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Datasheet

Brontispa longissima (coconut hispine beetle)

Summary

  • Last modified
  • 22 June 2017
  • Datasheet Type(s)
  • Pest
  • Natural Enemy
  • Invasive Species
  • Preferred Scientific Name
  • Brontispa longissima
  • Preferred Common Name
  • coconut hispine beetle
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Brontispa longissima (coconut hispine beetle); adult. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
TitleAdult
CaptionBrontispa longissima (coconut hispine beetle); adult. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
Copyright©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
AdultBrontispa longissima (coconut hispine beetle); adult. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult.  Ventral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
TitleAdult
CaptionBrontispa longissima (coconut hispine beetle); adult. Ventral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
Copyright©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult.  Ventral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
AdultBrontispa longissima (coconut hispine beetle); adult. Ventral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult.  Lateral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
TitleAdult
CaptionBrontispa longissima (coconut hispine beetle); adult. Lateral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
Copyright©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult.  Lateral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
AdultBrontispa longissima (coconut hispine beetle); adult. Lateral view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult, head and pronotum. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
TitleAdult
CaptionBrontispa longissima (coconut hispine beetle); adult, head and pronotum. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
Copyright©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU
Brontispa longissima (coconut hispine beetle); adult, head and pronotum. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)
AdultBrontispa longissima (coconut hispine beetle); adult, head and pronotum. Dorsal view. Museum set specimen. (Specimen collected 10 October 1989 Broome, WA T. Fitspatrick ex. palm)©PaDIL/Cameron Brumley DAFWA - CC BY 3.0 AU

Identity

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Preferred Scientific Name

  • Brontispa longissima (Gestro) 1885

Preferred Common Name

  • coconut hispine beetle

Other Scientific Names

  • Brontispa castanea Lea
  • Brontispa froggatti Sharp
  • Brontispa longissima var. javana Weise
  • Brontispa longissima var. selebensis Gestro
  • Brontispa reicherti Uhmann
  • Brontispa simmondsi Maulik
  • Oxycephala longipennis Gestro
  • Oxycephala longissima Gestro

International Common Names

  • English: coconut leaf hispid; new hebrides coconut hispid
  • French: brontispe du cocotier

Local Common Names

  • Netherlands: tweekleurige klapperbladkever

EPPO code

  • BRONLO (Brontispa longissima)
  • BRONSE (Brontispa longissima var. selebensis)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Coleoptera
  •                         Family: Chrysomelidae
  •                             Genus: Brontispa
  •                                 Species: Brontispa longissima

Notes on Taxonomy and Nomenclature

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Maulik (1938) considered the species Oxycephala longissima, O. longipennis, Brontispa froggatti, B. javana [B. longissima var. javana], B. selebensis [B. longissima var. selebensis], B. castanea, B. simmondsi and B. reicherti as colour varieties and hence synonyms of B. longissima. Gressitt (1957) gave a full synonymy for the species. Several physiological races occur, as evaluated by hospitality to the parasite Tetrastichus brontispae (Mo, 1965). Takano et al. (2011, 2013) reported the presence of two cryptic species within B. longissima.

Description

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Eggs

The brown eggs are elliptical, about 1.5 mm long and 1 mm wide, with each end broadly rounded. The slightly convex upper surface has the chorion with a honeycomb sculpturation. They are fixed to the leaf surface by a cementing substance (Maulik, 1938).

Larvae

The larvae are whitish in colour. The first-, second- and final-instar larvae are described in detail by Maulik (1938). The head of the first stage larva is comparatively large compared with the body. The entire cuticle is more densely covered with minute spicules. A seta arises from about the middle of the lateral margin of each thoracic segment, with two setae on each of the abdominal lateral processes. Each process of the tail-shovel bears a large, sharp, curved spine at the inner angle, and a series of five or six setae along the dorsal and ventral margins. The larva is about the same length as the egg, but about 0.75 mm in width.

The second-instar larva resembles the full-grown larva more than the first instar. The lateral abdominal processes are longer, each bearing four setae which are comparatively longer than those of the fully developed larva and situated at different points around the periphery of the apex. There are eight setae on the prothorax (four on each side) and six on the meso- and metathorax (three on each side, two setae on the produced part and one posteriorly). The distinct spine at the inner angle of each prong of the tail-shovel is not prominent as in the first-instar.

The fully developed larva has the body moderately flat, almost parallel-sided, very slightly and gradually narrowed from the prothorax towards the apex, composed of 13 segments (one head, three thoracic and nine abdominal). It is almost 9 mm long and 2.25 mm wide. The anus is situated ventrally on the ninth segment, if the fold at the anal orifice is considered as representing a segment, then the abdomen should be regarded as 10-segmented. The distinct head bears a pair of 2-segmented antennae; a group of five ocelli, three in a line and two in another, situated behind the antennae; a pair of apically bidentate mandibles.

The thoracic segments are broader than they are long, the mesothorax very slightly broader than the prothorax and the metathorax very slightly broader than the mesothorax. The dorsal surface of the prothorax is more strongly sclerotized with a fine median suture and laterally rounded. The meso- and metathorax bears laterally a small knob in the middle, bearing two fine, short setae. There is a pair of well developed 3-segmented legs on each thoracic segment, each terminating in a single claw and fleshy pad-like structure. Each of the first seven abdominal segments is broader than it is long; each of the first three segments is somewhat shorter than the following segments; the eighth and ninth together form the terminal tail-shovel. Each segment bears laterally a moderately long process, ventral to the spiracles. A lateral process is a conical structure projecting horizontally nearly from the middle of the margin, except in the fifth, sixth and seventh segments, in which they appear to arise more from the posterior part. The apical part is subconical and distinctly thinner than the basal part. The entire surface is densely covered with spinules. The apical part bearing three or four setae. The tail-shovel is longer than broad, apically deeply concave, the prongs bent inwards and bluntly-pointed. The upper ridge on each side bearing nine spinules, three smaller on the basal part nearer the large spiracle, four larger on the middle part and two smaller on the bent apical part. There are 4 small spinules on the lower ridge more widely spaced than those on the upper ridge. The upper surface is concave, bearing many irregularly placed transparent areas and the ventral surface is flat. There are 9 pairs of spiracles, one thoracic, between the pro- and mesothorax, situated on a conical structure resembling a lateral process, and one pair dorsally situated on each of the first seven abdominal segments.

The larvae undergo five instars which can be distinguished on the morphometrics ( in mm) of the tail-shovel as follows: L1, 0.33, mean 0.13; L2, 0.47, mean 0.20; L3, 0.65, mean 0.29; L4, 0.82, mean 0.37; L5, 0.94, mean 0.45.

Dorsal habitus views of the first-, second- and final-instar larvae are provided by Maulik (1938).

Generic and specific keys to separate the larvae of Brontispa species are provided by Gressitt (1963). The key characters are: body not oval with a continuous margin; head visible from above; last abdominal segment with a caudal process; not leaf-mining; meso- and metathoracic segments lacking lateral processes; caudal process with arms widely separated basally, thickened or strongly angulate posteriorly; lateral abdominal processes short and blunt, rarely last two longer; spiracle of last segment elliptical (Generic); lateral abdominal processes subequal, eighth shorter than greatest width of an arm of caudal process; emargination of caudal process usually broader than long; arms of caudal process parallel-sided externally, at least in central part; emargination of caudal process reaching about half way from apices of arms to spiracles; emargination of caudal process not much broader than long, broadly oval, widest in middle; arm curved and subacute apically; eighth abdominal process shorter than preceding (Specific).

Pupae

The pupae of several Brontispa spp. including B. longissima are described by Maulik (1938). The head bears three processes, one median and one on each side. Each lateral process is fleshy, broadest basally, pointed apically, pre-apically bearing small blunt spine. Thorax with meso- and metanotum each bearing a pair of wings. The abdomen is 9-segmented, the eighth and ninth segments are fused. Each of the first six segments bears laterally a pair of spiracles opening dorsally. Each segment bearing spinules arranged as follows: no spinules on first segment; from segments 2 to 7, two groups of four in transverse line, one nearer the basal margin and the other nearer the apical, those of the basal group are comparatively larger and more widely-spaced, those of the apical line are situated in the intervals of the spinules of the basal group; on segment 8 only two in transverse line near the basal margin. Dorso-laterally bearing a longitudinal series of spinules, one on each of the first seven segments, situated a little posterior to and more inward than the corresponding spiracle (on the seventh in a similar position). Anterior to each spiracle on segments 3 to 6 and in a similar position on 7 is a spine. Laterally on segments 1-7 bearing a group of closely placed spinules bearing moderately long setae, the spinules being less prominent on segments 1-2, or 3. Ventrally, only segments 4-7 bearing spinules. The prongs of the tail-shovel are more slender, somewhat longer than those of the larva, and also lack lateral spines or setae. The last larval exuvium is always retained on the prongs of the tail-shovel.

Adults

The adults 8.5-9.5 mm long, 2.00-2.25 mm wide; length of antenna, 2.75 mm. The males are slightly smaller than females (Maulik, 1938). Their colour varies geographically from reddish-brown in Java, to almost black in the Solomon Islands and Irian Jaya. Some specimens have the elytra brown or black, or have a spindle-shaped black marking on the elytral suture. Considerable overlapping of these forms, which were long regarded as distinct species, occurs (Lever, 1969). Maulik (1938) examined the male genitalia, especially the median lobes of the different colour varieties, and found no differences. However, the median lobes of distinct species, for example B. linearis and B. longissima, exhibit obvious structural differences.

Maulik (1938) and Gressitt (1963) have provided keys to separate the adults of Brontispa species. The relevant key characters for B. longissima are: antennae not serrate; central portion of head usually parallel-sided, broader than long; rostrum more than half as long as first antennal segment; pronotum flattish, shiny, with several large impunctate areas; body quite flat and narrow; prothorax laterally distinctly concave; anterior lateral angles of pronotum expanded, expanded portion broadly-rounded, constricted behind, without a minute projection at inner angle.

Distribution

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The coconut leaf hispa was originally described from the Aru Islands. It is native to Indonesia, possibly to Irian Jaya, and also to Papua New Guinea, including the Bismarck Archipelago, where it seldom causes serious problems. It was reported from the Solomon Islands in 1929 and from Vanuatu in 1937 by Risbec (1942) who stated that it had been present in New Caledonia for several years. Cohic (1961) was the first to record B. longissima from New Caledonia (Tahiti), Long (1974) for American Samoa and Anon (1981b) for Western Samoa. It is also present in northern Australia (Fenner, 1984) and Taiwan (Shiau, 1982).

It is not known from the Cook Islands, Fiji, Kiribati, Niue, Tokelau, Tonga and Tuvalu (Waterhouse, 1985).

B. longissima was detected for the first time in Hong Kong in 1988 infesting 30 petticoat palms in a nursery (Lau, 1991). This outbreak was eradicated, but in late 1991, a few mature coconuts (>30 years old) were affected, indicating that the pest may be established in Hong Kong. Since most of the locally grown ornamental palms, including petticoat palms, king palms and dwarf date palms were imported from China in recent years, it is suspected that B. longissima was introduced from China, probably from the Shenzhen area of Guangdong Province.

Recently, B. longissima has spread to Singapore, Vietnam, Nauru, Cambodia, Laos, Thailand, Maldives, Myanmar and Hainan Island, China (Rethinam and Singh, 2007). It is feared that it will spread from the Maldives to Sri Lanka and southern parts of India.

The distribution map includes some records based on specimens of B. longissima from the collection in the Natural History Museum (London, UK): dates of collection are noted in the List of countries (NHM, various dates).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

Bismarck ArchipelagoPresentNativeRethinam and Singh, 2007
CambodiaPresentRethinam and Singh, 2007; EPPO, 2014
ChinaRestricted distribution, ; EPPO, 2014
-GuangdongPresent1999Zhang et al., 2004; EPPO, 2014
-GuangxiPresentEPPO, 2014
-HainanPresentRethinam and Singh, 2007; EPPO, 2014
-Hong KongPresent, ; Lau, 1991; EPPO, 2014
-MacauPresentMo et al., 2006
IndonesiaRestricted distributionNative, ; Rethinam and Singh, 2007; EPPO, 2014
-Irian JayaPresent, ; THOMAS, 1962; Waterhouse and Norris, 1987; EPPO, 2014
-JavaPresent, ; , ; Mo, 1965; Kalshoven and Laan, 1981; EPPO, 2014
-MoluccasPresent, ; , ; , ; Lepesme, 1947; EPPO, 2014
-Nusa TenggaraPresentEPPO, 2014
-SulawesiPresent, ; , ; Woodroof, 1979; Kalshoven and Laan, 1981; EPPO, 2014
JapanPresentEPPO, 2014
-Ryukyu ArchipelagoPresentTakasu et al., 2010; EPPO, 2014
LaosPresentRethinam and Singh, 2007; EPPO, 2014
MalaysiaAbsent, unreliable recordKalshoven and Laan, 1981; EPPO, 2014
MaldivesPresentRethinam and Singh, 2007; EPPO, 2014
MyanmarPresentRethinam and Singh, 2007; EPPO, 2014
PhilippinesPresent===, 2007; EPPO, 2014
SingaporePresent, ; APPPC, 1987; AVA, 2001; Rethinam and Singh, 2007; EPPO, 2014
TaiwanPresent, ; Shiau, 1982; Chiu and Chen, 1985; Chiu et al., 1985; EPPO, 2014
ThailandPresentRethinam and Singh, 2007; EPPO, 2014
VietnamPresentRethinam and Singh, 2007; EPPO, 2014

Africa

MadagascarAbsent, unreliable recordEPPO, 2014
MauritiusAbsent, unreliable recordEPPO, 2014
SeychellesAbsent, unreliable recordEPPO, 2014

Oceania

American SamoaPresent, ; Long, 1974; Waterhouse and Norris, 1987; EPPO, 2014
AustraliaRestricted distribution, ; EPPO, 2014
-Australian Northern TerritoryPresentAnon, 1981a; , ; Jones and Elliot, 1986; EPPO, 2014
-QueenslandPresentAnon, 1981a; , ; , ; Jones and Elliot, 1986; EPPO, 2014
French PolynesiaPresent, ; Cohic, 1961; EPPO, 2014
NauruPresentRethinam and Singh, 2007; EPPO, 2014
New CaledoniaPresent, ; Risbec, 1942; Cochereau, 1969; EPPO, 2014
Papua New GuineaPresentNative, ; Hassan, 1972; Waterhouse and Norris, 1987; Rethinam and Singh, 2007; EPPO, 2014
SamoaPresentAnon, 1981b; , ; Stechmann and Semisi, 1984; EPPO, 2014
Solomon IslandsPresent, ; Risbec, 1942; Brown and Green, 1958; EPPO, 2014
VanuatuPresent, ; , ; , ; Risbec, 1942; EPPO, 2014
Wallis and Futuna IslandsPresent, ; Waterhouse and Norris, 1987; EPPO, 2014

Hosts/Species Affected

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The host range of B. longissima includes various Palmae [Arecaceae]. In Papua New Guinea, coconut, sago palms, areca or betel palm (Areca catechu), royal palms (Roystonea regia), oil palm and ornamental palms are attacked. In northern Australia, hosts include areca palms (A. catechu), nicobar palm (Bentinckia nicobarica), carpentaria palm (Carpentaria acuminata) and fish tail palm (Caryota mitis). In Hong Kong, it is also reported from ivory nut palm (Phytelephas), petticoat palm (Washingtonia robusta), king palm (Archontophoenix alexandrae) and dwarf date palm (Phoenix roebelenii) (CSK Lau, Agriculture and Fisheries Department, Hong Kong, personal communication, 1992).

Host Plants and Other Plants Affected

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Plant nameFamilyContext
Areca catechu (betelnut palm)ArecaceaeOther
Cocos nucifera (coconut)ArecaceaeMain
Elaeis guineensis (African oil palm)ArecaceaeOther
Metroxylon sagu (sago palm)ArecaceaeOther
Phoenix roebelliniArecaceaeOther
Washingtonia robusta (mexican washington-palm)ArecaceaeOther

Growth Stages

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Symptoms

Top of page Both adults and larvae damage the leaflets of young unopened fronds. They graze away the leaf surface in streaks, which are typically parallel to the midrib. The narrow feeding scars enlarge to form irregular, brown blotches as the frond opens. The brown areas shrivel and curl, giving the leaf a characteristic scorched, ragged appearance.

Large areas of the leaflets break off leaving the foliage partially skeletonized and its effective photosynthetic tissue may in extreme cases be reduced to virtually nothing. As the leaflets separate when the fronds expand, the adults move to attack younger leaves. They leave narrow chewing marks, which are individually less damaging than larval feeding. O'Connor (1940) proposed that adults cause more damage than larvae due to the longer duration of this stage, but Tothill (1929) considered that the larvae were more destructive. Destruction of young leaf spike tissues restricts growth for a long time and heavy attack may cause death. Palms weakened by attack are more susceptible to drought and disease (Waterhouse and Norris, 1987).

The shorter interval between the opening of successive fronds on mature palms, and their larger size, contribute greatly to their relative freedom from attack. A mature palm may be expected to produce a new frond every 4-5 weeks. In young palms, after the rapid emergence of the first few seedling leaves, the fronds are produced less frequently and over the first 4-5 years of its life a young palm will, on average, produce a new frond only once in every 6-7 weeks; under adverse conditions the interval may exceed 8 weeks. Thus, on a young palm there is never more than one frond vulnerable to attack at any one time, and there is usually an interval between the opening of consecutive fronds when there is not one at a vulnerable stage. This results in all immigrant beetles becoming concentrated on those palms which do have a vulnerable frond, and the relatively small size of the fronds renders the damage more devastating.

The fronds are larger in mature palms and there may be more than one infested frond at any one time; so that, although the number of beetles may be greater, there is less damage. The rate of frond production gradually speeds up after the palm is about 5 years old, and, by the time it is 8-10 years old, the palm will be producing new fronds at the normal interval of one every 4-5 weeks. It is significant that, at this age, healthy, vigorous palms achieve relative immunity to severe beetle attack.

List of Symptoms/Signs

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Leaves

  • external feeding

Biology and Ecology

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Waterhouse and Norris (1987) briefly reviewed the biology of B. longissima from the studies of Maulik (1938), O'Connor (1940), Froggatt and O'Connor (1941), Smee (1965) and Maddison (1983). Kalshoven (1981) provided biological and ecological information based on the work of Franssen and Mo (1952) in Indonesia.

The eggs are laid in groups of one to four, end to end, in a furrow chewed in the leaf by the adult, between or inside the tightly folded leaflets. The beetle covers each egg with excreta. The eggs hatch after an incubation period of about 5 days.

The newly hatched larva begins to feed between and inside unopened leaflets. The number of instars varies from five to six. The larvae are fairly sedentary and avoid light. The larval period is 30-40 days, followed by a prepupal period of 3 days and a pupal period of 6 days. The pupa lies freely between the apposed surfaces of the developing folded leaflets. The development from egg to adult takes 5-7 weeks.

The beetles, which also seem to avoid light, are nocturnal and fly well (Kalshoven, 1981). The adults feed among the young unopened leaflets and, since they live up to 220 days, their cumulative damage greatly exceeds that of the larvae. There is a preoviposition period of 1-2 months and 100 or more eggs may be laid.

There are about three overlapping generations per year.

Dry periods favour the development of Brontispa populations. This conclusion is based on the number of damaged leaves in coconut crowns, which represents the extent of infestation to the central axis (Kalshoven, 1981). Strong monsoon winds reduce the influence of parasitic wasps. No damage occurred in regions of West Java with high rainfall, although the beetle has been found in these areas (Kalshoven, 1981).

The beetle is only capable of weak flight, so new infestations spread slowly, unless aided by human activity (Maddison, 1983).

In Papua New Guinea, Brontispa is usually controlled by natural enemies, but sometimes it can become a pest (Smee, 1965).

Natural enemies

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Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
Asecodes hispinarum Parasite Larvae Maldives, Nauru, Thailand, Vietnam, Western Samoa Cocos nucifera
Beauveria bassiana Pathogen
Chelisoches morio Predator Larvae/Pupae
Chrysonotomyia Parasite Larvae
Hispidophila brontispae Parasite Eggs
Metarhizium anisopliae Pathogen Adults/Larvae/Pupae
Oecophylla smaragdina Predator Larvae/Pupae
Ooencyrtus pindarus Parasite Eggs
Ooencyrtus podontiae Parasite
Tetrastichus brontispae Parasite Larvae/Pupae American Samoa; Australia; Marquesas Islands; New Caledonia; New Guinea; Papua New Guinea; Solomon Islands; Sulawesi; Tahiti; Taiwan; Vanuatu; Western Samoa Cocos nucifera
Trichogrammatoidea nana Parasite Eggs Solomon Islands Cocos nucifera

Notes on Natural Enemies

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Three wasp parasites of B. longissima are known in Java. Two of these are egg parasites: the trichogrammatid Hispidophila brontispae; and the encyrtid Ooencyrtus pindarus. One H. brontispae wasp develops per Brontispa egg, producing about 15-17% parasitism (Kalshoven, 1981; Waterhouse and Norris, 1987), and O. pindarus produces about 10% parasitism (Kalshoven, 1981). The eulophid, Tetrastichus brontispae, which is found in 60-90% of the pupae (Awibowo, 1934) and 10% of the larvae, develops in 18 days; about 20 specimens emerge from one Brontispa pupa.

Parasitized larvae may die before pupation, but parasites still emerge. However, the level of parasitization by T. brontispae is not always this high and Lange (1950) recorded an average of only 16% in pupae. The life cycle of T. brontispae is 16-21 days; at least two generations may be produced to each one of B. longissima (Lever, 1936a, b; Lange, 1953).

Two native wasp parasites are known in the Rabaul district of Papua New Guinea: the non-specific egg parasite, Trichogrammatoidea nana, and the eulophid larval parasite, Chrysonotomyia sp. [possibly Asecodes hispinarum]. A large percentage of Brontispa eggs are attacked by T. nana, which has also been bred from Brontispa eggs in the Solomon Islands. Chrysonotomyia sp. is comparatively rare.

Occasionally, Brontispa larvae have been killed by a bacterial disease (Froggatt and O'Connor, 1941; O'Connor, 1940) or by the fungus Metarhizium anisopliae in Papua New Guinea (Waterhouse and Norris, 1987). A fungus which was found to affect larvae, pupae and adult Brontispa (Maddison, 1983) may have been M. anisopliae, which is known to attack Brontispa. M. anisopliae caused 15-20% mortality of both adults and larvae in American Samoa (Waterhouse and Norris, 1987).

In Australia large numbers of torn, empty egg shells of Brontispa have been found in a nest of the ant, Tetramorium simillimum, but the significance of this ant in influencing numbers of the pest is unknown (Fenner, 1984). The ant Pheidole megacephala attacks T. brontispae on New Caledonia and both young larvae and parasitized pupae of Brontispa (Cochereau, 1965).

The earwig Chelisoches morio has been reported as a predator of B. longissima in Vanuatu (Risbec, 1933).

Impact

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Brontispa attacks palms of all ages, although it is most damaging to young palms in nurseries and for the first 4-5 years after planting out in the field, especially in dry areas. Neglected palms are more heavily attacked than those kept free from undergrowth (Froggatt and O'Connor, 1941; Maddison, 1983).

Coconut plantations in South Sulawesi, Indonesia, which were in poor condition due to poor soil conditions, infestations by aleurodids and inadequate maintenance, were more susceptible to attack by B. longissima. Severe Brontispa attacks were reported in nearly all regions of south-east Sulawesi in 1929. The hispid sometimes occurred together with Aleurodicus, Oryctes and palm weevils, which together killed numerous palms, while other trees were in such poor condition that they did not produce fruit for many years. The outbreaks continued until between 1935 and 1937 when the situation greatly improved due to the plants having older, fully developed, more resistant foliage. In later years, the incidence of Brontispa was comparable with that for Java; 10-15% of the trees in certain localities showing injury. The effect of the severe infestations was felt for a number of years because growers ceased to maintain their plantations during the outbreak years (Kalshoven, 1981).

In North Sulawesi, the main damage caused by Brontispa occurs on young, 2-3-year-old palms, which are not yet fruiting. This infestation may supersede earlier attacks by Plesispa. Progressively smaller numbers of Brontispa are found in 5-10-year-old trees. When 8-10-year-old trees start to fruit, traces of previous infestation are still visible, but there is generally no further damage. It is thought that the heart leaves of older trees become firmer and gradually less suitable as a breeding place for the pest and are, therefore, no longer penetrated by the beetles. Palms grown in poor conditions, which have a less compact heart, are more susceptible to attack (Kalshoven, 1981).

Light attacks result in minor leaf injury, and a slight decrease in fruiting at the axils of the damaged leaves. Fruit production is significantly reduced if eight or more leaves are destroyed. Under prolonged outbreak conditions, such as those which occurred in South Sulawesi for several years, fruit-shedding takes place, newly-formed leaves remain small, the trees appear ragged and may ultimately die.

Outbreaks of Brontispa occur quite regularly in east Java, especially near Blitar, where about 55,000 trees were damaged in three districts during the dry season of 1940. Brontispa is also a regular pest in Besuka, especially north of Banjuwangi (near Giri) where the climate is rather dry. Outbreaks have also been reported from Madura (Kalshoven, 1981). B. longissima causes widespread and serious damage to unopened leaves of coconut in Java (Mo, 1965).

Brontispa attacks in North Sulawesi were usually less severe than those in the south of the island. Damage was also reported from the Sangihe Islands, on Ceram, on the islands Banggai and Labolo of the Banggai group and on the Aru islands (in association with Plesispa). Brontispa additionally occurred in Bali, but was of little significance. In 1936, Adonaria Island, east of Flores, Indonesia, became a focus for Brontispa attacks, from which the pest migrated to Flores in later years causing particular damage to coconuts in valleys under humid conditions. No trees died, but there was loss of yield. The outbreak ended in 1939.

B. longissima is the most serious pest of young palms in the Solomon Islands; at a time when considerable areas of young palms existed, it was estimated that damage to the value of £65,000 had been suffered in less than 10 years on Lever's Estates alone (Tothill, 1929). In extreme cases, damage can completely arrest the development of young palms and may even kill them (Brown and Green, 1958). Only occasionally are mature palms attacked on a large enough scale in the Solomon Islands to cause serious damage.

Detection and Inspection

Top of page Young coconut palms should be inspected for eggs between or inside the tightly folded leaflets and early feeding damage of the larvae between and inside unopened leaflets, where browning and death of the surrounding tissues can be seen.

Prevention and Control

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Cultural Control

A surgical method of control has been attempted in the Solomon Islands; this involved cutting out and destroying the central unopened frond which harbours the pest (Brown and Green, 1958). This procedure must be conducted over a large area at one time to reduce re-infestation from neighbouring palms and must be repeated fairly often to be effective. Palms which were 3-6 years old could stand the loss of one leaf every 6 months, but younger palms could not as this caused too great a reduction in growth rate (Tothill, 1929). However, this method is expensive and will not greatly affect the Brontispa population as a whole unless mature palms are also treated (Tothill, 1929).

Mechanical control of the pest by removing affected heart leaves is laborious and has very little effect (Kalshoven, 1981).

Chemical Control

Chemicals used to control the pest must reach the insect in the narrow crevices between the leaflets and chemical treatment must be maintained throughout the year because B. longissima breeds continuously, with several generations a year. If beetles start breeding as soon as they reach the young palms, larvae will probably start appearing after about 1 week; if there has been any residual effect from insecticide treatment, or if the invasion rate is slow, it will be considerably longer before the larval population presents a serious risk (Brown and Green, 1958).

The application of chemicals at 10-day intervals was more effective than 3-weekly applications. Frequent applications of high volumes of spray, in excess of that required to provide satisfactory levels of control, caused slight phytotoxicity; growth and the production of new fronds was retarded. Satisfactory control can be achieved at low cost using a fine, low-volume spray applied from above to the central spike of each individual palm (Brown and Green, 1958).

Damage caused by Brontispa is considered serious enough for chemical control measures to be used on young palms. Chemicals recommended are carbaryl and trichlorfon (Maddison, 1983; Smee, 1965; Stapley, 1972, 1980a; Wu and Tao, 1976). The pesticide is applied to the central spike of the palm. Trichlorfon eliminated Brontispa from isolated areas of young palms in Western Samoa (Bourke, 1981). Monthly spraying of young coconuts with permethrin is also advocated in Western Samoa (Hollingsworth et al., 1986; Peters et al., 1984).

In Australia, Jones and Elliot (1986) recommended spraying the unopened fronds thoroughly with carbaryl; repeated applications were necessary as new fronds emerged.

Insecticide resistance in B. longissima has been documented by Georghiou and Lagunes-Tejeda (1991).

Biological Control

The history of biological control of this pest in Indonesia and the Pacific from 1932 to 1984 is reviewed by Waterhouse and Norris (1987). The prospects for control are also discussed.

An outbreak of B. longissima in Sulawesi, Indonesia, in 1932, led to the importation of Tetrastichus brontispae from Java. Field recoveries made the following year and a few years later, showed substantially reduced injury to coconut plantations by the beetle (Franssen and Mo, 1952). Attempts to establish this parasite in east Java between 1932 and 1937, and in central Java in 1954, were unsuccessful (Rao et al., 1971). Mo (1965) provided one possible explanation for these failures and the variation in the effectiveness of the parasite in different areas; strains from east and central Java could only be propagated on the strain of B. longissima from which they originated. In one strain, almost all of the parasite larvae died within 3-4 days of hatching and the host also succumbed a few days later. Phagocytic encapsulation and melanization were occasionally noted but neither was the primary cause of death.

T. brontispae was introduced from the Solomon Islands to Papua New Guinea in 1939 because Brontispa occasionally caused damage. The parasite became established but the impact of this introduction has not been recorded (O'Connor, 1940).

T. brontispae was introduced and established in Australia in 1984 (Waterhouse and Norris, 1987).

Early attempts to establish T. brontispae in the Solomon Islands from Java and Sulawesi were unsuccessful (Lever, 1936 a,b). Later releases of T. brontispae and Trichogrammatoidea nana in the 1930s resulted in establishment, but adequate control was not attained (Johns, 1941). T. brontispae was re-introduced in 1968 and released in the Russell island group; it became established and widespread. The introduction reduced the incidence of Brontispa from 95% to 5% of newly planted Federal Malay States palms, and the spraying of young palms is now seldom necessary (MacFarlane, 1981; Stapley, 1971, 1972, 1980a). The parasitization rate reached 100% of the pupae found in some plantations (Stapley, 1971).

Attempts have been made to use the green tree ant, Oecophylla smaragdina, in biocontrol of B. longissima in the Solomon Islands. However, Oecophylla does not alleviate the Brontispa problem in highly susceptible young palms because they usually do not colonize palms before they flower: the honeydew produced by scale insects and mealybugs, which colonize palms after flowering, is a powerful attractant (Stapley, 1980b).

T. brontispae was first released in French Polynesia in 1962 and effective biological control was established throughout the Society Islands after several years, with parasitization rising to 38% (Stapley, 1971). Mass releases of T. brontispae began on Tubuai Island and Rangiroa Island in July, 1984; it is now well established and coconut trees are beginning to recover from serious damage caused by B. longissima in 1981 and 1983 (Gourves and Samuelson, 1979).

T. brontispae was introduced to New Caledonia from Saipan in the Northern Mariana Islands in 1963; it soon became well established with reports of up to 24% parasitization of B. longissima. However, even a combination of this parasite, a fungal disease, and the earwig predator, Chelisoches morio, did not provide satisfactory control of the pest (Cochereau, 1969).

After four unsuccessful attempts to establish T. brontispae in American Samoa, large numbers of the parasite were sent from Western Samoa in 1985; the species is now established (Waterhouse and Norris, 1987). However, no parasitoids were found in a survey of American Samoa conducted by Vögele and Zeddies (1990).

T. brontispae was introduced to Western Samoa from New Caledonia in 1981 and mass-reared on a wide scale (FAO, 1981). In 1981 and 1982, a strain of T. brontispae from the Solomon Islands (originally from Tahiti) was mass-reared and released, as was a strain from Papua New Guinea in 1983. In areas of severe attack in 1981, even 50-70-year-old coconut palms were seriously damaged; all the central leaves were brown and there were no young fronds. By late 1982, most of the trees had recovered substantially. However, Brontispa is still spreading slowly into new areas on both Upolu and Savai'i, Western Samoa, and is causing serious damage in these areas.

Four coconut plantations in Western Samoa were sampled in 1985 for natural mortality factors (Hollingsworth et al., 1988). The eulophid, Chrysonotomyia sp. [possibly Asecodes hispinarum], was the most important cause of larval mortality, parasitizing 75% of fourth-instar larvae collected from two plantations. The pupal parasite, T. brontispae, was recovered from one plantation but was not an important cause of mortality. An entomogenous fungus, thought to be Metarhizium anisopliae, was found on adults in three plantations sampled. The fungus infected approximately 65% of third- and fourth-instar larvae, and 27% of adults, in one plantation. Chelisoches morio was common in infected coconut spears (averaging 5.1 nymphs and 1.2 adults per tree) and fed on larvae and pupae of B. longissima in the laboratory.

In 1987, Vögele (1989) carried out a study of the mortality factors in 120 subpopulations of B. longissima in Western Samoa. Asecodes sp. [possibly A. hispinarum] was the dominant parasitoid in the field, present in 37% of the samples. T. brontispae was recovered from only 3% of samples and was not an important mortality factor. An extensive survey of damage to 37,000 trees showed that B. longissima was under control and did not cause any significant yield losses; initial production losses were estimated to be as high as 50-70%. The benefit:cost ratio of this project was 3.9:1 during the implementation period (1981-1986) and 9.9:1 for the period 1987-1990. The internal rate of return for 1981-1990 exceeded 40%; the project was highly successful.

A steady decline in damage was recorded in Western Samoa from 42.4% in 1984 to 15.4% in 1987, following the release of T. brontispae and Asecodes sp. [possibly A. hispinarum] during 1981-1986 (Vögele and Zeddies, 1990). In contrast, approximately 74% of all palms in American Samoa were infested compared with only 14.3% in Western Samoa. B. longissima infected by Metarhizium sp. were found on 12.5% of sampled trees.

An unidentified eulophid larval parasite, possibly A. hispinarum, from Papua New Guinea was released in 1982 in Western Samoa. In 1984, Brontispa larvae were found which were parasitized by another Chrysonotomyia sp.; it also occurred on the island of Savai'i where Brontispa was not present at the time of the Papua New Guinea introduction, and so it may be a native species. An additional, slightly larger eulophid larval parasite, with alternating black and white stripes on the abdomen, was discovered soon afterwards attacking Brontispa larvae. This parasite is less common than the Chrysonotomyia sp. and may be the species introduced from Papua New Guinea.

In Western Samoa, Chrysonotomyia parasitized an average of 96% of the fourth-instar larvae in one plantation and Metarhizium infected about 65% of third- and fourth-instar larvae, and 27% of adult beetles in another (Waterhouse and Norris, 1987).

The larval parasitoid, A. hispinarum was collected in Western Samoa in 2003 and introduced into Maldives, Nauru, Thailand and Vietnam (Rethinam and Singh, 2007).

A biological control programme for B. longissima using T. brontispae is currently in progress in Taiwan (Chiu and Chien, 1989).

In 1984, an undescribed heterocoptid mite from Papua New Guinea and the Solomon Islands was introduced to, and became established in, Western Samoa. It is thought to be a predator and many individuals are capable of killing a newly moulted adult Brontispa (Waterhouse and Norris, 1987).

Brontispa can be controlled on young palms by spraying suspensions of M. anisopliae. The fungus is capable of spreading rapidly during wet weather, killing more than half of the Brontispa larvae and adults present (Waterhouse and Norris, 1987).

Two field trials of microbial control of B. longissima were conducted in the Pingtung area of Taiwan in 1986 and 1987. The pest could not be detected after three applications of M. anisopliae-1, formulated as a homogeneous biomass, in granules or in a conidial suspension (Liu et al., 1989; Liu, 1994).

There is merit in considering the introduction of suitable strains of this parasite from Java or from other countries where they are now established, where B. longissima is an important pest and where T. brontispae is not present or produces low mortality. Experience in American Samoa, demonstrated that it is necessary to release substantial numbers of the parasite into suitable populations of Brontispa to ensure establishment. The compatibility of strains of Tetrastichus should be evaluated before release, using a strain or strains of Brontispa collected from the country which is contemplating introduction (Waterhouse and Norris, 1987).

The presence of two cryptic species within B. longissima, one distributed over a limited area (Australia, Papua New Guinea, East Timor and Sumba Island) and the other collected from a wide area of Asia and the Pacific region, should also be considered when natural enemies are introduced (JIRCAS, 2012).

Host-Plant resistance

Coconut varieties in the Solomon Islands vary considerably in their susceptibility to Brontispa attack and planting introduced, susceptible cultivars may have upset the previous balance in some areas. Rennell, a variety from the isolated Renell island in the Solomon Islands, is scarcely attacked, whereas varieties from Malaysia such as FMS (Federated Malay States) and dwarf red and yellow palms are highly susceptible. The susceptibility of Malaysian dwarfs is carried into their hybrids. Some varieties from the Ivory Coast and Fiji also show high degrees of resistance (Stapley, 1980a, 1981). Of six local coconut cultivars were tested in Western Samoa, five were highly susceptible, and green dwarf was fairly resistant (FAO, 1983).

Integrated Pest Management

Young palm trees are sprayed with a suspension of M. anisopliae and permethrin in Western Samoa (Waterhouse and Norris, 1987).

B. longissima has been controlled in Taiwan by the use of parasites in conjunction with insecticides (Chang, 1991).

References

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