CAB Reviews 2015 10, No. 007

A critical review of the literature on the pest spp. (, Hesperiidae): , 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: [email protected]

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 (Linnaeus), Evans and Erionota acroleuca (Wood-Mason and de Nice´ville) (=E. hiraca (Moore)). Erionota thrax and E. torus are shown to be pests of 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 , E. thrax and E. torus have not been distinguished in the economic literature. The 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, .

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 , which helped considerably in interpreting the literature for this review. The Erionota spp. material in the 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 (Figure 2), has a very similar biology, also occurs in South- East Asia and as an introduced in Mauritius, and Erionota thrax (Linnaeus) is well known as ‘the banana (Okinawa). Both have been the target of successful ’ or ‘banana leaf roller’ in South-East Asia, and biological control programmes where they have been as an introduced pest in various Pacific territories. It introduced. A third, Erionota acroleuca (Wood-Mason is amongst the largest members of the family Hesperiidae and de Nice´ville) (Figure 3) is similar in appearance but (Figure 1). A second species, Erionota torus Evans the larvae feed on palms. All three overlap in much

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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; life size, the scale bar is 1 cm. Male: SE , 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 University Museum). The specimens illustrated are less dark than those of the other two Erionota spp. illustrated because they have faded with age. Erionota torus often cannot be reliably distinguished from E. thrax thrax (Figure 1) without examination of the genitalia. of their distribution in South-East Asia and there is an Classification extensive literature which indicates considerable confu- sion as to the Erionota spp. concerned. In this review, Erionota Mabille is a genus in the subfamily Hesperiinae, I attempt to untangle this literature, correct some errors, family Hesperiidae. In his work on the Hesperiidae of and identify aspects needing further attention. Europe, Asia and Australia, Evans [1] grouped seven

http://www.cabi.org/cabreviews Matthew J. W. Cock 3

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

Oriental genera as the Erionota sub-group within the or cut sections of Musa leaves into a tube, either in a Plastingia group of Hesperiinae. In their work on the spiral (Gangara, ) or in a roll (Erionota). This genitalia of the Hesperiidae of South , Inoue´ and behaviour separates them from all other Hesperiinae Kawazoe´ [2] suggested affinities between Lotongus Distant familiar to the author, and supports their recognition as a ([1] Plastingia subgroup), Gangara Moore, Erionota, Matapa distinct group. Moore ([1] Erionota subgroup), Unkana Distant, There is information available on the early stages of Distant and Pirdana Distant ([1] Unkana subgroup) the additional genera that Inoue´ and Kawazoe´ [2] sug- based primarily on characters of the male genitalia. gested could be grouped with Erionota. Lotongus calathus Although Warren [3] treated Erionotini as a provisional (Hewitson) has an unusual biology as documented by tribe of Hesperiinae in his Ph.D. thesis on the classification Igarashi and Fukuda [11]. The distinctive ova are flattened of the Hesperioidea, this was a much enlarged group of with a grooved ring around the top. They were found laid genera, which in his subsequent co-authored publications singly, and the newly hatched larvae make individual [4, 5] were placed incertae sedis. leaf folds on their unidentified palm seedling food plant. Of the seven genera that Evans grouped in his Erionota The second instar larvae then aggregate to make a subgroup, no information has been located on the early communal shelter, which is always co-occupied by a small stages of species of Zela De Nice´ville, Ilma Swinhoe, Ge ant, Dolichoderus bituberculatus Mayr, which does not De Nice´ville or Pudicitia De Nice´ville, but there is infor- attack the larvae. The mature larva has a four-lobed black mation on Gangara, Erionota and Matapa. marking on the face, reminiscent of those of some African (Fabricius) feeds on various palms and the early Artitropa spp. [12, 13]. The larvae pupate in a stout stages have been illustrated in Igarashi and Fukuda [6] and cocoon within the shelter, and the pupa is pale brown Cock et al. [7] among others. (Moore) is with no striking features. Igarashi and Fukuda [11] docu- a bamboo-feeder [8], and the early stages are illustrated ment the life history of Unkana ambasa (Moore) and by Bascombe et al. [9] and Saji et al. [10]. Erionota spp. feed Unkana mytheca (Hewitson) on Pandanus odorifer (as on palms (Arecaceae), Musa spp. (, bananas and P. odoratissimus, Pandanaceae). The ova are smooth and plantains) and some related Zingiberales. Igarashi and domed, similar to those of many other Hesperiinae; Fukuda [6] illustrate the early stages of E. torus, E. thrax the larvae are translucent grey-green with dark, oval and E. grandis (Leech), while Bascombe et al. [9] also heads, and a dark anal plate in the case of U. ambasa; illustrate the early stages of E. torus. The leaf shelters the leaf shelters are cylindrical and made by rolling the of at least some members of all three genera are note- leaf along the axis of the mid-rib; the pupae are brown, worthy for being tubes made by rolling palm leaflets the proboscis reaching the cremaster. Rearing revealed

http://www.cabi.org/cabreviews 4 CAB Reviews that the all-female species Erionota harmachis (Hewitson) dissection. In particular, the strongly upturned, more is a synonym of the all-male species U. mytheca [14], heavily chitinized apical process of the valves of male indicating affinities in adult structure characters. Hidari E. torus, compared with the simple, blunt apical process irava (Moore) makes leaf shelters between leaflets of of E. thrax can often be distinguished. A recent student its palm hosts where small numbers of larvae cohabit thesis yet to be published [40] concluded that ‘the banana [11, 15, 16]. Pirdana hyela (Hewitson) and Pirdana distanti skippers, E. thrax and E. torus, are a monophyletic group (Staudinger) feed on spp. (Asparagaceae) which represent a single host shift from Arecaceae or [11]. The ovum is strongly ribbed; the head and body of Zingiberaceae to Musaceae’. the larva of P. hyela superficially resembles that of an There is some confusion regarding the correct name Artitropa sp. [12, 13], while that of P. distanti has a for E. acroleuca [19]. It was first described by Wood- similar head, but the body is dark with a yellow dorsal Mason and de Nice´ville [25] in August 1881 as Telegonus line and yellow dorsolateral transverse bars; leaf shelters acroleucus in the abstract of a paper which was not pub- are made by rolling the food plant leaf longitudinally; the lished in full until December that year when the combi- pupa has a short, blunt, frontal spike. The rather diverse nation Hesperia acroleuca was used [26]. In the interim, in life histories of representatives of these genera do not September, Moore [27] independently described the same suggest any obvious affinities with Erionota, unlike that species as Hesperia hiraca. The description in the abstract pointed out for Gangara and Matapa above. [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’, Taxonomy i.e. it meets the requirements for the publication of the new name and so has precedence over Moore’s name. The checklist in Table 1 presents the current position. Gender agreement has been applied under the ICZN [41], The species of immediate concern as pests are E. torus, i.e. E. acroleuca is correct rather than E. acroleucus fol- E. thrax and E. acroleuca. However, as some or all the lowing the treatments of De Jong and Treadaway [17] and other species also feed on Musacaeae and Arecaceae as Xue and Lo [19]. The male of E. acroleuca has the apex larvae, they are all of concern as potential quarantine of the forewing upper side white, which distinguishes it pests. from the males of E. thrax and E. torus. The female is Linnaeus [22] described Papilio thrax from Java, referring generally smaller than those of E. thrax and E. torus, but to an un-named image in Clerck (36, plate 42.2). Honey the white patch at the forewing apex is variable, and and Scoble [37] explain that there are two specimens some specimens are not reliably distinguishable except of E. thrax in the Linnean Collection, one of which has by examination of the genitalia. Fortunately, Inoue´ and been designated the lectotype, but as Mabille [20] also Kawazoe´ [2] again provide good drawings of both male pointed out, Clerck’s [36] image is of a different, unre- and female genitalia of this species (ssp. apicalis). lated species (a South American species of the genus The authorship of ssp. apicalis also needed clarification. Thracides). Xue and Lo [19] showed that when Evans [29] published Evans [21] described E. torus from Sikkim and gave the name as var. apicalis, this made it an unavailable characters to separate the adults from E. thrax based infraspecific name under Article 45.5.1 of the International on appearance and male genitalia. In his catalogue of the Code of Zoological Nomenclature [41], and it should Hesperiidae of Europe, Asia and Australia [1] he added his take the authorship from the first publication to use it as diagrammatic figures of the male genitalia. Males of E. torus a valid name, i.e. De Jong and Treadaway [17], as listed in (Figure 2) can usually but not always be distinguished Table 1. from those of E. thrax (Figure 1) by the more rounded Astictopterus harmachis Hewitson was described from distal margin of the forewing in E. torus compared to Sumatra [42], and was subsequently treated as a species the straighter margin and apically more pointed forewing of Erionota [1, 21]. However, it is now known to be the of E. thrax. Separation of the two species, particularly female of U. mytheca, which had previously only been females, is only reliable if based on examination of the known from the male [11, 14]. 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 Distribution and Spread documented and drawn by Inoue´ and Kawazoe´ [2] and available on the internet. Corbet et al. [39] include There are four subspecies of E. thrax recognized in figures of the male genitalia of both species, and the male South-East Asia: E. thrax thrax widespread from north-east genitalia and female venation of E. torus are illustrated to the Philippines, Sulawesi and Java, E. thrax mindana by Bascombe et al. [9] from . The diagnostic Evans in the southern Philippines but overlapping with characters of both male and female genitalia can usually be E. thrax thrax in the northern Philippines, E. thrax alexandra seen by brushing off the scales of the abdomen tip using a Semper from upland northern Luzon and E. thrax has- short, fine paint brush, obviating the need for traditional drubal Fruhstorfer from Kep. Sula (Mangole, Sanana),

http://www.cabi.org/cabreviews 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]

Genus Species Original genus of description Subspecies (or species if described Synonym as subspecies) Type locality1 Erionota Mabille, 1878 [20] torus Evans, 1941 [21] Erionota Kurseong, Sikkim [India] thrax Linnaeus, 1767 [22] Papilio Java [] ssp. thrax Linnaeus, 1767 [22] ssp. mindana Evans, 1941 [21] Erionota Mindanao [Philippines] http://www.cabi.org/cabreviews 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 Nice´ville, 1881 [25, 26] Telegonus S[outh] Andaman [Islands], India ssp. acroleuca Wood-Mason & De Nice´ville, 1881 [25, 26] =acroleucus Wood-Mason & De Nice´ville, 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, Philippines ssp. apicalis De Jong & Treadaway 1992 [17] Erionota acroleuca Bassein, Burma [= Pathein, ] =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, Philippines tribus 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] China ssp. grandis Leech, 1890 [32] ssp. alsatia Evans, 1939 [33] Erionota Likiang, Yunnan, China sybirita Hewitson, 1876 [34] Hesperia =mabillei Staudinger, 1889 [35] Erionota Palawan, Philippines 5 Cock W. J. Matthew 1Information in square brackets has been added where it was not included in the original description. 6 CAB Reviews North Moluku, Indonesia [1, 18, 43]. Of these, only thrax was recently reported from Palpa District, central E. thrax thrax is known to have spread beyond its indi- southern Nepal [62], but this needs confirmation in genous range. view of Smith’s [63] statement that only E. torus oc- In the Pacific E. thrax thrax spread to by 1957 curs in Nepal (R. Muniappan, personal communication, [44] and Saipan in the Northern Mariana Islands by 2014). the late 1960s [45], to Oahu, by 1973 [46], to Erionota torus was more restricted in distribution, but is Irian Jaya, east Indonesia some years before 1983 [47], now widely introduced. It is indigenous in northern India: to mainland Papua (PNG) in 1983 [45], an early report of E. thrax as common around Dehradun to New Britain, PNG, in 1991 [48], to Duke of York (=Dehra Dun) at the base of the Western Himalaya [64] and New Ireland islands, PNG, by 1994, and possibly to is considered to refer to E. torus since only specimens of Bougainville, PNG, about this time [47]. Muniappan [49] this species are present in the NHM from Dehra Dun [1]. refers to the original introduction into Guam as being It is also found in Nepal [63, 65] and Northeastern from the Philippines, based on both localities having India to Southern China and Peninsular Malaysia [1], but it American air force and naval bases, and the heavy civilian spread to Mauritius around 1968 (reported as E. thrax, but traffic between the Philippines and Guam (R. Muniappan, see above), Southern Japan in 1971 [66], Taiwan in 1986 personal communication, 2014). Within the Hawaiian [67, 68], the southern Philippines probably in the early Islands it spread from Oahu to Kauai and Maui in 1974 and 1980s (Cebu, Dinagat, Leyte, Mindanao, Negros, Samar; Hawaii, Molokoi and Lanai in 1975 [50, 51]. [18]), and has just been found in the Western Ghats The records from Guam, Saipan and Hawaii have been of India [69]. There is a single male specimen labelled referred to incorrectly as E. torus, e.g. Igarashi and Fukuda Sarawak in the NHM, but the absence of any other [11], Opler and Warren [52] and perhaps others. The records from Borneo suggests its presence there should origin of this error and its correction are outlined in be considered unconfirmed. It does not seem to be pre- Pelham [53]. Kawazoe´ and Wakabayashi [54] illustrated sent in Java or the rest of Indonesia. E. thrax as E. torus and stated that it occurs in Hawaii Erionota acroleuca occurs in the Andaman Islands and Guam. Snyder and Kumon [55] and Fukuda and Nicho as the nominate subspecies. Although Swinhoe [28] [56] clarified that E. thrax is indeed the species occurring described a synonym from the Nicobar Islands, Evans [1] in Hawaii, not E. torus. treats this as an erroneous locality. Erionota acroleuca apex Although Parsons [57] treats the population that is restricted to the Philippines [17, 18]. Erionota acroleuca spread into PNG as E. thrax hasdrubal, the specimen that apicalis was formerly treated as a synonym of E. acroleuca he illustrates is E. thrax thrax, based on the characters in apex, but De Jong and Treadaway [17] established that it is Evans [1, 21], and is treated as such by Waterhouse and a separate subspecies found from Myanmar and Northern Norris [45] and Sands et al. [58]. Apart from the records Vietnam, through the Malaysian peninsula to Borneo, from New Guinea, it is unclear to what extent E. thrax of Sumatra, Java and east to West Nusa Tenggara. Finally, either subspecies thrax or subspecies hasdrubal has spread E. acroleuca sakita is found on Sulawesi, and the islands to through eastern Indonesia, either naturally, or with acci- the east of Central Sulawesi [1, 43]. dental human assistance. It likely seems that the major translocations of E. thrax Erionota thrax was first reported in Mauritius in 1970 and E. torus were mediated by aircraft flights carrying [59], although it was suspected to have been introduced in infested plants or planting materials, infested leaves or 1968 [45]. The author has examined the three Mauritius adult butterflies. Ova occasionally occur on fruit in bun- specimens in the Natural History Museum, London ches, and so may also be transported internationally [70], (NHM), collected by P. M. H. Davis at Black River in 1979, but the challenge for the hatching larvae to reach Musa and reported in Davis and Barnes [60], and finds that all spp. leaves after arrival is very considerable. Military flights are E. torus, based on the visible external genitalia. The have been implicated or suggested as the route for original identification reported in Monty [59] was made the spread to Mauritius and Hawaii: E. torus is ‘thought to at the NHM, but only female specimens seem to have have been introduced in military aircraft based in Malaysia been provided, and at that time the distinctive female in 1968’ [45], and E. thrax was discovered in Hawaii at genitalia of E. thrax and E. torus had not been documented. Hickam Air Force Base, Oahu [46]. The spread of E. thrax Although it is not impossible that both species were thrax to Guam has been linked to American military bases accidentally introduced into Mauritius, in the absence on Guam and the Philippines, as well as heavy civilian of any specimens of E. thrax, it seems safe to assume that traffic between the two (R. Muniappan, personal com- only E. torus is present. munication). The fact that Guam and Saipan both had Veenakumari and Mohanraj [61] documented the strong military bases in the 1950s suggests how the sub- first record of E. thrax thrax from the Andaman Islands, sequent introduction to Saipan may have been facilitated, although it would be desirable to confirm this identifica- rather than to other islands in the region. During its tion by examination of the genitalia (K. Veenakumari, outbreak phase in PNG, E. thrax spread throughout the personal communication, 2014). The author was unable PNG mainland within 6 years, moving several hundred to locate any specimens in the NHM to do so. Erionota km per year [58]. This would have been due to a mixture

http://www.cabi.org/cabreviews Matthew J. W. Cock 7 of natural dispersal of gravid females, and accidental man- indications of errors. Accordingly, I present my analysis assisted transport of planting materials, leaves and adult in some detail below. Erionota thrax thrax and E. torus are butterflies. similar in adult appearance [1, 21] and life history [6, 39], What is less clear is the mechanism of spread between and where their ranges overlap, have almost certainly relatively nearby islands, e.g. through the Hawaiian Islands been confused – with each other and with other Erionota and from New Guinea to New Britain. Waterhouse spp., particularly E. acroleuca. Reports from localities and Norris [45] speculate that ‘the water gaps between where only E. thrax or only E. torus occur are more likely Southern PNG and Northern Australia are probably well to be reliable, but where both occur there will be within the flight range of adults. ... Southwest Pacific confusion between the two. The situation is rendered islands will be at risk if inter-island movement of fresh more difficult to interpret by the habit of many authors banana leaves is not prohibited. In any case adults may be of compiling food plant records from multiple sources able to fly to nearby islands and establish themselves.’ This without attribution, so that it is not clear whether any speculation by Waterhouse and Norris [45] has subse- particular observation, such as a food plant record, is an quently been presented in a pest risk assessment as original observation or repeated from an earlier source, ‘E. thrax has spread from Oahu to the other Hawaiian which may be misidentified, represent a different sub- Islands flying over water for distances of up to 150 km species, or be from a different geographical area [72]. [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 Food plants of E. thrax females flying these distances has been found. The fact that E. thrax and E. torus have not spread more widely I have found reports on the food plants and biology of already, e.g. within the parts of Indonesia where they E. thrax thrax only, and none on the other subspecies of do not occur naturally, or within the Northern Mariana E. thrax, although it seems likely that they will have a simi- Islands, suggests that such long distance flights over sea do lar food plant range. In its indigenous range, many refer- not occur. Similarly, the evolution of isolated subspecies ences (e.g. [15, 39, 73–77]) indicate that E. thrax feeds of E. thrax (above) indicates a species that is not highly primarily on Musa spp. (Musaceae), especially banana mobile. (M.  paradisiaca, = M. sapientum and M. paradisiaca) and Inter-island movement of boats seems a more likely M. textilis (abaca or Manila hemp). Sands et al. [58] sur- pathway of introduction, as Sands et al. [48] discuss: veyed E. thrax in PNG when it first spread through the ‘Lights on board boats are considered to be a means for island of New Guinea. They report collecting immature attracting the crepuscular adult E. thrax and allowing them stages of E. thrax from Musa  paradisiaca, M. textilis, other to be carried from one island to another. A specimen in Musa spp. (including sections Australimusa and Eumusa) the Natural History Museum, London, was collected and several varieties. Waterhouse and Norris [45] point on board a ship in 1929, near Seram (less than 200 km out that in its introduced range, E. thrax thrax is known west of New Guinea), outside of its native range. From only from cultivated and wild species of banana (based on the distinctive pattern of spots beneath the hind wings, personal communications of experienced entomologists this specimen probably came from Java. A specimen from who worked on the introduced pest), although their Port Moresby, PNG in 1961 and referred to by Parsons observation to this effect from Mauritius is actually based [57] probably travelled to PNG on board a ship since this on E. torus. However, records from other recorded food specimen pre-dated first establishment of E. thrax in plants, especially in the older literature, need critical North-western PNG by 22 years.’ examination. It is possible that different species of Erionota In view of the foregoing discussion and the confusion are responsible for the records on bamboo and palms. that has arisen, it is worth recognizing the possibility that Probably the oldest record of E. thrax as a pest in the last 65 years or so since Evans [1] completed his of banana (pisang, Musa paradisiaca) is that of Horsfield & review of the Hesperiidae of the region, E. torus could Moore [78] in A Catalogue of the Lepidopterous in the have spread to areas where only E. thrax thrax was known Museum of the Hon. East-India Company. As these obser- (or vica versa), and easily been overlooked. This possibi- vations were from Java, where E. torus is not reported, lity merits checking with recent material, for example they should be reliable. Soon afterwards, Distant [79] in in the island of Borneo where there is already a single Rhopalocera Malayana, repeats Horsfield & Moore’s [78] record of E. torus, and Sumatra and Java which are in observation on the food plant, although his illustration of relatively easy reach of Peninsular Malaysia. 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 Food Plants species illustrated is E. thrax, the observations could relate to either E. thrax or E. torus or both. Duport [81] refers to I have reviewed much of the literature on food plants E. thrax as a banana pest in Indochina, and this could refer of Erionota spp. and find considerable confusion and to both E. thrax and E. torus. Williams [82] describes the

http://www.cabi.org/cabreviews 8 CAB Reviews biology on banana on Luzon Island, Philippines, which sagu (sago palm), pinnata (as A. saccharifera) (sugar should be correctly associated with E. thrax. Beller and palm), and based on a verbal report guineensis (West Bhenchitr [83] include E. thrax thrax as a banana pest in African oil palm) as food plants. Keuchenius [92] and their list of insect pests of (Siam); this could be E. Leefmans [93] also report E. thrax as a pest in thrax or E. torus. Indonesia (Dutch East Indies). However, since Piepers There are many other food plant references compiled and Snellen [73] treat E. acroleuca as a synonym of E. thrax, in Robinson et al. [84] with sources, and in Vane-Wright it is suggested here that the records from palms are and De Jong [43] and CABI CPC [85] without sources. more likely to relate to E. acroleuca, which has since These include various palms (Arecaceae), bamboos been documented as a palm pest (below). Specimens from and grasses (), Heliconia spp. (Heliconiaceae), P. C. T. Snellen’s collection (which includes material from Curcuma angustifolia (Zingiberaceae) and traveller’s palm M. C. Piepers) are in the Naturalis Biodiversity Centre, (Ravenala madagascariensis, Strelitziaceae), which are Leiden, Netherlands but they cannot be examined assessed below. at present during rearrangements of the collection Robinson et al. [84] cite Bhumannavar et al.’s [86] (R. De Jong, personal communication, 2014); perhaps in Insects of Agricultural Importance in Andaman and Nicobar the future examination of this collection will clarify some Islands that E. thrax feeds in rolled leaves on the intro- of these records. It is furthermore suggested that Piepers duced R. madagascariensis. Veenakumari et al. [87] made and Snellen’s records are the basis of the continuing new observations and reported the food plants of E. thrax references to palms as food plants of E. thrax in the lit- thrax in the Andaman Islands as only MusaÂparadisiaca erature. Seitz [94] lists sugar-cane, coconut, Rhapis and and R. madagascariensis. As Veenakumari et al. [87] sepa- in addition to Musa spp.; the similarity to the rate their food plant records of E. thrax thrax from those species listed in Piepers and Snellen [73] suggests that of E. acroleuca, and the leaf structure of R. madagascariensis their record of A. pinnata (as A. saccharifera) was mistaken is suitable for leaf rolling as practiced by E. thrax, this for sugar-cane. record is accepted here, even though Sands et al. [58] did P. B. Richards [95] and R. M. Richards [96] reported not find immature stages of E. thrax on this food plant in H. irava Moore and E. thrax as hesperiid pests of coconut PNG. Mau and Kessing [88] give Strelitzia (Strelitziaceae) in West Malaysia, but Corbett [74] in his Insects of Coco- as a food plant genus in Hawaii, but this does not seem to nuts in Malaya considers the two have been confused, that have been formally published. Nevertheless, it is a possible H. irava is the coconut pest and E. thrax is only a banana food plant, although perhaps as a spill-over effect (below). pest. Corbet et al. [39] in The Butterflies of the Malay Similarly, Mau and Kessing [88] list the genus Heliconia Peninsula highlight Musa spp. as the food plants of E. thrax as a food plant in Hawaii. In PNG, the ornamental native thrax and E. torus, point out that the two do not differ in species Heliconia papuana is a widespread host for E. thrax life history, and include coconut and sugar palm (i.e. thrax (F. M. Dori and D. P. A. Sands, personal commu- A. pinnata) as reported food plants. nication, in [47]), although the introduced Neotropical In Les Insectes des Palmiers, Lepesme [75] considers Heliconia bihai is not, and is not in the laboratory [58]. E. thrax to be very common on banana, but rarer on Elaeis, Igarashi and Fukuda [6] list C. angustifolia (wild or Indian Cocos, Metroxylon, , [Arecaceae], arrowroot; Zingiberaceae) as a food plant. No other sugar-cane and bamboo [Poaceae]’, citing De Joannis source has been located for this record (which is repeated (in [81]), Richards [95], Seitz [94], Williams [82], in 43), but given that C. angustifolia has large leaves, similar Dammerman [97], Corbett [74] and Beller and Bhenchitr to those of Musa, Ravenala and Heliconia, it seems a pos- [83]. As noted above, Duport [81], Williams [82] and sible food plant. A record of Canna (Cannaceae) as a food Beller and Bhenchitr [83] report E. thrax (and/or E. torus) plant genus in Hawaii [88] could be interpreted in this as a banana pest. Richards [95], Seitz [94] and Corbett way, but could also be due to a spill-over effect. [74] are also discussed above. Dammerman [97] treated At this point it is worth noting that Musaceae E. thrax as a banana pest, also found on ‘Manila hemp, and Strelitziaceae are closely related families in the bamboo, coconut, oil palm and other palms’ but does not order Zingiberales, while Heliconiaceae, Cannaceae and indicate his sources. None of these sources listed by Zingiberaceae although also placed in Zingiberales are Lepesme [75] are the source of records on Calamus and less closely related [89, 90]. Oligophagous species are Cyrtostachys. mostly, but not always, more likely to feed on species Hutacharern and Tubtim [98] listed E. thrax thrax from closely related to their main food plant, than on those less Calamus sp. (rattan) in Thailand, and Johari and Che Aziz closely related (e.g. [91]). The remaining food plant [99] listed it as a pest of Calamus trachycoleus in Peninsular records are of species from other plant orders, and Malaysia. Erionota thrax was listed as a pest of C. manan correspondingly less likely. in Peninsular Malaysia by Norani et al. [100], although The original records of E. thrax [thrax] feeding on in a subsequent treatment of rattan pests in Peninsular Arecaceae seem to be those of Piepers and Snellen [73] Malaysia, Maziah et al. [101] refer to and illustrate from Java, who in addition to Musa spp., list Cocos nucifera the larva as E. torus, rather than E. thrax. In their survey (coconut), Rhapis excelsa (as R. flabelliformis), Metroxylon of hesperiids associated with C. manan, Steiner and

http://www.cabi.org/cabreviews Matthew J. W. Cock 9 Aminuddin [102] also record E. torus, but later correct placed on M. Â paradisiaca (as M. sapientum) as controls, this to E. acroleuca apicalis (as E. hiraca apicalis) [103]. This began feeding. Furthermore, the leaf shelters that E. thrax correction was based on the advice of L. G. Kirton, who and E. torus make on banana (see below) involve beha- indicates that E. acroleuca (as E. hiraca) is the Erionota sp. viour patterns that would not be readily transferable to that feeds on Calamus spp. and other palms. Hence, in his a pinnate leaved palm. Hence, the evidence indicates recent Naturalist’s Guide, Kirton [77] refers to E. thrax that E. thrax thrax does not use palms as food plants, and it thrax feeding on bananas only. is anticipated that all the records from Arecaceae, should In ‘Pests of Oil Palms in Malaysia and their Control’ Wood be referred to E. acroleuca. It is unfortunate, therefore, [104] includes E. thrax as a minor pest, but the illustration that the records from palms have led to E. thrax being of the adult (plate XIVb above) is of a male E. acroleuca mentioned repeatedly as a pest of palms in the pest lit- apicalis with the distinctive white fore wing tips. There is a erature (e.g. [75, 110–115]). It is equally unfortunate that relatively recent record of E. thrax hasdrubal as a pest of the common name ‘palm redeye’ has been applied to this oil palm in New Britain [105]. The adult illustrated in the species. The oldest use of this name appears to derive paper is not E. thrax hasdrubal, but it is not clear whether from Evans [116, 117], and is assumed to be based on the it represents E. thrax thrax, E. torus, an individual of food plant records referred to above, ignoring the well- E. acroleuca lacking the pale forewing apex, or some other documented damage to bananas. Although Brigadier taxon of Erionota. Of these, only E. thrax thrax is known W. H. Evans was an excellent morphological taxonomist, from New Britain, but above we have discounted all there is almost no reference to food plants and early records of palms as food plants for this subspecies. It will stages in any of his works, so it seems likely that he was be necessary to examine and dissect voucher specimens not that well-informed on this aspect, hence the use of to confirm or clarify this record. this inappropriate name. I recommend that the use of this There are further records from Arecaceae catalogued common name should be discontinued for all species of by Robinson et al. [84]. In their ‘List of economic pests, Erionota to avoid future confusion. host plants, parasites and predators in West Malaysia There are also records from Poaceae. Robinson et al. (1920–1978)’ Yunus and Ho [106] include Caryota and [84] attribute a record of Chrysopogon to Sevastopulo’s as food plants of E. thrax, but do not specify [118] The Food Plants of Indian Rhopalocera, but this is the source for this record, and Pholboon [107] and incorrect, as Sevastopulo does not list this grass. Hutacharern and Tubtim [98] in their compilations of (sugar cane) is listed as food plant by plant pests in Thailand, list grandis as a food plant of Sevastopulo [118] based on Seitz [94], which it was sug- E. thrax. These records from palms have not been tracked gested above is an error for sugar palm. Robinson et al. back to their original source for this review, but below [84] attribute a record of Saccharum to Corbet et al. [39], we document that in West Malaysia E. acroleuca apicalis is but the food plant listed there is sugar palm, i.e. A. pinnata recorded to feed on Caryota spp. and E. sybirita on a Licuala as originally listed by Piepers and Snellen [73] and dis- sp., indicating that these records are likely to be based on cussed above. Robinson et al. [84] also cite A Manuscript misidentifications. Kitamura [108] reared a single Erionota List of Butterfly Hostplants in the Western Ghats and Southern sp. from a leaf shelter on coconut on Panay Island, India by H. Gaonkar (2000), which does not seem to have Philippines, and illustrated the larva, pupa and shelter been subsequently published, that Saccharum officinarum (a longitudinally rolled leaflet). He identified the resultant is a food plant of E. thrax thrax. Whether this is a repeat adult female as E. thrax. Could it have been another of one of the earlier errors or a new record, it is anyway species such as E. acroleuca or E. surprisa De Jong unlikely to be an original record as E. thrax is not reported and Treadaway? Kitamura [109] subsequently reared from this part of India, and E. torus has only recently been E. acroleuca apex from a palm and compares the larva found there. It is concluded that none of the published with his coconut-reared specimen, so clearly considered records of Poaceae as food plants for E. thrax should be them different. Nevertheless, given the concerns raised accepted. here, and the similarity of adult females, this record based Finally, Robinson et al. [84] cite Hutacharern and on a single female merits checking by examination of the Tubtim [98] that Butea monosperma (Fabaceae) is a food genitalia before accepting. plant of E. thrax thrax in Thailand, but this is an error for In their survey of E. thrax in New Guinea, Sands the record of Calamus sp. on the same page ([98], p. 41), et al. [58] specifically report that no immature stages and this record should not be perpetuated. were found on Ensete glaucum (Musaceae), H. bihai Interpreting these food plant records, it is worth (Heliconiaceae), R. madagascariensis (Strelitziaceae), remembering two facts documented in weed biological N. fruticans, Metroxylon sp., Calamus spp., C. nucifera, Areca control. Firstly, oligophagous species are mostly, but not catechu or E. guineensis (Arecaceae), even when infested always, more likely to feed on species closely related to Musa spp. plants were growing close by. Furthermore, their main food plant, than on those less closely related in the laboratory, groups of ten newly-hatched of E. thrax [91]. On this basis, Musaceae and Strelitziaceae are part of died without feeding when placed on fresh foliage of one clade of Zingiberales, while , Cannaceae H. bihai, C. nucifera and E. guineensis, whereas all larvae and Heliconiaceae are in the other [89]. Families from

http://www.cabi.org/cabreviews 10 CAB Reviews outside the order Zingiberales, i.e. Arecaceae in this case, and Aminuddin [103] subsequently indicate that this are much less likely to be suitable food plants. Secondly, record should be treated as E. acroleuca (=E. hiraca). when a successful weed biological control agent is first re- In a laboratory study in Taiwan, Tsai et al. [131] fed leased, there is often a population explosion, during which cohorts of each of the five instars of larvae of E. torus with there can be spill-over effects where feeding damage cut portions of leaves of banana, Strelitzia reginae (bird occurs on other plants, usually because insects have of paradise flower; Strelitziaceae), C. nucifera (coconut; strayed or fallen from the normal food plant onto adjacent Arecaceae), areca nut (A. catechu; Arecaceae), plants. The same thing may happen with any newly oldhamii (bamboo; Poaceae) and sugar cane (Saccharum introduced species, such as E. thrax and E. torus in their officinarum; Poaceae). Feeding was most extensive on exotic range, so that food plant records during a popu- banana, but almost as heavy on Strelitzia reginae; there was lation explosion will include species that are not normally very little feeding on coconut (none for the cohort of suitable, on which eggs will not normally be laid, or which instar 1), less on areca nut and none on bamboo or sugar will not sustain the full life cycle. Some of the observations cane. Average survival for all instar cohorts was 50% on S. reported, e.g. from Hawaii [88], may be of this nature. reginae, 5% on coconut and 0% on bamboo and sugar To summarize, it is concluded that the food plants of cane. This is a measure of the survival of larvae trans- E. thrax are restricted to Musa spp., perhaps some other ferred to a plant other than banana, and does not reflect Musaceae and a few related, similarly broad-leaved species what would happen following oviposition on that plant in in Heliconiaceae, Strelitziaceae and possibly Zingiberaceae the field. If there was no feeding in instar 1 on coconut, all (all Zingiberales). No convincing food plant records have larvae would die in the field, while the data does not allow been found of subspecies of E. thrax apart from the us to interpret what total mortality would be on S. regi- nominate subspecies. 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 Food plants of Erionota torus 21% of individuals completed their metamorphosis on banana at 20 and 30 C, respectively, most mortality As has become evident during this analysis of E. thrax, occurring in instars 1 and 2. It is difficult to know how to there has been confusion between E. thrax and E. torus interpret these two apparently conflicting results. How- where the two co-occur in mainland South-East Asia. ever, we can conclude that sugar cane, bamboo, areca nut Accordingly, food plant records without voucher speci- and coconut are unsuitable food plants, whereas S. reginae mens can only be accepted for E. torus with full confidence is less suitable than banana. where E. thrax does not occur. It is clear that like E. thrax, There are internet records of E. torus using Canna sp(p). E. torus is primarily a pest of Musa spp. Only E. torus (Cannaceae) as food plants (e.g. [132]), but these have not occurs in Hong Kong and adjacent China, and here it has been traced to any formal publication, and may well been documented to feed on Musa spp. but not on be derived from the records for E. thrax, when the name other plants since the late nineteenth century as E. thrax E. torus was misapplied to that species. [119–121], and more recently as E. torus (e.g. [9, 122]). As noted above and discussed below, the early stages Similarly, only E. torus appears to be present in the and leaf shelters of E. thrax and E. torus are very similar. Western Himalayas [1], so Mackinnon & de Nice´ville’s The shelters can only be built using a leaf with a large flat [64] report of E. thrax as common on cultivated plantains lamina, e.g. Musa spp. Heliconia spp., some Strelitziaceae, around Dehradun is assumed to refer to E. torus. Marantaceae, Zingiberaceae, etc. (all in the Zingiberales), Since the early stages of E. thrax and E. torus cannot and unlikely to be used on pinnate palms. It is concluded be distinguished, and they are both common pests of that all food plant records of E. torus from palms should Musa spp., it follows that where both species co-occur in be disregarded, and although the possibility of other mainland South-East Asia and the Southern Philippines, Zingiberales being used as food plants cannot be discoun- then any studies on the pest status, natural enemies, ted, none should be considered confirmed at this time. 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 Food plants of E. acroleuca J. Okolle and colleagues in Peninsular Malaysia on the population dynamics, within-field and within-plant dis- Erionota acroleuca acroleuca has been recorded using tribution of the banana skipper and its parasitoids palms as food plants in the Andaman Islands: C. nucifera, [70, 123–129]. Pinanga kuhlii, Calamus sp. [87]. Erionota acroleuca apicalis There are a few published records of palms as food from South-East Asia also feeds on palms. In Peninsular plants for E. torus. Robinson et al. [84] list Caryota Malaysia, it is recorded from ‘oil palm’ ([104] as E. thrax, and regia. As noted above, in their treatment of [76]), Caryota sp. and Orania sylvicola (=macrocladus) [106], pests of rattan palm in Peninsular Malaysia, Maziah et al. Calamus manan, C. ornatus, C. scipionum, Caryota mitis, [101] include E. torus, although Steiner [130] and Steiner Areca triandra and Adonidia merrillii [130, 133]. Recently,

http://www.cabi.org/cabreviews Matthew J. W. Cock 11 Xue and Lo [19] report rearing this subspecies from shelter making behaviour cannot be used to separate Caryota maxima (as C. ) in Yunnan Province E. thrax and E. torus. In contrast, although hardly docu- and Arenga westerhoutii in Guangxi Province, China, and mented, the leaf shelter of E. acroleuca on palm leaves may an unidentified palm in Singapore; they conclude that be constructed similarly to those of E. thrax and E. torus, E. acroleuca is primarily a palm feeder. Kitamura [109] but is also made by longitudinally rolling a leaflet or pulling reports Saribus rotundifolia (= rotundifolia) to be a together two leaflets or parts of a larger leaf. This latter food plant of E. acroleuca apex on Samar Island, Philippines. type of shelter seems typical for palm-feeding Hesperiinae L. G. Kirton (in 103, 130) has suggested that all records [7], and contrasts with the shelters of E. thrax and E. torus of Erionota spp. from rattan palms (Calamus spp.) should where the leaf is rolled transversely. 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 Life history of E. thrax records of E. thrax from C. nucifera, R. excelsa, M. sagu, A. pinnata and E. guineensis in Java [73] and the various Horsfield and Moore [78] provide the first illustrations of repetitions of these in the literature (see above) are the larva and pupa. As these observations were from Java, considered to be in error for E. acroleuca. L. G. Kirton where E. torus is not reported, they should be reliably (personal communication, 2014) transferred larvae of associated with E. thrax. The figure portrays the larva as E. acroleuca apicalis of various sizes from palms to Musa sp. white, with long erect setae on the body, and a dark and they all died. The record of E. thrax from oil palm in brown-black head. The pupa is incorrectly shown, in that New Britain, PNG [105] needs clarification by examina- it has a short, strong, pointed, slightly upturned frontal tion of voucher specimens; it might prove to be E. acro- spike. Horsfield’s emerged pupa is preserved in the NHM, leuca. In conclusion, E. acroleuca is the Erionota species that and is accurately drawn, so is here assumed to belong to is commonly found on a wide range of palms, and until some other species. Distant [79] reproduces these such time as there are authoritative records of E. thrax figures of the larva and pupa in Rhopalocera Malayana, or E. torus breeding on palms, all such existing records perpetuating this error regarding the pupa. Semper [24] should be considered misidentifications, most probably illustrates the larva, correct pupa and a large leaf shelter for E. acroleuca. 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 Food plants of other Erionota spp. 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 The Chinese species, E. grandis feeds on palms [6], as does head overgrown with very short hairs and wholly covered E. sybirita (Hewitson). The former is a gregarious species with a white waxy powder, that also entirely fills its and so unlikely to be mistaken for other Erionota spp. abode’. Although Piepers and Snellen treat E. acroleuca as Steiner [130, 133] records the later feeding on Licuala a synonym of E. thrax, it seems likely given the description kunstleri in Peninsular Malaysia. It is a rare species found of the shelter on Musa in Java that they are dealing with from Southern Myanmar and Thailand to Borneo and the immature stages of E. thrax. Their painting of the pupa Palawan [1, 18], so is unlikely to account for many of the also appears to represent this species, having no frontal records of other Erionota spp. from palms. The food plants spike, and a long proboscis extending beyond the cre- of the remaining Erionota spp. are unknown. master. In Lepidoptera Indica Vol. X, Swinhoe [80] gives the food plants as ‘wild or cultivated Musa’ and states that the larva Immature Stages ‘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 Erionota thrax, E. torus and E. acroleuca are similar in adult enormous leaves of the wild or cultivated Musa,or appearance [1, 21]. The life history and food plants of plantains, to obtain a suitable segment to be rolled up.’ E. thrax and E. torus are very similar [6, 39]. However, The larva is figured from ‘Grote’s original drawing’. while the early stages of E. acroleuca are also similar, the Arthur Grote prepared the first catalogue of the butterfly food plants and shelters are not. Hence, at this time, the collection of the Asiatic Museum in 1857–1859; he also early stages cannot be reliably separated by diagnostic took notes on the life histories, making use of a Munshi characters, and E. acroleuca can only be provisionally Zynulabdin, a local artist [134], so the material for these separated by its food plants and shelters. figures was likely to have been collected in the vicinity The larvae of E. thrax and E. torus make their leaf of Kolkata (Calcutta), and could represent either E. thrax shelters by cutting and rolling the lamina of their food or E. torus. Although not acknowledged, the figure of a plant leaves. More detailed observations of shelter for- pupa is copied from [78] and so is incorrect. Evans [1] mation are needed for both species, with information on indicates that the adult illustrated by Swinhoe [80] is individual variation, feeding habits, etc. At present, the E. thrax; however, the observations on early stages and

http://www.cabi.org/cabreviews 12 CAB Reviews 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 Figure 4 Shelters of E. thrax thrax and E. torus on Musa sources. Ashari and Eveleens [135] provide drawings of sp. 1, E. thrax, Maui, Hawaii, 13 Aug 2002 (g Forest and the life cycle and provide some additional information. Kim Starr); 2, E. thrax outbreak, River, Papua New g More recent observations are similar, but some have Guinea ( C. Dewhurst); 3, E. torus, Serdang, Malaysia, 6 been made from areas where E. thrax and E. torus occur Feb 2003, M. J. W. Cock. Differences observed between individual shelters are not considered to be diagnostic, but together and so cannot be allocated to species with to reflect individual differences. 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 from below ... Feeding occurs only during the night Fukuda [6] refer to the species that has spread to the Átime. The bottom of the nest is usually closed, and often Pacific and Asia as E. torus, rather than E. thrax. As they do filled with a large amount of frass pellets. ... the larva not refer to the diagnostic differences in the genitalia, one moves and nests are renewed several times. Moving takes cannot be sure that their identifications were thus con- place in the direction towards the basal part of the leaf. It firmed. However, although both E. torus and E. thrax occur takes a few days to make a new nest. Some larvae, in Sikkim, India, and the possibility of confusion cannot however, may show behaviour patterns exceptional to be ignored, the adults that they illustrate are correctly the habits described above.’ Leaf rolls made by the mature associated (the difference in male forewing shape is very larvae extend all the way from the edge to the leaf midrib, clear), so it is assumed that the early stages are also which leads to considerable loss of leaf area (Figure 4.2). correctly associated. Although there are photographs and The leaf rolls in Figure 4.1 appear to have five rolls of leaf, brief descriptions of the biology of E. thrax in its intro- and dangling from the bottom is a narrow strip of leaf duced range [45, 136] no detailed accounts have been margin which has not been eaten. This can be interpreted located in this review. as the larva feeding on the leaf edge within its shelter, but Eggs are laid singly or in groups, mostly on the under- not the section furthest from the midrib, which would side of the leaves, but also on the upper side. Eggs are form the distal end of the leaf roll, where a feeding larva 2.3 mm wideÂ1.1 mm high, with about 22 fine ribs stop- might be vulnerable to predators or disturbance. This may ping short of the micropyle. be the reason for the two lines in Williams’ [82] drawing Detailed descriptions of the leaf shelter building beha- mentioned above. viour are not available. The newly hatched larvae form a There are five larval instars and the final instar grows to little leaf roll from the edge of the leaf, inside which they 50 mm or more. All instars have dark brown to black feed [135]; it is not clear whether the first shelters heads and pale bodies. After the second instar there is an are simple leaf folds from the leaf edge, or the transverse increasingly dense cover of white powder on the body. leaf roll observed later – or both. Igarashi and Fukuda The body has fairly inconspicuous pale, erect setae; the [6] note that ‘the larva rolls a part of a food plant leaf head is dark brown, rounded, indent at vertex, widest inwards ...to make a nest in which to rest with its head near base, with setae similar to those on the body; the directed upwards. ... The larva eats the wall of the nest pronotum, spiracles, true legs and prolegs are all pale,

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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). more or less concolorous with the body. It can be seen pubescence, and with a black head. It is easily discovered, that there is minimal white powder on the head, thorax as it gnaws out of the gigantic banana leaves, by two and ventral part of the abdomen of the individual shown in parallel-cuts a longitudinal piece which it rolls up like a Figure 5.1, but the covering becomes more extensive as cigar, in the centre of which it lives. These characteristic the larva becomes fully grown. The pupa (Figure 6.1) is cuts you can see in the Musa-plantations from a distance pale white-brown, nearly cylindrical but deeper and wider of 20 or more paces, so that the larvae, and still more at the thorax; no frontal projection and the proboscis easily the pupae, may be collected in numbers ad lib. The sheath projects beyond the cremaster. The pupa and the pupa is of a dingy pale yellow with a very long case of the inside of the pupal shelter are lined with white waxy proboscis projecting beyond the end of the body like a powder. 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 Life history of E. torus make the shelter are not clear, but may parallel those shown by Williams [82] for E. thrax, and perhaps explained Walker [119] wrote about the butterflies of Hong Kong, by the feeding behaviour of the larvae inside the shelters and was the first to comment on the leaf-roll shelters, as extrapolated above for E. thrax based on Figure 4.1, i.e. stating that E. thrax (i.e. E. torus) has been reared ‘in 1892 the cut nearer the midrib is used to make the shelter, but from larvae found in rolled-up leaves of banana’. Mack- the second ‘cut’ is actually where the larva stops feeding innon and de Nice´ville [64] include a brief description of within the shelter in order not to expose itself at the the larva in northern India, and mention that pupation is in bottom. a portion of a rolled-up leaf. In the Palaearctic butterflies Based on his observations at Hong Kong and Canton section of the Macrolepidoptera of the World, Seitz [137] (Guangdong, China), Hoffmann [121] provided a quite wrote ‘The larvae of Erionota ... feed on the gigantic detailed description of the early stages and their biology, leaves of Musa, which are yards long. In forming their together with drawings of the ovum, larva and pupa, and shelter they commence by making two incisions from the photographs of adults and shelters. Eggs are laid singly or edge of the leaf towards the midrib at a distance of about in irregular groups on the upper and lower surfaces of 8–10 cm one from the other, and then proceed to roll this leaves (more frequently the latter). They measured 2 mm piece of the leaf up like a cigar, the larva living in the in diameter and 1.13 mm in height, with the flat area on cylindrical hollow within this roll.’ When dealing with the top 0.45 mm wide; there are 24–29 fine ribs. They are Indo-Australian butterflies, Seitz [94] expands on this ‘The usually orange in colour, but there is considerable varia- larva is snow-white, covered with a very short wax-like tion over time and within batches. There are five larval

http://www.cabi.org/cabreviews 14 CAB Reviews instars. The first instar is 3 mm long when recently hat- from above, or one with the leaf under surface on the ched, pale green with the head and pronotum black, and inside of the shelter and the other with the leaf under no wax powder. The width of the head capsules of the five surface on the outside of the shelter. In any case, they instars were 1.08, 1.45, 2.08, 2.9 and 4.26 mm. After the seem to be incorrect, as both species roll the leaf flap first moult, the larva is covered with an increasingly clockwise or anticlockwise depending on which side of dense layer of white powder. The fifth instar measures the leaf the shelter is formed, and the shelters normally about 44 mm long; the surface of the blackish-brown head have the leaf underside inside the shelter for both species. is ‘more or less granulose’, with a moderate covering of Igarashi and Fukuda [6] also indicate that the top end of weak, silky setae, thicker on the front. The pupa measures the larval leaf shelter is open and that the larva comes out 40–46 mm, and is about 7 mm wide and deep at the of this opening to feed on the leaf by night; this also seems thorax and is covered with white powder. It is pale dirty to be incorrect as photographs of the leaf shelters indicate yellow apart from the dark brown cremaster; the pro- no feeding outside the shelters [9, 66, 121, 138], and the boscis extends 5 mm beyond the cremaster; spiracles indications are that the larvae feed inside the tube, as brown; the eyes become reddish about halfway through discussed below. Working in Japan, Makibayashi [139] the development period. provides a detailed description with clear diagrams to After the early reports considered above, there show how a final instar caterpillar makes a new leaf seems to be little in the way of original observations, as shelter. His diagrams show that the leaf roll is pulled into opposed to repeating information from the literature, position by silk strands between the leaf roll and the until late last century, when accounts illustrated with adjacent unaffected part of the leaf, which is in due course, photographs started to appear. There is a brief summary is pulled into the shelter in turn. from Mauritius in Monty [59] as E. thrax. Bascombe et al. The following notes on E. torus in Mauritius were pre- [9] describe and illustrate the early stages of from Hong pared by T. C. E. Congdon (personal communication, Kong, and Igarashi and Fukuda [6] include the life history 2014). The larva makes a cut in the leaf, and rolls the cut from Japan. Additional information is provided from the portion, extending the cut across the leaf as it grows. No observations of T. C. E. Congdon (personal communica- evidence of eating can be seen outside the shelter, and tion, 2014) and the author’s unpublished observations. when the tube is unrolled, the unrolled leaf does not refill The following is largely based on Bascombe et al.’s [9] the space it originally occupied. It appears that the larva account, except as indicated. The egg ([9], Plate 103.1–2) feeds by eating the inner layers of the tube, thus avoiding is 1.8–2.2 mm in diameter, dome-shaped with 22–24 fine the dangers inherent in emerging from its shelter. One ribs stopping short of the micropyle; they are pink or effect of this is that the inside diameter of the tube variegated pink and yellowish-white when laid, becoming becomes much greater than is usual in Hesperiinae tube paler as they develop. They are usually laid on the leaf shelters. The bottom end of the shelter is loosely sealed underside, and may be laid singly, but more often in with silk, retaining the frass, thus hindering access by clusters of up to 30, rarely 50. In contrast, Igarashi and predators. The top of the shelter is crudely closed with Fukuda [6] state that the ova are 2.2 mm in diameter, and silk, but must be opened when the larva extends the cut, normally laid singly on the leaf upper side, although bat- and rolls more of the leaf. Old strands of silk are cut and ches of 3–36 are recorded; this may reflect a gen- remain visible along the cut on the section not incorpo- eralization on a relatively small number of observations. rated into the shelter. This detail is at variance with the On hatching the larva eats the egg shell, and then moves description and diagrams of Makibayashi [139] above, and to the edge of the leaf to make its first shelter, which is the author’s unpublished observations from western either a simple fold parallel and close to the edge of the Malaysia that the silk strands used to form the shelter leaf ([9], Plate 103.3), or a cone shaped roll formed by are attached to parts of the leaf that are subsequently making a curved cut from the edge of the leaf and rolling incorporated into the shelter. Further observations are the resultant flap under. This first cut may be curved needed to clarify whether these are local or individual towards the leaf base or away from it, but all subsequent variation in behaviour. The larva pupates within the shelters are made with the cut curved towards the base of shelter, head up and without a girdle. the leaf. Larvae feed on the inner parts of the roll, which is From an early age, the larva is covered with an closed at the inner end (nearest midrib) and open at increasingly thick layer of white powder, so that the body the distal end, where frass may accumulate. As the larva appears white with erect pale setae partially covered with grows, it extends the cut and rolls more of the leaf into the powder, and the uniformly dark head is partially the shelter. Additional shelters are constructed if the obscured with the white powder (Figure 5.2). The larva shelter is damaged, unable to roll further or encounters grows to 55 mm in length [6] and is covered with white another shelter. flocculence as it prepares for pupation. Igarashi and Fukuda [6] indicate that the leaf shelters of Pupation takes place in the final leaf roll shelter which is E. torus and E. thrax are rolled in opposite directions, 15–20 mm long, lined with a thin layer of silk and closed although it is not clear from the text and diagrams whe- at the distal end by a thin mesh, which is easily broken by ther this means clockwise and anticlockwise when seen the emerging adult. Ito and Nakamori [140] refer to the

http://www.cabi.org/cabreviews Matthew J. W. Cock 15 pupal shelters in Japan as ‘trumpet shaped’, 1.5–2.0 cm earlier observations. To make the larval leaf shelter, larvae diameter at the smaller [basal] opening, and 3.0–3.5 cm at cut the leaflets near the base about two-thirds of the the terminal opening. This does not seem to be typical. distance to the midrib, and wrap the distal part in several Igarashi and Fukuda [6] note that the pupal shelter is turns to form a roll (Steiner refers to this as a longitudinal simpler than the larval shelter, having 3–4 layers rather roll, but I would interpret this term as parallel to the mid- than 5–6, and the pupa is formed with the head upwards, rib, which is not the case here). The larva rests and feeds although the adult butterfly emerges from the bottom of inside the leaf roll. Prior to pupation, the larva creates a the shelter. similar kind of leaf roll, which is heavily lined with wax. The pupa (Figure 6.2) is 45 mm long, pale brown with a L. G. Kirton (personal communication, 2014) confirms variable and patchy covering of white powder. The pro- these observations, and has also observed that on Caryota boscis extends almost to the cremaster. It is attached at sp(p)., which have laminate fishtail pinnae, the larva makes the cremaster, and unlike many Hesperiinae does not a shelter in a similar manner to those of E. thrax and construct a silk girdle. 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 Life history of E. acroleuca some additional information regarding E. acroleuca apicalis. Larvae, hatching from egg clusters develop in separate Wood [104] treats E. acroleuca apicalis as E. thrax pinnae on the same rachis, rolling the pinnae into a ‘cone- and includes a photograph of a hesperiid larva and shape shelter’. They illustrate in colour a larva similar to shelter (plate XIVa) without explicitly stating whether it those above, with a dark brown head with short setae, represents E. thrax or Cephrenes chrysozona (Plo¨tz), the conspicuous long setae on the body, and the head and two named species of Hesperiidae treated. The angle of body covered with white waxy powder. view does not permit a definitive identification to be Maziah et al. [101] record that pupation is in the final made, but it is more likely to be E. acroleuca than larval shelter and ‘the pupa is long, slender and also C. chrysozona. The colouring of the larva suggests an covered with the same whitish powdery substance pre- Erionota sp., rather than a Cephrenes sp., which has a green sent on the larva. When disturbed the pupa wriggles body and white head with dark lines [9, 39]. The leaf violently in the shelter’. Xue and Lo [19] indicate that the shelter appears to have been formed by making a diagonal pupa is 33 mm, pale cream in colour with brown spiracles, fold across the oil palm leaflet, but since this may have the proboscis extending beyond the cremaster. It is been constructed in captivity, it is not necessarily indica- ‘sealed in a cocoon’ in a shelter formed by ‘rolling the tive of what happens in the field. Mature C. chrysozona undersurface of the apex half of the rachis [pinna?] lat- larvae roll palm leaflets parallel to the mid rib (author’s erally’. This description of the shelter is not entirely clear, unpublished observations). and although their illustrations of the shelter and pupa Steiner [133] records observations on the ova of show a silk cocoon within a leaf shelter, the structure of E. acroleuca apicalis (as E. hiraca). The ova are white, with the shelter cannot be interpreted, although it appears to a smooth surface and hemispherical in shape. They can be aligned longitidinally with the pinna veins. be found on all leaves of C. manan, but the majority are Finally, Kitamura ([109], Figure 18, upper photo) found on the youngest leaf, and although found on both includes a photograph of the final instar of E. acroleuca surfaces of the leaf, they are mostly found on the upper apex in his report on this butterfly feeding on L. rotundifolia surface. Xue and Lo [19] indicate that the ova of on Samar Island, Philippines. The larva is similar to those E. acroleuca apicalis are laid in a cluster on the frond reported above (and to other Erionota spp.), but the leaf underside. shelters are not documented. Maziah et al. [101] treat E. acroleuca apicalis as E. torus The shelter building behaviour of the larvae seems in their illustrated account of the pests of rattan, and to vary with the leaf structure of the food plant palm. illustrate the larva with a photograph. They note that the Further observations from different food plants and from larva ‘rolls up the leaf from the tip parallel to the veins and different parts of the range of this species and its sub- lives and feeds from one end of the leaf in this shelter’. species, supported with voucher specimens, are needed This wording is not completely clear, but in view of sub- to clarify the biology of this species. 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 Life history of E. sybirita midrib), made by most palm-feeding Hesperiinae [7]. They describe the larva as ‘pale green covered with short silky There are also some details on the life history of E. sybirita hairs and a whitish powdery substance and has a dark on L. kunstleri in Steiner’s [133] thesis. The eggs are white brown head capsule’. Their photograph shows a larva to grey, hemispherical with a smooth surface; almost similar to those of E. thrax and E. torus. Steiner’s [133] all were found on the underside of the leaflets. The description and photograph of a 34 mm larva match larvae have a whitish green body, but unlike the other

http://www.cabi.org/cabreviews 16 CAB Reviews Erionota spp. treated above, no wax layer. The head [150] point out that O. pallidipes is the senior synonym. capsule is dark brown to black. The pupae are brownish, Ashmead [173] described O. pallidipes from the Philippines covered with wax, and about 36 mm long. The very long from ‘two specimens bred by Father W. A. Stanton from proboscis sheath extends beyond the abdomen by up to an undetermined aphid’. This now seems to be a host 40 mm. The pupae are attached only by the cremaster and error, perhaps because a mummified aphid may be similar do not have a silk girdle. Like the larvae of E. hiraca, the in size, shape and colour to a parasitized ovum of E. thrax. larvae of E. sybirita cut the leaflets near the base and wrap Ooencyrtus pallidipes is a common and widespread egg the distal part into a roll with several turns (again referred parasitoid of E. thrax (Table 2) that also parasitizes ova of to as longitudinal, but this term should be avoided for the E. torus (Table 4). It is reported parasitizing E. thrax (or leaf rolls of Erionota spp.). The larva stays in the roll and E. torus) from Nepal, Thailand, Malaysia, Sumatra, Java, eats it from the inside. Prior to pupation the lower end of PNG and Guam, although the last two may represent the tube is sealed with silk. It appears that the absence of a accidental introductions. It has been introduced as a covering of white waxy powder on the larva (and perhaps biological control agent in Mauritius and Hawaii for the also on the pupa) as well as the specific host plant should control of E. thrax (although the pest in Mauritius is now facilitate separation of the early stages from E. acroleuca. 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]. Natural Enemies of E. thrax and E. torus Females avoid parasitized ova for oviposition [127]. In Sumatra, it is reported to cause 10–27% mortality Waterhouse and Norris [45] summarized the available [148, 151] and Hasyim et al. [147] reared an average of information on the natural enemies of E. thrax, many only 3.8 adults per parasitized egg. It has been recorded from identified to genus or family, but as has become apparent several Lepidoptera, including Eupterotidae, Gracillariidae, above this summary would not have separated natural Nymphalidae, Papilionidae and Pieridae [141], but the ova enemies of E. thrax from those of E. torus where they of Gracillariidae are most probably too small to support occur together. This table is updated as three separate this species. A record from ova of a buprestid represents tables below, broken up into records that can be un- a misidentification [150]. 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 Pediobius erionotae Kerrich (Eulophidae) records in Gold et al. [171], Tinzaara and Gold [172], and Muniappan et al. [115] are not repeated here. Key This species was described from specimens reared from information on what appears to be the most common ova of E. thrax (or E. torus) in peninsular Malaysia, and the and important parasitoids (all ) is sum- type series includes material from Java, and R. A. Syed’s marized in the following paragraphs. material from Sabah, Malaysia reared from ova, larvae and cocoons of Cotesia erionotae (Wilkinson) [149, 155]. Brachymeria spp. (Chalcididae) Kerrich [155] does not seem entirely convinced by the records from E. thrax larvae and even less so by the Several Brachymeria spp. that attack pupae of Erionota spp. record from a C. erionotae cocoon. This is another com- appear to be polyphagous species, including B. lasus mon egg parasitoid of E. thrax (or E. torus), which has (Walker), B. albotibialis (Ashmead), B. euploeae (West- also been recorded Sumatra, where it causes 16–39% egg wood) and B. thracis (Crawford). Okolle et al. [127] found mortality [148, 151]. 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. Cotesia erionotae (Wilkinson) (Braconidae) [127] report seven per host for B. albotibialis. The incon- sistency of names used in different publications, even from This gregarious larval parasitoid was described from the same area, suggests that at least some material has E. thrax (or E. torus) in West Malaysia [157], and is known been misidentified in the past. 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 Ooencyrtus pallidipes (Ashmead) () Mauritius (Table 4). It was originally described in the genus Apanteles, but Austin and Dangerfield [174] trans- This species was better known as O. erionotae Ferrie`re, ferred it to Cotesia. Larvae of instars 2–4 are parasitized which was described from Peninsular Malaysia from ‘eggs and the mature parasitoid larvae emerge mainly from of E. thrax L. and of an unknown Lepidopteron on wild instar 5 [127]. Hasyim et al. [147, 148] record an average plantain leaf’ [169]. Given the locality, this could have of 58 adults per parasitization, although cocoons are often been from either E. thrax or E. torus. Huang and Noyes hyperparasitized, while Okolle et al. [127] record an

http://www.cabi.org/cabreviews Matthew J. W. Cock 17 average of 87 per larva. This species has been released and egg parasitoids Ooencyrtus pallidipes and Agiommatus and established for biological control of E. torus in Maur- sp. nr. sumatraensis to Mauritius in 1971–1973. All four itius and Taiwan and E. thrax in Hawaii, Guam, Saipan and species were released in Mauritius. Cotesia erionotae PNG. It was initially thought to be already present in and O. pallidipes became established, and although there Taiwan, attacking folus (Cramer) (Hesperiidae, were recoveries of Agiommatus sp. it did not persist. The Hesperiinae, incertae sedis) which feeds on parasitoids quickly provided effective biological control. (Zingiberaceae), but closer examination showed differ- In 1975 a cyclone severely damaged banana plants and ences in microsculpture and colouring [175], suggesting drastically reduced banana skipper and parasitoid popu- this is a different species. In light of this, a recent record lations which, nevertheless, built up again in 1976. None of C. erionotae parasitizing U. folus in India [176, 177] may of the parasites established earlier was recovered in the need verification. 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 sp. () specimens of E. torus (as E. thrax) were taken on the west coast of Mauritius in 1979, but it was not seen elsewhere This is a common larval parasitoid of E. thrax in Sumatra between the years 1976 and 1980, so that its status and Java. In Java, 10–80% larval parasitism is reported, and was considered unclear. Williams [189] considered it (as it is commonly hyperparasitized by Brachymeria lasus, and E. thrax) ‘uncommon for reasons that are not clear’. rarely by Eurytoma sp. B and Theronia zebra [147]. It is Collectors from the African Butterfly Research Institute possible that the records of a Scenocharops sp. a common found early stages in 2014 (T. C. E. Congdon, personal larval parasitoid of E. thrax in Sabah [149] represent the communication, 2014), so E. torus continues as an same species as the two genera are similar and closely uncommon species of no significant pest status in related in the tribe Campoplegini. Mauritius, almost certainly kept under effective biological control by its introduced parasitoids. None of the other species of Hesperiidae found on Mauritius are known Xanthopimpla gampsura Krieger (Ichneumonidae) to be parasitized by any of the parasitoid species intro- duced against E. torus (J. Monty, personal communication, This is a common pupal parasitoid of E. thrax in Sumatra 1987 in 45). and Java. It was described from Borneo and Sumatra Based on the programme already carried out [178], and has also been recorded as a parasitoid of the for Mauritius (purportedly on E. thrax but actually on palm-feeding hesperiids H. irava and Cephrenes chrysozona. E. torus) when E. thrax appeared in Hawaii, a biological Earlier records from Java of Xanthopimpla sp. are referable control programme was immediately started [45]. In to this species, but it is not clear whether the record 1973, O. pallidipes was introduced from Guam where of X. regina from Sabah represents a different species or a it seemed to be an accidental introduction itself, and in misidentification. 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. Biological Control of E. thrax and E. torus Tables 3 and 4), the population of C. erionotae from Sabah would have been collected from E. thrax only, since There have been several programmes of biological control E. torus does not occur there, whereas that from Thailand against E. thrax (Guam, Saipan, Hawaii, PNG) and E. torus is likely to have been reared from both E. thrax and (Mauritius, Taiwan). Waterhouse and Norris [45] provide E. torus. Control in Hawaii was rapid, and the strain of a review of the earlier programmes against E. torus C. erionotae from Thailand was probably already estab- and E. thrax (summarized below and in Table 5). There lished and bringing the pest under control before that are numerous natural enemies in the indigenous range of from Sabah was released. In 1974, C. erionotae cultures of E. thrax (previous section, Tables 2–4), and several have the Thailand strain were sent from Hawaii to Guam, and been used as biological control agents (Table 5). from Guam to Saipan in 1974–75, in both cases giving Erionota torus was first reported in Mauritius in 1970 as good control [45, 49]. Both parasitoids were also sent E. thrax [59], although it was suspected to have been from Hawaii to Taiwan for the control of E. torus; they introduced with military air traffic from Malaysia in 1968 were released in 1987 and both became established [45]. Once established in Mauritius, it soon became a [182]. The impact has not been fully reported, although serious pest on tall banana varieties in urban areas, but Teruya [138] found E. torus to have ‘a low pest status’, not in rural commercial plantations of dwarf Cavendish- infestation ranging from 0.1 to 7% with an average of 4%, type bananas [45, 59]. R. A. Syed of the Commonwealth but he did not find the introduced parasitoids. Institute of Biological Control (part of CABI) studied the Erionota thrax may have spread into Irian Jaya before parasitoids of E. thrax in Sabah [188] and sent shipments 1983 as it appeared by the border in north-western of larval parasitoids Scenocharops sp. and Cotesia erionotae, mainland PNG in 1983, and then spread rapidly through

http://www.cabi.org/cabreviews Table 2 Parasitoids and diseases of Erionota thrax updated from Waterhouse and Norris [45]. Taxonomy adjusted from Noyes [141] and Yu [142]. Records of egg Reviews CAB 18 parasitoids 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 Philippines Pupa L. B. Uichanco Recorded from several families of Lepidoptera (Ashmead) editorial footnote in [144] Chalcidoidea, Chalcididae Brachymeria euploeae Indonesia, Pupa [15, 135, 145] Widespread in South-east Asia; highly polyphagous on Lepidoptera (Westwood) unspecified Chalcidoidea, Chalcididae B. euploeae (Westwood) Indonesia, Pupa [135] Common; 40–60% pupal parasitism in combination with Xanthopimpla west Java sp. This is likely to represent a mixture of B. lasus and B. thracis, judging from 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 of Casinaria sp. Chalcidoidea, Chalcididae B. lasus (Walker) USA, Hawaii Pupa [136] A widespread and polyphagous species (=B. obscurata (Walker)) Chalcidoidea, Chalcididae Brachymeria sp. Papua New Guinea, Pupa [58] < 5% pupal parasitism mainland Chalcidoidea, Chalcididae Brachymeria sp. nr Malaysia (Sabah) Pupa [149] This is not a valid species; perhaps an error for B. marginata http://www.cabi.org/cabreviews marginicollis (i.e. B. lasus) Chalcidoidea, Chalcididae Brachymeria sp. nr. Malaysia (Sabah) Pupa [149] This is likely to be B. lasus, of which obscurata is a synonym. obscurata (Walker) Brachymeria spp. cause 16% pupal parasitism Chalcidoidea, Chalcididae Brachymeria thracis Indonesia, Java Pupa [146] Close to Brachymeria euploeae; earlier records of that species may well (Crawford) refer to this Chalcidoidea, Chalcididae B. thracis (Crawford) Indonesia, Sumatra Pupa [147, 148] Gregarious, nine adults per host; 9% parasitism Chalcidoidea, 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) Indonesia, Sumatra Ovum [147, 148, 151] Gregarious, 2.8 adults per egg. Common and widespread; together (= Ooencyrtus erionotae Ferrie` re) with Pediobius erionotae 10–40% egg parasitism Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Indonesia, Ovum [15, 145] (= Ooencyrtus erionotae Ferrie` re) unspecified Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Malaysia (Sabah) Ovum [149] 4% egg parasitism (= Ooencyrtus erionotae Ferrie` re) Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Papua New Guinea, Ovum [58, 150] 26–33% egg parasitism (= Ooencyrtus erionotae Ferrie` re) mainland Chalcidoidea, Encyrtidae Ooencyrtus sp. (not erionotae Papua New Guinea, Ovum [58] Ferrie` re) mainland Chalcidoidea, Eulophidae Elasmus brevicornis Gahan Indonesia, Java Larva [97, 152, 153] Polyphagous, facultative hyperparasitoid Chalcidoidea, Eulophidae Elasmus philippinensis Ashmead Philippines Larva [153, 154] Chalcidoidea, Eulophidae Elasmus sp. Indonesia, Sumatra L2 [147] Gregarious, 12 adults per host. Only one record Chalcidoidea, 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; Ovum [155] Original description; holotype peninsular Malaysia Indonesia, Java Chalcidoidea, Eulophidae P. erionotae Kerrich Malaysia, Sabah Ovum [149] 30% egg parasitism. Syed’s material is included in the type series (= Pediobius sp.) Chalcidoidea, Eulophidae P. erionotae Kerrich Malaysia, Sabah Larva [149] Syed’s material is included in the type series, although Kerrich [155] (= Pediobius sp.) would seem to have reservations that this species attacks ova and larvae 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 Chalcidoidea, Eupelmidae Anastatus sp. Indonesia, unspecified Ovum [145] Chalcidoidea, Eupelmidae Anastatus sp. Papua New Guinea, Ovum [58] mainland Chalcidoidea, Eupelmidae Anastatus sp. USA, Hawaii Ovum [136] Chalcidoidea, Pteromalidae Agiommatus sp. Indonesia, unspecified Ovum [15, 145] Chalcidoidea, Pteromalidae Agiommatus sp. nr Malaysia (Sabah) Ovum [149] 2% egg parasitism sumatraensis Crawford Chalcidoidea, Pteromalidae Agiommatus sumatraensis Crawford Indonesia, Java Ovum [146] Chalcidoidea, Pteromalidae A. sumatraensis Crawford Indonesia, Sumatra Ovum [147, 151] Localized, causing 0–9% mortality Chalcidoidea, Trichogrammatidae Trichogramma sp. Guam Ovum [156] Chalcidoidea, Trichogrammatidae Trichogramma sp. USA, Hawaii Ovum [136] Ichneumonoidea, Braconidae Cotesia erionotae Indonesia, Java L4–L5 [146] (Wilkinson) 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 by Pediobius elasmi (Ashmead) Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Indonesia, unspecified Larva [15, 135, 145] (=Apanteles erionotae) Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Indonesia, west Java Larva [135] Up to 50% larval parasitism (=Apanteles erionotae) Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Malaysia (Sabah) Larva [149] 25% larval parasitism; heavily hyperparasitized by Eurytoma sp., (=Apanteles erionotae) which in turn is hyperparasitized by Pediobius sp. and P. elasmi http://www.cabi.org/cabreviews (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 attacks ova, larvae and parasitoids of E. thrax Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Indonesia, Sumatra Larva [157] (=Apanteles erionotae). 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 and Theronia zebra Ichneumonoidea, Ichneumonidae sp. Indonesia, Java L3–L4 [146] Ichneumonoidea, Ichneumonidae Charops sp. Indonesia, Sumatra L3 and L4 [147] Ichneumonoidea, Ichneumonidae Echthromorpha agrestoria USA, Hawaii Larva [136] (= E. fuscator) Ichneumonoidea, Ichneumonidae Scenocharops sp. Malaysia (Sabah) Larva [149] 17% larval parasitism Ichneumonoidea, Ichneumonidae Theronia zebra-zebra Vollenhoven Indonesia, Java Pupa [146] Polyphagous on pupae of several families of Lepidoptera Ichneumonoidea, 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 Hesperiidae Ichneumonoidea, Ichneumonidae X. gampsura Krieger Indonesia, Sumatra Pupa [147, 148] Common and widespread Ichneumonoidea, Ichneumonidae Xanthopimpla regina Malaysia (Sabah) Pupa [149] 7% pupal parasitism (=Xanthopimpla sp.) Ichneumonoidea, Ichneumonidae Xanthopimpla sp. Indonesia, unspecified Pupa [15, 145] Most probably this represents Xanthopimpla gampsura Krieger from Java Ichneumonoidea, Ichneumonidae Xanthopimpla sp. Indonesia, west Java Pupa [135] Common; 40–60% pupal parasitism in combination with Brachymeria euploeae. Almost certainly this represents Xanthopimpla gampsura 19 Cock W. J. Matthew Krieger Ichneumonoidea, Ichneumonidae Xanthopimpla sp. ‘Malay Archipelago’ Not stated [97] Probably this represents Xanthopimpla gampsura Krieger from Java Platygastroidea, Scelionidae Telenomus sp. nr. attaci Malaysia (Sabah) Ovum [149] Diptera, Phoridae Unidentified Indonesia, Sumatra Pupa [147] Uncommon Diptera, Sarcophagidae Unidentified Indonesia, Sumatra L5 [147] Uncommon Diptera, Sarcophagidae Unidentified Malaysia (Sabah) Pupa [149] 5% pupal (or larval–pupal) parasitism Diptera, Unidentified Indonesia, unspecified Pupa [135] Diptera, Tachinidae Unidentified Malaysia (Sabah) Larva [149] Diptera, Tachinidae Blepharipa sp. Indonesia, Sumatra L5 [147] Uncommon Diptera, Tachinidae Palexorista solensis (Walker) Indonesia, Sumatra L3–L5 [147] Uncommon Fungus Cordyceps sp. Malaysia (Sabah) Larva [149] 20 CAB Reviews the island over five years, and started to colonize the other islands of PNG [58]. It spread together with its occurs occurs 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 E. torus E. torus [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 anticipated only anticipated only 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 Mauritius Larva [45, 159] Introduced to an area where it is MauritiusChina, Fujian Ovum Ovum [45, 159] [150, Introduced 162] to an area where it is 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 re) ` production in Australia. Assuming similar levels of damage

Subba Rao China, Hong Kong Ovum [9, 158] Solitary species in Australia as in PNG, these benefits were estimated to Ferrie

group China, Hong Kong Pupabe [9] A$223 million to the year 2020. The assessment Ashmead Taiwan Larva [131, 160] Ectoparasitoid ). (Ashmead) of benefits did not consider the social and cultural uses

(Wilkinson) of banana leaves, nor the possible benefits in Irian Jaya, to regina (Townsend) Japan, Okinawa Pupa [161] which the parasitoid almost certainly spread. . Taxonomy adjusted from Noyes [141] and Yu [142] sp. Bauer et al. [190] considered the impact of the

sp. Japan, Okinawa LarvaPNG [161] biological control programme on poverty. Poverty through income deprivation is a significant problem in

Ooencyrtus erionotae PNG. Estimates from a 1996 household income survey Apanteles erionotae = = ( (

Erionota torus suggested that around 30% of the PNG population have Cotesia erionotae Xanthopimpla Anastatus Agiommatus erionotae Elasmus philippinensis Exorista japonica Ooencyrtus pallidipes Leurocerus hongkongensis 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 Parasitoids and diseases of 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- Chalcidoidea, TrichogrammatidaeChalcidoidea, unspecifiedIchneumonoidea, Unidentified Braconidae Ichneumonoidea, Ichneumonidae Unidentified Japan, Okinawa Ovum China, Hong Kong Ovum [161] [121] Chalcidoidea, Pteromalidae Chalcidoidea, Eupelmidae Chalcidoidea, Eulophidae Unspecified disease Japan, Okinawa Larva, pupa [161] Diptera, TachinidaeDiptera, Tachinidae Unidentified Taiwan Pupa [138] Chalcidoidea, Encyrtidae Table 3 FamilyChalcidoidea, Encyrtidae Speciesject has also Location contributed to lower urban Reference banana Comment prices,

http://www.cabi.org/cabreviews Table 4 Parasitoids and diseases of Erionota thrax or torus updated from Waterhouse and Norris [45]. Taxonomy adjusted from references given, Noyes [141] and Yu [142].

Family Species Location Reference Comment Chalcidoidea, Chalcididae Brachymeria albotibialis India Pupa [163] (Ashmead) 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 parasitoid Chalcidoidea, Chalcididae Brachymeria euploeae (Westwood) Thailand Pupa [45, 164–166]1 Chalcidoidea, Chalcididae B. euploeae (Westwood) India Pupa [163] Chalcidoidea, Chalcididae Brachymeria lasus (Walker) Malaysia, peninsular Malaysia Pupa [167, 168] Susainathan is incorrect to refer to this (= Chalcis marginata (Cameron)) pupal parasitoid as a larval parasitoid Chalcidoidea, Chalcididae B. lasus (Walker) (=Brachymeria Thailand Pupa [45, 164–166]1 marginata (Cameron)) Chalcidoidea, Chalcididae Brachymeria thracis (Crawford) India Pupa [163] Chalcidoidea, Encyrtidae Ooencyrtus pallidipes (Ashmead) Malaysia, peninsular Malaysia Ovum [125–128, 169] This is the original description from (= Ooencyrtus erionotae Ferrie` re) ova of E. thrax and an unidentified lepidopteran on wild plantain. The commonest egg parasitoid (51%) http://www.cabi.org/cabreviews Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Nepal Ovum [150] Chalcidoidea, Encyrtidae O. pallidipes (Ashmead) Thailand Ovum [45, 164–166]1 (= Ooencyrtus erionotae Ferrie` re) Chalcidoidea, Eulophidae Elasmus philippinensis Ashmead ‘Malay Archipelago’ [Malaysia Not stated [97] Parasitic on at least three families of and/or Philippines] Lepidoptera Chalcidoidea, Eulophidae Elasmus sp. Malaysia, peninsular Malaysia Larva [125] < 1% parasitism Chalcidoidea, Eupelmidae Anastatus sp. Thailand Ovum [166] Chalcidoidea, Eulophidae Pediobius erionotae Kerrich Malaysia, peninsular Malaysia Ovum [155] Original description; holotype peninsular Malaysia Ichneumonoidea, Braconidae Cotesia erionotae (Wilkinson) Malaysia, peninsular Malaysia Larva [125–128, The commonest larval parasitoid (=Apanteles erionotae). 157, 168] Ichneumonoidea, Braconidae C. erionotae (Wilkinson) Malaysia, peninsular Malaysia Larva [170] This record of ‘Apanteles sp.?’ (=Apanteles sp.?) is assumed to be C. erionotae, which was not described until 1928 Ichneumonoidea, Braconidae Cotesia erionotae Thailand Larva [45, 164–166]1 (Wilkinson)(=Apanteles erionotae) 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 [125] < 1% larval parasitism, 1% pupal ate .W ok21 Cock W. J. Matthew (L3, L4), pupa parasitism 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]. 2CBReviews CAB 22

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. 1971–1972 Malaysia (Sabah) Recoveries, but not [149, 179], J. Monty, personal nr. sumatraensis established communication, 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, http://www.cabi.org/cabreviews 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 to [136, 180, 184] distinguish from Thailand strain already becoming established)1 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 C. erionotae 1990 Thailand via Hawaii Established [48] Guinea (mainland) and Guam 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 cruegeri (Butler) (), 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 been reported to parasitize E. thrax in South-East Asia (Table 4). 4Reference [186] not examined, but based on citation in Waterhouse and Norris [45]. Matthew J. W. Cock 23 in that the fall in supply of bananas caused by the pest Research Gaps would have forced banana prices up. Banana prices are likely to have risen by about 15% had the biological In light of the foregoing, the following research questions control of the skipper not been achieved. Preventing this and gaps are identified: increase in market prices for bananas of about 15% by the introduction of the biological control measures resulted  Molecular methods, particularly barcoding, are needed in about 28 000 purchasers of bananas living just above to confirm identifications of early stages and develop the poverty line when they otherwise would have been identification tools. below it. The biological control program increased  Further collections and reliable identification of effective incomes across all income levels of banana pro- adults (by examination of the genitalia and/or barcod- ducers and banana consumers. 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. Discussion  Material from the Snellen collection should be ex- amined to see if some of their food plant records can The available information on the classification, taxonomy, be aligned with modern taxonomy of Erionota spp.; in food plants, natural enemies and biological control of two particular should their records of E. thrax from palms species of Erionota that feed on Musa spp. and one that be associated with E. acroleuca, which they considered a feeds on palms have been presented. It is concluded that synonym of E. thrax. E. thrax and E. torus feed as larvae on Musa spp., and to a  The life history and food plants of the other three limited extent some related species of Zingiberales, but subspecies of E. thrax should be established, which may records of these species from palms and other plant help to clarify their taxonomic status. groups are in error. The records from palms are now  As yet, there is no reliable way to separate the early attributed to E. acroleuca. This distinction is important in stages of E. thrax, E. torus and E. acroleuca morpholo- order to understand the ecology of these crop-feeding gically. Careful study and documentation of reared butterflies so that management strategies are appropriate. material, supported by reared vouchers and barcoding Where E. thrax and E. torus occur together, economic of individuals, is needed to see if diagnostic criteria can entomologists have hitherto treated them as a single be found. species. There is no published information to show  Careful observation and behavioural studies of how whether the occasional outbreaks of banana skipper in the larvae construct their shelters on different food this region where both species are indigenous are due plants may provide additional diagnostic characters. to one or both of these species. Similarly, it is not  Once the larvae of E. thrax and E. torus can be dis- known whether the associated parasitoids attack both tinguished, the host-parasitoid relations for the genus species equally or show preference for one or the other. can be clarified; do parasitoids accept both equally or is Nevertheless, biological control has been successful there specialization for one or the other? without knowing about the preferences of parasitoids for  The identity of E. thrax rather than E. torus in the the two species, or whether the material used was col- Andaman Islands may need confirmation. It does not lected from one or both Erionota spp. in Thailand. As seem to be a significant pest in the Andaman Islands. If E. torus in Mauritius was misidentified as E. thrax,it this is correct, is it because of indigenous or ac- was controlled using parasitoids of E. thrax from Sabah. cidentally introduced parasitoids, perhaps also attacking There are plenty of examples where a lack of under- E. acroleuca? standing of the taxonomy of the target pest of a biological  Recognizing the possibility that E. torus may have spread control programme has led to a mis-match of parasitoids into areas where only E. thrax thrax was known, it and target (see examples in [191]), and the author firmly would be worth making fresh collections to assess the supports the view that good taxonomy is critical to suc- situation, particularly in Borneo, Sumatra and Java. cess in many biological control programmes, but in the  It is considered that E. torus is most probably the case of Erionota spp. this does not seem to have been a only Erionota sp. in Mauritius, but further collection critical factor. should be made to confirm this. Erionota torus is present For the foreseeable future, it would be a sensible pre- in Mauritius at low densities, and seems to cause caution to ensure that voucher specimens are preserved no economic damage. It seems most likely that this and documented for all observations on Erionota spp. All is due to the continuing action of the introduced three species should be considered quarantine risks for parasitoids, but this should be investigated. The tropical areas where they do not naturally occur or have reasons for the low population density of E. torus in not already spread, and an improved understanding of Mauritius have implications for the quarantine risk their ecology should improve pest risk assessments that it presents to La Re´union and other Afrotropical relating to their spread in future. areas.

http://www.cabi.org/cabreviews 24 CAB Reviews  The new report of E. torus devastating bananas in the Department for International Development, the Swiss Western Ghats of India suggests that biological control Agency for Development and Cooperation and others. 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 References which would be most at risk. 1. Evans WH. A Catalogue of the Hesperiidae from Europe,  Further collections are required in New Britain, PNG, Asia and Australia in the British Museum (Natural History). to clarify the record of an Erionota sp. feeding on oil British Museum (Natural History), London; 1949, 502 pp, palm. 53 plates. 2. Inoue´S, Kawazoe´A. Hesperiid butterflies from South Vietnam (5). Tyo ta Ga 1970;21(1–2):1–14. Available from: URL: http://ci.nii.ac.jp/naid/110007706966. 3. Warren AD. The Higher Classification of the Hesperiidae Acknowledgements (Lepidoptera: Hesperioidea) [Ph.D. thesis]. Oregon State University, Corvalis, OR, USA; 2006, 458 pp. Dr Hideyuki Chiba (Research Associate, Bishop Museum) 4. Warren AD, Ogawa JR, Brower AVZ. Phylogenetic and Dr Laurence G. Kirton (Forest Research Institute of relationships of subfamilies and circumscription of tribes in Malaysia, retired) generously shared information, views the family Hesperiidae (Lepidoptera: Hesperioidea). and documentation. Dr Chiba and Dr Rienk De Jong Cladistics 2008;24:1–35. (National Museum of Natural History, The Netherlands) 5. Warren AD, Ogawa JR, Brower AVZ. Revised classification kindly reviewed parts of the manuscript paper prior to of the family Hesperiidae (Lepidoptera: Hesperioidea) based submission. I thank Dr Feng Zhang (CABI China Regional on combined molecular and morphological data. Systematic Entomology 2009;34:467–523. Centre), Dr Hiroko Kamei (University of Dundee), and Dr Tim Haye (CABI Europe-Switzerland) for translation 6. Igarashi S, Fukuda H. The Life Histories of Asian Butterflies, and interpretation of Chinese, Japanese and German texts Volume 2. Tokai University Press, Tokyo, Japan; 2000, 711 pp. respectively. In addition to the library resources of the 7. Cock MJW, Congdon TCE, Collins SC. Observations on African Butterfly Research Institute (ABRI), the the biology of Afrotropical Hesperiidae (Lepidoptera). Part 6. Hesperiinae incertae sedis: palm feeders. Zootaxa Biodiversity Heritage Library, the British Library, 2014;3831(1):1–61. CABI, and the Royal Entomological Society, I have been helped by a variety of people with reprints and copies 8. De Jong R. Revision of the Oriental genus Matapa Moore (Lepidoptera, Hesperiidae) with discussion of its phylogeny of literature, including Dr Rienk De Jong, Dr Ahsol and geographic history. Zoologische Mededelingen Hasyim (Indonesia), Suli Khing, Dr Lum Keng Yeang and 1983;57:243–70. Dr Soetikno S Sastroutomo (CABI SE Asia), Dr Banpot 9. Bascombe MJ, Johnston G, Bascombe FS. The Butterflies Napompeth (Thailand), Dr Justin Okolle (IRAD/CARBAP/ of Hong Kong. Academic Press, London; 1999, xiii + 422 pp, UB, Cameroon), Dr Don Sands (CSIRO Australia, 222 plates. retired), Dr K. Veenakumari (India), Dr Janny Vos 10. Saji K, Kale P, Kane A. Matapa aria Moore, 1865 – Common (CABI Netherlands), as well as all those scientists and Branded Redeye. In: Kunte K, Kalesh S, Kodandaramaiah U, organizations who have generously made their editors. Butterflies of India, v. 2.10. Indian Foundation for publications and information available via the internet. Butterflies; 2014. Available from: URL: http:// I thank Dr Charles Dewhurst (PNG Oil Palm Research www.ifoundbutterflies.org/sp/599/Matapa-aria [Last accessed 1 December 2014]. Association, retired), Colin Congdon and Forest and Kim Starr (http://www.starrenvironmental.com/) for 11. Igarashi S, Fukuda H. The Life Histories of Asian butterflies, Volume 1. Tokai University Press, Tokyo, Japan; 1997, 547 pp. allowing me to use their images. The images of adult E. torus (Figure 2) are of specimens in Hope Entomological 12. Riley ND. A revision of the continental African species of Collections, Oxford University Museum, and I thank the genus Artitropa (Lep. Hesperiidae). Transactions of the Entomological Society of London 1925;1925:281–8, plate 35. Dr James Logan and Ms Molly Carter who facilitated my visit. The images of adults of E. acroleuca (Figure 3) 13. Dickson CGC, Kroon DM (editors). Pennington’s Butterflies of Southern Africa. A.D. Donker, Johannesburg, South Africa; are of specimens in the collection of the Forest Research 1978, 670 pp. Institute of Malaysia and I thank Phon Chooi-Khim for facilitating my visit there. I also thank Dr Blanca Huertas 14. 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