed descriptions of the leaf shelter building beha-

viour are not available. The newly hatched larvae form a

little leaf roll from the edge of the leaf, inside which they

feed [135]; it is not clear whether the first shelters

are simple leaf folds from the leaf edge, or the transverse

leaf roll observed later – or both. Igarashi and Fukuda

[6] note that ‘the larva rolls a part of a food plant leaf

inwards . . . to make a nest in which to rest with its head

directed upwards. . . . The larva eats the wall of the nest

from below . . . Feeding occurs only during the night

�time. The bottom of the nest is usually closed, and often

filled with a large amount of frass pellets. . . . the larva

moves and nests are renewed several times. Moving takes

place in the direction towards the basal part of the leaf. It

takes a few days to make a new nest. Some larvae,

however, may show behaviour patterns exceptional to

the habits described above.’ Leaf rolls made by the mature

larvae extend all the way from the edge to the leaf midrib,

which leads to considerable loss of leaf area (Figure 4.2).

The leaf rolls in Figure 4.1 appear to have five rolls of leaf,

and dangling from the bottom is a narrow strip of leaf

margin which has not been eaten. This can be interpreted

as the larva feeding on the leaf edge within its shelter, but

not the section furthest from the midrib, which would

form the distal end of the leaf roll, where a feeding larva

might be vulnerable to predators or disturbance. This may

be the reason for the two lines in Williams’ [82] drawing

mentioned above.

There are five larval instars and the final instar grows to

50 mm or more. All instars have dark brown to black

heads and pale bodies. After the second instar there is an

increasingly dense cover of white powder on the body.

The body has fairly inconspicuous pale, erect setae; the

head is dark brown, rounded, indent at vertex, widest

near base, with setae similar to those on the body; the

pronotum, spiracles, true legs and prolegs are all pale,

Figure 4 Shelters of E. thrax thrax and E. torus on Musasp. 1, E. thrax, Maui, Hawaii, 13 Aug 2002 (

gForest and

Kim Starr); 2, E. thrax outbreak, Fly River, Papua NewGuinea (

gC. Dewhurst); 3, E. torus, Serdang, Malaysia, 6

Feb 2003, M. J. W. Cock. Differences observed betweenindividual shelters are not considered to be diagnostic, butto reflect individual differences.

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12 CAB Reviews

more or less concolorous with the body. It can be seen

that there is minimal white powder on the head, thorax

and ventral part of the abdomen of the individual shown in

Figure 5.1, but the covering becomes more extensive as

the larva becomes fully grown. The pupa (Figure 6.1) is

pale white-brown, nearly cylindrical but deeper and wider

at the thorax; no frontal projection and the proboscis

sheath projects beyond the cremaster. The pupa and the

inside of the pupal shelter are lined with white waxy

powder.

Life history of E. torus

Walker [119] wrote about the butterflies of Hong Kong,

and was the first to comment on the leaf-roll shelters,

stating that E. thrax (i.e. E. torus) has been reared ‘in 1892

from larvae found in rolled-up leaves of banana’. Mack-

innon and de Niceville [64] include a brief description of

the larva in northern India, and mention that pupation is in

a portion of a rolled-up leaf. In the Palaearctic butterflies

section of the Macrolepidoptera of the World, Seitz [137]

wrote ‘The larvae of Erionota . . . feed on the gigantic

leaves of Musa, which are yards long. In forming their

shelter they commence by making two incisions from the

edge of the leaf towards the midrib at a distance of about

8–10 cm one from the other, and then proceed to roll this

piece of the leaf up like a cigar, the larva living in the

cylindrical hollow within this roll.’ When dealing with the

Indo-Australian butterflies, Seitz [94] expands on this ‘The

larva is snow-white, covered with a very short wax-like

pubescence, and with a black head. It is easily discovered,

as it gnaws out of the gigantic banana leaves, by two

parallel-cuts a longitudinal piece which it rolls up like a

cigar, in the centre of which it lives. These characteristic

cuts you can see in the Musa-plantations from a distance

of 20 or more paces, so that the larvae, and still more

easily the pupae, may be collected in numbers ad lib. The

pupa is of a dingy pale yellow with a very long case of the

proboscis projecting beyond the end of the body like a

spear . . .’. These seem to be Seitz’ own observations as

he refers to collecting many specimens in Hong Kong, i.e.

they are E. torus not E. thrax. The reference to two cuts to

make the shelter are not clear, but may parallel those

shown by Williams [82] for E. thrax, and perhaps explained

by the feeding behaviour of the larvae inside the shelters

as extrapolated above for E. thrax based on Figure 4.1, i.e.

the cut nearer the midrib is used to make the shelter, but

the second ‘cut’ is actually where the larva stops feeding

within the shelter in order not to expose itself at the

bottom.

Based on his observations at Hong Kong and Canton

(Guangdong, China), Hoffmann [121] provided a quite

detailed description of the early stages and their biology,

together with drawings of the ovum, larva and pupa, and

photographs of adults and shelters. Eggs are laid singly or

in irregular groups on the upper and lower surfaces of

leaves (more frequently the latter). They measured 2 mm

in diameter and 1.13 mm in height, with the flat area on

top 0.45 mm wide; there are 24–29 fine ribs. They are

usually orange in colour, but there is considerable varia-

tion over time and within batches. There are five larval

Figure 5 Final instar larva of E. thrax thrax and E. torus..1, E. thrax, Maui, Hawaii (g Forest and Kim Starr); 2, E. torus,Mauritius (g T. C. E. Congdon). The apparent difference in head colour is not considered diagnostic.

Figure 6 Pupa of E. thrax thrax and E. torus on banana, lateral view. 1, E. thrax, Papua New Guinea (g C. Dewhurst);2, E. torus, Mauritius (g T. C. E. Congdon).

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Matthew J. W. Cock 13

instars. The first instar is 3 mm long when recently hat-

ched, pale green with the head and pronotum black, and

no wax powder. The width of the head capsules of the five

instars were 1.08, 1.45, 2.08, 2.9 and 4.26 mm. After the

first moult, the larva is covered with an increasingly

dense layer of white powder. The fifth instar measures

about 44 mm long; the surface of the blackish-brown head

is ‘more or less granulose’, with a moderate covering of

weak, silky setae, thicker on the front. The pupa measures

40–46 mm, and is about 7 mm wide and deep at the

thorax and is covered with white powder. It is pale dirty

yellow apart from the dark brown cremaster; the pro-

boscis extends 5 mm beyond the cremaster; spiracles

brown; the eyes become reddish about halfway through

the development period.

After the early reports considered above, there

seems to be little in the way of original observations, as

opposed to repeating information from the literature,

until late last century, when accounts illustrated with

photographs started to appear. There is a brief summary

from Mauritius in Monty [59] as E. thrax. Bascombe et al.

[9] describe and illustrate the early stages of from Hong

Kong, and Igarashi and Fukuda [6] include the life history

from Japan. Additional information is provided from the

observations of T. C. E. Congdon (personal communica-

tion, 2014) and the author’s unpublished observations.

The following is largely based on Bascombe et al.’s [9]

account, except as indicated. The egg ([9], Plate 103.1–2)

is 1.8–2.2 mm in diameter, dome-shaped with 22–24 fine

ribs stopping short of the micropyle; they are pink or

variegated pink and yellowish-white when laid, becoming

paler as they develop. They are usually laid on the leaf

underside, and may be laid singly, but more often in

clusters of up to 30, rarely 50. In contrast, Igarashi and

Fukuda [6] state that the ova are 2.2 mm in diameter, and

normally laid singly on the leaf upper side, although bat-

ches of 3–36 are recorded; this may reflect a gen-

eralization on a relatively small number of observations.

On hatching the larva eats the egg shell, and then moves

to the edge of the leaf to make its first shelter, which is

either a simple fold parallel and close to the edge of the

leaf ([9], Plate 103.3), or a cone shaped roll formed by

making a curved cut from the edge of the leaf and rolling

the resultant flap under. This first cut may be curved

towards the leaf base or away from it, but all subsequent

shelters are made with the cut curved towards the base of

the leaf. Larvae feed on the inner parts of the roll, which is

closed at the inner end (nearest midrib) and open at

the distal end, where frass may accumulate. As the larva

grows, it extends the cut and rolls more of the leaf into

the shelter. Additional shelters are constructed if the

shelter is damaged, unable to roll further or encounters

another shelter.

Igarashi and Fukuda [6] indicate that the leaf shelters of

E. torus and E. thrax are rolled in opposite directions,

although it is not clear from the text and diagrams whe-

ther this means clockwise and anticlockwise when seen

from above, or one with the leaf under surface on the

inside of the shelter and the other with the leaf under

surface on the outside of the shelter. In any case, they

seem to be incorrect, as both species roll the leaf flap

clockwise or anticlockwise depending on which side of

the leaf the shelter is formed, and the shelters normally

have the leaf underside inside the shelter for both species.

Igarashi and Fukuda [6] also indicate that the top end of

the larval leaf shelter is open and that the larva comes out

of this opening to feed on the leaf by night; this also seems

to be incorrect as photographs of the leaf shelters indicate

no feeding outside the shelters [9, 66, 121, 138], and the

indications are that the larvae feed inside the tube, as

discussed below. Working in Japan, Makibayashi [139]

provides a detailed description with clear diagrams to

show how a final instar caterpillar makes a new leaf

shelter. His diagrams show that the leaf roll is pulled into

position by silk strands between the leaf roll and the

adjacent unaffected part of the leaf, which is in due course,

is pulled into the shelter in turn.

The following notes on E. torus in Mauritius were pre-

pared by T. C. E. Congdon (personal communication,

2014). The larva makes a cut in the leaf, and rolls the cut

portion, extending the cut across the leaf as it grows. No

evidence of eating can be seen outside the shelter, and

when the tube is unrolled, the unrolled leaf does not refill

the space it originally occupied. It appears that the larva

feeds by eating the inner layers of the tube, thus avoiding

the dangers inherent in emerging from its shelter. One

effect of this is that the inside diameter of the tube

becomes much greater than is usual in Hesperiinae tube

shelters. The bottom end of the shelter is loosely sealed

with silk, retaining the frass, thus hindering access by

predators. The top of the shelter is crudely closed with

silk, but must be opened when the larva extends the cut,

and rolls more of the leaf. Old strands of silk are cut and

remain visible along the cut on the section not incorpo-

rated into the shelter. This detail is at variance with the

description and diagrams of Makibayashi [139] above, and

the author’s unpublished observations from western

Malaysia that the silk strands used to form the shelter

are attached to parts of the leaf that are subsequently

incorporated into the shelter. Further observations are

needed to clarify whether these are local or individual

variation in behaviour. The larva pupates within the

shelter, head up and without a girdle.

From an early age, the larva is covered with an

increasingly thick layer of white powder, so that the body

appears white with erect pale setae partially covered with

the powder, and the uniformly dark head is partially

obscured with the white powder (Figure 5.2). The larva

grows to 55 mm in length [6] and is covered with white

flocculence as it prepares for pupation.

Pupation takes place in the final leaf roll shelter which is

15–20 mm long, lined with a thin layer of silk and closed

at the distal end by a thin mesh, which is easily broken by

the emerging adult. Ito and Nakamori [140] refer to the

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14 CAB Reviews

pupal shelters in Japan as ‘trumpet shaped’, 1.5–2.0 cm

diameter at the smaller [basal] opening, and 3.0–3.5 cm at

the terminal opening. This does not seem to be typical.

Igarashi and Fukuda [6] note that the pupal shelter is

simpler than the larval shelter, having 3–4 layers rather

than 5–6, and the pupa is formed with the head upwards,

although the adult butterfly emerges from the bottom of

the shelter.

The pupa (Figure 6.2) is 45 mm long, pale brown with a

variable and patchy covering of white powder. The pro-

boscis extends almost to the cremaster. It is attached at

the cremaster, and unlike many Hesperiinae does not

construct a silk girdle.

Life history of E. acroleuca

Wood [104] treats E. acroleuca apicalis as E. thrax

and includes a photograph of a hesperiid larva and

shelter (plate XIVa) without explicitly stating whether it

represents E. thrax or Cephrenes chrysozona (Plotz), the

two named species of Hesperiidae treated. The angle of

view does not permit a definitive identification to be

made, but it is more likely to be E. acroleuca than

C. chrysozona. The colouring of the larva suggests an

Erionota sp., rather than a Cephrenes sp., which has a green

body and white head with dark lines [9, 39]. The leaf

shelter appears to have been formed by making a diagonal

fold across the oil palm leaflet, but since this may have

been constructed in captivity, it is not necessarily indica-

tive of what happens in the field. Mature C. chrysozona

larvae roll palm leaflets parallel to the mid rib (author’s

unpublished observations).

Steiner [133] records observations on the ova of

E. acroleuca apicalis (as E. hiraca). The ova are white, with

a smooth surface and hemispherical in shape. They can

be found on all leaves of C. manan, but the majority are

found on the youngest leaf, and although found on both

surfaces of the leaf, they are mostly found on the upper

surface. Xue and Lo [19] indicate that the ova of

E. acroleuca apicalis are laid in a cluster on the frond

underside.

Maziah et al. [101] treat E. acroleuca apicalis as E. torus

in their illustrated account of the pests of rattan, and

illustrate the larva with a photograph. They note that the

larva ‘rolls up the leaf from the tip parallel to the veins and

lives and feeds from one end of the leaf in this shelter’.

This wording is not completely clear, but in view of sub-

sequent observations, it is clear that the leaflet is rolled

transversely (more or less at right angles to the mid rib),

unlike the longitudinally rolled shelter (parallel to the

midrib), made by most palm-feeding Hesperiinae [7]. They

describe the larva as ‘pale green covered with short silky

hairs and a whitish powdery substance and has a dark

brown head capsule’. Their photograph shows a larva

similar to those of E. thrax and E. torus. Steiner’s [133]

description and photograph of a 34 mm larva match

earlier observations. To make the larval leaf shelter, larvae

cut the leaflets near the base about two-thirds of the

distance to the midrib, and wrap the distal part in several

turns to form a roll (Steiner refers to this as a longitudinal

roll, but I would interpret this term as parallel to the mid-

rib, which is not the case here). The larva rests and feeds

inside the leaf roll. Prior to pupation, the larva creates a

similar kind of leaf roll, which is heavily lined with wax.

L. G. Kirton (personal communication, 2014) confirms

these observations, and has also observed that on Caryota

sp(p)., which have laminate fishtail pinnae, the larva makes

a shelter in a similar manner to those of E. thrax and

E. torus on Musa spp., i.e. by cutting an arc from the edge

of the lamina and rolling the resultant flap into a tube, in

which the larva hides and feeds. Xue and Lo [19] provide

some additional information regarding E. acroleuca apicalis.

Larvae, hatching from egg clusters develop in separate

pinnae on the same rachis, rolling the pinnae into a ‘cone-

shape shelter’. They illustrate in colour a larva similar to

those above, with a dark brown head with short setae,

conspicuous long setae on the body, and the head and

body covered with white waxy powder.

Maziah et al. [101] record that pupation is in the final

larval shelter and ‘the pupa is long, slender and also

covered with the same whitish powdery substance pre-

sent on the larva. When disturbed the pupa wriggles

violently in the shelter’. Xue and Lo [19] indicate that the

pupa is 33 mm, pale cream in colour with brown spiracles,

the proboscis extending beyond the cremaster. It is

‘sealed in a cocoon’ in a shelter formed by ‘rolling the

undersurface of the apex half of the rachis [pinna?] lat-

erally’. This description of the shelter is not entirely clear,

and although their illustrations of the shelter and pupa

show a silk cocoon within a leaf shelter, the structure of

the shelter cannot be interpreted, although it appears to

be aligned longitidinally with the pinna veins.

Finally, Kitamura ([109], Figure 18, upper photo)

includes a photograph of the final instar of E. acroleuca

apex in his report on this butterfly feeding on L. rotundifolia

on Samar Island, Philippines. The larva is similar to those

reported above (and to other Erionota spp.), but the leaf

shelters are not documented.

The shelter building behaviour of the larvae seems

to vary with the leaf structure of the food plant palm.

Further observations from different food plants and from

different parts of the range of this species and its sub-

species, supported with voucher specimens, are needed

to clarify the biology of this species.

Life history of E. sybirita

There are also some details on the life history of E. sybirita

on L. kunstleri in Steiner’s [133] thesis. The eggs are white

to grey, hemispherical with a smooth surface; almost

all were found on the underside of the leaflets. The

larvae have a whitish green body, but unlike the other

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Matthew J. W. Cock 15

Erionota spp. treated above, no wax layer. The head

capsule is dark brown to black. The pupae are brownish,

covered with wax, and about 36 mm long. The very long

proboscis sheath extends beyond the abdomen by up to

40 mm. The pupae are attached only by the cremaster and

do not have a silk girdle. Like the larvae of E. hiraca, the

larvae of E. sybirita cut the leaflets near the base and wrap

the distal part into a roll with several turns (again referred

to as longitudinal, but this term should be avoided for the

leaf rolls of Erionota spp.). The larva stays in the roll and

eats it from the inside. Prior to pupation the lower end of

the tube is sealed with silk. It appears that the absence of a

covering of white waxy powder on the larva (and perhaps

also on the pupa) as well as the specific host plant should

facilitate separation of the early stages from E. acroleuca.

Natural Enemies of E. thrax and E. torus

Waterhouse and Norris [45] summarized the available

information on the natural enemies of E. thrax, many only

identified to genus or family, but as has become apparent

above this summary would not have separated natural

enemies of E. thrax from those of E. torus where they

occur together. This table is updated as three separate

tables below, broken up into records that can be un-

equivocally associated with E. thrax (Table 2), E. torus

(Table 3), or because of their location could be associated

with either species (Table 4). Summaries of parasitoid

records in Gold et al. [171], Tinzaara and Gold [172],

and Muniappan et al. [115] are not repeated here. Key

information on what appears to be the most common

and important parasitoids (all Hymenoptera) is sum-

marized in the following paragraphs.

Brachymeria spp. (Chalcididae)

Several Brachymeria spp. that attack pupae of Erionota spp.

appear to be polyphagous species, including B. lasus

(Walker), B. albotibialis (Ashmead), B. euploeae (West-

wood) and B. thracis (Crawford). Okolle et al. [127] found

that B. albotibialis also attacks prepupae. Where recorded,

they are gregarious; Hasyim et al. [147, 148] record an

average of nine parasitoids per host, and Okolle et al.

[127] report seven per host for B. albotibialis. The incon-

sistency of names used in different publications, even from

the same area, suggests that at least some material has

been misidentified in the past.

Ooencyrtus pallidipes (Ashmead) (Encyrtidae)

This species was better known as O. erionotae Ferriere,

which was described from Peninsular Malaysia from ‘eggs

of E. thrax L. and of an unknown Lepidopteron on wild

plantain leaf’ [169]. Given the locality, this could have

been from either E. thrax or E. torus. Huang and Noyes

[150] point out that O. pallidipes is the senior synonym.

Ashmead [173] described O. pallidipes from the Philippines

from ‘two specimens bred by Father W. A. Stanton from

an undetermined aphid’. This now seems to be a host

error, perhaps because a mummified aphid may be similar

in size, shape and colour to a parasitized ovum of E. thrax.

Ooencyrtus pallidipes is a common and widespread egg

parasitoid of E. thrax (Table 2) that also parasitizes ova of

E. torus (Table 4). It is reported parasitizing E. thrax (or

E. torus) from Nepal, Thailand, Malaysia, Sumatra, Java,

PNG and Guam, although the last two may represent

accidental introductions. It has been introduced as a

biological control agent in Mauritius and Hawaii for the

control of E. thrax (although the pest in Mauritius is now

known to be E. torus). Females are reported to prefer

younger ova for oviposition [45] and once the larvae start

to form within the ovum, they are not parasitized [127].

Females avoid parasitized ova for oviposition [127].

In Sumatra, it is reported to cause 10–27% mortality

[148, 151] and Hasyim et al. [147] reared an average of

3.8 adults per parasitized egg. It has been recorded from

several Lepidoptera, including Eupterotidae, Gracillariidae,

Nymphalidae, Papilionidae and Pieridae [141], but the ova

of Gracillariidae are most probably too small to support

this species. A record from ova of a buprestid represents

a misidentification [150].

Pediobius erionotae Kerrich (Eulophidae)

This species was described from specimens reared from

ova of E. thrax (or E. torus) in peninsular Malaysia, and the

type series includes material from Java, and R. A. Syed’s

material from Sabah, Malaysia reared from ova, larvae

and cocoons of Cotesia erionotae (Wilkinson) [149, 155].

Kerrich [155] does not seem entirely convinced by the

records from E. thrax larvae and even less so by the

record from a C. erionotae cocoon. This is another com-

mon egg parasitoid of E. thrax (or E. torus), which has

also been recorded Sumatra, where it causes 16–39% egg

mortality [148, 151].

Cotesia erionotae (Wilkinson) (Braconidae)

This gregarious larval parasitoid was described from

E. thrax (or E. torus) in West Malaysia [157], and is known

to attack one or both Erionota spp. in Thailand, West

and East Malaysia, Sumatra and Java (Table 3); it attacks

E. thrax in Sabah, Malaysia (Table 2) and E. torus in

Mauritius (Table 4). It was originally described in the

genus Apanteles, but Austin and Dangerfield [174] trans-

ferred it to Cotesia. Larvae of instars 2–4 are parasitized

and the mature parasitoid larvae emerge mainly from

instar 5 [127]. Hasyim et al. [147, 148] record an average

of 58 adults per parasitization, although cocoons are often

hyperparasitized, while Okolle et al. [127] record an

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16 CAB Reviews

average of 87 per larva. This species has been released

and established for biological control of E. torus in Maur-

itius and Taiwan and E. thrax in Hawaii, Guam, Saipan and

PNG. It was initially thought to be already present in

Taiwan, attacking Udaspes folus (Cramer) (Hesperiidae,

Hesperiinae, incertae sedis) which feeds on gingers

(Zingiberaceae), but closer examination showed differ-

ences in microsculpture and colouring [175], suggesting

this is a different species. In light of this, a recent record

of C. erionotae parasitizing U. folus in India [176, 177] may

need verification.

Casinaria sp. (Ichneumonidae)

This is a common larval parasitoid of E. thrax in Sumatra

and Java. In Java, 10–80% larval parasitism is reported, and

it is commonly hyperparasitized by Brachymeria lasus, and

rarely by Eurytoma sp. B and Theronia zebra [147]. It is

possible that the records of a Scenocharops sp. a common

larval parasitoid of E. thrax in Sabah [149] represent the

same species as the two genera are similar and closely

related in the tribe Campoplegini.

Xanthopimpla gampsura Krieger (Ichneumonidae)

This is a common pupal parasitoid of E. thrax in Sumatra

and Java. It was described from Borneo and Sumatra

[178], and has also been recorded as a parasitoid of the

palm-feeding hesperiids H. irava and Cephrenes chrysozona.

Earlier records from Java of Xanthopimpla sp. are referable

to this species, but it is not clear whether the record

of X. regina from Sabah represents a different species or a

misidentification.

Biological Control of E. thrax and E. torus

There have been several programmes of biological control

against E. thrax (Guam, Saipan, Hawaii, PNG) and E. torus

(Mauritius, Taiwan). Waterhouse and Norris [45] provide

a review of the earlier programmes against E. torus

and E. thrax (summarized below and in Table 5). There

are numerous natural enemies in the indigenous range of

E. thrax (previous section, Tables 2–4), and several have

been used as biological control agents (Table 5).

Erionota torus was first reported in Mauritius in 1970 as

E. thrax [59], although it was suspected to have been

introduced with military air traffic from Malaysia in 1968

[45]. Once established in Mauritius, it soon became a

serious pest on tall banana varieties in urban areas, but

not in rural commercial plantations of dwarf Cavendish-

type bananas [45, 59]. R. A. Syed of the Commonwealth

Institute of Biological Control (part of CABI) studied the

parasitoids of E. thrax in Sabah [188] and sent shipments

of larval parasitoids Scenocharops sp. and Cotesia erionotae,

and egg parasitoids Ooencyrtus pallidipes and Agiommatus

sp. nr. sumatraensis to Mauritius in 1971–1973. All four

species were released in Mauritius. Cotesia erionotae

and O. pallidipes became established, and although there

were recoveries of Agiommatus sp. it did not persist. The

parasitoids quickly provided effective biological control.

In 1975 a cyclone severely damaged banana plants and

drastically reduced banana skipper and parasitoid popu-

lations which, nevertheless, built up again in 1976. None

of the parasites established earlier was recovered in the

two years following the cyclone, but damage to banana by

the skipper was recorded as being very low in 1978 and it

remained uncommon. Davis and Barnes [60] report that

specimens of E. torus (as E. thrax) were taken on the west

coast of Mauritius in 1979, but it was not seen elsewhere

between the years 1976 and 1980, so that its status

was considered unclear. Williams [189] considered it (as

E. thrax) ‘uncommon for reasons that are not clear’.

Collectors from the African Butterfly Research Institute

found early stages in 2014 (T. C. E. Congdon, personal

communication, 2014), so E. torus continues as an

uncommon species of no significant pest status in

Mauritius, almost certainly kept under effective biological

control by its introduced parasitoids. None of the other

species of Hesperiidae found on Mauritius are known

to be parasitized by any of the parasitoid species intro-

duced against E. torus (J. Monty, personal communication,

1987 in 45).

Based on the programme already carried out

for Mauritius (purportedly on E. thrax but actually on

E. torus) when E. thrax appeared in Hawaii, a biological

control programme was immediately started [45]. In

1973, O. pallidipes was introduced from Guam where

it seemed to be an accidental introduction itself, and in

1974 C. erionotae was introduced and released, initially

from Thailand (early 1974) and then from Sabah (late

1974). Based on the information presented above (e.g.

Tables 3 and 4), the population of C. erionotae from Sabah

would have been collected from E. thrax only, since

E. torus does not occur there, whereas that from Thailand

is likely to have been reared from both E. thrax and

E. torus. Control in Hawaii was rapid, and the strain of

C. erionotae from Thailand was probably already estab-

lished and bringing the pest under control before that

from Sabah was released. In 1974, C. erionotae cultures of

the Thailand strain were sent from Hawaii to Guam, and

from Guam to Saipan in 1974–75, in both cases giving

good control [45, 49]. Both parasitoids were also sent

from Hawaii to Taiwan for the control of E. torus; they

were released in 1987 and both became established

[182]. The impact has not been fully reported, although

Teruya [138] found E. torus to have ‘a low pest status’,

infestation ranging from 0.1 to 7% with an average of 4%,

but he did not find the introduced parasitoids.

Erionota thrax may have spread into Irian Jaya before

1983 as it appeared by the border in north-western

mainland PNG in 1983, and then spread rapidly through

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Table 2 Parasitoids and diseases of Erionota thrax updated from Waterhouse and Norris [45]. Taxonomy adjusted from Noyes [141] and Yu [142]. Records of eggparasitoids of E. thrax in the Philippines [143] are not included, as the host eggs are listed from coconut and so probably are not E. thrax.

Family Species Location Reference Comment

Chalcidoidea, Chalcididae Brachymeria albotibialis(Ashmead)

Philippines Pupa L. B. Uichancoeditorial footnotein [144]

Recorded from several families of Lepidoptera

Chalcidoidea, Chalcididae Brachymeria euploeae(Westwood)

Indonesia,unspecified

Pupa [15, 135, 145] Widespread in South-east Asia; highly polyphagous on Lepidoptera

Chalcidoidea, Chalcididae B. euploeae (Westwood) Indonesia,west Java

Pupa [135] Common; 40–60% pupal parasitism in combination with Xanthopimplasp. This is likely to represent a mixture of B. lasus and B. thracis, judgingfrom the results of Erniwati and Ubaidillah [146]

Chalcidoidea, Chalcididae Brachymeria lasus (Walker) Indonesia, Java Pupa [146]Chalcidoidea, Chalcididae B. lasus (Walker) Indonesia, Sumatra Pupa [147, 148] Gregarious, nine adults per host. Common and widespread; together

with B. thracis 5–20% pupal parasitism. Common hyperparasitoid ofCasinaria sp.

Chalcidoidea, Chalcididae B. lasus (Walker)(=B. obscurata (Walker))

USA, Hawaii Pupa [136] A widespread and polyphagous species

Chalcidoidea, Chalcididae Brachymeria sp. Papua New Guinea,mainland

Pupa [58] < 5% pupal parasitism

Chalcidoidea, Chalcididae Brachymeria sp. nrmarginicollis

Malaysia (Sabah) Pupa [149] This is not a valid species; perhaps an error for B. marginata(i.e. B. lasus)

Chalcidoidea, Chalcididae Brachymeria sp. nr.obscurata (Walker)

Malaysia (Sabah) Pupa [149] This is likely to be B. lasus, of which obscurata is a synonym.Brachymeria spp. cause 16% pupal parasitism

Chalcidoidea, Chalcididae Brachymeria thracis(Crawford)

Indonesia, Java Pupa [146] Close to Brachymeria euploeae; earlier records of that species may wellrefer to this

Chalcidoidea, Chalcididae B. thracis (Crawford) Indonesia, Sumatra Pupa [147, 148] Gregarious, nine adults per host; 9% parasitismChalcidoidea, Encyrtidae Leurocerus ovivorus Crawford Malaysia (Sabah) Ovum [149] Described from two spp. of Nymphalidae. 0.5% egg parasitism [149]Chalcidoidea, Encyrtidae Ooencyrtus pallidipes (Ashmead) Indonesia, Java Ovum [146, 150]Chalcidoidea, Encyrtidae O. pallidipes (Ashmead)

(= Ooencyrtus erionotae Ferriere)Indonesia, Sumatra Ovum [147, 148, 151] Gregarious, 2.8 adults per egg. Common and widespread; together

with Pediobius erionotae 10–40% egg parasitismChalcidoidea, Encyrtidae O. pallidipes (Ashmead)

(= Ooencyrtus erionotae Ferriere)Indonesia,unspecified

Ovum [15, 145]

Chalcidoidea, Encyrtidae O. pallidipes (Ashmead)(= Ooencyrtus erionotae Ferriere)

Malaysia (Sabah) Ovum [149] 4% egg parasitism

Chalcidoidea, Encyrtidae O. pallidipes (Ashmead)(= Ooencyrtus erionotae Ferriere)

Papua New Guinea,mainland

Ovum [58, 150] 26–33% egg parasitism

Chalcidoidea, Encyrtidae Ooencyrtus sp. (not erionotaeFerriere)

Papua New Guinea,mainland

Ovum [58]

Chalcidoidea, Eulophidae Elasmus brevicornis Gahan Indonesia, Java Larva [97, 152, 153] Polyphagous, facultative hyperparasitoidChalcidoidea, Eulophidae Elasmus philippinensis Ashmead Philippines Larva [153, 154]Chalcidoidea, Eulophidae Elasmus sp. Indonesia, Sumatra L2 [147] Gregarious, 12 adults per host. Only one recordChalcidoidea, Eulophidae Pediobius erionotae Kerrich Indonesia, Java Ovum [146]Chalcidoidea, Eulophidae P. erionotae Kerrich Indonesia, Sumatra Ovum [147, 148, 151] Common and widespread; together with Ooencyrtus pallidipes

10–40% egg parasitism [147]; 16–39% parasitism [151]Chalcidoidea, Eulophidae P. erionotae Kerrich Malaysia, Sabah;

Indonesia, JavaOvum [155] Original description; holotype peninsular Malaysia

Chalcidoidea, Eulophidae P. erionotae Kerrich(= Pediobius sp.)

Malaysia, Sabah Ovum [149] 30% egg parasitism. Syed’s material is included in the type series

Chalcidoidea, Eulophidae P. erionotae Kerrich(= Pediobius sp.)

Malaysia, Sabah Larva [149] Syed’s material is included in the type series, although Kerrich [155]would seem to have reservations that this species attacks ova andlarvae

Chalcidoidea, Eulophidae Pediobius sp. Malaysia, Sabah Pupa [149]Chalcidoidea, Eulophidae Sympiesis sp. Indonesia, Java L2 [146]Chalcidoidea, Eulophidae Sympiesis sp. Malaysia (Sabah) Larva [149]Chalcidoidea, Eupelmidae Anastatus sp. Indonesia, Java Ovum [146]Chalcidoidea, Eupelmidae Anastatus sp. Indonesia, Sumatra Ovum [147, 151] Local and rare; 0.2–2.6% mortality

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Chalcidoidea, Eupelmidae Anastatus sp. Indonesia, unspecified Ovum [145]Chalcidoidea, Eupelmidae Anastatus sp. Papua New Guinea,

mainlandOvum [58]

Chalcidoidea, Eupelmidae Anastatus sp. USA, Hawaii Ovum [136]Chalcidoidea, Pteromalidae Agiommatus sp. Indonesia, unspecified Ovum [15, 145]Chalcidoidea, Pteromalidae Agiommatus sp. nr

sumatraensis CrawfordMalaysia (Sabah) Ovum [149] 2% egg parasitism

Chalcidoidea, Pteromalidae Agiommatus sumatraensis Crawford Indonesia, Java Ovum [146]Chalcidoidea, Pteromalidae A. sumatraensis Crawford Indonesia, Sumatra Ovum [147, 151] Localized, causing 0–9% mortalityChalcidoidea, Trichogrammatidae Trichogramma sp. Guam Ovum [156]Chalcidoidea, Trichogrammatidae Trichogramma sp. USA, Hawaii Ovum [136]Ichneumonoidea, Braconidae Cotesia erionotae

(Wilkinson)Indonesia, Java L4–L5 [146]

Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Indonesia, Sumatra L4–L5 [147, 148] Gregarious, 57 adults per host. Common and widespread.Hyperparasitized commonly by Eurytoma sp. A and once byPediobius elasmi (Ashmead)

Ichneumonoidea, Braconidae C. erionotae (Wilkinson)(=Apanteles erionotae)

Indonesia, unspecified Larva [15, 135, 145]

Ichneumonoidea, Braconidae C. erionotae (Wilkinson)(=Apanteles erionotae)

Indonesia, west Java Larva [135] Up to 50% larval parasitism

Ichneumonoidea, Braconidae C. erionotae (Wilkinson)(=Apanteles erionotae)

Malaysia (Sabah) Larva [149] 25% larval parasitism; heavily hyperparasitized by Eurytoma sp.,which in turn is hyperparasitized by Pediobius sp. and P. elasmi(Ashmead) (= P. lividiscutum (Gahan)). Presumably the Pediobius sp.is P. erionotae Kerrich; Syed’s material is included in the type series,although Kerrich [155] has strong reservations that this species attacksova, larvae and parasitoids of E. thrax

Ichneumonoidea, Braconidae C. erionotae (Wilkinson)(=Apanteles erionotae).

Indonesia, Sumatra Larva [157]

Ichneumonoidea, Ichneumonidae Casinaria sp. Indonesia, Java L3–L4 [146]Ichneumonoidea, Ichneumonidae Casinaria sp. Indonesia, Sumatra L2–L4 [147, 148] Common and widespread; 10–80% parasitism. Commonly

hyperparasitized by B. lasus, and rarely by Eurytoma sp. B andTheronia zebra

Ichneumonoidea, Ichneumonidae Charops sp. Indonesia, Java L3–L4 [146]Ichneumonoidea, Ichneumonidae Charops sp. Indonesia, Sumatra L3 and L4 [147]Ichneumonoidea, Ichneumonidae Echthromorpha agrestoria

(= E. fuscator)USA, Hawaii Larva [136]

Ichneumonoidea, Ichneumonidae Scenocharops sp. Malaysia (Sabah) Larva [149] 17% larval parasitismIchneumonoidea, Ichneumonidae Theronia zebra-zebra Vollenhoven Indonesia, Java Pupa [146] Polyphagous on pupae of several families of LepidopteraIchneumonoidea, Ichneumonidae T. zebra-zebra Vollenhoven Indonesia, Sumatra Pupa [147, 148] Local and rare; facultative hyperparasitoid of Theronia zebra.Ichneumonoidea, Ichneumonidae Xanthopimpla gampsura Krieger Indonesia, Java Pupa [146] Known from at least three palm-feeding HesperiidaeIchneumonoidea, Ichneumonidae X. gampsura Krieger Indonesia, Sumatra Pupa [147, 148] Common and widespreadIchneumonoidea, Ichneumonidae Xanthopimpla regina

(=Xanthopimpla sp.)Malaysia (Sabah) Pupa [149] 7% pupal parasitism

Ichneumonoidea, Ichneumonidae Xanthopimpla sp. Indonesia, unspecified Pupa [15, 145] Most probably this represents Xanthopimpla gampsura Krieger from JavaIchneumonoidea, Ichneumonidae Xanthopimpla sp. Indonesia, west Java Pupa [135] Common; 40–60% pupal parasitism in combination with Brachymeria

euploeae. Almost certainly this represents Xanthopimpla gampsuraKrieger

Ichneumonoidea, Ichneumonidae Xanthopimpla sp. ‘Malay Archipelago’ Not stated [97] Probably this represents Xanthopimpla gampsura Krieger from JavaPlatygastroidea, Scelionidae Telenomus sp. nr. attaci Malaysia (Sabah) Ovum [149]Diptera, Phoridae Unidentified Indonesia, Sumatra Pupa [147] UncommonDiptera, Sarcophagidae Unidentified Indonesia, Sumatra L5 [147] UncommonDiptera, Sarcophagidae Unidentified Malaysia (Sabah) Pupa [149] 5% pupal (or larval–pupal) parasitismDiptera, Tachinidae Unidentified Indonesia, unspecified Pupa [135]Diptera, Tachinidae Unidentified Malaysia (Sabah) Larva [149]Diptera, Tachinidae Blepharipa sp. Indonesia, Sumatra L5 [147] UncommonDiptera, Tachinidae Palexorista solensis (Walker) Indonesia, Sumatra L3–L5 [147] UncommonFungus Cordyceps sp. Malaysia (Sabah) Larva [149]

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the island over five years, and started to colonize

the other islands of PNG [58]. It spread together with its

egg-parasitoid, O. pallidipes. However, control was not

adequate with the egg parasitoid alone, and so C. erionotae

from Guam was introduced after quarantine safety tests

[48]. This rapidly resulted in good biological control of

the pest. The programme in PNG was supported by

ACIAR (Australian Centre for International Agricultural

Research), who subsequently funded assessments of the

benefits to PNG and Australia [47] and the impact on

poverty in PNG [190].

Waterhouse et al. [47] extrapolated from several

studies on defoliation and fruit loss indicating that more

than 16% defoliation will cause loss of yield, and 50%

defoliation will cause 28% yield loss. As the banana skipper

spread, it destroyed an average of some 60% of banana

leaves, leading to both a serious delay in fruit maturation

and reduced weight of banana bunches. After C. erionotae

was introduced and established, there was a reduction in

the estimated average leaf damage from 60 to 5%. The

losses due to E. thrax each year were calculated at 30% of

crop, of which 95% was saved by the biological control

introduction. Using a discount rate of 5% per year over

the 30 year period, the net present value of lost pro-

duction due to E. thrax amounts to approximately

A$301.8 million. The value of damage prevented by bio-

logical control is estimated at A$201.6 million.

The reduction of banana skipper abundance by 90%

in Southern PNG has correspondingly reduced the

chance of adults invading not only Australian islands in

the Torres Strait but also the Australian mainland.

The successful biological control programme in PNG

has therefore provided significant benefits to banana

production in Australia. Assuming similar levels of damage

in Australia as in PNG, these benefits were estimated to

be A$223 million to the year 2020. The assessment

of benefits did not consider the social and cultural uses

of banana leaves, nor the possible benefits in Irian Jaya, to

which the parasitoid almost certainly spread.

Bauer et al. [190] considered the impact of the

PNG biological control programme on poverty. Poverty

through income deprivation is a significant problem in

PNG. Estimates from a 1996 household income survey

suggested that around 30% of the PNG population have

household incomes below the poverty line (i.e. income is

not sufficient to sustain 2200 calories per day of food

consumption and cover the cost of essential non-food

items). The biological control of E. thrax has had a sig-

nificant beneficial impact on the effective incomes of most

PNG households by improving the supply of bananas

and by lowering the price of bananas to purchasers.

The improvement in income due to biological control

measures against the banana skipper would put between

6000 and 15 000 rural farmers (depending on assump-

tions) above the poverty line who would otherwise have

been below it. The successful implementation of the pro-

ject has also contributed to lower urban banana prices,Table

3ParasitoidsanddiseasesofErionota

torus.Taxonomyadjustedfrom

Noyes[141]andYu[142]

Family

Species

Location

Reference

Comment

Chalcidoidea,Encyrtidae

Leurocerushongkongensis

SubbaRao

China,HongKong

Ovum

[9,158]

Solitary

species

Chalcidoidea,Encyrtidae

Ooencyrtuspallidipes(Ashmead)

(=OoencyrtuserionotaeFerriere)

Mauritius

Ovum

[45,159]

Introducedto

anareawhere

itis

anticipatedonly

E.torusoccurs

Chalcidoidea,Eulophidae

Elasmusphilippinensis

Ashmead

Taiwan

Larva

[131,160]

Ectoparasitoid

Chalcidoidea,Eupelm

idae

Anastatussp.

Japan,Okinawa

Larva

[161]

Chalcidoidea,Pteromalidae

Agiommatuserionotae

China,Fujian

Ovum

[150,162]

Chalcidoidea,Trichogrammatidae

Unidentified

Japan,Okinawa

Ovum

[161]

Chalcidoidea,unspecified

Unidentified

China,HongKong

Ovum

[121]

Ichneumonoidea,Braconidae

Cotesia

erionotae(W

ilkinson)

(=Apanteleserionotae).

Mauritius

Larva

[45,159]

Introducedto

anareawhere

itis

anticipatedonly

E.torusoccurs

Ichneumonoidea,Ichneumonidae

Xanthopim

pla

sp.reginagroup

China,HongKong

Pupa

[9]

Diptera,Tachinidae

Unidentified

Taiwan

Pupa

[138]

Diptera,Tachinidae

Exorista

japonica(Townsend)

Japan,Okinawa

Pupa

[161]

Unspecifieddisease

Japan,Okinawa

Larva,pupa

[161]

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Table 4 Parasitoids and diseases of Erionota thrax or torus updated from Waterhouse and Norris [45]. Taxonomy adjusted from references given, Noyes [141] andYu [142].

Family Species Location Reference Comment

Chalcidoidea, Chalcididae Brachymeria albotibialis(Ashmead)

India Pupa [163]

Chalcidoidea, Chalcididae B. albotibialis (Ashmead) Thailand Pupa [45, 164]1

Chalcidoidea, Chalcididae B. albotibialis (Ashmead) Malaysia, peninsular Malaysia Pupa [125, 126, 128] The commonest pupal parasitoidChalcidoidea, Chalcididae Brachymeria euploeae (Westwood) Thailand Pupa [45, 164–166]1

Chalcidoidea, Chalcididae B. euploeae (Westwood) India Pupa [163]Chalcidoidea, Chalcididae Brachymeria lasus (Walker)

(= Chalcis marginata (Cameron))Malaysia, peninsular Malaysia Pupa [167, 168] Susainathan is incorrect to refer to this

pupal parasitoid as a larval parasitoidChalcidoidea, Chalcididae B. lasus (Walker) (=Brachymeria

marginata (Cameron))Thailand Pupa [45, 164–166]1

Chalcidoidea, Chalcididae Brachymeria thracis (Crawford) India Pupa [163]Chalcidoidea, Encyrtidae Ooencyrtus pallidipes (Ashmead)

(= Ooencyrtus erionotae Ferriere)Malaysia, peninsular Malaysia Ovum [125–128, 169] This is the original description from

ova of E. thrax and an unidentifiedlepidopteran on wild plantain. Thecommonest egg parasitoid (51%)

Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Nepal Ovum [150]Chalcidoidea, Encyrtidae O. pallidipes (Ashmead)

(= Ooencyrtus erionotae Ferriere)Thailand Ovum [45, 164–166]1

Chalcidoidea, Eulophidae Elasmus philippinensis Ashmead ‘Malay Archipelago’ [Malaysiaand/or Philippines]

Not stated [97] Parasitic on at least three families ofLepidoptera

Chalcidoidea, Eulophidae Elasmus sp. Malaysia, peninsular Malaysia Larva [125] < 1% parasitismChalcidoidea, Eupelmidae Anastatus sp. Thailand Ovum [166]Chalcidoidea, Eulophidae Pediobius erionotae Kerrich Malaysia, peninsular Malaysia Ovum [155] Original description; holotype

peninsular MalaysiaIchneumonoidea, Braconidae Cotesia erionotae (Wilkinson)

(=Apanteles erionotae).Malaysia, peninsular Malaysia Larva [125–128,

157, 168]The commonest larval parasitoid

Ichneumonoidea, Braconidae C. erionotae (Wilkinson)(=Apanteles sp.?)

Malaysia, peninsular Malaysia Larva [170] This record of ‘Apanteles sp.?’is assumed to be C. erionotae,which was not described until 1928

Ichneumonoidea, Braconidae Cotesia erionotae(Wilkinson)(=Apanteles erionotae)

Thailand Larva [45, 164–166]1

Ichneumonoidea, Ichneumonidae Scenocharops sp. Thailand Larva [45, 164–166]1

Ichneumonoidea, Ichneumonidae Xanthopimpla sp. Malaysia, peninsular Malaysia Pupa [168]Ichneumonoidea, Ichneumonidae Xanthopimpla sp. Thailand Pupa [166]Platygastroidea, Scelionidae Unidentified ‘scelionid’ Thailand Ovum [45, 164]1

Diptera, Phoridae Melaloncha sp. Malaysia, peninsular Malaysia Larva(L3, L4), pupa

[125] < 1% larval parasitism, 1% pupalparasitism

Diptera, Tachinidae Bessa remota (Aldrich) Thailand Larva [45, 164]1

Diptera, Tachinidae Exorista sp. Thailand Larva [45, 164]1

Diptera, Tachinidae Unidentified Thailand Larva [45, 164]1

Bacteria Unidentified Thailand Larva [45, 164]1

Virus Unidentified Thailand Larva [45, 164]1

1Napompeth [164] has not been examined and the information presented here is based on that given in Waterhouse and Norris [45].

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Table 5 Introductions for the biological control of E. torus and E. thrax, updated from Waterhouse and Norris [45]

Target Country Species Released Source Result References

E. torus Mauritius Agiommatus sp.nr. sumatraensis

1971–1972 Malaysia (Sabah) Recoveries, but notestablished

[149, 179], J. Monty, personalcommunication, 1987 (in [45])

Cotesia erionotae 1971–1974 Malaysia (Sabah) Established [149, 159, 179, 180]Ooencyrtus pallidipes 1971–1972 Malaysia (Sabah) Established [149, 159, 179]Scenocharops sp. 1971–1974 Malaysia (Sabah) Not established [149, 179, 180, 181], J. Monty,

personal communication (in [45])E. torus Taiwan C. erionotae 1987 Thailand via Hawaii Established [182]

O. pallidipes 1987 Guam via Hawaii Established [182]E. thrax Hawaii C. erionotae 1973–1974 Thailand Established [136, 183, 184]

1974 Malaysia (Sabah) Unknown (no way todistinguish from Thailandstrain already becomingestablished)1

[136, 180, 184]

O. pallidipes 1973 Guam2 Established [183, 184]Scenocharops sp. 1974 Malaysia (Sabah) Not established [136, 180, 184]

E. thrax Guam3 C. erionotae 1974 Thailand via Hawaii Established [49, 156, 185, 186]4

E. thrax Saipan C. erionotae 1974–1975 Thailand via Guam Established [49, 156]E. thrax Papua New

Guinea (mainland)C. erionotae 1990 Thailand via Hawaii

and GuamEstablished [48]

1This could be done today using molecular methods.2Ooencyrtus pallidipes seems to have spread accidentally to Guam without deliberate human assistance.3A Brachymeria sp. was introduced from Papua New Guinea for the biological control of Pericyma cruegeri (Butler) (Erebidae), became established and has been reared from pupae of E. thrax[49]. It has been referred to as Brachymeria sp. [49], B. albotibialis [156, 186], B. euploeae [45] and B. lasus [187]; all three Brachymeria names are currently valid species [141], and all have beenreported to parasitize E. thrax in South-East Asia (Table 4).4Reference [186] not examined, but based on citation in Waterhouse and Norris [45].

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in that the fall in supply of bananas caused by the pest

would have forced banana prices up. Banana prices

are likely to have risen by about 15% had the biological

control of the skipper not been achieved. Preventing this

increase in market prices for bananas of about 15% by the

introduction of the biological control measures resulted

in about 28 000 purchasers of bananas living just above

the poverty line when they otherwise would have been

below it. The biological control program increased

effective incomes across all income levels of banana pro-

ducers and banana consumers.

Discussion

The available information on the classification, taxonomy,

food plants, natural enemies and biological control of two

species of Erionota that feed on Musa spp. and one that

feeds on palms have been presented. It is concluded that

E. thrax and E. torus feed as larvae on Musa spp., and to a

limited extent some related species of Zingiberales, but

records of these species from palms and other plant

groups are in error. The records from palms are now

attributed to E. acroleuca. This distinction is important in

order to understand the ecology of these crop-feeding

butterflies so that management strategies are appropriate.

Where E. thrax and E. torus occur together, economic

entomologists have hitherto treated them as a single

species. There is no published information to show

whether the occasional outbreaks of banana skipper in

this region where both species are indigenous are due

to one or both of these species. Similarly, it is not

known whether the associated parasitoids attack both

species equally or show preference for one or the other.

Nevertheless, biological control has been successful

without knowing about the preferences of parasitoids for

the two species, or whether the material used was col-

lected from one or both Erionota spp. in Thailand. As

E. torus in Mauritius was misidentified as E. thrax, it

was controlled using parasitoids of E. thrax from Sabah.

There are plenty of examples where a lack of under-

standing of the taxonomy of the target pest of a biological

control programme has led to a mis-match of parasitoids

and target (see examples in [191]), and the author firmly

supports the view that good taxonomy is critical to suc-

cess in many biological control programmes, but in the

case of Erionota spp. this does not seem to have been a

critical factor.

For the foreseeable future, it would be a sensible pre-

caution to ensure that voucher specimens are preserved

and documented for all observations on Erionota spp. All

three species should be considered quarantine risks for

tropical areas where they do not naturally occur or have

not already spread, and an improved understanding of

their ecology should improve pest risk assessments

relating to their spread in future.

Research Gaps

In light of the foregoing, the following research questions

and gaps are identified:

� Molecular methods, particularly barcoding, are needed

to confirm identifications of early stages and develop

identification tools.

� Further collections and reliable identification of

adults (by examination of the genitalia and/or barcod-

ing) are needed to test the conclusion that E. thrax

and E. torus feed only on Musa spp. and a few similar

species, whereas E. acroleuca feeds only on various palm

species.

� Material from the Snellen collection should be ex-

amined to see if some of their food plant records can

be aligned with modern taxonomy of Erionota spp.; in

particular should their records of E. thrax from palms

be associated with E. acroleuca, which they considered a

synonym of E. thrax.

� The life history and food plants of the other three

subspecies of E. thrax should be established, which may

help to clarify their taxonomic status.

� As yet, there is no reliable way to separate the early

stages of E. thrax, E. torus and E. acroleuca morpholo-

gically. Careful study and documentation of reared

material, supported by reared vouchers and barcoding

of individuals, is needed to see if diagnostic criteria can

be found.

� Careful observation and behavioural studies of how

the larvae construct their shelters on different food

plants may provide additional diagnostic characters.

� Once the larvae of E. thrax and E. torus can be dis-

tinguished, the host-parasitoid relations for the genus

can be clarified; do parasitoids accept both equally or is

there specialization for one or the other?

� The identity of E. thrax rather than E. torus in the

Andaman Islands may need confirmation. It does not

seem to be a significant pest in the Andaman Islands. If

this is correct, is it because of indigenous or ac-

cidentally introduced parasitoids, perhaps also attacking

E. acroleuca?

� Recognizing the possibility that E. torus may have spread

into areas where only E. thrax thrax was known, it

would be worth making fresh collections to assess the

situation, particularly in Borneo, Sumatra and Java.

� It is considered that E. torus is most probably the

only Erionota sp. in Mauritius, but further collection

should be made to confirm this. Erionota torus is present

in Mauritius at low densities, and seems to cause

no economic damage. It seems most likely that this

is due to the continuing action of the introduced

parasitoids, but this should be investigated. The

reasons for the low population density of E. torus in

Mauritius have implications for the quarantine risk

that it presents to La Reunion and other Afrotropical

areas.

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Matthew J. W. Cock 23

� The new report of E. torus devastating bananas in the

Western Ghats of India suggests that biological control

should be considered in this situation. However, a

careful risk analysis with regard to indigenous species of

Hesperiinae would be needed first. To support this, a

phylogenetic classification of the Hesperiinae would be

necessary, to identify the most closely related genera

which would be most at risk.

� Further collections are required in New Britain, PNG,

to clarify the record of an Erionota sp. feeding on oil

palm.

Acknowledgements

Dr Hideyuki Chiba (Research Associate, Bishop Museum)

and Dr Laurence G. Kirton (Forest Research Institute of

Malaysia, retired) generously shared information, views

and documentation. Dr Chiba and Dr Rienk De Jong

(National Museum of Natural History, The Netherlands)

kindly reviewed parts of the manuscript paper prior to

submission. I thank Dr Feng Zhang (CABI China Regional

Centre), Dr Hiroko Kamei (University of Dundee), and

Dr Tim Haye (CABI Europe-Switzerland) for translation

and interpretation of Chinese, Japanese and German texts

respectively. In addition to the library resources of the

African Butterfly Research Institute (ABRI), the

Biodiversity Heritage Library, the British Library,

CABI, and the Royal Entomological Society, I have been

helped by a variety of people with reprints and copies

of literature, including Dr Rienk De Jong, Dr Ahsol

Hasyim (Indonesia), Suli Khing, Dr Lum Keng Yeang and

Dr Soetikno S Sastroutomo (CABI SE Asia), Dr Banpot

Napompeth (Thailand), Dr Justin Okolle (IRAD/CARBAP/

UB, Cameroon), Dr Don Sands (CSIRO Australia,

retired), Dr K. Veenakumari (India), Dr Janny Vos

(CABI Netherlands), as well as all those scientists and

organizations who have generously made their

publications and information available via the internet.

I thank Dr Charles Dewhurst (PNG Oil Palm Research

Association, retired), Colin Congdon and Forest and

Kim Starr (http://www.starrenvironmental.com/) for

allowing me to use their images. The images of adult

E. torus (Figure 2) are of specimens in Hope Entomological

Collections, Oxford University Museum, and I thank

Dr James Logan and Ms Molly Carter who facilitated

my visit. The images of adults of E. acroleuca (Figure 3)

are of specimens in the collection of the Forest Research

Institute of Malaysia and I thank Phon Chooi-Khim for

facilitating my visit there. I also thank Dr Blanca Huertas

and Mr John Chainey who facilitated a visit to the Natural

History Museum, London. Preparation of this review was

partially funded by the CABI Development Fund, which

is supported by contributions from the Australian Centre

for International Agricultural Research, the UK’s

Department for International Development, the Swiss

Agency for Development and Cooperation and others.

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